Revision of the (Coleoptera: Cerambycidae)

Steven W. Lingafelter and E. Richard Hoebeke

Published by The Entomological Society of Washington Washington, D.C. 2002 Author's addresses:

Steven W. Lingafelter Systematic Entomology Laboratory, Plant Sciences Institute, ARS, USDA National Museum of Natural History Washington, D.C. 20560-0168

E. Richard Hoebeke Department of Entomology, Comstock Hall Cornell University Ithaca, New York 14853-0901

ISBN 0-9720714-1-5 Published by: The Entomological Society of Washington, Washington, D.C., U.S.A. David R. Smith, Editor. Printed by Allen Press, Inc., Lawrence, Kansas, U.S.A. Cover Design: Kent Loeffler, Cornell University Endpaper Design: Frances Fawcett, Ithaca, New York Recommended format for citation: Lingafelter, S. W. and E. R. Hoebeke. 2002. Revision of Anoplophora (Coleoptera: Cerambycidae). Entomological Society of Washington, Washington, D.C., U.S.A. 236 pp. 41 color plates. Date issued: July 15, 2002.

Front cover: Anoplophora glabripennis (Motschulsky) and map of Anoplophora distribution. Back cover: See back endpaper for legend. LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 7

TABLE OF CONTENTS

Preface ______------···--·- ...... ·--··············-·····-· .. ·-- 9 Acknowledgments ------11 Introduction ------···· ··--··--··-- 13 Summary of Taxonomic Changes ______.... __ ...... ·------13 Methods ------14 Biology of Anoplophora ------16 Historical Review of Anoplophora and its Tribal Assignment . ·······------26 Systematics ------.... ---·------..... ··-· ··------···· .. ------28 Description of adult Anoplophora _.- _ ..... ------·-- .... ····------28 Description and diagnosis of Anoplophora larvae ------29 Discussion of characters supporting monophyly of and Anoplophora ______30 Diagnosis of adult Anoplophora ------32 Distinction of genera related to Anoplophora ------32 Checklist of Anoplophora Species -- -- ·-- ...... ·-·-····--· .. ------33 Key to Species of Anoplophora ------· .. _------35 Species Accounts Anoplophora albopicta (Matsushita) ------·--- .... ------··- ·····------40 Anoplophora amoena (Jordan) ------··········-----·--·-·------41 Anoplophora asuanga Schultze ------·-·------42 Anoplophora beryllina (Hope) ---- ·····-················--·------.... ------43 Anoplophora birmanica Hiidepohl ______.. . .. ______46 Anoplo pho ra bow ring ii (White) ------····------4 7 Anoplophora chiangi Hua and Zhang -----·· .. ·····------48 Anoplophora chinensis (Forster) ------49 Anoplo pho ra coe ruleoante nnata (Breuning) ------············ ...... ········-···········-- -- 5 8 Anoplophora davidis (Fairmaire) ------······ ------59 Anoplophora decemmaculata Pu ------61 Anoplophora elegans (Gahan) ______...... ------104 Anoplophora fiavomaculata (Gre sitt) ______...... ·····------105 Anoplophora freyi (Breuning) ------106 Anop lop ho ra f ruhsto rfe ri (A uri villi us) ------·· ------108 Anoplophora glab ripennis (Motschulsky) ______------109 Anoplophora graafi (Ritsema) ------···· ------·· .. ·------·······--··· .. ······· -- -- 113 Anoplophora granata Holzschuh ------·-- ·---- -···----··------· ·· ------115 Anoplophora horsfieldii (Hope) ------······------·--·------116 Anop lop ho ra imitator (White) ------· . ·------118 Anoplophora irregularis (Auri villi us) ------..... ______120 Anoplophora jianfenglingensis Hua ------··--·------· ...... ------·· ---- 120 Anop lop ho ra leec hi (Gahan) ------121 Anoplophora longehirsuta Breuning ------123 Anoplophora lucipor Newman ------124 Anoplophora lurida (Pascoe) ------··------125 Anop lop ho ra macularia (Thomson) ------12 7 Anoplophora mamaua Schultze ------131 Anoplophora medenbachii (Ritsema) ------132 Anoplophora ogasawaraensis Makihara --- ·---····--········ ·------134 Anoplophora similis (Gahan) ------· ...... ------134 Anoplophora sollii (Hope) ------· ...... ______136 Anoplophora stanleyana Hope ------137 Anoplophora tianaca Schultze ------139 Anoplophora viriantennatus Wang and Chiang ------140 Anoplophora zonator (Thomson) ------140 Species Designated as incertae sedis ------. ·· ------142 8 ENTOMOLOGICAL SOCIETY OF WASHINGTON

Species Transferred from Anoplopho ra ------.... ·-· ------143 Literature Cited ------143 Table 1. Comparison of character states in Anoplophora macularia/chinensis complex ------·------150 Figures 4-67 ------150 Plates 1-21 ------.. ····-··· ------·------62 Plates 22- 34 . . . .. ···· ------······ . ------208 Maps 1-14 ------·...... ··------221 Index ------235 About the Authors ------· --· ------236 LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 9

