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CERAMBYCIDAE
OF THE WORLD
Biology and Pest Management
 CONTEMPORARY TOPICS in
ENTOMOLOGY SERIES

THOMAS A. MILLER EDITOR

Insect Symbiosis
Edited by Kostas Bourtzis and Thomas A. Miller

Insect Sounds and Communication: Physiology,

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Insect Symbiosis, Volume 2

Edited by Kostas Bourtzis and Thomas A. Miller

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Food Exploitation by Social Insects: Ecological,

Behavioral, and Theoretical Approaches
Edited by Stefan Jarau and Michael Hrncir

Molecular Biology and Genetics of the Lepidoptera

Edited by Marian R. Goldsmith and František Marec

Honey Bee Colony Health: Challenges and

Sustainable Solutions
Edited by Diana Sammataro and Jay A. Yoder

Forensic Entomology: International Dimensions

and Frontiers
Edited by Jeffery Keith Tomberlin and
Mark Eric Benbow
Greenhouse Pest Management
Edited by Raymond A. Cloyd
Cerambycidae of the World: Biology and Pest
Management
Edited by Qiao Wang
CERAMBYCIDAE
OF THE WORLD
Biology and Pest Management

edited by Qiao Wang

Boca Raton London New York

CRC Press is an imprint of the

Taylor & Francis Group, an informa business
CRC Press
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Boca Raton, FL 33487-2742

© 2017 by Taylor & Francis Group, LLC

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Version Date: 20161019

International Standard Book Number-13: 978-1-4822-1990-6 (Hardback)

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Library of Congress Cataloging‑in‑Publication Data

Names: Wang, Qiao, 1960 November 16- , editor.

Title: Cerambycidae of the world : biology and pest management / [edited by]
Qiao Wang.
Description: Boca Raton : Taylor & Francis, 2017. | Series: Contemporary
topics in entomology series
Identifiers: LCCN 2016028247 | ISBN 9781482219906 (hardback : alk. paper) |
ISBN 9781315313252 (e-book) | ISBN 9781315313238 (e-book) | ISBN
9781315313245 (e-book) | ISBN 9781315313221 (e-book)
Subjects: LCSH: Cerambycidae. | Insect pests--Control.
Classification: LCC QL596.C4 C465 2017 | DDC 595.76/48--dc23
LC record available at https://lccn.loc.gov/2016028247

Visit the Taylor & Francis Web site at

http://www.taylorandfrancis.com

and the CRC Press Web site at

http://www.crcpress.com
The editor dedicates this book to the late Professor Shu-nan Chiang (1914–2013),

a cerambycidist, who led him to the world of Cerambycidae.

Contents

Preface ...................................................................................................................................................... ix
Editor ........................................................................................................................................................ xi
Contributors ............................................................................................................................................xiii

1. General Morphology, Classification, and Biology of Cerambycidae ......................................... 1

Marcela L. Monné, Miguel A. Monné, and Qiao Wang

2. Life History and Population Dynamics of Cerambycids ............................................................ 71

Robert A. Haack, Melody A. Keena, and Dominic Eyre

3. Feeding Biology of Cerambycids................................................................................................. 105

Robert A. Haack

4. Reproductive Biology of Cerambycids ....................................................................................... 133

Lawrence M. Hanks and Qiao Wang

5. Chemical Ecology of Cerambycids ............................................................................................. 161

Jocelyn G. Millar and Lawrence M. Hanks

6. Cerambycids as Plant Disease Vectors with Special Reference to Pine Wilt ......................... 209
Süleyman Akbulut, Katsumi Togashi, and Marc J. Linit

7. Laboratory Rearing and Handling of Cerambycids................................................................. 253

Melody A. Keena

8. Natural Enemies and Biological Control of Cerambycid Pests ............................................... 291

Timothy D. Paine

9. Cultural Control of Cerambycid Pests ....................................................................................... 305

Qiao Wang

10. Chemical Control of Cerambycid Pests ..................................................................................... 329

Qiao Wang

11. Cerambycid Pests in Forests and Urban Trees ......................................................................... 351

Robert A. Haack

12. Cerambycid Pests in Agricultural and Horticultural Crops ................................................... 409

Qiao Wang

13. Invasive Cerambycid Pests and Biosecurity Measures ............................................................ 563

Dominic Eyre and Robert A. Haack

Index ...................................................................................................................................................... 619

