Forest Entomology

Forest entomology

To: A. H. MacAndrews Who introduced me to the world of forest insects

Forest entomology A global perspective

By William M. Ciesla

This edition first published 2011 Ó 2011 by William M. Ciesla Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell. Registered office: John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Ciesla, William M. Forest entomology : a global perspective / by William M. Ciesla. p. cm. Includes bibliographical references and index. ISBN 978-1-4443-3314-5 (cloth) 1. Forest insects. I. Title. SB761.C537 2011 634.9'67—dc22 2010048037 A catalogue record for this book is available from the British Library. This book is published in the following electronic formats: ePDF 9781444397871; Wiley Online Library 9781444397895; ePub 9781444397888 Set in 9/11 PhotinaMT by Thomson Digital, Noida, India 1 2011

Contents

Acknowledgements, vii

9. BARK AND AMBROSIA BEETLES, 173

The author, ix

10. LARGE CAMBIUM AND WOOD BORING INSECTS, 203

Preface, xi

11. SUCKING INSECTS, 233

1. THE WORLD’S FORESTS AND THEIR DYNAMICS, 1

12. GALL INSECTS, 253

2. FOREST INSECT DYNAMICS, 15

13. TIP, SHOOT AND REGENERATION INSECTS, 273

3. FOREST INSECT AND HUMAN INTERACTIONS, 31 4. MONITORING FOREST INSECTS, THEIR DAMAGE AND DAMAGE POTENTIAL, 49 5. FOREST INSECT MANAGEMENT, 73

14. INSECTS OF TREE REPRODUCTIVE STRUCTURES, 295 15. INSECTS OF WOOD IN USE, 313 References, 329

6. FOREST INSECT ORDERS AND FAMILIES, 97

Subject and Taxonomic Index, 355 Host index, 371

7. FOLIAGE FEEDING INSECTS – LEPIDOPTERA, 113

Colour plate sections appear between pp. 100–101, 132–133, 180–181, 228–229 and 276–277

8. OTHER FOLIAGE FEEDING INSECTS, 149

COMPANION WEBSITE This book has a companion website: www.wiley.com/go/ciesla/forest with Figures and Tables from the book for downloading

Acknowledgements A half-century of work as a forester, forest entomologist, remote sensing specialist and forest health program manager is reflected in the pages of this text. My chosen profession has provided me with opportunities to work and travel across much of the USA and also Mexico, portions of Central and South America, Europe, Asia and Africa. I have also had the good fortune to meet many outstanding people along the way who became my mentors, colleagues and friends. Aubrey H. MacAndrews, Professor of Forest Entomology at the State University of New York, College of Environmental Science and Forestry at Syracuse University introduced me to the world of forest insects when I was an undergraduate forestry student. As a young forest entomologist, I had the good fortune to work with people like Gene Amman, A.T. Drooz and Hoover Lambert, USDA Forest Service. Robert C. Heller, USDA Forest Service and later, University of Idaho, first introduced me to color aerial photos as a tool to map and assess forest insect damage. Russell K. Smith, USDA Forest Service, provided me the opportunity, guidance and confidence to manage forest health programs in several Forest Service regions and at the national level. In partnership with contemporaries, including J. W. Barry, W. H. Klein, W. B. White, F. P. Weber, J. D. Ward and many others, we evaluated and integrated remote sensing, geographic information systems and the technology of pesticide application into forest health protection programs. Jorge Macias (Mexico), Attilio Disperati, Edson Tadeu Iede, Yeda Oliviera and Augusta Rosot (Brazil), Aida Baldini Urrutia, Osvaldo Ramierez and Patrico Ojeda (Chile), Joseph Mwangi (Kenya), Muhammad Yousuf Khan and Hafeez Buzdar (Pakistan), Zhou Jian Sheng and Wang Gaoping (China), Cernal Akesen and Musa Erkanat (Cyprus), Heinrich Schmutzenhofer and Edwin Donaubauer (Austria), Mihai Barca (Romania) and many other international colleagues introduced me to the cultures and forests of their respective countries.

Rainier P€ ohlmann, Director of Public Affairs, Bavarian National Park, Germany, shared his insights on bark beetles, forest dynamics and people and how they affect management policy of a national park. Many friends and colleagues provided photos of subjects not available from my own archives. These included Ronald F. Billings, Texas Forest Service, S. Sky Stephens, Colorado State Forest Service, E. Richard Hoebecke, Cornell University, Paula Klasmer, Instituto Nacional de Tecnologia Agropecuaria, Argentina, David Cappaert, Michigan, R. Scott Cameron, International Paper Company, Larry Barber, Sheryl Costello, Jerald E. Dewey, Jose Negrón and Brian Howell, USDA Forest Service. Several of the images in this text were accessed via the www.forestryimages.org website of the University of Georgia, which is the work of Extension Entomologist, G. Keith Douce. William Jacobi, Department of Bioagricultural Sciences and Pest Management, Colorado State University, made available samples of damage caused by ambrosia beetles, carpenter ants, termites and wood borers, photos of which are included in this text. The team at Wiley-Blackwell took the title of this work “A Global Perspective” seriously. They arranged for constructive and supportive peer review of the manuscript by Robert G. Foottit, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada. Ward Cooper, Senior Commissioning Editor and Delia Sandford, Managing Editor of Wiley-Blackwell, Oxford, UK provided overall leadership to the production and Kelvin Matthews, Production Manager, also in Oxford, arranged for peer review and design of the cover. Maggie Beveridge, currently residing in Lusaka, Zambia, served as copy editor and was instrumental in transforming a still unfinished draft into a polished document, ready for printing. Jessminder Kaur, Production Editor, Wiley-Blackwell, Singapore, supervised the layout and printing. Prakash Naorem, Thomson Digital, Noida, India provided leadership to

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Acknowledgements

the typesetting and layout. Therefore, this project was indeed one of “global” proportions, with people representing six countries and four continents working together to make it a reality. Finally, special thanks go to my wife and life partner, Pat Ciesla, who spent many long days with me during field assignments in Brazil, China, Chile,

Cyprus, Kenya, Indonesia, the USA and many other places and provided encouragement and support as I researched and wrote this text. Several of her photos are also included. William M. Ciesla Fort Collins, Colorado, USA

The author William M. Ciesla earned a Bachelor of Science Degree in Forestry from the State University of New York, College of Environmental Science and Forestry at Syracuse University in 1960. He later specialized in forest entomology at the same institution and earned a Master of Science Degree in 1963. He began his professional carrier in Asheville, North Carolina with USDA Forest Service and subsequently held assignments as a forest entomologist and forest health program manager in several locations throughout the USA. From 1990 to 1995 he served as Forest Protection

Officer for the Food and Agriculture Organization of the United Nations in Rome, Italy and in 1995, chartered Forest Health Management International, an international forest health consulting service based in Fort Collins, Colorado. He has worked with forest insects in over 30 countries and is author or co-author of over 160 publications. He is also author of Stranieri: An Italian Odyssey, a light hearted account of life and travels in Italy. In 2005 he received the Western Forest Insect Work Conference - Founder’s Award for outstanding contributions to forest entomology in the West.

Preface Insects are by far the most dominant group of animals on the Earth’s surface. Over a million species have been described and new species are continuously being discovered. They are an integral part of the biodiversity of our planet and presently comprise about 80–85% of the total number of animal species known to science. Some experts estimate that the total number ultimately recognized may eventually approach 30 million species. More than 1000 species can occur in a relatively simple ecosystem such as a back yard and can number many millions per hectare of land surface. In addition to being abundant, insects are extremely diverse and adaptable organisms. They are found in virtually every ecosystem. Insects are classified into about 30 orders. Each order is further subdivided and classified into families, genera and species. In North America alone, some 698 families, with an estimated 95,553 species are recognized (Erwin 1982, Chapman 2006, Triplehorn & Johnson 2005). Insects are an integral part of all of the world’s forest ecosystems. Many species serve beneficial and even critical functions in forests. Some visit flowers and pollinate plants. Others function as agents in the breakdown of dead vegetation. Insects, such honeybees or the lac insect, provide products beneficial to humans. Others kill old, mature trees and make way for the establishment of young, more vigorous forests. When insects become excessively abundant, they can damage trees and forests and thus compete with humans for the goods and services that forests provide. Some groups of insects, such as foliage feeders and bark beetles, are

considered major forest pests and their prevention and control is an integral part of forest management. Worldwide, approximately 68 million ha of forests suffer some type of insect damage each year. This is more than twice the area burned by wildfires (FAO 2005). This text examines forest insects in a global context and addresses the species of major concern in the world’s forest ecosystems. It is divided into two parts: Chapters 1–5 examine the role of insects in forests, their dynamics and their effects on natural forests, plantations, agroforestry systems, urban forests, wood and non-wood products. Approaches to forest insect monitoring are reviewed and alternatives for management of damaging forest insects within the framework of integrated pest management (IPM) are presented. The basis for classification of forest insects into orders and families is reviewed in Chapter 6. Chapters 7–15 address the main thrust of this text and provide descriptions of important forest insects, their distribution, hosts, life histories and economic, social and ecological impacts. These chapters are organized according to the damage the insects cause, rather than by taxonomic groups. This text is written for foresters, forest entomologists and forest health specialists engaged in management and protection of forests, worldwide. It is intended to call attention to the importance of insects in the world’s forest ecosystems, the need to consider management of insect pests as an integral part of sustainable forest management and to serve as a guide to the identification of the world’s major pest species.

Chapter 1

The World’s Forests and their Dynamics

INTRODUCTION Forests cover 3.952 billion ha or 30.3% of the Earth’s surface. Other wooded lands cover another 1.3 billion ha.1 They provide habitat for many living organisms. Moreover, they provide a wealth of goods and services for humans. Forests are a source of both wood and nonwood products, including lumber, pulpwood, fuel wood, resin and food items such as nuts, fruits, mushrooms, edible plants and game. In addition, they provide protective cover for watersheds, range for domestic animals and are an important source of recreation and spiritual refreshment. A more recently appreciated value of forests is their ability to remove excess carbon from the Earth’s atmosphere, much of which is produced by humans through burning of fossil fuels or clearing of forests, and store it in woody biomass. The 1

Forests are defined as land spanning 0.5 ha with trees higher than 5 m and a canopy of more than 10%. Other wooded land is defined as land spanning more than 0.5 ha with trees higher than 5 m and a canopy cover of between 5% and 10% (FAO 2005, 2009a).

world’s forests presently store an estimated 240 gigatonnes (Gt) of carbon in woody biomass and a total of 683 Gt of carbon in forest ecosystems as a whole (FAO 2005, 2009a). The world’s forests have been subject to human pressure since the beginning of civilization. Large areas have been deforested to make room for agriculture, communities, industrial sites, roads and highways. Additional areas have been degraded as trees of the most desirable species and quality were harvested, forests were overgrazed by domestic livestock or burned for land clearing or to drive game. Presently, the rate of forest loss due to land use change is estimated at 13 million ha/year, resulting in a net reduction of forest area of about 7.3 million ha/year. Only 36% of the world’s forests are regarded as “primary forests.” These are defined as forests of native species in which there is no clearly visible indication of human activity and ecological processes are not significantly disturbed (FAO 2005, 2009a). Forests have the capacity to regenerate and produce goods and services for humans on a continuing basis, provided they are managed in a sustainable manner.

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Forest entomology: a global perspective

The concept of sustainability evolved as a result of the United Nations Brundtland Commission (United Nations 1987), which defined sustainable development as “development that meets the needs of the present without compromising the ability of future generations to meet their own needs.” Sustainable development focuses attention on finding strategies to promote economic and social development in ways that avoid environmental degradation, overexploitation or pollution. Forests managed under the concept of sustainable development, therefore, should provide needed goods and services for present as well as future generations. The world’s forests are dynamic. They are in a continuous state of flux and, in addition to human activities, are subject to disturbance by wind, water, fire, insects and disease. This chapter provides an overview of the world’s forests and the factors that characterize their dynamics.

FOREST ECOSYSTEMS The world’s forests are highly varied and complex and many classification systems have been used to categorize them. A system proposed by Olson et al. (2001) subdivides the Earth’s vegetation into eight biogeographic regions (Fig. 1.1) and 14 biomes. The biomes are further subdivided into 867 ecoregions. Eight of the

14 biomes proposed by Olson et al. (2001) are forest biomes and include: .

Tropical and subtropical forests: tropical and subtropical moist broadleaf forests; * tropical and subtropical dry broadleaf forests; * tropical and subtropical coniferous forests; * mangroves. Temperate forests: * temperate broadleaf and mixed forests; * temperate conifer forests; * Mediterranean forests, woodlands and scrub. Boreal forests/taiga. *

.

.

Tropical and Subtropical Forests The portion of the Earth that lies roughly between 23.5 north and south latitude, or between the Tropics of Cancer and Capricorn, is regarded as the tropics. The tropics are characterized by having consistently warm temperatures and are frost free. Annual and monthly mean temperatures are above 18–20 C and there is a difference of no more than 5 C between the warmest and coolest month of the year. These temperatures allow for biological activity to take place throughout the year, except in areas with seasonal droughts. The subtropics, on the other hand, are two bands around the earth adjacent to the tropics, from about 10 north and south latitude to 23.5 north and south

Fig. 1.1 Biogeographic regions of the world (redrawn from Olson et al. 2001).

The world’s forests and their dynamics

3

Fig. 1.2 Moist tropical forests are richest in species diversity (Parque Nacional Foz do Igua¸cu, Brazil).

latitude. While the climate is generally warm, subtropical regions are subject to occasional frosts and plant communities are more tolerant of cold temperatures than those found in the tropics. Species composition in tropical forests varies according to moisture, soil types and geological history. The richest species diversity is in Latin America, followed by Southeast Asia and Africa (Fig. 1.2). There is little similarity in species between these regions, although they do share some common plant families and genera. The tropical forests of Southeast Asia are dominated by members of the plant family Dipterocarpaceae. These are broadleaf trees that include many valuable timber species (Nair 2007). Tropical and Subtropical Moist Broadleaf Forests Most tropical and subtropical moist broadleaf forests have no discernable dry season. They are composed of broadleaf evergreen trees and are rich in terms of both plant and animal diversity. The largest area of moist tropical forest occurs in South America, in the Amazon Basin. Other areas of moist tropical forest occur in portions of Africa and Southeast Asia. Tropical and subtropical moist broadleaf forests are characterized by having dense, luxuriant, multistoried plant growth. Woody lianas or vines are common. Monocots, such as bamboos or canes, are also common in some areas.

Tree branches often provide habitat for luxuriant growth of ferns, orchids, bromeliads, mosses and lichens. Many of the world’s moist tropical forests occur at elevations of DþE If conditions are unfavorable, death and emigration rates exceed the rates of birth and immigration and populations decline: BþI 13 cm) trees with thick bark, provide a virtually unlimited food supply for the development of mountain pine beetle, Dendroctonus ponderosae, outbreaks. Other factors that can alter host quality include stocking levels (number of trees/ha), available soil nutrients, soil moisture or presence of mechanical injury. These are independent of population numbers or density. Changes in the quality of suitable host material or host depletion by insect outbreaks, on the other hand, are a function of population numbers and densities and are, therefore, density dependent factors.

Climate Climate is an important factor that regulates insect numbers and can have both favorable and detrimental effects. Periods of warm, dry weather can be favorable for development and survival of many foliage feeding insects. As temperatures cool or moisture levels increase, insects may cease feeding or become more susceptible to infection by fungi or virus (Fig. 2.4). Late spring frosts can kill large numbers of early instar larvae that have just hatched or emerged from overwintering sites. Moreover, frosts can damage food sources such as foliage. Cold winter temperatures can kill overwintering stages of insects and keep their numbers at low levels. Low winter temperatures, for example, cause extensive mortality of broods of mountain pine beetle, Dendroctonus ponderosae, overwintering under the bark of infested trees. Early autumn or mid-spring temperatures of about 18 C and winter temperatures below 37 C can cause sufficiently high levels of overwintering mortality to influence the course of outbreaks. In areas where low temperatures occur more or less regularly, such as near the northern limits of its natural range or at high elevations, the probability of outbreaks has been low (Amman et al. 1990).

Climate–Host Interactions Climatic events such as droughts, excess precipitation or severe storms can have a profound effect on trees. Drought stresses trees and lowers their resistance to bark beetle and wood borer attacks. Bark beetle outbreaks, for example, often follow extended droughts. When levels of precipitation increase, host resistance

Forest insect dynamics

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Fig. 2.4 Adult of the bark beetle, Ips typographus, killed by the fungus Beauvaria bassiana. High moisture levels favor fungi and increase their ability to attack and kill insects (southern Bavaria, Germany, photo by Patricia M. Ciesla).

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Forest entomology: a global perspective

increases and populations of some species of bark beetles decline. Lightning strikes in trees due to summer storms create focal points for bark beetle attacks that could spread to neighboring trees. Severe storms that cause extensive windthrow also create suitable host material for several bark beetle species. They can build up to high population densities in the windthrow and subsequent generations invade surrounding standing trees. Interactions between Insects and other Biotic Agents The activities of one insect population may have a strong influence on host quality for an unrelated insect population. In forest entomology, the classic example is outbreaks of foliage feeding insects that weaken their hosts and increase their susceptibility to attacks by bark beetles and/or wood borers. Defoliator outbreaks in mixed oak forests, by Lymantria dispar and other species, weakens trees and increases their susceptibility to attacks by flat headed wood borers of the genus Agrilus, i.e. A. bilineatus in North America and A. pannonicus in Europe. Outbreaks of conifer defoliators, such as western spruce budworm, Choristoneura occidentalis, and Douglas-fir tussock moth, Orgyia pseudotsugata, in western North America and Siberian silk moth, Dendrolimus sibirica, in Asia are also known to set the stage for increased activity by bark beetles and wood borers. Infection by pathogenic fungi, especially root pathogens, such as Armillaria spp. and Heterobasidium annosum, also reduces host resistance and increases susceptibility to attack by bark beetles and wood borers. Interactions between insects, pathogens, climate and host condition are the cause of a group of complex diseases known as diebacks and declines, which are discussed in greater detail later in this chapter. FOREST INSECT OUTBREAKS Most forest inhabiting insects consistently remain at low levels and are of little or no concern. However, others can go through periods of extremely high numbers and alternating low periods where they are difficult to detect. Insects that undergo periods of high population levels and become damaging are those of greatest concern, especially in natural forests. Several classes of outbreak cycles have been described (Berryman 1986). The mechanisms that favor development of insect

outbreaks are often complex and involve interactions of several density dependent and density independent factors. These relationships are often difficult to identify and there is often considerable disagreement among investigators as to their causes.

Sustained Gradient Outbreaks Some environments have conditions that are consistently favorable for development of high numbers of insects. While population densities may vary somewhat from year to year, these insects are almost always present and, in some cases, damaging year in and year out. An example of a sustained gradient outbreak is that of the larch gall midge, Dasineura rozhkovi (¼ D. laricis) in southern Siberia. This insect infests buds of various species of larch, Larix spp., where it produces an artichoke-shaped gall. The infested buds are killed and affected trees do not produce flowers, cones or seeds during the following year. Infestation levels capable of damaging up to 90% of the buds have been reported. Affected trees are not killed and damaged branches produce new shoots the following year. These provide an ample food source for the next generation of gall midges. This insect has become an important pest in larch seed orchards in the region. Infestation levels are consistently high from year to year and chemical control is required in larch seed orchards to ensure flower production (Isaev et al. 1988). Another insect capable of sustained gradient outbreaks is the western balsam bark beetle, Dryocoetes confusus, found in high elevation forests of subalpine fir, Abies lasiocarpa, in western North America. Some level of activity by this insect can be detected almost every year. A high incidence of root disease, caused by several fungi, stresses these trees and provides a continuous supply of suitable host material for invasion by the beetles. Many insects that are pests of forest plantations tend to remain at high levels because of the virtually unlimited host material provided by large areas of single species, even-aged stands characteristic of most plantations. Examples include white pine weevil, Pissodes strobi, in North America, several species of shoot and tips moths, Rhyacionia spp., in conifer plantations and the mahogany shoot borers, Hysipyla grandella and H. robusta, in mahogany plantations in tropical regions of Africa, the Americas and southeastern Asia.

Forest insect dynamics Cyclic Outbreaks Periodic and, occasionally, predictable oscillations in the abundance of animals, known as population cycles, are one of the most fascinating characteristic of population dynamics. These cycles are not present in all species but occur in a number of forest insects. During peaks, populations may reach extremely high numbers and during low periods the same species may be difficult to locate. In high-elevation forests of European larch, Larix decidua, in the Alps, the larch bud moth, Zeiraphera diniana, has exhibited a high degree of variation in numbers on a more or less regular cycle. Population densities are known to vary from 1 to > 30,000 larvae/ tree. During years of high population density, larvae may cause complete defoliation. Historical records of defoliation indicate that outbreaks have occurred in high-elevation larch forests with a high degree of regularity every 8–9 years. These cycles do not occur in lower elevation mixed species forests where larch is a component. Analysis of the tree ring history of larch forests in the Alps indicates that outbreaks have occurred on an average of once every 9.3 years over the past 1173 years but have not occurred since 1981 (Baltensweiler & Fischlin 1988, Esper et al. 2007). Numbers of the North American Douglas-fir tussock moth, Orgyia pseudotsugata, can fluctuate greatly between generations. A study by Watt (1968) indicates that this species had the most extreme population fluctuations of any of several hundred species of Canadian forest Lepidoptera. There is some evidence that tussock moth populations may possess characteristics of both cyclic and eruptive outbreaks (see following section). Historical records in the Blue Mountains of eastern Oregon, USA indicate that outbreaks occurred at about 9-year intervals (1928–9, 1937–9, 1946–8, 1963–5, 1972–4, 1992–3 and 2000–1) (Brookes et al. 1978, Powell 2008).

Eruptive Outbreaks Some forest insect populations remain at low population levels for many years, then suddenly explode to extremely high levels, often with little or no warning, over large areas of forest and cause extensive damage. These are referred to as population eruptions or eruptive outbreaks. Many examples of eruptive outbreaks of bark beetles and defoliating insects have

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been documented from the forests of Asia, Europe and North America. The southern pine beetle, Dendroctonus frontalis, undergoes eruptive and highly destructive outbreaks throughout its natural range. While this insect is usually present at epidemic levels somewhere within its range, there are periods at any given location where its levels are extremely low. In Central America, devastating outbreaks occurred in Honduras during the mid1960s and again during the early 1980s. A regional outbreak, which encompassed many pine forests in Central America, occurred from 2000 to 2003. During years between outbreaks, the insect was difficult to detect (Billings et al. 2004). The pine forests of east Texas, USA also have a history of devastating outbreaks of D. frontalis. However, the insect has not been found in east Texas since the early 1990s. A startling and fascinating aspect of eruptive outbreaks is the synchronous buildup of insect populations over large areas within a relatively short time. One scenario or hypothesis is that populations build to high levels in localized areas known as epicenters, where there is an abundance of suitable feeding and breeding sites. Subsequent generations disperse to surrounding areas where they too become damaging. A second scenario is the synchronous buildup of populations over large areas. This was first observed in populations of the Canadian lynx by Australian statistician P.A.P. Moran (Moran 1953). Moran’s hypothesis was that if two or more populations of the same species are influenced by the same density dependent factors, then correlated density independent factors, usually weather induced, would bring the population’s fluctuations into synchrony. This phenomenon is known as the “Moran Effect” (Hudson 1999, Ripa 2000). Spruce budworm, Choristoneura fumiferana, a major defoliator of boreal forests across North America, may remain at low levels for long periods ranging from 30 to 100 years. When outbreaks develop, they typically cover large areas, sometimes millions of hectares. The outbreaks of the 1910s, 1940s and 1970s covered 10, 25 and 55 million ha, respectively. Outbreaks are often associated with heavy crops of male flowers in stands of mature balsam fir, Abies balsamea, the insect’s favorite host. During outbreaks, adults disperse and invade regions as distant as 600 km from their points of origin. Tree ring analysis in eastern Canada, where this insect has been a historical pest, suggests occurrence of outbreaks over the past 200–300 years and indicates that outbreaks occurred more frequently

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Forest entomology: a global perspective

during the 20th century than previously. Reasons for the increase in frequency, extent and severity may be the result of human activities, including timber harvesting, fire protection and budworm spraying, that favor development of spruce–fir forests over other species in the region and make the forest more prone to budworm attack (Blais 1985). Buildup of spruce budworm populations, initially in relatively small areas, coupled with the mass migrations of adults, suggests that dispersal of large numbers of insects into uninfested areas is a key factor in the dynamics of this insect. However, an analysis of the dynamics of spruce budworm populations in New Brunswick, Canada disputes this. This analysis indicates that local populations tend to oscillate in unison. Local population processes are composed of two parts, a basic oscillation and secondary fluctuations about the basic oscillation. The basic oscillation is the result of several density dependent mortality factors that affect larval instars III–VI and cause increases in budworm numbers. Secondary oscillations are the result of local population buildups supplemented by an influx of female moths dispersing from other areas. These are highly correlated with local meteorological conditions. This study concluded that moth invasions can only accelerate an increase in local populations to an outbreak level when the local population is already in an upswing phase. Therefore, localized increases in spruce budworm numbers brought about by density dependent factors are brought into synchrony by density independent factors such as weather, the so called Moran Effect (Royama 1984). Other analyses of both cyclic and eruptive forest insect outbreaks confirm the role of the Moran Effect in their dynamics. For example, analyses of outbreak patterns associated with six damaging forest insects indigenous to Europe and North America: spruce budworm, Choristoneura fumiferana, forest tent caterpillar, Malacosoma disstria, larch bud moth, Zeiraphera diniana, mountain pine beetle, Dendroctonus ponderosae, and gypsy moth, Lymantria dispar, indicate a spatial synchrony consistent with the Moran Effect (Peltonen et al. 2002).

Declines and Diebacks Declines and diebacks of trees and forests are complex conditions caused by interaction of several biotic and abiotic agents, including insects. While these may not

be considered as outbreaks in the classic sense, many declines are episodic and insects often play a significant role in their occurrence. Causes of declines are difficult to diagnose. In many cases, a scenario or “working hypothesis” is the only realistic diagnosis that can be made. Symptoms associated with these conditions are often similar, regardless of the cause(s) and include growth loss, thin crowns, yellow (chlorotic) foliage, dieback and tree death. Forest decline episodes have been reported worldwide (Ciesla & Donaubauer 1994). The interaction of several biotic and abiotic agents as causes of decline is described by Manion (1991) as an array of predisposing, inciting and contributing factors. Predisposing factors are present in a forest for long periods, cause stress but at a level too low to produce visible symptoms. These include occurrence of large areas of mature forests, which have reduced growth rates and increased susceptibility to diseases such as root pathogens, or exposure to low levels of air pollutants. Inciting factors are short-term events that weaken the already stressed trees and cause growth loss, crown thinning and/or dieback. Examples include defoliator outbreaks, late spring frosts, prolonged drought or excess precipitation. Contributing factors are those that cause death of declining trees and are usually secondary bark beetles and/or wood boring insects that are unable to attack healthy, vigorous trees. Outbreaks of forest defoliators have been implicated as inciting factors in a number of forest decline events. In parts of Europe and eastern North America, for example, a condition known as oak decline has caused extensive forest damage. Outbreaks of several defoliating insects, including Lymantria dispar, Malacosoma disstria and Tortrix viridana, have functioned as inciting factors. Moreover, two species of wood borers, Agrilus bilineatus in North America and A. pannonicus in Europe, have been implicated as contributing factors to oak decline. Several scenarios that involve an interaction of predisposing, inciting and contributing factors have been formulated to explain oak decline on both sides of the Atlantic Ocean (Table 2.2). While Manion’s (1991) scenario considers declines as a disease, Mueller-Dumbois (1983, 1986, 1982) considers them an integral part of forest dynamics and succession. He developed this hypothesis while working with a decline of ohia, Metrosideros polymorpha, in the Hawaiian Islands, USA. While he concurs that declines are a complex of interacting factors, he argues that the primary predisposing cause of decline is the occurrence of even-aged forests (cohorts) growing on marginal soils

Forest insect dynamics

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Table 2.2 Predisposing, inciting and contributing factors to oak decline in Europe and North America. Factor

Europe

North America (Appalachian Mountains, USA)

North America (Ozark Mountains, USA)

Predisposing factors

Root pathogens, Armillaria spp.

Root pathogens, Armillaria spp.

Fire exclusion

Inciting factors

Extreme cold Drought Insect defoliation

Drought Insect defoliation

Drought

Contributing factors

Wood borer (Agrilus pannonicus) Fungi

Wood borer (Agrilus bilineatus)

Wood borers (Enaphalodes rufulus)

Sources: Haack & Acciavatti 1992, Hartman and Blank, 1992, Gibbs & Grieg 1997, Spencer & Sutton 2001.

that have reached a stage of senescence, a condition he refers to as synchronous cohort senescence. When exposed to an inciting factor, such as drought, excess rainfall or defoliation, they decline. While some trees may recover, others die as a result of attack by secondary insects. A decline of quaking aspen, Populus tremuloides, in western North America seems to fit the scenario proposed by Mueller-Dumbois. This condition, referred to as “sudden aspen decline” (SAD), involved rapid death of aspen climax stands growing at the lower elevation limits of tree growth. These are pure, even-aged clonal stands regenerated from sprouts and therefore have limited genetic diversity. P. tremuloides reaches maturity at about 60 years and many stands in the affected area were in excess of 100 years. During the late 1990s and into the first years of the 21st century, the region was subject to a prolonged drought, which may have incited decline. Several species of secondary wood borers and bark beetles were associated with dying aspens. Most affected stands had a dense ground cover of aspen regeneration, which released as older trees died. Aspen climax stands that lack understory regeneration often have been overgrazed by either wild or domestic ungulates (Author’s observations, Worrall et al. 2008). A common error when diagnosing causes of forest declines and diebacks is that the contributing factor, which is often an insect, is the agent to which the cause is attributed. This is because evidence of secondary insects, such as bark beetles and/or woodborers, is most obvious and is often present for long periods. A classic example is the case of a decline of Acacia nilotica, an important riparian species in the Blue Nile River Basin

of the Sudan. This species grows in pure, even-aged stands in old oxbow lake beds along the Blue Nile and is often regenerated by direct seeding. Initially, decline and mortality of A. nilotica was attributed to a stem infesting wood borer, Sphenoptera chalcicroa, which was present on affected trees in moderate numbers. A later evaluation suggested that other stem boring insects also occurred in the affected trees and were most likely contributing factors of a complex condition. It was hypothesized that the decline may have been due to senescence of even-age forests coupled with almost annual inundations of water and silt in the oxbow lakes from annual flooding of the Blue Nile, which, over time, fill the lakes with silt and render them too dry to support tree growth (Ciesla 1993). Insect outbreaks are the result of many of the same factors that cause decline. A forest may be predisposed to a bark beetle outbreak because of its age and structure. Occurrence of drought, which adds an additional stress to the forest, or mild winter temperatures that increase survival if overwintering insects may serve as the inciting factors. The key difference is that the insect outbreak may be able to infest and kill large numbers of trees even after climatic conditions become more favorable for growth and survival of the host trees simply because of the massive numbers of insects present.

CLIMATE CHANGE AND FOREST INSECT DYNAMICS Global climate change, due to increased levels of carbon dioxide (CO2) and other greenhouse gases in

28

Forest entomology: a global perspective

the Earth’s atmosphere, has emerged as the primary environmental issue of the 21st century. CO2 accounts for less than 1% of the Earth’s atmosphere, yet its presence is critical because it causes thermal radiation from the sun to be reflected back to the Earth’s surface, thus increasing the Earth’s temperature. As the level of CO2 and other greenhouse gases (e.g. methane, carbon monoxide, nitrous oxides) increases, there is more reflection of thermal radiation and temperatures increase. Conversely, if levels of these gases decrease, the Earth’s temperature cools. Without the presence of greenhouse gases, the Earth’s temperature would be about 33 C cooler than it is today and too cold to support life as we know it. Levels of atmospheric CO2 and other greenhouse gases have been increasing gradually since the mid19th century. Many scientists have attributed this to increasing human activities such as burning of fossil fuels (coal, natural gas, oil) for energy, industry and transportation, and release of carbon sequestered in trees due to deforestation and land use change. Concurrently, global surface temperatures have increased by about 0.74 C over the past century. Eleven of the 12 years from 1995 to 2006 ranked among the 12 hottest years on record since 1850, when worldwide temperature measurements began. Consistent with rising global temperatures, episodes of cold temperatures and frosts have declined over the past 50 years. Mountain glaciers have shrunk in size, changes in precipitation patterns have been recorded and there is evidence of intensified tropical cyclone activity since about 1970 (IPCC 2007). Forests as well as insects and other organisms that inhabit forests can respond to a changing climate. Natural ranges of trees and forest ecosystems could shift to higher latitudes or higher elevations as temperatures rise. Insects, with their short life cycles and ability to disperse, could react to a changing climate much more rapidly than their host plants. In an earlier part of this chapter, the effect of short-term climatic fluctuations such as drought, excess precipitation or severe storms on insect population dynamics has already been discussed. The following sections address how insects might be affected by long-term climate change and provide some examples of insects that may have already responded to a changing climate. A comprehensive review of potential impacts of climate change on forest health is provided by Moore and Allard (2008).

Shifts in Natural Ranges and New Host Associations Long-term warming of the Earth’s temperature could cause insects to shift ranges north in the northern hemisphere, south in the southern hemisphere or upwards in elevation. As ranges expand, insects could come in contact with new hosts. Several examples of changes in natural ranges of forest insects, some with new host associations have been reported. In Europe, two species of processionary caterpillars, oak processionary, Thaumetopoea processionea, and pine processionary, Thaumetopoea pityocampa, have been found north of their known ranges. The “traditional” range of the oak processionary is central and southern Europe and Israel. Populations have been detected further north and this insect is now known to occur in Belgium, northern France, the Netherlands, Sweden and the UK (Evans 2007, FAO 2009b). Pine processionary caterpillar occurs in pine forests of the Mediterranean Basin where larvae feed during winter. Studies in France indicate that they can survive in areas that receive 1800 hours of sun/year with an average minimum January temperature of above 4 C (Huchon & Demolin, 1971). There are indications of a northward and upward altitudinal expansion in the range of this species, both in France and Italy (Battisti et al. 2005). During the summer of 2003, the warmest summer in Europe during the past 500 years, this insect was able to expand its range into high-elevation pine forests in the Italian Alps (Battisti et al. 2006). Some bark and ambrosia beetles and their associated fungi are also expanding their ranges. In western North America, walnut twig beetle, Pityophthorus juglandis Blackman, has been indigenous to portions of Arizona, California and New Mexico, USA, and northern Mexico, where it infests stems of several species of walnuts, Juglans spp., and causes occasional twig dieback. Beginning in 2002, infestations were discovered further north, in portions of Colorado, Idaho, Oregon, Washington and Utah, USA, in a new host, black walnut, Juglans nigra. This bark beetle has developed an association with a fungus, Geosmithia sp., which is highly pathogenic to black walnut and causes “thousand cankers” disease. This fungus, vectored by P. juglandis, has killed hundreds of black walnut trees and a potential exists for the beetle and the fungus to spread east into the natural range of black walnut

Forest insect dynamics where its lumber is highly prized for furniture, gunstocks and other wood products (Wood 1982, NAPPO 2008, Tisserat et al. 2009). The ambrosia beetle, Platypus quercivorus, occurs in temperate, subtropical and tropical forest ecosystems across Asia. During the early 1980s, mortality of Quercus mongolica forests was reported from western Japan. This condition was attributed to a fungus, Raffaelea quercivora, which is the fungal symbiont of P. quercivorus. Oaks resistant or tolerant to R. quercivora may have co-evolved under a stable relationship between the tree, fungus and beetle during a long evolutionary process. Q. mongolica may not have been part of this coevolution. The present epidemic may be the result of a warming trend that began in the late 1980s, which allowed the insect to extend its range to more northerly latitudes and higher elevations where Q. mongolica occurs (Kamata et al. 2002, Kubono & Ito 2002). Factors other than climate change that can cause insects to expand their natural ranges include accidental introductions via human trade and travel or climatic anomalies such as high winds that can carry insects over long distances. Provided they find suitable hosts the insects can become established in their new locations.

Additional Generations Ability of insects to undergo additional generations and, therefore, become more damaging, is another potential consequence of a warming climate. This could be especially true for species that already undergo several generations/year. Depending on latitude and elevation, southern pine beetle, Dendroctonus frontalis, may undergo from three to seven generations/year (Thatcher et al. 1980). A warmer climate could cause this species to undergo additional generations, especially in the more northerly portions of its range. The North American spruce beetle, D. rufipennis (Kirby), typically requires 2 years to complete a generation but at lower, warmer elevations it can complete a generation in a year (Holsten et al. 1999). Warmer climates could, therefore, favor a 1-year life cycle, thus increasing the destructive potential of an already damaging species. Some forest defoliators also have the potential to have additional generations in warmer climates. For example, an Asian pine caterpillar, Dendrolimus punctatus Walker, currently has two and a partial third

29

generation/year in east central China and four generations/year in Vietnam (Dao Xuan Truong 1990). Warmer temperatures could potentially increase the number of generations in a given location.

Outbreaks in New Locations Low winter temperatures are known to kill overwintering populations of insects and reduce the probability of outbreaks. This was the case with mountain pine beetle, Dendroctonus ponderosae, near the northern limits of its range or at high elevations. Moreover, in northerly latitudes and high elevations, this insect requires 2 years to complete a generation as opposed to one generation/ year across most of its range (Amman et al. 1990). In 1994, an outbreak of mountain pine beetle developed in the central-interior region of British Columbia, Canada. The outbreak expanded dramatically and by 2007, covered over 10 million ha. In addition, since 2006, this insect extended its range into the Peace River area of northern Alberta. Several factors are believed to be responsible for this outbreak. The predominant forest cover is even-age stands of Pinus contorta, a favorite host of mountain pine beetle. Many of these stands are older than 60–80 years and contain large-diameter, thick-barked trees. In 1910, an estimated 2.5 million ha of British Columbia’s 14.9 million ha of lodgepole pine forests were classified as mature. By 1990, an estimated 8 million ha were classified as mature, a threefold increase. Climate change may have also contributed to the unprecedented extent and severity of this outbreak. A combination of hot dry summers and mild winters caused reduced overwintering brood mortality and increased the potential for a higher portion of insects to complete a generation in 1 year. This increased the insect’s damage potential and allowed it to reach epidemic levels at higher elevations and more northerly latitudes. Climatic data indicate that average minimum temperatures in the affected areas have increased by 2.2–2.6oC over the past 100 years. There is increasing concern in Canada that this insect could spread into areas where the ranges of Pinus contorta and P. banksiana, a component of Canada’s transcontinental boreal forest, overlap. This could provide mountain pine beetle an opportunity to attack a new host and expand its range east across Canada (Logan & Powell 2001, British Columbia Ministry of Forests, Lands and Mines 2003).

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Forest entomology: a global perspective

In portions of Colorado and Wyoming, USA, another mountain pine beetle outbreak of unprecedented proportions began during the late 1990s. In Colorado, most lodgepole pine forests occur at elevations at or above 2500 m. Milder temperatures at these elevations, coupled with a high proportion of mature forests susceptible to beetle attack, are believed to have incited this outbreak. A mountain pine beetle phenology model predicts the observed northward and upward elevation shift of populations that could undergo one generation/year. This model also indicates that even under a conservative climate change scenario, the area where mountain pine beetle could complete one generation/year will shift considerably north, to the point where it may be maladapted over much of its traditional range. However, a cold tolerance model suggests that winter survival of mountain pine beetle will continue to remain low throughout Canada’s boreal forests and thus limit its eastward spread (Logan & Powell 2001, Regniere and Bentz, 2009).

Outbreaks of Insects Previously Considered Non-Damaging A changing climate could create conditions favorable for occurrence of outbreaks of little known or previously considered innocuous insects. The geometrid, Nepytia janetae, was first described in 1967 and considered a curiosity (Rindge 1967). From 1996 to 1999, an outbreak of this defoliator erupted on 4000 ha of high-elevation ( > 3000 m) forests of Engelmann spruce, Picea engelmannii, and corkbark fir, Abies lasiocarpa var. arizonica, in the White and Pinaleño Mountains of Arizona, USA. Defoliation, in combination with secondary attacks of several species of bark beetles, resulted in extensive tree mortality, especially in the Pinaleño Mountains. This insect has an unusual life cycle for a high-elevation species. Adults appear in late June and July and deposit eggs, which hatch in late September. Larvae feed throughout the winter and following spring. Milder winter temperatures could favor this insect by increasing larval survival rates. Another outbreak occurred in portions of the Sacramento Mountains in New Mexico between 2006 and 1

Personal communication, Ann M. Lynch, USDA Forest Service, Rocky Mountain Research Station, Tucson, Arizona, USA.

2008 (Fairweather et al. 2006, USDA Forest Service 2007, 2009).1 In the Southern Alps, an outbreak of a web spinning sawfly, Cephalica arvensis, between 1985 and 1992 is believed to be the result of warming temperatures. Sawflies of the genus Cephalica generally have a low fecundity and undergo an extended diapause of several years stimulated by low temperatures at the time of pupation. The outbreak corresponded to a period of high temperatures, low precipitation and severe water stress resulting in reduced mortality, faster development and higher rates of feeding (Marchisio et al. 1994, Battisti 2004). Changes in Population Cycles There is evidence that climate change can alter the cyclic nature of some forest insects. The cyclic nature of larch bud moth, Zeiraphera diniana, outbreaks in the Alps was described earlier in this chapter. These cycles apparently continued despite climatic changes such as the Medieval Optimum, a period of warm temperatures, and the Little Ice Age, a period of cooler temperatures, across Europe. Beginning in 1981, these cycles ended. Absence of outbreaks of this insect, beginning during the late 20th century, corresponds to a period of regional warming that is unprecedented over the past 1000 years (Esper et al. 2007). Forest Insect Outbreaks as Carbon Sources Outbreaks that cause extensive tree mortality reduce carbon uptake by forests and increase CO2 emissions caused by decay of dead trees, a positive feedback for climate change. According to a study by Kurz et al. (2008), between 2002 and 2020, trees killed by mountain pine beetle outbreak in British Columbia, Canada, will release some 270 megatonnes (Mt) of carbon over 374,000 km2 of forest. During the worst year, impacts of this outbreak, in terms of carbon release, will be equivalent to 75% of the average annual direct forest fire emissions from all of Canada during 1959–99. They conclude that massive insect outbreaks can undermine the ability of boreal forests to take up and store atmospheric carbon.

Chapter 3

Forest Insect and Human Interactions

INTRODUCTION

FOREST INSECTS AS PESTS

Trees and other forest vegetation provide food and breeding sites for many insect species. Virtually every part of a tree: foliage, stems, branches, bark, cambium layer, woody tissue, roots, flowers, seeds or cones, is utilized by insects. In doing so, they can compete with humans for food and fiber either because they feed on trees and cause growth loss, loss of seed crops or tree death, or because they serve as vectors of fungi, viruses or other agents that cause plant disease. How we relate to the insects that occupy forests depends on our system of values and needs for the goods and services that forests provide. Humans may view a forest insect as beneficial, damaging or simply innocuous, depending on how that insect affects the forest and, in turn, how that affects the resources we receive from the forest.

When is an insect a pest? The answer lies in the age-old question “If a tree falls in the forest and no one is around to hear it, does it make a sound?” A similar question can be asked about insects. “If trees are attacked and killed by insects and there are no humans around to make use of them, are those insect pests?” The answer is no. In the absence of humans, an insect outbreak is simply an ecological event, part of the dynamics of the forest. If, however, the trees under attack are a source of lumber, fiber, food, fuel or other product of human value, the insects in question are, at least by some members of society, considered pests. If the damage they cause is significant, steps may have to be taken to reduce those losses so that humans dependent on these resources may continue to exist in comfort.

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Forest entomology: a global perspective

Fortunately, only a small proportion of the insect species found in the world’s forests competes with humans for forest resources. However, that relatively small number can have a wide range of ecological, social or economic impacts. Their damage can be significant, sometimes accounting for millions of US$ in resource losses and pest management costs.

Tree Damage Caused by Insect Feeding or Breeding Many insects feed on foliage of trees and other woody plants. During outbreaks, they can consume most or all of the foliage, sometimes over millions of hectares. Defoliation, especially if it occurs over several successive years, causes reduced seed yields, growth loss, branch dieback and, in extreme cases, tree death. Insects that breed in the inner bark and cambium of living trees cause tree mortality and, during outbreaks, kill trees over large areas. Wood boring insects reduce structural integrity of trees they infest, thus increasing their susceptibility to windthrow. Insects that feed in wood in use can cause structural damage to poles, fences and wooden buildings. Sucking insects, such as aphids and scales, feed on plant juices and heavy infestations can kill stems and foliage. In addition, species that establish colonies and feed on the bark of trees cause drying and cracking of the bark. This creates sites where fungi can invade and cause further damage. Gall making insects can kill branches, reduce cone and seed production and make trees in urban settings unsightly. Shoot and stem infesting insects kill growing portions of trees and cause growth reduction and tree deformity. Repeated infestations of shoot and stem insects can reduce young fast-growing trees to bushy shrubs. Insects that use flowers, cones, nuts and seeds of trees as a source of food or breeding sites can reduce seed yields. Moreover, some forest trees provide edible fruits and nuts, which humans have relied on as a staple food source.

Insects as Vectors of Tree Disease Some forest inhabiting insects have developed a symbiotic relationship with fungi, viruses, nematodes or other agents that cause tree disease. Insects carry spores or

other life stages of disease agents on their bodies, which are transferred to trees during feeding. In some cases, the disease weakens or kills the tree and makes it a suitable site for breeding or feeding by the insect vector. Bark beetles and ambrosia beetles are vectors of wood staining fungi. They carry the spores of these fungi in structures on the thorax of adults known as mycangia and introduce the fungi into the tree during breeding attacks. Many species of non-aggressive bark beetles breed in recently killed trees. This is true of the elm bark beetles, Scolytus multistriatus and S. scolytus. Both species can carry spores of the fungi, Ophiostoma ulmi and O. novo ulmi, which cause Dutch elm disease. When the new generation of adults emerges they feed in branch crotches of elms before seeking breeding sites. If the beetles are carrying spores of the Dutch elm disease fungi on their bodies, they inoculate the feeding sites. After infected trees die, they serve as breeding sites for the same beetles. Ambrosia beetles breed in woody tissue of host trees and have developed a symbiotic relationship with fungi known as ambrosia fungi. Female adults construct a network of galleries and cradles in the woody tissue. During gallery construction, they inoculate the wood with fungus spores. These fungi provide food for developing larvae. They also discolor or stain wood and can reduce lumber quality. In some cases, these fungi are pathogenic, especially if the insect vector should expand its range and attack new hosts. Other wood boring insects also serve as vectors of either fungi or nematodes. Larvae of most wood boring species cannot digest cellulose, which is their major source of nutrition. They live in a symbiotic relationship with wood decay fungi that can break down cellulose. The woodwasp, Sirex noctilio, is associated with a wood decay fungus, Amylostereum areolatum, which is injected into trees by females during egg deposition. This fungus, along with mucus produced by the insect, is pathogenic to species of North American pines that have been planted in the southern hemisphere and has caused widespread tree mortality. Longhorn beetles of the genus Monochamus are found throughout conifer forests of the northern hemisphere where they breed and bore in the inner bark, cambium and wood of host trees. These beetles are vectors of nematodes of the genus Bursaphelenchus. Most species of Bursaphelenchus cause no damage to conifers with which they have co-evolved. However, if they are introduced into a new location, they have the potential of becoming important forest pests. The pinewood

Forest insect and human interactions

33

nematode, B. xylophilus, is native to North America and the cause of a pine wilt disease. It has been introduced into China, Japan, Korea, and, more recently, Portugal, where it has caused extensive tree mortality of indigenous pines. Monochamus larvae become infested with nematodes while feeding in nematode infested trees. The nematodes are spread by adults when they feed on twigs and branches of pines. The nematode “dauerlarvae” emerge from the spiracles of adult beetles during feeding and invade trees via the feeding wound produced by the beetles. When the trees die from the nematode infection, they become suitable breeding sites for Monochamus (Kobayashi et al. 1984, Sousa et al. 2001). Pitch canker, caused by the fungus Fusarium subglutinans f. spp. pini is a pest of pines in southeastern USA, several Caribbean islands and has been introduced into New Zealand and South Africa. This disease was discovered in California, USA in 1986 in native forests of Monterrey pine, Pinus radiata, other pines and Douglas-fir, Pseudotsuga menziesii. Studies in California indicate that several insects are vectors of this fungus including twig beetles of the genus Pityophthorus, the cone beetle, Conopthorus radiatae, a dry twig and cone beetle, Ernobius punctulatus, and three engraver beetles of the genera Ips and Pseudips (Fig. 3.1, Storer et al. 1994). Indigenous vs Introduced Insects Forest insect pests may be either indigenous or exotic. Indigenous insects are an integral part of the ecosystems they occupy and have co-evolved with their hosts and environment. While some indigenous species reach epidemic levels and have a significant effect on forest dynamics, they are subject to checks and balances imposed by natural enemies including parasitoids, predators and disease, host condition and climatic factors. On the other hand, non-native or exotic species, which are introduced from other locations and become established in a new habitat, have the potential to be exceptionally damaging. Their new hosts may lack effective defense mechanisms against these insects and/or the new habitat has few or no natural enemies to help regulate populations. Indigenous Insects Most forests insects that are considered pests are indigenous. In temperate and boreal forests, insect outbreaks are a more or less regular occurrence in natural forests.

Fig. 3.1 Pitch canker, a damaging disease of pines caused by the fungus Fusarium subglutinatus f. spp. pini, is spread from tree to tree by several insects associated with pines (Point Lobos State Reserve, California, USA).