PREFACE longhorned . In an attempt to eradicate this de­ structive woodboring pest from these New York The impetus for this work is directly attributable neighborhoods, federal, state, and local regulatory to the first-time discovery of an invasive woodboring agencies began removing and destroying all trees beetle of the cerambycid genus Anoplophora in New with damage symptoms of this beetle-i.e., circular York City in 1996. Below is a chronology of the exit holes in trunks and branches and accumulations incident and its effects. of coarse sawdust at the base of trees and in branch In early August 1996, Ingram Carner, a long-time crotches. This cut-and-destroy mission began in resident of Greenpoint, Brooklyn, began noticing that March 1997, first with infested trees of McCarren many of his Norway trees along McGuinness Park, Greenpoint, Brooklyn. A venue were showing signs of distress. The trees had In early July 1998, Garry Luka, an employee of numerous large circular holes, 3/8-3/4 inch in di­ the Skokie Park District, just north of Chicago, sawed ameter, in the trunks and branches. At the base of the off a dead tree branch that was overhanging his prop­ trunks and in the crotches of the larger branches were erty in the 4300 block of North Winchester A venue, amounts of coarse sawdust. Mr. Carner believed Ravenswood, and offered the wood to his friend Bar­ that these street trees were being damaged by vandals ry Albach, to use for an upcoming camping trip. Mr. using cordless drills. However, on August 19, while Albach placed the firewood in the back of his pickup again examining these damaged trees, he noticed a truck, where it sat under a fiberglass cover for a large, shiny, black beetle emerging from one of the week. Then on July 9, his mother and a neighbor circular holes in the trunk. This time he guessed that spotted a large black beetle crawling on Albach's rear the damage might be due to these . He notified view mirror. When Mr. Albach uncovered the wood the New York City Department of Parks and Recre­ in the back of his truck, he discovered more of the ation and they dispatched a forestry inspector, HmTy beetles crawling about. Curiosity brought him to his Rothar, who collected a beetle and returned it to his computer, where he did a simple Internet search on office. Unable to identify the beetle, it was sent to "beetles" and luck would have it that the very first Cornell University's Department of Entomology. site he found was one on the Asian longhorned beetle There, it arrived at the Diagnostic Lab on Au­ in New York. What was illustrated in this pest alert gust 26, and diagnostician Carolyn Klass, who did by the USDA was exactly what Mr. Albach saw in not recognize it, sent it along to E. R. Hoebeke for the back of his truck. He decided immediately to con­ identification. Shortly thereafter, on September 2, tact the local USDA office and notify them of his Hoebeke identified the mystery woodboring beetle as find. After a site visit by USDA inspectors, it was Anoplophora glabripennis, a native of China and Ko­ determined that another major infestation of Asian rea. On September 5, he traveled to Greenpoint, longhorned beetle was present in the northside Chi­ Brooklyn, met with Harry Rothar, and went directly cago community of Ravenswood. In the next few to the site where the beetle was first discovered. More weeks, local Chicago television, radio, and newspa­ street trees in the vicinity of McGuinness A venue pers provided outstanding coverage of this newly dis­ were discovered to be heavily infested, and additional covered beetle infestation. As a result of this public­ beetles were collected. Upon his return to Cornell, ity, local citizens were responsible for the discovery Hoebeke sent specimens to the USDA Agricultural of two additional, but smaller, areas of infestation­ Research Service's Steven W. Lingafelter, a specialist one near Addison, DuPage County, about 5 miles of the Cerambycidae, and to Allan G. Samuelson of southeast of O'Hare International Airport, and anoth­ the Bernice Bishop Museum, Honolulu, Hawaii, an­ er in Summit, a suburb about 15 miles southwest of other cerambycid expert, for their confirmation. By downtown Chicago. With the detection of the Asian September 13, both systematists had confirmed Hoe­ longhorned beetle in Chicago, the federal govern­ beke 's identification of Anoplophora glabripennis. ment was compelled to take some action. On Septem­ A month later, in October 1996, another infested ber 11, Secretary of Agriculture Daniel Glickman area was found in Amityville, New York, about 40 held a news conference in Chicago to announce in­ miles east of Brooklyn, on Long Island. A tree cut­ terim U.S. trade restrictions with China. The newly ting company of this community had been pruning announced regulations-set to go into effect on De­ and removing dead or dying trees from the metro­ cember 17, 1998-would require that goods from politan New York area in previous years and much China be packed in non-wood material or wood that of this wood was stored on their property in Amity­ had been treated with heat or chemical fumigation to ville where it was sold as firewood. Many of the ki11 . Removal of beetle-infested trees in Chi­ street trees bordering this property and elsewhere in cago began on February 4, 1999. Infested trees con­ the immediate vicinity were infested by this Asian tinue to be found in New York and Chicago. As of 10 ENTOMOLOGICAL SOCIETY OF WASHINGTON

August 4, 2001, a total of 5,324 infested trees in New anese that were destroyed. Intensive surveys York and 1 infested trees in Chicago have been there are being coordinated by state and federal agen­ detected and destroyed. cies. This species has also been introduced into Eng­ No one knows exactly how long the Asian long­ land (Cooter 1998) and established recently in Italy horned beetle has been present in the United States, (Colombo and Limonta 2001). or when it first began infesting trees in New York Because the volume of commodities imported an­ City or Chicago. Fortuitously, a property owner in nually from China associated with solid wood pack­ Bayside, Queens, came forth shortly after the discov­ ing material is estimated to be in the millions of ship­ ery of infested trees in this neighborhood in February ments, it is expected that other non-native woodbor­ 1999 with a Polaroid print of the Asian longhorned ing beetles will continue to find their way into N011h beetle, dated from 1992. There is little doubt that this American urban landscapes using this pathway. dangerous tree-infesting beetle gained entry into USDA port officials have been placed on higher alert, North America in solid wood packing material, such and continue to inspect high-risk cargoes from Asia as crating, pallets, and dunnage, originating from for species of Anoplophora and other woodboring China, and used extensively in the international trade pests. industry for packaging goods and merchandise. Prior The U.S. Department of Agriculture's Agricultural to the Asian longhorned beetle's discovery in August Research Service (ARS) continues to support an 1996, there were only two known interceptions of Asian longhorned beetle research program that pro­ Anoplophora glabripennis in North America. One vides necessary research leading to environmentally was from Chinese crating in Loudenville, Ohio, in and publicly acceptable technologies for mitigating 1992, and another on wood crating and dunnage, also beetle populations. Based on recommendations of a from China, in Vancouver, British Columbia, in 1992. Science Advisory Panel, ARS was advised in Octo­ Since August, 1996, numerous interceptions of A. ber 1996 that top priorities should include beetle sys­ glabripennis have been made at major U.S. ports and tematics, determination of basic biology, pheromone in numerous warehouses countrywide. Other coun­ tries are also dealing with this as a new invasive pest. identification and isolation, field detection and trap­ Recently, its first occurrence in Europe was indicated ping technologies, evaluation of beetle reproductive in July, 2001, in Braunau, Austria (Tomiczek 2001). biology, and identification and testing of potential However, A. glabripennis is not the only species of natural enemies. the genus to be intercepted in the United States. Oth­ In early 1998, USDA-ARS awarded Steven W. er serious tree pests of Asia also have been found as Lingafelter and E. Richard Hoebeke a grant to un­ unintentional hitchhikers in 1999. Several live spec­ dertake a comprehensive taxonomic revision of the imens of Anoplophora chinensis, one of Asia's most Oriental genus Anoplophora and to prepare an illus­ serious pests, were found in April in a green­ trated handbook for the identification of species in house in Athens, Georgia, where Crepe Myrtle bon­ the genus. The identification guide is intended to be sai trees from China were infested, and another in­ part of a national strategy to eradicate this woodbor­ terception of a single live specimen of the white-spot­ ing pest and to help port-of-entry regulatory officials ted longicorn beetle, Anoplophora chinensis (form nationwide distinguish this Asian pest and its close malasiaca), was made by a homeowner in June on relatives from the hundreds of native species of Cer­ an infested, potted dwarf Japanese maple in Delavan, ambycidae. We trust that this monograph will be a Wisconsin. Very recently, in August, 2001, 4 speci­ positive contribution to that end. mens of Anoplophora chinensis (form malasiaca) were found in two nurseries in the Seattle area, Wash­ E. R. Hoebeke ington state (Tukwila and These specimens S. W. Lingafelter were also associated with a shipment of bonsai Jap- November, 2001 LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 11

