/ y 7 t; otates ill Department of Agriculture A Revision of the Agricultural Research Subfamily Criocerinae Service Techrrical (Chrysomelidae) of Bulletin 1805 Nortli America Nortli of iViexico r- --"I United States Department of A Revision of tiie Subfamily Agriculture Agricultural Criocerinae (Ciirysoiinelidae) Research Service of North America Technical Bulletin 1805 Nortli of i\Aexico May 1993 Richard E. White Systematic Entomology Laboratory, Plant Sciences Institute Agricultural Research Service, USDA c/o U.S. National Museum of Natural History Washington, DC 20560 Abstract White, Richard E. 1993. A Revision of the Subfamily Criocerinae (Chrysomelidae) of North America North of Mexico. U.S. Department of Agriculture, Technical Bulletin No. 1805, 158 pp., 32 plates. This revision recognizes 44 species of Criocerinae in America north of Mexico. For each species there is a description, drawings, and a map showing the range. There are scanning electron micrographs showing some of the most significant morphological features of selected species. Twelve species are described as new: Lema maderensis, L. melanqfrons, Neolema adunata, N. cordata, N. ovalis, N. quadriguttata, Oulema coalescens, O. elongata, O. laticoUis, O. melanoventris, O. minuta, and O. variabais. New synonymy includes the following: Lema albini Lacordaire [=N. sexpunctata (Olivier)l, L. baiteata equesiris Lacordaire (=L. faaíteaía LeConte), L. trûinea datwaphila Kogan and Goeden (=L. triiinea White), Oulema gaspensis Brown [=0. palustns (Blatchley)], O. palustrisßoridana Schaeffer [=0. palustris (Blatchley)], and O. coloradensis Linell [=0. texana (Crotch)]. Lema circumvittata Clark had been ranked as a subspecies of L. corguncta Lacordaire; it is here raised to the species level. Lema trabeata Lacordaire is also raised to the species level; it has to the present been listed as a subspe- cies of L. confusa Chev. New combinations include Neolema ephippium (Lacordaire), N. gundlachiana (SuflFrian), N. Jacobina (Linell), N. sexpunctata (Olivier), Oulema concoior(LeConte), O. comuia (Fabricius), O. maculicollis (Lacordaire), O. morfiiineimpressa (Schaeffer), O. soy i (Crotch), O. simulans (Schaeffer), and O. texana (Crotch). In addition, N. gundlachiana (Suflfrian) is newly recorded from the United States. Neotypes are designated for five species: Lema trivittata Say, L. trilineata (Olivier), Oulema collaris, (Say), Oulema comuta (Fabricius), and Lilioceris lüii (Scopoli). Lema omata Gravenhorst is treated as a forgotten name, for it has remained unused in the literature as a senior synonym for over 50 years. It is probable that it equals Lema trûinea Whitt. One name is treated as of uncertain status: L. intermedia Guerin. Keywords: Chrysomelidae, Coleóptera, Criocerinae, Crioceris, Lema, Lilioceris, Neolema, Oulema, taxonomic revision. While supplies last, single copies of this publication may be obtained, at no cost, on request from USDA-ARS-BA, Systematic Entomology Laboratory, Bldg. 046, Beltsuüle Agricultural Research Center-West, Beltsville, MD 20705. Copies of this publication may also be obtained from the National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161. Trade names are used in this publication solely for the purpose of providing specific information. Mention of a trade name does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture or an endorsement by the Department over other products not mentioned. May 1993 in Acknowledgments My thanks are extended to the following individuals, who have loaned speci- mens for this study or aided in other ways. R.C. Bechtel, Division of Plant Industry, Reno, Nevada; Nicole Berti, Museum National d'Histoire Naturelle, Paris; Susann Braden and Walter Brown, National Museum of Natural His- tory, Washington, DC; H.R. Burke, Department of Entomology, Texas A&M University; N.M. Downie, Lafayette, Indiana; W.R. Enns, University of Mis- souri; Bryan Farrell, University of Maryland; D.H. Kavanaugh, California Academy of Sciences; Laurent LeSage, Biosystematics Research Center, Ottawa, Canada; A.F. Newton, Museum of Comparative Zoology, Harvard University; Diether Peschken, Agriculture Canada, Regina, Saskatchewan; E.G. Riley, Department of Entomology, Texas A&M University; E.P. Rouse, University of Arkansas; Sharon Shute, British Museum of Natural History; J.M. Sullivan, St. Louis, Missouri; C.A. Triplehom, The Ohio State University; Ken Walker, Museum of Victoria, Australia; Robert Ward, Washington, DC; R.E. Woodruff, Florida Department of Agriculture, Gainesville. My appreciation is expressed to Allen Norrbom, Edward Balsbaugh, Jr., and Ingolf Askevold for reviewing the manuscript. Contents Introduction 1 Historical review 2 Materials 2 Methods 3 Nomenclature 5 Quoted