These forests have less species diversity than do tropical and subtropical forests and, under certain conditions, can provide an unlimited supply of suitable host material. Natural forests in both Eurasia and North America are subject to widespread outbreaks of indigenous defoliating caterpillars. Defoliators such as Dendrolimus sibirica, D. spectabilis, Euproctis kargalika and Lymantria dispar asiatica have a history of outbreaks across portions of Asia. In central and northern Europe, indigenous defoliators including Dendrolimus pini, Lymantria dispar, L. monacha and Tortrix viridana cause extensive defoliation. In the Mediterranean forests of southern Europe, the Near East and northern Africa, repeated outbreaks of the processionary caterpillars, Thaumetopoea pityocampa and Th. wilkinsonii, have defoliated pine

34

Forest entomology: a global perspective

forests. Conifer forests across Canada and portions of the USA are subject to periodic outbreaks of the budworms, Choristoneura fumiferana and C. occidentalis. Other significant North American forest defoliators include forest tent caterpillar, Malacosoma disstria, in mixed broadleaf forests and Douglas-fir tussock moth, Orgyia pseudotsugata, in forests of Abies and Pseudotsuga menziesii. These outbreaks have, on occasion, been the subject of massive, and often controversial, campaigns to reduce population levels and subsequent damage via aerial and ground applications of insecticides. Outbreaks of indigenous bark beetles kill trees over large areas. These insects are especially damaging across portions of North and Central America and also Eurasia. In North America, the most damaging species are members of the genus Dendroctonus. Of the 19 recognized species in this genus, 17 are indigenous to the western hemisphere (Wood, S.L. 1982). D. frontalis, for example, is a pest of pine forests from eastern and southern USA, west to Arizona and New Mexico and south through Mexico to northern Nicaragua. Outbreaks of D. ponderosae cause extensive tree mortality in pine forests across portions of western North America. D. rufipennis causes extensive tree mortality in mature spruce forests across all of North America and D. pseudotsugae is an important pest of mature forests of its host Pseudotsuga menziesii. Several bark beetles of the genus Ips are also important forest pests and, during outbreaks, cause extensive tree mortality. Examples include I. acuminatus and I. typographus in Eurasia and I. avulsus, I. confusus, I. pini and others in North America. Other damaging bark beetles include Orthotomicus erosus and Tomicus destruens in the Mediterranean region of Europe, North Africa and the Near East and Scolytus ventralis in western North America. Introduced Insects Insects are opportunistic organisms and will use any method available to disperse and seek new hosts. Humans are effective vectors for long-distance spread of insects, either intentionally or inadvertently, and have been responsible for introduction of many species that have had severe ecological, social and economic consequences when established in new locations. In the USA alone, estimated environmental and economic costs associated with all alien invasive forest species, including insects, is estimated at US$ 2.1 billion annually (Pimentel et al. 2005).

Globally, humans provide opportunities for introduction and establishment of damaging forest insects in two ways. First, international trade and travel or within-country movement of plant products provides pathways for the movement of insects. Second, establishment of plantations of trees exotic to an area provides additional hosts that increase their chances of survival once they arrive at a new location. Some key pathways for movement of forest and tree pests include: .

live plant materials: nursery stock house plants scion material for grafting bonsai; unprocessed logs (Fig. 3.2); lumber, especially lumber containing bark strips; dunnage – wood used to stabilize containers in transit; fuel wood (Fig 3.3).

    . . . .

Increased human travel and international trade is causing an alarming increase in rates of insect introduction and establishment worldwide. Wood borers and bark beetles feature prominently among invasive forest insects because they are easily transported inside wood products, wood crating and dunnage where they are concealed and protected. According to records available from the United States Department of Agriculture (USDA) Animal Plant Health Inspection Service (APHIS) between 1985 and 2000, bark and ambrosia beetles were intercepted at ports of entry in the USA 6825 times from 117 countries. Overall, 73% of these interceptions were from wooden packing materials (Haack 2003). During the same period, 25 species of wood borers, bark and ambrosia beetles became established and several have caused severe damage. A similar study in New Zealand indicates that between 1950 and 2000, 1500 interceptions of bark and ambrosia beetles, representing 103 species were made (Brockerhoff et al. 2006). A summary of key forest insect introductions, which have affected natural forests worldwide, is given in Table 3.1.

Insect Pests of Forests of Human Origin Forest Plantations Forest plantations occupy about 187 million ha worldwide, or about 5% of the world’s total forest area.

Forest insect and human interactions

35

Fig. 3.2 A variety of wood products, including unprocessed logs, a potential pathway for spread of bark beetles and wood boring insects, await shipment to foreign destinations (Concepción, Chile).

Forests and trees are planted for many purposes. Industrial forest plantations, established for wood and fiber production, account for 48% of the total plantation area. Protective forest plantations, established for conservation of soil and water, account for 26% and the

objectives of the remaining world’s forest plantation area are unspecified. Significant areas of forest plantations are found on all of the world’s continents, except Antarctica. Asia has the largest area of forest plantations, about 116 million ha (FAO 2001).

Fig. 3.3 Transport of fuel wood over long distances poses a hazard of spread of a variety of wood infesting insects (Fort Collins, Colorado, USA).

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Forest entomology: a global perspective

Table 3.1 Examples of introduction and establishment of damaging forest insects into natural forests, worldwide. Type of damage

Species

Native range

Introduced range

Mode of introduction

Foliar damage

Lymantria dispar Gypsy moth

Eurasia

North America

Hyphantrea cunea Fall webworm Gilpinia hercyniae European spruce sawfly

North America Europe

China, eastern Europe, Japan, Korea, Russia North America

Purposely introduced as a potential source of silk Unknown

Bark and ambrosia beetles

Dendroctonus valens Red turpentine beetle Tomicus piniperda

North America

China

Eurasia

North America

Wood products, crating

Wood borers

Agrilus planipennis Emerald ash borer Anopolphora glabripennis Asian longhorn beetle Tetropium fuscum

Asia

North America

China, Japan

North America, Europe

Wood products and crating Wood products and crating

Europe

Eastern Canada

Wood products and crating

Adelges piceae Balsam woolly adlegid Adelges tsugae Hemlock woolly adlegid Pineus boerneri Pine woolly aphid

Europe

North America

Nursery stock

Asia

North America

Eurasia?

Cryptococcus fagisuga

Europe

Africa, Australia, New Zealand, North America, South America Eastern North America

Unknown, probably live plant materials Nursery stock

Coptotermes formosana Formosan termite

China, Japan, Taiwan

Sucking insects

Wood products insects

Many of the world’s forest plantations are composed of fast-growing exotic trees. Extensive plantations of eucalypts, pines, poplars, teak and other exotic trees have been established in tropical, subtropical and temperate regions of the southern hemisphere (Fig. 3.4). Even in Europe and North America, where native species are preferred, there are some plantations of exotic species. For example in the British Isles, plantations of the North America conifers Picea sitchensis and Pinus contorta ssp. contorta have been established. In the Galicia region of Spain, plantations of the North American Pinus radiata have been established and in Portugal there are extensive plantations of Eucalyptus. Moreover, these plantations are often established with genetically improved seedlings, bred specifically for certain

North America, South Africa, Southeast Asia

Nursery stock? Unprocessed logs

Unknown Wood products and crating

desirable characteristics such as rapid growth, good form or disease resistance. Trees of the genera Eucalyptus and Pinus are the most extensively planted trees and account for 20% and 10%, respectively, of the world’s total forest plantation area (FAO 2001). Forest plantations typically, although not always, consist of extensive, even-aged stands of a single species. These can provide an unlimited food supply for insects. Insects that damage forest plantations can be classified into four categories: . . . .

indigenous insects in plantations of native trees; indigenous insects that have adapted to exotic trees; introduced insects in plantations of native trees; introduced insects in plantations of exotic trees.

Forest insect and human interactions

37

Fig. 3.4 Extensive plantations of exotic trees, such as Pinus taeda, which is widely planted in southern Brazil, can provide a favorable habitat for establishment of damaging exotic insects (Santa Catarina State, Brazil).

Indigenous Insects in Plantations of Native Trees Over much of Asia, Europe and North America, forest plantations tend to be established with indigenous species. These include Cunninghamia lanceolata, Pinus massoniana and Populus spp. in China, Picea abies, Pinus sylvestris and Populus spp. in Europe, Pinus taeda and Pinus elliotii in southeastern USA and Pseudotsuga menziesii in western Canada and the USA. Plantations of trees indigenous to a region are susceptible to their own complex of indigenous insect pests. Moreover, since the plantations provide a large amount of suitable host material, they can favor buildup of insects that would be less damaging in surrounding natural forests composed of several tree species. In China and Vietnam, the defoliating caterpillar Dendrolimus punctatus is an important pest of indigenous pine plantations. In eastern and central China Pinus massoniana plantations are defoliated. In Vietnam outbreaks have occurred in plantations of P. merkusii, often established on steep deforested slopes where they are an important source of lumber, pulpwood, fuel wood and resin (Dao Xuan Truong 1990). A sawfly, Nesodiprion biremis, is one of several species of concern in pine nurseries and plantations in northern Thailand. Nun moth, Lymantria monacha, is a defoliating caterpillar of both Picea abies and Pinus sylvestris in central

Europe. This insect was originally a pest of natural P. abies forests but readily adapted to pine plantations established in this region. In Poland, P. sylvestris plantations are established on low nutrient, sandy soils or abandoned marginal agricultural lands. Widespread outbreaks of this insect occurred in these plantations between 1978 and 1983, and again during the early to mid-1990s. Other defoliating insects of P. sylvestris plantations in Europe include Bupalis piniarius, Dendrolimus pini, Panolis flammea and sawflies, Gilpinia spp. (Speight & Wainhouse 1989, Ciesla 1994). In North America, several pine sawflies and tip and shoot borers such as the Nantucket pine tip moth, Rhyacionia frustrana, and the white pine weevil, Pissodes strobi, are pests of indigenous conifer plantations. Reproduction weevils, such as Hylobius pales, are damaging to pine seedlings in plantations (Drooz 1985). Poplars, Populus spp., are important plantation species in many parts of Asia, Europe and North America because of their rapid growth and excellent wood quality and are subject to damage by a number of insects. In central China, for example, the Asian longhorn beetle, Anopolphora glabripennis, damages older poplar plantations (Schmutzenhoffer et al. 1996, Lingafelter & Hoebeke 2002). Mahogany shoot borers, Hysipyla grandella and H. robusta, are pests of young mahogany, Cedrela, Khaya,

38

Forest entomology: a global perspective

Swietenia, etc., throughout the tropics wherever these trees are grown in plantations and can cause significant growth loss and tree deformity (Browne 1968, Horak 2001, Wagner et al. 2008).

Indigenous Insects That have Adapted to Exotic Trees Many indigenous species have adapted to the presence of a new host in their environment due to establishment of forest plantations with fastgrowing exotics. In Colombia, where extensive plantations of exotic species of Cupressus, Eucalyptus and Pinus have been established, several indigenous insects have found the plantations to be suitable host material. According to one report, 29 indigenous foliage feeding insects are associated with plantations of exotic conifers, including caterpillars of the family Geometridae, walkingsticks and leaf cutting ants. Several geometrid caterpillars, Cargolia arena, Chrysomima semilutearia, Glena bisulcata and Oxydia trychiata, have caused at least moderate levels of defoliation (Rodas P. 1998). In Chile, larvae of Ormiscodes cinnamomea, which normally feed on foliage of broadleaf trees of the genus Nothofagus, have adapted to plantations of Pinus radiata and can defoliate young pines. Ohmart and Edwards (1991) provide a worldwide review of indigenous insects that have found species of Eucalyptus to be suitable host material. This includes 223 species in Brazil, 96 in China, 94 in India, 31 in New Zealand, 105 in Papua New Guinea and 62 on the island of Sumatra, Indonesia. Termites and leaf cutting ants are among the more damaging insect pests. In northern Scotland, the pine beauty moth, Panolis flammea, a defoliator of Scotch pine, Pinus sylvestris, has adapted to plantations of the Pacific Coast subspecies of the North American lodgepole pine, Pinus contorta ssp. contorta, and has defoliated 10–19-year-old plantations (Speight & Wainhouse 1989). A tussock moth, Orgyia mixta, which normally feeds on Acacia and other broadleaf trees in eastern Africa, was discovered feeding on the foliage of Pinus radiata in plantations established in Kenya during the 1950s and has caused localized defoliation in plantations of this species in both Kenya and Zambia (Schabel 2006).

Exotic Insects in Plantations of Native Trees Several cases are known of exotic insects becoming

pests of plantations of indigenous trees. In North America, two species of sawflies indigenous to Eurasia, the European pine sawfly, Neodiprion sertifer, and the European spruce sawfly, Gilpinia hercyniae, have damaged plantations. The European pine shoot moth, Rhyacionia buoliana, is a pest of plantations of indigenous pine plantations in parts of North America (Drooz 1985).

Exotic Insects in Exotic Tree Plantations Exotic, invasive insects introduced and established in plantations of exotic forest trees have the potential to cause extensive and widespread damage. A classic example of an exotic insect in exotic forest plantations is the wood wasp, Sirex noctilio, in the southern hemisphere. Monterrey or radiata pine, Pinus radiata, was first planted in the late 19th century in southern Australia, Chile, New Zealand and South Africa. The excellent growth and form of this tree encouraged widespread planting. These provided raw material for thriving lumber and paper industries, especially in Chile and New Zealand, which each presently have in excess of 1.5 million ha of P. radiata plantations. S. noctilio first appeared in New Zealand during the early 1900s where it was probably introduced via unprocessed pine logs imported from Europe. Plantations established in New Zealand during 1920–30 stagnated because there was no market for small logs from thinning operations. This made the plantations susceptible to S. noctilio and its associated fungus, Amylostereum areolatum. By 1947, extensive tree mortality occurred, primarily in unthinned plantations. S. noctilio subsequently appeared in P. radiata plantations on the Australian island of Tasmania and on the Australian mainland in 1952. In 1980, infestations were detected in South America where this insect damaged plantations of P. elliotii and P. taeda in Argentina, southern Brazil and Uruguay. In 1994, infestations were discovered in P. radiata plantations in the Cape Peninsula of South Africa and in Chile in 2001, again on P. radiata (Ciesla 2003a). Several other insects have been introduced into exotic pine plantations throughout the tropics and in the temperate regions of southern hemisphere (Table 3.2). Eucalypts, which comprise about 20% of the world’s forest plantations, have been victims of introduction and establishment of many insects from Australia. These include several defoliators, sucking insects and wood borers (Table 3.3, Ohmart & Edwards 1991).

Forest insect and human interactions

39

Table 3.2 Examples of introduction and establishment of damaging forest insects into exotic plantations of Pinus worldwide. Mode of introduction

Type of damage

Species

Native range

Introduced range

Bark and ambrosia beetles

Hylurgus lignipera Red haired pine beetle Ips grandicollis

Europe

Chile, New Zealand

North America

Australia

Buprestis novomaculata Sirex noctilio Sirex wood wasp

Europe

Chile

Eurasia

Australia, New Zealand, South Africa, South America

Cinara maritima

Mediterranean Europe, Near East Eurasia?

Argentina, Brazil, Chile Africa, Australia, New Zealand, North America, South America China, Korea, northeastern USA China

Unknown

North America: Canada, USA South America: Argentina, Chile

Unknown

Wood borers

Sucking Insects

Pineus boerneri

Shoot borers

Matsucoccus matsumurae Oracella acuta Loblolly pine scale

Japan

Rhyacionia buoliana European pine shoot moth

Europe

North America

Plantations of Cupressus lusitanica were established in eastern and southern Africa where it was favored because of its rapid growth and excellent form. In 1986, a giant conifer aphid, initially identified as Cinara cupressi, was discovered in Malawi. Infestations spread into neighboring countries and caused extensive tree mortality. In Kenya, where 46% of the forest plantations were composed of C. lusitanica, damage was especially severe. This insect was ultimately redescribed as a new species, C. cupressivora, which is believed to be indigenous to the eastern Mediterranean region of Europe and the Near East. The severe impact of this insect on cypress plantations in Africa has forced foresters to shift to other plantation species, especially eucalypts (Ciesla 1991, Ciesla et al. 1995, Watson et al. 1999). The sawfly, Neomatus oligospilus, has a circumpolar distribution in the northern hemisphere. This species has been introduced into southern Africa, Argentina, Australia and New Zealand, where it has become a pest

Probably wood containing bark strips Unprocessed logs, wood products Unprocessed logs, wood products Unprocessed logs, wood products

Nursery stock

Ornamental pines Infested scion material

of both poplar and willow plantations (Dapoto & Giganti 1994, Ede 2006). Agroforestry Agroforestry, the practice of integrating trees with agricultural crops, has been used in some areas of the world for thousands of years. Examples include interplanting of leguminous trees that have nitrogen fixing bacteria on their roots, growing crops under the shade of trees (understory intercropping) and establishment of plantations of trees to protect crops from wind damage. While some crops may be grown under shade of a natural forest, most agroforestry involves planted trees (Wilkinson & Elevitch 2000). One of the most widely used trees in agroforestry in the tropics is the leucaena, Leucaena leucocephala. This tree is native to Central America, southern Mexico and northern South America. It has an exceptionally rapid growth rate and a wide range of uses. Leucaena psyllid,

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Forest entomology: a global perspective

Table 3.3 Examples of introduction and establishment of damaging forest insects into exotic plantations of Eucalyptus worldwide Type of damage

Species

Native range

Introduced range

Mode of introduction

Foliar damage

Gonipterus scutellatus Eucalyptus weevil

Australia

Africa, Asia, Europe, North America (California), South America

Unknown

Wood borers

Phoracantha semipunctata, Eucalyptus longhorn borer

Australia

Wood products

P. recurva Eucalyptus longhorn borer

Australia

Africa, Europe, Near East, New Zealand, North America (California), South America Africa, Europe, New Zealand, North America (California), South America

Glycaspsis brimblecombei Red gum lerp psyllid

Australia

Ctenarytaina eucalypti Blue gum psyllid

Australia

Leptocycbe invasa Blue gum chalcid Ophelimus eucalypti Eucalyptus gall wasp

Australia?

Sucking insects

Gall insects

Australia?

Heteropsylla cubana, feeds on new shoots of leucaena and occurs throughout the native range of this tree. In 1983, leucaena psyllid was detected in Florida, USA and a year later, in the Hawaiian Islands. Between 1984 and 1993, the spread of this insect across the Pacific Islands into Asia, Australia and, ultimately, Africa, was of spectacular proportions. Newly established infestations caused extensive drying and dieback of new shoots, growth loss and some tree mortality. The future of this tree in agroforestry was threatened until pest management methods, which emphasized introduction of the pysllid’s natural enemies, were developed (Banpot Napompeth 1994). Urban Forests Trees contribute significantly to the quality of life of people who live in cities and urban areas. They add beauty and harmony to what would otherwise be a

Chile, Mauritius, Mexico, Portugal, Spain, USA (California) South America Ireland, Mexico, Sri Lanka, USA (California) Africa, Asia, Europe, North America Africa, Mediterranean Europe, Near East, New Zealand

Wood products

Infested planting stock?

Infested planting stock?

Unknown Unknown

harsh, cold landscape. They provide shade (Fig. 3.5), filter noise and break up “heat islands” created when the sun’s radiation is absorbed by tall buildings. Even in rural areas, planted ornamental trees along roadsides can add a special character to the landscape (Fig. 3.6). Urban forests are special ecosystems because they are composed almost entirely of planted trees, many of which are exotic. In some parts of the world, especially areas with semi-arid or arid climates, these trees are planted in areas that were previously steppe or desert vegetation. Since many of the trees established in urban forests are exotic, many of the insects that damage these trees are also exotic and have spread to new locations with their hosts. Moreover, in developed countries, where people are able to spend leisure time working in their gardens, they are more aware of insects whose damage may be largely cosmetic (e.g. species of gall insects). Damage by these insects would be largely unnoticed in forests.

Forest insect and human interactions

41

Fig. 3.5 Neem trees, Azadirachta indica, provide welcome shade from the hot African sun (Dogondoutche, Niger).

Several introduced insects have caused significant damage to urban forest landscapes. The smaller European elm bark beetle, Scolytus multistriatus, introduced into North America is a vector of Dutch elm disease, a pathogen with origins in Asia that has killed millions of elm shade trees. In many communities in northeastern and north central USA and adjoining Canada, American elm, Ulmus americana, was the dominant

shade tree. When the disease and its insect vectors arrived, large numbers of elms were killed and many communities lost all or most of their shade trees. Gypsy moth, Lymantria dispar, an introduced defoliator of broadleaf trees, has caused extensive defoliation in urban forests in many areas of eastern USA. Moreover, large numbers of larvae associated with outbreaks are a nuisance in gardens, parks and green belts.

Fig. 3.6 Columnar cultivars of the Mediterranean cypress, Cupressus sempervirens, planted along a rural road add a special character to the landscape of Tuscany, Italy.

42

Forest entomology: a global perspective

Emerald ash borer, Agrilus planipennis, native to Asia, was discovered in 2002 in Michigan, USA. Since its arrival, it has spread into several states and adjoining portions of Canada. This insect is an aggressive tree killer of ash, Fraxinus spp. Since its establishment, millions of ash trees have been killed. This beetle threatens trees in both natural and urban forests, where several species of ash are important shade trees. In Europe, horse chestnut, Aesculus hippocastanum, a tree with a dense crown, dark green foliage and cultivars that produce clusters of flowers in several colors, is a popular shade tree. In 1985, a leaf mining insect, Cameraria ohridella, was observed for the first time on this tree by Lake Ohrid in Macedonia. Heavy infestations produce brown blotches of discoloration on the leaves and are unsightly (Plate 1). In 1989, infestations were detected in Austria and within a few years the insect spread throughout most of Europe. The origin of this leaf miner is still unknown (Deshka & Dimic 1986, Busko 2006, Ivinskis & Rimðait€e 2006). Neem, Azadirachta indica, a tree native to the Indian subcontinent, has been widely planted to provide shade in cities and villages in the SudoSahelian region of Africa. It is one of a few trees that can thrive in the hot, semi-arid climate of western Africa and still produce a crown of dense foliage. This tree provides welcome shade under which local residents work and set up outdoor markets. Oriental scale, Aonidiella orientalis, an insect that feeds on the foliage, petioles and stems of host trees, became established across northern and central Nigeria, where neem is widely planted as a shade tree. Infestations cause a browning of the foliage and thinning of the crown. A long-term strategy to minimize impacts of damaging pests of urban trees and shrubs is to plant as great a variety of species as local conditions will allow. Most biotic pests (insects, mites, fungi, etc.) are relatively host specific. Therefore, should a new pest appear, only a portion of trees would be affected. This was a lesson learned, somewhat painfully, following introduction of the insect-vectored Dutch elm disease into both Europe and North America.

BENEFICIAL FOREST INSECTS

keep the so called “damaging” insects in check. Still others contribute to the breakdown of dead vegetative and animal matter. In addition, a number of insects that feed on forest trees produce direct benefits to humans because they are sources of food or other products of value or have been subjects of curiosity because of their striking appearance or unique habits. Products of Value to Humans Honey Before they were domesticated, honey bees, e.g. Apis mellifera, were forest dwellers. They produce honey, the oldest sweetener known to humans. Honey is produced from the nectar of flowering plants, was originally stored in wax combs in trees hollowed out by decay fungi, and used as a food to feed larvae throughout the year and adult bees during winter. Primitive human societies hunted for trees where bees stored honey to obtain the precious sweetener. Eventually beekeeping, or apiculture, the science and art of raising bees in artificial hives, where honey can easily be harvested, became a major agricultural enterprise. Today, domestic honey bees have become more important as pollinators of agricultural crops than for production of honey. Food Larvae of several forest insects are locally important food sources. Pandora moth, Coloradia pandora, larvae feed on several species of pine in western USA. They are a traditional food source for the Paiute, an indigenous tribe from Nevada and southern California. During outbreaks, larvae are collected as they drop from host trees to the ground to pupate and are roasted and dried on site (Blake & Wagner 1987). In southern Africa, larvae of the mopane worm, Imbrasia belina, feed on the foliage of the mopane tree, Colophospermum mopane, and are an important local food source. Roughly 65% of rural people in southern Africa collect the larvae for subsistence use and 35% sell the dried larvae at rural markets. Some are traded locally and internationally as snacks and canned products (Browne 1968, Ditlhogo et al. 1996, Makhado & Potgeiter 2009). Silk

A vast number of forest dwelling insects have beneficial functions in the forest ecosystem. Many species pollinate plants. Others, such as parasitoids and predators, help

Larvae of the silkworm, Bombyx mori (Lepidoptera: Bombycidae), feed on foliage of mulberry, Morus spp.

Forest insect and human interactions The larvae spin cocoons of silk in which they pupate. Silk is the most luxurious of fibers, with an unsurpassed beauty, luster and softness. It takes dyes well and, although soft, is one of the strongest, most durable fibers known. Silk has been produced commercially in China and India for centuries. The method of silk production was a carefully kept secret and silk became a major item of trade with Europe and the Near East. The “Silk Road” was a major trade route that began in China, passed through central Asia and ended in the eastern Mediterranean region. Silkworm larvae have been domesticated for so long and reared in flat baskets containing mulberry leaves that they have lost their ability to grasp leaves and branches of host trees and could no longer survive in the wild (Fig. 3.7).

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Several other forest caterpillars are sources of “wild” silk. Wild silk is coarser than cultivated silk, which makes it more durable and practical for clothing. Most wild silk producing species feed on the foliage of broadleaf trees and some have been at least semicultivated. In China, a wild silk known as tussah silk is produced by the tussar moth, Antheraea pernyi. Larvae feed on foliage of oaks, Quercus spp., and are reared on pruned oak shrubs 1.5–2 m high. A related species, A. yamamai, has been cultivated in Japan for over 1000 years (Ciesla 2002a). Dyes and Inks In Mediterranean Europe, a scale insect, Kermococcus vermilis, which infests the stems of several oaks, was highly prized because female adults were a source of red dye used for dyeing wool and silk before other dye sources were available. The Aleppo gall, Cynips gallae tinctoria, produces a gall on oaks in Mediterranean Europe and the Near East. These galls have a high tannic and gallic acid content and have been of commercial importance since the time of ancient Greece for manufacture of inks that did not fade (Ciesla 2002a). Lac In India and neighboring countries, the lac insect, Kerria (¼Laccifer) lacca, which feeds on the stems of broadleaf trees, is known for secretion of a substance known as lac. Lac is scarlet in color and used to dye wool and silk. It is also used as a cosmetic, a medicinal drug, as shellac to coat candies and give fresh fruits and vegetables a glossy finish. The lac insect is cultured in India and to a lesser extent, Myanmar and Thailand where it is considered one of the most valuable of all plant parasitic insects. Biological Control

Fig. 3.7 Silkworms, Bombyx mori, are reared in large baskets for production of silk (Hanoi, Vietnam).

Occasionally, an insect pest in one part of the world may serve as a biological control agent for a noxious plant in another. In western China, Central Asia and Russia, salt cedars, Tamarix spp., are indigenous and planted for shelterbelts and sand dune stabilization. When introduced into southwestern USA, these trees became invasive in riparian areas and displaced native vegetation. Larvae and adults of five species of diorhabda beetles, Diorhabda spp., feed on foliage of salt cedar and

44

Forest entomology: a global perspective

damage plantations established within the natural range of these trees. Diorhabda beetles have been introduced into the USA as a biological control agent for these invasive trees (DeLoach et al. 2003, Hudgeons et al. 2007, Tracy & Robbins 2009).

Curiosities Some forest insects are colorful, eye catching or have habits that arouse human curiosity. Some have even become subjects of various art forms. In North America, the monarch butterfly, Danaus plexippus, the larvae of which feed on milkweeds, Asclepias spp., is a migratory species. In autumn, the large, colorful orange and black butterflies travel thousands of miles to spend the winter in groves of trees along the coast of California or in dense forests of sacred fir (oyamel), Abies religiosa, in central Mexico. The migratory habits of this insect have intrigued amateur and professional entomologists alike and many people have tracked migration patterns to help locate overwintering sites. Overwintering sites in central Mexico were not discovered until 1975 and are now threatened by illegal logging (Plate 2, Brower 1977, Slayback et al. 2007). On the Greek island of Rhodes, the forest of the Petaloides Valley (petaloides is Greek for butterfly) is the site of an annual migration of the moth, Panaxia quadripunctaria. Moths mate in this valley and then depart to lay eggs in other locations in autumn. The valley, with its migrating butterflies, is an important tourist attraction (Petanidou et al. 1991). Other forest insects are favorites among collectors because of their brilliant coloring. Species of morpho butterflies, Morpho spp., of tropical forests in Mexico, Central and South America, with their bright metallic blue or violet wings, are a classic example. In addition, adults of wood boring beetles of the families Buprestidae (metallic wood borers or jewel beetles) and Cerambycidae (long horned beetles) are favorites among insect collectors. Striking colors and patterns on a few forest insects have made them subjects of various art forms. In China, longhorn beetles of the genus Anopolphora, several of which are damaging wood borers, have been the subject of ivory and wood carvings (Lingafelter & Hoebeke 2002). On Roatan Island, Honduras, females of a sawfly, Sericoceros mexicanus, that feed on the foliage of sea grape, Coccoloba uvifera, are the subject of weavings made by backstrap weavers (Fig. 3.8, Ciesla 2002b).

Fig. 3.8 A weaving by traditional backstrap weavers depicts female adults of the sawfly, Sericoceros mexicanus (Roatan Island, Honduras).

FOREST ENTOMOLOGY AS A CAREER Forest entomology can be an exiting and rewarding career and several paths are open to individuals interested in working with forest insects. The following sections outline educational requirements and alternative career paths for people who chose to work in forest entomology or its sister field, forest pathology, the study of forest diseases.

Educational Requirements Many of today’s professional forest entomologists begin their formal education in an undergraduate forestry program, develop an interest in forest insects (or diseases) and decide to specialize. Others take

Forest insect and human interactions undergraduate training in general biology, agricultural science or entomology. Regardless of their undergraduate training, graduate level training in forest entomology leading at least to a Master of Science (MSc) degree is essential. A high proportion of prospective forest entomologists now undertake a program of study leading toward a Doctor of Philosophy (PhD) degree with some continuing on with postdoctoral studies.

Additional Skills In addition to formal training in forest entomology and related courses, other skills are essential. Certainly, a working knowledge of forest entomology’s sister field, forest pathology, is important, especially for those who are frequently asked to diagnose tree pest problems that could be caused by insects, pathogens or other agents. The ability to write is a critical skill. Forest entomologists should document their observations and findings either as reports or formal publications in scientific journals for the benefit of resource managers, professional colleagues or in the popular media to inform the public on the status and management of damaging forest pests. The ability to draw is a desirable skill, especially for those who wish to specialize in insect taxonomy. An interest and skill in photography is essential to document insect damage, the insects themselves and activities undertaken to manage insect pests. Since many of the duties of a forest entomologist involve design of experiments or sampling insect populations, a basic knowledge of statistics is essential. While forest entomologists are not expected to be expert statisticians, they should possess a basic understanding of the science to allow them to work effectively with a statistician to design experiments and develop appropriate protocols for insect sampling. Perhaps most important is the ability to communicate and work effectively with people of other professions and the general public. As specialists, forest entomologists work with forest managers or forest landowners to communicate why an insect may have become a pest and recommend appropriate actions to address forest insect issues. They must also be able to work effectively with the public, including those who may be opposed to taking action against a pest, especially on public lands.

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Research Research on the biology, ecology and management of damaging forest insects is a critical activity needed to develop appropriate forest insect management strategies and tactics to effectively reduce losses. New approaches for management of forest insects are continuously needed to meet changing needs of forest managers or to develop environmentally friendly pest management tools. Moreover, new pests are continuously appearing and provide additional challenges for forest managers. Therefore, there is a need for a strong cadre of research scientists dedicated to gaining an improved knowledge of damaging forest insects, their interactions with local climate, their hosts and natural enemies and to develop new approaches to forest insect management that can be readily implemented in the field. Universities Most entomology or forestry faculties at universities will employ one or more forest entomologists. Usually the duties of these individuals involve a combination of teaching undergraduate and graduate level courses in forest entomology and related topics, providing guidance to graduate students and conducting research, often funded by external grants. If forest entomologists are employed at a public university, their duties may also involve extension; the process of putting new knowledge obtained through research into practice. Forest Research Institutes Most countries with a significant forest resource base have national forest research institutes. These may be part of the national forest service or function as a separate entity. In large countries with a diverse forest base, such as Canada, China and the USA, the research capacity may be decentralized into a network of regional forest research centers. In other countries (e.g. Austria, Brazil, Kenya, Romania and Tanzania) most or all of the forest research capacity is centralized in a single laboratory. Forest research institutes employ scientists to conduct research on the ecology, management and protection of forests. Forest entomologists and forest pathologists are an integral part of the research team. Research forest

46

Forest entomology: a global perspective

entomologists conduct studies on the biology, ecology and management of forest insects of concern to forest managers and may also provide technical assistance in their management. In addition, forest research entomologists may become part of interdisciplinary teams to address broader, more complex issues such as the overall management of a forest ecosystem. A continuing challenge for forest research is to obtain sufficient funding and support to adequately address the needs of a country’s forest sector. Even in developed countries, support for forest research is often inadequate. Therefore, scientists are unable to travel to the field to conduct studies and the research capacity becomes ineffective and disrespected by the people who desperately need the knowledge generated by these institutes.

Applied Forest Entomology Applied forest entomologists conduct surveys to detect insect pests, delineate areas affected by outbreaks and to obtain clues as to their underlying causes. They provide continuing education to foresters and forest workers in recognizing insect pests and their damage and provide technical assistance to forest managers and forest landowners in forest insect management. They may be called on to provide leadership to suppression projects designed to reduce insect populations and commensurate damage. One of the more challenging tasks required of applied forest entomologists is to put into practice new knowledge or technologies obtained through research. This may involve design and conduct of pilot studies, usually in collaboration with research counterparts, to determine how the new knowledge or technologies will perform under operational conditions.

Government Agencies National, state or provincial forestry departments often employ forest entomologists to conduct forest insect surveys, provide technical assistance and related duties. In the USA, the Forest Service of the USDA has a forest health protection organization staffed with specialists at the national and regional level. In most Forest Service regions, a network of zone or field offices that serve several National Forests, as well as other federally

administered lands (e.g. National Parks, Indian Reservations, Bureau of Land Management, Military Reservations) are staffed with forest entomologists and forest pathologists who work as a team to address forest pest issues. Most state forest services also employ one or more forest entomologists who perform the same duties on state and private forest lands. Since most forested areas in the USA are a mosaic of Federal, state and private lands, it is critical that technical specialists from the USDA Forest Service and state forest services work as partners to address pest issues on lands of mixed ownership. In Canada, applied duties related to forest entomology and pathology rest with the provincial forest services whereas the research function is the responsibility of the Canadian Forest Service. In many countries, the applied aspects of forest entomology and pathology are assigned to foresters who have a special interest and, in some cases, some specialized training in these areas. In Chile, each regional office of Corporacion Nacional Forestal (CONAF), the Chilean national forest service, has a forester specifically assigned to matters relating to forest health. At the national headquarters in Santiago, several staff specialists have graduate level training in forest entomology or forest pathology. Private Industry The two primary private sector employers of forest entomologists are companies that manufacture products to support forest insect and disease management and the forest industry. Companies who manufacture products to support forest pest management (e.g. chemical or biological insecticides, pheromones, insect traps, etc.) may employ forest entomologists as technical representatives whose primary duties are to promote and seek applications and markets for their products and to work with forest managers or public sector forest entomologists to evaluate product efficacy. Forest companies have a need for technical expertise in forest insect management on forest lands they own, which provide raw material for their mills. In many countries, forest companies assign forest insect and disease management responsibilities to an area or district forester as all or part of their duties and rely on expertise available in public resource management agencies such as national, state or provincial forest services, forest research institutes or universities for technical support. However, there are exceptions. In

Forest insect and human interactions Chile, Forestal Arauco, a large forest company, has its own research department, which includes expertise in forest entomology. Moreover, a Chilean consortium of forest companies has chartered Controladora Contra Plagas, a laboratory that mass rears parasitoids for classic biological control of European pine shoot moth, Rhyacionia buoliana, and is staffed by a team of specialists with training in forest entomology and techniques for mass rearing of insects under laboratory conditions (Controladora de Plagas Forestales 1997).

International Opportunities Opportunities also exist for international work in forest entomology. The ability to work effectively in other countries requires an additional skill, the ability to read, converse and write in more than one language. An ideal combination of language skills is English, French and Spanish, with a fluency in at least one and a working knowledge of the other two. The Food and Agriculture Organization of the United Nations (FAO) provides technical assistance in all aspects of forestry to its member countries and has a full-time forest health specialist on the staff of the Forestry Department at its international headquarters in Rome, Italy. This post could be filled by either a forest entomologist or forest pathologist. In addition, FAO may employ consultants on a short-term basis to provide technical assistance on matters dealing with forest insects. Other United Nations Agencies such as the United Nations Development Programme (UNDP), the United Nations Office of Project Services (UNOPS) or the World Bank (WB) fund international development projects via grants or lowinterest loans. These include forest sector development projects, which may include an element of protection from damaging forest insects and require the services of a forest entomologist, at least on a short-term basis. Many developed countries have organizations that provide bilateral financial and technical assistance to

47

developing countries. Examples include the Canadian International Development Agency (CIDA), The Danish International Development Agency (DANIDA), Gesselschaft für Technische Zussamenarbeit (GTZ) (Germany) and the US Agency for International Development (USAID). The European Union (EU) also funds international development assistance projects. If forest sector development projects are funded by these agencies, they could require technical expertise in forest entomology. National forest research institutes and universities may provide opportunities for short-time technical exchange visits between scientists from different countries for teaching and/or research. The Consultative Group in International Agricultural Research (CGIAR) has a number of international research institutes worldwide including two that address forestry issues: the International Center for Research in Agroforestry (ICRAF) based in Nairobi, Kenya and the Center for International Forestry Research (CIFOR) based in Bogor, Indonesia. ICRAF conducts research and disseminates information on appropriate agroforestry technologies and CIFOR is a center for research and global knowledge on forest conservation and improvement of the livelihoods of people in the tropics. Both institutes have dealt with forest health concerns and have need for forest entomologists and forest pathologists, at least on a short-term basis. Finally, there are volunteer organizations, such as the US Peace Corps, which provides opportunities for recent graduates in forest entomology, among many other disciplines, to gain international experience. Beginning in 1967, a team of Peace Corps volunteers worked for several years in Chile and established the groundwork for a viable and sustained program in forest entomology. They designed and taught courses in forest entomology at several universities, conducted surveys in both natural forests and forest plantations and identified a number of insect pests (Billings et al. 1973).

Chapter 4

Monitoring Forest Insects, their Damage and Damage Potential

INTRODUCTION

Is the Insect Present?

Conducting surveys to monitor forest insects and their damage is one of the most critical activities carried out by applied forest entomologists. Data acquired from these surveys keep forest managers and other stakeholders updated on the status of key pest species on a local, regional or country-wide basis. These data also provide, in part, the basis for decisions for or against pest management and the most appropriate strategies and tactics to be taken to reduce losses (see Chapter 5). While the decision to take action against a forest pest lies with the forest manager or landowner, it is the responsibility of forest health specialists, both entomologists and pathologists, to provide the best data possible on which to base these decisions.

This question is directed towards either established or potentially invasive insects. Approaches designed to ensure early detection of exotics include inspections of plant and wood products at ports of entry and/or warehouses and deployment of traps to monitor for presence or absence of the insect(s) in question.

OBJECTIVES Forest insect monitoring is designed to address the following questions.

How Abundant is the Insect? Estimation of insect abundance, or population density, addresses concerns such as: . .

Are populations increasing or decreasing? Is an outbreak imminent?

Insect numbers are typically expressed as the number of life stages (eggs, larvae, nymphs or adults) on a per unit basis such as per branch of a given length, unit of foliage area, bark surface or numbers of insects collected per trap.

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Forest entomology: a global perspective

Where are the Insects and their Damage? Occurrence of insects and their damage must be described in a spatial context, either with regard to political units affected (states, provinces, counties), land ownership (government, industrial private, non-industrial private) or landscape features (drainage basins, mountain ranges, vegetation types, elevations). What is the Severity of Damage? Severity of damage caused by insects can be expressed in a number of ways. Tree mortality caused by bark beetles is expressed as number of trees per spot or per hectare in a drainage basin or land ownership class. Foliar injury is usually classified by predetermined intensity classes (e.g. light, moderate, heavy) or area of aerially visible defoliation. Why is the Insect Abundant? Occurrence of an insect outbreak is usually a symptom of a deeper forest health issue. Therefore, it is important to explore site and stand conditions, past forest management practices, climatic anomalies or other factors that may be favoring increases in insect numbers. What is the Potential for Damage by Future Generations of the Insect? This is a critical question if actions are being considered to suppressaninsectoutbreak.Willtheoutbreakcontinueor will populations decline due to natural factors? Sampling methods and prediction models have been developed for many forest insects, usually based on systematic counts of an overwintering stage, such as egg masses, to predict damage during the next growing season. These surveys may be supplemented by observations on levels of disease or parasitism in the population. This information is used to help decide for, or against, direct control and which areas should be treated.

What is the Probability that an Outbreak will Occur? Forest conditions such as stocking levels, age, species diversity, soil conditions or other factors can

predispose forests to forest insect damage. Tree and stand hazard/risk rating systems are designed to assess the probability that an outbreak will occur in the future and the magnitude of loss that can be expected. This information helps forest land managers set priorities for cultural treatments well in advance of an impending outbreak.

When is the Best Time to Apply a Pest Management Tactic? If direct control is undertaken, a pest management tactic must be applied at the time the target insect is most vulnerable. This is especially true of insects that are cryptic and spend much of their life cycle inside a bud, shoot, fruit or cone. Monitoring techniques are available or can be developed to establish when the most vulnerable life stage is present to ensure that pest management tactics are applied at the time of optimum efficacy.

How Effective were the Pest Management Tactics? If one or more pest management tactics are applied to reduce an insect population and its damage, their effectiveness should be evaluated to determine if the objectives of the treatment(s) were achieved. These should consider both immediate effects, such as population reduction, and long-term effects such as reduced levels of defoliation and re-invasion of treated areas from surrounding untreated forests.

SURVEILLANCE AND REPORTING Surveillance and reporting, or field surveillance, is an informal but highly effective process for early detection of potentially damaging insects and other damaging agents in forests. Field surveillance is carried out by local foresters, forest workers or forest landowners who have some familiarity with the agents that could damage their forests. It is done in conjunction with other field work such as layout of timber harvesting operations, fire prevention, trail and road maintenance, wildlife habitat improvement, etc. Should field personnel encounter a pest or notably high levels

Monitoring forest insects, their damage and damage potential of damage, they report the condition to officials with the authority to assess and manage the pest. In the case of a forest company, that responsibility may lie with a district or area forester. On public or nonindustrial private forest lands, the authority may rest with officials in a state, provincial or national forest service or agriculture ministry. Field surveillance is especially effective in areas where local foresters and forest workers cover a relatively small area of forest, where forests are easily accessible or are intensively managed. In order for field surveillance to be effective, forest landowners and workers must recognize pests and early stages of damage. Therefore, periodic training in recognition of signs and symptoms of pests and mechanisms for reporting damage, provided by forest entomologists and pathologists, is an essential part of field surveillance. Leaflets or brochures that describe key pest species are also helpful to raise levels of awareness of forest pests and provide a handy reference to aid in the recognition of pest species and their damage. These materials are typically produced by forest services and/or extension services. The availability of the Internet increases the accessibility of this information and allows for its easy updating as new information becomes available. Timely reporting of pest occurrence to appropriate authorities is critical for field surveillance to be effective. In Canada and the USA, standardized forms are available for reporting of occurrence of damaging pests (Fig. 4.1). Many developing countries lack the capacity to conduct even the most informal of surveillance activities and, even more important, do not have a capacity to identify pest species should they be detected. In 2001, the Australian Centre for International Agricultural Research (ACIAR) initiated a project to increase capacity in forest health surveillance for several South Pacific island nations including Fiji, Samoa, Tonga and Vanuatu. This project uses the term “surveillance” in a broader context than what is described in the above sections. Activities include formal ground surveys in forest plantations and near ports of entry, use of pheromone traps to detect potential insect pests, development of a standardized reporting system, establishment of a capacity for timely identification of potential pest species, and a network to share information (ACIAR 2001, Sanjana Lai 2008).

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ESTIMATING INSECT NUMBERS Monitoring often involves estimating numbers of insects present to: . . .

predict potential for damage by the following generation; time application of pest management tactics; evaluate the efficacy of pest management tactics.

Obviously, it is physically impossible to count all of the insects on a hectare of forest land or even on an individual tree. Therefore sampling schemes are developed to provide reliable estimates of insect numbers that meet objectives of the monitoring. Development of effective methods of insect population sampling requires knowledge of the insect, its biology and ecology as well as some basic knowledge of statistics and sampling design. Considerations Once the objectives of the monitoring have been identified, the following considerations are addressed in development of protocols for sampling an insect population: . . . . .

sample unit (e.g. tree, branch, bole or trap); sample size; where to sample; when to sample; how often to sample.

Sample Unit The sample unit is the place from which the sample is drawn, such as: . . . .

. .

branches of a given length on which numbers of eggs, egg masses or larvae are counted; sections of bark of known area on which counts of bark beetle life stages are made; leaves on which counts of aphids or scales are made; branches containing cones or fruits to estimate levels of infestation by insects that attack reproductive structures; entire trees (seedlings or saplings); an area of soil or litter surface to estimate numbers of cocoons or pupae.

Selection of the sample unit for a given insect is defined by a combination of the biology and habits of

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Forest entomology: a global perspective

Fig. 4.1 USDA Forest Service, forest pest detection report form used by field personnel to report insect and disease occurrence.

Monitoring forest insects, their damage and damage potential the insect and where it is practical to draw the sample. Branch samples are often used for estimating population densities of defoliators. They are also effective sample units if the insect deposits its eggs on branches. Sections of bark of known size are logical sample units for estimating population densities of bark beetles or for insects that deposit their eggs or egg masses on the bark surface of host trees. Use of whole trees as a sample unit is limited to small trees, usually less than 3 m tall and where insect densities are relatively low or are concentrated in colonies. Insect numbers are expressed relative to the sample unit. Population densities of defoliators, for example, are expressed as numbers of eggs, egg masses or larvae per branch of given length or per unit area of foliage surface. Numbers of bark beetles are expressed as insects per square meter of bark surface. Densities of aphids or scales can be expressed as insects or insect colonies per leaf or per branch of known size. In cases where whole trees can serve as the sample unit, insect densities are expressed as numbers of egg masses, larvae or colonies of larvae per tree. Sample Size

Optimum sample size can be determined as follows (Freese 1967): n¼

t2 s E2

Where: n ¼ optimum sample size (e.g. number of trees to sample) t ¼ value of t at n  1 degrees of freedom s ¼ standard deviation of the mean E ¼ level of precision desired (e.g.  x egg masses/tree). Use of this formula is illustrated using field data on numbers of egg masses of pine processionary caterpillar, Thaumetopoea wilkinsonii, on young trees less than 3 m in height in Pinus brutia plantations in northern Cyprus (Ciesla 2003b). The objective of this work was to develop a method to predict defoliation by this insect using egg-mass counts to serve as a basis for decisions for, or against, control. Egg-mass counts were made on each of 30 randomly selected trees from each of two plantations. The following statistics were computed for each plantation and for the two plantations combined (Table 4.1): .

Sample size is a key consideration when designing survey methods to estimate population numbers. Obviously, taking too large a sample is inefficient and involves unnecessary work while under-sampling presents a risk of an inaccurate population estimate. Sample size is a function of the variability of the population, expressed as the standard deviation of the mean, and the desired precision of the estimate.

53

. . . .

average number of egg masses/tree; range of egg masses/tree; standard deviation; standard error; 95% and 99% confidence limits.

Optimum number of sample trees needed was determined for each plantation and the two plantations combined for the following levels of precision:

Table 4.1 Thaumetopoea wilkinsonii egg-mass densities and sampling errors for trees 75% of the foliage discolored.

Numerical defoliation ratings for each crown level were obtained by multiplying the damage rating by the crown level. Tree defoliation rating was the sum of the

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Forest entomology: a global perspective

Fig. 4.2 Overwintering larvae of larch casebearer, Coleophora laricella, attached to spur shoots of western larch, Larix occidentalis, are a sample unit that can be used to predict defoliation by this insect.

three crown level ratings and ranged from a low of 0 to a high of 72 (Table 4.3). Sample point defoliation rating was the mean of the 10 sample trees. Regression analysis was used to define the relationship between overwintering larvae and defoliation rating. This analysis indicated that the quadratic equation: y ¼ a þ bx þ cx2 Where: y ¼ defoliation index x ¼ number of overwintering larvae per 100 spur shoots provided the best fit for the data. A regression model: y ¼ 4.015 þ 0.4419x – 0.00104x2 was computed with

a coefficient of determination (r2) of 0.763 for 2 years of data (Table 4.4) (Ciesla & Bousfield 1974). Sequential Sampling Sequential sampling is a technique in which the sample size is flexible and counts of the attribute of interest are accumulated. As a sample is drawn and counted, a decision is made to either classify the population or continue sampling (Figs 4.3 & 4.4). This approach is best suited for sampling of insect populations and their damage when counts of the attribute of interest can be made in the field. It minimizes sampling time and effort where population densities are clearly high or low. In places where densities are moderate, or highly variable, more samples are required before a population is classified (Waters 1955).

Table 4.3 Matrix used to compute a larch casebearer damage index for three crown levels and five defoliation classes Crown level

Upper Mid Lower

Damage class Weighting factor

Negligible (0)

Light (2)

Moderate (4)

Heavy (6)

Severe (8)

1 3 5

0 0 0

2 6 10

4 12 20

6 18 30

8 24 40

Source: Ciesla & Bousfield 1974.