ACKNOWLEDGMENTS (NCSU), Kyu-Tek Park (KNUC), Jeong-Kyu Kim and Ohseok Kwon (KETU), Young Bok Cho We are grateful to the United States Department of (HUNM), Seong-Mo Lee (NSMK), Bong-Kyu Byun Agriculture, Agriculture Research Service, National (FRIS), Pu Fuji and Yu Peiyu (IZAS), Darren Mann Programs Staff (Raymond CaITuthers, Kevin Hackett, (OXUM), Al Samuelson and Karin Kami (BPBM), Walker Jones), Plant Sciences Institute, and System­ Hiroshi Makihara and Mitsuaki Shimazu (FRIC), Ish­ atic Entomology Laboratory (hereafter, SEL: Manya izue Adachi (FRIC), Wen-I Chou (NTUC), Hsien- Stoetzel and Michael Schauff, Research Leaders) for Shih, Man-Miao Yang and Chan providing and administering funds for fieldwork, mu­ (NMNS), Jung-Tai Chao and Wen-Chi Yeh (TFRI), seum work, illustrations, photography, and manu­ L. Y. Chou (TARI), Otto Merkl (HNHM), Masahiro script costs that led to completion of this revision. Ohara (EIHU), Jun-Im Song (EWUC), Bruce Gill For hosting us or otherwise helping us immensely (CNCI), Jean-Michel Maes (SEAN), Heiner Sch()n­ with innumerable logistical issues associated with our mann (NHMW), Bert Viklund and Thomas Pape foreign travel, we thank: Kunio Suzuki (Toyama Uni­ (NHRS), Carolus Holzschuh (CHPC), Larry Bezark versity, Gofuku, Japan), Nobuo Ohbayashi, Masahiro (LBPC), Jim Cope (JCPC), Dan Heffern (DHPC), Sakai, and students Kaori Kasahara and Shinji Yano Wayne Hsu, Luis M. Chong, and Huan-Chih Hsu (EUMJ), Hongyin Chen, Gangying Zhang, and Paul (Taipei, Taiwan), Robert Husband (Adrian ..._,,...,.,_,.....,"""""· Schaefer (Sino-American Biological Control Labo­ Michigan), Timothy L. McCabe (NYSM), David ratory, Beijing, China), Kee-Jeong Ahn (CNUC), Williams (USDA Forest Service, Newtown Square, Jong-Eun Lee (ANUC), Young-Chul Chang, Jun­ Pennsylvania), and Joseph V. McHugh and Cecil L. Seok Lee, and Kyu-Tek Park (KNUC), Wu Weiwen Smith (UGCA). and Chiang Shunan (SACA), Hua Lizhong (ICRI), For help in entering locality data for Anoplophora Dhruba Manandhar (Kathmandu, Nepal), Mark Vol­ in a database, we thank Alexey Tishechkin (Louisi­ kovitsh (ZMAS), Masataka Sato (Nagoya Women's ana State University, Baton Rouge). For help in cu- University, Nagoya, Japan), Shigehiko Shiyake the Smithsonian Institution collection of Cer­ (OMNH), Junichi Yukawa, Teruhisa Ueno, Hiroaki ambycidae instrumental to this project, we thank: Kojima (KUEC), John Whitman (Cornell University, Dan Heffern (Houston, Texas), Jenny Schade (Ijams­ on leave at University of Tsukuba, Japan), Kazuhiko ville, Maryland), Alex Harman (Silver Spring, Mary­ Konishi (ITLJ), Ritzuo Nishita (Kyoto University, land), and Elisabeth Roberts (SEL). Kyoto, Japan), Shfihei Nomura (NSMT), Noboru Ito For the habitus photographs and preparation of the (Kawanishi City, Hyago Pref., Japan), Tateo Ito (Ya­ plates, we thank the enthusiastic and talented Kent wata, Kyoto Japan), Akiko Saito (Natural Loeffler (Cornell University). For photographs of History Museum, Chiba, Japan), Tatsuya Niisato (To­ some types, we thank Tatsuya Niisato (Tokyo, Japan). kyo, Japan), Keith Kuan Lee (Pei Kuan Resort, Ilan, For the photograph of the Netsuke, we thank Alex­ Taiwan), Wayne Hsu, Luis M. Chong, and Huan­ ander Konstantinov (SEL). For many of the line il­ Chih Hsu (Taipei, Taiwan), Al Samuelson (BPBM), lustrations and the half-tones of elytral maculations, Baode Wang (USDA-APHIS-PPQ, Otis, Massachu­ we thank Mary Van Slyke (Erie, Pennsylvania). For setts), Robert Haack (Michigan State University, the carbon dust rendering of Anoplophora USDA-FS, East Lansing, Michigan), and Gao Rui­ nis used in the endpaper design, we thank Frances tong (CFRB). Fawcett (Ithaca, New York). For access to collections and or loans of material, For miscellaneous logistical help and/or providing we thank: Nobuo Ohbayashi (EUMJ), Sharon Shute specimens we thank: Paul Schaefer (USDA, Newark, (BMNH), Jean Menier (MNHN), John Chemsak and Michael Smith (USDA, Newark, DE), Thomas Cheryl Barr (EMEC), David Kavanaugh and Roberta Dubois and Ann Hajek (Cornell University). Brett (CASC), Eva Sprecher-Uebersax (FCBS), Jun­ translation services, we thank: Wenhua Lu (USDA- ichi Yukawa (KUEC), Vic Mastro (USDA-FS-Otis, Ansonia, Connecticut), Bruce Gill (CNCI), Akito Massachusetts), Phil Perkins and Brian Farrell Kawahara, who also provided specimens (Cornell (MCZC), Lee Herman (AMNH), Don Azuma and University), Xiuyu (Julie) Cong (Cornell University), Daniel Otte (ANSP), John Rawlins and Robert L. Zhimou Wen (Cornell University), Xin Wang Davidson (CMNH), N. Nikitsky (MSUC), D. Chris­ (Cornell University), and Kuniko Arakawa (San Jose, topher Darling (ROME), Igor Lobanov (ZMAS), Al Costa Rica). For assistance with public dissemination Newton and Philip Parrillo (FMNH), Brett Ratcliffe and popular publications, we thank Hank Becker and Mary Liz Jameson (UNSM), Steve Ashe and (USDA, ARS) and Blaine Friedlander (Cornell Uni­ Rob Brooks (SEMC), Mike Thomas (FSCA), Wayne versity News Service). Gall (BMSC), David L. Stephan, Robert L. Blinn Norman Woodley (SEL) helped in innumerable 12 ENTOMOLOGICAL SOCIETY OF WASHINGTON ways, constructive advice, finding obscure lit­ (SEL). For help in obtaining literature, we also thank erature, and carefully reviewing and testing the key. Chris Thompson and Elisabeth Roberts (SEL) and For thorough reviews and comments on the manu­ Marty Schlabach (Cornell University). And, finally, script, we especially thank Chris Thompson (who the second author (ERH) thanks his wife, Maureen greatly improved the bibliography and nomencla­ Carter, for her unflagging support and encouragement ture), John Chemsak (EMEC), and Allen Norrbom throughout the course of this work. LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 13

