The Bionomics of Blister Beetles of the Genus Meloe and a Classification of the New World Species
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SELANDER ILLINOIS BIOLOGICAL MONOGRAPH 42 UNIVERSITY OF ILLINOIS PRESS 252 001 The Bionomics of Blister Beetles of the Genus Meloe and a Classification of the New World Species JOHN D. PINTO and RICHARD B. SELANDER ILLINOIS BIOLOGICAL MONOGRAPHS UNIVERSITY OF ILLINOIS PRESS ILLINOIS BIOLOGICAL MONOGRAPHS Volumes 1 through 24 contained four issues each and were available through subscription. Beginning with number 25 (issued in 1957), each publication is numbered consecutively. No subscriptions are available, but standing orders are accepted for forthcoming numbers. Prices of previous issues still in print are listed below, and these may be purchased from the University of Illinois Press, Urbana, Illinois. Microfilm and photo-offset copies of out-of-print titles in the Illinois Biological Monographs are available from University Microfilms, Inc., 313 North First Street, Ann Arbor, Michigan 48107, and the Johnson Re- print Corporation, 111 Fifth Avenue, New York, New York 10003. Balduf, W. V. (1959): Obligatory and Facultative Insects in Rose Hips. 12 pis. No. 26. $3.50. Brandon, Ronald A. (1966): Systematics of the Salamander Genus Gryinophilus. 23 figs. No. 35. $4.50. Campbell, John M. (1966): A Revision of the Genus tofaopoda (Coleoptera: Alleculidae) in North America and the West Indies. 174 figs. No. 37. $5.75. Levine, Norman D., and Ivens, Virginia (1965): The Coccidian Parasites (Protozoa, Sporozoa) of Rodents. 2 figs. 48 pis. No. 33. $7.50. List, James Carl (1966): Comparative Osteology of the Snake Families Typhlopidae and Leptotyphlopidae. 22 pis. No. 36. $3.75. Morgan, Jeanne (1959): The Morphology and Anatomy of American Species of the Genus Psaronius. 82 figs. No. 27. $3.00. Paolillo, Dominick J., Jr. (1963): The Developmental Anatomy of Isoetes. 26 figs. 19 pis. No. 31. $2.50. Ray, James Davis, Jr. (1956): The Genus Lysimachia in the New World. 20 pis. 11 maps. Vol. 24, Nos. 3-4. $2.50. Selander, Richard B. (I960): Bionomics, Systematics, and Phylogeny of Lytta, a Genus of Blister Beetles (Coleoptera, Meloidae). 350 figs. No. 28. $4.50. Selander, Richard B., and Mathieu, Juan M. (1969): Ecology, Behavior, and Adult Anatomy of the Albida Group of the Genus Epicauia (Coleoptera, Meloidae). 60 figs. No. 41. $5.95. Stannard, Lewis J., Jr. (1957): The Phylogeny and Classification of the North American Genera of the Suborder Tubulifera (Thysanoptera). 14 pis. No. 25. $2.50. Unzicker, John D. (1968): The Comparative Morphology and Evolution of the Internal Female Reproductive System of Trichoptera. 14 pis. No. 40. $3.95. The Bionomics of Blister Beetles of the Genus Meloe and a Classification of the New World Species Adult female of Meloe laevis feeding on a stem of Tradescantia. The Bionomics of Blister Beetles of the Genus Meloe and a Classification of the New World Species JOHN D. PINTO and RICHARD B. SELANDER ILLINOIS BIOLOGICAL MONOGRAPHS 42 UNIVERSITY OF ILLINOIS PRESS URBANA, CHICAGO, AND LONDON Board of Editors-. James E. Heath, Willard Payne, H. H. Ross, Richard B. Selander, and P. W. Smith. This monograph is a contribution from the Department of Entomology, University of Illinois. Issued January, 1970. © 1970 by the Board of Trustees of the University of Illinois. Manufactured in the United States of America. Library of Congress Catalog Card No. 71-94400. 252 00081 1 Dedication This work is dedicated to the memory of the late distinguished ento- mologist, Dr. Edwin C. Van Dyke, who studied Meloe four decades ago. CONTENTS BIONOMICS 6 TAXONOMY 92 LITERATURE CITED 181 ADDITIONAL FIGURES 191 INDEX 213 INTRODUCTION The genus Meloe, with 146 described species, is one of the larger genera of blister beetles or Meloidae. In the adult stage blister beetles are soft-bodied, long-legged, often brightly colored insects of generally moderate size for Coleoptera. All are heteromerous and have a hypog- nathous head that is distinctly delimited from the pronotum by a narrow neck region. Most have long elytra covering fully developed, functional hind wings, but in Meloe and several other taxa the elytra are abbreviated and the hind wings have been lost altogether. Loss of the organs of flight has apparently evolved independently in several lines of blister beetles. Besides Meloe, there are several flightless genera or species groups in the subfamily Meloinae and others in the subfamily Nemognathinae. Most blister beetles are phytophagous as adults; a few do not feed in the adult stage. Flowers and leaves of a wide variety of forbs are the main sources of food, but grasses and woody plants are also at- tacked. In the larval stage most genera of blister beetles, including Meloe, feed on the provisions and immatures of wild bees. A few genera attack the eggs of grasshoppers. Although in some cases the active first instar larva searches for its food source directly, in Meloe and many other genera associated with bees the first instar larva reaches its feeding site by phoresy on the adult of the host insect. 