Monitoring forest insects, their damage and damage potential

57

Table 4.4 Overwintering larch casebearer population density and predicted defoliation based on the quadratic equation y ¼ 4.015 þ 0.4419x – 0.00104x2. Number of overwintering larvae/100 spur shoots (x)

Defoliation index (y)

Predicted defoliation

0–8.9 9.0–26.9 27.0–44.9 > 45.0

Negligible Light Moderate Heavy

0–11.5 11.6–60.4 60.5–136.5 136.6–236.75* *Highest observed population density. Source: Ciesla & Bousfield 1974.

potential damage as either light ( 3.3 egg clusters/ tree), moderate (8.3–14 egg clusters/tree) or severe ( 26 egg clusters/tree). Dominant or co-dominant jack pines, Pinus banksiana, are selected at a sample point, felled and all egg clusters occurring on the needles are counted in the field. A maximum of 10 trees per site are felled and egg clusters counted. Counts are compared with the sequential table (Table 4.5) to decide if the area can be classified or to continue sampling. If no decision can be made after 10 trees, then trees with a cumulative egg cluster count of less than 51 are considered as light, 51–188 as moderate and more than 188 as severe (Tostowaryk and McLeod, 1972). Sequential sampling has also been used for purposes other than damage prediction. In Brazil, a binomial (infested/not infested) sequential sampling plan has been developed to estimate infestation levels and tree mortality in pine plantations caused by the wood wasp, Sirex noctilio. Three sample lines are established in a

Cumulative number of insect life stages

A challenge associated with development of sequential sampling plans is that the spatial distribution of the population of interest (e.g. binomial, negative binomial or Poisson) must be determined. Most epidemic insect populations have a contagious or negative binomial distribution, where the occurrence of one individual of the insect of interest increases the probability that other individuals will be found. Negative binomial distributions are defined by a parameter “K,” which is difficult to compute. Despite these challenges, a number of sequential sampling plans have been developed for forest insects to predict damage and establish a basis for decisions for or against control and several are summarized by Fettig et al. (2001). Procedures for development of sequential sampling plans for forest insects are detailed by Waters (1955). A sequential sampling plan was developed in Canada to predict damage by the Swaine’s jack pine sawfly, Neodiprion swainei, a defoliator of pine forests in both Canada and the north central USA. This classifies

Fig. 4.3 General model for a two-class sequential sampling plan.

Heavy

g

in

pl

m

ue tin

on

sa

Moderate

C

ling

amp

es tinu Con

Light

Number of samples (n)

Fig. 4.4 General model for a three-class sequential sampling plan.

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Forest entomology: a global perspective

Table 4.5 Sequential sampling decision table for Swaine’s jack pine sawfly in Canada based on the number of egg clusters per tree Number of trees

1 2 3 4 5 6 7 8 9 10

Cumulative number of egg clusters

Light









– – 2 7 12 17 22 27 33 36

– – – – – 44 49 54 59 64

– – – – – 49 68 86 105 124

83 102 120 139 158 177 196 214 233 252

Continue sampling

Moderate

Continue sampling

Severe

Source: Tostowaryk & McLeod 1972.

plantation and 40 trees are examined in each line. Each tree is tallied as infested or uninfested. The minimum sample size is 48–57 trees, depending on the desired sample error. Maximum number sample size is 200–250 trees. A sequential sampling plan has also been developed in Brazil to estimate the level of parasitism achieved by the nematode, Deladenus siricidicola, on Sirex noctilio (Penteado et al. 1993, 2008).

.

for contact insecticides and 14 and 21 days for materials that must be ingested (e.g. the bacterial insecticide Bacillus thuringiensis). Count all live larvae and buds on each branch sample in the laboratory. Larval densities were initially expressed as larvae per 100 buds and later as numbers of larvae per branch. Both larval density and percent population reduction for each post-spray sample period was computed (Flavell et al. 1978, 1979).

Evaluation of Treatment Effects TRAPS AND SEMIOCHEMICALS Sampling plans have been developed to evaluate efficacy of chemical and biological insecticides against forest insects. In 1975, a team of forest entomologists and statisticians developed a set of protocols for testing aerial applications of insecticides under semi-operational conditions (pilot control projects) against western spruce budworm, Choristoneura occidentalis, in western USA. These tests usually consist of one or two materials or dosage rates. The protocols consist of the following: . . .

.

.

Knowledge of the chemical ecology of insects (see Chapter 2) has provided a number of tools to monitor for occurrence of damaging forest insects and their abundance. Traps baited with semiochemicals (either pheromones, host attractants or a combination of the two) are used to monitor for: . .

Establish three blocks/treatment and three untreated checks. Size of each block is  1200 ha. Select 25 clusters of three trees each. Trees should be relatively open grown, 11–18 m in height and full crowned. For each sample period, remove one branch about 45 cm long from each of the four cardinal directions (north, east, south and west) from each tree using pole pruners. Sample periods are: pre-spray – 24 h prior to application and post-spray – 7 and 14 days after treatment

. .

presence of invasive insects detection of increases in numbers of insects timing of treatments effectiveness of treatment tactics.

Trapping systems have several advantages for insect monitoring. They are relatively easy to deploy, maintain and collect. Moreover, cost is low.

Presence or Absence of Invasive Insects Several trapping systems using either semiochemicals or trap trees have been developed for early detection of

Monitoring forest insects, their damage and damage potential

59

potentially invasive insects, including defoliators, bark beetles and wood borers. Gypsy Moth Disparlure, a synthetic version of the sex attractant of both the European and Asian forms of gypsy moth, Lymantria dispar, is a powerful attractant for male moths. Traps baited with disparlure are widely used throughout areas not known to be infested by this insect in Canada and the USA. Disparlure baits are deployed in delta traps coated with a sticky substance during the adult flight period, collected at the end of the flight period and examined for presence of male moths. Traps are usually deployed in urban areas, initially at a low trap density of one trap/mile2 (one trap/2.59 km2). If male moths are trapped, trap density is increased the following year to 32–64 traps/2.59 km2. This level of trap density is known as delimitation trapping and is designed to define areas in need of eradication. Following treatment, trap densities are maintained at 32–64 traps/2.59 km2 for 1 year. If no male moths are trapped, density is reduced to 16 traps/2.59 km2 the following year. If results continue to be negative, trap density is reduced back to one trap/2.59 km2 the third year following treatment (British Columbia Ministry of Forests and Range n.d.). Trapping for early detection of gypsy moth is also conducted in other countries. For example, disparlurebaited traps are used to monitor for possible introduction of Asian gypsy moth in the Pacific Island of Fiji (Sanjana Lai 2008).

Bark Beetles and Wood Borers Traps baited with a combination of pheromones and host attractants have been used with varying degrees of success for detection of exotic bark beetles. The most widely used trap for bark beetle monitoring is the Lindgren funnel trap. These consist of a column of funnels that collect beetles as they fly into the attractant source located at the bottom of the trap. They are available in 4, 8, 12 and 16 funnel sizes (Fig. 4.5, Lindgren 1983). In the USA, the USDA Animal Plant Health Inspection Service (APHIS) has worked with several state departments of agriculture to survey for exotic bark beetles as part of the Cooperative Agricultural Pest Survey. In most cases, Lindgren funnel traps, baited

Fig. 4.5 Lindgren funnel trap baited with bark beetle aggregation pheromones is a valuable tool for detection, assessment and mass trapping of several bark beetles (photo by Jose Negrón, USDA Forest Service).

with various combinations of ethanol, turpentine and commercially available bark beetle lures, such as IpslureÒ or ChalcopraxÒ, have been used. Traps are deployed near ports of entry, warehouses that receive goods from foreign countries packed in wooden crating and disposal sites for wooden crating and dunnage from international sources. A joint USDA Forest Service/APHIS Rapid Detection and Response Pilot Project for Exotic Bark Beetles was initiated in 2001 using Lindgren funnel traps baited with several combinations of semiochemicals, including a-pinene and ethanol and ethanol and a threecomponent aggregation pheromone for the Eurasian bark beetle, Ips typographus. This project resulted in the initial detection of banded elm bark beetle, Scolytus schevyrewi, in the USA in 2003 (Negrón et al. 2005). In New Zealand, a semiochemical-based early warning system was initiated in 2002 for detection of exotic bark beetles and wood boring insects. Approximately

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Forest entomology: a global perspective

580 funnel traps baited with a-pinene and ethanol, b-pinene and ethanol or frontalin and ethanol or ipsdienol were placed around the country at high-risk sites, including major seaports, international airports and forests near high-risk sites. Additional traps were used to compare these lures with unbaited control traps to assess their effectiveness in catching established exotic bark beetles and wood borers, and to assess the effects of placing traps at varying distances from host trees. Over 27,000 beetles were captured between 2002 and 2004, but no new insect establishments were detected (Brockerhoff et al. 2006). In Anhui Province, China, a system was developed as part of an FAO/UNDP forestry development project to monitor the spread of pinewood nematode, Bursaphelenchus xylophilus, by trapping its vector, the sawyer beetle, Monochamus alternatus. The trapping system used a-pinene/ethanol baits placed in flight traps. The objective was to collect beetles ahead of the known area of nematode infestation to determine if local populations of sawyer beetles carried the nematode. Traps were established near log decks, sawmill sites and forested areas where high numbers of M. alternatus would be expected to occur (Fig. 4.6, Author’s observation). Trap trees stressed by injection of a weak herbicide are used to monitor the occurrence and spread of infestations of the wood wasp, Sirex noctilio, in pine plantations wherever this insect has become established outside of its natural range. This insect is attracted to stressed trees for egg deposition and, if populations are present in an area, they will readily attack the herbicide-treated pines.

Population Monitoring Defoliating Insects Traps baited with sex attractant pheromones are used to monitor population numbers of forest defoliators including: the spruce budworms, Choristoneura fumiferana and C. occidentalis, in North America; the processionary caterpillars, Thaumetopoea pityocampa and Th. wilkinsonii, in the Mediterranean Basin, and Dendrolimus punctatus in Asia (Fig. 4.7). One of the most widely used and successful semiochemical-based defoliator population monitoring schemes is the Douglas-fir tussock moth Early Warning System (DFTM–EWS). This defoliator has a cyclic

Fig. 4.6 A flight trap baited with a-pinene and ethanol is used to collect specimens of the sawyer beetle, Monochamus alternatus, to determine if they are infected by pine wood nematode, Bursaphelenchus xylophilus (Anhui Province, China).

pattern of outbreaks, which tend to occur in any one location at 7–10 year intervals (see Chapter 2). DFTM–EWS is used throughout much of insect’s range in western North America. The objective of the DFTM–EWS is to detect population increases indicative of impending outbreaks and identify areas where more intensive larval or egg-mass sampling is needed to delineate areas to be considered for direct control. Each EWS plot consists of five traps placed along a line at 23-m intervals and at least 23 m away from roads in stands dominated by Douglas-fir. Traps are placed near the ends of branches about 2.75 m above ground on relatively open-grown host trees. Traps are modified 2.25-litre milk cartons cut to a delta shape

Monitoring forest insects, their damage and damage potential

61

conducted to predict defoliation and to identify areas that should be treated (Ravlin et al. 1991).

Bark Beetles

Fig. 4.7 An entomologist inspects a trap baited with the female sex attractant of the pine caterpillar, Dendrolimus punctatus, to monitor population levels of this defoliator (Anhui Province, China).

with interiors lined with adhesive and, within the trap, a small pellet containing the synthetic pheromone is suspended above the adhesive by a long pin. Traps are deployed between late July to mid-August and collected the following mid-October to early November. Plots are evenly distributed over the host type at a density of at least one plot for every 2115 ha of susceptible host type. Capture of a minimum of 25 male moths per trap serves an “early warning” of an impending outbreak. During over 20 years of monitoring, this system provided advance warnings of 1–3 years for seven of nine outbreaks (Daterman et al. 2004). In eastern USA, numbers of male gypsy moths, Lymantria dispar, caught in pheromone traps are used to identify areas where egg-mass surveys should be

A practical and reliable system to monitor population trends of the southern pine beetle, Dendroctonus frontalis, using pheromone traps has been implemented throughout the range of this bark beetle in southeastern USA. The South-wide SPB Prediction System was developed by the Texas Forest Service and involves monitoring numbers of southern pine beetle and its major predator, the clerid beetle, Thanasimus dubius. Three Lindgren funnel traps baited with the southern pine beetle aggregation pheromone, frontalin, and a rapid-release container of turpentine are established in each county or National Forest to be surveyed. The traps are monitored for 4 consecutive weeks during spring, at peak bloom of flowering dogwood, Cornus florida, which coincides with southern pine beetle long-range dispersal flights. Insects are collected weekly. Average number of southern pine beetles/trap/day and the ratio of southern pine beetle to T. dubius is used to predict population trends for the remainder of the year. This system has proven to be practical and reliable for forecasting general trends (increasing, declining or static) and expected infestation levels (low, moderate, high or outbreak) throughout the region. It serves as an early-warning system that predicts pending outbreaks and/or collapses of southern pine beetle populations (Billings & Upton 2008).

Regeneration Weevils Weevils that breed in roots of freshly cut conifers and feed as adults on seedlings (e.g. Hylobius spp., Pachylobius picivorus) are strongly attracted to resin odors of host trees. Several traps have been developed to estimate population levels of these largely nocturnal insects. Radial discs, ranging in size from 12 to 18 cm in diameter and about 5 cm thick cut from green pines and placed overnight in areas where population level estimates are needed are an effective survey tool in southeastern USA. An average of eight adults per trap have been collected in areas such as recently burned or harvested areas where weevils would be expected to congregate and as many as 42 adults have been collected in a single trap (Ciesla & Franklin 1965). Other trapping techniques include use of pine billets treated with insecticide or polyvinyl chloride (PVC) pitfall traps

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Forest entomology: a global perspective

baited with a 5 : 1 ethanol : turpentine mixture (Rieske & Raffa 1993, Fettig & Salom 1998). Timing of Treatments Pheromone traps are an effective means of timing applications of chemical or biological pesticides. In forestry, this approach is used primarily in intensively managed sites such as nurseries, seed orchards or in urban forests. Pheromones have been used to determine need for and timing of pesticide application for several coneworms of the genus Dioryctria in seed orchards (Turgeon et al. 1994, Meeker 2004). In addition, sex pheromones of several wood boring species of clearwing moths, including lilac ash borer, Podesesia syringae, have been identified and synthesized. Traps baited with the pheromone are deployed 2 weeks prior to adult flight. Insecticides are applied 10–14 days after the first males are captured in traps. Weekly monitoring of insects in traps during the flight period may indicate need for a second treatment if the flight period is extended (Nielsen 1978). REMOTE SENSING Remote sensing, defined as “collection and interpretation of data based on measurement of electromagnetic energy reflected or emitted from objects of interest,” has broad applications in natural resource assessment. Damage caused by many forest insects, especially those that cause tree mortality, defoliation or foliar discoloration, is visible from long distances (Plates 3–6). This lends itself to assessment using various remote sensing technologies ranging from aerial observation (sketchmapping), aerial photos, airborne video and satellite imagery (Ciesla 2000). Remote sensing provides the best set of tools to record the location and intensity of forest insect damage in a spatial context.

application. For example, maps showing the location of outbreaks can be obtained simply by flying over an infested area in a small aircraft and mapping damage. If more detailed data are required, such as estimates of numbers of infested trees and/or timber volume affected, then aerial photos or high-resolution satellite imagery, in combination with a small ground sample, may be needed. Biology of the Target Insect Knowledge of the target pest, its damage and when damage is most visible is critical to ensure that reliable data are acquired. Moreover, individuals responsible for data interpretation must be familiar with the target insect, its damage and vegetation types affected to recognize and accurately classify the damage. Acquisition of Ground Data Use of remote sensing for assessment of forest damage does not eliminate need for some level of field or ground data. Data obtained by remote sensing should be supplemented by ground data to verify that the information obtained via remote sensing is in fact true. Depending on the technique used and the data requirements, acquisition of ground data may range from informal ground checks to establishment of a subsample of ground plots linked to the remote sensing imagery using multi-stage sampling.

Design Remote sensing often involves sampling of large areas of forest with aerial imagery in combination with a small ground sample. Several multi-stage sampling techniques have been developed for natural resource management applications that can be applied to assessment of damage by forest insects and other agents. These are reviewed by Ciesla (2000).

Considerations Effective use of remote sensing for assessment of forest insect damage requires careful consideration of several factors. Data Requirements Data needed to meet the objective of the survey should define the remote sensing tools used for a specific

Aerial Forest Health Surveys Aerial forest health surveys or aerial sketchmapping is a widely used technique for assessment of forest damage caused by insects and other agents. It consists of one or two trained aerial observers flying in a high-wing aircraft who record the location of areas affected by forest damage on topographic maps, usually of a scale of

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Fig. 4.8 An aerial observer briefs a pilot on a mission to map damage caused by cypress aphid, Cinara cupressivora (Nairobi, Kenya).

1 : 100,000 or, more recently, using touch-screen computers (Fig. 4.8). This technique has been used to map damage caused by forest insects and other agents as early as the 1920s in Canada and the USA. In the USA, systematic annual aerial forest health surveys were begun in 1947 with the passage of the Federal Forest Pest Control Act and are a cooperative effort between USDA Forest Service and state forest services. In the USA, approximately two thirds of the forest area is flown annually and between 2002 and 2006, an average of 1.8 million km2 of forest was surveyed each year (Klein et al. 1983, Ciesla 2000, McConnell et al. 2000, Johnson & Wittwer 2008). In Canada, aerial forest health surveys are conducted by provincial forest services. In New South Wales, Australia, aerial surveys are conducted over Pinus radiata plantations by Forest New South Wales, primarily for assessment of tree mortality caused by two introduced insects: the wood wasp, Sirex noctilio, and the bark beetle, Ips grandicollis (Carnegie et al. 2005, 2008). During the 1990s, aerial surveys were used in Kenya to map tree mortality in Cupressus lusitanica plantations caused by cypress aphid, Cinara cupressivora (Ciesla et al. 1995). Aerial sketchmap surveys have also been conducted in southern Brazil, initially for

detection and assessment of tree mortality in pine plantations caused by the woodwasp, Sirex noctilio, but also for other applications (Malheiros de Oliviera et al. 2006). In central Mexico, aerial sketchmap techniques have been used to locate overwintering colonies of the monarch butterfly, Danaus plexippus, in Abies religiousa forests (Slayback et al. 2007). Advantages of aerial sketchmapping are that large areas of remote forest, an average of 80,000–85,000 ha/ day can be surveyed in a relatively short time and at a low cost (less than US$ 0.025/ha). In addition, resultant data are available to forest managers and other stakeholders within 1–2 days. A disadvantage is that data are subjective and their accuracy depends on the experience and skill of individual aerial observers, weather and light conditions and other factors. Uses of Aerial Forest Health Survey Data The primary purpose of aerial forest health surveys is to provide data on location and intensity of forest damage in a timely fashion so that forest managers can plan and conduct appropriate management activities. These data can be used to plan and conduct timber salvage operations or conduct additional surveys to predict next

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season’s damage and begin planning for appropriate pest management actions, including drafting of environmental impact analyses. Another use of aerial forest health survey data is to report regional and/or country-wide pest conditions and for development of historical databases on occurrence of outbreaks of major forest pests. These data are a source of information on trends and cyclic patterns of outbreaks and may be helpful for prediction of future outbreaks. Aerial forest health survey data, stored in a geographic information system (GIS), have been used to evaluate long-term effects of insecticide treatments against at least one forest defoliator; western spruce budworm, Choristoneura occidentalis. An evaluation of the effects of insecticide treatment directed against this insect from 1982 to 1992 on subsequent defoliation was conducted in Oregon and Washington, USA. For each treatment, the extent and severity of defoliation was calculated for the treated area and a set of four nested rings surrounding the treated area for up to 8 years (3 years prior to treatment, the year of treatment, and 4 years following treatment). Insecticide treatments coincided with reduced percentages of defoliation during the year following treatment. However, the percentage of defoliation usually returned to pre-treatment levels by the second year, and severity of defoliation in treated and adjacent untreated areas was nearly identical (Sheehan 1996). Operational Parameters Aerial surveys are flown at an average altitude of 300–450 m above the terrain at an average airspeed of 160–190 km/hour. Flights are generally made between the hours of 08:00 and 14:00. A variety of small aircraft are used for aerial surveys. The Cessna 182, 185, 205, 206 and 210 high-wing aircraft are most widely used. Other aircraft that have been used include the twin engine Partnavia P-68, the Aero Commander 500 and the DeHaviland Beaver. Helicopters have also been used effectively but are considerably more expensive to operate than fixed-wing aircraft. Timing of aerial surveys is critical to their success. Aerial surveys must be conducted when damage is at its peak. In addition, phenomena that could mask damage signatures, such as fall coloring of broadleaf trees, and some conifers (Larix, Metasequoia, Taxodium), in temperate zone forests, should be at a minimum. Over most of western North America, aerial forest health surveys are

conducted during July and August. This time frame effectively captures damage caused by most bark beetles and foliage feeding insects. Aerial observers must be able to: . . .

read maps and know their precise location at all times; recognize aerial signatures of vegetation types and forest damage; have normal color vision and little or no susceptibility to motion sickness.

Roughly 100 hours of flight experience are needed before an aerial observer becomes proficient. Two mapping techniques are used. In mountainous terrain, contour surveys are conducted where the flight path conforms to drainages and ridges in the survey area. In level or gentle terrain, grid surveys of straight flight lines, either east–west or north–south and spaced at about 5-km intervals are flown (Fig. 4.9). Damage types most frequently mapped include defoliation, foliage discoloration, tree mortality or mechanical injury, such as windthrow caused by severe storms. Integration of New Technologies Technologies that have been integrated into aerial forest health surveys over the past 20 years include GIS, global positioning systems (GPS), use of touchscreen computers for data recording and automated flight following (AFF). These have improved observer safety, made data recording more efficient and provide for improved analysis and storage of the data.

Geographic Information Systems (GIS) The ability of GIS to store, analyze and retrieve spatial data on the status of forest insect and disease outbreaks was first evaluated in the USA by the USDA Forest Service during the late 1970s and early 1980s. This technology became operational during the late 1980s and today is used country-wide to store information on the status of major forest pests. GIS is used to develop historical records of pest outbreaks and record changes in their status from year to year. In addition, GIS allows for integration of data on location of pest outbreaks with other resource information such as vegetation type, elevation, slopes, aspect, etc. GIS maps, showing the location of insect and disease damage by year, are now available on the Internet for anyone to study (e.g. www. fs.fed.us/r2/resources/fhm/aerialsurvey/).

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Fig. 4.9 Aerial forest health survey flight patterns. A: contour flight over mountainous terrain. B: grid flight over level or rolling terrain with east-west flight lines.

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Forest entomology: a global perspective

Global Positioning Systems (GPS) The network of orbiting satellites known as GPS, which can be used to triangulate the position of a person, a vehicle, ship or aircraft, anywhere on the Earth’s surface, has revolutionized navigation. This technology has proven to be a valuable tool in the conduct of aerial forest health surveys. GPS receivers, installed in survey aircraft, can be used to locate the starting and ending points of a day’s aerial survey. During grid surveys, the starting and ending points of each flight line can be entered in the GPS receiver and used to help the aerial survey pilot keep the aircraft on the flight line. Touch-Screen Computers Touch-screen computers, linked to GPS receivers, have replaced paper maps for recording location and intensity of forest damage (digital aerial sketchmapping). Products such as the Motion Tablet or Hammerhead, loaded with mapping software (geoLink) and linked to a GPS receiver via the Bluetooth wireless technology, are widely used. An advantage of these systems is that recorded data can be converted into shape files, which are entered directly into a GIS without additional scanning or digitizing. In addition, the system display shows the survey aircraft’s locations. This eliminates the need for aerial observers to continuously locate themselves relative to terrain features (Schrader-Patton 2002).

Automated Flight Following In western USA, where a high proportion of forest land is administered by federal agencies such as the Forest Service (Department of Agriculture) or the Bureau of Land Management (Department of Interior), a protocol has been in place for the survey pilot to advise a local fire dispatch center via radio of the aircraft’s location at 15-minute intervals. AFF triangulates the aircraft’s location using a GPS receiver and transmits these data to a communications satellite, which, in turn, transmits the aircraft’s location to the local dispatch center every 2 minutes. If a signal is not received, a warning message is transmitted. AFF significantly reduces response time in case the survey aircraft develops a problem and must make an emergency landing in a remote area. Aerial Observer Training As mentioned previously, aerial observers require about 100 hours of in-flight training before they become

proficient. Guides for recognition of aerial signatures of forest damage are available that cover most forest ecosystems in the USA (Ciesla 2006, Ciesla et al. 2008). In addition, in the USA, aerial observers are required to attend aviation safety training every 3 years. Training consists of on-line courses and a 1-week classroom session that qualify an observer to manage a fixed-wing aviation program. In-flight training of new aerial observers has become more formalized. Initially, an aerial observer trainee flies with an experienced aerial observer. Later, the trainee receives “front seat” training and has the opportunity to work with the pilot to direct the survey. Hours of aerial observer experience are logged in a manner similar to how a pilot’s flight hours are logged. Aerial Photography Of all of the remote sensing tools currently available, aerial photographs, acquired either by film or digital cameras, provide the highest resolution imagery. Aerial photos have been used for assessment of forest damage where more detailed information is needed than can be acquired from aerial forest health surveys (Ciesla 2000). Photographic Parameters Image Types Forest damage caused by insects or other agents usually first appears as a change in color of tree crowns. This is difficult, if not impossible, to resolve on panchromatic (black and white) film. Therefore, either color or color infra-red (CIR) aerial films or digital images are used for assessment of forest damage. True color aerial images record information in colors as seen by the human eye and, for this reason, are relatively easy to interpret. CIR film is a false color film, sensitive to visible light and the near infrared region (to 0.9 m) of the electromagnetic spectrum. This film has been of value in forestry and other natural resource applications because, in combination with a medium yellow (minus-blue) filter, it can easily penetrate atmospheric haze and there is a greater contrast between certain vegetation types, such as conifer vs deciduous broadleaf forests than can be seen by the human eye or on a true color image. Deciduous broadleaf forests often appear as a bright red color and conifers appear dark red brown or magenta (Plate 7). Color films are available that can be processed to either negatives for

Monitoring forest insects, their damage and damage potential production of photo prints or to positive transparencies. For forest health applications, color transparencies have been preferred because they have higher resolution than prints. Digital camera systems have largely replaced film cameras for virtually all photographic applications. Digital aerial mapping cameras, both small and large format, are now available and widely used. Some digital aerial mapping cameras will record data in both the visible and near infrared regions of the electromagnetic spectrum from whichbothnaturalcolorandCIRimages.Therefore,aerial films have become obsolete. As of December 2009, Eastman Kodak, the sole producer of CIR aerial film, ceased production of this product. Formats A variety of image formats have been used for aerial photographic assessments of forest damage. These include: . . .

small format – 35 or 70 mm aerial photos; mapping format – 230 mm; panoramic format – 115  965 mm.

The most widely used film for forest damage assessment has been the standard 230 mm mapping format.

Scales A range of photo scales have been used with varying degrees of success. These range from 1 : 500 to 1 : 2000 for small format photography, from 1 : 4000 to 1 : 8000 for mapping format photography and from 1 : 30,000 for high-resolution, panoramic format photos. A photo scale of 1 : 8000 is generally considered optimum for forest damage assessments where individual tree crowns must be resolved.

Applications Aerial images have been used with varying degrees of success for a number of diverse applications relating to detection and assessment of damage caused by forest insects.

Inventory of Bark Beetle Damage Multi-stage sampling techniques, using a combination of aerial photos and a small ground sample, have been developed to inventory losses incurred by bark beetle outbreaks (Ciesla 2000). These techniques provide data on:

. . .

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location and area affected; number of trees infested; timber volume affected.

One approach is three-stage sampling where the area affected is stratified by aerial sketchmapping. This area is then sampled with aerial photo stereo pairs or triplets, of a scale of 1 : 4000, 1 : 6000 or 1 : 8000, on which the number of dying or “fading” trees is counted. A small subsample of the aerial photo plots is examined on the ground to correct the aerial photo sample and obtain data for volume loss estimates. Another approach involves a two-stage sample where aerial photo sampling is done over a forested area, the aerial images are interpreted and a small ground sample is taken and used to correct the aerial estimate using statistical procedures such as double sampling with regression. In Germany, complete aerial photo coverage with CIR film has been acquired over the Bavarian National Park, an area of about 24,500 ha, to map areas of windthrow and subsequent damage to adjacent stands caused by the bark beetle, Ips typographus.1 Mapping Forest Defoliation CIR aerial photos, taken from a flying height of approximately 18,300 m, have been used effectively to map defoliation of broadleaf forests by gypsy moth, Lymantria dispar, in eastern USA. This work was done during the mid1980s when defoliation occurred over several eastern states to an extent that was too large an area to be mapped by conventional aerial forest health surveys. The aircraft used to acquire the images was a National Aeronautics and Space Administration (NASA) ER-2 reconnaissance aircraft. The aerial photos were acquired over a 2–3-day period over a multi-state area when defoliation was at its peak, interpreted monoscopically and the location of defoliated areas was transferred to topographic maps (Acciavatti 1990).

Assessment of Forest Pest Management Tactics Both color and CIR aerial photos have been used to assess effectiveness of pest management tactics directed against forest insects. CIR film has been used to assess the degree of foliage protection achieved by aerial application of both biological and chemical insecticides 1

Information in this paragraph is based on an interview with Rainier P€ohlmann, Director of Public Affairs, Bavarian National Park, 12 October 2009.

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Forest entomology: a global perspective

directed against forest defoliators (Plate 8). Color aerial photos have been used to demonstrate effectiveness of thinning pine forests to prevent attacks by the mountain pine beetle, Dendroctonus ponderosae (Ciesla 2000).

confusing. One set of generic definitions, provided by the General Accounting Office (GAO) of the USA, is used in at least one set of guidelines for risk/hazard assessment and is presented below (Schmitt & Powell 2005). Risk Assessment

Other Remote Sensing Tools Other remote sensing tools, such as airborne video, radar, lidarand dataacquired fromairborne satellites, areusedin natural resource management and have been used to some extent for assessment of forest insect damage. To date, their applicability for assessment of forest insect damage is limited by their resolution. Imagery from airborne satellites such as Landsat TM or SPOT can resolve cumulative damage caused by several years of bark beetle activity butcannot separate current mortalityfrom that of previous years. Use of these approaches has, to data, been largely experimental. Moreover, the probability of a satellite occurring over an area of forest damage when the damage signature is at its peak and skies are sufficiently cloud free to capture the damage is low.

Risk assessment is the process of evaluating natural hazards, such as disturbance agents, including insect outbreaks, the probability of a hazardous event occurring and the resulting consequences or losses if the event occurs. Hazard Hazard is defined as a potential event, such as fire, insect or disease outbreak, and the conditions causing it. Hazard-rating systems are used to determine infestation or infection potential and where the heaviest damage is expected based on certain biotic and abiotic conditions. The terms hazard and susceptibility are often used interchangeably.

RISK/HAZARD RATING

Risk

In the previous chapter the point was made that outbreaks are a function of host condition, climate and the dynamics of the insect. Risk/hazard rating examines underlying causes of outbreaks, especially host condition, and attempts to predict the probability of an outbreak occurring in the near future. Some of the variables considered in rating susceptibility to insect outbreaks include:

Risk is the likelihood or probability that an event will occur. In the context of insect risk assessment, for example, risk depends on both stand hazard and insect population densities. The terms risk and vulnerability are used interchangeably.

. . . .

species composition; age; stocking levels; elevation/latitude.

Risk/hazard rating schemes provide forest managers with an opportunity to set priorities and undertake management actions, such as thinning and timely harvesting, to create conditions less hospitable for development of outbreaks well in advance of their occurrence.

Susceptibility Susceptibility is the probability of an infestation occurring, based on stand characteristics (species composition, tree density, etc.). Vulnerability Vulnerability is the probability that a tree or stand will suffer damage. Note that susceptibility reflects the influence of forest or stand conditions on hazard, whereas vulnerability refers to the probability that damage will occur.

Definitions Much of the terminology associated with risk/hazard rating of forests for their susceptibility to pest damage has been used interchangeably and may be somewhat

Values Values are the losses that might be incurred should the damaging event occur. In a human context, social or

economic values might be lost or compromised during an outbreak. In an environmental or ecological context, wildlife habitat and other values could be either damaged or improved as a result of an outbreak. Risk/Hazard Rating Systems In North America, risk/hazard rating systems have been developed for both forest insects and diseases. These range from relatively simple approaches used to rate individual stands without the aid of a computer to regional, even country-wide, systems that integrate capabilities of mathematical models, geographic information systems and forest inventory databases. In the USA, many of these systems have been compiled in a database known as the Forest Insect and Pathogen Hazard Rating Systems Database by the USDA Forest Service, Forest Health Technology Enterprise Team and can be downloaded from the Internet (www.fs.fed.us/ foresthealth/technology/haz_rating_database.shtml). Mountain Pine Beetle in Lodgepole Pine, USA Amman et al. (1977) describe a simple procedure for estimating susceptibility of lodgepole pine forests in the Rocky Mountains of western USA to mountain pine beetle, Dendroctonus ponderosae. This is based on the following assumptions: .

.

.

Stands occurring at low elevations and lower latitudes will have a climate more suitable for overwintering brood survival and, therefore, are subject to greater losses during outbreaks. Trees of large diameters (> 20 cm) have a thicker phloem and will produce more brood than smaller diameter trees. Stands age 60 years and older tend to have a higher proportion of large-diameter trees.

This system uses tree diameter, stand age and location to predict risk or susceptibility to mountain pine beetle damage. Numerical ranks are assigned to each variable. Rankings for each variable are added to provide a stand rating from 3 (low) to 27 (high) (Fig. 4.10, Table 4.6). It rates stand hazard or susceptibility but does not consider the presence or absence of a mountain pine beetle population in or near the stands being evaluated. Therefore, it does not consider risk or vulnerability in the context of the presence or absence of the insect in question.

Elevation (thousand feet)

Monitoring forest insects, their damage and damage potential

12 11 10 9 8 7 6 5 4 3

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Lo Mo w (1) de 5 % 0% mo mor ta r ta lity lity

40

42 44 46 48 Latitude (degrees)

50

Fig. 4.10 Interaction of elevation and latitude on stand susceptibility to mountain pine beetle, Dendroctonus ponderosae (redrawn from Amman et al. 1977).

Mountain Pine Beetle in Lodgepole Pine, Canada A two-step mountain pine beetle risk index has been developed in Canada that considers both stand characteristics and beetle pressure. The first step involves computation of a susceptibility index, which considers several stand variables: (i) proportion of susceptible pine basal area; (ii) age; and (iii) location (latitude and elevation). Beetle pressure index is a function of the size and proximity of a mountain pine beetle population that may affect the stand being rated. The final risk index combines the susceptibility and beetle pressure indices and provides a continuous scale index for any lodgepole pine stand. Since the mountain pine beetle is dynamic, the authors caution that the beetle pressure index should be recalculated at regular intervals (Shore & Safranyik 1992). Multiple Pest Agents, Eastern Oregon, USA Schmitt & Powell (2005) describe a procedure that provides stand-level hazard ratings for six individual insect and disease pests and three pest groups in the Blue Mountains of eastern Oregon, USA. This system makes use of stand-attribute data that can be obtained from interpretation of aerial photographs and use of stand-level databases. Seven stand attributes are used in various combinations to rate susceptibility of stands to each of these pests or pest groups. The ratings are done on a form, without a computer (Fig. 4.11, Tables 4.7 & 4.8).

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Forest entomology: a global perspective Table 4.6 Stand hazard or susceptibility rating system for mountain pine beetle in lodgepole pine stands in the Rocky Mountains, USA (see also Fig. 4.10) Elevation/latitude

Age (years)

Average diameter breast height (dbh) (cm)

< 60 (1) 60–80 (2) > 80 (3)

< 18 (1) 18–20 (2) > 20 (3)

High (1) Medium (2) Low (3) Source: Amman et al. 1977.

Southern Pine Beetle Hazard Regional Rating System, Southeastern USA

Plain, Piedmont and Mountains. Each model was constructed at a 30-m resolution within a GIS environment using a set of forest parameter layers that include basal area, diameter, stand density index and a multicriteria modeling framework (USDA Forest Service 2006a).

A regional map of hazard of southern pine beetle, Dendroctonus frontalis, has been developed by USDA Forest Service for southeastern USA. This map is a compilation of eight mathematical models run across 15 ecological zones. Southern pine beetle hazard is defined as “degree of vulnerability of a stand to southern pine beetle, based on stand and physiographic attributes.” In several of the models, southern pine beetle presence was considered a precursor to hazard and separate susceptibility and vulnerability models were constructed. Models were developed from variations of existing southern pine beetle hazard rating systems developed for three major physiographic regions of southeastern USA: the Coastal

Pine Needle Gall Midge Hazard Rating System, South Korea A tree damage risk assessment system has been developed for pines damaged by the pine needle gall midge, Thecodiplosis japnensis, in South Korea. This system is based on the use of artificial neural networks. These are non-linear data modeling tools used to model complex

STAND SUSCEPTIBILITY RATING FORM (worksheet for one stand) Stand No._____________________________ T. _________ R. _________ Sec.__________ Observers ____________________________

Location _______________________________ Aerial Photos ___________________________ Date __________________________________

SCORES FOR RATING FACTORS INSECT OR DISEASE AGENT Defoliators Douglas-fir beetle Fir engraver Spruce beetle Bark beetles in ponderosa pine Mountain pine beetle in lodgepole pine Douglas-fir dwarf mistletoe Western larch dwarf mistletoe Root diseases

A _____ _____ _____ _____ _____ _____ _____ _____ _____

B _____ _____ _____ _____ _____ _____ _____ _____ _____

C _____ _____ _____ _____ _____ _____ _____ _____ _____

D _____ _____ _____ _____ _____ _____ _____

SUSCEPTIBILITY RATING* E

F

_____ _____ _____ _____ ____ _____ ____

TOTAL _____ _____ _____ _____ _____ _____ _____ _____ ______

LOW

≥5 ≤6 ≤7 ≥9 ≤6 ≤ 10 ≤5 ≤3 ≤4

MODERATE

HIGH

6-8 7-10 8-10 8-10 10-12 11-13 6-7 4-6 5-6

≥9 ≥ 11 ≥ 11 ≥ 11 ≥ 13 ≥ 14 ≥8 ≥7 ≥7

* This section shows how the total score (Total column) can be used to assign a categorical rating (low, moderate, high). Fig. 4.11 Worksheet used for stand-level hazard rating for multiple pest agents in the Pacific northwest region of the USA (Schmitt & Powell 2005).

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Table 4.7 Stand attributes and their corresponding database fields used to rate stand susceptibility to insect and disease pests in the Blue Mountains of eastern Oregon, USA Stand attribute

Database field

Existing tree species composition Tree species composition by layer Overstory tree size class Intra-stand variability Tree (stand) density Number of tree canopy layers Physiography/slope position

Forest cover type Canopy class composition Layer A size classes Clumpiness Tree canopy cover Canopy layering Slope curvature

Source: Schmitt & Powell, 2005.

relationships between inputs and outputs or to find patterns in data sets. They are adaptive systems that change their structure based on external or internal information that flows through the network during a “learning phase.” Variables used to predict probability of tree mortality or survival include: (i) diameter at breast height (dbh); (ii) tree height; and (iii) length, ratio, width, volume and density of tree crowns. Two approaches were developed. The first approach used an unsupervised neural network known as a self organizing map (SOM), which classifies trees according to their characteristics. The second approach used a multi-layer perceptron (MLP) to predict the probability of survival of affected trees following gall midge damage. The trained MLP model showed a 96% level of correct predictability (Park & Chung 2005).

National Pest Risk Map for the USA A National Insect and Disease Risk Map (NIDRM) was first developed for the USA in 2006. The threshold of mapping risk is the expectation that, without pest management, 25% or more of the standing live basal area of trees > 2.54 cm in diameter will die over the next 15 years due to insects and/or disease. The map represents the integration of 188 risk models that predict how certain tree species react to various mortality agents. The models, in turn, are the interactions of predicted agent behavior with known forest parameters. The most widely used forest parameters for the NIDRM are stand basal area (BA), stand density index (SDI), and tree diameter or its surrogate, quadratic mean diameter (QMD). Plot data were interpolated to create uniform “surfaces” that capture natural variations in forest parameters.

Table 4.8 Database fields used to rate susceptibility for each of six forest insects and diseases and three pest groups using aerial photographs in the Blue Mountains of eastern Oregon, USA Pest agent(s)

Database field Forest Canopy Layer A Clumpiness Tree Canopy Slope cover species size canopy layering curvature type composition classes cover

Defoliators Douglas-fir beetle Fir engraver Spruce beetle Bark beetles in ponderosa pine Mountain pine beetle (lodgepole pine) Douglas-fir dwarf mistletoe Western larch dwarf mistletoe Root diseases Source: Schmitt & Powell 2005.

A A A A A A A A A

B B B B B B B B B

– C C C C C C C C

– D D – D D – – –

C E E D E E – – –

D – – – F F D – –

– – – E – – – – –

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Forest entomology: a global perspective

Each of the 188 models, which predict the reaction of 42 mortality agents, including 13 recently introduced species, acting on 57 tree species, were documented using a spreadsheet template. The model results for each agent/host interaction are represented by a pair of maps showing: . .

likelihood (on a scale of 0–10) of an agent/host interaction resulting in mortality; percent contribution to total basal area loss attributed to that model.

The 2006 NIDRM displays projected risk of morality on a national scale at a 1-km spatial resolution and can be updated as new models and data become available. In its present form, NIDRM estimates that 23.5 million ha of the 303 million ha of forest land in the USA, including Alaska, are at risk from insect and disease (USDA Forest Service 2006b, Krist et al. 2007). An improved version of NIDRM is scheduled to be available by 2012. This map is expected to display areas at risk at a resolution of 240 m at the country-wide level and 30 m at a regional level.

Chapter 5

Forest insect management

Protection of forests from damaging insects and other pests is a complex and often costly process. Moreover, many activities related to forest pest management are controversial, especially when tactics such as aerial application of chemical insecticides or harvesting infested or dead trees are undertaken on public lands. This chapter provides a framework for activities designed to protect forests from damaging insects with minimal adverse impacts on other resources. This framework considers maintenance of forest health and vitality as the objective of forest pest management and integrated pest management (IPM) as a set of tools to achieve this objective.

HEALTHY FORESTS – THE OBJECTIVE The objective of managing forest pests is to help keep forests in a healthy, productive condition. The concept of a healthy forest evolved during the mid- to late 1980s. In economic terms, a “healthy” forest can be defined as “a forest in which pests and diseases remain at low levels and do not interfere with

management objectives.” In ecological terms, a healthy forest is considered “a fully functional ecosystem; one in which all of its parts can interact in a mutually beneficial way.” Healthy forests, therefore, are those that can meet economic, social and ecological functions now and in the future. The healthy forest concept directs forest managers to focus on the forest rather than its pests and takes into account the natural role of insects, fungi, fire and other so called “damaging agents” and their interactions in the dynamics of forest ecosystems. Pest outbreaks are viewed as a symptom of an unhealthy forest rather than as the problem. This directs forest managers and forest protection specialists to focus on underlying causes of insect or disease occurrence, such as overstocking, overmaturity, poor site/species matching, excessive fuels or presence of single species, even-aged forests with little diversity. Striving for healthy forests also involves anticipation of pest outbreaks based on historical records of their occurrence and knowledge of forest, climatic, soil and other factors that favor their occurrence. Ideally, this should allow time to implement management practices that will help make forests

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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inhospitable for buildup of damaging pests before outbreaks occur (Ciesla 1998). The notion of maintaining healthy forests may, initially, sound like an idyllic approach. This is especially true in cases where an introduced insect is causing damage in single species plantations in the absence of its natural enemies. Yet even some of the most damaging of exotic pests are known to respond to stand and site conditions or local climatic variations and there is some evidence that many exotic species focus their activities on unhealthy or stressed forests. An example is the wood wasp, Sirex noctilio, in exotic pine plantations in the southern hemisphere. In New Zealand, it was noted that stressed trees in unthinned plantations were the first to be attacked. Therefore, by its action, the insect was thinning overstocked pine plantations, albeit not to the same standards that a forester would use (McLean 1998). Similarly, when cypress aphid, Cinara cupressivora, appeared in eastern and southern Africa, the most severe damage occurred in plantations that were either overmature or had been established on lownutrient soils (Claude & Fanstin 1991, Obiri 1994, Ciesla et al. 1995).

INTEGRATED PEST MANAGEMENT The concept or philosophy of IPM as a “rational” approach to pest control was formalized during the 1960s as agricultural crop protection specialists became aware of adverse side effects of chemical pesticides including insect resistance to pesticides, occurrence of secondary pests, environmental damage and human health hazards. This led to a realization that alternative approaches, including cultural, biological and genetic tactics, used either alone or in combination, were also needed to provide long-term, effective protection against pests. IPM has many definitions. Between 1959 and 2000, some 67 definitions of either “integrated control” or IPM appeared in the worldwide literature (Bajwa & Kogan 2002). Smith et al. (1976) refer to IPM as “a process based on ecological principles and integrates multi-disciplinary methodologies in developing agroecosystem management strategies that are practical and effective and protect both public health and the environment.” FAO defines IPM as “the careful consideration of all available pest control techniques and subsequent integration of appropriate measures that discourage the development of pest populations and

keep pesticides and other interventions to levels that are economically justified and reduce or minimize risks to human health and the environment. IPM emphasizes the growth of a healthy crop with the least possible disruption to agro-ecosystems and encourages natural pest control mechanisms” (FAO n.d. (a)). Pimentel (1986) describes IPM as a “pest control method that includes judicious use of pesticide and non-chemical technologies – all of which are based on sound ecological principles.” IPM consists of two basic elements: a decision process and an action process. The decision process establishes the basis for pest management actions to be undertaken, including no action. The action process may consist of one or more ecologically, economically and socially acceptable tactics designed to reduce pest populations to non-damaging levels (Ciesla 1982).

THE DECISION PROCESS The forest owner or manager is the person who must ultimately decide for or against pest management and which, if any, pest management tactics to use. Selection of the most appropriate pest management approach is often the most time consuming and complex aspect of IPM. It requires careful consideration of the pest, its hosts, resource management objectives and ecological, economic and social consequences of the pest and of the available pest management tactics. Data on forest insect populations and their potential to cause damage in the immediate future are a critical input to the IPM decision process (see Chapter 4). The value of anticipated resource losses is estimated, as is the cost of treatment and its anticipated benefits. If treatment costs exceed losses, a rational decision may be to not treat and accept the loss. Other questions that should be addressed include: . .

Will natural controls take over within a short enough time so that artificial controls will be unnecessary? Will the ecological impacts of proposed treatments be so adverse that they outweigh the benefits of treatment?

The depth of analyses needed to support pest management decisions varies depending on the nature of the pest and its damage, the pest management tools being considered and the country and local laws and regulations regarding pest management. In some cases, surveys to determine if the insect has reached

Management of forest insect pests population levels that exceed a predefined economic threshold may be sufficient. In others, the decision process may be significantly more complex. The cost of proposed treatment is compared with the projected benefits in a cost–benefit analysis. Consideration is given to potential adverse impacts of treatments on non-target organisms with special consideration to impacts on rare or threatened species. In many countries, if a pest management action is considered on public lands, such as national or state forests, a formal environmental impact analysis with public review and comment may be required before any action is taken.

THE ACTION PROCESS Pest management actions consist of strategies and tactics. Strategies are the overall approaches to pest management and tactics are the tools. In IPM, there are three basic strategies: . . .

no action; prevention; direct control or suppression.

Depending on conditions, elements of one or more of these strategies may be part of an IPM project or system.

No Action The decision processes described in the preceding section may lead forest owners or resource managers to take no action either on all, or a portion of, the area affected by a forest pest and simply accept the losses. Selection of this option may be driven by one or more ecological, policy, economic or social considerations. Ecological Considerations Ecological factors that may drive a “no action” pest management decision include: .

.

Insect population monitoring may indicate that population levels of the target insect are declining and/or natural factors such as parasites and predators will exert enough pressure on the next generation that damage is expected to be minimal. An outbreak has become so massive that attempts at pest management are futile.

.

.

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Limited access, steep terrain or poor weather conditions constrain implementation of available pest management tactics. Implementation of pest management might adversely affect non-target species that may occur in all, or a portion of, the affected area.

Policy Considerations Land management objectives and policies may constrain use of certain, or all, pest management tactics. National parks and reserves or other lands with special classifications (e.g. designated wilderness in the USA) are usually managed on a “let nature take its course” or “no intervention” policy. A disturbance, such as an insect outbreak would, therefore, be considered part of the dynamics of the ecosystem and viewed as a means of replacement of old, mature forests with young, vigorous forests. Economic Considerations One of the key elements of the pest management decision process is to weigh the costs of treatment vs anticipated benefits (benefit/cost analyses). Benefits of pest management in terms of reduced rates of tree mortality, avoidance of growth loss or loss of wood quality are relatively easy to quantify on lands where timber production is one of the resource management objectives. Benefits of pest management on scenic and recreational values, improved wildlife habitat or water yields from forested watersheds, on the other hand, are more difficult to quantify. If benefit/cost analysis indicates a marginal or negative benefit of a proposed pest management action, the decision to not treat a pest problem is valid, at least in economic terms. Social Considerations Many pest management tactics are controversial. Use of chemical pesticides, with their potential adverse effects, is almost always a subject of controversy. Even proposed use of relatively host specific direct control measures, such as the bacterial insecticide, Bacillus thuringiensis, against a forest defoliator can be controversial because non-target Lepidoptera are also affected. In some countries, there is a strong public sentiment against timber harvesting and road construction on public forest lands. Therefore, proposals to salvage trees

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Forest entomology: a global perspective

killed by insects may be met with strong opposition. Public opposition to use of some, or all, available pest management tactics against a forest pest may cause public land forest resource managers to decide to take no action against a forest pest.

pest populations. The broad classes of IPM tactics include: . . . .