INTRODUCTION herein), can taxonomic stability be achieved, and we have approached this project from that perspective. Anoplophora is a moderate-sized genus of 36 spe­ cies of woodboring beetles that occur throughout SUMMARY OF TAXONOMIC CHANGES Asia, with the highest diversity in the tropical and The Asian longhorned beetles of the genus Ano­ subtropical regions. Most species have beautiful col­ plophora include 36 species as a result of this study. ors on the elytra, pronotum, antennae, tarsi, and ven­ The following 20 new synonyms are made: Melan­ ter. These colors superficially seem like mere patterns auster granulipennis Breuning 1938 and Anoplopho­ of the integument, but under high magnification are ra subberyllina Breuning 1964 are new synonyms of revealed to be due to very dense patches of short Anoplophora beryllina (Hope); Anoplophora stanley­ pubescence. In many species, these colored pubes­ ana var. grisea Tippmann 1953, is a new synonym cent patches on the elytra are arranged in round spots, of Anoplophora birmanica Htidepohl; Calloplophora in others they are arranged in transverse bands. Most abbreviata Thomson 1865, Calloplophora malasiaca species have very long antennae (about 1.3-1.6 times Thomson 1865, and Melanauster perroudi Pie 1953 the body length in males, 1.0-1.5 times the body are new synonyms of Anoplophora chinensis (For­ length in females of most species) and most are large, ster); Melanauster adonis Pie 1925b, Melanauster some over 50 mm. In the larval stage, in this adonis var. vitalisi Pie 1936, and Anoplophora al­ genus develop in and consume wood of many species boapicalis Chiang 1951 are new synonyms of Ano­ of trees. plophora davidis (Fairmaire); Cyriocrates depressi­ Anoplophora is a familiar genus to many Asians, notatus Chiang 1951 is a new synonym of Anoplo­ attested to through the abundance of common names phora fruhst01feri (Aurivillius); Melanauster nobilis in the literature, and their in art Two Ganglbauer 1890 is a new synonym of Anoplophora common names are used widely by laypersons in glabripennis (Motschulsky); Melanauster pirouletii Asia for various species: The Sky Ox Beetle and Fairmaire 1889 is a new synonym of Anoplophora Starry Night Sky Beetle. Many Japanese netsuke imitator (White); Anoplophora parelegans Chiang carvings from ivory and wood feature Anoplophora 1981 is a new synonym of Anoplophora longehirsuta (see Frontispiece). The recent introductions of Ano­ Breuning; Anoplophora lucipor lumawigi Htidepohl plophora glabripennis in the United States have es­ 1989 is a new synonym of Anoplophora lucipor tablished its notoriety in the New World (Becker Newman; Anoplophora malasiaca tokunoshimana 1998, Becker 2000, Cole 1999, Milius 1999, Stimmel Samuelson 1965, Melanauster oshimanus Fairmaire 1999). 1895, and Anoplophora oshimana ryukyensis Breun­ Despite the wide familiarity of the genus, much ing and Ohbayashi 1964 are new synonyms of Ano­ confusion has exisled concerning the identity and def­ plnphora macularia (Thomson); Anoplophora tonki­ inition of the various species. This study was done nea Breuning l 944a, Anoplophora stanleyana var. gloriosa Tippmann 1953, and Anoplophora stanley­ to provide a resource for the identification of all spe­ ana var. melancholica Tippmann 1953 are new syn­ cies in the genus Anoplophora. Thorough studies of onyms of Anoplophora stanleyana Hope. external and internal features, and their variation, for The following four taxa are transferred to Anoplo­ adults of all species were made and are presented phora and represent new combinations: Anoplophora here. We have provided numerous illustrations, many albopicta (Matsushita 1933); Anoplophora fruhst01feri in color, to allow even the non-specialist the ability (Aurivillius 1902); Anoplophora graafi (Ritsema to make identifications with confidence. We include 1880); and Anoplophora sollii (Hope 1839). The fol­ an identification key and distribution maps for each lowing taxa are removed from Anoplophora and rep­ species. A general description of larvae is presented, resent new or revised combinations: Pseudonemophas although this is necessarily brief since larvae of most versteegii (Ritsema 188 lc), new combination; Nemo­ species are unknown. phas ammiralis (Schwarzer 1931), new combination; One important goal was to stabilize the bennigseni Aurivillius 1908, restored com­ of this group by designating synonyms where we dis­ bination; itzingeri (Breuning 1938), new covered multiple names being used for the same tax­ combination; and rondoniana (Breuning on. This can only be done in a project such as this 1964), new combination. The following taxa are des­ which is revisionary in scope, treats all the fauna, ignated as incertae sedis: Anoplophora wusheana and focuses on examination of all type specimens. Chang 1960, as wusheana Chang, incertae sedis and Only by studying all species and numerous speci­ Calloplophora tonkinea