Bark Beetles Biology and Ecology of Native and Invasive Species Bark Beetles Biology and Ecology of Native and Invasive Species Edited by Fernando E Vega Sustainable Perennial Crops Laboratory, United States Department of Agriculture, Agricultural Research Service, Beltsville, MD, USA Richard W Hofstetter School of Forestry, Northern Arizona University, Flagstaff, AZ, USA AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 32 Jamestown Road, London NW1 7BY, UK 525 B Street, Suite 1800, San Diego, CA 92101-4495, USA 225 Wyman Street, Waltham, MA 02451, USA The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK First published 2015 Copyright © 2015 Elsevier Inc All rights reserved No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher Details on how to seek permission, further information about the Publisher’s permissions policies and our arrangement with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein) Notices Knowledge and best practice in this field are constantly changing As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress ISBN: 978-0-12-417156-5 For information on all Academic Press publications visit our website at store.elsevier.com Printed and bound in the United States 14 15 16 17 10 Dedication We dedicate this book to Dr Donald E Bright, in recognition of his outstanding contributions to systematics, biology, zoogeography, and the evolution of bark and ambrosia beetles Don was born in 1934 in Columbus, Ohio, and received his Bachelor of Science degree from Colorado State University in 1957 He served as an entomologist in the U S Army from 1957-1959, and in 1961 received his Master of Science degree from Brigham Young University in Utah, where he worked with Stephen L Wood In 1965 he was awarded a Doctor of Philosophy degree from the University of California at Berkeley and in 1966 he started working as a Research Scientist at the Canadian National Collection of Insects, in Ottawa Don retired in 2003 and moved to Fort Collins, Colorado, in 2006, where he joined Colorado State University as a Faculty Affiliate at the C P Gillette Museum of Arthropod Diversity in the Department of Bioagricultural Sciences and Pest Management Don has published nearly 100 bark and ambrosia beetlerelated publications (below), including “A Catalog of Scolytidae and Platypodidae” with Steve Wood (Wood and Bright, 1987, 1992), as well as three supplements to the catalog (Bright and Skidmore, 1997, 2002, Bright, 2014) Don’s contributions have been instrumental in gaining a better understanding of bark and ambrosia beetles Publications of D E Bright (in chronological order): Bright, D.E., 1963 Bark beetles of the genus Dryocoetes Eichhoff (Coleoptera: Scolytidae) in North America Ann Entomol Soc Am 56, 103–115 Bright, D.E., 1964 Descriptions of three new species and new distribution records of California bark beetles Pan-Pac Entomol 40, 165–170 Bushing, R.W., Bright, D.E., 1965 New records of hymenopterous parasites from California Scolytidae Can Entomol 97, 199–204 Bright, D.E., 1966 New species of bark beetles from California with notes on synonymy (Coleoptera: Scolytidae) Pan-Pac Entomol 42, 295–306 Bright, D.E., 1966 Support for suppression of Xyleborus Bowdich, 1825 Bull Zool Nom 23, 132 Bright, D.E., 1967 Catalogue of the Swaine types of Scolytidae (Coleoptera) with designations of lectotypes Can Entomol 99, 673–681 Bright, D.E., 1967 Lectotype designations for Cryphalus amabilis and C grandis (Coleoptera: Scolytidae) Can Entomol 99, 681 Bright, D.E., 1967 A review of the genus Cactopinus, with descriptions of two new species and a new genus (Coleoptera: Scolytidae) Can Entomol 99, 917–925 Bright, D.E., 1968 Review of the genus Leiparthrum Wollaston in North America, with a description of one new species (Coleoptera: Scolytidae) Can Entomol 100, 636–639 Bright, D.E., 1968 Three new species of Pityophthorus from Canada (Coleoptera: Scolytidae) Can Entomol 100, 604–608 Bright, D.E., 1968 Review of the tribe Xyleborini in America north of Mexico (Coleoptera: Scolytidae) Can Entomol 100, 1288–1323 Bright, D.E., 1969 Biology and taxonomy of bark beetle species in the genus Pseudohylesinus Swaine (Coleoptera: Scolytidae) University of California Publications in Entomology 54, 1–46 Thomas, J.B., Bright, D.E., 1970 A new species of Dendroctonus (Coleoptera: Scolytidae) from Mexico Can Entomol 102, 479–483 v vi Bright, D.E., 1970 A note concerning Pseudohylesinus sericeus (Mannerheim) (Coleoptera: Scolytidae) Can Entomol 102, 499–500 Bright, D.E., 1971 New species, new synonymies and new records of bark beetles from Arizona and California (Coleoptera: Scolytidae) PanPac Entomol 47, 63–70 Bright, D.E., 1971 Bark beetles from Newfoundland (Coleoptera: Scolytidae) Ann Soc Entomol Que 16, 124–127 Bright, D.E., 1972 The Scolytidae and Platypodidae of Jamaica (Coleoptera) Bulletin of the Institute of Jamaica 21, 1–108 Bright, D.E., 1972 New species of Scolytidae (Coleoptera) from Mexico, with additional notes Tribes Xyleborini and Corthylini Can Entomol 104, 1369–1385 Bright, D.E., 1972 New species of Scolytidae (Coleoptera) from Mexico, with additional notes II Subfamilies Scolytinae and Hylesininae Can Entomol 104, 1489–1497 Bright, D., 1972 New species of Scolytidae (Coleoptera) from Mexico, with additional notes III Tribe Pityophthorini (except Pityophthorus) Can Entomol 104, 1665–1679 Bright, D.E., 1973 Xyleborus howdenae, new name, and some corrections to “The Scolytidae and Platypodidae of Jamaica” Coleopterists Bull 27, 18 Bright, D.E., Stark, R.W., 1973 The bark and ambrosia beetles of California (Coleoptera: Scolytidae) California Insect Survey Bulletin 16, 1–169 Bright, D.E., 1975 Comments on the proposed conservation of four generic names of Scolytidae (Insecta: Coleoptera) Z.N.(S.) 20692072 Bulletin of Zoological Nomenclature 32, 135 Bright, D.E., 1976 The Insects and Arachnids of Canada, Part The Bark Beetles of Canada and Alaska (Coleoptera: Scolytidae) Agriculture Canada Publication No 1576, pp 1–241 Bright, D.E., 1976 Biological notes and new localities for three rare species of North American Trogositidae (Coleoptera) Coleopterists Bull 30, 169–170 Bright, D.E., 1976 Lectotype designations for various species of North American Pityophthorus Eichhoff (Coleoptera: Scolytidae) Coleopterists Bull 30, 183–188 Bright, D.E., 1976 New synonymy, new combinations, and new species of North American Pityophthorus(Coleoptera: Scolytidae) Part II Great Basin Nat 36, 425–444 Bright, D.E., 1977 New synonymy, new combinations, and new species of North American Pityophthorus(Coleoptera: Scolytidae) I Can Entomol 109, 511–532 Bright, D.E., 1978 New synonomy, new species, and taxonomic notes of North American Pityophthorus (Coleoptera: Scolytidae) Part III Great Basin Nat 38, 71–84 Bright, D.E., 1978 International voucher specimen collection of Scolytidae Entomol Soc Can Bull 10, 42 Campbell, J.M., Ball, G.E., Becker, E.C., Bright, D.E., Helava, J., Howden, H.F., Parry, R.H., Peck, S.B., Smetana, A., 1979 Coleoptera In: Danks, H.V (Ed.), Canada and its Insect Fauna.In: Memoirs of the Entomological Society of Canada, 108, pp 357–387 Bright, D.E., 1981 Studies on West Indian Scolytidae (Coleoptera) I New species, new distribution records and taxonomic notes Studies on Neotropical Fauna and Environment 16, 151–164 