Risks Associated with No Action There are certain risks associated with a decision to take no action against a forest pest on public lands. National parks and reserves, for example, receive large numbers of visitors. Presence of dead trees caused by insect outbreaks, especially in heavily used areas such as campgrounds, picnic areas, vistas or hiking trails, are a safety hazard, especially as stems decay and become subject to windthrow. Another risk is liability associated with spread or conceived spread of insects and resultant damage into neighboring forest lands where management objectives may be quite different. Presence of damaging populations of indigenous insects in a national park or reserve may be looked upon as a “natural” event and, therefore, not treated. However, introduction and establishment of exotic species may require special consideration. These could have devastating impacts on the very values under protection by the land use classification. An example is the introduction of the hemlock woolly adelgid, Adelges tsugae, into the Great Smoky Mountains National Park in the southern Appalachian Mountains of the USA. This insect is causing extensive mortality of eastern hemlock, Tsuga canadensis, a key component of riparian forest ecosystems, and has adversely affected the park’s biodiversity. As a result, officials are using a combination of chemical and biological control tactics in an attempt to reduce losses (USDI National Park Service n.d.).

. . .

regulatory; cultural; genetic; mechanical; biological control; semiochemicals; chemical insecticides.

Several of these can be used either to prevent or suppress damage. For example, cultural tactics, such as thinning, may create conditions inhospitable for development of pest populations. Salvage logging, on the other hand, is oriented towards the removal of trees that are infested or have been killed by insects to reduce insect populations and/or recover some of the losses incurred (Fig. 5.1). Similarly, semiochemicals can be deployed to prevent attacks through use of antiaggregating pheromones or to reduce insect populations by mass trapping or mating disruption. Details of tactics available in forest insect management are described in the following sections.

PEST MANAGEMENT TACTICS Regulatory Tactics The objective of regulatory tactics is to prevent introduction of exotic pests and to prevent or reduce their spread in areas where they have become established. Three areas of activity fall under the general heading of regulatory tactics: . . .

inspections of imported goods; conduct of pest risk analyses; establishment of quarantine zones.

Prevention and Suppression Pest management actions, designed to reduce losses incurred by forest insects, involve two strategies: prevention or suppression. Prevention is designed to reduce the probability of occurrence of damaging insects by creating environmental conditions inhospitable for their buildup. Suppression or direct control consists of actions directed against the insect to reduce population levels and subsequent losses to the resource. Tactics are the tools used to either prevent damage or reduce populations and subsequent losses from existing

Inspections of Imported Goods Inspections of agricultural commodities and forest products at ports of entry are designed to intercept and destroy exotic species that may be associated with these products. Considering the high risk of introduction of wood infesting insects, it has become necessary to not only inspect the commodity itself but also wooden containers that contain the product, dunnage and pallets, especially wood containing bark strips, for presence of potentially damaging wood borers and bark beetles.

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Fig. 5.1 Salvage logging of trees killed by southern pine beetle, Dendroctonus frontalis (Honduras, photo by R.F. Billings, Texas Forest Service).

Pest Risk Analyses

International and Regional Standards

Pest risk analyses are undertaken when new agreements are negotiated between international trading partners. These analyses identify key species with a potential for becoming pests should they be accidentally introduced, their potential pathways of entry and how to recognize the insect and infested material. During the late 1990s, teams of USDA APHIS and Forest Service experts conducted several pest risk analyses for importation of unprocessed logs and chips from Siberia and the Russian Far East, Mexico, New Zealand and South America (USDA Forest Service 1991, 1993, 1998, 2001).

Prevention of introductions of exotic pests involves a high level of international cooperation. At the global level, the International Plant Protection Convention (IPPC) is an international treaty that establishes standards designed to prevent introduction, establishment and spread of plant and plant product pests and to promote appropriate measures for their control. The IPPC Secretariat coordinates the activities of the Convention and is housed at the headquarters of FAO, Rome, Italy (www.ippc.int). IPPC activities are oriented largely toward agricultural crop pests but forest pests are also addressed. For example, International Standards for Phytosanitary Measures No. 15 (ISPM 15) establishes standards for wood packing materials used in international trade to prevent movement of cambium and wood boring insects. This standard applies to wood materials of a thickness greater than 6 mm, used to ship products between countries. It affects all wood packaging material (pallets, crates and dunnage) and requires that they be treated with heat or fumigated with methyl bromide and

Quarantines Quarantines are designed to prevent spread of an introduced pest once it has become established. In the case of forest insects, quarantine procedures focus on restricting movement of potentially infested fuel wood, other wood or non-wood products and nursery stock from infested to uninfested areas.

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Forest entomology: a global perspective

branded with a seal of compliance. Products exempt from ISPM-15 are made from alternative materials such as paper, plastic or wood panels. A 2009 revision of ISPM-15 requires that wood used to manufacture ISPM-15 compliant wood packaging must be made from debarked wood not to be confused with “bark free wood.” This was done to incorporate bark restriction regulations proposed by the EU. Several regional plant protection organizations operate under the umbrella of the IPPC. The European Plant Protection Organization (EPPO) is a regional intergovernmental organization responsible for cooperation in plant protection in the European and Mediterranean regions. Founded in 1951, it has grown from 15 to 50 member countries and now includes nearly every country in these regions (http://www.eppo.org/ ABOUT_EPPO/about_eppo.htm). EPPO’s objectives are to: .

.

. .

develop an international strategy against introduction and spread of pests that damage cultivated and wild plants, in natural and agricultural ecosystems (including invasive alien plants); encourage harmonization of phytosanitary regulations and all other areas of official plant protection action; promote use of modern, safe and effective pest control methods; provide a documentation service on plant protection.

Cultural Tactics Cultural tactics designed to reduce the probability of pest buildup include matching tree species selected for planting to suitable growing sites, controlling stocking through intermediate harvests (thinning) to maintain tree vigor and timely harvesting of forests when they reach maturity. Thinning and Timely Harvesting In western North America, mountain pine beetle, Dendroctonus ponderosae, outbreaks in ponderosa pine, Pinus ponderosae, forests are favored by high stand densities. Stands can be protected through regular thinning to basal areas below 23 m2/ha (Sartwell & Dolph 1976). Lodgepole pine forests, P. contorta, which also suffer devastating outbreaks of this insect, are most susceptible when they exceed age 60 years and a high proportion of trees have a diameter at breast height

(dbh) of > 20 cm. Since lodgepole pine forests typically occur as pure, even-age stands over large areas and readily regenerate following clearcuts, a practice of creating a mosaic of age classes over the landscape, with a high proportion of young stands, less than age 60 years, can reduce the overall impact of an outbreak (Amman et al. 1977, 1990). Factors that may limit this practice include lack of access, limited markets for lodgepole pine products and opposition to timber harvesting and road construction on public lands by environmental groups. In countries such as Australia, Brazil and New Zealand, where the woodwasp, Sirex noctilio, has been introduced and become established, thinning of pine plantations to increase vigor of residual trees will reduce infestation levels and subsequent losses (Haugen et al. 1990, Iede & Ciesla 1993, Iede et al. 1998).

Species Selection and Biodiversity One of the important silvicultural tactics to reduce impact of damaging insects is to maintain as high a level of biodiversity as possible. Unfortunately, forest plantation programs in many countries rely on relatively few tree species, often exotics, which happen to be well adapted to local climatic and soil conditions, are fast growing and produce a variety of useful wood products. In Brazil, for example, there are 6 million ha of forest plantations: 4 million ha of eucalypts and 2 million ha of pines. In the country’s three southernmost states there are 1.2 million ha of pine plantations, of which roughly 80% are Pinus taeda. In Chile, as of 2005, there were 2.078 million ha of forest plantations, of which 68% were of Pinus radiata. An additional 24% of the country’s forest plantation estate is composed of eucalypts, primarily Eucalyptus globulus (INFOR 2005). In Kenya during the 1990s, 46% of forest plantations were composed of Cupressus lusitanica. Reliance on one or two forest plantation species puts them at high risk should either a native species adapt to the large area of monoculture or should a damaging invasive pest become introduced and established. Introduction of cypress aphid, Cinara cupressivora, into Kenya and other countries of eastern and southern Africa caused extensive tree mortality in Cupressus lusitanica plantations. This forced forest departments to seek alternative species for their plantation programs. Maintenance of a mix of species or a mosaic of age classes where the natural forest is composed of just a

Management of forest insect pests few species, as is the case over much of the Rocky Mountain region of North America, helps increase biodiversity and reduces the overall volume of suitable host material available for development of outbreaks.

Mechanical Tactics Mechanical tactics include removal and destruction of all, or a portion of, infested trees to destroy the pest. Low-level populations of tent making defoliating caterpillars, such as Malacosoma spp. throughout the northern hemisphere and the pine processionary caterpillars, Thaumetopoea pityocampa and Th. wilkinsonii of Mediterranean Europe and northern Africa, can be effectively controlled by pruning and destroying branches containing tents and larval colonies. Hand pruning is also a viable tactic for control of gall insects on small ornamental trees and shrubs. A technique known as “cut and leave” is an effective treatment for southern pine beetle, Dendroctonus frontalis, in areas where opportunities to salvage log infested trees are limited. This consists of cutting all trees with fresh attacks or containing beetle broods and leaving them on site. Beetle survival in cut trees is reduced and, by disrupting pheromone production, spot size growth is prevented. This has been used effectively in outbreak areas in southeastern USA and Central America (Fig. 5.2, Billings et al. 2004). Solar radiation is a technique used to increase brood mortality of bark beetles. Trees containing brood are cut

Fig. 5.2 Strip of pines cut as part of a “cut and leave” technique to limit spot growth of southern pine beetle, Dendroctonus frontalis (Honduras, photo by R.F. Billings, Texas Forest Service).

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and bucked into workable lengths, placed in an open area with direct exposure to sunlight and wrapped in plastic sheeting. This produces temperatures high enough to cause brood mortality and has been used successfully to reduce mountain pine beetle broods in the Rocky Mountain region of the USA (Negrón et al. 2001). When pine woolly aphid, Pineus boerneri, was first discovered in Kenya, the initial response was to cut and destroy infested trees. However, this insect is easily carried on air currents and it was soon realized that infestation had spread far beyond the designated areas where trees were being destroyed (Owour 1991).

Genetic Tactics Genetic tactics make use of varieties of host plants that are either more tolerant to damage or less palatable to the pest. Identification and testing of varieties of Leucaena leucocephala, a fast-growing tree widely planted in the tropics as an agroforestry species, for resistance or tolerance to the introduced leucaena psyllid, Heteropsylla cubana, was a major line of investigation following this insect’s introduction into the Asia-Pacific region (Banpot Napompeth 1994).

Biological Control Biological control involves use of natural enemies (parasitoids, predators or diseases) of an insect to help

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keep its numbers in check. Several approaches to biological control have been used: . . .

classic biological control; augmentative biological control; microbial insecticides.

Classic Biological Control Importation of natural enemies to help control a damaging pest, usually an introduced species, is known as classic biological control (Fig. 5.3). This tactic has been widely used against a wide range of forest insects and there are many examples of its successful application. Release of a parasitic wasp, Pauesia bicolor, resulted in control of a giant conifer aphid, Cinara cronartii, which

Fig. 5.3 A screen cage is placed over a branch of a cypress, Cupressus lusitanica, for release of parasitoids as part of a classic biological control program for cypress aphid, Cinara cupressivora (western Kenya).

is native to North American and accidentally introduced into pine plantations in South Africa (Kfir et al. 1985, Mills 1990). In Chile, state-of-the-art mass rearing facilities have been established by forest industry for production of Orgilis obscurator, a parasitoid of the European pine shoot moth, Rhyacionia buoliana, which has become an important pest of that country’s Pinus radiata plantations. One such laboratory, Controladora de Plagas Forestales, was established in 1992. In 1996, this facility produced 1.2 million parasitized larvae and 22,558 adult female parasitoids for field release (Controladora de Plagas Forestales 1997). Attempts at classic biological control of forest pests in Europe, where, to date, there have been relatively few introductions of exotic species, have involved the movement of indigenous natural enemies from one area to another. In the UK, the natural enemy complex of many forest insects is considerably less diverse than in continental Europe. This suggests that there are opportunities to move natural enemies from the central portions of their distributions to outlying areas. When natural enemies are absent in certain areas, it is important to determine if this due to an unsuitable environment or simply separation of the pest and its enemy in certain parts of their natural ranges. In southern Sweden, infestations of adelgids occurred on plantations of Abies alba, a tree exotic to northern Europe. Transfer of a predaceous ladybird beetle, Scymnus impexus, from Germany led to its establishment in Sweden (Speight & Wainhouse 1989). Classic biological control is not necessarily restricted to exotic pests. In Colombia, for example, indigenous foliage feeding Geometridae that adapted to exotic plantations were successfully controlled via the introduction and release of a North American egg parasitoid, Telenomus alsophilae (Bustliio & Drooz 1977). Several factors should be considered before using classic biological control as a pest management tactic. One is the possibility that the introduced natural enemy might also attack innocuous or beneficial insects in the ecosystem. It is, therefore, necessary to thoroughly evaluate candidate species prior to their release to determine their host specificity. Another concern is the hazard of accidentally introducing hyperparasites, natural enemies of the biological control agents, which might eventually affect the agent’s efficacy. For example, colonies of the leucaena psyllid predator, Olla v-nigrum, released on the Indian Ocean island of Reunion, were subsequently discovered infested by three species of hyperparasites (Quilici et al. 1995).

Management of forest insect pests Augmentative Biological Control Tactics to increase abundance and efficiency of natural enemies already in place are known as augmentative biological control and involves two approaches: . .

mass rearing and inundative releases of parasitoids and predators; altering site conditions to create a more favorable habitat for natural enemies.

In China, mass rearing and release of an indigenous egg parasitoid, Trichogramma dendrolimi, for control of the defoliating caterpillar, Dendrolimus punctatus, has been used with reasonable success (Yan & Liu 1992). Birds are known predators of damaging forest insects, although their effectiveness is not well understood. In Europe, augmentation of insectivorous birds in forests through establishment of nesting boxes has been practiced for about 80 years. This tactic is based on the assumption that nesting site availability is limited. While this may be true for cavity nesting birds in areas where dead trees are removed, there is no certainty that an increase in bird density increases mortality of important pest species. When food supply is a limiting factor in bird population dynamics, an attempt to increase bird density by installation of nesting boxes during times when populations of damaging insects are at low levels may not be effective (Speight & Wainhouse 1989). In China and Vietnam, it is believed that elimination of livestock grazing in pine plantations will encourage growth of flowering plants, which serve as a source of nectar for parasitic wasps of the pine caterpillar, Dendrolimus punctatus, thus increasing parasitoid numbers. While there is some logic to this hypothesis, no data are available to substantiate it (Author’s observation). Microbials Use of agents that cause disease in insects (e.g. fungi, viruses, bacteria and nematodes) as a control agent is known as microbial control. One bacterial insecticide is available commercially for insect control and several viruses have been registered by government agencies responsible for pesticide regulation against forest insects.

Bacteria Bacillus thuringiensis (Bt) is a naturally occurring soil bacterium that causes disease in insects. Bt is considered ideal for pest management because of

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its relative host specificity and its lack of toxicity to humans or the natural enemies of many insect pests. There are several subspecies of Bt, each with a specific toxicity to a particular group of insects. B.t. israelensis (Bti) is effective against mosquitoes, blackflies and some fungus gnats. B.t. kurstaki (Btk) controls lepidopterous insects, including forest defoliators such as tent caterpillars and spruce budworms. B.t. san diego/tenebrionis, is effective against certain species of beetles (Martin 1994, Cranshaw 2009). Bt produces proteins in the form of a toxic crystal, known as the delta-endotoxin, which reacts with the cells of the gut lining of susceptible insects. They paralyze the digestive system, and infected insects stop feeding, usually within hours. Bt affected insects generally die from starvation, which can take several days. B.t. kurstaki is currently the most widely used microbial insecticide in the world and has been applied over thousands of hectares of forested lands in Asia, Europe and North America for control of forest defoliators (Speight & Wainhouse 1989, Reardon et al. 1994, Cranshaw 2009, Fig. 5.4, Table 5.1).

Viruses Epizootics caused by viruses are often seen in insect populations. Several groups of viruses infect and kill insects. Baculoviruses replicate in the nuclei of insect cells. Nucleopolyhedrosis viruses (NPVs) are by far the most common type of baculovirus and have a characteristic structure. Individual virus particles, known as virons, are contained within a protein matrix known as a polyhedron. Granulosis viruses (GVs) are less common and found only in the larval stages of moths and butterflies. With GVs, virons are individually encapsulated and not grouped together in polyhedra. Cytoplasmic viruses (CPVs) are similar to NPVs except in their site of action and that they belong to a different group of viruses, the Retroviridae. This group also contains viruses capable of infecting vertebrates and plants. To cause disease, virus particles must be ingested by larvae. After ingestion, the virus multiplies and death occurs quickly (Speight & Wainhouse 1989). Larvae of some forest caterpillars and sawflies are highly susceptible to baculovirus diseases (Fig. 5.5, Table 5.2). They are considered promising organisms for insect control because they can spread through a population quickly and persist in the environment for long periods. Viruses also have the advantage of being host specific. Therefore their use in the forest should not affect other organisms. Host specificity also works

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Forest entomology: a global perspective

Fig. 5.4 A spray aircraft applies the microbial insecticide, Bacillus thuringiensis, over a forested area infested by western spruce budworm, Choristoneura occidentalis (Mt Hood National Forest, Oregon, USA).

against these materials because private industry is reluctant to invest in their development and registration because they only offer limited markets. Several insect viruses have been produced and registered for use against forest defoliators, either by ground

or aerial application. Insect viruses must be mass produced by rearing large numbers of host insects in the laboratory, infecting them with virus and recovering virus particles from the cadavers. In the USA, the first two NPVs registered by the US Environmental

Table 5.1 Examples of forest defoliators treated with Bacillus thuringiensis kurstaki in Asia, Europe and North America. Scientific name

Common name

Where treated

Choristoneura fumiferana Choristoneura occidentalis Dendrolimus punctatus Dendrolimus sibiricus Lymantria dispar Lymantria monacha Malacosoma disstria Orgyia pseudotsugata Thaumetopoea pityocampa

Spruce budworm Western spruce budworm Pine caterpillar Siberian silk moth Gypsy moth Nun moth Forest tent caterpillar Douglas-fir tussock moth Pine processionary caterpillar

Thaumetopoea wilkinsonii

Eastern pine processionary caterpillar

Canada, USA Canada, USA China, Vietnam Mongolia, Russia Asia, Europe, North America Lithuania, Poland USA Canada, USA Mediterranean regions of Europe, North Africa and the Near East Cyprus

Sources: Speight & Wainhouse, 1989 and the Author’s experience.

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Enough refined virus was produced at this facility to treat 203,305 ha of forest at the recommended dosage of 2.681  108 activity units (AU)/ha. In 2000, some 12,145 ha of forests in Oregon and Washington were treated with this material. An additional 4048 ha were treated the following year. The virus production facility was closed in late 1992. However, detailed plans are available to produce additional virus if needed (Scott 2001).

Fig. 5.5 Larva of gypsy moth, Lymantria dispar, killed by a virus (South Mountain, Pennsylvania, USA).

Protection Agency (EPA) were the Douglas-fir tussock moth virus in 1976 (TM Biocontrol-1) and the gypsy moth virus in 1978 (Gypchek). Because of their host specificity and relative infrequency of use, especially the Douglas-fir tussock moth NPV, registration of these products was done by two USDA agencies: Forest Service and Agricultural Research Service (ARS) instead of a private company. As of October 2002, the US EPA registered five NPVs and GVs as pesticides and 10 pesticide products containing these active ingredients. Between 1980 and 1992, the Douglas-fir tussock moth NPV was produced in laboratory facilities in Corvalis, Oregon, USA by the USDA Forest Service.

Fungi All major groups of fungi contain species lethal to insects and most insect orders are susceptible. Fungi infect insects through the digestive tract and the integument. Larvae, pupae and adults are subject to attack. Weather plays an important role in determining effectiveness of fungi as biocontrol agents. Fungi need moisture and high humidity to germinate. Frequent rainfall during May and June, for example, can create conditions favorable for development of fungi that attack defoliating caterpillars. Fungi also thrive in warm temperatures. One of the most commonly occurring insect fungi is Beauvaria bassiana, sometimes referred to as the white muscadine fungus. This fungus is often seen as a mass of white mycelium covering the body of a dead insect (see Fig. 2.4). B. bassiana has been propagated in the laboratory for control of pine caterpillar, Dendrolimus punctatus, in China and Vietnam (Fig. 5.6) and applied in a unique manner. Fungus spores are inserted into large firecrackers from which a portion of the gunpowder has been removed. The firecracker is lit, tossed into the crowns of infested pines and the spores are released as the firecracker explodes. Unfortunately, data on the effectiveness of this method are lacking (Author’s observation). The fungus Entomophaga maimaiga causes a disease of gypsy moth, Lymantria dispar, in Japan. This fungus was first released in the USA near Boston, Massachusetts in 1910, as part of a program to introduce natural enemies of gypsy moth in areas where this insect was introduced. Scientists could find no evidence that the fungus had become established and the project was abandoned several years later. The fungus appeared unexpectedly in several northeastern states in 1989 and caused high mortality of many gypsy moth populations. Although scientists have several theories, the strange reappearance of this fungus is still a mystery. E. maimaiga passes the winter as a tough, thick-walled “resting spore” in the soil and on tree bark. In May or

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Table 5.2 Examples of baculoviruses known to affect forest defoliators. Insect species

Common name

Virus type

Host trees affected

Country(s)

Bupalis piniarius Gilpinia hercyniae Lymantria dispar

Pine looper Spruce sawfly Gypsy moth

NPV NPV NPV

Netherlands North America Europe, North America

Lymantria monacha Neodiprion sertifer Orgyia pseudotsugata

Nun moth European pine sawfly Douglas-fir tussock moth

NPV, CPV NPV NPV NPV

Pines Spruce Oaks and other broadleaf species Fir

Thaumetopoea pityocampa Zeiraphera diniana

Pine processionary caterpillar Larch bud moth

Lymantria fumida

CPV

Pine, spruce Pine Douglas-fir, true fir Pines

GV

Larch, pines

Japan Northern Europe Canada, Europe, USA Western North America Mediterranean regions of Europe, North Africa and the Near East Switzerland

CPV, cytoplasmic virus; GV, granulosis virus; NPV, nucleopolyhedrosis virus. Source: Bird 1953, Podgewaite et al. 1984, Speight & Wainhouse 1989 and the Author’s experience.

June, resting spores germinate and produce sticky spores at the end of a stalk that grows just above the soil surface (Reardon & Hajek 1997).

Nematodes Nematodes are microscopic, unsegmented worms with cylindrical bodies and occupy several ecological niches. Many are soil inhabiting. Others, such as the pinewood nematode, Bursaphelenchus xylophilus, are plant pests. Some are parasites of mammals, including humans, and others parasitize insects. The nematode, Deladenus siricidicola, is a parasite of the wood wasp, Sirex noctilio. It usually feeds on the fungus Amylostereum areolatum, which is introduced into trees when the female wood wasps deposit eggs. It also has the capacity to infect ovaries of the female wood wasps and render them incapable of reproducing (Zondag 1962). Infection levels can reach 100% and lead to a population collapse. Methods for mass rearing the nematode and inoculating infested trees have been developed and used with varying degrees of success in all areas where S. noctilio has become established (Haugen et al. 1990, Fig. 5.7).

Semiochemicals Use of semiochemicals (pheromone, kairomones, etc.) as a tool for monitoring forest insects was described in Chapter 4. These products are also effective insect pest

management tools. Approaches that involve semiochemicals for insect management include mass trapping, anti-aggregation and mating disruption. Mass Trapping Mass trapping of flying insects using pheromones to reduce populations has been used against both bark and ambrosia beetles. A major outbreak of the bark beetle, Ips typographus, occurred in Norway and Sweden between 1971 and 1981. In Norway, some 5 million m3 of timber were attacked and killed over about 140,000 km2. The outbreak was triggered by a massive windstorm in 1960 that affected about 4 million trees, followed by a drought from 1974 to 1976. Until 1979, the primary method of control was use of trap trees that consisted of mature fresh trees, felled in spring and left in the forest during the beetle’s flight period. When trees were colonized, they were removed. In 1979, pheromone traps, baited with components of the beetle’s aggregation pheromone, were used to mass trap beetles. From 1979 to 1980 over 600,000 pheromone baited stovepipe traps were deployed at a density of 20–30 traps/ha in the most heavily infested areas. During 1979, 1980 and 1981, several billion beetles were trapped. In 1980 alone, 4,500,000,000 beetles were trapped. After 1982, following a significant decline in beetle activity, the number of traps deployed was reduced to 100,000. The mass trapping program was part of an integrated

Management of forest insect pests

Fig. 5.6 A technician prepares a culture of the insectivorous fungus Beauvaria bassiana to be masse reared and used for control of pine caterpillar, Dendrolimus punctatus (Vinh, Vietnam).

program that also included harvesting of mature spruce and prohibition of storage of fresh cut logs with bark in forested areas. A similar control effort was also conducted in Sweden against I. typographus using pheromone baited traps during the same time period (Baake 1989, 1991, Wainhouse 2005). In western Canada, the ambrosia beetle, Trypodendron lineatum, can build up to large numbers in areas where freshly harvested logs are stored and sorted prior to distribution to sawmills. Breeding attacks and associated blue stain fungi cause significant loss in lumber quality. Mass trapping, using Lindgren funnel traps baited with the pheromone lineatum, was used on an operational trial to reduce beetle numbers and subsequent damage. During a sustained period of trapping

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Fig. 5.7 A suspension of the parasitic nematode Deladenus siricidicola is injected into a loblolly pine, Pinus taeda, infested with the wood wasp Sirex noctilio. The nematode attacks female wood wasps and renders them sterile (Santa Catarina State, Brazil).

over several years, millions of beetles were trapped and degradation of stored logs declined significantly. The decline in loss of wood quality was attributed to a combination of the mass trapping and improved management of both the timber supply and log storage at the sorting area (Lindgren & Fraser 1994, Wainhouse 2005). Anti-Aggregation Several pheromones produced by bark beetles have an anti-aggregating function and are emitted when trees have been saturated by colonizing beetles. They include the pheromone 3-methyl-2-cyclohexanone (MCH)

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produced by two North American bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, and the spruce beetle, D. rufipennis. Other anti-aggregating pheromones include ipsenol and verbenone. These have been used with varying degrees of success to repel bark beetle attacks. Application of anti-aggregating pheromones can be done in two ways, either by aerial application of microencapsulated pheromones or by hand distributed dispensers. MCH has been used to repel attacks by both Douglasfir beetle and spruce beetle in windthrown trees using microencapsulated pheromones (Furniss et al. 1977). A technique that involves use of bubble caps, which contain a slow-release formulation of anti-aggregating pheromones such as MCH or verbenone, has been developed. Bubble caps can be stapled to susceptible trees that are either standing or down. In the case of the Eurasian Ips typographus, two pheromone components have been shown to have an anti-aggregating effect: ipsenol and verbenone. In Norway, trees baited with formulations of both pheromones have effectively

repelled attacks by this beetle (Baake 1991) and in Germany, a slow-release formulation of verbenone, known as a verbenone strip, wrapped around spruce trees appears to protect them from attack (Vite & Baader 1990). Verbenone is also known to function as an anti-attractant for several North American pine infesting species of Dendroctonus, including the mountain pine beetle, D. ponderosae. A commercial verbenone pouch, which can be attached to susceptible trees, has been developed to repel attacks (Fig. 5.8). Unfortunately, results have to date been erratic (Negrón et al. 2006). A problem associated with use of anti-aggregating pheromones for bark beetle management is that, in the absence of supplemental aggregating lures, beetles are not trapped and killed. Therefore, treatments may protect the trees that contain the anti-aggregating pheromone but they may divert attacks into surrounding trees and stands, with little or no reduction in the overall numbers of trees attacked. This is most likely to occur during outbreaks when beetle population densities are high because there is an abundant source of

Fig. 5.8 Pouches containing the anti-aggregating pheromone verbenone are stapled to the bark of a pine to repel mountain pine beetle, Dendroctonus ponderosae (photo by Sheryl Costello, USDA Forest Service).

Management of forest insect pests aggregation pheromones due to pioneer beetles that initiate attacks (Wainhouse 2005).

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HerconÒ flakes at 10–20 g of active ingredient/ha, reduced shoot damage to pines caused by larvae by 76–88% (Sower et al. 1982).

Mating Disruption Sex attractant pheromones are an effective tactic for disruption of reproductive cycles of damaging insects. The objective is to reduce or eliminate populations of the target insect from a given area. This tactic is most effective when the target species are at low to moderate population densities. As is the case with antiaggregating pheromones, application of pheromones for mating disruption may involve either microencapsulated pheromones or hand distributed dispensers. One of the more successful uses of mating disruption as a pest management tactic for a forest insect is the Slowthe-Spread of the Gypsy Moth Project (STS) in the USA. This is a coordinated effort by USDA and 10 states to slow the rate of spread of gypsy moth, Lymantria dispar, into uninfested areas. A grid of pheromone traps is used to detect and delimit gypsy moth populations along the leading edge of the known area of infestation. If new infestations are detected along this edge, they are treated with either insecticides or by mating disruption. Initially, most areas were treated using Bacillus thuringiensis. By 2000, mating disruption using the gypsy moth sex pheromone, disparlure, became the primary treatment tactic. Application rates of 74 g of active ingredient/ha were used exclusively until 2000. However, field tests indicated high levels of efficacy at lower doses. Therefore, mating disruptants were applied at 37 g/ha starting in 2001, and at both 14.8 g/ha and 7 g/ha starting in 2002. Based on criteria developed for use in the STS decision support system, an analysis of treatment results for the years 1993–2001 (prior to the use of the 14.8 g/ha rate) showed that the success rate for blocks treated with mating disruption was greater than for blocks treated with Bt. The success rate of all treatments combined from 1994 to 2003 ranged from 85% to 95%. Using the same criteria, treatment success ranged from 88% for Bt to 100% for mating disruption at 14.8 g/ha. Treatment success was 93% and 95% for mating disruption at 37 g/ha and 74 g/ha, respectively. A plastic laminated flake formulation of the gypsy moth pheromone, disparlure, for mating disruption (HerconÒ Disrupt II) is registered by the US EPA for either ground or aerial application (Plimmer et al 1982, Thorpe 2005). In western USA, a synthetic version of the pheromone of a pine shoot borer, Eucosma sonomana, applied by air for mating disruption, as ConRelÒ fibers or

Chemical Insecticides Chemicals, applied either from the ground or by lowflying aircraft, are still widely used for insect control. Their use is considered an integral part of IPM. However, under the philosophy of IPM, chemical insecticides are usually considered the tactic of last resort, to be used when no other effective pest management alternatives are available. Moreover, chemical insecticides should be used sparingly, applied at reduced intervals and with as high a degree of precision as possible. Because of the relatively high cost of chemical control and the low rates of return from forest products, forest managers have not relied as heavily on chemical pesticides as their agricultural counterparts. However, there have been cases, especially in North America, where millions of hectares of forest were treated with aerial applications of chemicals for control of defoliators such as spruce budworm, Choristoneura fumiferana, and gypsy moth, Lymantria dispar. Today, microbial agents such as Bacillus thuringiensis, or insect growth regulators (IGRs) such as diflubenzuron, have largely replaced these materials. Today, most uses of chemicals in forest insect management are confined to special use areas such as tree nurseries, seed orchards or high-use recreation sites. Types of Chemical Insecticides Chemical insecticides can be classified by their chemical nature and/or mode of action. The classification given in the following sections is based largely on their chemical nature. Inorganic Insecticides Inorganic1 compounds of arsenic, such as lead arsenate, have been used for control of many insects. While highly effective, they 1 The terms “organic” and “inorganic” used in this discussion refer to chemical compounds that either contain carbon (organic compounds) or lack carbon (inorganic compounds). The alternative definition of “organic” is agricultural crops produced without use of chemical pesticides, chemical fertilizers, bioengineering or ionizing radiation.

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are also toxic to other organisms and are known to persist in the environment for long periods. Use of most of these materials has been discontinued.

Botanicals Botanical insecticides are derived from plant extracts. One of the most widely used botanical insecticides is pyrethrum or pyrethrins. They are extracts of a chrysanthemum-like flower, Dendranthema (¼ Chrysanthemum) cinearariaefolium. The insecticidal nature of pyrethrins has been known for a long time. The Chinese traded pyrethrum daisies in dried form along the Silk Road and into Europe where it became popular as a louse and flea repellent. By the 1800s, crushed pyrethrum flowers were found in most European pharmacies. Pyrethrins are highly toxic to insects but are expensive and break down so rapidly in sunlight that they have little or no value for protection of crops or forests. They are used primarily as household insecticides. Azadirachtin, an extract of the neem tree, Azadirachta indica, also has insecticidal properties. Neem extracts have been known to disrupt or inhibit insect development, interfere with mating and sexual communication, repel feeding and deter egg laying. They have been shown to affect a wide range of insects (NRC 1992).

Chlorinated Hydrocarbons Chlorinated hydrocarbons were the first of several organic chemicals shown to have insecticidal properties. Examples include dichlorodiphenyltrichloroethane (DDT), methoxchlor, lindane, endrin and dieldrin. DDT was introduced during World War II and was soon proven effective against vectors of human diseases, including malaria and yellow fever. It is also effective against forest defoliators and was applied over large areas for control of defoliator outbreaks. Soil treatments using several other chlorinated hydrocarbons, including chlordane and dieldrin, were once used to provide long-term protection to wooden structures from termites. Chlorinated hydrocarbons are persistent and harmful to many non-target organisms including aquatic insects, fish and rapturous birds. Moreover, they kill natural enemies of other pests, such as scales and mites. Repeated applications of DDT have resulted in increases in numbers of scale insects. DDT has been banned in most countries of the world but is still used on a limited basis for control of malaria mosquitoes. Most of the

other chlorinated hydrocarbons have either been banned or have only limited uses.

Organophosphates Organophosphates function as nerve poisons. They react irreversibly with the enzyme acetylcholinesterase, a chemical responsible for inactivating acetylcholine at the neuromuscular junctions and at synapses in the central and peripheral nervous systems. Examples of organophosphate insecticides are malathion, diazinon, parathion and methyl parathion. Several of these materials are extremely toxic and easily absorbed through the skin. They tend to be relatively short lived in the environment. Malathion, which has a relatively low mammalian toxicity, has been used for control of forest defoliator outbreaks and is still widely used for insect control in urban forest ecosystems.

Carbamates Carbamate insecticides also inhibit production of acetylcholinesterase. The most widely used carbamate in forest insect management is carbaryl (SevinÒ), which has been applied aerially for control of forest defoliators and is used as a preventative spray to protect trees from bark beetle attack. Carbamates biodegrade relatively rapidly in the environment and warm blooded animals can detoxify and excrete these materials rapidly. One drawback of carbaryl is that it is damaging to honeybees.

Synthetic Pyrethroids Synthetic or stabilized pyrethroids are chemically similar to the pyrethrum plant extracts but are somewhat more persistent in the environment. Examples include resmethrin, permethrin and deltamethrin. They are broad spectrum insecticides that affect many insects, both harmful and beneficial. They have a low mammalian toxicity and break down relatively rapidly in the environment. Several synthetic pyrethroids have been used for control of forest defoliators and to protect trees from bark beetle attack.

Systemic Insecticides Systemic insecticides or systemics are products absorbed by one part of the plant and translocated to other portions of the plant, thus rendering them toxic to insects (Bennett 1957). Several organophosphate and carbamate insecticides have systemic properties. Systemics can be applied to the soil

Management of forest insect pests around the root collars of plants or trees, injected into the stem or sprayed on the plant. They are especially effective against sucking insects, stem and shoot borers and some cambium borers. Many systemic insecticides are highly toxic and can only be purchased and applied by licensed pesticide applicators. An exception is acephate (OrtheneÒ), which is used to protect ornamental trees and shrubs from insect attack.

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sooty mold from plants caused by aphid and scale infestations. These products are most widely used for insect control in urban settings but have also been used against the introduced hemlock woolly adelgid, Adelges tsugae, in National Parks in the USA (USDI National Park Service n.d).

Insecticide Resistance Insect Growth Regulators IGRs are a group of insecticides that interfere with insect development. They interfere with the synthesis of chitin, the material that makes up the exoskeletons of insects, or disrupt molting. They have little or no effect on adults but can be effective when applied to larvae or nymphs. The insecticide diflubenzuron (DimilinÒ) is an example of a chitin inhibitor and has been used for control of the larval stages of several forest defoliators including gypsy moth, Lymantria dispar.

Horticultural Oils Oils have been used for insect control for many years. Early products were most often referred to as “dormant oils” because they could only be applied to plants when dormant because they could damage foliage, shoots and buds. Modern horticultural oils have been developed as an alternative to traditional chemical insecticides. They are highly refined petroleum products that can be applied to plants for insect control at any time of year without injury to the plant and are safe to humans and non-target organisms. They are not poisons and work by suffocation. In order to be effective, target insects must be totally covered with the oil. They are only effective against insects with limited or no mobility, such as aphids and scales, and are used to control this group of insects on shade and ornamental trees and shrubs.

Insecticidal Soaps Insecticidal soaps are another safe and effective alternative to conventional insecticides. They are made from fatty acids of either plant or animal oils and are a highly refined version of products such as liquid soaps. However, conventional liquid soaps are not a substitute for insecticidal soaps because they can cause plant damage. Insecticidal soaps are effective against soft bodied insects such as aphids, thrips and the crawler (motile) stages of scale insects. They are also effective for removing honeydew and

Repeated exposure of insects to a pesticide can result in development of populations with significantly reduced susceptibility to that insecticide, even after just a few generations. Individuals in the population most resistant to a pesticide survive and pass the trait for resistance on to their offspring. Occurrence of insecticide resistance was noted as early as 1897 when difficulties were reported in attempts to control San Jose scale, Quadraspidiotus perniciosus, and codling moth, Cydia pomonella. Insecticide resistance to DDT was observed in 1946 when populations of resistant houseflies were discovered. As of 1984, the list of resistant species involved 14 orders and 83 families, and 428 different insects and related organisms. Of these, 61% were of agricultural importance and the remainder of medical/ veterinary concern (Forgash 1984). Resistance has even been observed to the biological insecticide Bacillus thuringiensis. This was first reported in 1985 in populations of a stored grain insect, the Indian meal moth, Plodia interpunctella (McGaughey 1985). To date, there have only been a few isolated cases of insecticide resistance reported in forest insect populations. This is due to the relatively low rates of exposure of forest insects to chemical insecticides when compared to insects of agricultural or medical/veterinary importance.

Pesticide Registration Pesticide use is highly regulated. Most countries now require that pesticides, including insecticides, be registered by an appropriate government agency for specific uses. In the USA, the Federal Insecticide, Fungicide, and Rodenticide Act (FIFRA), first passed in 1972 and amended several times since, provides control of pesticide distribution, sale and use. All pesticides used in the USA must be registered (licensed) by the US EPA. Registration assures that pesticides are properly labeled and, if used in accordance with specifications, will not cause unreasonable harm to the environment. Use of each registered pesticide must be consistent with

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directions on the label. Data required to register a pesticide are provided by the manufacturer and include product chemistry, environmental fate, residue chemistry, dietary and non-dietary hazards to humans, and hazards to domestic animals and non-target organisms (US EPA 2010). Microbial insecticides and semiochemicals are usually classified as “biological pesticides” as distinct from chemical pesticides. Registration requirements for these materials in Europe and the USA are generally less stringent than those of chemical pesticides, which reflect the fact that they occur naturally in the environment, are non-toxic in their mode of action and do not have a narrow range of activity (Wainhouse 2005).

APPLICATION TECHNOLOGY Microbials, semiochemicals and chemical insecticides can be applied either from the ground or air, depending on local conditions and the magnitude of the project.

Successful application of these materials requires careful consideration of the application equipment and its calibration and characterization, droplet size if liquids are applied and how the application is affected by meteorological conditions such as air currents, temperature, relative humidity and precipitation.

Ground Application Equipment A range of equipment is available to apply materials for insect control from the ground. These include hydraulic sprayers, mist blowers and foggers. Hydraulic sprayers are used for high-volume applications such as prevention of bark beetle attack on selected trees near home sites, developed recreation sites and other high-use areas (Fig. 5.9). In this case, the spray must reach a minimum top diameter limit and is applied to the point of runoff. Mist blowers and foggers apply sprays in smaller droplet sizes and tend to simulate applications of insecticides applied by aircraft.

Fig. 5.9 Preventative sprays are applied to high-value trees to prevent bark beetle attack in a forested homesite (Colorado, USA, photo by Sheryl Costello, USDA Forest Service).

Management of forest insect pests Aircraft Aircraft have been widely used for insecticide application in both agriculture and forestry and much of the technology used in forestry has been adapted from agricultural uses of aircraft. An excellent global review of the use of aircraft in agriculture is provided by Akesson and Yates (1974) although there have been a number of innovative changes since this work was published. In forestry, both chemical and microbial insecticides and, to a lesser extent, semiochemicals have been applied aerially. Both fixed-wing and rotorwing (helicopters) aircraft have proven to be valuable tools for application of these materials over large areas of remote, rugged and often inaccessible areas of forest. Fixed-Wing Aircraft Most fixed-wing aircraft presently in use for application of insecticides in forestry are low-wing monoplanes, capable of lifting a maximum payload of 900–1133 kg. These are most effective in mountainous terrain. Larger multi-engine aircraft can be used in forested areas with level or gentle terrain. Working speeds of these

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aircraft range from 145 to 182 km/hour (Akesson & Yates 1978a). Many types of fixed-wing aircraft have been adapted for application of pesticides, ranging from military aircraft such as the Grumman TBM to larger cargo transport aircraft such as the Douglas DC-3 or C-47. In eastern Europe, China and Russia, the Antonov AN-2, a bi-wing multi-purpose aircraft, has been used extensively for aerial application of insecticides (Fig. 5.10). More recently, a line of single-engine aircraft, specifically designed for aerial application of these materials, has become available. Examples include the Americanmade Piper Pawnee, Grumman Ag Cat and Ayres Thrush, and the Polish Pezetel M-18 Dromadier. These aircraft are designed to carry heavy payloads and have the power and maneuverability to apply materials from low altitudes and in mountainous terrain with relative safety. Some models are equipped with turbine engines that extend their maneuverability and load capabilities (Akesson & Yates 1978a, Quantick 1985a). Helicopters Helicopters have increased maneuverability and can operate from heliports established in close proximity to

Fig. 5.10 A Russian Antonov AN-2 configured for aerial application of insecticides over forests (Vilinius, Lithuania).

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the target area. However, they are considerably more expensive to purchase, operate and maintain than fixed-wing aircraft. Helicopters are widely used in forestry, especially in remote, mountainous regions. They usually operate at airspeeds of 80–145 km/hour, about two thirds of the speed of fixed-wing aircraft. Therefore, their productivity per flight hour is lower. Improved downwash of air from the rotor can result in improved deposition of spray particles at airspeeds below 32–40 km/hour. At airspeeds higher than 40 km/hour, the downwash effect of the rotor is lost (Akesson & Yates 1978b, Quantick 1985b). As is the case with fixed-wing aircraft, many models and sizes of helicopters are used in application of insecticides over forests. Some popular helicopters used in application of insecticides over forests include the Bell 47 B-1, a small agricultural helicopter, the military Bell 204 (Huey) or its civilian counterpart, the Bell 205, and the Bell 206-Jet Ranger.

Spray Systems Aircraft spray systems consist of several components. For application of liquids they consist of a tank to hold the material, filters to remove matter likely to cause blockage in the system, a pump to supply the liquid under pressure to atomizing equipment and a power source to drive the pump. In fixed-wing aircraft, the power source may be either a hydraulic or mechanical pump driven by the aircraft’s engine or a windmill mounted in the propeller slipstream. The latter system is cheap and simple but sensitive to changes in airspeed and power. In helicopters, it is impractical to use windmill driven pumps because of their relatively low airspeeds (Quantick 1985b). Spray aircraft should also be equipped with an emergency dump system to empty the spray tank in 10 seconds or less, should the aircraft encounter problems when in flight. While sudden release of a chemical pesticide may result in environmental contamination, the release of the aircraft’s payload during an emergency could save the pilot’s life. Two types of atomizing equipment are used to emit liquid sprays: a boom and nozzle system or rotary atomizers. Both are designed to break up spray into small droplets. Boom and nozzle systems consist of a hollow tube, streamlined in cross-section with plugs at the outer ends to facilitate cleaning. Generally between 20 and 50 nozzles are placed along the boom. Each nozzle is designed to deliver a given volume of liquid per minute

Fig. 5.11 A Micronaire AU 5000 rotary atomizer used for aerial application of both microbial and chemical insecticides (Vilinius, Lithuania).

at a specified boom pressure, usually 1.4 or 2.8 kg/cm2. Rotary atomizers consist of rotating cages, cylinders or discs driven by windmills using the aircraft’s slipstream or by electric motors (Quantick 1985b). Examples include the MicronairÒ AU series (Fig. 5.11) and the BeecomistÒ atomizers. Rotary atomizers have the advantage of delivering spray droplets of a given range of sizes independent of airspeed.

Calibration and Characterization Calibration of spray aircraft ensures the correct volume of material is applied over the target area. Aircraft characterization establishes the proper swath or lane separation of the aircraft. Both should be verified for each aircraft prior to the start of a project. Flow Rate Calibration involves computation of the aircraft’s flow rate, which is a function of the required volume rate, the effective swath width and the aircraft ground speed. This can be computed from one of the following two equations (Quantick 1978b): Metric :

FR ¼ VWS=600

Where: FR = flow rate in litres/minute V = volume in litres/ha

Management of forest insect pests W = swath width in m S = groundspeed in km/hour. Imperial or English :

FR ¼ VWS=495

Where: FR = flow rate in gallons/minute V = volume in gallons/acre W = swath width in feet S = groundspeed in miles/hour. Flow rate can be adjusted on boom and nozzle systems by adding or removing nozzles to achieve the desired flow rate. For spray systems equipped with rotary atomizers, system pressure is adjusted to achieve the desired flow rate. Actual flow rate of a spray aircraft can be verified by filling the spray tank with a known amount of spray and measuring the time required for the aircraft to release the spray with a stopwatch. For aircraft equipped with hydraulic or electrical pumps, calibration can be verified on the ground. Aircraft equipped with windmill pumps require that spray release time must be checked while the aircraft is flying.

Swath Width Swath width or lane separation is another critical variable to determine prior to start of a spray operation. If lanes are too widely separated, a portion of each lane will not receive the proper dosage and can result in ineffective treatment (see Plate 8). Lanes spaced too closely can result in overtreatment and inefficient aircraft use. Operational swath width can be determined by establishing a line of spray deposit cards (e.g. white cards made of KromekoteÒ paper) at known intervals in an open field and flying the aircraft over the card line while releasing spray. Oil based sprays can be sized by the stain they leave on a card stained with a material that reacts with the oil. Some water based formulations, such as several commercial formulations of Bacillus thuringiensis, have enough color to leave a stain on a white card. Other formulations may require the addition of a dye to stain cards for both swath width determination and droplet sizing (see following section). A square centimeter grid is then placed over each card and through counts of a subsample of grids on each card an estimate of the number of stains/cm2 is made across the card line. Effective swath is generally defined as the portion of the area card line where a droplet

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density greater than or equal to 20 droplets/cm2 is achieved. Ideally, the droplet density should be relatively even across the card line. Most modern spray aircraft are equipped with GPS to help maintain proper swath lane separation. Starting and ending points for each flight line can entered into the GPS receiver prior to the day’s flight and displayed on the unit’s screen for the pilot to follow. Actual flight track is recorded by the system and can be plotted on paper maps using GIS to monitor pilot performance and quality of application.

Droplet Size The droplet size spectrum delivered by the spray system is critical to the efficacy of any liquid formulation applied. In insect control, only a small portion of the material released from an aircraft may reach the intended target, i.e. insect or the host foliage. The remainder is lost to drift, defined as that portion of the spray that leaves the target area, or is deposited on the forest floor. Decreasing droplet or particle size increases the number of particles available to contact the target. Unfortunately, it also results in increased losses due to drift (Akesson & Yates 1978b, Barry & Ciesla 1981). Droplet size spectra are usually measured in micrometers (mm) and expressed as volume median diameter (VMD) or number median diameter (NMD) (Quantick 1985b). These are defined as: . .

VMD ¼ half of the volume of spray is above or below droplets of this size. NMD ¼ half of the number of spray droplets is above or below this size.

The smaller droplets within a spray cloud are the ones that impinge on the intended target, i.e. either the insect or the foliage. For example, a study in Montana, USA, indicated that 93% of the larvae of western spruce budworm, Choristoneura occidentalis, had been contacted by spray droplets no larger than 50 mm during application of an experimental spray. This study concluded that since most of the spray applied to forests by current application systems consists of droplets larger than 50 mm, the biologically active portion of the drop spectrum is low (Himel & Moore 1967). Most aerial spray systems available today deliver droplets with a VMD ranging from 100 to150 mm. Droplet size calculations can be somewhat complex because they require collecting and sizing of a large

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number of droplets, usually collected on spray deposit cards. Droplets are sized by the stain they leave on the card. The size of the stain left on a spray deposit card is converted to droplet size by computation of a spread factor. Spread factors are usually linear or quadratic regression equations developed from application of droplets of known sizes to spray deposit cards in the laboratory and measuring stain size and are a function of both the spray formulation and paper used for spray deposit cards. An abbreviated approach, known as the D-max method, has been developed, which involves measuring the five largest stains collected across the spray swath. The largest stain in the series with a diameter difference of less than or equal to 32 mm to the next droplet in the series is selected. The stain diameter is converted to droplet size using the appropriate spread factor. VMD is determined by dividing the size of the selected droplet by 2.2 for slow-flying aircraft (133 km/hour) and 2.5 for medium-speed aircraft (280 km/hour) This can be used for both oil and water based sprays (Maksymiuk 1964, Ciesla & Livingston 1980).

wind velocities low. Typically, operations begin when there is enough light for the pilot to see the terrain. In June in the northern hemisphere, this is approximately 05:00 hours. On a day with exceptionally favorable weather, it may be possible to continue operations until 10:00 hours. In some areas late afternoon spraying, roughly 1–2 hours before sunset, may also be feasible.