vii
Preface

There are more than 36,000 described species in the family Cerambycidae worldwide. Although only a
small proportion of these species are pests in agriculture, forestry, or horticulture, their economic impact
is enormous, costing billions of dollars in production losses, damage to landscapes, and management
expenditures. A number of cerambycid species are important pests of various field, vine, and tree crops
as well as forest and urban trees in their native regions. However, with the substantial increase of inter-
national trade in recent decades, many cerambycid species have become established outside their natural
range of distribution, with the potential for causing enormous damage in these novel habitats. So far,
no comprehensive work dealing with all aspects of cerambycid biology and management from a global
viewpoint has been published.
This volume attempts to address that void by covering the entire spectrum from cerambycid classifi-
cation, biology, ecology, plant disease transmission to biological, cultural, and chemical control tactics,
to the world’s major agricultural and tree pests, invasive pests, and biosecurity measures. It is intended
to provide an entrance to the scientific literature on Cerambycidae for scientists in research institutions,
primary industries, and universities, and an essential reference for quarantine officers in governmental
departments charged with detection, exclusion, and control of cerambycids throughout the world. It is
hoped that this book will serve as a valuable reference work for many years to come.
This book is divided into 13 chapters, each of which covers a particular topic consisting of our cur-
rent knowledge and the gaps to be filled. Hundreds of examples, graphs, and photos are presented. The
book begins with an introductory chapter dealing with morphology of adults and immature stages, the
current classification system, the identification of adults and immatures to subfamily, and biology, global
diversity, and distribution of subfamilies. Chapter 2 discusses the types of habitats commonly occupied
by cerambycids; oviposition, fecundity, and egg development; voltinism, overwintering, quiescence, and
diapause; adult dispersal and longevity; and population dynamics in relation to environmental condi-
tions. Chapter 3 focuses on adult and larval feeding habits and wood digestion; flight, pollination, and
plant disease transmission in relation to adult feeding; larval host plant range and conditions, parts
and tissue utilized, and voltinism in relation to development and nutrition. Chapter 4 discusses adult
phenology and diet in association with host and mate location, mating and oviposition behavior, lar-
val development, and reproductive strategies. Historically, it was thought that cerambycids did not use
semiochemicals to mediate reproductive behaviors, but research over the past 15 years suggests that this
was erroneous, and that semiochemical use is very common if not ubiquitous within the family. Thus,
Chapter 5 summarizes recent research on cerambycid pheromones and their chemistry, the role of plant
volatiles as pheromone synergists, mechanisms for maintaining reproductive isolation, and applications
of pheromones and kairomones in pest management and detection of invasive pest species. Chapter 6
describes the biology and control of cerambycids as vectors of pathogens (nematodes) of the pine wilt
disease and plant–beetle–nematode interactions. Chapter 7 presents a thorough review of laboratory
rearing and handling of both cerambycid adults and immature stages with artificial and natural diets.
These are followed by three chapters on pest control tactics. Chapter 8 describes natural enemies in
relation to cerambycid life history, taxonomic range of natural enemies, impact of natural enemies on
cerambycid population dynamics, biological control approaches, and case studies. Using a number of
examples, Chapter 9 covers cultural control measures, including mechanical and sanitary techniques,
irrigation, plant density management, adjusting planting and harvest times, physical barriers, traps,
crop rotation and intercropping, plant resistance, and pest management in relation to climate change.
Chapter 10 discusses chemical control of cerambycid pests and provides a number of examples, cover-
ing the main classes of chemical insecticides and their field applications, including field sprays, bark
treatment, trunk injection and insertion, and soil and root treatment. Chapter 11 presents 43 selected
cerambycid species that illustrate the wide range of life history strategies found among cerambycids

ix
x Preface

infesting forest and urban trees throughout the world; information is provided on the identification of
adults, native and introduced geographic range, larval hosts, life history, economic impact, and control
options. In Chapter 12, 90 cerambycid species of economic importance in field crops, tree crops, and
vine crops from around the world are discussed along with their adult diagnoses, native and introduced
geographic range, damage, biology, and management measures. Chapter 13 deals with invasive ceram-
bycid pests and biosecurity measures, providing detailed information on interceptions and pathways
of invasive pests, inspection and detection methods, pest risk assessments, eradication programs, and
establishment and outbreaks of nonnative species.
Because of the nature of multiauthored contributions, it has not been possible to keep strict uniformity
in all chapters. The editor has tried, however, to adopt a uniform nomenclature for all cerambycid species
throughout the book. This has not been easy because the taxonomy of the Cerambycidae is still in flux,
and the recent synonymizations of several species are reflected in a few chapters. Although chapters are
logically linked, each represents an independent topic. Therefore, to keep the integrity of each chapter,
there is some overlap in subject matter in a few chapters.
The editor is indebted to many people for their advice during the preparation of this book, in particular
to Dr. J. Sulzycki, Dr. M. C. Thomas, Dr. T. A. Miller, J. J. Jurgensen, Jennifer Blaise, and to all of the
book’s contributors. This volume could not have been completed without the generosity of numerous
photographers, reviewers, and copyright holders, whose help is gratefully acknowledged in individual
chapters. I thank all of my family for their love and support, which have kept me going.

Qiao Wang
Massey University

More than 200 full color illustrations, which will be useful for identification purposes, are
available from the CRC Press website under the Downloads tab: https://www.crcpress.com/
Cerambycidae-of-the-World-Biology-and-Pest-Management/Wang/p/book/9781482219906
Editor

Qiao Wang, PhD, is a professor of entomology at the Institute of Agriculture and Environment, Massey
University, Palmerston North, New Zealand. He earned his MSc under Professor Shu-nan Chiang from
Southwest Agricultural University, Chongqing, China, PhD under Professors Ian W.B. Thornton and
Tim R. New from La Trobe University, Melbourne, Australia, and postdoctoral experience under Professor
Jocelyn G. Millar from the University of California, Riverside, before joining Massey University. He
has studied cerambycid beetles since 1982. His research team currently focuses on plant protection,
insect behavior, biological control, and evolutionary biology. Dr. Wang’s experience in Australia, China,
New Zealand, and the United States is reflected in his more than 300 publications; work with more
than 70 postgraduate, postdoctoral, and visiting scientists from around the world; service on editorial
boards of a number of international journals and international expert panels; and chairmanship of inter-
national conference sessions. Dr. Wang was awarded the 2012 Distinguished Scientist Award by the
Entomological Society of America for his outstanding contributions to entomological science during
his career.