Bright, D.E., 1981 Afrotrypetus, a new genus of bark beetles from Africa (Coleoptera: Scolytidae) Coleopterists Bull 35, 113–116 Bright, D.E., 1981 Taxonomic monograph of the genus Pityophthorus Eichhoff in North and Central America (Coleoptera: Scolytidae) Mem Entomol Soc Can 118, 1–378 Dedication Bright, D.E., 1980 Studies on the Xyleborini Three new species of Schedlia from New Guinea (Coleoptera: Scolytidae) Coleopterists Bull 34, 369–372 Bright, D.E., 1981 Eye reduction in a cavernicolous population of Coccotrypes dactyliperda Fabricius (Coleoptera: Scolytidae) Coleopterists Bull 35, 117–120 Bright, D.E., 1981 A new synonym of Agrilus sayi (Coleoptera: Buprestidae) Can Entomol 113, 871 Bright, D.E., 1982 Studies on West Indian Scolytidae (Coleoptera) New distribution records and descriptions of a new genus and species Studies on Neotropical Fauna and Environment 17, 163–186 Bright, D.E., 1982 Scolytidae (Coleoptera) from the Cocos Islands, Costa Rica, with description of one new species Coleopterists Bull 36, 127–130 Bright, D.E., Stock, M.W., 1982 Taxonomy and geographic variation In: Mitton, J.B., Sturgeon, K.B (Eds.), Bark Beetles in North American Conifers A System for the Study of Evolutionary Biology University of Texas Press, Austin, pp 46–73 Stewart, W.E., Bright, D.E., 1982 Notes on Pissodes fiskei (Coleoptera: Curculionidae) with a redescription of the species Coleopterists Bull 36, 445–452 Bright, D.E., 1985 New species and records of North American Pityophthorus (Coleoptera: Scolytidae), Part IV: The Scriptor Group Great Basin Nat 45, 467–475 Bright, D.E., 1985 New species and new records of North America Pityophthorus (Coleoptera: Scolytidae), Part V: The Juglandis Group Great Basin Nat 45, 476–482 Bright, D.E., 1985 Studies on West Indian Scolytidae (Coleoptera) Checklist of Scolytidae of the West Indies, with descriptions of new species and taxonomic notes Entomologische Arbeiten aus dem Museum G Frey 33 (34), 169–187 Bright, D.E., 1986 A Catalog of the Coleoptera of America North of Mexico: Family Mordellidae United States Department of Agriculture, Agriculture Handbook Number 529–125 Bright, D.E., 1987 New species and new records of North American Pityophthorus (Coleoptera: Scolytidae), Part VI The Lautus group Great Basin Nat 46, 641–645 Bright, D.E., 1987 New species and new records of North American Pityophthorus (Coleoptera: Scolytidae), Part VII Great Basin Nat 46, 679–684 Bright, D.E., 1987 The metallic wood-boring beetles of Canada and Alaska (Coleoptera: Buprestidae) The Insects and Arachnids of Canada, Part 15 Agriculture Canada Publication 1810, pp 1–335 Bright, D.E., 1987 A review of the Scolytidae (Coleoptera) of the Azores with description of a new species of Phloeosinus Bocagiana 107, 1–5 Flores, J.L., Bright, D.E., 1987 A new species of Conophthorus from Mexico: descriptions and biological notes (Coleoptera: Scolytidae) Coleopterists Bull 41, 181–184 Wood, S.L., Bright, D.E., 1987 A Catalog of Scolytidae and Platypodidae (Coleoptera), Part 1: Bibliography Great Basin Nat Mem 11, 1–685 Bright, D.E., 1988 Notes on the occurrence of Xyleborinus gracilis (Eichhoff) in the United States Coleopterists Bull 41, 338 Bright, D.E., 1988 Polydrusus cervinus (Linnaeus), a weevil new to Canada (Coleoptera: Curculionidae) Coleopterists Bull 42, 337 Bright, D.E., 1989 Two new species of Phloeosinus Chapuis from Mount Kinabalu, Borneo, with taxonomic notes (Coleoptera: Scolytidae) Coleopterists Bull 43, 79–82 Dedication Bright, D.E., 1989 New synonymy in North American Sitona (Coleoptera: Curculionidae) Coleopterists Bull 43, 77–78 Bright, D.E., 1989 Additions to the Scolytidae fauna of the Azores (Coleoptera) Bocagiana 129, 1–2 Bright, D.E., 1989 1) Scolytidae; 2) Platypodidae In: Stehr, F.W (Ed.), An Introduction to Immature Insects of North America Kendall/Hunt Publ Co., Dubuque, pp 613–616 Bright, D.E., 1990 A new species of Liparthrum from Borneo with notes on its generic placement (Coleoptera: Scolytidae) Coleopterists Bull 44, 485–488 Atkinson, T.H., Rabaglia, R.L., Bright, D.E., 1990 Newly detected exotic species of Xyleborus (Coleoptera:Scolytidae) with a revised key to species in eastern North America Can Entomol 122, 93–104 Bright, D.E., 1991 A note concerning Sitona tibialis (Herbst) in North America (Coleoptera: Curculionidae) Coleopterists Bull 45, 198–199 Bright, D.E., 1991 Studies in Xyleborini Review of the genus Sampsonius Eggers (Coleoptera: Scolytidae) Studies on Neotropical Fauna and Environment 26, 11–28 Bright, D.E., 1991 Family Derodontidae In: Bousquet, Y (Ed.), Checklist of Beetles of Canada and Alaska Research Branch, Agriculture Canada Publication 1861/E, pp 195–196 Bright, D.E., 1991 Family Melyridae In: Bousquet, Y (Ed.), Checklist of Beetles of Canada and Alaska Research Branch, Agriculture Canada Publication 1861/E, pp 211–213 Bright, D.E., Skidmore, R.E., 1991 Two new records of Scolytidae (Coleoptera) from Canada Coleopterists Bull 45, 368 Bright, D.E., 1992 Synopsis of the genus Hemicryphalus Schedl with descriptions of four new species from Borneo (Coleoptera: Scolytidae) Koleopterologische Rundschau 62, 183–190 Bright, D.E., 1992 The Insects and Arachnids of Canada Part 21 The Weevils of Canada and Alaska Vol (Coleoptera: Curculionoidea, excluding Scolytidae and Curculionidae) Agriculture Canada Publication 1882, pp 1–217 Bright, D.E., 1992 Systematics research In: Hayes, J.L., Robertson, J.L (Eds.), Proceedings of a Workshop on Bark Beetle Genetics: Current Status of Research U.S Department of Agriculture, Forest Service, Pacific Southwest Research Station, p 25, General Technical Report PSW–GTR–135 Bright, D.E., Skidmore, R.E., Thompson, R.T., 1992 Euophryum confine (Broun), a new weevil record for Canada and the New World (Coleoptera: Curculionidae) Coleopterists Bull 46, 143–144 Wood, S.L., Bright, D.E., 1992 A Catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index, Vols A and B Gt Basin Nat Mem 13, 1–1553 Bright, D.E., 1993 Systematics of bark beetles In: Schowalter, T.D., Filip, G.M (Eds.), Beetle-Pathogen Interactions in Conifer Forests Academic Press, London, pp 23–36 Peschken, D.P., Sawchyn, K.C., Bright, D.E., 1993 First record of Apion hookeri Kirby (Coleoptera: Curculionidae) in North America Can Entomol 125, 629–631 Bright, D.E., Skidmore, R.E., Dunster, K.E., 1994 Scolytidae (Coleoptera) associated with the dwarf hackberry, Celtis tenuifolia, in Ontario, Canada Coleopterists Bull 48, 93–94 Bright, D.E., 1994 New records and new species of Scolytidae (Coleoptera) from Borneo Koleopterologische Rundschau 64, 257–274 Bright, D.E., Poinar Jr., G.O., 1994 Scolytidae and Platypodidae (Coleoptera) from Dominican Republic amber Ann Entomol Soc Am 87, 170–194 vii Bright, D.E., 1994 A revision of the genera Sitona Germar (Coleoptera: Curculionidae) of North America Ann Entomol Soc Am 87, 277–306 Hobson, K.E., Bright, D.E., 1994 A key to Xyleborus of California, with faunal comments (Coleoptera: Scolytidae) Pan–Pac Entomol 70, 267–268 Coˆte´, S., Bright, D.E., 1995 Premie`res mentions Canadiennes de Phyllobius intrusus Koˆno (Coleoptera: Curculionidae) et tableaux de de´termination des espe`ces de Phyllobius et Polydrusus au Canada Fabreries 20, 81–89 Bright, D.E., 1996 Notes on native parasites and predators of the European pine shoot beetle, Tomicus