Meteorology

.

Weather conditions during the specified application period are critical to the success of the application. Low relative humidity and high temperatures can cause spray droplets to evaporate before they reach their intended target. Insufficient wind or turbulence can cause spray droplets to hang in the air and not impinge on target insects or foliage. High winds in excess of 10 km/hour will cause the spray to drift away from the target area. Occurrence of inversion layers can result in poor settling of smaller spray droplets. Rain or wet foliage will cause the spray to run off the target foliage, especially if oil based insecticide formulations are applied. Meteorological monitoring is, therefore, a critical aspect of any aerial application project. For small projects, deployment of observers equipped with field weather kits that monitor relative humidity, temperature and windspeed and portable radios is needed to determine if conditions are suitable for the application. For large projects, field weather stations coupled with observers in the target areas may be needed. Weather conditions are usually most favorable for aerial application during the early morning hours when relative humidity is high and temperatures and

Modeling Behavior of Aerial Applications The USDA Forest Service has developed an aerial spray drift and deposition model, known as the Forest Service Cramer-Barry-Grim (FSCBG) model, which predicts dispersion of material released from aircraft into the atmosphere and downwind from the release point. Effects of aircraft wake structure, droplet evaporation, local meteorology, penetration through forest or agricultural canopies and prediction of ground or canopy deposition, air dosage and concentration are all included in the model (Teske 1996, Teske et al. 1996). Applications of this model include:

. . . .

establishment of protective buffer zones based on defined spray scenarios, non-target areas and sensitive species; prediction of productivity, biological effects and operational costs of a spray project; definition of options to achieve a specified buffer zone; prediction of sensitivity on drift and productivity of changes in spray scenario; integration and correlation of this information into understandable graphics.

INTEGRATED PEST MANAGEMENT SYSTEMS IPM systems are a set of decision-making and pest management tools directed against a pest or pest complex. The use of various pest management tactics to address a pest may be determined by factors such as accessibility, terrain features and land use classification. In addition, some components of an IPM system may be in various stages of development. An example of an evolving IPM system is management of the wood wasp, Sirex noctilio, in pine plantations in the southern hemisphere. Early detection is accomplished by baiting suppressed trees in plantations

Management of forest insect pests with a low dosage of a herbicide. This stresses trees and attracts attacking wasps. Infestations are treated either through inoculation of the parasitic nematode, Deladenus siricidicola, and thinning to reduce stocking and maintain tree vigor. Remote sensing technologies have been evaluated to locate small, non-industrial private pine plantations that might serve as reservoirs of this insect. Introduction of parasitic insects is another potential tactic for managing this insect (Haugen et al. 1990, Iede & Ciesla 1993, Disperati et al. 1998, Iede et al. 1998). In the Great Smoky Mountains National Park, located in the southern Appalachian Mountains of the southeastern USA, a combination of tactics is used to reduce the impact of the introduced hemlock woolly adelgid, Adelges tsugae (USDA National Park Service n.d.): .

.

Foliar treatments. Hemlocks in developed areas accessible by road are treated with insecticidal soaps or horticultural oils. This approach is effective against insects present on the tree at the time of application. Re-treatment is needed at 6-month to 1-year intervals. Systemic treatments. Hemlocks in, or near, campsites too tall to be sprayed are treated with a systemic insecticide (imidacloprid), either by soil drenching or

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by injecting the insecticide directly into the trunk. Treatment may remain effective for up to 3 years. Biological control. A classic biological control program that involves release of predaceous beetles, which feed exclusively on hemlock woolly adelgid, was initiated in 2002. Predaceous beetles are released in areas not treated with insecticides.

INTEGRATION OF NEW TECHNOLOGIES An important part of IPM is that no matter how advanced, sophisticated or effective an IPM system for a specific insect pest or pest complex may be, there is always room for introduction of new technologies that may allow for more accurate pest monitoring and prediction or new tactics, which provide more efficacious treatments with fewer undesirable side effects. Therefore, it is essential for forest health practitioners (both forest entomologists and pathologists) to have strong ties to scientists in research institutes charged with the development of new approaches to the management of forest insects and other pests and be ready and willing to implement new technologies as they become available.

Chapter 6

Forest Insect Orders and Families

INTRODUCTION The chapters that follow provide profiles of insects that utilize parts of trees for food and/or breeding sites. Most of the species described are of socioeconomic concern and some are extremely damaging. Others cause damage that is primarily of a cosmetic nature and may be of concern in urban settings but not in forests. A few of the species described are beneficial to humans or, simply, curiosities. The profiles are organized according to the portion of the plant that the insects utilize, i.e. foliage, bark and cambium, wood, stems, reproductive structures, etc. Those species that may utilize more than one part of the plant, such as insects that feed both in shoots and reproductive structures (flowers, nuts or cones), are included in the chapter that addresses the part of the plant where they are most frequently found. Within each chapter, insects are organized by taxonomic groups, i.e. orders, families, etc. This chapter addresses some of the basics about insects, including metamorphosis and how insects

are classified, and provides overviews of the major insect families that contain species of interest in forestry.

INSECTS DEFINED What are insects? Insects are a group of animals of the phylum Arthropoda, known as class Insecta. Arthropods are invertebrates with distinct body regions, an exoskeleton or hard exterior shell made of chitin and segmented appendages. Insects have three body regions, i.e. head, thorax and abdomen. The adult stages have three pairs of legs, a pair of antennae and, if winged, two pairs of wings. Related arthropods include the arachnids (spiders and mites), which have two body regions and four pairs of legs. Crustaceans (shrimp, crabs, lobsters, etc.) have two body regions, two pair of antennae and at least five pairs of legs. Chilopods or centipedes have two body regions, i.e. head and trunk, and one pair of legs per

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Insect development ranges from simple to extremely complex. Some insects undergo relatively little change as they develop, whereas others undergo rather remarkable changes over the course of their lives. The process of change as insects develop is known as metamorphosis. There are two overall types of metamorphosis: simple and complete. Insects that undergo simple metamorphosis have three life stages, i.e. egg, nymph and adult (Fig. 6.1). Eggs deposited by adults hatch into nymphs, which undergo several growth stages known as instars. In most cases nymphs resemble the adult stage but are smaller and do not have fully developed wings. The feeding habits of nymphs and adults of insects that

undergo simple metamorphosis tend to be similar and nymphs and adults are often seen feeding together in colonies. Adults are the reproductive stage and, if winged, disperse to new feeding sites. Examples of tree feeding insects that undergo simple metamorphosis are walking sticks, termites and sucking insects such as aphids, scales and true bugs (Plate 9). Insects that undergo complete metamorphosis have four distinct life stages, i.e. egg, larva, pupa and adult (Fig. 6.2). The appearance of each of these life stages is quite different. An example of complete metamorphosis is that of beetles (Coleoptera). Adults deposit eggs, which hatch into larvae. The larvae, usually referred to as grubs, may feed for several weeks or months and molt or shed their exoskeleton as they grow. The period between molts is called an instar. When feeding is completed, larvae may construct a cocoon and pupate. The pupal stage is a resting stage during which transformation into adults takes place. When the transformation is completed, adults emerge. Other examples

Fig. 6.1 Partial or incomplete metamorphosis: egg, nymph and adult as exemplified by the box elder bug, Boisea trivittata (Hemiptera: Rhopalidae).

Fig. 6.2 Four stages in the life history of a bark beetle of the genus Ips (Coleoptera: Curculionidae: Scolytinae): egg, larva, pupa and adult are an example of complete metamorphosis.

body segment. Diplopods or millipedes have two body regions and two pairs of legs per body segment.

METAMORPHOSIS

Forest insect orders and families of insects that undergo complete metamorphosis are moths and butterflies, the larval stage of which are typically called caterpillars, and flies, which have a larval stage often called “maggots.” When insects that undergo complete metamorphosis become adults, they stop growing. Some adults may continue feeding while others feed only on a limited basis or not at all. The primary function of adults is mating, reproduction and dispersal to new feeding sites. The life span of adults of many insects is short, just long enough for them complete these functions. Adults of other species may live for longer periods, with some species passing the winter months as adults. Some insects have developed the ability to reproduce by parthenogenesis, where all adults are females that reproduce without mating. Others may have both sexual and asexual generations. Insects that reproduce either partially or entirely by parthenogenesis include species of aphids, scales and wasps. While the adults of many insects are winged and can disperse over distances of several kilometers, others have either never developed wings or have lost their capacity to fly. Some aphids may develop both winged and non-winged adults. Many female moths are weak fliers because their bodies are heavily laden with eggs. Some, such as the female European gypsy moth, Lymantria dispar, have fully developed wings but do not fly. Females of other species of moths, such as some tussock moths, have wings reduced to stubs. They simply emerge from their pupal cases, emit pheromones and await arrival of a male. After mating, they deposit an egg mass on, or near, the pupal case and die.

TAXONOMY Classification Taxonomy is the classification of living organisms into recognizable entities and provides names by which they can be recognized. The fundamental unit of taxonomy is the species. The concept of a species is sometimes vague but most scientists agree that species are biological entities that: . .

are capable of breeding and producing viable offspring; under natural conditions, are reproductively isolated from organisms that may be similar in structure and appearance.

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In a formal biological classification, species are grouped into a hierarchy of categories known as taxa. The typical hierarchy of categories or levels of classification used in zoology and in the classification of insects is as follows: Phylum Subphylum Class Subclass Order Suborder Superfamily Family Subfamily Tribe Genus Subgenus Species Subspecies

Not all of these categories are required to classify a species. The primary levels are: phylum, class, order, family, genus and species. Scientific Names The scientific or Latin names of all living organisms are designated by genus and species. This system, known as the binomial system of nomenclature, was first introduced in 1749 by Carolus Linnaeus, a Swedish botanist. In his “Species Plantarum,” published in 1753, Linnaeus attempted to describe all known plants and assigned to each a two-part Greek or Latin name, which consisted of genus followed by species. His 1758 edition of Systema Naturae extended binomial classification to animals. For example, the European spruce bark beetle is designated as Ips typographus (Linnaeus) (Coleoptera: Curculionidae: Scolytinae). The representation of this species in the above hierarchy is as follows: Phylum Class Order Family Subfamily Tribe Genus Species

Arthropoda Insecta Coleoptera Curculionidae Scolytinae Ipinae Ips typographus

The name (Linnaeus) that appears immediately behind the genus and species is the author of the

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species; the person that originally described this species. I. typographus was described by Carolus Linnaeus. His name appears in parenthesis, indicating that the species typographus has since been placed in a genus other than the one under which the species was originally described. In the case of Ips typographus, the species was originally described as Dermestes typographus Linnaeus. If no parentheses appear behind the author’s name, the classification of the species has remained unchanged since its original description.

Name Changes Taxonomy, especially insect taxonomy, is an entity as dynamic as the insects themselves. New species are continuously being described. Moreover, families, genera and species, are re-examined by specialists and new classifications are proposed and published. For example, for many years, the bark beetles were classified as a distinct family of beetles, the Scolytidae. In 1995, they were re-classified as the subfamily Scolytinae under the family Curculionidae, the weevils (Lawrence & Newton 1995). In a similar revision, the tussock moths, a group containing many species of damaging forest defoliators, were once classified as the family Lymantriidae. A revision of this group now places them in the subfamily Lymantriinae under the family Noctuidae (Lafontaine & Fibiger 2006). Today, tools such as electron microscopes and analysis of mitochondrial deoxyribose nucleic acid (DNA) provide better insights into the classification of species.

Common Names In addition to scientific or Latin names, most wellknown plants and animals have common names by which they are referred to on an informal basis. Depending on the organism’s natural and introduced range, species may have common names in several languages. The bark beetle, Ips typographus, which ranges across the spruce forests of Eurasia, has common names in several languages: . . .

English – European spruce bark beetle; French – Gran scolyte de l’epicea, Le typographe de l’epicea; German – Buchdrücker, Grosser 8 - z€ahniger Fichtenborkenk€ afer;

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Norwegian – Granbarkbillen.

In most places, assignment of common names to living organisms is an informal process and names evolve over time. However, in Canada and the USA, the assignment of common names to insects has been formalized by the Entomological Society of Canada and the Entomological Society of America, respectively. Both societies have established processes for proposing common names and each publishes a list of “approved” common names on the Internet (www.entsoc.org/ Pubs/Common_Names/search.asp, www.esc-sec.ca/ commonnames.html). Common names used in this text are a combination of “approved” common names and those in common usage.

ORDERS AND FAMILIES The following sections provide brief overviews of insect families that include species of importance in forestry and profiled in the following chapters. They represent only a fraction of the total families and subfamilies recognized by taxonomists. In most cases, the classification of insects and nomenclature used follows that of Triplehorn and Johnson (2005) unless significant changes have occurred since this text was published.

Phasmatoidea, Walkingsticks Walkingsticks are a medium-sized order of insects, many of which look like sticks or leaves. About 2700 species are known and most occur in the tropics. A few species occur in temperate forests and defoliate trees. Some species are popular as pets. Four families of walkingsticks are recognized. The family that contains species of importance in forestry is the Heteronemiidae.

Isoptera, Termites Termites are a small but diverse order of cellulose feeding insects. They are unable to digest cellulose directly but have developed symbiotic relationships with protozoa and bacteria that do break down cellulose in wood. Approximately 2600 species in seven families and 281 genera are known worldwide (Kambhampati & Eggelston 2000). Many are damaging to wooden structures. They live in highly organized

Forest insect orders and families societies or colonies with distinct morphological forms or castes. These include reproductives, workers and soldiers. Each caste performs a different and critical function in the colony and can be so morphologically distinct that, to the untrained eye, they can appear to be separate species. Termites represent an ancient society and have been on Earth for a long time. They evolved some 200 million years ago, during the late Permian Period (Triplehorn & Johnson 2005). Mastotermitidae This family consists of a single species, Mastotermes darwiniensis, found only in northern Australia. It is considered the most primitive termite alive today. Unlike other termites, queens lay their eggs in small masses instead of singly.

Kalotermitidae The Kalotermitidae consists of several distinct groups: drywood, dampwood and powder post termites. They have no worker caste and immature forms of the other castes perform the work of the colony. Unlike other termite families, they do not construct earthen tubes. Drywood termites invade sound dry wood. They do not have a ground contact for a moisture source and may be found in buildings, furniture, utility poles and piled lumber. Dampwood termites invade moist, dead wood and tree roots. Powder post termites usually attack dry wood, without a ground contact and reduce it to powder.

Termopsidae, Dampwood Termites Approximately 85% of the world’s known termites are in the family Termopsidae. They invade dead wood but do not require a ground contact. A true worker caste is not produced. Some moisture is required in the wood they invade. Some species cultivate fungus gardens, which provide the food for the colony (Kambhampati & Eggleston 2000).

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tend to be small. Adults range from 6 to 8 mm long and the wingless forms are pale colored. Winged reproductives are dark brown-black. Species of the Rhinotermitidae always maintain contact with the soil and may construct earthen tubes to connect to wood that is not in direct contact with the soil.

Termitidae Termites of the family Termitidae tend to occur in arid and semi-arid environments. Some species lack a soldier caste. Some burrow under logs or cow dung and are not economically important. Another group, known as “desert termites,” is subterranean and may damage fence posts, poles and wooden buildings. Other species attack desert shrubs or other objects that have direct contact with the ground. Many Termitidae cultivate fungus gardens for a food source (Triplehorn & Johnson 2005).

Hemiptera, Aphids, Scales, Bugs The order Hemiptera (¼ Homoptera) is a large and diverse group of insects. The single characteristic that unifies this order is their mouthparts. They are of a piercing/sucking type and consist of four stylets enclosed in a slender, flexible sheath. These mouthparts are usually used for sucking plant juices but a few are blood sucking. Insects of the order Hemiptera undergo simple metamorphosis. In a few species, such as the scale insects, development somewhat resembles complete metamorphosis in that the last nymphal instar resembles a pupa. Several families include species that are important forest pests. Sucking insects that infest foliage and stems are discussed in Chapter 11, those that produce galls are discussed in Chapter 12 and those that feed on and damage tree reproductive structures are reviewed in Chapter 14.

Lygaeidae, Seed Bugs Rhinotermitidae The Rhinotermitidae consists of both subterranean and dry wood termites and contains species that cause severe damage to homes and other structures. They

The Lygaeidae are characterized by having dorsal abdominal spiracles. Many species are brightly colored. They feed chiefly on seeds and some are pests of agricultural crops. A few species feed on seeds and cones of forest trees.

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Coreidae, Leaf-Footed Bugs Coreids are a moderate-sized group of Hemiptera whose members have well-developed scent glands and most species will emit distinct odors when disturbed. Most are plant feeders but a few are predators of other insects. Some have hind legs that are expanded and resemble a leaf, hence the name “leaf-footed bugs.” Members of the genus Leptoglossus feed on seeds inside developing cones or conifers. Rhopalidae, Scentless Plant Bugs Thescentlessplant bugsarea smallfamilyofplantfeeding Hemiptera. They differ from the closely related Coreidae in that they lack well-developed scent glands. The box elder bugs, Boisea spp., which feed on seeds of boxelder, Acer negundo, are members of this family (Plate 9). Psyllidae, Jumping Plant Lice Psyllids are small insects, 2–5 mm long and often resemble miniature cicadas in body shape. They have strong legs for jumping and relatively long antennae. Nymphs of many species construct a white cover of sugar or lerp, and feed concealed under this shelter. In Australia, they are known as lerp insects. Aphididae, Aphids Aphids are a large group of small, soft-bodied insects and many are important forest pests. They usually occur in colonies, which can be composed of all life stages, on stems and leaves of plants. Aphids have pearshaped bodies, relatively long antennae and a pair of projections known as siphunculi (cornicles) at the posterior end of the abdomen. Adelgidae, Adelgids Adelgids are a small family of sucking insects related to aphids. About 70 species have been described and 50 are relatively well known. All are native to the northern hemisphere, where they infest boles and branches of conifers of the genera Abies, Larix, Picea, Pinus, Pseudotsuga and Tsuga. They have complex life cycles and individual species may be holocyclic, with sexual and asexual generations or anholocyclic with only asexual generations. A typical holocyclic species

Fig. 6.3 Life cycle of adelgids (Hemiptera: Adelgidae). The center circle depicts a host alternating holocycle and the lower circles depict anholocycles on the primary host (left) and the secondary host (right). Plain circles represent wingless generations and circles with lines represent winged generations (redrawn from Havill & Foottit 2007).

undergoes five generations, three on the primary host, which includes sexual reproduction and gall formation, always on species of Picea, and two asexual generations on the secondary host. The secondary host is always another conifer genus. The five generations require 2 years to complete (Havill & Foottit 2007, Fig. 6.3). Several species of Adelges and Pineus are important forest pests. They are easily transported on plant materials and some have been introduced into new locations and caused severe damage. Species of concern because they infest foliage, twigs or boles of secondary hosts are discussed in Chapter 11; those that produce galls on Picea are discussed in Chapter 12. Margarodidae, Giant Coccids or Bast Scales The Margarodidae are large scale insects, sometimes reaching lengths of 25 mm. Some are pests of

Forest insect orders and families agricultural crops and one genus, Matsucoccus, is a pest of pines in the northern hemisphere. Pseudococcidae, Mealybugs Mealybugs get their name from the copious amounts of mealy or waxy secretion that covers their bodies. Females are elongate to oval shaped, segmented and have well-developed legs. Some species lay eggs and others give birth to live young. They can be found on almost any part of the host plant.

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Kermesidae, Gall Scales Kermes scales, or gall-like scales, consist of several genera (Allokermes, Eriokermes, Kermes, Kermococcus, Nanokermes) found on oaks, Quercus spp., throughout the northern hemisphere. Feeding by some species can cause growth loss, branch dieback, leaf distortion and their honeydew is a medium for development of sooty mold fungi. Several are pests of ornamental trees and one species, Kermes vermilio, indigenous to the Mediterranean region, was once a highly prized source of red dye. Mature scales are tan, globular and hard and easily mistaken for galls or buds (Fig. 6.4).

Ericococcidae, Felt Scales This group of scale insects is somewhat similar to the Pseudococcidae but their bodies are only lightly covered with wax. Females and their eggs are often entirely closed in a felt-like sac. These insects also produce copious amount of honeydew, which provides a medium for growth of black sooty molds.

Kerriidae, Lac Scales These insects have a globular form, are legless and live in cells of resin. Most species occur in tropical climates. The honeydew or resin of one species, the lac insect, Kerria (¼ Laccifer) lacca, is harvested and used in production of shellac or varnish.

Fig. 6.4 Female of an unidentified species of oak kermes (Hemiptera: Kermesidae) with a typical gall-like shape on Gambel oak, Quercus gambelii (Near Moab, Utah, USA).

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Diaspididae, Armored Scales

Buprestidae, Flat Headed Wood Borers

Armored scales are the largest family of scale insects and consist of many species of plant pests. They develop waxy or scale-like coverings over themselves. Female nymphs are motile only during their first instar. During molting, the cast skin is incorporated into the protective covering with waxy secretions from their bodies. Females develop to maturity under the covering and deposit eggs or give birth to live young. Males develop much like females but emerge as motile, winged adults. Several are pests of shade and ornamental trees.

The Buprestidae are a large family with 15,000 known species in about 450 genera. Adults are called “jewel beetles” or “metallic wood borers” because of their metallic, iridescent hues (Plate 10). Many are popular with insect collectors. Adults are cylindrical, elongate or ovoid. Some species are small, around 3 mm in length, others are larger and over 100 mm long. Average size is 20–25 mm. Many species are sun lovers and are often seen on logs or tree stems exposed to direct sunlight. Larvae are known as flat headed wood borers because most species have a large flattened thorax that looks like a head (Fig. 6.5). Most species bore in logs, roots and stems but a few are leaf miners. Some are attracted to recently burned trees and forests in search of suitable breeding sites and often arrive in large swarms. Larvae of some species can survive in wood for long periods. One record from Australia indicates adults emerging from wood in use after 25 years. Therefore, they can be transported to new locations via international trade in wood products (Hill 1997, Evans et al. 2004).

Coleoptera (beetles) The order Coleoptera, or beetles, is the largest order of insects and make up about 40% of all known insect species (Erwin 1982). Beetles have two pairs of wings but the forewings are thickened and hard and usually meet in a straight line down the middle of the back. They are known as elytra and cover the hind wings, which are used for flight. Beetles undergo complete metamorphosis and have chewing mouthparts both in the larval and adult stages. Beetles can cause a wide range of damage to forest trees. Some species are defoliators (see Chapter 8) while others bore in the cambium and wood (see Chapters 9 & 10). Still others infest twigs or stems, are pests of young trees (see Chapter 13), reproductive structures (see Chapter 14) and wood in use (see Chapter 15).

Scarabaeidae, June or Cockchafer Beetles The Scarabaeidae are large, heavy bodied and variable. Adults of many species are brightly colored in metallic, iridescent hues. All are characterized by having antennae in which the last three segments are expanded into plate-like structures that can be spread apart and are known as lamellate antennae. Larvae are white, C-shaped, have well-developed legs and are known as “white grubs.” Scarab beetles are variable in their habits. Some feed on dung, decomposing plant materials or fungi. Many feed on living plants, including foliage, flowers and root systems, and are important pests of crops, forests, nurseries and golf courses. The larvae of some scarab beetles live in the soil and feed on roots of plants.

Bostrichidae, Twig Borers and Powder Post Beetles These beetles are elongate and somewhat cylindrical in shape. The head is bent downward and is scarcely visible when viewed from above. The antennae are straight with a loose club. Adults are from 3.5 to12 mm long. Most species bore in branches of live trees or seasoned lumber. Subfamily Lyctinae Members of the subfamily Lyctinae bore into dry, seasoned wood and are known as powder post beetles because they reduce the wood to a fine powder. They damage furniture, tool handles, beams and hardwood floors. Anobiidae, Death Watch and Spider Beetles These are small beetles, 1–9 mm long, cylindrical and covered with pubescence. The head is concealed from above by a hooded pronotum. Most live on dry vegetation such as twigs and logs or under bark of dead trees. The larvae of some species feed on fungi or in seeds and stems of plants. They are called “death watch” beetles because they make a ticking sound as they bore in the

Forest insect orders and families

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Fig. 6.5 Larva of a flat headed wood borer (Coleoptera: Buprestidae).

wood. Some species bore into and damage timbers, woodwork and furniture.

result, several species have been established in new locations and become pests.

Cerambycidae, Round Headed Wood Borers

Chrysomelidae, Leaf Beetles

The Cerambycidae are a large family with approximately 20,000 known species. Adults have long antennae and are known as longhorn beetles or, more archaically, longicorn beetles. Many are striking, colorful insects and a favorite of collectors (Plate 11). Larvae are white, legless grubs with an amber or light brown head capsule (Fig. 6.6). Larvae of most Cerambycidae bore in the cambium and wood of living trees, logs or wood in use. Others bore in stems and roots of herbaceous plants. They are known as round headed wood borers because they produce rounded or slightly oval exit holes when emerging from infested trees or logs. Some infest weakened or recently dead trees and others can infest living trees. Many are considered pests because they weaken the structural integrity of living trees or wood in use, reduce the quality of lumber, or function as vectors of tree diseases. Some are known to kill trees. Some species spend long periods in the larval stage in wood and can be transported long distances via logs, wooden crating, pallets, dunnage or fuel wood. As a

The chrysomelid beetles are closely related to the Cerambycidae but are smaller, usually less than 12 mm, have shorter antennae and are oval shaped. They feed on foliage and flowers of plants. In some cases, both adults and larvae can be seen feeding on the same plant. Larvae have a variety of habits. Some skeletonize foliage, others are leaf miners and a few are stem borers. Several species are tree pests and species of at least one genus, Diorhabda, are cultivated as biological control agents for control of salt cedar, Tamarix spp., an invasive tree in southwestern USA.

Curculionidae, Weevils Weevils have their heads prolonged into a beak or snout. Their mouthparts are small and hidden and the mandibles are located at the tip of the snout. All are plant feeders and can attack virtually any part of a plant, from roots to stems, foliage and fruits. Many are important pests of agricultural and tree crops. The

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Fig. 6.6 Larva of a round headed wood borer (Coleoptera: Cerambycidae).

Curculionidae are subdivided into a number of subfamilies. Two of these, the Platypodinae and Scolytinae include many important forest pests. Species in these two subfamilies do not have snouts characteristic of most Curculionidae.

Subfamily Scolytinae, Bark Beetles and Ambrosia Beetles Beetles of the subfamily Scolytinae are cylindrical, small (usually no more than 6–8 mm long), and brown-black in color. Antennae are short and the outermost segments are clubbed. Scolytinid beetles are either bark beetles or ambrosia beetles. Both groups have symbiotic relationships with fungi. Bark beetles breed in the inner bark of trees, often introducing wood staining fungi into the trees they attack. Other species are vectors of pathogenic fungi, which adults spread from tree to tree as they feed on twigs or branches. Attacking adults and larvae construct an elaborate network of galleries in the inner bark, which are often referred to as engravings. Most bark beetles breed in dying or recently dead or severely weakened trees. However, representatives of several genera are major

tree killing insects and constitute the single most destructive group of forest insects. Ambrosia beetles construct galleries in wood of dying, weakened or, occasionally, live trees. They inoculate the galleries with spores of ambrosia fungi, which serve as food for developing larvae. Ambrosia beetles do not kill trees unless the fungi with which they are associated are pathogenic.

Subfamily Platypodinae, Pinhole Borers or Ambrosia Beetles The Platypodinae are elongate, slender and cylindrical beetles with heads slightly wider than the pronotum. They are brown and range from 2 to 8 mm long. Antennae are short and clubbed. Adults bore into wood of trees and construct galleries and cradles in which brood are raised. Larvae feed on fungi introduced into the galleries by the invading adults. Most species confine their attacks to weakened, dying or recently dead trees and fresh cut logs. However, a few species are damaging because they and their associated fungi discolor or damage the structural integrity of wood.

Forest insect orders and families Hymenoptera (bees, wasps and ants) The order Hymenoptera, which includes such forest pests as sawflies, woodwasps, gall wasps, carpenter bees and carpenter ants, is a large and diverse group of insects. Adults typically have four membranous wings. Hind wings are smaller than forewings and have a row of tiny hooks on their anterior margin by which they attach to a fold on the posterior edge of the forewing. For most species, mouthparts are of a chewing type but in some families, such as bees, they form a tongue-like structure through which liquids, such as nectar from flowers, are taken. Metamorphosis is complete. The ovipositor, or egg laying mechanism, of females is usually well developed. In some cases it is modified into a stinger, which is also used as a weapon. Stings can be painful and some people have developed severe allergic reactions to bee and wasp stings. From the human perspective, the order Hymenoptera includes some of the most beneficial of all insects. Many species are parasitoids or predators of harmful insects. Others are pollinators of plants and the honey bee, Apis mellifera, which has been domesticated by humans for several hundred years, is the source of a healthy and nutritious sweetener. Some are pests of plants and trees of importance in agriculture and forestry and cause several kinds of damage. Species that feed on foliage are reviewed in Chapter 8. Those that bore in wood are reviewed in Chapter 10. Gall making species are described in Chapter 12, species that damage tree reproductive structures in Chapter 14 and pests of wood in use in Chapter 15.

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recognized by the unique antennae of males, which tend to have a U- or Y-shaped last antennal segment that resembles a tuning fork and makes them appear as though they have two sets of antennae. Some adults are dark but a number of tropical and subtropical species are brightly colored. Larvae typically feed in colonies on tree foliage. Diprionidae, Conifer Sawflies The Diprionidae are a small family of about 11 genera and 125 species and include a number of important forest defoliators. They occur throughout the conifer forests of the northern hemisphere and some are found as far south as Cuba, Mexico, Thailand, northern India and north Africa (Smith 1993, Cibri an Tovar et al. 1995). Tenthredinidae, Tenthredinid or Common Sawflies The Tenthredinidae is a large family of sawflies, subdivided into about 18 genera. About 6000 species have been described worldwide. Adults are small to medium in size and up to 20 mm long. They are wasp-like insects, often brightly colored and are often seen on foliage or flowers. They are found worldwide and feed on many species of plants. Larvae of most species are external feeders on foliage but some are leaf miners, stem borers or gall makers. Siricidae, Woodwasps

Pergidae The Pergidae are a moderate-sized family of sawflies. About 400 species in 60 genera have been described. TheyoccurprimarilyinAustralia andSouthAmerica and do not occur in Africa. Larvae typically feed in colonies on foliage of host trees but a few Australian species are leaf miners on Eucalyptus. Their morphology, habits and food habits are diverse. They are among the most common sawflies in the subtropical and tropical regions of the western hemisphere (Schmidt & Smith 2006).

Argidae The Argidae are a group of stout-bodied sawflies. About 800 species are known worldwide. They are easily

Woodwasps or horntails consist of about 85–100 species of primitive wasps. They are widely distributed in the forests of the northern hemisphere and extend south to Cuba, northern Central America, New Guinea, the Philippines, Vietnam, northern India and northern Africa. One genus occurs in tropical Africa. Larvae bore into weakened and dying trees and may take 1–3 years to complete development. Most are of minor importance but can reduce structural integrity and quality of lumber. Larvae are easily introduced to new locations via logs, lumber, wooden pallets or dunnage, where some species have caused severe damage. Adults are large robust insects that range from 25 to 38 mm in length. They have a spear-shaped plate, known as a cornus, at the end of the abdomen, which gives them the name “horntail.” Females have a long ovipositor

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under the cornus. Males are typically smaller than females. Larvae are legless grubs, creamy white in color, up to 30 mm long with a distinctive dark spine at the rear of the abdomen (Smith & Schiff 2002, Haugen & Hoebeke 2005). Eulophidae Eulophids are a large family of wasps and consist of about 4300 species in 300 genera worldwide. Adults of most species are 1–3 mm in length. Most are parasitoids and considered beneficial. A few are plant feeders and several produce galls on species of Eucalyptus and other plants. Torymidae The Torymidae are slightly elongate wasps with long ovipositors. Most are parasitoids of gall insects and caterpillars. Members of the genus Megastigmus infest and destroy seeds and some are important forest pests. Cynipidae, Gall Wasps Wasps of the family Cynipidae produce galls on woody plants. Adults are small, 1–8 mm in size, dark wasps and are known over much of the world. Many species produce galls on the foliage and stems of oaks, Quercus spp. Others produce galls on plants of the family Rosaceae. In North America, approximately 76% of the roughly 700 known species induce galls on oaks. Many Cynipidae have alternating sexual and asexual generations. The adults and galls produced by each form are morphologically distinct to the point that some have been regarded as separate species (Browne 1968, Drooz 1985). Formicidae, Ants Ants are an extremely large and well-known group of insects and are found almost everywhere in terrestrial habitats. They are social insects and most colonies contain at least three castes: queens, males and workers. Feeding habits of ants are extremely varied. Many are carnivores and feed on flesh of animals, living or dead. Others feed on plants, fungi, sap, nectar and honeydew produced by sucking insects. A few ants are

defoliators of forest trees and one group, the carpenter ants, build nests in wood.

Lepidoptera, Moths, Skippers and Butterflies The Lepidoptera are the second largest order of insects and include butterflies, skippers and moths. They undergo complete metamorphosis. Adults have membranous wings covered with scales. The scales can be either extremely colorful, as is the case with many butterflies, or they may be dull to blend in with their surroundings, such as the bark of trees. The larvae have strong mandibles, adapted for chewing, and adults have highly specialized mouthparts adapted for sucking nectar from flowers. More than 160,000 species are known. Many Lepidoptera, especially butterflies, are popular with insect collectors. Larvae of several families are damaging forest pests. Many species feed on tree foliage, can reach outbreak levels and cause widespread damage over millions of hectares (see Chapter 7). Larvae of others invade wood, cause structural damage to trees, or degrade lumber (see Chapter 10). Still others bore in shoots of trees or cones of conifers (see Chapters 13 & 14). Psychidae, Bagworms The bagworms include about 1000 described species in 300 genera distributed worldwide. Larvae construct portable cases in which they live as they feed. Female adults are often wingless and remain inside the larval case to mate and lay eggs (Rhainds et al. 2009). Gracillariidae, Leaf Blotch Miners The Gracillariidae are a large family and consist of about 98 known genera distributed worldwide. They are small moths with lanceolate wings. Larvae are leaf miners and several species can cause severe discoloration of the foliage of their host plants. Coleophoridae, Casebearers Adults are small moths with narrow, sharply pointed wings. Larvae are leaf miners when young and later live inside a case made from the foliage of host plants. Several species are of importance in agriculture and

Forest insect orders and families one species, larch casebearer, Coleophora laricella, is an important defoliator of larch, Larix spp. Gelechiidae, Leaf Miners The Gelechiidae are a large family of microlepidoptera and contain many species of agricultural importance. Members of the genus Coleotechnites are leaf miners of conifers. Several can reach epidemic levels and cause widespread damage.

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are held roof-like over the body. Larvae have a variety of feeding habits. They may feed on foliage and many species roll leaves with silken webbing and feed inside the rolled leaf. Others bore in buds, stems or fruiting structures of trees. When disturbed, larvae tend to move about rapidly in a twitching motion, presumably to ward off enemies. Many are important agricultural pests and some of the most damaging forest defoliators, shoot borers and cone insects are members of this family. Pyralidae, Snout Moths

Sesiidae, Clearwing Moths The Sessidae, or clearwing moths, consist of about 1000 known species. Adults are long legged, with a slender, dark body with bright red or yellow markings. The wings frequently lack scales and are transparent. Unlike other moths, the front and back wings are hooked together by a series of curved spines. Many species practice a form of protective mimicry and resemble wasps. Larvae are wood borers and several species are damaging pests of ornamental trees and forests.

The snout moths are so named because their mouthparts (labial palps) often project into a snout-like structure. This is a large family and larvae have a wide range of feeding habits. Larvae of many species feed on dry plant matter and are important pests of stored cereal grains. Cactoblastis cactorum feeds on prickly pear cactus, Opuntia spp., and has been introduced into Australia for biological control of this plant. Larvae of species of concern in forestry feed in the buds, shoots, stems and reproductive structures of host trees. Geometridae, Inchworms

Cossidae, Carpenter or Goat Moths The Cossidae consist of over 110 genera and about 700 known species. Adults tend to be nocturnal, are usually gray in color and have a wingspan ranging from 90 to 240 mm. Wing patterns and colors often mimic bark or leaves. Larvae are typically tree borers and up to 3 years may be required to complete a generation. The larvae of some species have an unpleasant odor and are called goat moths. Those that do not smell badly are edible. For example, larvae of Endoxyla leucomolcha, are known as “witchetty grubs” or “witjui grubs” and were an important food for Aboriginal women and children in central Australia. In central Chile, larvae of Chilecomadia moorei, known as butterworms, are harvested and used as pet food or fish bait. A few species are forest pests (Issacs 2002).

Tortricidae, Leaf Rollers and Budworms The Tortricidae are a large family of moths and consist of about 6300 described species. Adults are small, usually gray, tan or brown and may have dark bands or mottled areas on the wings. When at rest, the wings

The Geometridae are a large family of about 26,000 known species. The family’s name is derived from “geometer” or Earth measurer and refers to how larvae, which have prolegs on only the last two segments of their bodies, move about (Plate 12). They clasp with their front legs and draw up with the hind end, then clasp with the hind end (prolegs) and reach out for a new front attachment. This creates the impression that they are measuring their journey. Larvae are called inchworms, loopers, measuring worms or spanworms due to their characteristic loping gait. Many are forest defoliators. Lasiocampidae, Lappet Moths, Tent Caterpillars Known as eggars, snout moths, tent caterpillars or lappet moths, over 2000 species are known worldwide. Adults are large bodied with broad wings. Females lay many eggs, which are flat and either smooth or slightly pitted. In the case of tent caterpillars, eggs are deposited in masses and covered with a material that hardens in air. Adults are typically brown or gray, with hairy legs and bodies. Females are larger than males, but both sexes are similar in overall appearance. Mature larvae

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are large and hairy, especially on their sides. Larvae feed on foliage of many trees and shrubs. They feed gregariously during the early instars and often live in nests spun of silk. Thaumetopoeidae, Processionary Caterpillars This is a small family of medium sized, dull colored moths found in the Mediterranean Basin of southern Europe, North Africa and the Near East. Several species are important forest defoliators. Larvae of most species are gregarious and march in head to tail processions to new feeding and pupation sites. Mature larvae have urticating hairs that can cause severe skin irritation. Most authorities consider this family to contain a single genus, Thaumetopoea. However, some taxonomists have placed the species under three genera: Helianthocampa, Thaumetopoea and Tramatocampa (Halperin 1983, Doganlar & Ave 2001).

Saturniidae, Giant Silk Moths The saturniids include some of the world’s largest moths. Some have wingspans ranging from 150 to 250 mm. Many species are brightly colored and marked with translucent eyespots. The larvae are also large and many species are armed with conspicuous tubercles or spines. Most species pupate in silken cocoons. Larvae of several species feed on the foliage of forest trees, reach epidemic levels and cause extensive defoliation.

Sphingidae, Sphinx or Hawk Moths, Hornworms Sphinx or hawk moths are medium to large, heavybodied moths with long, narrow forewings. Some species may have a wingspan of up to 160 mm. Moths are typically strong fliers with a rapid wing beat. Some feed like hummingbirds, hover in front of flowers and extend their proboscis into the flower to gather nectar (Plate 13). Larvae have a conspicuous horn or spine at the dorsal surface of the 8th abdominal segment (Plate 14). They feed on the foliage of a variety of plants, including some of importance in agriculture and forestry.

Distribution is worldwide. Most species are robust and have drab forewings, although some, such as the underwing moths, may have brightly colored hind wings. Moths of most species are nocturnal and attracted to light. Larvae of several species live in soil, feed on bases of agricultural plants and are known as cutworms.

Subfamily Lymantriinae, Tussock Moths Moths are moderate sized and not colorful, having evolved to be camouflaged against the background of tree bark, lichens or leaves on which they alight. Adults have non-functional mouthparts and do not feed. Females of many species have either rudimentary or nonfunctional wings. Larvae are often strikingly colored and armed with clusters of long setae formed into hair brushes, hair pencils or radiating clusters protruding from raised warts. Some larvae have urticating hairs that cause allergic reactions. Larvae also have two medial dorsal glands on the sixth and seventh abdominal segments, which are often brightly colored in hues of red, orange or yellow. These characteristics often make larvae easier to identify than adults. Larvae are foliage feeders and several are major forest pests (Schaefer 1989, Lafontaine & Fibiger 2006). Arctiidae, Tiger Moths Moths of the family Arctiidae are known as tiger moths. They are small to medium stout-bodied moths with broad and often brightly colored wings with striking patterns of spots or stripes. They are largely nocturnal and, when at rest, hold their wings roof-like over their body. Larvae are robust and hairy. Those with uniformly dense hairs are called woolly bears and those with hairs in bristles and tufts are known as tussock moths. The North American woolly bear caterpillar, Pyrrharctia isabella, which has a brown band in the middle and black at each end, is a subject of American folklore. The amount of black on the larvae of the fall generation is thought to vary with the severity of the oncoming winter. Several species are forest defoliators.

Diptera, Flies Noctuidae, Owlet or Miller Moths The Noctuidae is a the largest family of Lepidoptera. It consists of about 35,000 species in 4200 genera.

The Diptera or flies are another large order of insects. They are characterized by having one pair of functional wings, i.e. the forewings. The hind wings are reduced to

Forest insect orders and families small, knobbed structures, referred to as halters. These function as organs of equilibrium. Metamorphosis is complete. The insects in this order occupy a wide range of habitats. Some are parasitoids and others feed on, and breed in, carrion. Still others, such as the mosquitoes, are vectors of diseases such as dengue, malaria and yellow fever. A few Diptera are plant pests and several are of concern in forestry. Cecidomyiidae, Gall Midges Gall midges are a large family of delicate flies with long legs and antennae and wings with reduced venation. They are small and usually range from 1 to 5 mm in

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length. Roughly two thirds of the species are gall makers. Others feed on plants without producing galls or live in decaying vegetation. Several species produce galls on trees.

Agromyzidae, Leaf-Miner Flies Members of the family Agromyzidae are small, yellow or black flies. Adults are usually 2–3 mm long. Approximately 2500 species are known worldwide. Larvae of most species are leaf miners that make serpentine or blotch mines in the leaves of host trees. Others infest stems and produce galls.

Chapter 7

Foliage Feeding Insects – Lepidoptera

INTRODUCTION Foliage feeders are the largest single group of damaging forest insects. Several thousand species feed on foliage of trees and other woody plants. Some species reach epidemic proportions and defoliate large areas of forest. Successive years of defoliation cause growth loss, branch dieback and tree mortality, especially when affected trees occur on poor soils or dry sites. Trees are usually not killed outright by defoliation but are weakened and have increased susceptibility to bark beetles and woodborers. Insect defoliation is known to be an inciting factor of forest decline. Insect feeding on foliage can take a variety of forms. The most common is external feeding by larvae and/or adults. All leaf tissue except the veins and midribs on broadleaf trees and entire needles of conifers are consumed. Other insects feed only on tender leaf tissue, leaving the veins. This gives damaged foliage a lacy appearance, known as skeletonizing. Other insects mine internally in foliage and cause dead spots and discoloration. Some leaf mining species feed inside the needles of conifers.

Many foliage feeding insects have evolved creative mechanisms to protect them from weather and natural enemies. Larvae of some species roll leaves, tie them with silken threads and feed inside the rolled leaf. Others build nests of webbing, bits of foliage, and/or frass. They live inside the nest, bag or case for some, or all, of their larval cycle. Larvae of many defoliators feed in colonies, especially during their early instars. Others are solitary feeders. This chapter addresses foliage feeding insects of the order Lepidoptera, the moths and butterflies. Foliage feeding insects of other orders of insects are addressed in Chapter 8.

Psychidae (Bagworms) Thyridopteryx ephemeraeformis (Haworth), Bagworm Distribution eastern USA.

This insect is indigenous to most of

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Hosts Hosts include several conifers and broadleaf trees. Favorite hosts are Thuja occidentalis and Juniperus virginiana. Other hosts include species of Acer, Picea, Pinus, Platanus occidentalis, Populus, Quercus, Robinia pseudoacacia, Salix, Taxodium distichum, Tilia and Ulmus.

Importance Bagworm is a pest of urban trees. Defoliation is unsightly and heavy damage can kill trees.

Life History There is one generation/year. Males emerge in autumn and fly to females to mate. Females remain in their bags and deposit eggs inside their pupal case. Eggs overwinter in the bags and hatch the following May–June. Newly hatched larvae spin bags of silk, mixed with bits of foliage and spent their entire larval period enclosed in this bag (Fig. 7.1). They feed on the surface of foliage at first, and later entire leaves are consumed. When mature, larvae attach their bags to twigs with silk and pupate.

Description of Stages Male moths are sooty black and have a wingspan of approximately 25 mm. Females are wingless and lack functional eyes, legs or antennae. They are maggot-like in appearance, yellow-white and nearly naked except for a circle of hairs at the posterior of the abdomen. Mature larvae are dark brown and 18–25 mm long. Head and thoracic plates are yellow, spotted with black.

Fig. 7.1 Bagworm, Thyridopteryx ephemeraeformis, on foliage of Taxodium distichum (southern Louisiana, USA).

Gracillariidae (Leaf Blotch Miners) Pest Management Infestations on ornamental trees and shrubs are difficult to control because they are often not detected until larvae are mature or have pupated. Bags containing larvae can be hand picked and destroyed. Applications of Bacillus thuringiensis or chemical insecticides are effective against heavy infestations provided they are detected early.

Related Species Giant or pawlonia bagworm, Clania variegata Snellen, is indigenous to Asia where larvae feed on foliage of several trees including pines, Acacia spp., Bischofia javanica, Pawlonia tomentosa and Pinus spp. (Drooz 1985, FAO 2007a).

Cameraria ohridella Deschka & Dimic, Horse-Chestnut Leaf Miner Distribution C. ohridella was first observed in 1985 by Lake Ohrid, Macedonia and was described as a new species in 1986. Its origin is still uncertain. In 1989, infestations appeared in Austria and spread throughout Europe, including Belgium, Denmark, Germany, the Netherlands, Poland, southern Sweden and central Europe. It has since spread west through France and south into Italy. Since 2002, it has been reported from Albania, Belarus, Moldova, western Russia, Spain, Turkey, UK and Ukraine.

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Fig. 7.2 Foliage of horse chestnut, Aesculus hippocastanum, damaged by horse chestnut leaf miner, Cameraria ohridella (Munich, Germany).

Hosts The host tree is horse chestnut, Aesculus hippocastanum.

Damage Larvae are leaf miners and make irregular blotch mines on the upper surface of leaves (Plate 1, Fig. 7.2). In Europe, horse chestnut is a popular ornamental and street tree. In Germany, this tree is often planted adjacent to outdoor restaurants and beer gardens. Leaf mining is unsightly, especially when infestations are heavy.

Life History C. ohridella has two to four generations/ year, depending on climatic conditions. Some pupae enter an extended diapause and may remain viable for up to three winters. Adults emerge in early morning and fly to tree trunks where they mate. A female produces 20–40 eggs. Eggs are deposited on the upper epidermis of leaves. Larvae have five feeding instars and complete development in about 3.5 weeks. Instar VI larvae do not feed but construct a cocoon made of silken

threads inside the mine. In summer, pupation is about 12–16 days and in winter pupae remain dormant for about 6 months.

Description of Stages The adult is a moth 6–7 mm long with a wingspan of 7.0–9.5 mm. The head has red hair and the antennae are almost as long as the forewing. Ground color of the thorax and forewings is red-brown. Forewings have a short white basal streak, two outwardly curved white narrow fasciae edged with black and two pairs of narrow costal and dorsal streaks. The apical part of the forewing is covered with rough blackish scales. Hind wings and abdomen are gray and legs are white with dark spots.

Pest Management Available tactics include removal of leaves containing pupae and either burning or composting them at high temperatures. It is critical to treat all foliage prior to April 1. The systemic insecticide imidacloprid can be applied to the boles of infested trees.

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Fig. 7.3 Leaf of quaking aspen, Populus tremuloides, with serpentine mines caused by aspen leaf miner, Phyllocnistis populiella (Roosevelt National Forest, Colorado, USA).

Related Species Two North American species of Cameraria, C. hamadryadella (Clemens), the solitary oak leaf miner, and C. cincinnatiella (Chambers), the gregarious oak leaf miner, cause blotch mines on foliage of white oaks including Quercus alba and Q. macrocarpa. They occur throughout eastern USA and occasional heavy infestations cause browning and premature dropping of foliage over large areas. Life history and habits are similar to C. ohridella (Deshka & Dimic, 1986, Solomon et al. 1999, Buszko 2006, Ivinskis & Rimðait€e 2006).

Phyllocnistis populiella Chambers, Aspen Leaf Miner Distribution This leaf miner has a transcontinental distribution across much of North America, including virtually all forested regions of Alaska, Canada, northeastern, north central and western USA.

Hosts

The host is quaking aspen, Populus tremuloides.

Importance Larvae construct serpentine mines in leaves of quaking aspen that turn brown (Fig. 7.3). Primary damage is cosmetic and mined foliage does not turn brilliant yellow-gold in autumn. Outbreaks have been reported from Alaska and western Canada since the 1950s and at least two outbreaks that exceeded 200,000 ha have occurred in interior Alaska since 1970. In 2000, aspen foliage discoloration was detected on over 120,000 ha. The outbreak expanded to over 266,000 ha in 2006. Outbreaks have also occurred in western USA.