xi
Contributors

Süleyman Akbulut Jocelyn G. Millar

Department of Forest Engineering Department of Entomology
Faculty of Forestry University of California
Düzce University Riverside, California
Düzce, Turkey
Marcela L. Monné
Dominic Eyre Departamento de Entomologia
Animal and Plant Health Museu Nacional, Universidade Federal do
Defra, Department for Environment, Rio de Janeiro
Food & Rural Affairs Rio de Janeiro, Brasil
Sand Hutton, York, UK
Miguel A. Monné
Robert A. Haack Departamento de Entomologia
Northern Research Station Museu Nacional, Universidade Federal do
USDA Forest Service Rio de Janeiro
U.S. Department of Agriculture Rio de Janeiro, Brasil
Lansing, Michigan
Timothy D. Paine
Lawrence M. Hanks Department of Entomology
Department of Entomology University of California
University of Illinois at Urbana-Champaign Riverside, California
Urbana, Illinois
Katsumi Togashi
Melody A. Keena
Department of Forest Science
Northern Research Station
Graduate School of Agricultural and
USDA Forest Service
Life Sciences
U.S. Department of Agriculture
The University of Tokyo
Hamden, Connecticut
Tokyo, Japan
Marc J. Linit
College of Agriculture Qiao Wang
Food and Natural Resources Institute of Agriculture and Environment
University of Missouri Massey University
Columbia, Missouri Palmerston North, New Zealand

xiii
1
General Morphology, Classification,
and Biology of Cerambycidae

Marcela L. Monné and Miguel A. Monné

Universidade Federal do Rio de Janeiro
Rio de Janeiro, Brasil

Qiao Wang
Massey University
Palmerston North, New Zealand

CONTENTS
1.1 Introduction ...................................................................................................................................... 2
1.2 Definition and Morphology of the Family Cerambycidae ............................................................... 8
1.2.1 Definition ............................................................................................................................. 8
1.2.2 General Morphology ........................................................................................................... 8
1.2.2.1 Adult..................................................................................................................... 8
1.2.2.2 Immature Stages ................................................................................................ 28
1.3 Key to Subfamilies of the Family Cerambycidae .......................................................................... 29
1.3.1 Adults................................................................................................................................. 36
1.3.2 Larvae ................................................................................................................................ 38
1.4 Diagnosis, Biodiversity, Distribution, and Biology of Subfamilies ............................................... 38
1.4.1 Subfamily Cerambycinae Latreille, 1802 .......................................................................... 40
1.4.1.1 Diagnosis............................................................................................................ 40
1.4.1.2 Comments .......................................................................................................... 46
1.4.1.3 Diversity and Distribution.................................................................................. 46
1.4.1.4 Biology ............................................................................................................... 46
1.4.2 Subfamily Dorcasominae Lacordaire, 1868 ...................................................................... 46
1.4.2.1 Diagnosis............................................................................................................ 46
1.4.2.2 Comments .......................................................................................................... 47
1.4.2.3 Diversity and Distribution.................................................................................. 47
1.4.2.4 Biology ............................................................................................................... 47
1.4.3 Subfamily Lamiinae Latreille, 1825.................................................................................. 47
1.4.3.1 Diagnosis............................................................................................................ 47
1.4.3.2 Comments .......................................................................................................... 55
1.4.3.3 Diversity and Distribution.................................................................................. 56
1.4.3.4 Biology ............................................................................................................... 56
1.4.4 Subfamily Lepturinae Latreille, 1802 ............................................................................... 56
1.4.4.1 Diagnosis............................................................................................................ 56
1.4.4.2 Comments .......................................................................................................... 58
1.4.4.3 Diversity and Distribution.................................................................................. 58
1.4.4.4 Biology ............................................................................................................... 58

1
2 Cerambycidae of the World

1.4.5 Subfamily Necydalinae Latreille, 1825 ............................................................................. 59

1.4.5.1 Diagnosis............................................................................................................ 59
1.4.5.2 Comments .......................................................................................................... 59
1.4.5.3 Diversity and Distribution.................................................................................. 59
1.4.5.4 Biology ............................................................................................................... 59
1.4.6 Subfamily Parandrinae Blanchard, 1845........................................................................... 60
1.4.6.1 Diagnosis............................................................................................................ 60
1.4.6.2 Comments .......................................................................................................... 61
1.4.6.3 Diversity and Distribution.................................................................................. 61
1.4.6.4 Biology ............................................................................................................... 61
1.4.7 Subfamily Prioninae Latreille, 1802 ................................................................................. 61
1.4.7.1 Diagnosis............................................................................................................ 61
1.4.7.2 Comments .......................................................................................................... 63
1.4.7.3 Diversity and Distribution.................................................................................. 63
1.4.7.4 Biology ............................................................................................................... 63
1.4.8 Subfamily Spondylidinae Audinet-Serville, 1832 ............................................................. 64
1.4.8.1 Diagnosis............................................................................................................ 64
1.4.8.2 Comments .......................................................................................................... 65
1.4.8.3 Diversity and Distribution.................................................................................. 65
1.4.8.4 Biology ............................................................................................................... 65
Acknowledgments.................................................................................................................................... 66
References ................................................................................................................................................ 66