piniperda (Linnaeus) in Canada (Coleoptera: Scolytidae) Proc Entomol Soc Ontario 127, 57–62 Cognato, A.I., Bright, D.E., 1996 New records of bark beetles (Coleoptera: Scolytidae) from Dominica, West Indies Coleopterists Bull 50, 72 Bright, D.E., 1997 Xyleborus fornicatus Eichhoff Crop Protection Compendium for Southeast Asia, Data Sheet CABI Electronic Database, 20 p Bright, D.E., 1997 Xylosandrus compactus Eichhoff Crop Protection Compendium for Southeast Asia, Data Sheet CABI Electronic Database, 20 p Bright, D.E., 1997 Xyleborus spp and related genera (Southeast Asia) Crop Protection Compendiumfor Southeast Asia, Data Sheet CABI Electronic Database, 13 p Bright, D.E., Skidmore, R.E., 1997 A Catalog of Scolytidae and Platypodidae (Coleoptera), Supplement (1990–1994) NRC Research Press, Ottawa, 368 p Bright, D.E., Peck, S.B., 1998 Scolytidae from the Gala´pagos Islands, Ecuador, with descriptions of four new species, new distributional records, and a key to species Koleopterologische Rundschau 68, 223–252 Bright, D.E., Rabaglia, R.J., 1999 Dryoxylon, a new genus for Xyleborus onoharaensis Murayama, recently established in the southeastern United States (Coleoptera: Scolytidae) Coleopterists Bull 53, 333–337 Bright, D.E., 2000 Scolytidae (Coleoptera) of Gunung Mulu National Park, Sarawak, Malaysia, with ecological notes and descriptions of six new species Serangga 5, 41–85 Vandenberg, N.J., Rabaglia, R.J., Bright, D.E., 2000 New records of two Xyleborus (Coleoptera: Scolytidae) species in North America Proc Entomol Soc Wash 102, 62–68 Bright, D.E., Skidmore, R.E., 2002 A Catalog of Scolytidae and Platypodidae (Coleoptera), Supplement (1995-1999) NRC Research Press, Ottawa, 523 p Bright, D.E., 2004 Scolytinae In: Cordo, H.A., Logarzo, G., Braun, K., Di Iorio, O (Eds.), Cata´logo de Insectos Fito´fagos de la Argentina y sus Plantas Asociadas” South American Biological Control Laboratory and Sociedad Entomolo´gica Argentina, Buenos Aires, pp 155–162 Schiefer, T.L., Bright, D.E., 2004 Xylosandrus mutilatus (Blandford), an exotic ambrosia beetle (Coleoptera: Curculionidae: Scolytinae: Xyleborini) in North America Coleopterists Bull 58, 431–438 Bright, D.E., Torres, J.A., 2006 Studies on West Indian Scolytidae (Coleoptera) A review of the Scolytidae of Puerto Rico, U S A with descriptions of one new genus, fourteen new species and notes on new synonymy (Coleoptera: Scolytidae) Koleopterologische Rundschau 76, 389–428 Bright, D.E., Bouchard, P., 2008 The Insects and Arachnids of Canada Part 25 The Weevils of Canada and Alaska Vol (Coleoptera: Curculionidae: Entiminae) NRC Research Press, Ottawa, 327 p viii Bright, D.E., 2010 Stevewoodia minutum, a new genus and species of Scolytidae (Coleoptera) from the West Indies Studies on West Indian Scolytidae (Coleoptera) ZooKeys 56, 45–48 Burbano, E., Wright, M., Bright, D.E., Vega, F.E., 2011 New record for the coffee berry borer, Hypothenemus hampei, in Hawaii J Insect Sci 11, 117 Bright, D.E., Kondratieff, B.C., Norton, A.P., 2013 First record of the “Splendid Tamarisk Weevil”, Coniatus splendidulus (F.) (Coleoptera: Dedication Curculionidae: Hyperinae), in Colorado, USA Coleopterists Bull 67, 302–303 Goldarazena, A., Bright, D.E., Hishinuma, S.M., Lo´pez, S., Seybold, S.J., 2014 First record of Pityophthorus solus (Blackman, (1928) in Europe Bulletin OEPP/EPPO 44, 65–69 Bright, D.E., 2014 A Catalog of Scolytidae and Platypodidae (Coleoptera), Supplement (2000–2010), with notes on subfamily and tribal reclassifications Insecta Mundi 0356, 1–336 Contributors Numbers in parentheses indicate the pages on which the authors’ contributions begin Thomas H Atkinson (41), University of Texas Insect Collection, University of Texas at Austin, Austin, TX, USA Matthew P Ayres (157), Department of Biological Sciences, Dartmouth College, Hanover, NH, USA Barbara J Bentz (157, 533), USDA Forest Service, Rocky Mountain Research Station, Logan, UT, USA Peter H.W Biedermann (85), Research Group Insect Symbiosis, Max Planck Institute for Chemical Ecology, Jena, Germany Ryan Bracewell (305), Department of Ecosystem and Conservation Sciences, University of Montana, Missoula, MT, USA Jiri Hulcr (41, 495), School of Forest Resources and Conservation and Department of Entomology, University of Florida, Gainesville, FL, USA Francisco Infante (427), El Colegio de la Frontera Sur (ECOSUR), Carretera Antiguo Aeropuerto Km 2.5, Tapachula, Chiapas, Mexico Andrew J Johnson (427), School of Forest Resources and Conservation, University of Florida, Gainesville, FL, USA Anna Maria J€onsson (533), Department of Physical Geography and Ecosystem Science, Lund University, Lund, Sweden Bjarte H Jordal (41, 85), University Museum of Bergen, University of Bergen, Bergen, Norway Anthony I Cognato (41, 351), Department of Entomology, Michigan State University, East Lansing, MI, USA Lawrence R Kirkendall (85), Department of Biology, University of Bergen, Bergen, Norway Thomas S Davis (209), Plant, Soil and Entomological Sciences, University of Idaho, Moscow, ID, USA Kier D Klepzig (209), United States Department of Agriculture Forest Service Station, Southern Research Station, Asheville, NC, USA Jamie Dinkins-Bookwalter (209), United States Department of Agriculture Forest Service Station, Southern Research Station, Asheville, NC, USA Massimo Faccoli (371), Department of Agronomy, Food, Natural Resources, Animals and Environment (DAFNAE), Agripolis, Legnaro (PD), Italy Christopher J Fettig (555), Invasives and Threats Team, Pacific Southwest Research Station, USDA Forest Service, Davis, CA, USA Jean-Claude Gre´goire (1, 585), Biological Control and Spatial Ecology Laboratory, Universite´ Libre de Bruxelles, Bruxelles, Belgium Matthias Herrmann (247), Max Planck Institute for Developmental Biology, Department of Evolutionary Biology, Tuebingen, Germany Jacek Hilszcza nski (555), Department of Forest Protection, Forest Research Institute, Se˛kocin Stary, Raszyn, Poland Richard W Hofstetter (209), School of Forestry, Northern Arizona University, Flagstaff, AZ, USA Paal Krokene (177), Norwegian Forest and Landscape ˚ s, Norway Institute, A Bo La˚ngstr€om (371), Swedish University of Agricultural Sciences, Department of Ecology, Uppsala, Sweden Franc¸ois Lieutier (371), Laboratoire de Biologie des Ligneux et des Grandes Cultures, Universite´ d’Orle´ans, Orle´ans, France B Staffan Lindgren (1, 585), Natural Resources and Environmental Studies Institute, University of Northern British Columbia, Prince George, BC, Canada Duane D McKenna (41), Department of Biological Sciences, University of Memphis, Memphis, TN, USA Kenneth F Raffa (1, 585), Department of Entomology, University of Wisconsin-Madison, Madison, WI, USA Diana L Six (305), Department of Ecosystem and Conservation Sciences, University of Montana, Missoula, MT, USA xv xvi Sarah M Smith (495), Department of Entomology, Michigan State University, East Lansing, MI, USA Fernando E Vega (427), Sustainable Perennial Crops Laboratory, United States Department of Agriculture, Agricultural Research Service, Beltsville, MD, USA Aaron S Weed (157), Department of Biological Sciences, Dartmouth College, Hanover, NH, USA Contributors Rudolf Wegensteiner (247), University of Natural Resources and Life Sciences, BOKU–Vienna, Department of Forest and Soil Sciences, Institute of Forest