Life History There is one generation/year. Adults overwinter under bark scales of both broadleaf trees and conifers. They emerge, feed and mate for about 2 weeks in early spring, just before buds break on host trees. Eggs are deposited singly on edges of newly emerged leaves and adults fold the leaf edge over the egg to form a protective shelter. Usually only one or two eggs are deposited per leaf but up to seven eggs per leaf have been observed during outbreaks. Larvae hatch, bore into the leaf and begin feeding within the epidermal

Foliage feeding insects – Lepidoptera tissue. They undergo five instars and, when feeding is completed, pupate inside leaf mines. Adults emerge from mines prior to leaf senescence in late August/ September and seek overwintering sites.

Description of Stages Adults are minute white moths with subtle brown or black markings on the lanceolate-shaped wings. Larvae are small, white and flat and about 5 mm long when mature.

Pest Management No effective direct control measures are available for this leaf miner (Kruse et al. 2007).

Coleophoridae (Casebearers) Coleophora laricella (Hübner), Larch Casebearer Distribution Larch casebearer is native to Europe and was introduced into North America about 1886. It was discovered in western USA in 1957.

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damage occurs. Pupation takes place in late May–early June.

Description of Stages Adults are silver to graybrown moths with no conspicuous markings. Their narrow wings are fringed with slender, hair-like scales. When at rest, with their wings folded, they are about 6 mm long. Mature larvae are about 6 mm long.

Pest Management In North America, classic biological control with two species of European parasitoids, Agathis pumila and Chrysocharis laricinellae, has helped reduce populations. Studies indicate that either species can parasitize over 90% of the population in an area. By 1981–2, the parasitoids, coupled with needle diseases and other mortality factors, reduced populations in western North America to a point where insects have been difficult to detect (Drooz 1985, Tunnock & Ryan 1995).

Gelechiidae (Leaf and Needle Miners) Coleotechnites

Hosts In Europe, Larix decidua is the host tree. In North America, L. laricina and L. occidentalis are hosts. All species of Larix are potential hosts.

Importance Successive and severe feeding, with up to 85–100% of the needles killed, can reduce growth potential by 95%, cause branch dieback, weaken trees and make them susceptible to root pathogens and wood boring insects.

Life History There is one generation/year. Adults emerge from pupal cases during late May–early July. Females deposit 50–70 eggs singly, usually on the undersides of needles. Larvae hatch in July and instars I–II mine inside needles. Between late August and midOctober, larvae convert mined needles into cases, in which instars III and IV live, feed and pupate. They attach their cases to needles using a pad of silk and continue to feed. In October, larvae attach their cases to twigs and spur shoots and overwinter (see Fig. 4.2). They resume feeding in spring when larch, a deciduous conifer, refoliates. This is when the most severe feeding

Coleotechnites is a genus of leaf mining caterpillars. Forty-nine species are recognized, all nearctic in distribution. Hosts include both broadleaf trees and conifers. Conifer infesting species are most damaging and several have caused extensive damage (Table 7.1). One species, C. piceaella (Kearfott), has been introduced into Europe and is now found in Austria, the Czech Republic, Germany, Great Britain, Hungary, Italy and Slovakia (Furniss & Carolin 1977, Hubrik 2007, Lee & Brown 2008). Coleotechnites milleri (Busck), Lodgepole Needle Miner Distribution Lodgepole needle miner occurs in California, west of the crest of the central and southern Sierra Nevada.

Hosts The host is lodgepole pine, Pinus contorta. During outbreaks, other pines such as red fir, Abies magnifica, and mountain hemlock, Tsuga mertensiana, may be attacked.

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Table 7.1 Representative species of Coleotechnites (Lepidoptera: Gelechiidae) in Eurasia and North America: their distribution and hosts. Species

Distribution

Hosts

C. apicitripunctella (Clemens)

Canada: Quebec USA: northeastern states USA: California

Taxodium distichum, Tsuga canadensis Pinus contorta

Canada: maritime Provinces to Alberta USA: Maine to Colorado Europe: introduced around 1962 USA: Colorado Canada: Alberta, British Columbia, Saskatchewan USA: Montana Canada: New Brunswick to Saskatchewan USA: northeastern states

Picea spp.

C. milleri (Busck) Lodgepole needle miner C. piceaella (Kearfott)

C. ponderosae Hodges & Stevens C. starki (Freeman)

C. thujaella (Kearfott)

Pinus ponderosa Pinus contorta

Thuja occidentalis

Sources: Furniss & Carolin 1977, Hodges & Stevens 1978, Drooz 1985, Hubrik 2007, Lee & Brown 2008.

Importance This needle miner has a history of long, sustained and destructive outbreaks in lodgepole pine forests in Yosemite National Park, California. Outbreaks occurred in 1903–21, 1933–41 and 1947–63. These caused extensive tree mortality resulting in a phenomenon known as a “ghost forest.” Outbreaks tend to be most severe in mature trees.

Life History Two seasons are required to complete a generation and adults are active during July and August of odd years. Females deposit eggs in small groups in needles hollowed out by previous generations of needle miner larvae. Instar I larvae mine into needle tips and overwinter. During the following season, they feed in the interior of several needles. In late summer early autumn, they become instar IV larvae and overwinter for a second winter. The fifth and final instar develops the following spring and causes considerable feeding damage. When feeding is completed, usually in June, larvae pupate inside mined needles.

Description of Stages Adults are mottled light gray moths with a wingspan of 8–13 mm and strongly fringed hind wings. Larvae are naked with black heads and body color ranging from uniform hues of lemon yellow to orange, pink and red.

Related Species An undescribed species of Coleotechnites, which resembles C. milleri, but has a 1-year life cycle, periodically defoliates lodgepole pine in central Oregon, USA. Coleotechnites starki (Freeman) occurs in the Canadian Rockies and Montana and is similar to C. milleri in life history and appearance. C. ponderosae Hodges & Stevens, damages ponderosa pine in Colorado. Localized outbreaks occur in ponderosa pine forests and cause a yellow discoloration of the needles and premature needle fall (Stark 1954, Koerber & Struble 1971, Mason & Tigner 1972, Stevens et al. 1996).

Tortricidae (Budworms and Leaf Rollers) Choristoneura Choristoneura consists of about 39 species worldwide, distributed over Africa, Asia, Europe and North America (Baixeras et al. 2008). Hosts are either conifers or broadleaf trees and several are important defoliators (Table 7.2). Two North American species, spruce budworm, C. fumiferana, and western spruce budworm, C. occidentalis, have defoliated millions of hectares of conifer forests in North America and have been the target of large suppression projects using aerial applications of biological and chemical insecticides. The life history of most species of Choristoneura is similar (Fig. 7.4). There is one generation/year. Adults

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Table 7.2 Representative species of Choristoneura (Lepidoptera: Tortricidae): their distribution and hosts. Species

Distribution

Hosts

C. biennis Freeman 2-year cycle budworm C. carana carana Barnes and Busck C. carana californicum Powell

Canada: British Columbia USA: Yukon Territory USA: California, San Gabriel Mountains USA: California, central Sierra Nevada, southern Cascades and northern Coast Range Eastern Canada and northeastern USA west to Alaska and south to Arizona and New Mexico Central and southern Europe east to the Near East, Mongolia, China and Japan Eastern Canada and USA west to British Columbia, Yukon Territory and Alaska USA: northern California, southern Oregon Western Canada and USA Western USA Canada: British Columbia USA: Oregon, Washington Europe and Near East

Abies lasiocarpa, Picea engelmannii, P. glauca Pseudotsuga macrocarpa Ps. menziesii

C. conflictana (Walker) Large aspen tortrix C. diversana Hu¨bner

C. fumiferana (Clemens) Spruce budworm C. lambertiana lambertiana (Busck) C. lambertiana subretiniana Obraztsov C. lambertiana ponderosana Obraztsov C. lambertiana (uncertain status) Western pine tortix C. murinana (Hu¨bner) Fir budworm C. occidentalis Freeman Western spruce budworm

C. pinus pinus Freeman Jack pine budworm

C. orea Freeman C. retiniana (Walsingham) Modoc budworm

Populus tremuloides, P. grandidentata Various broadleaf trees Abies sachalinensis (Japan) Abies balsamea, Picea glauca, P. rubens, P. mariana Pinus lambertiana P. contorta P. ponderosa, P. flexilis P. contorta, P. sylvestris Abies

Alberta and British Columbia, Canada south to northern California, Arizona and New Mexico, USA

Abies, Larix, Picea, Pseudotsuga, occasionally Pinus

Canada: maritime provinces east to Saskatchewan and eastern Alberta USA: Michigan, Minnesota, Wisconsin and Nebraska (plantations only) Canada: north coast of British Columbia USA: Alaska USA: northeastern California and south central Oregon

Pinus banksiana, P. resinosa

Abies amabilis, Picea sitchensis Abies concolor

Sources: Kamijo 1973, Powell 1995, Sarikaya & Avci 2006, Ovsyannikova & Grichanov 2009.

are active in early to late summer and deposit eggs in masses on the foliage of host trees. Eggs hatch in 10–14 days. Larvae pass through six instars. For conifer feeding species, instar I larvae do not feed. They seek a shelter in bark crevices or under lichen, construct a hibernaculum, molt and overwinter. Species that feed on broadleaf trees may skeletonize foliage prior to

overwintering. Larvae become active in spring as buds of host plants expand. Conifer feeding species may mine some needles, then bore into and destroy expanding buds. Later instar larvae feed on new foliage and shoots. Some larvae spin down on silken threads and may be dispersed by air currents. Several broadleaf feeding species (e.g. C. conflictana) are leaf rollers. Feeding is

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Forest entomology: a global perspective it is reportedly a pest of Abies sachalinensis and outbreaks have been reported on that species since 1965.

Eggs

Larvae (overwintering)

Importance C. diversana is considered an important pest of orchards, parks and forests. Outbreaks are most common in floodplains.

Larvae (active)

Pupae

Adults

J

F

M

A

M

J

J

A

S

O

N

D

Fig. 7.4 Generalized life cycle of budworms and leaf rollers of the genus Choristoneura.

usually completed by mid-summer and instar VI larvae tie tips of twigs or leaves with silk, form a nest and pupate. Pupation lasts about 10–15 days. Adults emerge, mate and lay eggs. Their life span ranges from a few days to about 1 week. Adults of at least one species, C. fumiferana, are known to travel, on their own and/or via prevailing winds, for long distances. Longdistance movement of adults by storm systems is a considered major dispersal factor. Adults can cover up to 450 km in one night. The one notable exception to this generalized life cycle is C. biennis, the 2-year cycle budworm, which requires 2 years to complete a generation. The second winter is passed as instar IV larvae (Greenbank et al. 1980, Kucera & Orr 1981, Fellin & Dewey 1982, Dobersberger et al. 1983, Duncan 2007, Ciesla & Kruse 2009). Choristoneura diversana (Hübner) Distribution This species ranges from central and southern Europe east across Russia, the Near East to Mongolia, northern China, Korea and Japan.

Hosts Throughout most of its range, larvae feed on various broadleaf plants including fruit trees, grasses, low woody plants and forest trees. Genera of plants that serve as hosts include Alnus, Betula, Fagus, Fraxinus, Lespedeza, Malus, Populus, Prunus, Quercus, Rhamnus, Rhododendron, Salix and Ulmus. In Japan,

Description of Stages Adults have a wingspan of 15–20 mm for males and 19–23 mm for females. Background wing color of males is brown with areas of yellow or gray. Wing pattern on forewings is darker than the background, usually brown. Hind wings are gray. Females are olive brown in color with a more pronounced wing pattern. Larvae range in color from light-green to brown-green. Head capsule varies in color from brown to red-brown.

Pest Management Direct control of infestations is accomplished by removal of infested branches at low population levels. During outbreaks, biological or chemical insecticides are applied in spring after blossoming of fruit trees (Kamijo 1973, Ovsyannikova & Grichanov 2009).

Choristoneura fumiferana (Clemens), Spruce Budworm Distribution Spruce budworm has a transcontinental distribution across North America from Nova Scotia, Canada, south to portions of the northeastern and north central states of the USA, across the boreal forests of Canada and west to Alaska (Fig. 7.5).

Hosts Abies balsamea is the species most severely damaged in eastern North America. Spruces, Picea glauca, P. mariana and P. rubens, are also hosts and, during outbreaks, some feeding can occur on associated conifers. Spruce mixed with balsam fir is more likely to suffer damage than spruce in pure stands. Further west, beyond the natural range of A. balsamea, P. glauca is the favorite host.

Foliage feeding insects – Lepidoptera

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Fig. 7.5 Distribution of two species of Choristoneura (Lepidoptera, Tortricidae), spruce budworm, C. fumiferana, and western spruce budworm, C. occidentalis, in North America (redrawn from Fellin & Dewey 1982 and Natural Resources Canada 2007).

Importance Larvae feed in expanding buds, male flowers and on new foliage. Spruce budworm is considered the most destructive forest defoliator of boreal and sub-boreal spruce–fir forests in Canada and the USA. Outbreaks often cover millions of hectares and persist for 8–10 years. Outbreaks are associated with maturing of balsam fir and increased staminate flower production, which are a favorite food of young larvae. Successive years of bud damage and defoliation cause growth loss, top kill and tree mortality.

Description of Stages Adults have a wingspan of about 20 mm, are gray with dark brown markings or red-brown with gray markings. Eggs are light green, 1 mm long and 0.2 mm wide and laid in elongate

masses of 2–60 eggs that overlap. Instar I larvae are about 2 mm long with a yellow-green body and brown head. Instar II larvae are yellow with a dark brown or black head. During the next four instars, larval body color changes from pale yellow to dark brown with light-colored spots along the back. Instar VI larvae average 25 mm long with a dark brown-black head. Pupae are pale green at first and later change to redbrown, marked with dark bands and spots.

Pest Management Reduction of the proportion of Abies balsamea, especially in mature forests, can lower susceptibility to attack. Outbreaks have been treated with aerial applications of both chemical and biological insecticides. Spruce budworm has been the target of

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some of the largest aerial campaigns directed against a forest defoliator, especially in eastern Canada and Maine, USA (Kucera & Orr 1981).

Choristoneura murinana (Hübner), Fir Budworm Distribution This budworm occurs over much of central and southeastern Europe and the Near East.

Hosts Primary hosts are species of Abies. In Europe, Abies alba is the major host. In 2002, infestations were reported for the first time in forests of A. cilicia in Turkey. Other hosts are species of Cedrus, Juniperus, Picea and Pinus.

Importance Larvae feed on needles, buds and shoots of host trees. Heavy defoliation weakens trees and increases susceptibility to attack by bark beetles such as Pityokteines curvidens and Cryphalus piceae. Outbreaks have occurred in fir forests across Europe and Turkey.

Description of Stages Average wingspan of adult moths is 22.8 mm for females and 18.5 mm for males. Forewings are gray and yellow with variable brown markings and a broad red-brown transverse band. Hind wings are gray-brown with yellow tips. Newly hatched larvae are yellow-green with a brown head capsule and are about 1.5 mm long. Mature larvae are 16–22 mm long, have a shiny black head and a pale gray-green body color with lighter colored lateral and ventral segments. They have an orange anal shield and are covered with tubercles with black hairs (Canadian Food Inspection Agency 2006, Kimoto & Duthie-Holt 2006, Sarikaya & Avci 2006).

Choristoneura occidentalis Freeman, Western Spruce Budworm Distribution Western spruce budworm is the most widely distributed conifer defoliator in western North America. It ranges from Alberta and British Columbia, Canada south into western USA from Washington, Idaho and Montana to northern California, Arizona and New Mexico (Plates 15–18, Fig. 7.5).

Hosts Preferred hosts are Abies grandis, A. concolor, A. lasiocarpa, Larix occidentalis, Picea engelmannii, P. glauca, Pseudotsuga menziesii and P. pungens. Larvae also feed on Abies amabilis, Pinus albicaulis, P. contorta var. latifolia, P. flexilis, P. ponderosa, P. monticola, Tsuga mertensiana and T. heterophylla. Importance Damage is similar to that of spruce budworm except on western larch where larvae mine and sever the terminal and lateral shoots. Outbreaks can cover large areas and persist for over 10 years. No typical pattern or trend in epidemics is apparent. Most early epidemics lasted a few years and subsided. An epidemic in the northern Rocky Mountains began in 1949, persisted until 1994, and defoliated over about 1.2 million ha annually. In the Pacific Northwest, two outbreak cycles have occurred since 1947, i.e. from 1948 to 1963 and from 1971 to 1997. Outbreaks have also occurred in British Columbia, Canada. Defoliation causes growth loss, top kill and tree mortality and damaged trees are susceptible to bark beetle attack. In addition to foliage, larvae feed on staminate flowers and developing cones of host trees. Decline in seed production has a serious impact in seed orchards, seed production areas, and forest sites that are difficult to regenerate naturally. Description of Stages Adults are about 12.7 mm long with a wingspan of 22–28 mm. Both sexes are similar in appearance, although the females are more robust. Forewings are gray or orange-brown, banded or streaked, and some have a conspicuous white dot on the wing margin. Eggs are oval, light green, about 1.2 mm long and overlap like shingles. Young larvae are yellow-green with brown heads. Instar II–IV larvae have black heads and thoracic shields and orange or cinnamon-brown bodies. Instar V larvae have reddishbrown heads marked with black triangles, black thoracic shields and pale olive-brown bodies marked with small white spots. Instar VI larvae are 25–32 mm long, with tan or light chestnut-brown heads and thoracic shields and olive or red-brown bodies with large ivorycolored areas. Pupae are 13–16 mm long, broad at the head and narrower toward the tail. They are initially yellow-brown or green-brown and later red-brown. Pest Management Cultural tactics include reduction of the portion of host trees, especially Abies spp., in

Foliage feeding insects – Lepidoptera forests and favoring non-host trees such as Pinus ponderosa. Outbreaks can be treated with aerial or ground applications of either chemical insecticides or Bacillus thuringiensis (Dewey 1970, Fellin & Dewey 1982, Ciesla & Mason 2005).

Tortrix viridana Linnaeus, Green Oak Tortrix Distribution Green oak tortrix occurs throughout Europe, including the British Isles, north Africa and the Near East including Cyprus, Iran and Israel.

Hosts Primary hosts are oaks, Quercus spp. Other hosts are species of Acer, Betula, Carpinus, Fagus and Populus. Green oak tortrix also feeds on shrubs, including species of Urtica and Vaccinium.

Importance This insect is a defoliator of oak forests and larvae prefer upper crowns of host trees. Successive defoliation causes growth loss, weakens trees and predisposes them to attack by other organisms. Green oak tortrix has been implicated as one of several inciting factors in European oak decline.

Life History Green oak tortrix has one generation/ year. Adults are active from late June–early August. Females deposit 50–60 eggs in cement-like masses on branches, leaves and branch forks, throughout the crown. Eggs hatch the following spring and larvae first bore into opening buds and later feed on foliage and flowers. They roll leaves with silken threads and feed within the rolled leaf. Late-instar larvae may also feed on the bark of new shoots. Pupation occurs in late spring within rolled leaves.

Description of Stages Adults have pale green or yellow-green forewings with brown-gray to gray hind wings. Both wings have a frayed outer edge. Wingspan is 18–23 mm. Head is yellowish and the body is gray. Eggs are round, 0.7 mm in diameter, initially light yellow in color, changing to brown. Young larvae are gray with dark heads. Mature larvae are gray-green and 15–19 mm long (Browne 1968, Canadian Food Inspection Agency 2010).

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Zeiraphera diniana Guen ee, Gray Larch Bud Moth Distribution This insect occurs in the Alps of central Europe, including portions of France, Italy and Switzerland. It has been introduced into the UK.

Hosts Primary host is European larch, Larix decidua. During outbreaks, other conifers, including Pinus cembra and P. sylvestris, are also damaged. Populations introduced into the UK have adapted to exotic plantations of P. contorta. Several “host races” of this insect have been identified. One has a host preference for larch and another for pines.

Importance Gray larch bud moth is an example of an insect that undergoes outbreak cycles at regular intervals. Outbreaks have occurred on average every 9 years in high mountain valleys of Switzerland at elevations above 1800 m. Outbreaks cause widespread defoliation and discoloration of larch forests in areas that are important for tourism. Defoliation seldom kills trees. Since 1981, outbreak cycles have declined in intensity.

Life History This tortricid has one generation/year. Adults tend to be nocturnal, are active from July to October, depending on elevation, and live for about 30 days. Period of egg deposition at any one location is about 20 days. Most females deposit from 20 to 80 eggs, although some may produce up to 350 eggs. Eggs are deposited in small masses under lichen on twigs and branches or between cone scales of host trees. Diapausing embryos in the eggs overwinter. Larvae hatch from May to June, about the time of bud burst on larch, and begin to feed. Instar IV larvae live inside needle cases that consist of several needles webbed together with silk and feed on the foliage. Larval instar V is the most destructive and lasts for 10–14 days. When feeding is completed, larvae drop to the litter and pupate. Pupation lasts from 25 to 36 days.

Description of Stages Adults have a wingspan of between 18 and 20 mm. Forewings are gray-brown with irregular red-brown markings. Instar IV larvae are yellow with orange head capsules and instar V larvae are gray-purple with black head capsules. Mature

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larvae of the larch feeding host race tend to be darker in color than pine feeding forms. Pest Management Aerial application of chemical pesticides was used to suppress outbreaks during the years following World War II.

Related Species Zeiraphera improbana (Walker) occurs in the Rocky Mountains of western Canada and the USA, where it feeds on foliage of western larch, Larix occidentalis. A larch bud moth, Zeiraphera sp., has been reported from L. laricina in the Alaskan interior, USA and 240,000 ha were defoliated during 1975–6 (Werner 1980, Baltensweiler & Fischlin 1988, Abgrall & Soutrenon 1991, Baltensweiler 1998, Hagle et al. 2003).

Geometridae, Loopers and Inchworms Many Geometridae are forest defoliators and some are important forest pests. Individual species feed on either broadleaf trees or conifers (Table 7.3). Erannis tiliaria (Harris), Linden Looper Distribution Linden looper is a North American geometrid found from eastern Canada and the USA west to Alberta and south to Missouri and Utah.

April–May and larvae feed until July, then pupate in soil and litter. Adults are active from October to December. Females crawl up the boles of trees, mate and deposit eggs singly or in small groups in bark crevices. Description of Stages Female adults are wingless, light gray to brown and about 12 mm long. Males have fully developed wings with an average wingspan of 42 mm. Forewings are buff and marked with two transverse wavy brown bands and a sprinkling of brown spots. Mature larvae are about 37 mm long and have rusty brown heads. Body color may be entirely yellow or yellow with a series of 10 dark wavy longitudinal lines on the dorsal surface.

Related Species Mottled umber moth, E. defoliara (Harris), is native to Europe and the Near East, where it feeds on a variety of broadleaf trees. Jacobson’s spanworm E. jacobsoni (Djakonoff) is native to Mongolia and Russia and feeds on Larix (Browne 1968, Drooz 1985, Epova & Pleshanov 1995).

Evita hyalinaria Blandaria (Dyar) Distribution This geometrid is native to central Mexico and is known from the states of Hidalgo, Mexico, Michoac an and Puebla.

Hosts This insect is a defoliator of broadleaf trees including species of Acer, Betula, Carya, Malus, Quercus, Tilia and Ulmus.

Hosts Primary host is sacred fir, Abies religiosa. During outbreaks it may also occur on species of Pinus, Prunus and Quercus.

Importance Linden looper occasionally reaches epidemic levels, often in association with other Geometridae including spring and fall cankerworms, Paleacrita vernata (Peck) and Alsophila pometaria (Harris), respectively, elm spanworm, Ennomos subsignarius (Hübner) and Phigalea titea (Cramer). Outbreaks can cause defoliation for several years but are usually short lived.

Importance E. hyalinaria blandaria is a defoliator of A. religiosa forests in central Mexico. Defoliation typically occurs from lower to upper branches. During outbreaks, trees may be stripped of their foliage, causing growth loss and tree mortality. Mature forests are most susceptible to outbreaks although young trees are also defoliated.

Life History Linden looper has one generation/year and winter is spent in the egg stage. Eggs hatch during

Life History Typically, there is one generation/year although two generations may occur at low elevations. Adults are active in May and during outbreaks,

Foliage feeding insects – Lepidoptera

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Table 7.3 Representative species of forest defoliating Geometridae: their distribution and hosts. Species

Distribution

Hosts

Alsophila pometaria (Harris) Fall cankerworm

Eastern North America west to Alberta, Canada and California, Colorado, Montana, USA Europe

Acer, Betula, Populus, Quercus, Salix,Ulmus

Bupalus piniaria Linnaeus Pine looper Erannis defoliaria (Harris) Mottled umber moth E. jacobsoni (Djakonoff) Jacobson’s spanworm E. tiliaria (Harris) Linden looper Epirrita autumnata (Borkhausen) Autumnal moth Evita hyalinara blandaria (Dyar) Glena bisulca Rindge Lambdina fiscellaria fiscellaria e) (Guene Hemlock looper L. fiscellaria lugubrosa (Hulst) Western hemlock looper Operopthera bruceata (Hulst) Bruce spanworm O. brumata (Linnaeus) Winter moth Nepytia freemani Munroe False hemlock looper N. janetae Ringe

Pinus sylvestris, Pinus spp., Larix decidua, Pseudotsuga menziesii Betula, Fagus, Quercus, Ulmus and other broadleaf trees

Europe including the British Isles, Canada. Introduced into British Columbia Asia: Russia, Mongolia.

Larix gmelinii, L. sibirica

Canada: Nova Scotia to Alberta USA: eastern states west to Utah Northern and central Europe

Acer, Betula, Carya, Malus, Quercus, Tilia, Ulmus Alnus, Betula

Central Mexico South America: Colombia Eastern North America

Abies religiosa Cupressus lusitanica, Pinus spp. Tsuga canadensis, other conifers

Canada: British Columbia USA: Alaska, Oregon, Washington Canada: coast to coast USA: northeastern, north central states Europe, Asia, Near East North America: introduced Canada, USA: western provinces and states USA: Arizona, New Mexico

Tsuga heterophylla, other conifers Acer, Alnus, Amalanchier, Betula, Fagus, Prunus, Populus, Quercus, Salix Acer, Betula, Malus, Populus, Prunus, Quercus, Salix, Tilia, Ulmus, Vaccinium Pseudotsuga menziesii Abies lasiocarpa, Picea engelmannii, other conifers

Sources: Browne 1968, Drooz & Bustliio 1972, Furniss & Carolin 1977, Drooz 1985, Tenow et al. 2004,2007, Fairweather et al. 2006.

extensive moth flights can obscure visibility. Eggs are laid in small masses on fir foliage, although during outbreaks they may also be deposited on foliage of other trees and shrubs. Eggs hatch in July, young larvae feed on the undersides of needles until autumn and defoliation is not conspicuous. During winter, larvae remain at rest on the foliage and resume feeding in April of the following year. Pupation occurs in cocoons on branches of trees, shrubs and blades of grass.

wing. Eggs are barrel-shaped, 1 mm in diameter, graygreen, later changing to pale yellow. Larvae are inchworms with prolegs on the sixth and tenth abdominal segments. Head is light brown with dark punctures and slightly broader than the body. Body color ranges from pale gray-brown to gray with two darker subdorsal bands (Fig. 7.7). Pupae are brown and about 21 mm long (Cibri an Tovar et al. 1995). Glena bisulca Rindge

Description of Stages Adults are fragile moths with a wingspan of 26–31 mm. Body and wings are covered with yellow-white scales (Fig. 7.6). Two medial lines occur on the forewings that continue to the edge of the

Distribution This species occurs in portions of Colombia and is one of a complex of indigenous defoliators in areas where extensive plantations of exotic conifers have been established.

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Fig. 7.6 Adult the geometrid, Evita hyalinaria blandaria (Mexico State, Mexico).

Fig. 7.7 Mature larva of Evita hyalinaria blandaria (Mexico State, Mexico).

Foliage feeding insects – Lepidoptera Hosts Primary host is Cupressus lusitanica, which has been planted extensively in the Department of Antioquia. It also feeds on Pinus patula.

Damage Outbreaks cause heavy defoliation and C. lusitanica is sensitive to even a single defoliation. In Colombia, outbreaks cause three periods of defoliation/ year and can cause extensive tree mortality.

Life History There are three generations/year and the period from adult to adult is about 17 weeks. Adult activity varies by location and prolonged emergence and oviposition causes overlapping of life stages and larval instars. Adults spend considerable time at rest on basal portions of tree boles with wings outstretched and antennae turned under the wings. Females lay eggs in bark crevices of host trees. Each female deposits over 300 eggs, which hatch in 11–12 days. Larvae undergo five to six instars with females tending to have the greater number of instars. They are wasteful feeders that notch or cut through the foliage they eat, large quantities of which fall to the ground. Larvae feed 33–35 days, shorten and drop to the litter to pupate. Pupation lasts 16–22 days. Moths emerge and move to boles of trees where they expand their wings.

Description of Stages Both males and females have a white background with a scattering of dark spots. Females are slightly darker. Wingspan is 40–50 mm. Male antennae are pectinate and female antennae are filiform. Eggs are light green, oval and turn red within 3 days. Eggs are 0.45  0.84 mm. Larvae are about 3 mm long when they hatch and 40–45 mm when mature. Bodies are tan at first but develop brown or red markings. Pupae are brown and glossy.

Pest Management Establishment of pine or eucalypt plantations instead of cypress in areas that are most susceptible to defoliation can prevent damage. Aerial application of pesticides is dangerous because of steep terrain in the affected areas and frequent low cloud cover. Classic biological control, using an egg parasite, Telenomus alsophilae, of the North American geometrid Alsophila pometaria has been undertaken

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(Drooz & Bustliio 1972, Bustliio & Drooz 1977, Rodas P. 1998).

Lasiocampidae, Lappet Moths, Tent Caterpillars Dendrolimus Dendrolimus is a large genus of conifer feeding Lepidoptera, several of which can reach epidemic levels and cause severe defoliation. All are Asian except for D. pini, which is found in Europe (Table 7.4). Dendrolimus pini Linnaeus Distribution D. pini is distributed across Europe, except the UK and western portions of Asia. Hosts Larvae feed on foliage of pines. Pinus sylvestris is the preferred host.

Importance Larvae defoliate pine forests and, during outbreaks, also feed on bark of young shoots after all foliage is consumed. In Poland, D. pini prefers 30–60year-old forests on poor sites, especially those suffering from moisture stress.

Life History This species normally has one generation/year but some individuals take 2 years to complete their life cycle. Adults are present from late June to early August and fly after sunset. After mating, females lay 150–300 eggs on needles, branches and boles of host trees in clusters of up to 100 eggs. Egg hatch in about 2 weeks. Larvae first feed on remains of their eggs and then move to pine needles. In mid-October–early November, after the first killing frosts, larvae migrate to the litter to overwinter. The following spring, they climb back into crowns and resume feeding. Pupation occurs from late June to August in yellowgray cocoons long attached to bark crevices, needles and branches.

Description of Stages Adults are light gray to brown-black. Females have a wingspan of 80 mm, males 60 mm. Forewings have a small white spot and a brown band. Mature larvae are 50–89 mm long, gray

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Table 7.4 Representative species of Dendrolimus (Lepidoptera: Lasiocampidae): their distribution and hosts. Species

Distribution

Hosts

D. houi Lajonquiere Yunnan pine caterpillar D. kikuchii Matsumara D. pini Linnaeus D. punctatus Walker Pine caterpillar D, sibiricus Tschetwerikov Siberian silk moth D. spectabilis Walker

Southwestern China

Pinus

China, Taiwan Europe, western Asia Southern China, Taiwan, Vietnam

Pinus Pinus sylvestris Pinus elliottii , P. merkusii, P. massoniana, P. taeda Abies, Larix, Picea, Pinus, Tsuga

D. tabulaeformis Tsau et Lui

Russia, except extreme northern areas, Kazakhstan, northern Mongolia, northern China, North and South Korea China, South Korea, Japan, southeastern Siberia China

Pinus densiflora, P. taeda, P. tabulaeformis, P. thunbergeriana, P. strobus P. armandi, P. massoniana, P. tabulaeformis, P. thunbergiana, 

 Indicates an exotic host. Sources: Dao Xuan Troung 1990, Zhang Zhi-Zhong 1990, Vorontsov 1995, Kolk & Starzyk 1996, Zhang Chao-ju 2002.

to brown, hairy with steel-blue hairless bands on the second and third abdominal segments and a V-shaped spot on the eighth abdominal segment (Plate 19). Eggs are oval, 2 mm long and gray or brownish-gray in color. Pest Management Mixed species stands are encouraged in areas susceptible to repeated defoliation. Predaceous ants and insectivorous birds are protected to encourage natural control. Aerial and ground applications of chemical and biological insecticides are used to treat outbreaks (Kolk & Starzyk 1996). Dendrolimus punctatus Walker, Pine Caterpillar Distribution D. punctatus occurs in southern China, Taiwan and Vietnam.

Hosts Larvae feed on foliage of pines, Pinus spp. In China, the primary host is Mason pine, Pinus massoniana. Slash pine, P. elliotii, and loblolly pine, P. taeda, native to southeastern USA and widely planted in central and southern China, are also attacked. In Vietnam, P. massoniana and P. merkusii are hosts.

Importance Pine caterpillar is an important pest of pine in China and Vietnam. Outbreaks are common in young plantations and pure natural stands ranging in age from 8 to 15 years. Older needles are preferred. The major impact in young plantations is growth loss. Tree mortality is minor, although trees weakened by defoliation could become susceptible to attack by secondary insects. Larvae have urticating hairs that cause skin rash and eye irritation in some individuals.

Life History The number of generations/year increases in the southern part of this insect’s range. In Hunan Province, China, there are two complete generations/year and a partial third. In Vietnam, there are four generations/year and larvae are present from March to May, June to July, August to September and October to March. Females lay 300–400 eggs, with a maximum of 800 eggs, in groups on pine needles. Larvae feed on needles and pass through six instars. Pupation occurs in cocoons attached to needles and small branches. Winter is usually spent as diapausing larvae. Natural control is brought about by several parasitoids and diseases. In Vietnam, they include egg parasitoids, Telenomus sp. and Anastus sp. Larvae and pupae are killed by the fungus, Beauvaria bassiana, and virus diseases. Larval parasitoids include Pimpla spp. and a fly of the family Tachinidae.

Foliage feeding insects – Lepidoptera

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Fig. 7.8 Eggs of Dendrolimus punctatus on foliage of Pinus merkusii (Vinh, Vietnam).

Description of Stages Adults have a wingspan of 50–80 mm with females slightly larger than males. Wings are medium dull gray-brown and forewings have two dark lines. Mature larvae are 55–70 mm long, abdominal and thoracic segments have alternating light gray and black bands. The black bands contain orange markings. Larvae are covered with fine hairs (Plate 20). Eggs are rose to light brown in color and deposited in rows on pine needles (Fig. 7.8).

Pest Management Prohibiting livestock grazing in pine plantations is done to encourage growth of flowering plants. This is believed to increase nectar for adult parasitic wasps and increase survival rates. Direct control includes application of chemical insecticides, Bacillus thuringiensis and the fungus, Beauvaria bassiana. In addition, all life stages may be hand picked from trees and adults caught in light traps. Large firecrackers, with some gunpowder removed and replaced with fungal spores, have been used to deploy B. bassiana (Browne 1968, Dao Xuan Troung 1990).

Dendrolimus sibiricus Tschetwerikov (¼ D. superans sibiricus Tschetwerikov), Siberian Silk Moth Distribution D. sibiricus is indigenous to northern China, Kazakhstan, North and South Korea, northern Mongolia, and Russia, except extreme northern areas. It apparently is spreading westward into European Russia.

Hosts Larvae feed on conifers of the genera Abies, Larix, Picea, Pinus and Tsuga.

Importance This insect is an important defoliator of conifers in Russia and Kazakhstan and one of the most important defoliators of larch in China. Outbreaks can occur over thousands of hectares and lead to tree mortality or predispose trees to attack by secondary insects. Over a 90-year period (1855–1945), Siberian silk moth killed about 4 million ha of Russian forests. From 1932 to 1957, this insect defoliated 7 million ha

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of forests in western Siberia and China and caused tree mortality on over half of this area. Outbreaks occur with a periodicity of 10–11 years and are often preceded by 2–3 years of dry weather.

countries where this insect is an important pest (Florov 1948, Kolomiets 1958, Epova & Pleshanov 1995, Vorontsov 1995, Yang & Gu 1995, Canadian Food Inspection Service 2006). Malacosoma

Life History The number of generations varies with location. In most areas, 2 years are required to complete a generation. Further north, 3 years may be required and in southern areas a generation can be completed in 1 year. Adults are usually active in mid-July. After mating, females lay eggs in the needles in the lower crown. Eggs are laid in masses of up to 200 eggs. A female typically lays 200–300 eggs but some can lay up to 800 eggs. During outbreaks, eggs may be laid throughout the tree or on the ground. Eggs usually hatch within 13–15 days (maximum 20–22 days). Larvae have 6–8 instars. Instar I larvae feed on edges of needles. Instars II and III consume more foliage and in September, instar III larvae migrate to the soil and litter to overwinter. The following April, larvae return to the crowns and continue to feed on foliage, bark of young shoots and cones. In autumn, they again migrate to the soil to overwinter a second time. In May–June of the following year, larvae emerge and resume feeding. During this period, they consume about 95% of the food they need for development and major defoliation occurs. Pupation occurs in cocoons in June and adults emerge in July.

Description of Stages Adults are yellow-brown, light gray, dark brown or almost black. Front wings have two dark transverse bands and a white spot in the center. Hind wings are the same color as the forewings but lack markings. Females are slightly larger than males, about 40 mm long, with a wing span of 60–80 mm. Males are about 30 mm long with a wing span of 40–60 mm. Larvae are black to dark brown with numerous spots and long reddish hairs. The second and third abdominal segments are marked with blue-black stripes. Mature larvae are 55–70 mm long. Eggs are oval, 2.2 mm long and 1.9 mm wide, initially light-green but turn creamy-white, then darken and become spotted.

Pest Management Direct control with aerial applications of chemical and biological insecticides is conducted by the Federal Forest Service of Russia and other

Malacosoma consists of 26 species worldwide. Six are native to North America, the remainder are Eurasian (Table 7.5). Several species can reach outbreak levels, strip host plants of their foliage, then wander across open ground in search of additional food sources, including plants not normally fed upon. The life history and habits of all species are similar (Fig. 7.9). There is one generation/year. Adults are active in mid-summer and lay a single egg mass on young branches of host trees. Egg hatch occurs shortly after they are laid but larvae remain in the egg and winter is spent as diapausing larvae. Larvae appear in early spring as soon as buds break, usually construct silken tents for shelter from weather and for molting and feed on foliage. They feed in colonies and gradually expand the tent as the larvae grow. Pupation occurs in silken cocoons on the bark of trees, fence, brush and weeds. Malacosoma californicum (Packard), Western Tent Caterpillar (Plates 21–23) Distribution Western tent caterpillar occurs across western North America, east to western Quebec and north to Alaska. There are six recognized subspecies and a highly variable “central population.” Each occupies a well-defined geographic area. This species also occurs in three states of northern Mexico. Hosts Western tent caterpillar has a broad host range and includes both trees and low woody plants. Tree hosts include species of Alnus, Arbutus, Betula, Corylus, Crataegus, Fraxinus, Malus, Quercus, Populus, Prunus and Salix. Woody shrub hosts include Ceanothus spp., Cercocarpus ledifolius, Purshia tridentata, Physocarpus spp., Rhus trilobata, Ribes spp. and Rosa spp. Importance Outbreaks can cause complete stripping of trees resulting in growth loss, branch dieback and top kill, reduced fruit production and, in some cases, tree mortality. Heavy infestations on Purshia tridentata in eastern Oregon and northern California have killed large areas of this plant, which is an important browse

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Table 7.5 Representative species of Malacosoma (Lepidoptera: Lasiocampidae): their distribution and hosts. Species

Distribution

Principal hosts

M. americanum Fabricius Eastern tent caterpillar M. californicum (Packard) Western tent caterpillar

Eastern Canada and USA

M. constrictum (Henry Edwards) Pacific tent caterpillar M. disstria (Hu¨bner) Forest tent caterpillar M. incurvum (Henry Edwards) Southwestern tent caterpillar M. indica Walker Indian tent caterpillar M. nuestria Linnaeus Lackey moth, European tent caterpillar, Japanese tent caterpillar. M. tigris (Dyar) Sonoran tent caterpillar

Mexico: Baja California USA: California, Oregon, Washington Southern Canada and continental USA, except Alaska Mexico USA: Arizona, Colorado, Nevada, Utah Northern India, Pakistan

Malus, Prunus and other broadleaf plants Alnus, Betula, Ceanothus Prunus, Purshia, Populus, Quercus, Salix and other broadleaf plants Quercus

Western and northern Canada and USA

Alnus, Betula, Nyssa, Populus, Prunus, Quercus and others Populus, Prunus, Salix Malus, Quercus dilatata, Q. incana

North Africa, Europe, China, Near East, Japan, Korea, Mongolia, Russia, Taiwan

Wide range of forest and fruit trees

Mexico USA: Arizona, Colorado, New Mexico, west Texas

Quercus

n Tovar et al. 1995, Canadian Food Inspection Agency 2006. Sources: Browne 1968, Furniss & Carolin 1977, Cibria

Description of Stages Adults have a wingspan of 25–38 mm for males and 38–51 mm for females. Color of adults of both sexes varies from dark red-brown to yellow and gray with intermediate hues. Forewings are marked with two lines, which may be either lighter or darker than the base color. Larvae vary widely in color. Head capsules are dark blue to black and the body color patterns are mixtures of black, orange and blue. A broken blue or blue-white stripe usually occurs longitudinally along the dorsum (Stehr & Cook 1968, Cibri an Tovar et al. 1995, Fitzgerald 1995, Ciesla & Ragenovich 2008).

Eggs

Larvae (diapausing in egg)

Larvae (active)

Pupae

Adults

J

F

M

A

M

J

J

A

S

O

N

D

Fig. 7.9 Generalized life history of tent caterpillars of the genus Malacosoma.

species for deer and livestock. Defoliated trees and tents are unsightly and large hordes of larvae are a nuisance in residential and recreation areas. Migrating larvae can be a road hazard.

Malacosoma disstria (Hübner), Forest Tent Caterpillar Distribution Forest tent caterpillar is the most widely distributed of the North American tent caterpillars and occurs across southern Canada and most of the USA, except Alaska.

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Hosts Hosts vary according to location. In eastern Canada and northeastern USA, sugar maple, Acer saccharum, northern red oak, Quercus rubrum, and ash, Fraxinus spp., are preferred. In the Great Lakes region, oaks, Quercus spp., quaking aspen, Populus tremuloides, and sugar maple are most frequently damaged. In the Appalachian Mountains and central USA, oaks are the preferred hosts. In the Atlantic and Gulf Coast regions, low-lying forests of water tupelo, Nyssa aquatica, sweetgum, Liquidambar styraciflua, and other species are defoliated. In the Mississippi Valley, eastern cottonwood, Populus deltoides, and elms, Ulmus spp., are damaged and in eastern Texas, oak forests are defoliated. In the west, quaking aspen is preferred, although other broadleaf species are also attacked. Importance Forest tent caterpillar is considered the most destructive of the North American tent caterpillars. Outbreaks normally last 2–3 years and defoliation can occur over thousands of square kilometers. Life History The larvae of this species do not construct tents. Instead, they form a silken mat on the trunk or branch, where they congregate when at rest or during molting. When high populations cause complete defoliation, larvae move around in search of food, often crossing highways and impeding traffic. Description of Stages Adults are buff colored and have a wingspan of 25–38 mm. The forewings have two darker oblique lines near the middle. Newly hatched larvae are almost uniformly black, about 3 mm long, and have conspicuous hairs. Mature larvae have pale blue lines along the sides of a brown body and a row of mid-dorsal keyhole or footprint-shaped, whitish spots on a black background (Batzer & Morris 1978, Fitzgerald & Webster 1993).

Larix, Malus, Morus, Populus, Prunus, Pyrus, Quercus, Rosa, Rubus, Salix, Sorbus, Syringa, Ulmus and, occasionally, Fraxinus and Tilia. Importance This insect is primarily a pest of fruit trees but can also damage forest trees and shrubs. Description of Stages Adults vary in color. There are two morphs, i.e. light and dark, with less frequent intermediate colors. Basic color is brown, but can range from yellow-ochre to red-brown. Males have a wingspan of 30–40 mm. Females are more robust with a wingspan of 36–40 mm. Forewings have a brown, narrow, oblique, central transverse band. Instar I larvae are black and about 2 mm long. Mature larvae are 40–55 mm long, slender and covered with fine hairs. Larvae are marked with a distinct white dorsal line and blue or gray-blue and red-yellow lateral bands separated by black edging (Browne 1968, Shiga 1977). Thaumetopoeidae, Processionary Caterpillars Thaumetopoea Thaumetopoea consists of about nine species native to Europe, North Africa and the Near East and can be divided into two groups. Winter processionary moths have a summer pupal diapause and larvae feed in winter. Summer processionary moths do not have a pupal diapause and larvae feed in spring and summer (Fig. 7.10). The summer diapause causes events in the Th. pityocampa Th. wilkinsonii

Adults Eggs Larvae Pupae

Malacosoma nuestria (Linnaeus) Th. processionea

Adults

Distribution This insect has several subspecies and is widely distributed across Eurasia, from Europe, including southern UK, south to north Africa and east through China, Iran, Japan, North and South Korea, Russia, Syria, Taiwan and Turkey. Hosts M. nuestria has a wide host range including species of Acer, Alnus, Amygdalus, Betula, Carpinus, Castanea, Cotoneaster, Corylus, Crataegus, Fagus, Juniperus,

Eggs Larvae Pupae

J

F

M A

M

J

J

A

S

O

N

D

Fig. 7.10 Generalized life history of winter and summer processionary moths of the genus Thaumetopoea.

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Fig. 7.11 Heavy defoliation of a young Pinus brutia plantation by pine processionary caterpillar, Thaumetopoea wilkinsonii (northern Cyprus).

life cycle of winter processionary moths, such as adult flight and egg hatching, to occur earlier at higher latitudes and elevations. Larvae of most species have urticating hairs that can cause allergic reactions in humans as well as domestic animals. Thaumetopoea pityocampa Dennis and Schiffermüller, Pine Processionary Caterpillar Thaumetopoea wilkinsonii Tams, Eastern Pine Processionary Caterpillar Distribution Th. pityocampa and Th. wilkinsonii are sibling species and have been considered synonymous under Th. pityocampa. However, analysis of mitochondrial DNA of populations throughout the Mediterranean present compelling evidence that they are closely related but separate species. They are found throughout the pine forests of the Mediterranean Basin. Studies in France indicate that they can survive in areas that receive 1800 hours of sun/year and an average minimum January temperature of above 4 C. Th. pityocampa occurs in Europe from southern France, Portugal and Spain west to Italy, Greece and the Balkan Peninsula, and in

north Africa from Morocco west to Algeria and Tunisia.1 Th. wilkinsonii is known from Cyprus, Israel, Lebanon, Syria and Turkey.

Hosts Larvae feed on foliage of Pinus and Cedrus. Pine hosts include P. brutia. P. halepensis, P. nigra, P. nigra var. maritima, P. pinaster, P. sylvestris and, to a lesser degree, P. pinea. P. radiata, native to California, USA, and planted in the Galicia region of northwestern Spain, is also damaged.

Importance Natural forests, plantations and ornamental trees are subject to outbreaks. Young forests, especially plantations on poor sites, are most susceptible. During outbreaks, needles can be stripped from pines in less than a week (Fig. 7.11). Few trees are killed directly but defoliated trees suffer from reduced growth and increased susceptibility to bark beetles. Larvae have urticating hairs. 1

Populations in North Africa on Cedrus have also been referred to as Th. bonjeani (Powell).

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Fig. 7.12 Egg mass of pine processionary caterpillar, Thaumetopoea wilkinsonii, on needles of Pinus brutia (northern Cyprus).

Fig. 7.13 Instar IV pine processionary caterpillar, Thaumetopoea wilkinsonii, larval nest and defoliation on Pinus brutia (northern Cyprus).

Life History There is one generation/year with a summer pupal diapause and larvae feed in winter. Adults emerge in late summer and lay eggs on needles of host trees. Pines growing in the open or at the forest edge are preferred for egg laying. Eggs are laid in rows at the base or mid-point of two or three needles and held together by a white gummy substance secreted by the female (Fig. 7.12). Larvae hatch in late summer– autumn and begin to feed in colonies. Beginning with instar III, larvae construct silken tents that provide protection from natural enemies and insulation against cold temperatures (Fig. 7.13). Some feeding may occur on sunny days in December–January but feeding begins in earnest from February to March. During spring, larvae form single-file, head to tail processions and

crawl down boles of host trees to pupate in the soil. A portion of the pupae emerges the following year. Some remain in diapause for 5 or more years.

Description of Stages Males have a wingspan of 28–36 mm. Forewings are medium brown and the wings are white. Females have a slightly larger wingspan, 36–59 mm and lighter colored forewings wings than males. Eggs are pearl to light yellow. Average egg mass length is 28 mm, average width is 4.7 mm. Larvae are black with bright orange markings that are conspicuous by instar III and are covered with tufts of fine hairs. When mature, larvae are about 35 mm long (Plate 24).

Foliage feeding insects – Lepidoptera Pest Management Aerial and ground applications of chemical insecticides or Bacillus thuringiensis are used to suppress outbreaks. Low-level populations in plantations can be treated by pruning and destruction of branches containing larval colonies (Huchon & Demolin 1971, Buxton 1983, Speight & Wainhouse 1989, Abgrall & Soutrenan 1993, Salvato et al. 2002).

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Pest Management Direct control tactics include removal and destruction of egg masses before they hatch and aerial and ground applications of chemical or biological insecticides (Halperin & Sauter 1999, Thomas et al. 2002, Evans 2007, FAO 2009b, UK Forestry Commission n.d.).