Pie 1907, as tonkinea Pie, in­ mens of a taxon to assess variation and constancy of ce rtae sedis. The following new synonyms are made characters (on which we base our species concepts for these taxa excluded from Anoplophora: Monocha- 14 ENTOMOLOGICAL SOCIETY OF WASHINGTON mus glabronotatus Pie l 934a, and Monochamus al­ wan, and Nepal in an effort to gain fresh material bescens v. subun~formis Pie l 934a are new synonyms and biological and distributional data. More impor­ of Pseudonemophas versteegii (Ritsema 1881 c ). tantly, we have carefully examined museum hold­ Lectotypes are designated for the following names: ings, some of which have been accumulated for over Monochamus beryllinus Hope; Calloplophora luc­ 200 years. We have visited nearly 50 museums in tuosa Thomson; Calloplophora malasiaca Thomson; Europe, Asia, and North America, and received over Melanauster davidis Fairmaire; Melanauster macro­ a thousand specimens from them and other institu­ spilus Gahan; glabripennis Motschulsky; tions and individuals listed below and in the Ac­ Melanauster pirouletii Fairmaire; Calloplophora knowledgments. macularia Thomson. Types for the following species A list of institutional collections follows. Acro­ were not located and neotypes are herein designated: nyms are from Arnett et al. (1993), and are in alpha­ Anoplophora asuanga Schultze; Anoplophora rna­ betical order by acryonym. Curators for institutional maua Schultze; Anoplophora tianaca Schultze. and private collections are listed in the Acknowledg­ ments. METHODS We present an overview of the nomenclatural his­ AMNH American Museum of Natural History, tory of Anoplophora and its tribal association. We pro­ New York City, New York, USA vide a discussion of biology and distribution of Ano­ ANSP Academy of Natural Sciences, Philadel­ plophora. We then provide generic descriptions and phia, Pennsylvania, USA diagnoses of larval and adult Anoplophora. A catalog ANUC Andong National University, Andong, for all currently recognized species of Anoplophora, South Korea with complete lists of synonyms and abbreviated ref­ BMNH The Natural History Museum, London, erences, is included. Detailed descriptions and diag­ United Kingdom noses of each species, based on all material at hand BMSC Buffalo Museum of Science, Buffalo, New and examined from museums and collections around York, USA the world, are presented. Morphological characters are BPBM Bernice P. Bishop Museum, Honolulu, Ha­ thoroughly illustrated and an atlas of features for Ano- waii, USA plophora is provided as a reference 4 ). CASC California Academy of Sciences, San Fran­ Illustrations and photographs are provided as fol­ cisco, California, USA lows: 34 full color photographic plates of all species CFRB Chinese Academy of Forestry, Forest Re­ of Anoplophora and available holotype, lectotype, search Institute, Beijing, China syntype, or neotype specimens are presented. The CHPC Carolus Holzschuh Private Collection, Vi­ first 18 plates are of mostly pristine examples of both enna, Austria sexes for most species and are arranged to facilitate CMNH Carnegie Museum of Natural History, identification, with similar species located together. Pittsburgh, Pennsylvania, USA Following these are three plates (Plates 19-21) show­ CNCI Canadian National Collection of Insects, ing examples of genera related to Anoplophora. Leg­ Ottawa, Ontario, Canada ends for Plates 1-21 also show actual sized silhou­ CNUC Chungnam National University, Daejon, ettes of the beetles photographed. These 21 plates are South Korea located prominently in the middle of the book. The CUIC Cornell University Insect Collection, Itha­ last 13 plates (Plates 22-34) show most available pri­ ca, New York, USA mary type specimens, mostly arranged alphabetically DHPC Dan Heffern Private Collection, Houston, by specific epithet and are located at the end of the Texas, USA book, immediately following the figures. EIHU Entomological Institute, Hokkaido Univer­ Over 60 figures are provided (each as a plate with sity, Sapporo, Japan numerous individually labelled elements) of Anoplo­ EMEC Essig Museum of Entomology, University phora in situ, morphological dissections, and line of California, Berkeley, California, USA drawings of various features. These are all together EUMJ Ehime University and Ohbayashi Collec­ at the end of the book, immediately before the re­ tion, Matsuyama, Japan maining Plates and Maps, with the exception of the EWUC Ewha Women's University, Seoul, South full color biological figures 1-3) that are lo­ Korea cated within the chapter on biology. FCBS Frey Collection and Natural History Mu­ A comprehensive revision of this nature depends seum, Basel, Switzerland on a thorough study of specimens. We have con­ FMNH Field Museum of Natural History, Chica­ ducted field work in Japan, China, South Korea, Tai- go, Illinois, USA LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 15