1 THE BIONOMICS OF BLISTER BEETLES The family Meloidae is best developed in dry temperate and tropical regions, where large aggregations of adults often form a conspicuous element of the fauna. Meloe is represented in these regions but is rather exceptional in possessing an additional, distinctly boreal element. The present work treats the bionomics of the genus Meloe and the taxonomy of the New World species. In addition to presenting an extensive survey of the literature, the bionomic portion incorporates the results of original investigations. These were concentrated largely on five species of Meloe occurring in Illinois. General areas of investi- gation were adult nonsexual and sexual behavior, interspecific and other ecological relationships, and postembryonic ontogeny. The taxonomic portion of the work is devoted largely to the recog- nition and characterization of the New World species of the genus. Unfortunately, the Old World representation of Meloe has never been satisfactorily described or classified. Thus, while we have attempted to achieve generality in our classification, inadequacies of the literature and the fact that relatively small amounts of material were available from the Old World have prevented our formulating a comprehensive, definitive classification of the genus. Although the species of Meloe occurring in Canada and the United States were revised taxonomically by E. C. Van Dyke in 1928, a dearth of material at that time and unsatisfactory association of type speci- mens with working material resulted in much confusion concerning the identity of the species. Additional taxonomic difficulties arose from the fact that several of the anatomical characters on which Van Dyke relied heavily are highly variable intraspecifically. In the present work seven species are described as new and five names are placed in synonymy, yielding a total of 23 recognized species in the New World. The classification is based largely on the anatomy of adults and first instar larvae. Behavioral characters have been utilized also, but these are generally associated with obvious structural modifications, and only in exceptional cases can it be said that the behavioral characters provide taxonomic information not already implicit in the anatomy. The classification presented is, of course, tentative. We anticipate that many changes will be made in it as knowledge of the genus increases and, in particular, as the larvae of additional species are discovered. For some years the genus Meloe has been segregated from other genera of Meloidae in either a separate subfamily (MacSwain, 1956) or tribe (Selander, 1964). In this work it is treated as a genus of the tribe Meloini, which is defined so as to include, besides Meloe, the genus Spastonyx. The latter taxon was formerly considered to be a INTRODUCTION subgenus of Eupompha in the tribe Lyttini. It is herein elevated to generic rank and assigned to a position near Meloe primarily on the basis of larval anatomy. ACKNOWLEDGMENTS The study was supported by an award to the senior author of a Public Health Service Pre-doctoral Fellowship (5-F1-GM-25-454-03) from the National Institute of General Medical Sciences and research grants from the National Science Foundation (GB-2437 and GB-5547; Richard B. Selander, Principal Investigator) We are deeply indebted to many of our colleagues and friends for their generous assistance. Loans of specimens from institutional col- lections now or formerly in their charge were made by Norman L. Anderson, Jr., Montana State University; George E. Ball, University of Alberta; William F. Barr, University of Idaho; Ross T. Bell, Uni- versity of Vermont; Horace R. Burke, Texas Agricultural and Me- chanical University; George W. Byers, University of Kansas; Leland Chandler, Purdue University; Joan B. Chapin, Louisiana State Uni- versity; Arthur Cole, University of Tennessee; P.. J. Darlington, Jr., and John F. Lawrence, Museum of Comparative Zoology, Harvard University; Ralph Dawson, Washington State University; L. Dieck- mann, Deutsches Entomologisches Institut; William A. Drew and Don C. Arnold, Oklahoma State University; Henry S. Dybas and Rupert L.