1.1 Introduction
Cerambycidae Latreille, 1802, commonly known as longicorns, longhorns, longicorn beetles, longhorned
beetles, longhorned borers, round-headed borers, timber beetles, or sawyer beetles, are among the most
diverse and economically important families of Coleoptera. Taxonomic interest in the family has been
fairly consistent for the past century, but the description of new taxa has accelerated in recent decades.
The number of described cerambycid species in the world is about 36,300 in more than 5,300 genera
(Tavakilian 2015). The adult body length ranges from less than 2 mm in Cyrtinus pygmaeus (Haldeman)
(Linsley 1961) to greater than 170 mm in Titanus giganteus (L.) (Williams 2001). Cerambycids are widely
distributed around the world—from sea level to 4,200 m above—wherever their host plants are found.
Distribution and generic diversity of the world’s cerambycid subfamilies and tribes are shown in Table 1.1.
The longicorn adults are free-living beetles that may or may not need to feed. They can live for a few days
to a few months depending on whether they feed (Hanks 1999; Wang 2008). Cerambycids usually reproduce
sexually but, in very rare cases—such as in some species of Kurarus Gressitt (Cerambycinae) (Goh 1977)
and Cortodera Mulsant (Lepturinae) (Švácha and Lawrence 2014), they can reproduce parthenogenetically.
Švácha and Lawrence (2014) suggested that at least in Cortodera, parthenogenesis probably is of recent
origin because the female has a distinct spermatheca with a spermathecal gland. Mate location depends on
the occurrence and status of larval hosts, adult food sources, and/or pheromones. Hanks (1999) predicted
that the absence of feeding in the adult stage of many species is associated with the production of long-range
pheromones, but the current knowledge shows that the use of volatile pheromones is widespread in ceramby-
cids (see Chapter 5). The females lay their eggs on or near their hosts. The larvae of most cerambycid species
feed on woody plants, but some select herbaceous hosts. The vast majority of species at the larval stage are
living and feeding inside the plants although small minorities are free-living in soil and feed on plant roots.
Many cerambycid larvae are dead plant feeders and play a major role in recycling dead plants; others
attack living plants of different health states, ranging from stressed to healthy plants. To date, there are
about 200 cerambycid species worldwide that have some economic impact on agriculture, forestry, and
horticulture, causing billions of dollars of damage in production losses, environmental disasters, and
management costs. They may damage plants by direct feeding and/or transmission of plant diseases.
General Morphology, Classification, and Biology of Cerambycidae 3

TABLE 1.1
Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes
Subfamilies and Tribes Biogeographic Regions No. Genera
Cerambycinae Latreille, 1802 All biogeographic regions 1,757
Acangassuini Galileo & Martins, 2001 Neotropical 1
Achrysonini Lacordaire, 1868 All biogeographic regions 20
Agallissini Le Conte, 1873 Neotropical 3
Alanizini Di Iorio, 2003 Neotropical 1
Anaglyptini Lacordaire, 1868 All biogeographic regions 12
Aphanasiini Lacordaire, 1868 Afrotropical and Australian 6
Aphneopini Lacordaire, 1868 Australian 5
Auxesini Lepesme & Breuning, 1952 Afrotropical 8
Basipterini Fragoso, Monné & Campos Seabra, 1987 Neotropical 2
Bimiini Lacordaire, 1868 Australian and Neotropical 7
Bothriospilini Lane, 1950 Neotropical 11
Brachypteromatini Sama, 2008 Palaearctic 1
Callichromatini Swainson, 1840 All biogeographic regions 178
Callidiini Kirby, 1837 All biogeographic regions 38
Callidiopini Lacordaire, 1868 All biogeographic regions 62
Cerambycini Latreille, 1802 All biogeographic regions 99
Certallini Fairmaire, 1864 Palaearctic, Afrotropical, and Australian 9
Chlidonini Waterhouse, 1879 Afrotropical (Madagascar) 2
Cleomenini Lacordaire, 1868 Afrotropical and Oriental 23
Clytini Mulsant, 1839 All biogeographic regions 83
Compsocerini Thomson, 1864 All biogeographic regions 33
Coptommatini Lacordaire, 1869 Australian 1
Curiini LeConte, 1873 Neotropical 1
Deilini Fairmaire, 1864 Palaearctic and Australian 3
Dejanirini Lacordaire, 1868 Oriental 2
Diorini Lane, 1950 Neotropical 1
Distichocerini Pascoe, 1867 Australian 2
Dodecosini Aurivillius, 1912 Neotropical 4
Dryobiini Arnett, 1962 Nearctic and Neotropical 3
Eburiini Blanchard, 1845 Neotropical 23
Ectenessini Martins, 1998 Neotropical 12
Elaphidiini Thomson, 1864 Nearctic and Neotropical 91
Eligmodermini Lacordaire, 1868 Neotropical 5
Erlandiini Aurivillius, 1912 Neotropical 1
Eroschemini Lacordaire, 1868 Australian 2
Eumichthini Linsley, 1940 Nearctic 2
Gahaniini Quentin & Villiers, 1969 Afrotropical 1
Glaucytini Lacordaire, 1868 Oriental and Australian 18
Graciliini Mulsant, 1839 All biogeographic regions 22
Hesperophanini Mulsant, 1839 All biogeographic regions 85
Hesthesini Pascoe, 1867 Australian 1
Heteropsini Lacordaire, 1868 Neotropical and Australian 29
Hexoplini Martins, 2006 Neotropical 22
Holopleurini Chemsak & Linsley, 1974 Nearctic 1
Hyboderini Linsley, 1940 Nearctic and Neotropical 4
Hylotrupini Zagajkevich, 1991 Palaearctic 1
Ideratini Martins & Napp, 2009 Neotropical 1
(Continued)
4 Cerambycidae of the World

TABLE 1.1 (Continued)

Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes
Subfamilies and Tribes Biogeographic Regions No. Genera
Lissonotini Swainson, 1840 Neotropical 1
Luscosmodicini Martins, 2003 Neotropical 1
Lygrini Sama, 2008 Afrotropical 1
Macronini Lacordaire, 1868 Australian 4
Megacoelini Quentin & Villiers, 1969 Afrotropical 2
Methiini Thomson, 1860 Oriental, Afrotropical, and Neotropical 19
Molorchini Gistel, 1848 All biogeographic regions 26
Mythodini Lacordaire, 1868 Oriental 4
Necydalopsini Lacordaire, 1868 Neotropical 12
Neocorini Martins, 2005 Neotropical 7
Neoibidionini Monné, 2012 Neotropical 55
Neostenini Lacordaire, 1868 Australian 4
Obriini Pascoe, 1871 All biogeographic regions 43
Ochyrini Pascoe, 1871 Australian 1
Oedenoderini Aurivillius, 1912 Afrotropical 1
Oemini Lacordaire, 1868 All biogeographic regions 101
Opsimini LeConte, 1873 Nearctic and Palaearctic 3
Oxycoleini Martins & Galileo, 2003 Neotropical 2
Paraholopterini Martins, 1997 Neotropical 1
Phalotini Lacordaire, 1868 Australian 4
Phlyctaenodini Lacordaire, 1868 Australian and Neotropical 17
Phoracanthini Newman, 1840 Australian 22
Phyllarthriini Lepesme & Breuning, 1956 Afrotropical 4
Piesarthriini McKeown, 1947 Australian 4
Piezocerini Lacordaire, 1868 Neotropical 19
Platyarthrini Bates, 1870 Neotropical 1
Plectogasterini Quentin & Villiers, 1969 Afrotropical 8
Plectromerini Nearns & Braham, 2008 Neotropical 1
Pleiarthrocerini Lane, 1950 Neotropical 1
Plesioclytini Wappes & Skelley, 2015 Nearctic 1
Proholopterini Monné, 2012 Neotropical 3
Protaxini Gahan, 1906 Oriental 1
Prothemini Lacordaire, 1868 Oriental 3
Psebiini Lacordaire, 1868 Afrotropical and Neotropical 24
Pseudocephalini Aurivillius, 1912 Australian and Neotropical 4
Pseudolepturini Thomson, 1861 Oriental 6
Psilomorphini Lacordaire, 1868 Australian 3
Pteroplatini Thomson, 1861 Afrotropical and Neotropical 10
Rhagiomorphini Newman, 1841 Australian 4
Rhinotragini Thomson, 1861 Neotropical 82
Rhopalophorini Blanchard, 1845 Nearctic, Neotropical, and Australian 29
Sestyrini Lacordaire, 1868 Oriental 2
Smodicini Lacordaire, 1868 Afrotropical, Nearctic, and Neotropical 8
Spintheriini Lacordaire, 1869 Australian 2
Stenhomalini Miroshnikov, 1989 Oriental 2
Stenoderini Pascoe, 1867 Australian and Oriental 10
Stenopterini Gistel, 1848 Palaearctic and Oriental 14
Strongylurini Lacordaire, 1868 Australian 6
(Continued)
General Morphology, Classification, and Biology of Cerambycidae 5

TABLE 1.1 (Continued)

Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes
Subfamilies and Tribes Biogeographic Regions No. Genera
Tessarommatini Lacordaire, 1868 Australian 1
Thraniini Gahan, 1906 Oriental 3
Thyrsiini Marinoni & Napp, 1984 Neotropical 1
Tillomorphini Lacordaire, 1868 Nearctic, Neotropical, Oriental, and Australian 31
Torneutini Thomson, 1861 Neotropical 16
Trachyderini Dupont, 1836 All biogeographic regions 154
Tragocerini Pascoe, 1867 Australian 1
Trichomesiini Aurivillius, 1912 Australian 1
Trigonarthrini Villiers, 1984 Afrotropical 2
Tropocalymmatini Lacordaire, 1868 Australian 1
Typhocesini Lacordaire, 1868 Australian 4
Unxiini Napp, 2007 Neotropical 8
Uracanthini Blanchard, 1853 Australian 6
Vesperellini Sama, 2008 Palaearctic 1
Xystrocerini Blanchard, 1845 Afrotropical and Australian 2
Dorcasominae Lacordaire, 1868 Afrotropical, Oriental, and Palaearctic 95
Apatophyseini Lacordaire, 1869 Afrotropical, Oriental, and Palaearctic 90
Dorcasomini Lacordaire, 1868 Afrotropical, Oriental, and Palaearctic 5
Lamiinae Latreille, 1825 All biogeographic regions 2,964
Acanthocinini Blanchard, 1845 All biogeographic regions 386
Acanthoderini Thomson, 1860 All biogeographic regions 66
Acmocerini Thomson, 1864 Afrotropical 6
Acridocephalini Dillon & Dillon, 1959 Afrotropical 1
Acrocinini Swainson, 1840 Neotropical 1
Aderpasini Breuning & Teocchi, 1978 Afrotropical 1
Aerenicini Lacordaire, 1872 Neotropical 26
Agapanthiini Mulsant, 1839 All biogeographic regions 84
Amphoecini Breuning, 1951 Australian 2
Ancitini Aurivillius, 1917 Australian 1
Ancylonotini Lacordaire, 1869 Afrotropical, Oriental, and Palaearctic 36
Anisocerini Thomson, 1860 Neotropical 26
Apomecynini Thomson, 1860 All biogeographic regions 240
Astathini Thomson, 1864 Australian, Afrotropical, Oriental, and Palaearctic 23
Batocerini Thomson, 1864 Australian, Oriental, and Palaearctic 10
Calliini Thomson, 1864 Neotropical 40
Ceroplesini Thomson, 1860 Afrotropical, Oriental, and Palaearctic 88
Cloniocerini Lacordaire, 1872 Afrotropical 1
Colobotheini Thomson, 1860 Neotropical 12
Compsosomatini Thomson, 1867 Neotropical 13
Cyrtinini Thomson, 1864 Australian and Neotropical 16
Desmiphorini Thomson, 1860 All biogeographic regions 319
Dorcadionini Swainson, 1840 Palaearctic and Oriental 14
Dorcaschematini Thomson, 1860 Oriental and Australian 9
Elytracanthinini Bousquet, 2009 Neotropical 1
Enicodini Thomson, 1864 Australian and Oriental 27
Eupromerini Galileo & Martins, 1995 Neotropical 5
Forsteriini Tippmann, 1960 Neotropical 16
Gnomini Thomson, 1860 Australian, Oriental, and Palaearctic 4
(Continued)
6 Cerambycidae of the World