Entomology, Forest Pathology and Forest Protection, Vienna, Austria Beat Wermelinger (247), Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Forest Dynamics, Birmensdorf, Switzerland Preface This is the first book broadly dedicated to the ecology, phylogeny, and management of bark beetles (Coleoptera: Curculionidae: Scolytinae) on a global scale The ecological and economic impact of bark beetles on trees is a global issue that often surpasses all other disturbances including fire and storms Bark beetles are of economic importance in forests, orchards, and urban areas as well as agricultural crops, and wood commodities Despite these powerful impacts, most of the approximately 6,000 described species colonize stressed or dead tree tissues only Most bark beetles feed on the phloem or fungi in the inner bark, but a minority specializes on other plant tissues such as cones and seeds The association of bark beetles with microbes has led to a variety of symbioses, and these relationships have led to the great success and diversification of bark beetles Such symbioses may account for the majority of the world’s most recent invasive tree and crop pests Climate change and human-facilitated movement of bark beetles has also contributed to the explosive increase and range expansion of bark beetles, as is the case with the red turpentine beetle in China, the mountain pine beetle in Canada, and the coffee berry borer in tropical regions Recent genomic data on bark beetles and associated microbes have increased our knowledge of the evolution and ecology of these complex communities The present volume includes chapters on ecology, morphology, taxonomy, phylogenetics, evolution, population dynamics, tree defense, symbioses, natural enemies, climate change, management strategies, and the economy and politics of bark beetles In addition, individual chapters are dedicated to bark beetles in the genera Dendroctonus, Ips, Tomicus, Hypothenemus, and Scolytus The editors have brought together an international team of authors, in an effort to combine the vast amount of literature and a diversity of viewpoints into one volume We thank all the authors for their excellent contributions We hope that this book’s information and illustrations are valuable to entomologists, ecologists, foresters, land managers, and students interested in bark beetles We thank Pat Gonzalez and Kristi A.S Gomez at Academic Press for their help and support throughout this project Ann Simpkins cross-checked the references in many chapters, for which we are grateful We appreciate the patience and support of Wendy S Higgins, Ian G Vega, Karen B London, Brian J Hofstetter and Evan M Hofstetter during the creation of this book Fernando E Vega and Richard W Hofstetter xvii Economics and Politics of Bark Beetles Chapter 15 physiology with local biotic and abiotic conditions Because of this plasticity, the socioeconomic impacts of this insect and optimal management approaches vary widely In the western United States and Canada, I pini coincides with several outbreak pine-killing species, such as D ponderosae and D brevicomis LeConte In these regions, I pini is largely a secondary insect, orienting to plant and insect volatiles emitting from trees attacked by the more aggressive species, or colonizing severely stressed trees either alone or in a scramble competition (Rankin and Borden, 1991; Safranyik et al., 1996) Throughout much of the west, I pini is at least partially beneficial to humans, because it reduces reproductive success of primary bark beetles For example, when D ponderosae colonize fireinjured trees, competition with I pini is one of the factors that limit its population increase (Powell et al., 2012) However, this competitive effect can be reduced somewhat, by vertical partitioning of the resource, whereby I pini is often concentrated in the upper stems Under some conditions, I pini can be a pest in western forests, particularly during drought years or in highly dense stands (Kegley et al., 1997) During a chronic outbreak in Montana from 1974 to 1994 in ponderosa pine (Gara et al., 1999), slash management to promote rapid drying of host material was shown to be important For example, how slash is distributed and treated, and how the equipment is used affects colonization rate, rate of drying, and prevalence of natural enemies (Six et al., 2002) The timing of thinning operations can be optimized to minimize population buildup (Gara et al., 1999) Finally, providing a “green chain,” i.e., providing a continuous supply of fresh slash during beetle flight, was recommended to prevent spillover attacks into live trees (Kegley et al., 1997) In the midwestern and northeastern portions of North America, there are no landscape-scale aggressive bark beetles that attack pine In these regions, I pini fills the niche of a primary, tree-killing species However, the live trees this beetle selects almost always show at least moderate acute or chronic stress prior to attack In the Great Lakes region, stress caused by belowground herbivory and accompanying root infection provide a continuous but limited source of susceptible trees (Klepzig et al., 1991) In plantations having high populations of these predisposing agents, I pini can be problematic and sometimes requires direct control by sanitation or pheromone-based mass trapping However, unlike aggressive species, I pini populations not become selfsustaining and encompass entire landscapes after an initial population increase For example, during drought years, both the numbers of I pini and the proportion of trees it kills that did not have prior root infection increase markedly Unlike species such as D ponderosae and D rufipennis, after the drought subsides, I pini populations again become restricted to trees with previously colonized roots or lower stems, and populations decline (Aukema et al., 2010) Reliance on such 605 a predictably and spatially concentrated resource as rootinfested trees appears to facilitate predator impacts (Erbilgin et al., 2002), but the spatial separation of plantations can inhibit predator dispersal to new infestations (Ryall and Fahrig, 2005) Ips pini also shows high plasticity in its pheromone chemistry All I pini produce ipsdienol, but local populations vary in the stereochemistry of their signals (Lanier et al., 1972, 1975; Miller et al., 1989) Most western populations produce almost entirely (À)-ipsdienol In contrast, midwestern and eastern populations produce blends that are either racemic or biased toward (+)-ipsdienol Some areas of western Canada produce substantial amounts of (+)-ipsdienol In addition to enantiomeric differences, midwestern and eastern populations produce lanierone, which is not attractive by itself, but greatly increases attraction to ipsdienol (Teale and Lanier, 1991; Miller et al., 1997) Western populations not produce lanierone, although it is weakly to strongly synergistic to (À)-ipsdienol in Arizona and Montana, and British Columbia, Canada (Miller et al., 1997; Steed and Wagner, 2008) Furthermore, populations in both Arizona and Montana had seasonal shifts in preference (Steed and Wagner, 2008) These patterns appear to arise from local selective pressures, specifically avoidance of interspecific competition with sympatric Ips (Birch et al., 1980; Borden et al., 