FRIC Forestry and Forest Products Research In­ OMNH Osaka Natural History Museum, Osaka, stitute, Fruit Tree Sciences, Tsukuba, Japan Japan FRIS Forestry Research Institute, Seoul, South OXUM Hope Entomological Collections, Oxford, Korea United Kingdom FSCA Florida State Collection of , ROME Royal Ontario Museum, Toronto, Canada Gainesville, Florida, USA SACA Southwest Agricultural University, Beibei, HNHM Hungarian Natural History Museum, Bu­ Chongqing, China dapest, Hungary SEAN Servicio Entomol6gico Aut6nomo, Museo HUNM Hannam University Natural History Mu­ Entomol6gico, Leon, Nicaragua seum, Daejon, South Korea SEMC Snow Entomological Museum, University ICRI Zhongshan University Collection, Guang­ of Kansas, Lawrence, Kansas, USA zhou, China TARI Taiwan Agricultural Research Institute ITLJ National Institute of Agro-Environmental Collection, Taichung, Taiwan Sciences, Tsukuba, Japan TFRI Taiwan Forestry Research Institute, Taipei, IZAS Zoological Institute, Academia Sinica, Bei­ Taiwan jing, China UGCA University of Georgia Collection of Ar­ JCPC Jim Cope Private Collection, San Jose, thropods, Athens, Georgia, USA California, USA UNSM University of Nebraska State Museum, KEIU Korean Entomological Institute, Korea Lincoln, Nebraska, USA University, Seoul, South Korea USNM National Museum of Natural History, KNUC Center for Insect Systematics, Kangwon Smithsonian Institution, Washington, D.C., National University, Chuncheon, South USA Korea ZMAS Zoological Institute, Russian Academy of KUEC Kyushu University Insect Collection, Kyu­ Sciences, St. Petersburg, Russia shu, Japan LACM Natural History Museum of Los Angeles For our descriptions, at least one specimen of most County, Los Angeles, California, USA species was completely cleared and disarticulated to LBPC Larry Bezark Private Collection, Sacra­ carefully reveal morphological characters. Too few mento, California, USA specimens were available for several species, and LSUK Linnean Society Collection, Burlington therefore we could not make dissections. For others, House, Piccadilly, London, United King­ the genitalia were carefully dissected from both sex­ dom es, when available, and the internal sac was everted MCZC Museum of Comparative Zoology, Har- and studied. Characters were illustrated, mostly on vard University, Cambridge, Massachu­ scratchboard, using a Leica dissecting microscope setts, USA and camera lucida, and photographed through the MNHN Museum National d'Histoire Naturelle, same scope with a Sony DKC-5000® digital camera. Paris, France We have prepared, with much help from Kent Loeff­ MSUC Moscow State University, Moscow, Russia ler (Cornell University), color photographs of dorsal NCSU North Carolina State University, Raleigh, views of good specimens of both sexes for most spe­ North Carolina, USA cies using a Nikon® camera system body, I 05 NHMW Naturhistorisches Museum, Vienna, Aus­ Micro Nikkor® with bellows), scanning in slides us­ tria ing Nikon LS-2000@ and Umax Powerlook 3000@ NHRS N aturhistoriska Riksmuseet, Stockholm, scanners, and providing prints using a Codonics dye Sweden sublimation printer. The specimens are reproduced at NIAS National Institute of Agricultural Sciences, the maximum size allowable to fit on the page so that Suwon, South Korea features are easily visible. Therefore, specimens are NMNS National Museum of Natural Science, Tai­ not represented to scale, but we have included ranges chung, Taiwan of lengths for each species along with silhouettes of NSMK National Science Museum, Seoul, South actual size. Further, the back cover illustration shows Korea most species to approximate relative scale, so relative NSMT National Science Museum, Tokyo, Japan sizes can be easily discerned. NTUC National Taiwan University, Taipei, Tai­ Following each description and diagnosis wan section, we provide a discussion of all the types and NYSM New York State Museum Collection, Al­ include color photographs of most primary types bany, New York, USA (Plates For all new synonymies and many 16 ENTOMOLOGICAL SOCIETY OF WASHINGTON earlier synonymies, we provide justifications. We Jist ically important group of woodboring beetles. Be­ all the type material and other specimens examined cause most biological literature on these beetles is for each species. These data are sorted and subsorted published in Chinese, especially that treating Anoplo­ for each species by country, state or province, and phora glabripennis and the other major citrus often specific locality, and latitude/longitude to facil­ of the genus, it has been difficult to gain access to itate examination for specific locality data. This also past and present research on these woodborers. Those makes it easier for the user to relate a specific dot on papers published exclusively in Chinese, but with at a map to a record in the data (although in many cases, least an abstract, have been cited in the over­ a dot on a map may represent several proximal lo­ all discussion. However, in the following review, we calities). do not discuss or summarize the published literature We provide at the end of the book 14 maps show­ that deals mostly with the management and control ing the known collection localities for all species of (chemical, cultural, or biological) of the economical­ Anoplophora. Some discussion is warranted regard­ ly important Of the 36 species herein rec­ ing our map generation process. For most localities ognized in Anoplophora, the biology, habits, and host we have researched the currently accepted spelling of plants are known for only one-third. The majority of the place name along with latitude and longitude the published papers concern the economically im­ based on the National Imagery and Mapping Agen­ portant species (that A. chinensis including form cy's (www.nima.mil) Geonet Names Server. This is malasiaca, A. macularia, A. glabripennis including a crucial step since so many transliterations exist for form nobilis, and Pseudonemophas versteegii). Al­ many localities in Asia, especially China. Also, many though, we include coverage of the biology of P. names have simply changed and older specimen data versteegii here in this review, it is our opinion (and is not reconcilable with simple comparison to current N. Ohbayashi, pers. comm.), based on external and maps. The NIMA site was crucial to help us establish internal morphology, that this species belongs in localities. Still, in some cases, the locality remained Pseudonemophas. indecipherable or ambiguous. In the case of ambig­ Because A. glabripennis has been a critical focus uous localities, we either made our best guess (often of federal, state, and local agricultural officials since based on distributional data from other specimens), its detection in New York in the summer of 1996, or placed the locality with a "?" in the symbol on and since it has the potential to be a serious tree pest the map, or did not plot the locality. in urban and native forests of North America, we Once we established the locality, the NIMA site treat it first separately. The next section provides a provided latitude and longitude coordinates which we summary of biology for it and the other species for copied into our specimen database. Records from this which biological data are known. database were exported and reformatted into the dec­ Biology of Anoplophora glabripennis.-Since the imal format recognized by the United States Geolog­ late 1970s, the Asian longhorned beetle, Anoplopho­ ical Survey's Crusty Simple Map Generator (http:// ra glabripennis, has become a major forest pest in crusty.er.usgs.gov/mapit/index.html). By entering the mainland China. It is endemic to mainland China coordinate data into the web program, a simple map (Hua et al. 1992) and Korea (Kim et al. 1994 ). Al­ was produced for each We downloaded high though, a few specimen records from Japan have resolution postscript files of those maps and overlaid been seen, there are no known populations of A. gla­ them on detailed topographic maps from the Moun­ bripennis occmTing there. In China, it is found tain High Maps® series. Through layering manipu­ throughout a vast geographic range, 21 °N to 43°N lation using Photoshop® software, the final version latitude and 100°E to 127°E longitude (Yan 1985), of each map was produced. nearly equivalent to all of North America, from the Finally, we present a key to the recognized spt::CH~s Great Lakes and southern Canada to southern in Anoplophora. Where possible, the key has relied Mexico. Indeed, Peterson and Vieglais dem­ on easily seen characters which are defined in such onstrated through ecological modelling, its potential a way that variability is not a hindrance. For variable to survive throughout most of the United States. Its species, we have multiple entries in the key to facil­ prevalence and range in China has increased in the itate identification. past several decades as a result of widespread plant­ ing of susceptible poplar (Populus) hybrids (EPPO BIOLOGY OF ANOPLOPIIORA 2001 ). Yan ( 1985) provided a map showing A. gla­ Perspectives and Overview.-We provide a com­ bripennis to be most damaging in a zone of eastern prehensive review of the biology of species of the China extending from Liaoning to Jiangsu province Oriental and eastern Palearctic genus Anoplophora, and inland to Shanxi, Henan, and Hubei provinces; summarizing the published literature on this econom- it is also present further west to Gansu, Sichuan and LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 17