data 6 Hosts 6 Life history summaries 8 Species life history data 9 Diagnostic characters 12 Abbreviations 14 Criocerinae 14 List of Criocerinae of North America 18 Key to North American genera of Criocerinae 19 Lema Fabricius 19 Neolema Monros, new status 56 Oulema Des Gozis 78 Crioceris Müller 121 Lilioceris Reitter 128 A forgotten name 132 Uncertain status 133 References cited 134 Illustrations Maps Male genitalia A Revision of tlie Subfamily Criocerinae (Chrysomelidae) of Nortii America Nortii of IVIexico by Richard E. White Introduction The subfamily Criocerinae is a large and nearly worldwide group of roughly 1,500 described species. The NortJi American species are medium-sized leaf beetles (2.7-8.0 mm long) that feed on no less than seven different families of plants. Although some species are serious economic pests, this subfamily has never been thoroughly studied. Recent elevations of supposed subspecies to the species level (Balsbaugh and Hays 1972, White and Day 1979) provide evidence of our incomplete knowledge of the interrelationships of the taxa. The transfer by Monros (1960:229) of certain species traditionally placed in Lema to Oulema has been corroborated but found to be incomplete, for herein more species are similarly transferred. This study is intended to consolidate existing information on the North Ameri- can species of the subfamily; describe new taxa; provide a description for each species; correct generic assignments where necessary; establish new syn- onymy as needed; redefine known species; and provide keys, illustrations, and maps to aid in identifications. Twelve species are herein described as new, and 5 new synonyms and 11 new combinations are presented. One species is newly recorded from the United States. Two former subspecies are raised to the species level. This work brings to 44 the number of North American species. Illustrations of each species (figs. 1-45) are intended to show the extremes of variability in color pattern and (if it occurs) in melanism. The literature for many species is extensive, so to aid in its examination, an indication is given of the type of information provided by each reference cited in the species synonymy (e.g., "Monros, 1960:222 (world checklist), Kaufmann, 1967:363 (biology and larva)"). Biological notes on specimen labels and those found in the literature are summarized in the "Hosts" section of each species treatment. Four economically important species have been accidentally introduced into North America from Europe: Lilioceris íiíüScopoli, the lily leaf beetle (about 1944), Crioceris asparagi (Linnaeus), the asparagus beetle (about 1859), C. daodecimpunctata (Linnaeus), the spotted asparagus beetle (about 1881), and Oulema m^elanopus (Linnaeus), the cereal leaf beetle (about 1959). The most injurious native species is Lema trilinea White, the threelined potato beetle; it is not as injurious as it once was, and not as significant as the last three species named above. One European species. Lemapuncticollis (Curtis), has recently been introduced into Canada in hopes that it will help in the control of Canada thistle. There is as yet no proof that it has become established. The species currently of the greatest importance to agriculture is the cereal leaf beetle, Oulema melanopus (L.). Its significance and the interest in it are shown by the more than 400 papers published in economic literature of North America since the early 1960's (Battenfield et al. 1982). Research entomologist, Systematic Entomology Laboratory, U.S. Department of Agriculture, Agricultural Research Service, c/o U.S. National Museum of Natural History, Washington, DC 20560. Historical Review Three of the species included herein were described by Linnaeus (1758:376) as members of the genus Chrysomela: Crioceris asparagU C. duodecim- punctata. and Oulema melanopus. In 1763 Scopoli described Attelabus liliU a species now in the genus Lilioceris, Fabricius described two North American species: Lema solani (1798) and Lema comuta (1801, now Oulema). Two species were described in Crioceris by Olivier in 1808: Lema sexpunctata (here placed in Neolema) and L. trilineata (now íriíinea White). Say in 1824 de- scribed two species in Lema: L. trivittata and Oulema coUaris. Guérin- Méneville in 1829 (1844) described the single species Crioceris nigrovittata (now in Lema), Curtis in 1830 described Crioceris puncticoUis (now in Lema), In 1835 Chevrolat described the species Lema corifuscL The number of taxa increased significantly in 1845 when Lacordaire described Lema albini (here placed in NeolemcH. L balteata equestris, L. coryuncta, L. ephippium (here placed in Neolemdl, L. confusa trabeata, L. tnvirgata, L brunnicollis (now in Oulema), and L. maculicollis (here