1992), and escape from predators that exploit beetle pheromones in prey finding (Raffa and Dahlsten, 1995) That is, pheromone blends produced by I pini differ from those of sympatric congenerics, which in the west include other species producing ipsdienol but in the midwest produce ipsenol Likewise, local predators show mismatches from their prey in preferences for stereochemistry and derived components, suggesting time-lagged coevolution (Raffa et al., 2007) Regardless of their evolutionary origins, these variable mixtures, and the distinctions between local predator and prey preferences, provide opportunities to greatly improve both the efficacy and selectivity of pheromonally-based population monitoring and control methods (Dahlsten et al., 2003), but at the same time cause commercial challenges due to the need for locally specific blends, which increase cost of management In addition to the above distinctions between eastern and western populations, I pini also shows plasticity in its life history with latitude and elevation It has an apparently facultative diapause, showing variation in cold tolerance and voltinism, both between regions and between years within regions (Lombardero et al., 2000) The number of generations per year can range from one to five depending on regional temperatures This insect also shows plasticity in its overwintering behavior In northern regions of the Midwest, it overwinters as adults in the soil In other regions, it overwinters both during various life stages under the bark and as adults in soil 606 Bark Beetles From a management perspective, knowledge of the population dynamics of I pini, i.e., its responding to a resource pulse but not becoming self-driving, can be used to guide control strategies Specifically, losses to this insect can be reduced by controlling the predisposing agents, by controlling I pini directly, or both This can involve tactics such as seasonally timing thinning operations to avoid infestation by lower-stem colonizing beetles such as D valens, removing slash, sanitation clearing, or localized application of semiochemicals (Kegley et al., 1997) This contrasts with the eruptive bark beetle species, which can only be successfully managed by preventing initial population increases beyond a critical threshold Thus, from a policy standpoint, I pini does not 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Zhu, J.Y., Luo, X., Tian, S., Gleisner, R., Negro´n, J., Horn, E., 2011 Efficient ethanol production from beetle-killed lodgepole pine using SPORL technology and Saccharomyces cerevisiae without detoxification Tappi J 10, 9–18 Zimmermann, L., Moritz, K., Kennel, M., Bittersohl, J., 2000 Influence of bark beetle infestation on water quantity and quality in the Grosse Ohe catchment (Bavarian Forest National Park) Silva Gabreta 4, 51–62 Zolubas, P., Negro´n, J., Munson, A.S., 2009 Modelling spruce bark beetle infestation probability Baltic For 15, 23–27 Zycha, H., 1943 Die Buchenrindenfaule Der Deutsche Forstwirt 63, 1–2 Index Note: Page numbers followed by f indicate figures and t indicate tables A Acari (mites), 262, 263t Aggression alternate attractor model, 166f, 168–169, 170f capacity of adults, 162–163, 163f definition, 161 host defense tolerance, 163–164, 164f multiple traits, 164–165, 165t overwhelm tree defense, capacity to, 164, 165f single equilibrium abundance model (see Single equilibrium abundance model) Algae, 279 Alternate attractor model, 166f, 168–169, 170f Ambrosia beetles, 601–602 aggregation behavior, 11 definition, 42 evolution and diversity (see Bark and ambrosia beetles) Amplified length fragment polymorphism (AFLP) fingerprinting, 437 Araptus laevigatus, 111 Arthropod natural enemies Acari (mites), 262, 263t Coleoptera, 250–258, 252t dipteran flies, 250, 258–262, 259t Hemiptera and Heteroptera, 262, 263t impact of, 274–277 insect predators, 262, 263t parasitoids (see Parasitoids) Arthropod symbionts, 237 Aulonium ruficorne, 258 B Bacteria biodiversity, 215–216 B thuringiensis, 278–279 Dendroctonus species, 4, 4f, 321–322 ecology of, 216 in European bark beetles, 279 infection studies, 279 S marcescens, 279 Tomicus species, 386 Bark and ambrosia beetles blocking/plugging, role of, 122–123 branch killer, 105–106 delayed dispersal and alloparental care, 125–127 egg placement, 118–119 egg tunnel orientation, 118 evolution and diversity, 67f female fungus farming, 123–124 frass removal, 122 herbiphagy, 99–100 inbreeding, 132 intracellular bacteria, 130 larval cooperation, 127–128 larval feeding modes, 132 live plants, breeding, 106–107 male inbreeders, 124–125 mating behavior, 107–109 mating systems (see Mating systems) monocots, breeding in, 102–103 morphology, 88–91 mycophagy, 102 myelophagy, 101 paternal care, 124 phloeophagy, 94–97 phylogenetics, 87–88 reproductive altruism, 128–130 seeds and seedling killer, 105–106 sexual dimorphism, 91–92 sexual selection, 131–132 social behaviors and ecology, 119–120 social evolution theory, 120 spermatophagy, 101–102 tree-killing bark beetle, 103–105 ventilation holes, 123 xylomycetophagy, 97–99 xylophagy, 99 Bark beetles antennal characters, 59–61 character variation, 64 climate change (see Climate change) coffee berry borer (see Coffee berry borer) computer-based identification keys, 57 conifer defense (see Conifer defense) Curculionidae phylogeny, 65–68 definition, 42 Dendroctonus (see Dendroctonus) dichotomous printed keys, 57 digital catalogs, 57 elytra, 61–63 evolutionary pre-analysis, 44 eyes, 58–59 fossil, 53–56, 54t head, 58 Ips (see Ips) legs, 61 molecular approach, 45, 46–50 morphological approach, 45–46 mycangia, 63–64 origin and evolution, 56 pheromones, 53 population genetics and phylogeography, 50–53 pre-DNA sequence, 44–45 South American, 80 symbiotic interaction (see Symbiotic interaction, bark beetle) taxonomy (see Taxonomy) thorax, 61 Tomicus (see Tomicus) Beauveria bassiana, 280–284 Bigyny, 111 Bothrosternini, 64 Braconidae, 273 C Calcium oxalate crystals, 184 Canningia tomici, 386 Cephalonomia hyalinipennis, 468f, 471 Cephalonomia stephanoderis cage releases, 465 field releases, 465–466, 465t life cycle, 463–465, 464f, 465t rearing methods, 465 Chytridiopsis typographi, 386 Climate change coffee berry borer, 474 Dendroctonus species, 325–326 developed models, 537–545 ecosystem models, 536 insect traits, 533–534 management models, 536–537 northern habitats, 533–534 phenology models, 534–536 Coccotrypes species, 100 Coeloides vancouverensis, 272 Coffee berry borer adult size and mating, 436f, 440 ants, 471–472 bacteria, 455–456 berry borers infection, 439, 457f birds, 472–473 caffeine detoxification, 443 climate change, 474 colonizing female, 439 cultural control, 473–474 damage and losses, 436f, 437–439 distribution, 437, 438t eltyron, vestiture and texture, 432f, 435 endosulfan resistance, 455 entomopathogenic fungi (see Entomopathogenic fungi) 615 616 Index Coffee berry borer (Continued) flight muscles, 446 functional haplodiploidy, 442 fungal endophytes, 459 fungi, 443–444 generations per year, 440–441 genome, 444 hole bored, 436f, 439 host plants, 444 Hypothenemus (see Hypothenemus) immature stages, 435, 436f infestation process, 436f, 439 kairomones, 444 larval instars, 436f, 440 lateral and dorsal view, 431f, 435 life cycle, 436f, 440 longevity, 441–442 nematodes, 459–460 