Fig. 1. a, Anoplophora chine11sis (Forster) (form malasiaca) on Citrus near Matsuyama, Shikoku, Japan, June, 1998. b, A. glabripennis (Motschulsky) (form nobilis) on Salix in Langfang City, Hebei, China, June 26, 1998. c, Adult feeding damage of A. glabripennis on young Salix branchlets in Langfang City, Hebei, China, June 26, 1998.

Yunnan provinces, and farther south (but not in of Qinghai, Xinjiang, and Xizang ( = Tibet) in the southeast China) (EPPO 2001). Li and Wu (1993) west. Map 6 depicts all the locality records we have recorded A. glabripennis nearly throughout the coun­ gleaned from actual specimens. try, but found it absent from the Chinese provinces In China, larval feeding by A. glabripennis causes l8 ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. a, Egg . cars or A11oplop/iora glabripennis (Motschulsky) on Salix in Langfang City, Hebei, China, June 26, 1998 (note adult below). b, mating pair or A. c/iill(!llSis ( orsler) (form malasiaca) at ba.;;;c of Cirrus in Fukuoka Fruit Tree Re carch Station, Fukuoka City, Japan, July 8, 1998. c, same as b, on trunk of Citrus. widespread mortality to 24 deciduous tree species urban street trees, including Populus, Salix, Ulmus, (Yang et al. 1995), but particularly to Populus, Salix, Fraxinus, Hibiscus, Aesculus, Betula, but most im­ and Ulmus (Xiao 1980, Li and Wu 1993). Other Chi­ portant, Acer (Haack et al. 1997 ). Any species of nese woody plants also have been recorded as hosts, Acer is highly attractive to this exotic pest, with the including Acer, Alnus, Malus, Melia, Marus, Platan­ following species as confirmed hosts: Acer negundo, us, Prunus, Pyrus, Robinia, Rosa, and Sophora. An­ A. platanoides, A. saccharinum, A. saccharum, A. oplophora glabripennis is one of the most serious rubrum, and A. pseudoplatanus. Unlike most other pests of poplar (Populus spp.) in China. There, it pri­ Cerambycidae, A. glabripennis attacks not only ap­ marily attacJss species and hybrids in the Aegeiros parently healthy and vigorous trees, but also stressed section of Populus (P. nigra, P. deltoides, P. X can­ trees and recently cut logs (Hua et al. 1992, Gao et adensis, and the Chinensis hybrid P. dakhuanensis) al. 1993, He and Huang 1993). Young and old trees (Li and Wu 1993, Luo and Li 1999). Other sections are attacked indiscriminately. of the genus (Alba and Tacamahaca) are also at­ Throughout its native range, this woodborer usu­ tacked, but are only slightly susceptible (Li and Wu ally has a single generation annually (univoltine) but 1993). In the United States (New York and Chicago), occasionally some individuals require up to two years A. glabripennis attacks and oviposits on a number of (bivoltine) to complete their development (Hua et al. LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 19

Fig. 3. a, Anoplophora macularia (Thomson) (2 spotted form with H3 and H4 separated) on Citrus in Yangmingshan, Taipei County, Taiwan, June 11, 1999. b, A. chinensi (Forster) (form malasiaca) mating on Citrus in Fukuoka Fruit Tree Research Station, Fukuoka City, Japan, July 8, 1998 (note similarity to A. macularia). 20 ENTOMOLOGICAL SOCIETY OF WASHINGTON

1992). Summarizing from Hua et al. (1992), in the tissue, pass whitish feces and sawdust from the egg Chinese provinces of Liaoning, Shandong, Henan, site (Xiao 1980). Maturing larvae tunnel upward into and Jiangsu, A. glabripennis is univoltine or bivol­ a branch or trunk for 10-30 cm. Full-grown larvae tine. In Liaoning Province, over 80% of the popula­ can attain a size of 50-60 mm. A final instar larva tion is univoltine, with first to third instar larvae creates a chamber or cell near the outer bark and overwintering, while another 11-20% of the popu­ transform into a pupa. Newly emerging adults chew lation is bivoltine and with mature larvae overwin­ an exit hole through the inner bark, exiting the host tering. In Jiangsu Province, 98% are univoltine, with tree through 6-18 mm circular exit holes. 2% bivoltine, and the overwintering stage is the ma­ Although adult feeding can cause twig mortality, ture larva. In Henan Province, 14% are univoltine this is minor compared to the damage caused by lar­ and 86% bivoltine. In the spring, mature, overwin­ vae feeding as they tunnel through branches and tered larvae pupate, while other larvae feed until late trunks. If larval densities are high, individual branch­ March and make a pupal chamber in the larval gal­ es or entire trees can die. For heavily infested trees, leries from late April to early May. The prepupal strong winds or heavy snow pack can cause branches stage lasts 9-39 days (Mean: 21.8 days), while the and even whole trees to break or collapse. Larval pupal stage ranges from 13-24 days (mean: 19.6 feeding can reduce the quality of hardwood lumber, days). After emergence, adults remain in the pupal veneer, and wood fiber. The Asian longhorned bee­ chamber for 6-15 days (mode: 7 days) and then chew tle's preference for maple species (Acer) is consid­ their way to the outside. In China, adults begin to ered by most as an enormous threat to maple sugar appear in early April, with peak emergence from production in the northeastern United States if the mid-June to early July, and they are still active in beetle becomes widely established there. In addition, October. property values are reduced in areas where the beetle Newly emerged adults feed first (maturation feed­ is established, and areas where fall foliage tourism is ing) on the tender bark of small twigs and branches important likely will become severely impacted if in­ (Fig. le), and sometimes on tender leaves and leaf fested (Kucera 1996, Appleby 1999). petioles, and then mate on branches and trunks. It is General summary of known biology for Anoplo­ unclear whether long-range pheromones are utilized phora species.-Host trees: Anoplophora species at­ by the sexes to attract each other for mating purposes; tack and damage numerous hardwood trees, across recent investigations indicate that they do not exist, many families of angiosperms, and also some conif­ but that the sexes instead find each other on the host erous trees. Host trees, cited in the published litera­ tree after being attracted there by host volatiles. Once ture, are summarized in the species accounts, under mated, a female chews through the bark of the host the subheading "Hosts." Damage and Economic Im­ tree to the cambial layer, forming the "egg scars" portance: In general, members of the family Cer­ (Fig. then turns and inserts her ovipositor, and ambycidae are phytophagous in their feeding habits lays a single egg, just under the inner bark (phloem). (Linsley 1959). Their larvae usually bore into tissues Typically, females lay eggs in the upper trunk and of woody plants ranging from moribund to dead and along major branches where the bark is thin, smooth, decomposing, while some species (in the genera Aga­ and tender. Also, small branches in the upper canopy panthia and Tetraopes, for example) feed in the are attacked and used for oviposition sites as well. stems of living herbaceous plants (Hanks 1999). In As the tree crown begins to die, the beetle will attack contrast, some cerambycid species (especially in the and oviposit along the entire trunk and even on the subfamilies Cerambycinae and ) infest liv­ exposed roots of larger, older trees. Adults tend to ing but stressed woody angiosperms, and thus are return to the same tree to lay eggs if the tree remains considered important pests of forest, plantation, and viable. The literature above suggests that females lay street trees. The Cerambycinae and Lamiinae include 25-40 eggs, but some can lay as many as 100+ eggs many species that are among the world's most im­ during their life span. Females live 14-66 days while portant tree pests (Hanks 1999). males are shorter lived, at 3-50 days. In New York Longhorned beetles of the genus Anoplophora in­ and Chicago, females lay eggs from July to early clude species that oviposit in healthy and vigorous November. Eggs hatch in one to two weeks, and first­ host trees, but also in weakened trees. Larval feeding instar larvae feed exclusively in rotting cambium and may contribute to decline of host trees, particularly remain in this region for the first two molts, and enter when they are repeatedly attacked by multiple gen­ the sapwood only after the third instar. Second-instar erations of the borer. Other in the genus at­ larvae feed on healthy phloem and xylem, passing tack host trees that are alive and growing but whose brown feces and sawdust from the oviposition site; defenses have been compromised by chronically poor late third and fourth instars, feeding on the xylem growing conditions such as soil compaction, low soil LINGAFELTER AND HOEBEKE: REVISION OF ANOPLOPHORA 235