TABLE 1.1 (Continued)

Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes
Subfamilies and Tribes Biogeographic Regions No. Genera
Gyaritini Breuning, 1950 Australian and Oriental 14
Heliolini Breuning, 1951 Australian 1
Hemilophini Thomson, 1868 Neotropical and Nearctic 127
Homonoeini Thomson, 1864 Australian, Oriental, and Palaearctic 22
Hyborhabdini Aurivillius, 1911 Oriental 1
Lamiini Latreille, 1825 Afrotropical, Australian, Oriental, and Palaearctic 48
Laticraniini Lane, 1959 Neotropical 2
Mauesiini Lane, 1956 Neotropical 4
Megabasini Thomson, 1860 Neotropical 1
Mesosini Mulsant, 1839 All biogeographic regions 99
Microcymaturini Breuning & Teocchi, 1985 Afrotropical 3
Moneilemini Thomson, 1864 Nearctic and Neotropical 1
Monochamini Gistel, 1848 All biogeographic regions 263
Morimonellini Lobanov, Danilevsky & Murzin, 1981 Palaearctic 1
Morimopsini Lacordaire, 1869 All regions except Nearctic 47
Nyctimeniini Gressitt, 1951 Australian and Oriental 1
Obereini Thomson, 1864 All regions except Neotropical 3
Oculariini Breuning, 1950 Afrotropical 2
Onciderini Thomson, 1860 Neotropical and Nearctic 81
Oncideropsidini Aurivillius, 1922 Oriental 1
Onocephalini Thomson, 1860 Neotropical 3
Onychogleneini Aurivillius, 1923 Oriental 1
Parmenini Mulsant, 1839 All biogeographic regions 87
Petrognathini Blanchard, 1845 Afrotropical and Oriental 10
Phacellini Lacordaire, 1872 Neotropical 7
Phantasini Kolbe, 1897 Afrotropical 3
Phrynetini Thomson, 1864 Afrotropical, Oriental, and Palaearctic 14
Phymasternini Teocchi, 1989 Afrotropical 1
Phytoeciini Mulsant, 1839 All biogeographic regions 32
Pogonocherini Mulsant, 1839 All biogeographic regions 33
Polyrhaphidini Thomson, 1860 Afrotropical and Neotropical 2
Pretiliini Martins & Galileo, 1990 Neotropical 1
Proctocerini Aurivillius, 1922 Afrotropical 1
Prosopocerini Thomson, 1864 Afrotropical 18
Pteropliini Thomson, 1860 All biogeographic regions 256
Saperdini Mulsant, 1839 All regions except Neotropical 154
Stenobiini Breuning, 1950 Afrotropical 7
Sternotomini Thomson, 1860 Afrotropical 20
Tapeinini Thomson, 1857 Neotropical and Oriental 2
Tetraopini Thomson, 1860 Nearctic and Neotropical 3
Tetraulaxini Breuning & Teocchi, 1977 Afrotropical 2
Tetropini Portevin, 1927 Palaearctic 2
Theocrini Lacordaire, 1872 Afrotropical 8
Tmesisternini Blanchard, 1853 Australian and Oriental 12
Tragocephalini Thomson, 1857 Afrotropical 63
Xenicotelini Matsushita, 1933 Oriental 1
Xenofreini Aurivillius, 1923 Neotropical 3
Xenoleini Lacordaire, 1872 Australian, Oriental, and Palaearctic 3
(Continued)
General Morphology, Classification, and Biology of Cerambycidae 7

TABLE 1.1 (Continued)

Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes
Subfamilies and Tribes Biogeographic Regions No. Genera
Xylorhizini Lacordaire, 1872 Afrotropical, Australian, Oriental, and Palaearctic 10
Zygocerini Thomson, 1864 Australian and Oriental 9
Lepturinae Latreille, 1802 All biogeographic regions 210
Desmocerini Blanchard, 1845 Nearctic 1
Encyclopini LeConte, 1873 Nearctic and Palaearctic 2
Lepturini Latreille, 1802 All biogeographic regions 140
Oxymirini Danilevsky, 1997 Palaearctic 1
Rhagiini Kirby, 1837 All biogeographic regions 53
Rhamnusiini Sama, 2009 Palaearctic and Oriental 2
Sachalinobiini Danilevsky, 2010 Nearctic and Palaearctic 1
Teledapini Pascoe, 1871 Oriental 3
Xylosteini Reitter, 1913 Palaearctic and Oriental 7
Necydalinae Latreille, 1825 Nearctic, Palaearctic, and Oriental 2
Necydalini Latreille, 1825 Nearctic, Palaearctic, and Oriental 2
Parandrinae Blanchard, 1845 All biogeographic regions 19
Erichsoniini Thomson, 1861 Neotropical 1
Parandrini Blanchard, 1845 All biogeographic regions 18
Prioninae Latreille, 1802 All biogeographic regions 302
Acanthophorini Thomson, 1864 Afrotropical 7
Aegosomatini Thomson, 1861 Afrotropical, Oriental, and Australian 20
Anacolini Thomson, 1857 Afrotropical, Oriental, and Neotropical 33
Cacoscelini Thomson, 1861 Afrotropical and Australian 5
Callipogonini Thomson, 1861 Afrotropical, Palaearctic, and Neotropical 17
Calocomini Galileo & Martins, 1993 Neotropical 1
Cantharocnemini Thomson, 1861 Afrotropical and Australian 6
Closterini, Lacordaire, 1868 Afrotropical, Australian, and Oriental 8
Ergatini Fairmaire, 1864 Afrotropical, Palaearctic, and Nearctic 5
Eurypodini Gahan, 1906 Palaearctic and Oriental 4
Hopliderini Thomson, 1864 Afrotropical 5
Macrodontiini Thomson, 1861 Neotropical 5
Macrotomini Thomson, 1861 All biogeographic regions 78
Mallaspini Thomson, 1861 Neotropical 10
Mallodonini Thomson, 1861 Afrotropical, Oriental, Nearctic, and Neotropical 10
Meroscelisini Thomson, 1861 Afrotropical, Australian, and Neotropical 21
Prionini Latreille, 1802 All biogeographic regions 50
Remphanini Lacordaire, 1868 Oriental 6
Solenopterini Lacordaire, 1868 Neotropical 7
Tereticini Lameere, 1913 Afrotropical and Australian 3
Vesperoctenini Vives, 2005 Neotropical 1
Spondylidinae Audinet-Serville, 1832 All biogeographic regions 32
Anisarthrini Mamaev & Danilevsky, 1973 Palaearctic 4
Asemini Thomson, 1861 All biogeographic regions 12
Atimiini LeConte, 1873 Nearctic, Neotropical, and Palaearctic 3
Saphanini Gistel 1848 Afrotropical and Nearctic 10
Spondylidini Audinet-Serville, 1832 Neotropical, Nearctic, and Palaearctic 3
8 Cerambycidae of the World