olfactometer bioassays, 445 oviposition, 440 parasitoids (see Parasitoids) parthenogenesis, 442 pheromones, 442 plant resistance, 454–455 rearing, 451 repellents, 453–454 sampling methods, 451 sex ratio, 441, 441t shade (see Shade) taxonomic characters, 431f, 432f, 437 taxonomy and synonymies, 435–437 thrips, 473 traps and attractants, 451–453 vision, 442–443 Coleoptera, 250–258, 252t Colonial mating, 109 Colonial polygyny, 113 Communication acoustics, 13 aggregation behavior, 11 chemical, 13, 14t Dendroctonus, 306t, 308 functions and roles, 11–13, 12t intraspecific variation, 13–15 physical defense, 11 Conifer defense abiotic disturbances, 201 aggregation and mass attack, 198 anatomical layout, 181–186 bark beetle life cycle, 179 bark beetle symbionts, 199 biotic disturbance, 200–201 chemical defenses, 180 chemical traits, 186189 mass attack, 179 naăve host trees, 200 Norway spruce trees, 197–198 pioneer beetles, 197–198 preformed and induced defenses, 196–197 thinning effects, 201–202 tree colonization, 179 tree death, 198–200 tree-killing bark beetles, 178f tree phenology, 201 Conophthorus biology, 518–519 C radiatae, 159 description, 518, 518f diagnosis, 517–518, 518f distribution, 517 D ponderosae, 518f, 519 taxonomic history, 518 Copulation, 108–109 Cortex, 183 Corthylini, 64 Corthylus columbianus, 107 Corticeus C glaber, 258 C parallelus, 258 Cryphalini, 64 Cryptoxilos sp., 468f, 471 D Dendroctonus, 105 anthropogenic change, effects of, 324–326, 338 bacteria, 321–322 body size, 306–307, 306f communication and host location, 306t, 308 coniferous forests, 305 D adjunctus, 4, 320, 336–337 D approximates, 320, 337 D.armandi, 317, 326 D brevicomis, 214f, 321, 337 D frontalis (see Dendroctonus frontalis) D jeffreyi, 319–320, 335 D mexicanus, 320, 336 D micans, 3, 4f, 7f, 257, 318, 329–330 D murrayanae, 318, 330 D parallelocollis, 319, 332 D ponderosae (see Dendroctonus ponderosae) D pseudotsugae, 317, 327–328 D punctatus, 3, 318, 330 D rhizophagus, 319, 332–333 D rufipennis (see Dendroctonus rufipennis) D simplex, 317, 326–327 D.terebrans, 318, 331 D valens, 318–319, 331–332 D vitei, 320, 335 ecological impacts, 324 economic impacts, 324 filamentous fungi, 315–317, 316t fire-damaged trees, 312 genomics, 338 host range and specialization, 307–308 host tree, 313 interactions, 313–315 karyotypic diversity, 306t, 311 life history, 305 microbial symbioses, 315 mites, 323 moisture, effects of, 312 morphology, 305–306 natural enemies, 323–324 nematodes, 322–323 phylogeny and taxonomy, 306t, 308–309, 309f physiographic site, effects of, 312 population genetics and cryptic species, 309–311 sexual size dimorphism, 306t, 307 temperature, effects of, 311–312, 338 yeasts, 321 Dendroctonus frontalis, 210f ecosystem models, 538 fungi, 320–321 larval feeding chamber of, 212f larval feeding galleries of, 212f management models, 538 nitrogen supplementation, 320 outbreaks, 335–336 phenology models, 537–538 Pinus species, 335 population persistence in, 213f prothoracic mycangium of, 210f in southeastern USA, 335 temperature, 336 Dendroctonus ponderosae, 3–4, 4f, 319–320 development rate of, 334 diapause in, 334 ecosystem models, 541 genetic variability, 333 life cycle, 334 management models, 541 outbreaks, 333 overwintering, 334 phenology models, 538–541 Dendroctonus rufipennis, 328–329 ecosystem models, 542 management models, 542 phenology models, 542 Denim Pine, 604 Digital sketch-mapping systems, 558, 559f Dipteran flies, 250, 258–262, 259t DNA barcoding, 52–53 Dolichopodid flies, 258–261 Dryocoetes biology, 504–505 D confusus, 210f, 219f, 504f, 505 description, 503, 504f diagnosis, 503, 504f distribution, 503 taxonomic history, 504 E Economics computable general equilibrium framework, 587 environmental consequences, 586–587 estimation of, 587 mountain pine beetle, 603–604 outbreak incur costs, 587 silvicultural consequences, 585–586 social dimensions, 587–588 wood colonization, 587 Ecosystem models process-based ecosystem models, 536 statistical ecosystem models, 536 Index 617 Eight-toothed spruce bark beetle attitude, 599 environmental and social impact of, 599 features, 599 Greater Bohemian Forest Ecosystem, 599 I typographus, 590, 599 keystone species, 599 nitrate leaching, 599 pest monitoring, 600 political element, 600 quarantine dimension, 600–601 risk modeling, 590–599, 591t, 597t, 600 salvage logging, 600, 600f Sauber Forstwirtschaft, 599–600 storm damage, 590–599, 597t Endemic equilibrium (EnEq), 23–25, 23f Enhanced wetness difference index (EWDI), 558–559 Entomopathogenic fungi, 280–283, 281t artificial infestations, 458–459 artisanal production methods, 459 Beauveria bassiana, 457, 457f, 458 Botrytis rileyi, 456–457 enzymatic action, 458 Hirsutella eleutheratorum, 456 insect death, 456 laboratory bioassays, 458–459 Lecanicillium cillium, 456 Metarhizium anisopliae, 459 mode of action, 456 Paecilomyces fumosoroseus, 456 phylogenetics and systematics, 456 sexual/asexual reproduction, 456 shaded vs unshaded plots, 458 traditional spraying methods, 459 Epuraea species, 258 Eruptive threshold density (ErT), 23–25, 23f Eugregarinida, 285–286 Euwallacea biology, 509 description, 508, 508f diagnosis, 508, 508f distribution, 508 E destruens, 509 E fornicatus, 508f, 509–510 E validus, 510 taxonomic history, 509 F Fossil bark beetles, 53–56, 54t Fungi abiotic factors, 213 biodiversity of, 211 biotic agents, 213 degree of virulence, 213 Dendroctonus species, 3–4, 315–317, 316t detoxifying host chemicals, 213 ecology of, 211 entomopathogenic species (see Entomopathogenic fungi) microsporidian species, 283–284, 284t Tomicus species, 387–390, 387t as tree killers, 213 yeasts (see Yeasts) G Harem polygyny, 97, 111–113 Helicosporidium parasiticum, 279 Hemiptera, 262, 263t Herbaceous plants species, 160–161 Herbiphagy, 98, 99–100 Heteroptera, 262, 263t Heterospilus coffeicola, 470–471 Horizontally transferred bacterial gene encoding mannanase (HhMAN1), 454 Hylastinus obscurus, 160–161 Hylobius abietis, 542–543 Hypothenemus H areccae, 429f, 430 H birmanus, 429f, 430 H concolor, 429f, 430 H crudiae, 430–433, 431f, 432f H curtipennis, 429f, 433 H dissimilis, 429f, 433 H eruditus, 429f, 433 H hampei, 102, 159, 543–544 H interstitialis, 431f, 432f, 433 H javanus, 429f, 433 H obscurus, 102, 431f, 432f, 433–434 H opacus, 429f, 434 host plants, 430 H seriatus, 431f, 432f, 434 identification, 427, 428f, 429f, 431f, 432f life cycle, 430 molecular phylogenetics, 435 taxonomy, 427–429, 428f, 429f I cembrae, 362 I chinensis, 362 I confusus, 362–363, 362f I cribricollis, 362 I duplicatus, 362f, 363 I emarginatus, 363, 363f I grandicollis, 363 I hauseri, 363 I hoppingi, 363–364 I hunteri, 364 I integer, 364 I knausi, 364 I lecontei, 364 I longifolia, 364 I montanus, 364 I nitidus, 351, 352f, 364 I paraconfusus, 364 I perroti, 364–365 I perturbatus, 365 I pilifrons, 365 I pini, 6f, 360f, 365 I plastographus, 365 I schmutzenhoferi, 365 I sexdentatus, 363f, 365 I shangrila, 365 I stebbingi, 365, 366 I subelongatus, 366 I tridens, 6f, 366 I typographus (see Ips typographus) I woodi, 366 pheromones, 351–352 phylogeny of, 355f, 357, 358f population genetics, 357–359 species diagnosis, 352–354, 353t species groups, 356t subgenera, 354 taxonomy, 354–357 Ips typographus, 555, 556t biological control, 574 debarking, 574 diagnosis, 366 distribution, 366 insecticides, 