INDEX Valid names are in Roman boldface; synonyms, nomina nuda, and unjustified emendations are in italics. abbreviata Thomson 49 lumawigi ...... ,.._,.._,,,. ______124 adonis Pie 59 lurida (Pascoe) ______125 a.ffiicta Thomson 49 luteonotatus Pie------·------·------109 albescens Pie 143 mac rospilus Gahan ___ _ 104 alboapicalis Chiang 59 macularia (Thomson) ------·---­ 127 albonotatus Pie 104 mqjor 141 albopicta (Matsushita) 40 malachiticus Nonfried ------______132 ammiralis Schwarzer 143 malasiaca Thomson ______49 amoena (Jordan) 41 131 angustata Pie 137 131 angustatus Pie 109 meden1bachi \'-'aa•~UJLUJ------132 Anoplophora Hope 28 medenbachii (Ritsema) ------132 argentifer Pie 43 Me lanauste r Thomson------·------28 asuanga Schultze 42 melancholica Tippmann ______137 beryllina (Hope) 43 melanostictus White------·------­ 43 bennigseni Aurivillius 143 Mic romelanauste r Pie------·------­ 28,47 birmanica Hiidepohl 46 nankineus Pie 109 bowringii (White) 47 nig rotrifasciatus Pie------·------116

Calloplophora Thomson 28 nobilis "--'ULH,:o;,HJ•"-"-'~'- ---·------·---- 109 chapaensis Breuning 137 ogasawaraensis Makihara ------·------­ 134 chiangi Hua and Zhang 48 Oplopho ra Hope ------28 chinensis (Forster) 49 oshimanus Fairmaire ______127 coeruleoantennata (Breuning) 58 para!uc ipo r Breuning------­ 42 coloremutabilis Tippmann 104 pare le gans Chiang ------··------­ 123 Cyriocrates Thomson 28 perroudi Pie------··------­ 49 da vi dis (Fairmaire)------5 9 pirouletii Fairmaire ------·------­ 118 decemmaculata Pu______------·- 61 pulchricornis Voet ------­ 49 depressinotatus 108 p unc tato r F abriei us ------49 elegans (Gahan) 104 rondoniana (Breuning) ______143 elegans Kano 40 ryukyensis Breuning and Ohbayashi ______127 Falsocyriocrates Pie 28 sepulchralis Thomson------·------______49 farinosus Houttuyn 49 sepulcralis Thomson ______------49 fasciatus Pie 140 s iamens is (Breuning) ------··------·-----­ 143 ftavomaculata (Gressitt) 105 similis (Gahan) 134 freyi (Breuning) 106 sinensis Gmelin ------49 fruhstorferi (Aurivillius) 108 sollii (Hope) ______136 glabripennis (Motschulsky) 109 splendidus Gressitt ------­ 40 glabronotatus Pie 143 stanleyana Hope 137 gloriosa Tippmann 137 superbus Pie ______------104

1 1 1 graafi (Ritsema) 113 s uh be ry ll ina I-

About the Authors

Steven W. Lingafelter obtained his B.S. and M.S. de­ grees in Biology from Midwestern State University (Nor­ man Horner, Advisor), Wichita Falls, Texas in 1989 and 1991, respectively, and his Ph.D. in Entomology from the University of Kansas (J. Steve Ashe, Advisor), Lawrence, Kansas in 1996. He is a Research Entomologist for the Systematic Entomology Laboratory, Plant Sciences Insti­ tute, Agricultural Research Service of the United States Department of Agriculture and is based at the Smithsonian Institution, National Museum of Natural History, in Wash­ ington, D.C. He provides immediate identifications of long­ horned beetles and weevils intercepted at ports of entry in the US to prevent introduction of exotic pests. He is cura­ tor in charge of the National Collection of Cerambycidae and Curculionidae at the Smithsonian Institution which to­ tals about five thousand drawers and one million speci­ mens. The bulk of his time is spent on systematic research on longhorned beetles and weevils. In the last 10 years, he has traveled to Costa Rica, Panama, Brazil, South Ko­ rea, Japan, China, Russia, Armenia, Turkey, Nepal, Eng­ land, France, Canada, and throughout the United States in pursuit of beetles as part of his research program. He has published over 20 peer reviewed papers including one monograph and one book on various beetle families in­ cluding Cerambycidae, Chrysomelidae, Staphylinidae, and Silphidae.

E. Richard Hoebeke is a Senior Extension Associate in the Department of Entomology at Cornell University, and Assistant Curator of the Cornell University Insect Collec­ tion. As manager of the largest land-grant university col­ lection in North America, with nearly 6 million preserved specimens, he is responsible for the curation of all insect groups, but particularly the Coleoptera, as well as for pro­ viding identification support. He received a B.S. degree in Biological Sciences in 1971 and a M.S. degree in System­ atic Entomology in 1973 from Michigan State University, and continued Ph.D. work in Systematic Entomology at Cornell University from 1974-1977. Presently, he focuses on the detection of and survey for non-indigenous species among the North American entomofauna. Recognized as a national leader in exotic pest detection, he has discovered numerous immigrant insects new to the Western Hemi­ sphere, North America, or the United States. He was the first entomologist to identify the highly destructive Asian longhorned beetle, Anoplophora glabripennis, from an in­ fested Brooklyn neighborhood in New York City in 1996. He has authored or co-authored over 60 refereed papers on the recognition, distribution, seasonal history, and hab­ its of adventive insects, and has published more than a dozen papers on the taxonomy of beetles of the family Staphylinidae.