With the increase of international trade in recent years, many cerambycid species have been intercepted;
some have become established outside their natural distribution range, causing serious problems globally
(Haack et al. 2010; see Chapter 13).
Linsley (1961, 1962a) and Wang (2008) summarize the general morphology and biology of the
Cerambycidae. More recently, Švácha and Lawrence (2014) have made a very detailed treatment of the
morphology and a general account of the ecology of the Cerambycidae. Ślipiński and Escalona (2013) gave
a good introduction to the morphology and ecology of Australian cerambycids. In this chapter, we summa-
rize the current knowledge about this family, including the definition and morphology, and a brief introduc-
tion to the taxonomy, distribution, and general biology at the subfamily level. We aim to provide readers
with a fundamental knowledge of cerambycids as well as a guide for those who may wish to consult specific
chapters in this book where detailed treatments of Cerambycid biology and pest management are discussed.

1.2 Definition and Morphology of the Family Cerambycidae

1.2.1 Definition
Traditionally, the family Cerambycidae had wider scope, including nine subfamilies: Anoplodermatinae,
Aseminae, Cerambycinae, Lamiinae, Lepturinae, Parandrinae, Philinae, Prioninae, and Spondylidinae
(Napp 1994). In the current classification system (Bouchard et al. 2011; Monné 2012; Švácha and
Lawrence 2014), Oxypeltinae, Vesperinae, and Disteniinae are considered independent families. We use
the new system in this book and discuss eight subfamilies: Cerambycinae, Dorcasominae, Lamiinae,
Lepturinae, Necydalinae, Parandrinae, Prioninae, and Spondylidinae. Table 1.1 summarizes the distri-
bution and generic diversity of cerambycid subfamilies and tribes.

1.2.2 General Morphology

The general morphology of Cerambycidae is extracted from Ślipiński and Escalona (2013) and Švácha
and Lawrence (2014).

1.2.2.1 Adult
1.2.2.1.1 Diagnosis
General external morphology of cerambycid adults is illustrated in Figures 1.1 and 1.2. Antennae usually
filiform, elongate, and 11-segmented, rarely serrate and >12-segmented, usually inserted on pronounced
tubercles; eyes usually emarginate; prothorax without pleural sutures; tibia with two distinct tibial spurs;
tarsi usually pseudotetramerous with fourth tarsomere usually minute and concealed by third tarsomere;
elytra usually covering abdomen; hind wings with a spur on radio-medial crossvein; abdomen usually
with five visible sternites, fifth sternite entire.

1.2.2.1.2 Description
1.2.2.1.2.1 Head The head is prognathous and more or less horizontal in the Parandrinae (Figures 1.3
and 1.4). It is produced anteriorly to form a short to moderately long muzzle in some Lepturinae
(Figures 1.5 and 1.6), Dorcasominae, and Cerambycinae, inclined anteriorly in the Spondylidinae, and is
vertical or retracted, with the genal line directed posteriorly, in the Lamiinae (Figure 1.7). The eyes are
entire in the Parandrinae (Figure 1.3), most Lepturinae, and some Prioninae; feebly emarginate in the
Spondylidinae (Figure 1.8) and most Prioninae (Figure 1.9); emarginate to entire in the Dorcasominae;
and usually are deeply emarginate and reniform in the Cerambycinae (Figure 1.10) and Lamiinae
(Figure 1.7); although occasionally they are divided—as in Tetraopes Schönherr—or lacking the upper
lobe—as in Tillomorpha Blanchard. The facets of the eyes are large and coarse in the Parandrinae, most
Prioninae, and some Asemini and Cerambycinae; usually, they are finer in the Lepturinae, Lamiinae,
and more specialized Cerambycinae.
The antennae usually have 11 antennomeres (Figures 1.1 and 1.2) that are inserted near the base
of the mandibles in the Parandrinae (Figure 1.3), Prioninae (Figure 1.9), and in some Spondylidinae;
General Morphology, Classification, and Biology of Cerambycidae 9

Head

Scape
Pedicel
Eye Claw
Protarsomere V

Pronotum

Scutellum Mesofemur

Elytrum

Elytral
suture
Metafemur

Metatibia

Elytral Metatarsomeres
apex

FIGURE 1.1 General morphology, dorsal view of Trachyderes succinctus (L.) (Cerambycinae).