574 lateral habitus of, 362f pheromone-baited traps, 573–574, 574t risk and hazard rating and silviculture, 574–575, 575f sanitation salvage, 572, 573t trap trees, 572–573, 573f verbenone and aggregation pheromones, 574 Ips typographus entomopoxvirus (ItEPV), 278 I L Gas chromatography-mass spectrometry (GC-MS), 445 Glischrochilus lecontei, 258 Global positioning systems (GPS), 558, 559f Gnathotrichus biology, 520 description, 519, 520f diagnosis, 519, 520f distribution, 519 G sulcatus, 520f, 521 taxonomic history, 520 Gregarina typographi, 285–286 Grosmannia G clavigera, 319–320 G yunnanensis, 389, 390 Gymnosperm cone, 102 H Inbreeding polygyny, 113–115 Ips biology, 351 evolution of, 359–360 I acuminatus, 360–361, 360f I amitinus, 360f, 361 I apache, 360f, 361 I avulsus, 361 I bonanseai, 361 I borealis, 361, 362f I calligraphus, 361–362 Leptographium wingfieldii, 389, 390 Life history and ecology bacteria, 4, 27 chemical signaling, 27 communication (see Communication) dispersal flight, feeding substrate, 2–3 forest ecosystem services, 25, 26t forest fire risk, 25 fungi, 3–4, 27 gender role, 618 Index Life history and ecology (Continued) haplo-diploid ambrosia beetles, host defenses, 6–8, 7f host location and selection, 5–6, 6f host plant utilization, symbionts in, 9–10 host substrate quality, 8–9 invasive species, 26 mating and social strategies, 27 mites, natural enemies (see Natural enemies) nematodes, phoretic mites, population dynamics (see Population dynamics) reproductive strategy, resource partitioning, 10–11 saprophages, Scolytinae, 1, socioeconomic impacts, 25–26, 28 tree-killing bark beetles, Log Wizard™, 563f Lonchaea species, 261 M Malamoebia scolyti, 386 Management strategy acoustics, 562 aerial survey, 558–559, 559f biological control, 562–563 Dendroctonus, 555, 556t ecology, 556–557, 557f fire, 565–566 ground-based surveys, 559–560 insecticides, 564–565, 565f I typographus (see Ips typographus) landscape heterogeneity, 571 outbreaks development, 557–558 prescribed fire, 571 risk and hazard rating systems (see Risk and hazard rating systems) salvage, 564 sanitation, 563–564, 563f, 564f Scolytus, 555, 556t semiochemicals (see Semiochemicals) social acceptance, 571 thinning, 569–571, 570t Mating behavior copulation, 108–109 courtship, 108 fighting, 107–108 repeated mating, 109 Mating systems, 131 adult mating systems, 132 bigyny, 111 colonial mating/polygamy, 109 colonial polygyny, 113 female-initiated mating system, 110–111 harem polygyny, 111–113 inbreeding polygyny, 113–115 male residency, 109–110 monogyny, 110 parthenogenetic reproduction, 116–117 partial inbreeding, 115–116 pioneer sex, 110 Medetera M aldrichii, 261 M bistriata, 261 M dendrobaena, 261 M nitida, 258–261 M signaticornis, 261 M zinovjewi, 261 Microsporidia, 283–284, 284t Mites, 218f biodiversity, 216–217 Dendroctonus species, 4, 323 ecology of, 217–219 Ips species, phoretic bark beetle hosts, 209–211, 219t population dynamics, impact on, 219–233 Tomicus species, 392 Monarthrum biology, 522–523 description, 521, 521f diagnosis, 521, 521f distribution, 521 M fasciatum, 521f, 523 M mali, 521f, 523 M scutellare, 521f, 523 taxonomic history, 522 Monogyny, 98, 110 Mountain pine beetle annual allowable cut, 604 climate, 604 Denim Pine, 604 D ponderosae, 603 large-scale eruptions, 603 lodgepole pine, 603–604 susceptible age classes, 603, 603f Mutualism theory, 209 Mycangia, 209, 210f Mycophagy, 98, 102 N Natural enemies behavior and impact mitigation, 19 biological control program, 19 Dendroctonus species, 323–324 habitat characteristics, 17–18 impacts of, 18–19 limited resources, 17 monoterpene toxicity, 16–17 nematodes (see Nematodes) parasitoids (see Parasitoids) pathogens (see Pathogens) population dynamics (see Population dynamics) predators (see Predators) shifting prey, 17 Tomicus species, 400–401 tritrophic signaling, 19 vertebrate predators, 16 Nematodes biodiversity, 233 biology, 287–289 commensals, 233–234 Dendroctonus species, 4, 322–323 evolution, 287 mutualism, 234 parasites, 234–236 population dynamics, impact on, 236 taxa, 288t Tomicus species, 390–392, 391t Nemozoma elongatum, 257 Neogregarinida, 285t, 286 Non-pine conifers, 186 Norway spruce trees, 197–198 Nudobius lentus, 258 O Ophiostoma, 590–599, 598f O canum, 387–389 O minus, 212f, 213, 387, 389, 390 O tingens, 387 P Pagiocerus frontalis, 159 Parasitaphelenchus papillatus, 390–392 Parasitoids, 98 Braconidae, 273 C hyalinipennis, 468f, 471 Cryptoxilos sp., 468f, 471 C stephanoderis (see Cephalonomia stephanoderis) ecology, 262–266 Heterospilus coffeicola, 470–471 host location, 266–273 Hymenoptera, 274, 275t Phymastichus coffea (see Phymastichus coffea) prorops nasuta (see Prorops nasuta) Pteromalidae, 273–274 Tomicus species, 393–397, 394t wasps, 267t Pathogens algae, 279 bacteria (see Bacteria) fungi (see Fungi) horizontal transmission of, 286 incidence, 286–287 negative impacts, 286 protists (see Protista) vertical transmission, 286 viruses (see Virus) Periderm, 182–183 Phenolics, 188–189 Phenology models cold hardening, 536 development time, 535 diapause, 535–536 Phloem-feeding bark beetles, 160, 161, 165t, 166 Phloeophagy transporting fungi, 95–97 wood consumption, 94–95 Phloeotribus liminaris, 104 Phylogenetics definition, 42 Dendroctonus species, 306t, 308–309, 309f Ips species, 355f, 357, 358f Tomicus species, 374–375, 374f Phymastichus coffea cage releases, 469–470 Index 619 countries, 466, 468t dark brown wasps, 466, 467f females oviposit, 466–469, 468f field releases, 470 history, 466, 467f immature stages, 466 parasitization, 466–469 rearing methods, 469 Pine conifers, 186 Pine engraver, 604–606 Pinhole borers, 98 Pioneer beetles, 12–13 Pityogenes chalcographus, 257 Pityophthorus, 110–111 biology, 516–517 description, 515, 515f diagnosis, 515, 515f distribution, 514 P juglandis, 517 taxonomic history, 515–516 Placusa depressa, 258 Platypodinae See Bark and ambrosia beetles Platysoma P attenuatum, 257–258 P cylindricum, 257–258 P parallelum, 257–258 Plegaderus transversus, 257–258 Poecile gambeli, 249–250 Politics exotic species, 590 management, 589–590 mountain pine beetle, 603–604 predictive models, 590, 591t Polygamy, 98, 109 Polygraphus biology, 500–501 diagnosis and description, 500, 500f distribution, 500 P rufipennis, 501 taxonomic history, 500 Polyphenolic parenchyma (PP) cells, 183–184 Population dynamics diversity, 19–20 endogenous feedbacks, 158, 158f EnEq and ErT, 23–25, 23f exogenous effects, 158, 158f fungi, impacts of, 212–213 herbaceous feeding bark beetles, 160–161 mites, impacts of, 219–233 nematodes, 236 population growth, 157–158 seed-feeding species, 159 stability, 158 survival,development, and reproduction, 20–22, 21f Tomicus species (see Tomicus) tree infesting species (see Tree-infesting bark beetles) yeasts, impacts of, 215 Predators Acari (mites), 262, 263t ants, 471–472 birds, 472–473 bluebirds, 249–250 chickadees, 249–250 Coleoptera, 250–258, 252t competitors, 19 dipteran flies, 250, 258–262, 259t Hemiptera and Heteroptera, 262, 263t insect predators, 262, 263t jays, 249–250 juncos, 249–250 as mortality and potential regulating agents, 22 nuthatches, 249–250 snakeflies, 250, 262, 263t thrips, 473 Tomicus species, 393–397, 395t tree creepers, 249–250 tyrant flycatchers, 249–250 woodpeckers (see Woodpeckers) Prorops nasuta cage