Saturniidae, Giant Silkworm Moths Thaumetopoea processionea Linnaeus, Oak Processionary Caterpillar Distribution Oak processionary caterpillar occurs in central and southern Europe and Israel. Its range is expanding northward and populations have been detected in Belgium, northern France, the Netherlands, Sweden and the UK.

Coloradia Coloradia consists of eight known species. Four are found in western USA and the remainder in Mexico. Larvae feed on foliage of pines and have urticating hairs (Powell & Opler 2009). Coloradia pandora Blake, Pandora Moth

Hosts Larvae feed on foliage of broadleaf trees and shrubs. Oaks, Quercus spp., including Q. cerris, Q. petraea and Q. robur, are preferred but species of Betula, Carpinus, Castanea, Corylus and Fagus are also hosts.

Importance Defoliation causes growth loss and can incite oak decline. Larvae have urticating setae or hairs.

Life History There is one generation/year. Adults are active from July to early September. They are nocturnal and live for 1–2 days. Females lay 100–200 eggs in a mass covered with scales on twigs and branches of host trees. Eggs hatch the following spring and larvae are active from April to June. They feed in colonies and construct silken nests for protection. Larvae move in head to tail processions to new feeding sites. Pupation occurs in the nest in late June–early July.

Description of Stages Adults have a wingspan of 30–32 mm. Forewings are gray with white and dark gray markings. Early instar larvae are uniformly brown with a dark head. Mature larvae have a gray head capsule and an overall gray body color. They have a dark dorsal band with a white stripe on either side. The body is covered with short hairs and red-orange tubercles from which radiate tufts of long hairs.

Distribution Pandora moth occurs throughout pine forests in western USA.

Hosts Hosts are Coulter pine, Pinus coulteri, Jeffrey pine, P. jeffreyi, lodgepole pine, P. contorta, piñon pine, P. edulis, sugar pine, P. lambertiana, and ponderosa pine, P. ponderosa.

Importance Outbreaks occur in areas where soils are composed of pumice or decomposed granite and loose enough for larvae to bury themselves prior to pupation. Moderate to heavy defoliation has occurred in portions of northern Arizona, California, Colorado, southern and central Oregon, Utah and Wyoming, USA (Fig. 7.14). Larvae are a traditional food for the Paiute, an indigenous tribe in the Owens Valley/Mono Lake area of California. They are collected in trenches and gathered by hand once or twice a day. Collected larvae are processed on site by roasting and drying. A mound is made of sandy soil and a fire is built on and around it to heat the soil. When coals die down the mound is opened, live larvae are tossed in and mixed with the hot sand for 30 minutes to an hour. This removes larval setae. Larvae are then sifted from the hot sand, washed, sorted and checked to see if they are properly cooked. The dried larvae are stored in a cool dry place and keep for 1–2 years. Dried larvae are prepared by boiling in plain or salted water for about 1 hour.

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Fig. 7.14 Heavy defoliation of Pinus ponderosa by pandora moth, Coloradia pandora (Deschutes National Forest, Oregon, USA).

Life History Two years are required to complete a generation. Adults are active during late June and July. After mating, females deposit eggs in clusters of 2–50 on the needles or bark of host pines. Eggs hatch in August and larvae feed in small colonies on the new foliage. Larvae overwinter at the base of needles and may feed during warm, sunny days. Feeding resumes in spring and it is during this period that heaviest feeding damage occurs. In late June, larvae enter the soil to a depth of 25–125 cm, form elliptical cells and pupate. The second winter is spent as pupae and adults emerge the following spring and summer. Some pupae may enter an extended diapause for up to 5 years. Description of Stages Adults are heavy bodied, gray-brown with a wingspan of 70–110 mm. They have a small dark spot near the center of each wing. The base and anterior margins of the hind wings are covered by pink hairs, which may be slightly darker on males. Eggs are globular, white and about 3 mm long. Larvae are 5 mm long when first hatched and have shiny black heads and brown to black bodies covered with short black hairs. Mature larvae are 57 to76 mm

long, brown or black to dark green with segmental white lines or bands along each side and a subspiracular white stripe. Pupae are dark purple-brown, from 25 to 35 mm long and not enclosed in a cocoon. Related Species Black Hills pandora moth, Coloradia doris Barnes, occurs in Colorado, Montana, South Dakota and Wyoming, USA. Larvae feed on foliage of Pinus ponderosa. Larvae are longer and have more conspicuously branched spines than those of C. pandora. C. velda Johnson & Walter is closely related to C. doris and endemic to the San Bernardino Mountains of California, USA, where larvae feed on foliage of P. monophylla (Carolin & Knopf 1968, Blake & Wagner 1987, Powell & Opler 2009). Imbrasia belina (Westwood), Mopane Worm, Mopane Emperor Moth Distribution Mopane worm occurs in southern Africa including Botswana, the Democratic Republic of Congo, South Africa, Zambia and Zimbabwe. Hosts

Host tree is mopane, Colophospermum mopane.

Foliage feeding insects – Lepidoptera Importance Larvae feed on foliage and outbreaks cause severe defoliation. Field studies indicate that defoliation reduces seed production. Young trees appear to be more severely affected than mature trees. Larvae are an important food for people in Botswana, Zimbabwe and portions of South Africa. They are rich in protein, fats and amino acids. Sixty-five percent of rural people in southern Africa collect mopane worms for subsistence use and 35% sell dried larvae at rural markets. Some are processed and traded locally and internationally as snacks and canned products. There are concerns that this insect is becoming less abundant due to excessive harvesting of the larvae and unfavorable climatic conditions.

Life History Across most of its range, there are two generations and periods of defoliation/year. In arid regions there is only one generation/year. Adults are active from November to December and February to March and deposit a single cluster of 50–200 eggs on branches or foliage of host plants. Eggs hatch about 10 days later and larvae have five instars. Instars I–III are gregarious and feed in colonies of 20–200 individuals. Instar IV and V larvae are solitary. Feeding occurs over about a 6-week period after which larvae burrow in the soil to pupate. Adults emerge and live only a few days to mate and lay eggs.

Description of Stages Adults are large moths with orange-brown wings and large eye spots. Larvae are yellow with black spots and markings. They have conspicuous black spikes and white hairs and are 90–100 mm long when mature.

Related Species Outbreaks of I. nicitans Fabricius have been reported in plantations of Acacia mearnsii in Uganda. Maesopsis eminii is also reported as a host. Outbreaks usually last for a single generation and result in loss of vigor of host trees (Browne 1968, Ditlhogo et al. 1996, Makhado & Potgeiter 2009).

Ormiscodes cinnamomea (Feisthamel) Distribution O. cinnamomea is found in central Chile and probably adjoining portions of Argentina.

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Hosts Host plants include several indigenous broadleaf trees including Cryptocarya alba, Lithrea caustica, Nothofagus dombeyi, N. obliqua, Peumos boldos and Schinus latifolius. It has also adapted to Chile’s extensive Pinus radiata plantations.

Importance Outbreaks can cause heavy defoliation and growth loss. Heavy defoliation of P. radiata on trees aged less than 10 years can cause tree mortality.

Life History There is one generation/year. Adults are active in autumn (February–May). Sex ratio is roughly 80% males and 20% females. Females are strong fliers and capable of dispersing over long distances. They deposit egg masses of 150–300 eggs on branches of host trees. Depending on temperature, eggs incubate for 50–120 days and hatch from in midAugust until late December. Larvae are nocturnal, feed in colonies and are active in December–January. When feeding is complete they move en masse from tree crowns to the soil to pupate.

Description of Stages Males have a wingspan of 60–80 mm. Wings are light brown to rosy-brown. Forewings have light colored transverse veins. Antennae are plumose. Females are larger, wingspan is 66–95 mm and wing color is similar to males. Antennae are filiform. Eggs are oval-shaped, 2 mm long and 1 mm wide. Color is white to pale green. Mature larvae are 100 mm long and 12 mm wide. Body color is black with yellow longitudinal subdorsal lines. Each body segment has red tubercles with clusters of red-brown hairs (Fig. 7.15). Pupae are naked, black with an overall length of 30–40 mm.

Pest Management Pest management tactics have not been developed for this insect. However, a complex of natural enemies, mostly parasitoids, is known to help keep populations in check.

Related Species O. amphimone (Fabricius) also defoliates broadleaf trees and is found south to the southern tip of South America in both Argentina and Chile (Aguayo Silva et al. 2008).

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Fig. 7.15 Colony of Ormiscodes cinnamomea larvae migrating to a pupation site (near Lonquimay, IX Region, Chile).

Foliage feeding insects – Lepidoptera Sphingidae, Sphinx or Hawk Moths Pseudosphinx tetrio (Linnaeus), Frangipani Hawkmoth Distribution This moth occurs throughout the neotropics and subtropics. Its range extends from southern Brazil north through Central America, Mexico and the West Indies to south Florida, southern Mississippi, Arkansas, Texas, and southern Arizona in the USA.

Hosts Hosts are members of the family Apocynanceae or dogbanes. These include the frangipanis, Plumeria rubra, P. alba, P. obtusa and Adenium sp., Allamanda cathartica, Echites umbellata, Himatanthus sucuuba and Rhabdadenia biflora. The author has observed larvae feeding on giant milkweed, Calotropis procera, on the Caribbean island of Tobago.

Importance Larvae appear in urban settings and can defoliate frangipani trees in a few days. One larva can eat three large leaves per day. They start feeding from the leaf tip and work back and have been known to feed on tree stems if excessive feeding depletes leaf availability. Contact with the larvae reportedly causes eye irritation and, according to local legend on the Caribbean island of Tobago, coming in contact with larvae will cause “the fever.”

Life History In Florida, adults are active from March to September and attracted to species of Vinca for nectar. Females deposit 50–100 eggs in clusters on leaves of host plants. Larvae feed from September through January on plants that produce white toxic latex that they detoxify and possible store for defensive purposes. The larvae are aposmatic. They have a striking color pattern that warns predators that they have an unpleasant taste and/or may be toxic. When feeding is completed, larvae drop to the litter and pupate.

Description of Stages Female adults are larger than males and lighter in color. Adult dorsal forewings are brown with a dark spot at the base of the costal margin and blurry gray and white markings. Dorsal hind wings are dark brown with white along the inner margin and the lower half of the outer margin. Body is striped with

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transverse gray-white bands and wider black ones. Wingspan is 12.7–14 cm. Eggs are smooth except for minute punctures on the surface. They are pale green, ellipsoidal and measure 2.2–2.5 mm. Larvae are velvetblack with yellow rings and a red-orange head and can attain lengths of up to 15 cm. The black “horn” on abdominal segment eight is approximately 2.5 cm long and is located on an elevated orange “button”. Thoracic legs and prolegs are orange and speckled with black spots (see Plate 14). Newly molted larvae are light yellow and dark gray in alternating transverse rings and obtain their typical coloring several hours after molting (Author’s observations, Dunford & Barbara 2005).

Noctuidae, Subfamily Lymantriinae, Tussock Moths Dasychira (¼ Paraorgyia) grisefacta (Dyar), Western Pine Tussock Moth Distribution This tussock moth is indigenous to western North America. In occurs in British Columbia and southern Alberta, Canada, and Arizona, Colorado, Montana, Nebraska, New Mexico, North Dakota, Oregon, South Dakota and Wyoming, USA.

Hosts P. grisefacta feeds on conifers, including Douglas-fir, Pseudotsuga menziesii, western hemlock, Tsuga heterophylla, Engelmann spruce, Picea engelmannii, white spruce, P. glauca, and ponderosa pine, Pinus ponderosa. Several other conifers are minor hosts.

Importance In the Pacific Northwest, D. grisefacta is a relatively uncommon solitary defoliator. However, it occasionally reaches outbreak levels on individual open-grown trees in urban settings. This insect has reached epidemic proportions in ponderosa pine forests on the eastern slopes of the Rocky Mountains and Great Plains and has caused defoliation on several thousands of hectares.

Life History Western pine tussock moth has one generation/year. Adults emerge in late July–August. After mating, females lay eggs in small clusters on host foliage. Larvae emerge soon after, feed briefly and seek

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hibernation sites under bark scales, where they overwinter. Larvae resume feeding in spring on old foliage and continue until July when they pupate. Description of Stages Females are wingless. Males have a wingspan of about 42 mm. Forewings are light gray-black with a small white spot. Mature larvae are up to 35 mm long. The head capsule is black and unmarked and the body is covered with white or yellowish hairs radiating from black tubercles. There are four mid-dorsal tufts of dirty white hairs on abdominal segments one to four. Two long black pencil-tufts project forward from the first thoracic segment and three more extend from the rear of the body. Abdominal segments six and seven have bright red dorsal glands (Ferguson 1978, Duncan 2007).

Euproctis chrysorrhoea (Linnaeus), Browntail Moth Distribution Browntail moth is indigenous to southern and central Europe, the Near East and northern Africa. Its eastern limits of distribution are the Caucasus and northwestern Kazakhstan. It was introduced into North America and infestations were discovered in Massachusetts, USA in 1897. After its initial establishment, it spread throughout much of the New England states of the USA and east to Nova Scotia, Canada. Its range began to decline in the 1920s and it presently occurs only in a few locations along the coast of Maine and Massachusetts. Hosts Hosts are broadleaf trees and shrubs, including Castanea sativa and species of Acer, Populus, Quercus, Salix and Tilia. It also feeds on foliage of fruit trees of the family Rosaceae. Importance This insect can reach outbreak proportions in broadleaf forests within its natural range and cause heavy defoliation. In the UK, it has been a pest in urban forests. Larvae have urticating hairs, which can be irritating to humans. In North America it has caused localized defoliation in areas where it has been introduced. Life History There is one generation/year. Adults are active from June to August. Females deposit eggs in

clusters of 200–500 eggs, beginning in mid-July. Eggs hatch in early August and larvae feed in colonies of as many as 2,000 larvae. Early instars skeletonize leaves. In autumn, instar II and III larvae form nests of tightly webbed foliage where they overwinter. They become active again in spring as buds of broadleaf trees burst and resume feeding on buds and, later, foliage. In early spring, the larvae feed during the day and return to their nests at night. As temperatures warm, they abandon their nests. After 30–50 days of feeding, the larvae pupate, either individually or in small groups, in the lower portions of tree crowns, boles and branches.

Description of Stages Adults have a wingspan of 30–40 mm. Head, thorax and abdomen are white with a silky shimmer. The apex of the abdomen of females is covered with golden hairs and that of males is covered with brown hairs. Larvae are about 38 mm long when mature, brown in color with tubercles that contain clusters of yellow-brown setae. The dorsal surface has a broken white band and conspicuous red spots on the ninth and tenth abdominal segments.

Pest Management In winter, branches containing nests of overwintering larvae can be pruned and burned. In some cases, aerial and ground applications of chemical or biological insecticides may be needed to control outbreaks. A host specific nucleopolyhedrosis virus has been recovered from larvae and can be used to treat outbreaks.

Related Species The yellow-tail moth, E. similis (Fuessly) also occurs in Europe and feeds on broadleaf trees and shrubs. The abdomen of both sexes is covered with golden-yellow hairs (Cory et al. 2000, USDA Forest Service 2004, Grichanov & Ovsyannikova 2008b).

Euproctis kargalika Moore, Turkistan Browntail Moth Distribution This species is indigenous to Asia and from northwestern China, southern Siberia and Altai Kray, Russia, Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan and Uzbekistan. It has also been reported from Iran, in the vicinity of Teheran, where it may have been introduced.

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Hosts Larvae of E. kargalika feed on buds and foliage of many deciduous trees and shrubs, including species of Acer, Betula, Caragana, Cotoneaster, Crataegus, Malus, Pistacia, Pyrus, Quercus, Ribes, Rosa, Rubus, Salix and Ulmus.

overwintering larval colonies, light trapping of adults and aerial spraying with either chemical or biological insecticides. Natural enemies, including several parasitoids, predators and fungi, help bring about the natural collapse of outbreaks (EPPO 2002a).

Importance This insect is considered an important defoliator of deciduous forests in central Asia. Outbreaks occur periodically, usually in mountainous regions at altitudes between 900 and 1600 m, and last for 2 years. They can occur over large areas and cause 100% defoliation, resulting in tree mortality. Trees and forests in plains regions, shelterbelt plantings, fruit orchards and urban areas can also suffer damage. Most feeding occurs in spring. High populations of other forest defoliating caterpillars may occur in association with outbreaks of this insect. Larvae have urticating setae.

Leucoma salicis (Linnaeus), Satin Moth

Life History There is one generation/year and larvae overwinter in nests made of tightly woven silken threads. Moth flight can occur from early June to mid-September, depending on elevation. Eggs hatch between early July and September, again depending on elevation. Early instar larvae make a next of leaves tied together with webbing and feed inside. Larvae usually overwinter as instar II in nests and become active in spring, when they feed on buds and foliage and can cause severe defoliation. Larvae construct new nests in branches of host trees made up of webbing and feed in colonies until they reach instars V and VI, when they feed as individuals. Pupation can occur from late April to June but the individual period of pupation is about 15 days. Description of Stages Adults are white moths. Males have a wingspan of 30–35 mm and females 35–40 mm. Forewings have several black spots on the upper surface. The distal segments of the abdomen are covered with a thick layer of yellow-brown hairs. Mature larvae are 35–37 mm long, gray-white with black bands and an orange-brown pattern. Clumps of setae originate from tubercles on the sides of the body and are yellow-white in color.

Pest Management During outbreaks, direct control tactics include collection and burning of egg masses and

Distribution Satin moth is native to Europe and Asia. It was first reported in North America in 1920 in both the New England states of the USA and southwestern British Columbia, Canada. In the USA, this insect is now found in the northeast, the Pacific northwest and Wyoming.

Importance In Europe and North America, larvae feed on foliage of Populus and Salix. In China, species of Acer, Corylus and Fraxinus are known hosts.

Damage Outbreaks occur in both its native range and places where it has been introduced. It can cause complete defoliation resulting in growth loss, top-kill and tree mortality (Browne 1968, Humphreys 1996).

Life History L. salicis has one generation/year in northern Europe, North America and Inner Mongolia. In the southern parts of its European range and China, two to three generations may occur. During a 1-year life cycle, eggs are laid from early July to late August on leaves, twigs, branches and trunks of host trees, or indiscriminately on other objects. Eggs hatch about 2 weeks later. Instar I and II larvae skeletonize foliage for about 2 weeks, then spin hibernaculae in sheltered areas under moss or bark crevices, where they molt and overwinter (Fig. 7.16). Instar III larvae emerge the following April and feed on foliage until mid-June. Pupation occurs in a cocoon of loosely spun white silk within rolled leaves on twigs or in bark crevices. Adults emerge in early July.

Description of Stages Adults are large silvery white moths with a satin sheen and dark body showing through the hairs. Wingspan is 30–50 mm, body length is 15–20 mm. Eyes and legs are black. Antennae of male moths are plumose while those of females are

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Fig. 7.16 Early instar larvae of the satin moth, Leucoma salicis, on foliage of Populus tremuloides (British Columbia, Canada).

thread-like. Larvae are pale to medium gray-brown and about 40 mm long when mature. Larval head capsule is dark gray and the body’s dorsal surface has one row of large oblong, double, shiny yellowish-white blotches and two subdorsal broken yellowish lines. Larvae also have two lateral and two subdorsal rows of tufted brownish setae. Pest Management Outbreaks can be treated with aerial or ground applications of Bacillus thuringiensis (Browne 1968, Drooz 1985, Humphreys 1996, Schmutzenhofer et al. 1996). Lymantria Lymantria consists of about 31 species native to Asia and Europe and includes some of the world’s most damaging forest defoliators. One species, L. dispar, has been introduced into North America where it is a major defoliator of broadleaf forests (Table 7.6, Pogue & Schaefer 2007).

Lymantria dispar (Linnaeus), Gypsy Moth (Plates 25–27) Distribution Several subspecies are recognized (Table 7.6), including L. dispar dispar (Linnaeus), the European gypsy moth and L. dispar asiatica Vnukovskji, the Asian gypsy moth. L. dispar dispar occurs throughout Europe nearly to the Ural Mountains, the Near East, the Mediterranean islands of Corsica and Sardinia, Algeria, France, Italy and Morocco. This subspecies was introduced into North America in 1868 or 1869 and is established as far north as New Brunswick and Nova Scotia, Ontario, and Quebec, Canada, south to North Carolina and west to parts of Indiana, Illinois, Michigan, Ohio, West Virginia and Wisconsin, USA. Isolated populations have occurred in California, Colorado, Missouri, Oregon, Utah and Washington, USA and British Columbia, Canada. L. dispar asiatica occurs in Asia, generally east of the Ural Mountains into the Russian Far East, the northern two thirds of China, Mongolia, and North and South Korea. It does not occur south of the Himalayas (Fig. 7.17).

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Table 7.6 Representative species of Lymantria (Lepidoptera: Noctuidae: Lymantriinae): their distribution and hosts. Species

Distribution

Principal hosts

L. dispar dispar (Linnaeus) European gypsy moth

Europe, North Africa, introduced and established in eastern North America Temperate Asia, Russia east of the Ural Mountains, north of China, North and South Korea, Mongolia Japan. Honshu, Shikoku and Kyushu. Locally established on parts of Hokkaido. Asia. Japan, Russia, Taiwan and Vietnam, westward across China, Thailand, Nepal, India, Sri Lanka. Eurasia

Broadleaf trees including Alnus, Betula, Corylus, Crataegus, Quercus, Populus, Tilia and others. Also some conifers Wide range of broadleaf and coniferous trees

L. dispar asiatica Vnukovskji Asian gypsy moth.

L. dispar japonica (Motschulsky) L. mathura Moore Pink gypsy moth

L. monacha (Linnaeus) Nun moth L. obfuscata Walker Indian gypsy moth L. umbrosa Butler Hokkaido gypsy moth

Afghanistan, northern India, Pakistan Hokkaido, Japan, northeast into the Kuril Islands, Russia

Found on many of the same plants used by L. dispar dispar and L. umbrosa Wide range of broadleaf trees and shrubs

Wide range of broadleaf and coniferous trees, damaging to conifers in central Europe Broadleaf trees including Alnus, Cydonia, Juglans Malus, Morus, Populus, Prunus, Pyrus, Quercus, Robinia, Rosa Wide range of broadleaf and coniferous trees. Damaging to Larix leptolepis plantations

Source: Pogue & Schaefer 2007.

Hosts Hosts include well over 100 plants. L. dispar dispar prefers species of Alnus, Betula, Corylus, Crataegus, Populus, Quercus and Tilia. Late instar larvae can feed on conifers. L. dispar asiatica has a broader host range than its European counterpart and Siberian larch, Larix sibirica, is the dominant host over much of eastern Russia and Mongolia. Importance Outbreaks in the natural ranges of both subspecies and, in the case of L. dispar dispar, in North America have caused millions of hectares of defoliation. Outbreaks of European gypsy moth can incite oak decline (see Chapter 3). In urban forests, defoliation is unsightly and larvae on exterior walls, roads and lawn furniture are a nuisance. Life History Gypsy moth has one generation/year. Adults are active from July to August. The key difference between L. dispar dispar and L. dispar asiatica is that females of L. dispar dispar do not fly, even though they have fully developed wings. L. dispar asiatica females are capable of flight. After mating, egg masses are laid on branches and trunks of trees but can be deposited in

almost any sheltered location, including lawn furniture or inside the hubcaps of vehicles. Larvae hatch 4–6 weeks later but remain inside the eggs for the winter. Larvae emerge at budburst and begin feeding. Early instar larvae are subject to dispersal by air currents. They feed for about 6 weeks and then pupate in bark crevices, branches or on the ground. Pupation lasts 7–14 days.

Description of Stages Female adults are white with a wingspan of about 50 mm. Males are dark brown with black bands across the forewings with a wing span of about 37 mm. Mature larvae range from 37 to 60 mm long. Head has yellow markings and body is sooty colored and hairy. The dorsal surface has five pairs of blue spots followed by a double row of six pairs of red spots. Egg masses are buff when first laid but may bleach out over the winter when exposed to direct sunlight and weathering.

Pest Management Isolated populations in North America have been eradicated via spray programs and

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Fig. 7.17 Worldwide distribution of gypsy moth, Lymantria dispar (Lepidoptera: Noctuidae: Lymantriinae) (adapted from Orozumbekov et al. 2009).

mass trapping of males using pheromones. Outbreaks are treated with aerial or ground applications of chemical or biological insecticides. Biological insecticides include a nucleopolyhedrosis virus (Gypchek) and Bacillus thuringiensis. The fungus Entomophaga maimaiga causes a disease of gypsy moth, Lymantria dispar, in Japan and has been introduced into North America (Drooz 1977, McManus et al. 1989, Reardon & Hajek 1997, Pogue & Schaefer 2007). Lymantria monacha (Linnaeus), Nun Moth Distribution Nun moth has an almost continuous distribution across the Palearctic boreal conifer forests.

Hosts Larvae can feed on both conifers and broadleaf trees. Conifers are preferred and include species of Abies, Juniperus, Larix, Picea, Pinus and Tsuga. Broadleaf hosts include Acer, Betula, Carpinus, Prunus, Quercus, Salix and others.

Importance Nun moth is considered one of the most damaging forest defoliators in central Europe. Outbreaks have occurred in Picea abies and Pinus sylvestris forests in Austria, Belarus, Czech Republic, Germany, Poland, Romania, Russia and other countries, caused widespread defoliation and extensive tree mortality (Fig. 7.18). From 1978 to1983, an outbreak in Poland required aerial application of insecticides on over 2.5 million ha, representing 25% of the country’s forests. Populations were again high in Poland during the mid-1990s.

Life History Nun moth has one generation/year. Adults are active during July–August but may persist until September. They are nocturnal and males can fly long distances to search for females. Mated females deposit eggs in masses in bark crevices or under scales and lichens. One female can lay 200–250 eggs with 20–100 eggs per mass. Larvae hatch and overwinter inside the egg. During late April or early May of the

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length. Mature larvae have gray-yellow heads with black and brown spots. There are two lines of tubercles with tufts of hair on the sides of the body and light spots on the third, seventh and eighth abdominal segments. Pest Management Outbreaks have been treated with aerial and ground applications of chemical and biological insecticides (Speight & Wainhouse 1989, Ciesla 1994, Kolk & Starzuk 1996, Pogue & Schaefer 2007). Orgyia Orgyia consists of about 19 known species worldwide. Larvae feed on a range of trees, shrubs and herbaceous plants. One species, O. antiqua, has a holarctic distribution. Eight are native to Asia, Africa or Europe and 10 to North America. Several are pests of forests, ornamental trees, orchards and/or range plants (Table 7.7). Larvae have dense tufts of hair (tussocks) that protrude from the backs. Hairs of most species are urticating and cause tussockosis, an allergic skin irritation. Female adults have wings reduced to inconspicuous pads. They deposit a single mass on the cocoons from which they emerged. Eggs are usually covered with a frothy protective coating to which body hairs of females adhere (Perlman et al. 1976, Ferguson 1978). Fig. 7.18 Defoliation of Scotch pine, Pinus sylvestris, by nun moth, Lymantria monacha (central Poland).

following year, larvae emerge and remain in a group for several days. They then climb into the crown and feed on young foliage and male flowers. Instar II–VI larvae feed on old needles. Larval feeding takes from 40 to 80 days. Pupation occurs on tree trunks or crowns and understory plants. Description of Stages Adult females are larger than males, 15–20 mm long, with a wingspan of 45–55 mm. Males are 12–15 mm long, with a 35–45 mm wingspan. Male antennae are comb-shaped, while female antennae are thread-like. Color of adults varies considerably with light to dark forms occurring. Forewings of both sexes are white with wavy dark cross bands. Hind wings are gray-brown. Eggs are round-shaped, graybrown and about 1 mm in diameter. Instar I larvae are 3–5 mm long. They are dark with tufts of hair of varying

Orgyia antiqua (Linnaeus), Rusty Tussock Moth Distribution O. antiqua is the widest ranging species of Orgyia and occurs throughout Asia, Europe and much of North America. In the UK it is known as the common vapourer. Several subspecies are recognized.

Hosts Hosts include many broadleaf trees and shrubs, including species of Acer, Alnus, Betula, Malus, Salix and Vaccinium. Conifer hosts include species of Abies, Larix, Picea, Pseudotsuga menziesii and Tsuga heterophylla.

Importance Rusty tussock moth is a common and occasionally damaging solitary defoliator in urban forests. Localized, short-lived outbreaks have also occurred in forested areas.

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Table 7.7 Representative species of Orgyia (Lepidoptera: Noctuidae: Lymantriinae): their distribution and hosts. Species

Distribution

Hosts

O. antiqua (Linnaeus) Rusty tussock moth

Holarctic – Asia, Europe and North America

O. cana Henry Edwards

Western USA

O. leucostigma (J.E. Smth) White marked tussock moth

Eastern North America, west to Colorado, New Mexico, USA and British Columbia, Canada Africa: Kenya Asia: Japan, Korea, Taiwan. Introduced to Auckland, New Zealand 1996, eradicated 1999 Western North America

Broadleaf species: Acer, Alnus, Betula, Malus, Salix, Vaccinium, Conifers: Abies, Larix occidentalis, Picea, Pseudotsuga menziesii, Tsuga heterophylla Amelanchier, Ceanothus velutinus, Purshia tridentata, Quercus chrysolepis, Q. kelloggii, Rhamnus californica, Rosa, Salix and others Broadleaf trees and shrubs, occasionally feeds on conifers

O. mixta Snellen O. thyellina Butler

O. pseudotsugata (McDunnough) Douglas-fir tussock moth

Pinus radiata Larix, Malus, Quercus, Salix, Ulmus

Abies, Picea, Pseudotsuga menziesii, occasionally Pinus

Sources: Austara & Migunda 1971, Furniss & Carolin 1977, Drooz 1985, Glare 2009.

Life History Adults are active in late summer–early autumn. After mating, wingless females lay up to 200 completely exposed eggs on their cocoons. Larvae emerge the following late May–early June and feed until late July–early August. When feeding on conifers, larvae initially feed on the current year’s foliage, causing it to turn brown. Later they may feed on both current and older foliage. Defoliation occurs first in the upper crown, then in the outermost portion of the branches and finally in the lower crown and further back on branches. Mature larvae pupate in a cocoon on foliage or tree boles in August.

Description of Stages Males have rust brown wings and each forewing has a white, comma-shaped spot. Wingspan is 35–38 mm. Mature larvae are up to 30 mm long with a black, unmarked head capsule. The body is covered with yellow-white hairs radiating from orange tubercles and with mid-dorsal tufts of white hairs on the first to fourth abdominal segments. Two long, black pencil tufts project forward from the first thoracic segment. A similar tuft extends back from

the rear of the body (Browne 1968, Ferguson 1978, Duncan 2007). Orgyia pseudotsugata (McDunnough), Douglas-fir Tussock Moth Distribution Douglas-fir tussock moth is indigenous to western North America from British Columbia, Canada, south to Arizona and New Mexico, USA.

Hosts Three conifers are favorite hosts and preference depends on location. In the northern and eastern parts of its range, British Columbia, Canada, and Colorado, western Montana, northern Washington, USA, Douglas-fir, Pseudotsuga menziesii, is preferred. In the central part of its range, Idaho, Oregon and southern Washington, USA, Douglas-fir, white fir, Abies concolor, and grand fir, A. grandis, are equally acceptable. In the southern part of its range, Arizona, California, Nevada and New Mexico, USA, white fir is the preferred host. In urban forest ecosystems, blue spruce, Picea pungens, is a favorite host although it is usually not damaged in forests. After larvae have consumed all of the preferred

Foliage feeding insects – Lepidoptera foliage, they feed on other conifers and understory shrubs.

Importance This insect can reach outbreak levels at 7–10-year intervals and cause widespread and severe defoliation. Outbreaks develop explosively and subside abruptly after 1–3 years, often due to infection of larvae by a nucleopolyhedrosis virus. During outbreaks, trees can be stripped of their foliage in a single growing season. Defoliation causes growth loss, top kill and tree mortality. Weakened trees are often killed by bark beetles. During an outbreak in the 1970s in the Blue Mountains of Oregon and Washington, USA, 39% of all host trees were killed in heavily defoliated areas. Within these areas were patches where nearly all host trees died. Top kill in the heavily defoliated areas occurred in 10% of the grand fir and 33% of the Douglas-fir. Outbreaks tend to develop at low elevations on warm, dry sites. Larval setae are urticating and can cause severe skin irritation.

Life History There is one generation/year. Adults are active from late July to November, depending on location. Males are most active around midday. Wingless females emit a pheromone, which attracts males during flight, and mating occurs soon after emergence. Females remain on the cocoon and lay an egg mass mixed with a frothy substance and hairs. One egg mass may contain up to 350 eggs. Eggs overwinter and hatch in late May or early June, which coincides with bud burst and shoot elongation of host trees. One to seven days after egg hatch, larvae crawl to new needles and begin feeding. Dispersal is by windborne early instar larvae. When larvae drop from foliage, they produce silken threads and are carried by air currents. The distance most caterpillars travel rarely exceeds 500 m. Larvae feed on the current year’s foliage when young, but when half grown, they can complete development on old foliage. Pupation occurs from late July to late August inside a thin cocoon of silken webbing mixed with larval hairs. Pupation lasts 10–18 days.

Description of Stages Females have rudimentary wings, small thread-like antennae and a large abdomen filled with eggs. They are about 19 mm long, graybrown in color and hairy. Males are gray-brown to black-brown moths with feathery antennae with a

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wingspan of 25–31 mm. Forewings are gray brown and have two indistinct, irregular dark bars and two vague whitish spots. Hind wings are brown (Plate 28). Early instar larvae are 4–7 mm long and have fine body hairs. Mature larvae are 30 mm long with two long, dark tufts or pencils of hair located behind the head. Another pencil is located at the rear of the abdomen. Four buff-colored tussocks are located forward along the middle of the back. The remainder of the body is covered with short tufts of hair radiating from red centers (Plate 29).

Pest Management Management of outbreaks involves early detection and application of control measures before defoliation occurs. Population fluctuations are monitored using pheromone-baited traps to attract males. Early instar larvae can be treated with aerial or ground applications of Bacillus thuringiensis, the Douglasfir tussock moth virus or chemical insecticides. Identification of sites at high risk of outbreaks and favoring nonhost trees is a cultural tactic that can prevent outbreaks (Wickman 1978, Wickman et al. 1981). Sarsina Sarsina is a small genus of neotropical Lymantriinae and consists of 8–10 poorly defined species indigenous to Mexico, Central and South America.

Sarsina violascens Herrich-Schaeffer Distribution This species is known from Mexico south to Argentina. It is widely distributed in Brazil and known from the states of Bahia, Espirito Santo, Minas Gerias, Par a, Paran a, Rio Grande do Sul, Santa Catarina and S^ ao Paulo. In Mexico, it has been reported from the states of Tabasco and Veracruz. Hosts Hosts are guava, Psidium guajava, strawberry guava, Psidium cattelianum (Mrytaceae), guaco, Mikania spp. (Compositae), tea olive or sweet olive, Osmunthus fragans (Oleaceae), and species of Eucalyptus, including E. citriodora, E. cloenziana, E. grandis, E. nesophila and E. urophylla. Importance S. violascens is a defoliator of eucalypt plantations in Argentina, Brazil, Mexico, Paraguay and

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Peru. It is an important pest in central Brazil and portions of Mexico where eucalyptus plantations are established. Life History Data from Mexico suggest that this insect has three generations/year with active life stages present during 8 months of the year. The first two generations are most damaging with natural enemies reducing numbers of the third generation to low levels. In Brazil, adults are active from March to December depending on local climate. Both sexes fly and are active at night. Eggs are deposited singly or in strips of up to 40 eggs on foliage. Duration of the egg stage is about 11 days. Larvae are nocturnal and congregate on the lower third of boles of host trees during the day. Average duration of larvae is 37 days. When feeding is completed, larvae enter a prepupal stage for 2 days. Pupation occurs in foliage, tree trunks and understory vegetation and lasts about 9 days. Description of Stages Moths have robust abdomens with yellow and red-brown coloration. Upper wings are mottled brown or violet, with four dark bands and four lighter bands. Ventral surface of wings are yellow with three irregular brown transverse lines. Antennae are pectinate on both sexes. Wingspan is 42–53 mm and males are slightlysmaller than females. Eggs are spherical, about 1.25 mm in diameter and milky white in color. Larvae are light brown to beige and covered with urticating hairs. Pupae are humpbacked and red-brown with green highlights on both dorsal and ventral surfaces. Male pupae are smaller than females. Pupal length of males averages 17 mm, females 24 mm (Zanuncio et al. 1992, 1993, Izquierdo et al. 2000, Izquierdo & Gilli 2002) Arctiidae Hyphantria cunea (Drury), Fall Webworm (Plate 30) Distribution Fall webworm is native to North America and widely distributed across southern Canada, most of the USA and northern Mexico. It is established in Europe and parts of Asia. Fall webworm first arrived in Europe in the 1940s in Hungary and spread throughout the continent. It was reported from China in 1979 and is now established. Hosts This insect feeds on many broadleaf trees, including species of Acer, Alnus, Betula, Carya,

Diospyros, Fraxinus, Juglans, Liquidambar, Populus, Salix and Ulmus. In China, 175 different plant species in 49 families and 108 genera are known hosts, including agricultural crops such as cabbage, cotton, fruit trees and maize.

Damage Fall webworm is a common but minor defoliator in its native habitat where defoliation and webs in urban areas are considered unsightly. In places where it has been introduced, it is considered a major pest. For example, in China and Kyrgyzstan, fall webworm is regarded as a major pest of forest, fruit and agricultural crops. In Romania it is the only exotic forest insect that has, to date, caused severe damage.

Life History In North America, fall webworm has one generation/year in northern locations and two generations/year further south. Adults are active at night and lay eggs in flat masses of several hundred eggs on the undersides of leaves. Larvae feed in colonies and form large, tan-colored webs over trees. Heavy infestations can enclose most of a tree crown in webbing. When larvae finish feeding, they drop to the soil, spin transparent cocoons and pupate.

Description of Stages Adults have a nearly white body with an occasional scattering of black and orange markings on the body and legs, and a wingspan of about 30 mm. Eggs are light green or yellow and globose. Two races of larvae are recognized: a black-headed form in the northern part of its range and a red-headed form further south. Mature larvae are 30–35 mm long when mature. The northern strain has a pale yellow or green body with a dark dorsal stripe and white hairs growing out of orange tubercles. Mature larvae of the southern strain have a tan or yellow body, with red to orange tubercles and light brown hairs.

Pest Management In places where fall webworm has been introduced, pest management measures include regulatory tactics and application of Bacillus thuringiensis (Furniss & Carolin 1977, Drooz 1985, ISSG 2007).

Chapter 8

Other Foliage Feeding Insects

Insects representing orders other than the Lepidoptera are also important foliage feeders of forest trees. This chapter addresses foliage feeding insects of importance in forestry of the orders Phasmatoidea (walkingsticks or stick insects), Coleoptera (beetles) and Hymenoptera (bees and wasps). PHASMATOIDEA (WALKINGSTICKS) Heteronemiidae Diapheromera femorata (Say) Distribution This species is widely distributed across eastern North America as far west as the Great Plains and Texas. Hosts Hosts are broadleaf trees including species of Quercus, Prunus, Tilia and Ulmus. Damage D. femorata is the only North American walkingstick of economic importance. It occasionally

reaches outbreak levels and can defoliate thousands of hectares of broadleaf forests.

Life History In southern USA, there is one generation/year. Further north, 2 years may be required to complete a generation. Eggs overwinter and in northern locations, two winters are passed as eggs. Eggs hatch in May–early June and nymphs feed first on foliage of herbaceous plants and low shrubs. Later, they feed on the same trees as the adults. Adults appear in July–August and females lay eggs until the onset of winter by showering them on the ground. During outbreaks, the sound of falling eggs resembles that of rain.

Description of Stages Adults are about 75 mm long and resemble twigs. Body color is variable and ranges from brown to green (Fig. 8.1). Some individuals are mottled or multicolored in hues of gray, green, red or brown. Newly hatched nymphs are pale green and about 8 mm long (Drooz 1985).

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Fig. 8.1 Pair of walkingsticks, Diapheromera femorata, mating (Skyline Drive, Virginia, USA).

Didymuria violescens (Leach), Spurlegged Phasmatid Distribution Spurlegged phasmid is indigenous to mountain forests of southeastern Australia.

Hosts Several eucalypts including alpine ash, Eucalyptus delegatensis, and mountain ash, E. regnans, are hosts.

Importance Nymphs and adults feed on foliage. Nymphs cut crescent-shaped indentations into leaf margins when feeding. They feed only on soft, immature leaves and new shoots and typically consume an entire leaf before moving on to other leaves. In contrast, adults can feed on much tougher mature leaves and tend to be more wasteful, cutting off large pieces of foliage that drop to the ground. This insect usually causes little or no long-term damage to trees or forests. However, outbreaks that cover large areas can occur and cause complete defoliation. Outbreaks tend to occur in high-elevation forests, where there is a higher proportion of preferred hosts, and in even-aged forests

between the ages of 20 and 60 years. In some cases, two distinct populations may exist in the same area. This leads to situations where defoliation by peak population of one group is followed in the next year by defoliation by the second group. This causes continuous defoliation over a prolonged period. Repeated heavy defoliation can cause tree mortality. Life History Two years are usually required to complete a generation and, during outbreaks, defoliation typically occurs once every 2 years. However, the length of a generation can vary between 1 and 4 years. Females deposit eggs in the litter between late summer and late autumn, with peak egg production occurring in early autumn. Mating is not necessary and unfertilized eggs produce females. Eggs incubate over 18–20 months and hatch during spring and early summer. Immediately after hatching, nymphs move across the forest floor and climb the first vertical object in their path. Most spend their entire lives in the first eucalypt they reach, although some adult males glide to search for a mate. Roughly 2 weeks are required for these insects to pass through five instars before becoming adults.

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Description of Stages Nymphs are light green and soft bodied. Adults are about 80 mm long, excluding antennae. Males are brown-green and females are leaf green. Females have a wider abdomen and shorter antenna than males. Both sexes have wings but only males fly.

Description of Stages Adults are 19–26 mm long with a medium brown body color. The prothoracic shield is slightly darker than the elytra. The body and legs are clothed with short, dense white hairs. The elytra have faint longitudinal bands (Sabatinelli 1976, Triggiani & Tarasco 2008).

Pest Management Management of outbreaks includes reduction of understory vegetation. This creates a warmer, drier microclimate and causes eggs to desiccate. Prescribed burning is also used to kill eggs. Aerial spraying of insecticides has been used to treat outbreaks.

Hylamorpha elegans (Burmeister)

Related Species Two other walkingsticks defoliate eucalypt forests in Australia: Ctenomorphodes tessulatus (Gray) and Podacanthus wilkinsonii Macleay (Ohmart & Edwards 1991, Collett 2006).

COLEOPTERA (BEETLES) Scarabaeidae (June Beetles) Anoxia matutinalis matutinalis Castelnau Distribution A. matutinalis matutinalis is one of three recognized subspecies and is found in the Mediterranean region of Europe, including most of Italy, the island of Malta and portions of the former Yugoslav Republic.

Hosts Two Mediterranean pines, Pinus halepensis and P. pinea, are known hosts.

Importance Adults feed on foliage and can cause moderate to heavy defoliation. Outbreaks have been reported in coastal pine forests and woodlands in the South Apulia region of Italy.

Life History Relatively little is known about the life history and habits of this beetle. Adults emerge in large numbers from soil in early June and feed on needles. Larvae are known to occur in the soil and have been reared in the laboratory. However, they have not yet been found in the field.

Distribution This beetle occurs in central and southern Chile, from the VIII to X regions, including Chiloe Island, and adjoining Argentina including Nahuel Huapi National Park in northern Patagonia.

Hosts Hosts include several species of southern beeches, Nothofagus spp.

Importance Adults feed on foliage and during outbreaks large trees can be completely stripped. Larvae feed on roots. Repeated defoliation causes growth loss, branch dieback and, under rare circumstances, tree mortality. Feeding by larvae on roots can also lead to stress and weakening, especially in combination with drought.

Life History H. elegans has one generation/year. In Chile, adults are active during summer (midNovember–late February). Adults fly at night and are attracted to lights. Eggs are laid in the ground in pastures or nurseries and larvae, known as white grubs, occur in the soil from January to November where they feed on plant roots or, occasionally, decaying logs. During periods when soil moisture is high, larvae are found at the depth of plant roots, but as the soil dries, they move deeper into the ground. Pupation occurs in the soil and adults emerge in early summer.

Description of Stages Adults are attractive June beetles, 11.8–18.2 mm long. Color of head, pronotum and elytra is light to dark apple green with a metallic silver, bronze or, rarely, orange reflection usually present at the apex of the clypeus, the mid-point of each eye and the lateral margin of the pronotum and elytra. The color of the head, pronotum and elytra may vary from light brown to pale yellow-green. Larvae are white C-shaped grubs with a light brown head. Length of

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mature larvae is 25–27 mm (Duran 1952, Billings & Holsten 1969, Ratcliffe & Ocampo 2002, Aguayo Silva et al. 2008).

Importance This insect is considered one of Tasmania’s most damaging forest insect pests. It defoliates second growth stands and plantations in all age classes.

Chrysomelidae (Leaf Beetles) Calomicrus apicalis Demaison Distribution This species is found in the eastern Mediterranean region, including portions of Syria and Turkey.

Hosts Reported hosts are Cedrus libani, Pinus brutia and P. nigra ssp. pallasiana.

Importance Adults feed on new foliage, which turns yellow and dies. Outbreaks have been reported in young Cedrus libani forests in several areas of Turkey since 2000 with up to 3000 ha of defoliation.

Life History This insect has only recently been reported a pest and little is known of its life history and habits.

Description of Stages Adults are about 5–6 mm long. The thorax and elytra are yellow to yelloworange. The head and abdomen are dark brown. Males are more robust than females and the femora of males are slightly enlarged. Legs of females are entirely yellow and the basal portion of the femora is dark brown to black on males.

Pest Management Pest management methods are not available for this insect (Bezdek 2006, Aytar et al. 2008). Chrysophtharta bimaculata (Olivier), Tasmanian Eucalyptus Leaf Beetle Distribution This beetle is endemic to the Australian island of Tasmania.

Hosts Several species of eucalypts are the hosts, including Eucalyptus delegatensis, E. nitens and E. regnans.

Life History There is one generation/year. However, trees in any one area may experience more than one peak of egg laying and defoliation because of extended adult activity. Females deposit eggs in masses on young foliage in spring and summer. Eggs hatch in about 10 days and young larvae begin to feed in groups. They consume all of the foliage except the midribs and some of the leaf margin. Larvae feed for about 3–4 weeks. When feeding is complete, larvae drop to the ground, burrow into the soil and pupate in earthen chambers. Adults emerge in late summer–early autumn and feed before dispersing to overwintering sites under bark and in litter.

Description of Stages Adults are 5–10 mm long, pale green with two black marks on the prothorax. During winter, they turn deep red-brown but change back to pale green when they resume activity the following spring. Larvae are dark green and 15–20 mm long when fully grown.

Pest Management Management of this leaf beetle involves use of both chemical and biological controls in combination with intensive monitoring of population levels and development of trees genetically resistant to feeding.

Related Species A complex of related leaf beetles, including C. agricola (Chapuis), C. nobilitata (Erichson), C. varicollis (Chapuis), Paropsis atomaria Olivier and P. charybdis (Stål), defoliate eucalypts and are pests of plantations in portions of the Australian mainland as well as Tasmania. P. charybdis has been introduced into New Zealand, where it has become a pest (Ohmart & Edwards 1991, Phillips 1994, Nahrung 2006, Elliott & deLittle n.d.). Diorhabda spp., Diorhabda Beetles (Plates 31 & 32) Distribution Several species of Diorhabda feed on foliage of salt cedars, Tamarix spp. Their taxonomy is

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complex and the group was again revised in 2009. D. elongata (Brulle) is a Mediterranean species found from Italy to western Turkey. D. carinata (Falderman) occurs in grasslands, deserts, and forests from southern Ukraine south to Iraq and west to Kazakhstan. D. sublineata (Lucas) occupies Mediterranean woodlands from France to north Africa and subtropical deserts east to Iraq. D. carinulata (Desbrochers) (¼ D. elongata deserticola Chen) inhabits cold temperate deserts of Mongolia and China west to Russia and south to montane grasslands and warm deserts in southern Iran. D. meridionalis Berti & Rapilly occurs in maritime subtropical deserts of Iran, southern Pakistan and Syria.

Pest Management In China and Russia, infestations are treated with insecticides to prevent defoliation and plantation mortality (DeLoach et al. 2003, Lewis et al. 2003, Hudgeons et al. 2007, Tracy & Robbins 2009).

Hosts In their native habitats, these insects are associated with salt cedar, Tamarix spp., and occasionally species of Myricaria.

Hosts Primary hosts are black locust, Robinia pseudoacacia and honey locust, Gleditsia triacanthos. Species of Betula, Crataegus, Fagus, Malus, Prunus, Quercus and Ulmus are occasionally attacked.

Damage Larvae and adults damage foliage by removing and eating sections of the leaves and by scraping tissue from the leaves and green stems. This causes large sections of stems to desiccate and causes death of more plant tissue than is actually consumed. In China and Russia, where salt cedars are planted for shelterbelts and sand dune stabilization, they are considered pests. However, in southwestern USA, where salt cedars are exotic and invasive along riparian areas, these beetles have been successfully introduced as classic biological control agents.

Life History In China, D. carinulata normally has three generations/year. In southwestern USA, there are two generations/year in most sites where this insect has been released. Adults of the first generation emerge in early to mid-July and second generation adults emerge from mid-August to early September and overwinter. Overwintering adults emerge in spring and females lay eggs singly or in clusters of up to 14 on tender leaves. Eggs hatch 7–10 days later. Larvae have three instars and when finished feeding, drop to the ground and pupate in litter or loose soil.