Labium Mandible

Prosternum

Prosternal Procoxal
process cavity
Mesepisternum

Mesosternal Mesepimerum
process
Metepisternum
Metasternum
Metacoxal
Metacoxa cavity

Sternite

FIGURE 1.2 General morphology, ventral view of Trachyderes succinctus (L.) (Cerambycinae).
10 Cerambycidae of the World

1 mm
FIGURE 1.3 Head, lateral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permission
from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

1mm

FIGURE 1.4 Head, dorsal view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permission
from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)
1 mm

FIGURE 1.5 Head, dorsal view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho,
editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)
General Morphology, Classification, and Biology of Cerambycidae 11

1 mm
FIGURE 1.6 Head, lateral view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho,
editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

1mm

FIGURE 1.7 Head, lateral view of Estola obscura Thomson (Lamiinae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)
1 mm

FIGURE 1.8 Head, lateral view of Asemum striatum (L.) (Spondylidinae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)
12 Cerambycidae of the World

1 mm
FIGURE 1.9 Head, lateral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

1mm

FIGURE 1.10 Head, lateral view of Achryson surinamum (L.) (Cerambycinae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

are near the eyes in the Asemini and Lepturinae (Figure 1.6); and are usually more or less embraced
by the eyes in the Cerambycinae (Figure 1.10) and Lamiinae (Figure 1.7). In some diurnal Lamiinae
(e.g., Octotapnia Galileo & Martins and Pseudotapnia Chemsak & Linsley) and Prioninae, the
antennae may have fewer antennomeres. In some Lamiinae (e.g., Paratenthras Monné), the first
three antennomeres are long, whereas the remaining flagella are reduced and sometimes moniliform.
The number of antennomeres may be 12 in a number of unrelated groups and more than 12 in a few
Cerambycinae and Prioninae (up to 30 in some species of Prionus Müller). The antennal structure is
similar between sexes in the Parandrinae, Spondylidinae, and Lepturinae, and strikingly dissimilar
in many Prioninae and in most Cerambycinae and Lamiinae. In the Parandrinae and Spondylidinae,
differentiation of antennomeres is not well marked; the scape is short, the second antennomere is
not greatly reduced in size, half as long as, or subequal to the third antennomere, and the segments
that follow are subequal in length. In the remaining subfamilies, the scape is usually more elongate,
the second segment is greatly reduced, and the following antennomeres are unequal in length—
with the third usually greatly elongated and those that follow diminishing to the ultimate antenno-
mere. The antennal segments are glabrous in the Parandrinae, Prioninae, and Spondylidinae, and are
pubescent in other subfamilies.
The labrum is fused with the epistoma in the Parandrinae and Prioninae but free in other subfami-
lies. The mandibles are acute in all of the Cerambycidae; large and often toothed in the Parandrinae
(Figure 1.4) and Prioninae (Figure 1.11); long, slender, and untoothed in the Spondylidinae; shorter
in most other groups; and are provided with a dense fringe of hairs in the inner margin of the
Dorcasominae and Lepturinae. The maxillae are typically bilobed; the inner lobe is obsolete in
the Parandrinae (Figure 1.12) and Prioninae (Figure 1.13). The ultimate segment of the palpi (both
General Morphology, Classification, and Biology of Cerambycidae 13

1 mm
FIGURE 1.11 Head, dorsal view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

1mm

FIGURE 1.12 Maxilla, ventral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permis-
sion from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

maxillary and labial) is pointed at the apex in the Lamiinae (Figures 1.14 and 1.15) and truncated
(Figures 1.16 through 1.19) in other subfamilies. The submentum projects between the bases of
the maxillae in the Lepturinae; is short in many Cerambycinae; and is absent in the Parandrinae,
Prioninae, and Spondylidinae. The mentum is distinctly transverse in the Parandrinae (Figure 1.20),
Prioninae, and Spondylidinae, and trapezoidal in the Lepturinae, Cerambycinae (Figure 1.18), and
Lamiinae (Figure 1.14). The ligula is corneous in the Parandrinae and Spondylidinae, and membra-
nous or coriaceous in the Lepturinae, Cerambycinae (Figure 1.18) (except Oemini and Methini), and
Lamiinae (Figure 1.14).
14 Cerambycidae of the World

1 mm

FIGURE 1.13 Maxilla, ventral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)
1mm

FIGURE 1.14 Labium, ventral view of Estola obscura Thomson (Lamiinae). (Reprinted with permission from
C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.)

1.2.2.1.2.2 Thorax The prothorax bears lateral carinae in the Parandrinae (Figure 1.21) and Prioninae
(Figures 1.22 and 1.23), which are lacking in other subfamilies (Figures 1.24 through 1.26). The pro-
coxae are strongly transverse in the Parandrinae and Prioninae, less so in some Spondylidinae—such
as Asemini, subconical in the rest Spondylidinae, conical in the Lepturinae, and usually rounded in the
Cerambycinae and Lamiinae. The procoxal cavities are closed behind in some Parandrinae, in some
Spondylidinae, and in most Lamiinae (Figure 1.26); wide open in the Prioninae (Figure 1.23), Asemini,
and most Lepturinae (Figure 1.25); and open or closed in the Cerambycinae. The scutellum is visible, some-
times well developed (Figure 1.1) and usually is not abruptly elevated, anteriorly flat, or separated from
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