exclusion/inclusion techniques, 462 clypeus and antennal base, 460, 461f field releases, 462–463, 462t hatching eggs, 460–461, 461f idiobiont solitary parasitoid, 460–461 immature stages, 460, 461f incubation, 460–461, 461f laid eggs, 460–461, 461f larva, 460, 461f rearing method, 461–462 short median snout-like projection, 460, 461f Protista Eugregarinida, 285–286 field-collected bark beetles, 285t in I typographus, 284 microsporidia and protozoa, 284 neogregarine protist, 286 rhizopodan species, 285 Pseudohylesinus biology, 502–503 description, 500f, 502 diagnosis, 500f, 501–502 distribution, 501 P granulatus, 503 taxonomic history, 502 Pteromalidae, 273–274 Pytho depressus, 258 R Radial rays, 184 Resin ducts, 184 Rhizophagus R depressus, 257 R dispar, 257 R grandis, 257 Rhizopoda, 285 Rhopalicus tutela, 272f Risk and hazard rating systems attractant-baited multiple-funnel traps, 562 beetle pressure index, 560–561, 561t definition, 560 direct control, 562–569 geographic location, 561–562 probability rating, 560–561, 561t site quality and tree-diameter distributions, 561–562 stand density, 560–561, 560t Roptrocerus xylophagorum, 272–273, 272f S Salpingus planirostris, 258 Salvage industrial salvage, 589 silvicultural salvage, 588–589 Sapwood, 185 Sclerenchyma cells, 184 Scoloposcelis S flavicornis, 262 S pulchella, 262 Scolytinae See also Bark and ambrosia beetles classification, 68–70 genera and tribes, 71t morphology, 64–65 traditional classification of, 45 Scolytini, 64–65 Scolytus biology, 497–498 diagnosis and description, 495–496, 496f distribution, 495 S multistriatus, 498 S quadrispinosus, 499 S schevyrewi, 499 S ventralis, 498 taxonomic history, 496–497 Seed-feeding species, 159 Semiochemicals antiaggregation pheromones, 567 attractant-baited multiple-funnel trap, 566, 566f barriers, 567, 567t efficacy and cost effectiveness, 569 forest structure and abiotic factors, 569 formulation, 568–569 functional terminology of, 12–13, 12t habitat suitability, 567–568 host presence, 567–568 host suitability and susceptibility, 567–568 meteorological conditions, 568–569 3-methylcyclohex-2-en-l-one, 567 pouch release devices, 568–569, 568f redefining selection criteria, 569 relative ubiquity of, 13, 14t semiochemical performance assessment, 569 verbenone, 567 Serratia marcescens, 279 Sexual dimorphism bark and ambrosia beetles, 91–92 Dendroctonus, 306t, 307 Shade and ants, 450 benefits, 446–447 buffer temperatures, 450 effectiveness of insecticides, 450 effects, 447, 448t and fungal entomopathogens, 450 620 Index Shade (Continued) infestation levels, 449 pest management strategy, 449 shaded plantations vs non-shaded plantations, 447 Single equilibrium abundance model non-aggressive species, 166f, 167 opportunistically aggressive species, 166f, 167–168 population growth rate, 166–167, 166f Snakeflies, 250, 262, 263t Spermatophagy, 98 araptus, 101 coccotrypes, 101 cone breeders, 102 H hampei, 102 H obscurus, 102 pagiocerus, 101 Spruce beetle, 602–603 Symbiotic interaction, bark beetle arthropods, 237 bacteria (see Bacteria) fungi (see Fungi) mites (see Mites) multi-species interactions, 237 mutualism theory, 209 mutualistic species, 209 nematodes (see Nematodes) terms and definitions, 209, 210t, 211 viruses, 236 yeasts (see Yeasts) T Taxonomy, 41–44 coffee berry borer, 435–437 Conophthorus, 518 definition, 42 Dendroctonus species, 308–309 Dryocoetes, 504 Euwallacea, 509 Gnathotrichus, 520 Hypothenemus, 427–429, 428f, 429f Ips species, 354–357 Monarthrum, 522 Pityophthorus, 515–516 Polygraphus, 500 Pseudohylesinus, 502 Scolytus, 496–497 taxonomists, 41–42 Tomicus species, 371–374 Trypodendron, 513 Xyleborus, 506 Xylosandrus, 511 Temnochila T chlorodia, 257 T virescens, 257 Terpenes, 186–188 Thamnurgus euphorbiae, 160 Thanasimus T dubius, 251–257, 251f, 393 T formicarius, 251–257, 251f, 393 Tomicobia seitneri, 272f Tomicus anthropogenic dispersal, 405 attack density, 397–398 bacteria, 386 climate effects, 405, 414 density-independent factors, 401–402 Dermestes piniperda, 371 detection and survey, 409–410 ecological consequences, 409 egg gallery, 377–379, 379f fecundity and brood productivity, 398–399 geographic distribution and host range, 377, 378f insect-pathogenic fungi, 390 intraspecific competition, 399–401, 400f life traits, 371 mite species, 392 morphology and species separation, 375–377 natural dispersal, 404–405 natural enemies, 400–401 nematodes, 390–392, 391t non-pathogenic fungi, 387–390, 387t parasitoids, 393–397, 394t phylogenetic tree of, 374–375, 374f population regulating factors, 399 predators, 393–397, 395t shoot attacks, 371, 379, 380f shoot damage, 406–407 synonymies and host species, 373t taxonomy, 371–374 T brevipilosus, 373t, 385–386 T destruens, 373t, 383–384, 412–413 timber quality, deterioration of, 405, 409 T minor, 372f, 382–383, 412 T pilifer, 373t, 386 T piniperda, 372f, 379–382, 410–412 T puellus, 373t, 386 tree mortality, 405–406, 407–409 tree resistance, biotic and abiotic factors, 402–404 T yunnanensis, 372f, 385, 413 unicellular eukaryotes, 386 vertebrates, 397 viruses, 386 Tree colonization, 179 Tree-infesting bark beetles aggression (see Aggression) agriculture, pests of, 159–160 conifer forests, impact on, 159–160 host plant resistance, 161–162, 170–171 life history strategies, 161 mass attack, 159–160 non-native diseases, vectors of, 160 phloem-feeding bark beetles, 160 pine plantations, 159–160 tree mortality, 159–160 Tree phenology, 201 Trypodendron biology, 513–514 description, 510f, 513 diagnosis, 510f, 513 distribution, 513 taxonomic history, 513 T domesticum, 601 T lineatum, 2–3, 514 U Uganda wasp See Prorops nasuta Uricolytic bacteria, 216 Uropodoid mites, 233 V Vascular cambium, 185 Virus Dendroctonus species, 277–278 DNA/RNA, 277–278 ItEPV, 278 occlusion body, 277–278 symbiotic interaction, 236 Tomicus, 386 W Wasps, 267t Wolbachia outbreeding depression, 130 phenotypic effects, 130 sex ratios, 130 Wood-boring beetles, 88–89 Woodpeckers, 247–250, 248t, 397 X Xyleborini, 65 Xyleborinus saxesenii delayed dispersal, 125 hemicellulases, 128 larval cooperation, 127–128 Xyleborus biology, 506–507 description, 506, 508f diagnosis, 506, 508f distribution, 505 mass attack and weak pathogen, 507 systemic virulence, 507 taxonomic history, 506 water-stressed trees, 507 X glabratus, 507–508, 508f Xylomycetophagy, 97–99 Xylophagy, 98, 99 Xylosandrus biology, 511 description, 510, 510f diagnosis, 510, 510f distribution, 510 taxonomic history, 511 X compactus, 510f, 511–512 X crassiusculus, 512 X germanus, 512–513 Y Yeasts biodiversity, 214, 214f Candida sp., 283 Dendroctonus species, 321 ecology of, 214–215 Metschnikowia species, 283 Pichia sp., 283 population dynamics, impact on, 215 Tomicus species, 387 .. .Bark Beetles Biology and Ecology of Native and Invasive Species Edited by Fernando E Vega Sustainable Perennial Crops Laboratory, United States Department of Agriculture, Agricultural... gaining a better understanding of bark and ambrosia beetles Publications of D E Bright (in chronological order): Bright, D.E., 1963 Bark beetles of the genus Dryocoetes Eichhoff (Coleoptera: Scolytidae)... of resin (Chapter 5) The quantity and importance of this resin vary greatly among conifer genera (Berryman, 1972), and even among species within a genus, Natural History and Ecology of Bark Beetles