Description of Stages Adults are 5–6 mm long, yellow with two black stripes on their back. Young larvae are black but mature larvae have a broad yellow stripe on each side and are about 8 mm long.

Odontata dorsalis (Thunberg), Locust Leaf Miner Distribution The original range of this leaf beetle was eastern USA, including the central Appalachian, Ozark and Ouachita Mountains. Naturalized populations now occur throughout the USA and southern Canada.

Importance Adults skeletonize and eat holes in leaves and larvae mine leaf tissue. Leaf mining is most damaging (Fig. 8.2). Outbreaks are fairly common and turn entire stands of black locust gray or brown but are more spectacular than destructive. In combination with other stress factors, outbreaks can contribute to growth loss and occasional tree mortality.

Life History There are two generations/year. Adults overwinter in bark crevices or leaf litter and emerge as leaves unfold in spring. Eggs are deposited on the undersides of leaflets. They overlap like shingles in groups of three to five and are cemented together by excrement. When eggs hatch, larvae feed collectively in common blister-like mines. Later, they disperse and excavate individual mines and pupate within the mines.

Description of Stages Adults are small, elongated, flat beetles, 5–6 mm long. The head is black and the elytra are orange with a broad black or brown stripe down the center. Mature larvae are yellow-white, flat and slightly larger than adults.

Pest Management Direct control is usually not necessary. Individual trees near homesites can be

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Forest entomology: a global perspective adults overwinter in attics, barns and sheds and are a nuisance because they are active on warm days and crawl into living quarters. Life History Elm leaf beetle can have two and a partial third generation/year. Adults overwinter and become active in spring as buds burst and leaves unfold. They chew holes in unfolding leaves and deposit eggs. Each female can deposit 400–800 eggs. Eggs hatch a week later and larvae feed for 2–3 weeks on the undersides of leaves. Damaged leaves dry out and turn brown. Mature larvae crawl to the base of the tree and pupate in bark crevasses. Adults overwinter. Description of Stages Adult are about 6 mm long, yellow-metallic green in color, with a dark stripe along the sides of the elytra. Mature larvae are about 12 mm long, dull yellow with two rows of black tubercles on the dorsal surface. The head and legs are black and there is a broad yellow stripe on the dorsal surface of the abdomen. Pupae are about 5 mm long, bright orangeyellow with a few black bristles (Plates 33 & 34). Pest Management Infestations on urban trees can be treated with ground applications of contact insecticides (Drooz 1985).

Fig. 8.2 Leaf mining on foliage of Robinia pseudoacacia by locust leaf miner, Odontata dorsalis (northern Virginia, USA).

protected with systemic insecticides (Drooz 1985, USDA Forest Service 2001). Pyrrhalta luteola (Müller), Elm Leaf Beetle Distribution Elm leaf beetle is native to Europe and was introduced into North America during the late 1800s. It is now found throughout southern Canada and the entire USA. Hosts

All species of elms, Ulmus spp., are hosts.

Importance Both larvae and adults feed on foliage. Larvae skeletonize leaves and adults chew larger holes. This insect is primarily a pest of urban trees and heavy skeletonizing and defoliation is unsightly. Moreover,

Curculionidae (Weevils) Gonipterus scutellatus Gyllenhal, Eucalyptus Weevil Distribution This weevil is native to Australia but has been introduced into many regions where eucalypts are grown. In Africa, it is found in Kenya, Lesotho, Madagascar, Malawi, Mauritius, Mozambique, South Africa, St Helena, Swaziland, Uganda and Zimbabwe. In Asia, it has been introduced and established in Zhejiang Province, China. In Europe it is established in France, Italy, Portugal and Spain. In North America it has been introduced into California, USA and in South America, infestations are known from Argentina, Brazil, Chile and Uruguay. Hosts Hosts are species of Eucalyptus, including E. camaldulensis, E. maidenii, E. punctata, E. robusta, E. smithii and E. viminalis.

Other foliage feeding insects Importance This weevil causes little or no damage in Australia. However, in places where it has been introduced and become established it is an important defoliator of eucalyptus plantations. Both larvae and adults feed on host trees but larvae are most damaging. Defoliation causes growth loss and tree mortality. In 1940, infestations were discovered on the island of Mauritius and 4 years later it caused severe defoliation of E. viminalis.

Life History Life history and habits of this weevil vary with local climatic conditions. In Mauritius, there are four generations/year and they are active year long. In South Africa, there are two to two and a half generations/year and in Italy, two generations/year are reported. In Mauritius, eggs are attached to leaves in gray capsules that contain 8–10 eggs. Females mate several times and continue to lay eggs over their lifetime, about 91 days. A female can deposit from 21 to 33 egg capsules. Larvae feed on leaves and twigs and pupation occurs in cells in the soil about 5 cm deep.

Description of Stages Adults are 12–14 mm long and vary from gray to red-brown in color. They have a light transverse band on the back and are covered with pale brown hairs. Larvae are yellow-green with black marks and are about 14 mm long when mature.

Pest Management Classic biological control with the egg parasitoid Anaphes nitens has been undertaken France, Italy and the USA. Chemical control of infestations is not recommended because of the danger to honeybees that are attracted to Eucalyptus flowers. Related Species G. gibberus Boisduval is also native to Australia and causes similar damage. This species has been introduced into Argentina, Brazil and Uruguay. Adults of the two species are difficult to separate visually but can be identified by examination of the genitalia (EPPO 2005a, FAO 2009b).

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Hosts European ash, Fraxinus excelsior, is the primary host. Other hosts include Phillyrea latifolia and Syringa vulgaris. Olive, Olea europaea, is an occasional host. Importance Adults feed on unopened buds of host plants. This delays appearance of new growth. Both larvae and adults feed on foliage. Larvae feed on the surface of the leaves and cause window-like blotches. Adult feeding produces holes in the foliage. In Romania, this insect is a pest of forests where 40% of the tree component is ash and is often associated with other forest defoliators. One study suggests that ash saplings that have suffered root damage are more likely to suffer feeding damage by this weevil. Life History Stereonychus fraxini has one generation/ year and adults overwinter in the litter or other sheltered places. Adults emerge in spring, feed on buds and, later, leaves and leaf petioles. After mating, females deposit eggs on the undersides of expanded leaves. Larvae feed on the leaf surface. Feeding is completed in late summer and pupation occurs in oval yellow-brown or brown, parchment-like cocoons on the leaf surface. Description of Stages Adults are about 3 mm long, gray-brown to red-brown with a dark thoracic disk. Eyes are located at the top of the head. The body is covered with white or gray-white pubescence. Larvae are 4 mm long when mature, slug-like in appearance and yellow-green in color. The head capsule is black (Alford 2007, Foggo et al. 2008). HYMENOPTERA (BEES AND WASPS) Most defoliating insects of the order Hymenoptera are sawflies of the suborder Symphyta. Adult females have saw-like ovipositors and usually deposit eggs individually in slits or pouches cut into leaves (Fig. 8.3). A number of species, representing several families, are of economic importance worldwide. Pergidae

Stereonychus fraxini (DeGeer), Ash Weevil Distribution This weevil is widely distributed across much of south central and central Europe, the Near East and North Africa.

Cerospastus volupis Konow Distribution This sawfly is known from south central Argentina and Chile.

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Fig. 8.3 Eggs of the pine sawfly, Neodiprion nanulus contortae, deposited individually in slits in a pine needle (Gallatin Gateway, Montana, USA).

Hosts The primary host is Nothofagus alpina but feeding has also been reported on N. obliqua.

Importance Larvae feed on foliage in the upper third of the crowns. Early instar larvae are skeletonizers and mature larvae consume all of the leaf. Heavy defoliation causes growth loss.

are brown. Length is 12–14 mm. Males are similar in appearance but smaller. Antennae are shorter than females but have more segments (15–19 for females, 22–23 for males). Mature larvae are about 24 mm long with an amber head, pale green legs and black prolegs. Body color is green with two lateral yellow bands (Fig. 8.4, Aguayo Silva et al. 2008).

Perga affinis Riek, Spitfire Grubs or Sawflies Life History This sawfly has one generation/year. Adults are active from late October to late January. They are not strong fliers and their capacity for dispersal is low. Females deposit eggs in groups of three to four in perforations in the foliage and lay 40–70 eggs. Eggs hatch within several days and larvae are present from mid-December to mid-April. Early instar larvae feed in colonies and later individually. Pupation occurs in cocoons in the soil.

Description of Stages Females are yellow-brown with dark bands on the abdomen. Wings are transparent yellow with dark brown veins. Antennae and legs

Distribution Three subspecies are recognized: P. affinis affinis Kirby, the steel blue sawfly, P. affinis insularis Risk, the large green sawfly and P. affinis atrata. All are indigenous to southeastern Australia, including the island of Tasmania.

Hosts Subspecies of P. affinis feed on foliage of many species of eucalypts.

Importance Larvae feed during winter and early spring when foliage removal has a low impact on host

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Fig. 8.4 Larval colony of the sawfly, Cerospastus volupis, on foliage of Nothofagus alpina (X Region, Chile).

trees. Therefore, trees can usually withstand repeated defoliation. This sawfly can cause complete defoliation of individual trees but outbreaks over large areas are unusual.

Life History Individuals may require 1–4 years to complete a generation. This is dependent on the length of pupal diapause, which typically lasts from 1 to 2 years. Adults are active in late summer (February– March). Females deposit an average of 56 eggs in a single row under the leaf surface using their serrated ovipositor to cut into the leaf tissue. Larvae are present during winter (March–November) and feed at night. During daylight hours they cluster in large groups on branches or foliage. While feeding, they maintain contact with one another by tapping the foliage or branches. When disturbed, they display a defensive reaction that consists of raising their heads in unison and regurgitating a yellow fluid that smells of eucalyptus oil. This habit gives them the common name “spitfire grubs.” If total defoliation of host trees occurs before the larval cycle is finished, they migrate as a colony to neighboring trees. In spring (September), larvae crawl down the tree, form large masses of brown

cocoons in the soil and diapause as pre-pupal larvae. Diapause usually lasts for 1–2 years. However, some individuals may remain in this stage for up to 4 years. Pupation occurs in late summer (January–February) and adults emerge from the soil 2–3 weeks later.

Description of Stages Adults are robust sawflies, 22 mm long, with a wingspan of 40 mm. Males are slightly smaller than females. Eggs are 4 mm long, spindle shaped and blue green. Larvae are predominantly brown to black and have short white hairs in the later instars. They are about 70 mm long when mature.

Related Species Other sawflies of the family Pergidae that feed on eucalypt foliage in Australia include the pale brown sawfly, Pseudoperga lewisii (Westwood), Pergagrapta bella (Newman) and the cattle poisoning sawfly, Lophyrotoma interrupta (Klug). The latter species is so named because the larvae contain a toxin, lophyrotomin, and can poison livestock if eaten (Ohmart & Edwards 1991, Elliott & Bashford 1995, Schmidt 2006, Fletcher 2008, Elliot & DeLittle n.d.).

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Argidae Sericoceros About 20 species make up this neotropical genus of sawflies. They occur from southern Mexico south to Argentina with one species in the West Indies. Most feed on species of Coccoloba (family Polygonaceae); however, one species is reported from Lonchocarpus spp. and another from Triplaris caracasana (Smith 1992, Smith & Benitez Dias 2001). Sericoceros mexicanus (Kirby) (Plates 35 & 36) Distribution S. mexicanus occurs from Chiapas, southern Mexico south to Panama.

Hosts Hosts are species of Coccoloba (sea grape), including C. uvifera and C. venosa.

Importance This sawfly can cause moderate to heavy defoliation. However, it is more a curiosity than a damaging pest. On Roatan Island, Honduras, where infestations are relatively common, backstrap weavers have captured a likeness of adult females in their weavings (see Fig. 3.8).

Life History Information on life history and habits is incomplete. There are at least two generations/year. Adults are strong fliers and swarm around host trees on warm, sunny days to mate and deposit eggs. During cool or overcast days, they rest on foliage and branches. One period of adult activity is early January. Eggs are deposited in circular or slightly oval clusters on the undersides of leaves. Mean egg cluster size is 32 eggs and ranges from 7 to 85 eggs. Females remain with the egg cluster until they die. Eggs hatch within 2–3 days and larvae feed gregariously on edges of leaves, consuming all of the leaf tissue except the major veins. Larvae undergo five or six instars. Pupation occurs in parchment-like cocoons attached to leaves, leaf petioles and branches of sea grape. Occasionally, cocoons are attached to wooden surfaces, such as picnic benches.

Description of Stages Females are about 12 mm long with a bright red-orange thorax and abdomen.

Wings are clear with black veins. Head, antennae and legs are dark brown to black. Antennae are filiform. Males are slightly smaller, 8–10 mm long; the thorax and ventral surface of the abdomen are bright redorange and the dorsal surface of each abdominal segment is red-orange with a black band. Wings are similar to females but smaller. Head, antennae and legs are dark brown to black and antennae are forked at the third antennal segment. Eggs are bright red and about 2 mm long when first deposited, later turning to pale orange or pink. First instar larvae are orange at first but take on a green cast when they begin feeding. For later instars, the head capsule is pale yellow immediately after molting and changes to medium or dark brown. Black eyespots are conspicuous even on larvae with dark brown head capsules. Legs are brown and prolegs are white and non-functional. The body surface is shiny and yellow-green on the dorsal and lateral surfaces, turning to yellow on the ventral surface. Both thoracic and abdominal segments have rows of tubercles that become more conspicuous in the later instars. The last two abdominal segments are often yellow. Overall length of mature (instar VI) larvae is 25–28 mm.

Related Species S. krugii (Cresson) is reported from the Dominican Republic, Puerto Rico and St Thomas, US Virgin Islands, where it feeds on several species of Coccoloba. In Puerto Rico, infestations can be found from sea level to about 825 m (Martorell 1941, Smith 1992, Ciesla 2002b, Smith & Janzen 2002).

Diprionidae (Conifer Sawflies) Gilpinia Gilpinia is a genus of about 20 species distributed across Asia and Europe (Table 8.1). Several are pests of conifer forests in their native habitat and at least one species has been introduced and become a pest in North America. Gilpinia (¼ Diprion) hercyniae (Hartig), European Spruce Sawfly Distribution This sawfly is native to continental Europe. It was first recorded in the UK in 1906 and achieved pest status in Wales between 1970 and 1974. It was discovered in North America near Ottawa,

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Table 8.1 Representative species of Gilpinia (Hymenoptera: Diprionidae): their distribution and hosts. Species

Distribution

Hosts

G. abieticola (Dalla Torre) G. fennica (Forsius) G. frutetorum Fabricius

Europe

Picea spp.

Finland Central and eastern Europe, northwestern Asia. Introduced into eastern Canada Thailand Thailand Northern and central Europe Pakistan Central eastern and northern Europe Native to continental Europe, introduced into the UK and North America Northern Asia and Europe

Picea spp. Pinus sylvestris (native range) P. resinosa (introduced range) Pinus kesiya Pinus kesiya Pinus sylvestris Abies pindrow Picea Picea

G. leksawasdii Smith G. marshalli (Forsius) G. pallida (Klug) G. pindrowi Benson G. polytoma Hartig G. hercyniae (Hartig) European spruce sawfly G. virens Klug

Pinus sylvestris

a & Roller 2004. Sources: Browne 1968, Drooz 1985, Leksawasdi et al. 1990., Holuc

Canada, in 1922 and has since spread over much of southeastern Canada and northeastern and north central USA.

Hosts This species feeds on all species of spruce, Picea spp. In Europe, P. abies is the principal host and in the UK, P. sitchensis, native to the Pacific Coast of North America and widely planted in the UK, is damaged. P. glauca is the favorite host in eastern North America but other species of Picea are also attacked.

Importance During outbreaks, defoliation of variable severity can occur. Larvae feed mainly on old foliage and, on rare occasions, new needles. Severe defoliation reduces the tree’s growth and vigor, and can kill trees that are completely defoliated. A severe outbreak occurred on the Gaspe Peninsula of eastern Canada from 1932 to 1935 that encompassed 15,500 km2 and caused severe tree mortality. An outbreak in Wales reduced height, radial and volume growth by 24–49%, 30–57% and 32–56%, respectively.

Life History Depending on location, there may be one or two generations/year. Adults appear from May to June. Most first generation adults are females and reproduction is parthenogenic. Eggs are laid in slits cut in pine needles and they hatch within several days. Larvae of all ages feed singly. Pupation occurs in

cocoons in the soil or litter. Second generation adults appear in early July and lay eggs. Second-generation larvae drop to the soil, pupate and overwinter with onset of cool weather.

Description of Stages Females are stout bodied and 6.0–8.5 mm long. Head and body are black, except for a cream stripe above the eyes and on the thorax. The abdomen has black and yellow bands. Males are slightly smaller. The abdomen is mostly black and the pronotum is marked with yellow. Larvae are yellow green when first hatched. Mature larvae are dark green with five longitudinal white lines on the body and are about 22 mm long.

Pest Management Exotic parasitoids have been released in Canada and the USA to help control outbreaks. Several have become established. In 1942, a virus disease caused a collapse of outbreaks in eastern North America and the insect is now considered a minor pest.

Related Species G. ventralis feeds on Pinus sylvestris and damages pine forests in Sweden. The nursery pine sawfly, G. frutetorum (Fabricius), has a Eurasian distribution and also feeds on Pinus. This sawfly has also been introduced into North America (Hedqvist 1972, Billany & Brown 1977, Drooz 1985, Williams et al. 2003).

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Neodiprion Species of Neodiprion are the most damaging of all of the sawflies and during outbreaks cause extensive defoliation (Fig. 8.5). Larvae feed almost exclusively on old foliage of conifers of the family Pinaceae, especially Abies, Picea, Pinus and Tsuga. About 47 species are known and all but seven, the European pine sawfly, N. sertifer and 6 species in China, are native to North America (Table 8.2). Outbreaks are usually of short duration, lasting from 2 to 4 years. Occurrence of conifer forests on poor sites, outwash plains or shallow, infertile soils increases the probability of outbreaks. Forests most susceptible to outbreaks are open-grown stands or even-aged plantations located on poor

growing sites (Ross 1955, Xiao Gangrou et al. 1985, McMillin & Wagner 1993). Neodiprion autumnalis Smith & Wagner (Plates 37 & 38) Distribution N. autumnalis is a North American species widely distributed over western USA and throughout pine growing regions of Mexico. A population studied in California and reported as being of the “N. fulviceps complex” is believed to be N. autumnalis. Hosts In western USA, Pinus ponderosa is the host. In Mexico, this species feeds on foliage of P. arizonica, P. engelmannii and P. teocote.

Importance This sawfly causes moderate to heavy defoliation of pole-sized and larger trees. Repeated defoliation causes growth loss. Outbreaks have been reported in western South Dakota (1400 ha), eastern Wyoming (34,000 ha) and in ponderosa pine– grassland transition zones in central Colorado. Widely spaced trees on dry sites with poor soils tend to be most severely affected.

Life History N. autumnalis has one generation/year and overwinters as eggs deposited in pine needles. Larvae hatch in spring and feed until mid-summer, after which they drop to the ground, spin cocoons and pupate (Fig. 8.6). Adults emerge in autumn, hence the species name autumnalis, and deposit eggs individually in pine needles.

Description of Stages Female adults are 8.5–10 mm long with orange heads, black antennae and a pale orange thorax with black stripes. Males are 7.5–8.5 mm long and black. Mature larvae are 17–23 mm long. The head capsule is honey-yellow to orange with a black eyespot. Body color is yellow-green to green, usually with two dark longitudinal subdorsal stripes.

Fig. 8.5 Heavy defoliation of loblolly pine, Pinus taeda, by the loblolly pine sawfly, Neodiprion taedae linearis (near Tullos, Louisiana, USA).

Pest Management In some areas, forest landowners have used aerial applications of chemical insecticides to reduce defoliation.

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Table 8.2 Representative species of Neodiprion (Hymenoptera: Diprionidae): their distribution and hosts. Species

Distribution

Hosts

N. abietis (Harris) Balsam fir sawfly

Eastern North America, also Alberta and British Columbia, Canada, California, Oregon, USA Western North America

Abies spp., Picea spp.

N. autumnalis Smith N. dialingensis Xiao & Zhou Dailing pine sawfly N. edulicolus Ross Pin˜on sawfly N. fulviceps (Cresson) N. lecontei (Fitch) Red headed pine sawfly N. nanulus nanulus Schedl Red pine sawfly N. nanulus contortae

Northeastern China Southwestern USA: Arizona, Colorado, New Mexico, Nevada, Utah Southewestern USA: Arizona, Nevada Mexico: Chiapas, Hidalgo, Vera Cruz Eastern North America

N. pinetum (Norton)

Nova Scotia west to British Columbia, Canada and south to Pennsylvania, USA Canada: Alberta, British Columbia USA: Idaho, Montana, Oregon Southeastern Canada, eastern USA

N. pratti pratti (Dyar) Virginia pine sawfly N. sertifer (Geoffrey) European pine sawfly N. swainei Middleton

Eastern USA: Illinois Maryland, North Carolina, Virginia Europe Introduced into North America Eastern Canada, north central USA

N. taedae linearis Ross Loblolly pine sawfly

Southern USA: Arkansas, Illinois, Louisiana, Mississippi, Missouri, Ohio, South Carolina, east Texas Canada, USA: southeastern Alaska, British Columbia, Oregon and Washington China

N. tsugae Middleton Hemlock sawfly N. xiangyunicus Xiao & Zhou

Pinus ponderosa (USA), P. arizonica, P. engelmannii, P. teocote (Mexico) Pinus tabulaeformis Pinus edulis, P. monophylla Pinus ponderosa, Pinus spp. Pinus spp. Pinus banksiana, P. resinosa Pinus contorta, P. ponderosa Pinus cembra, P. echinata, P. resinosa, P. rigida, P. strobus Pinus echinata, P. virginiana Pinus spp. Pinus banksiana, P. resinosa, P. strobus, P. sylvestris Pinus taeda

Tsuga heterophylla Pinus yunnanensis

Sources: Ross 1955, Furniss & Carolin 1977, Wilson & Averill 1978, Drooz 1985, Sheehan & Dahlsten 1985, Xiao Gangrou et al. 1985, Dunbar n Tovar et al. 1995, Anderbrandt et al. 1997. & Wagner 1990, Hengxiao et al. 1993, Cibria

Related Species N. fulviceps (Cresson) is similar in appearance and also feeds on P. ponderosa. This species overwinters in cocoons in the soil and adults emerge in spring to lay eggs (Ross 1955, Dahlsten 1966, Smith & Wagner 1986, Dunbar & Wagner 1990, Pasek 1991a,b, Cibrian Tovar et al. 1995). Neodiprion lecontei (Fitch), Red Headed Pine Sawfly Distribution This sawfly occurs throughout eastern North America from eastern Canada, west to the Great Plains and south to Florida and Texas, USA.

Hosts N. lecontei feeds on many pines, both native and exotic. It prefers the hard or yellow pines and host preference varies with location. In Canada and northern USA, Pinus banksiana and P. resinosa are preferred. Further south, P. echinata, P. elliotii, P. palustris and P. taeda are preferred hosts.

Importance Young pine less than 5 m tall are preferred and epidemics often occur in plantations. This species became a pest during the 1930s following establishment of extensive pine plantations in eastern North America. Moderate to heavy defoliation causes growth loss and may cause branches in the upper

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Fig. 8.6 Cocoons of the pine sawfly, Neodiprion autumnalis (Elbert County, Colorado, USA).

crown to fork. Complete stripping of P. banksiana and P. resinosa can kill trees. However, southern yellow pines can survive heavy defoliation even in areas where several generations of sawflies occur in one season.

Life History The number of generations varies with location. In most of Canada and northern USA, there is one generation/year. At the latitude of Michigan and New York, there is either a partial or complete second generation. Further south, three to five generations/ year may occur. In places where there is more than one generation, they may overlap and larval colonies of different ages may be present at the same time. Pre-pupal larvae overwinter in cocoons spun in litter beneath host trees. Pupation occurs in early spring and adults emerge several weeks later. Some pupae may remain in extended diapause for several years. Females deposit about 120 eggs in the current or previous year’s needles. Each egg is deposited individually in a slit cut into the needle. All eggs laid by a single female are clustered on the needles of a single branch. Eggs hatch 3–5 weeks later and larvae feed in colonies for 25–30 days. When mature, they drop to the litter,

spin cocoons and pupate. Winter is spent as pre-pupal larvae in the soil. Description of Stages Adults are 5–10 mm long with males smaller than females. Females are robust, with a red-brown head and thorax and black abdomen with white sides. Males are slender, all black and have feathery antennae. Newly hatched larvae are about 3 mm long and have brown, transparent heads. When mature, they are nearly 25 mm long with a red head. Body color varies from pale yellow-white to deep yellow and is marked by two to four rows of black spots on each side of the abdomen. The last abdominal segment has a large black patch on each side (Fig. 8.7). Pest Management Risk of outbreaks can be minimized by not planting pines on sites with marginal soil moisture and nutrients. When only a few colonies of larvae are present on roadside or ornamental trees they can be picked or shaken off trees and destroyed. Outbreaks may require aerial or ground applications of chemical insecticides (Wilson & Averill 1978, Drooz 1985).

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Fig. 8.7 Mature larvae of red headed pine sawfly, Neodiprion lecontei (photo by R.F. Billings, Texas Forest Service).

Neodiprion sertifer (Geoffroy), European Pine Sawfly Distribution This sawfly is native to most of Europe and across northern Asia to Japan. In the Alps, it occurs as an alpine form with a smaller body size. It was first reported in North America in 1925 and is now widely distributed in eastern USA and portions of Ontario and British Columbia, Canada.

Hosts European pine sawfly has a wide host range that includes most hard or yellow species of Pinus. In Europe, the favorite host is Scotch pine, Pinus sylvestris.

Importance Outbreaks tend to be cyclic and last 2–3 years. In Belgium and Germany, trees less than 25 years old are preferred. In Austria and Finland, it has a preference for older trees. Defoliation weakens trees and predisposes them to bark beetles and wood borers.

In Finland, Italy and Sweden, it is considered the most damaging sawfly of the family Diprionidae.

Life History There is one generation/year over most of its range. In Norway, 2 years are required to complete a generation. Eggs overwinter and hatch from April to mid-May. Larvae are present until mid-July. They may drop to the ground or remain in the tree to pupate in golden-brown cocoons. Adults emerge from late August to September and lay eggs in pine needles from late September to October.

Description of Stages Mature larvae are about 25 mm long with a black, unmarked head capsule. Body color is gray-green with a dark bordered light gray mid-dorsal pinstripe, a gray-green dorsal stripe, a dark spiracular stripe bordered by a light greenish gray supraspiracular stripe and a light gray subspiracular stripe. Each abdominal segment is marked with a pair of

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black annules. Pre-pupal larvae are brown with a black mid-dorsal stripe.

Pest Management Outbreaks collapse due to a naturally occurring virus. In the USA, this virus has been mass produced in the laboratory and registered as Neocheck-SÒ for direct control of outbreaks (PschornWalcher 1982, Podgewaite et al. 1984, Drooz 1985, Duncan 2007).

Nesodiprion Nesodiprion is a genus of conifer infesting sawflies indigenous to Asia. Several species are important pests of pines, especially plantations.

Nesodiprion biremis (Konow) Distribution This species is known from Indonesia (Sumatra) and Thailand.

Hosts Hosts are Pinus spp., including P. kesiya and P. merkusii. Larvae can feed on both native and exotic pines.

Importance Larvae feed on old foliage of host trees and, unless foliage is in short supply, do not feed on the current year’s growth. In northern Sumatra, Indonesia, it reportedly causes only sporadic defoliation and is not considered a pest. However, it is one of several sawflies of concern in pine nurseries and young plantations in the mountainous regions of northern Thailand.

Life History There are five to six overlapping generations/year. Adults deposit eggs singly on the needles of host trees. Several eggs are deposited on a needle. Larvae feed almost entirely on mature ( > 1 year old) needles. Both sexes pass through six instars. About 6 weeks are required to complete a generation.

Description of Stages Adult males average 6.0 mm in length and females 7.5 mm. Both sexes are black in color. Larvae are almost 22 mm long when mature.

Related Species N. japonica (Marlatt) occurs in northern China and Japan, where it feeds on foliage of Larix and Pinus (Beaver & Laosunthorn 1975, Leksawasdi et al. 1990, Nair 2001, 2007).

Zadiprion Zadiprion is a small genus composed of five North American sawflies. Two species occur in western USA and the others in Mexico. They resemble the sawflies of the genus Neodiprion in appearance and habits (Smith 1971, 1974, Cibri an Tovar et al. 1995).

Zadiprion falsus Smith (¼ Z.Vallicola Rohwer) Distribution This sawfly is found in northern and central Mexico, including the states of Chihuahua, Durango, Jalisco, Michoac an and Mexico.

Hosts Several species of Mexican pines are hosts. These include Pinus arizonica, P. ayacahuite, P. engelmannii, P. durangensis, P. leiophylla, P. michoacana, P. montezumae, P. oocarpa, P. pseudostrobus, P. radiata and P. teocote.

Importance Outbreaks cause moderate to heavy defoliation of pine forests. Successive defoliation causes reduced tree vigor, lower resin production and increased susceptibility to other pests. In areas where defoliation is heavy and continuous over several years, extensive tree mortality may occur. Several outbreaks have occurred. In 1927, some 25,000 ha of P. ayacahuite were defoliated in Michoac an. In 1971, 60,000 ha of P. michoacana, P. montezumae and P. pseudostrobus forests were defoliated.

Life History There is one generation/year. Adults are active between mid-July and late September. Females deposit eggs in typical conifer sawfly fashion in slits cut into pine needles. They deposit an average of 47 eggs, all on a single fascicle of pine needles. Eggs hatch on average 43 days later. Young larvae feed in colonies of two to five on pine needles. Male larvae undergo five instars and females six. Most severe defoliation is caused in November, beginning with instar IV. Feeding is usually completed by late

Other foliage feeding insects December when larvae drop to the ground to form cocoons. Larvae remain in the cocoons until March of the following year when they pupate. Pupation requires about 1 month.

Description of Stages Male adults are 7–8.7 mm long and black except for legs and ventral portions of the abdomen, which are pale yellow. Wings are membranous and transparent. Females are slightly larger and average 9–10 mm in length. They are lighter in color than males with a brown head and mostly yellow abdominal segments. Mature larvae are 25–30 mm long. The head is brown with conspicuous darker eyespots. Body color varies from light green, brown or pink-violet. Abdominal segments have eight pairs of prolegs.

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Related Species Z. rohweri (Middleton) (Fig. 8.8) occurs in southwestern USA. Larvae feed on older needles of Pinus monophylla and P. edulis and sometimes completely defoliate small trees. In 1998, this species defoliated P. edulis of all age classes on 21,000 ha in Arizona. It also defoliated P. edulis in Mesa Verde National Park, Colorado from 2006 to 2008. Z. townsendi (Cockrell) (¼ grandis (Rohwer), sometimes known as the “bull pine sawfly,” feeds on P. ponderosa in western USA (Smith 1971, Dunbar & Wagner 1990, Cibri an Tovar et al. 1995, USDA Forest Service 1999).

Tenthredinidae (Tenthredinid or Common Sawflies) Fenusa pusilla (Lepeletier), Birch Leaf Miner

Pest Management Heavy infestations have been treated successfully with contact insecticides. The optimum time for application of pesticides is October after most eggs have hatched and before heavy feeding damage occurs.

Distribution Birch leaf miner is native to Europe. It was introduced into North America during the early 1900s and now occurs from Newfoundland, Canada, west to Washington and Oregon and north to Alaska, USA.

Fig. 8.8 Larvae of the sawfly, Zadiprion rohweri, feeding on foliage of Pinus edulis (Mesa Verde National Park, Colorado, USA).

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Hosts This leaf miner infests birches, Betula spp. Susceptible species include B. ermanii, B. glandulifera, B. maximowicziana, B. papyrifera, B. pendula, B. platyphylla variety japonica, B. populifolia and B. turkestanica. Species of intermediate and variable susceptibility are B. costata and B. davurica. Larvae are unable to survive in B. alleghaniensis, B. grossa and B. lenta.

Importance Larvae mine in leaves and cause discoloration. Outbreaks in forested areas can occur over thousands of hectares. In urban areas, discoloration due to leaf mining is unsightly. Repeated defoliation can weaken trees and, in North America, feeding damage makes trees susceptible to attack by bronze birch borer, Agrilus anxius.

Life History This sawfly can have up to four generations/year. Pupation occurs in early spring and adults appear in mid-May. Eggs are deposited singly in slits cut near the center of leaves. Larvae feed on leaf tissue between the surfaces. At first they feed singly and create a kidney-shaped mine near where the egg was deposited. Later the mines coalesce and cause much

of the leaf to turn brown. Mature larvae chew out of the leaf and drop to the litter where they form cells 2.5–5 cm below the surface. They overwinter as prepupae in cocoons.

Description of Stages Adults are small sawflies, 3.7 mm long. They are black, except for small areas of white on the legs. The wings have light brown bands and are darker near the body. Mature larvae are somewhat flattened, yellow-white and about 6 mm long. They have black spots on the lower surface of the thorax.

Pest Management Application of chemical insecticides to urban trees shortly after budburst can prevent infestation. Two palearctic parasitoids, Lathrolestes nigricollis (Thomson) and Grypocentrus albipes Ruthe, have been introduced and established in Canada.

Related Species Elm leaf miner, F. ulmi Sundevel, causes similar damage to foliage of Ulmus spp. (Fig. 8.9) and F. dohrni (Tischbein) causes similar damage to

Fig. 8.9 Larvae of the leaf mining sawfly, Fenusa ulmi, inside leaf of American elm, Ulmus americana (Fort Collins, Colorado, USA).

Other foliage feeding insects Alnus. Both species are native to Europe and have been introduced into North America (Drooz 1985, Hoch et al. 2000, Langor et al. 2002). Nematus oligospilus (¼ N. desantisi) Forster Distribution This sawfly has a holarctic distribution and is found in Europe from Ireland east to the Himalayas and North America from Alaska, USA, south to Mexico. It has been introduced in several areas of the southern hemisphere, including Argentina (1980–1), Australia (2004), New Zealand (1997) and southern Africa. Hosts Most species of willow, Salix spp., and poplar, Populus spp., are hosts.

Importance N. oligospilus rarely reaches epidemic proportions in its native range. In Argentina, infestations were initially detected in Chubut Province. They spread rapidly, covering an area of 3000 km2 over a period of 9–10 years. In Argentina, it causes heavy defoliation in plantations. Poplars are an important plantation species in the Patagonia region of Argentina and the potential for severe damage is considered high. In 1997, an estimated 15,000 ha of Salix plantations were defoliated in the Delta del Parana of northern Argentina. Severe defoliation can cause a 60% reduction in annual increment and some tree mortality. In New Zealand, willow plantings have been established along river banks where their extensive root systems limit soil erosion during flooding. Occurrence of a major defoliator could reduce the value of willow for stream bank stabilization.

Life History This sawfly can complete four to six generations/year and overwintering occurs in cocoons in the leaf litter near the base of host trees. Males occur in its natural range in the northern hemisphere but thus far only females have been found in the southern hemisphere. They lay eggs in pouches, usually on the upper surface of leaves and may lay more than one egg on the same leaf. Larvae undergo from five to seven instars and feed on edges of leaf margins.

Description of Stages Adults are 7–8 mm long and antennae have nine segments. The head and thorax are

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yellow or orange-brown and shiny and the abdomen is green. Large dark eyes are prominent on the head. Larvae are yellow-green with a light brown head and may have dark brown stripes on the body. The head is cream colored with black eyes. A brown stripe runs from just behind each eye to the top of the head in the late larval instars. They may have a brown triangle on the top of the head and pale brown stripes run down the center of the head. Mature larvae are about 20 mm long and often curl up into a distinctive S shape. Cocoons are of two types: one is thinly spun, translucent and pale brown while the other is tightly spun, opaque and dark brown.

Pest Management Chemical control of localized infestations or individual ornamental trees is considered feasible. Some attempts are underway to explore for potential natural enemies for classic biological control (Dapoto & Giganti 1994, Ede 2006).

Formicidae, Leaf Cutter Ants Leaf cutter ants of the genera Acromyrmex and Atta are social insects and have the highest social development among ants. They are the only animals other than humans and some termites (see Chapter 15) that cultivate their own food from fresh vegetation. They also have the ability to use sophisticated antibiotics against fungi in the subterranean gardens that they cultivate. They cut leaves from plants, including trees, and grow fungi on the cut fragments. The fungi provide food for the colony. Nests can contain up to 8 million ants. They are found in a variety of habitats ranging from arid, subtropical and tropical regions. All species of leaf cutter ants are found in the western hemisphere in portions of North, Central and South America and the Caribbean Islands. Several are forest pests (San Juan 2005).

Acromyrmex spp Leaf cutter ants of the genus Acromyrmex are distinguished from the closely related Atta by the presence of four pairs of spines and a rough exoskeleton on the upper surface of the thorax. About 23 species are known (H€ olldobler & Wilson 1990, Table 8.3).

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Table 8.3 Distribution of species of Acromyrmex (Hymenoptera: Formicidae) in the western hemisphere. Species

Distribution

Subgenus Acromyrmex A. ambiguus Argentina, Brazil A. aspersus Argentina, Brazil, Colombia, Peru A. coronatus Costa Rica south to Bolivia and Brazil A. crossispinus Argentina, Brazil, Paraguay A. diasi Brazil A. diseiger Brazil A. gallardoi Argentina A. hispidus Argentina, Bolivia, Paraguay A. lrystrix Brazil, Guyanas, Peru A. laticeps Bolivia, Brazil, Uruguay A. lobicornis Argentina, Bolivia, Paraguay A. lundi Argentina, Bolivia, Brazil A. niger Brazil A. octospinosus Mexico to northern South America, Caribbean A. rugosus Columbia south to Argentina A. subterraneus Argentina, Brazil, Peru Subgenus Moellerius A. heyeri Argentina, Brazil, Paraguay, Uruguay A. landolti Northern South America to Argentina A. mesopotamicus Argentina A. pulvercus Argentina A. silvestrii Argentina, Uruguay A. striatus Argentina, Bolivia, Brazil A. versicolor USA: Arizona, California, Texas Northern Mexico € lldobler & Wilson 1990. Source: Ho

Distribution Species of Acromyrmex are found from southwestern USA, south to Paraguay and northern Argentina with most species indigenous to South America (Table 8.3). A. versicolor (Pergande) occurs in southwestern USA. A. octospinosus Reich is native to Central and South America and the Caribbean Basin. A. ambiguus Emery occurs in southern Brazil, Paraguay and Uruguay, A. aspersus F. Smith and A. laticeps Forel are Brazilian species and A. lundii Guerin-Meneville occurs in southern Brazil and Paraguay.

Hosts Host plants include a wide range of grasses, trees shrubs and plants of agricultural importance. In San Jose, Costa Rica and Panama City, Panama,

A. octospinosus foragers are common on trees of the genus Cecropia in urban environments. A. octospinosus foragers typically cut the leaves of small herbaceous plants, fallen flowers and fallen fruit parts in both wet and dry forests of Costa Rica. The Brazilian species, A. laticeps nigrosetosus, forages on foliage of Eucalyptus camaldulensis.

Importance Worker ants remove large pieces of foliage from plants to serve as a medium for the fungi they cultivate. Members of this genus more or less continuously change food plants and thus avoid completely stripping off leaves and thereby damaging trees.

Life History Ants of the genus Acromyrmex cultivate fungi in subterranean chambers. Studies of the Brazilian subspecies A. laticeps nigrosetosus indicate that nests consist of a single chamber with average dimensions of 0.28  0.23  0.19 m (length  width  height). Chambers are linked to the soil surface by two to five channels. Externally, they are covered by dry leaves and in their early stages are indicated by a mound of loose soil. Nest size averages 12,231 individuals. Winged males and females fly from their parent colonies and mate.Before leaving parent colonies, females store a small section of the fungus from the chamber into their buccal pouches as seed for fungus gardens of incipient colonies. Mated females (queens) loose their wings and dig into the soil where they establish new colonies. Adult flights of the desert leaf cutter ant, A. versicolor, occur after late summer rains that wet the soil.

Description of Stages Adult foragers are redbrown to brown, have four pairs of spines and a rough exoskeleton. The spines help them maneuver and carry fragments of foliage and flowers on their backs (Wetterer 1991, 1998, Johnson & Rissing 1993, Araujo & Della Lucia 1997). Atta The genus Atta consists of about 15 species found from Louisiana and Texas, USA, south through Mexico, Central and South America (Table 8.4, H€ olldobler & Wilson 1990).

Other foliage feeding insects Table 8.4 Distribution of species of Atta (Hymenoptera: Formicidae) in the western hemisphere. Species

Distribution

A. bispherica A. capiguara A. cephalotes

Brazil Brazil, Paraguay Southern Mexico to Ecuador and Brazil, Caribbean Guatemala to Colombia Brazil Cuba Colombia and Guyanas, south to Paraguay Arizona, USA south to El Salvador Brazil Brazil Argentina, Bolivia, Paraguay Costa Rica south to Argentina and Paraguay Brazil USA: Lousiana, Texas Argentina, Bolivia, Brazil

A. A. A. A.

colombica goiania insularis laevigata

A. A. A. A. A.

mexicana opaciceps robusta saltensis sexdens

A. silvai A. texana A. vollenweider

€ lldobler & Wilson 1990. Source: Ho

Atta cephalotes (Linnaeus) Distribution This species is indigenous to southern Mexico, all of Central America, Trinidad and Tobago in the Caribbean and northern South America.

Hosts Tree hosts include Cedrela odorata, Citrus spp., Cupressus lusitanica, Gmelina arborea, Mangifera indicata, Pinus patula, Swietenia macrophylla and Tectona grandis.

Importance Colonies occur under a variety of conditions ranging from natural forests and jungles, forest plantations, orchards to urban areas. Under some conditions, they are simply part of a complex forest ecosystem but in others they can cause moderate to heavy defoliation and require control. In Colombia, this ant is a pest of exotic conifer plantations.

Life History A colony can exist for as long as 20 years. It begins with the arrival of a mated female, whose wings have been detached. She selects a site to develop a new colony, digs a small chamber and

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deposits her eggs. Within 1 month, eggs produce the first workers. Workers forage for suitable foliage and return to the colony with pieces of leaves that are deposited in subterranean fungus chambers to feed the colony. The colony gradually grows until it contains thousands of individuals. After 3 years, the first reproductives appear. They leave the nest to begin new colonies.

Description of Stages Winged reproductives are about 16 mm long, red-brown and have a spine on each occipital lobe of the head. They also have pairs of spines on the thorax and abdomen. Males are 14 mm long and lack spines on the head and thorax. Workers are differentiated into three castes. Worker soldiers have yellow bristles on the front, four pairs of spines on the thorax and two on the abdomen. Head and mandibles are large and ocelli (eyes) are present. Foraging workers are smaller and lack eyes. Their entire body is redbrown. The fungus cultivators never leave the nest. They are light colored and about 4 mm long. Larvae are white and legless.

Pest Management In instances where trees require protection from defoliation, colonies can be killed using insecticide-treated baits. Foliage of high-value plantation or urban trees can be sprayed to prevent defoliation (Cibri an Tovar et al. 1995, Rodas P. 1998). Atta texana (Buckley), Town Ant, Texas Leaf Cutting Ant Distribution Texas leaf cutting ant occurs in west central Louisiana, east Texas and Mexico, south to Vera Cruz.

Hosts This ant prefers to clip grasses, other herbaceous plants and foliage of broadleaf trees for its fungus gardens. Worker/foragers will feed on the foliage of pines, Pinus echinata and P. taeda, especially during the winter months when there is no other green foliage available.

Importance Pines can be severely defoliated, especially during winter. Defoliation on pines resembles that caused by pine sawflies of the genus Neodiprion.

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Fig. 8.10 Mounds constructed by Texas leaf cutting ant, Atta texana (east Texas, photo by R. F. Billings, Texas Forest Service).

Fig. 8.11 Worker forager of Texas leaf cutting ant, Atta texana (photo by R. Scott Cameron, International Paper Company, courtesy of forestryimages.org).

Other foliage feeding insects Life History Elaborate nests are built underground, usually in well-drained and loamy soils. Sandy soils are preferred for nesting sites. Nest interiors may be 6 m deep and contain more than 1000 entrance holes. Nesting areas are often marked by crescent-shaped mounds 12–35 cm high and 30 cm in diameter. Each mound surrounds an entrance hole (Fig. 8.10). They consist of cavities in which the fungus gardens are maintained and are connected by narrow tunnels. Above ground, there are sharply defined foraging trails that lead from nests to plants from which ants collect foliage. Trails may extend for several hundreds of meters. Ants move in processions along the trails and each carries a fragment of foliage back to the nest. Foliage fragments may be several times the size of the ants and are borne upright over the head like a parasol. Foliage bits are cut into small fragments and placed in the underground gardens to raise the fungus. Winged

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males and females appear in May and June, fly from the colony and mate. Mated females (queens) loose their wings and dig into the soil where they establish new colonies.

Description of Stages Reproductives and worker/ foragers are rusty red. The head is strongly bilobed and antennae are 11 segmented. Legs are long and the thorax bears three pairs of prominent dorsal spines. Reproductives are about 18 mm long and workers are from 1.5 to12 mm long (Fig. 8.11). Larvae are white and lack legs.

Pest Management Use of insecticide-treated baits to destroy colonies has met with some success (Moser 1967, Drooz 1985, Kulhavy et al. 1998).

Chapter 9

Bark and Ambrosia Beetles

INTRODUCTION Two of the most damaging groups of forest insects are bark beetles and ambrosia beetles. They comprise two subfamilies of the Curculionidae (weevils): the Scolytinae and Platyponinae. The Scolytinae, formerly family Scolytidae, consists of about 6000 species of small, cylindrical, beetles found worldwide (Wood S.L. 1982). When adults first emerge from their pupal cases, they are amber colored, and turn red-brown to black when mature. While many species confine attacks to recently dead material and are of minor importance, species of several genera (e.g. Dendroctonus, Ips, Scolytus) attack and kill live trees. Bark beetles breed in the cambium and inner bark of host trees. Pitch tubes, resin flow or fine red-brown boring dust on the bark surface are indicators of attack (Fig. 2.3, Plate 39). Most species have a symbiotic relationship with wood staining fungi (e.g. Leptographium, Ophiostoma), some of which are pathogenic. Beetles carry spores of these fungi on their bodies, often in a specialized tube-like structure known as a mycangium, and spread the fungi from tree to tree. Provided

that the beetles attack living trees, these fungi invade the tree’s vascular system, discolor the wood (Fig. 9.1) and hasten tree death. When bark beetles invade trees, they construct egg galleries and deposit eggs, usually in individual niches. With few exceptions, larvae feed in individual galleries that are more or less perpendicular to the egg galleries. The gallery patterns, produced by the attacking beetles and their brood, are often characteristic and easily recognized. While adults may be somewhat similar in appearance, gallery patterns coupled with tree species attacked is often sufficient for field identification of the beetle involved. Some bark beetles are known as engraver beetles because of the characteristic, often attractive galleries they construct. Galleries are referred to as the “signature” of the bark beetle involved. Many bark beetles feed before they mate or initiate attacks. Depending on the species, adult feeding may occur in the cambium layer, on branch tips and/or crotches of host trees. This kills portions of branches and makes ornamental and/or Christmas trees unsightly. Others either feed on the tender bark or bore into the pith of young seedlings and kill them (see Chapter 13).

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Fig. 9.1 Discoloration of wood of lodgepole pine, Pinus contorta, caused by blue stain fungi associated with mountain pine beetle, Dendroctonus ponderosae (Roosevelt National Forest, Colorado, USA).

Several bark beetles vector pathogenic fungi during adult feeding. Ambrosia beetles make breeding attacks in the wood of host plants. After mating, females construct a network of galleries and cradles in the wood. Entrance galleries are marked by piles of fine, granular, white boring dust in bark crevices or on the ground adjacent to infested trees (Plate 40). They deposit an egg in each cradle and inoculate wood with spores of ambrosia fungi that provide food for developing larvae. Unless ambrosia beetles are associated with fungi that are pathogenic, their activity does not kill trees. In addition, most temperate species confine attacks to weakened or recently dead trees. However, a number of tropical species are capable of attacking live trees. Ambrosia beetle infestations cause loss of wood quality but, in a few cases, the galleries and wood stain can be an attractive feature in wood carvings, paneling and furniture. One North American genus of Scolytinae, Conophthorus, attacks and destroys cones and seeds of pines (see Chapter 14).

Immature stages of the subfamily Scolytinae are similar in appearance and difficult, if not impossible, to separate. Eggs are tiny and pearly white in color. Larvae are legless C-shaped grubs with a white body color and an amber head capsule with darker mouthparts (Fig. 9.2). Pupae are white and have partially developed wings and legs. Beetles of the subfamily Platyponinae are elongate, slender and cylindrical, 2–8 mm long, brown in color and with a head slightly wider than the thorax. Immature stages are similar in appearance to the Scolytinae. All are ambrosia beetles and usually confine their attacks to weakened or dying trees. They are found principally in tropical and subtropical climates. The Platyponinae can be more damaging than ambrosia beetles of the subfamily Scolytinae because their galleries are more extensive and extend deeper into the sapwood and heartwood. Dying, weakened or recently felled trees are preferred but healthy trees can be attacked, especially if areas of dead bark are present (Drooz 1985, Triplehorn & Johnson 2005).

Bark and ambrosia beetles

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Fig. 9.2 Larval stage of mountain pine beetle, Dendroctonus ponderosae.

Curculionidae (Subfamily Scolytinae – Bark Beetles) Dendroctonus Nineteen species comprise this genus, 17 from North and Central America, one from China and one from Eurasia (Fig. 9.3 & Table 9.1). They generally breed in conifers greater than 15 cm in diameter. Collectively, bark beetles of the genus Dendroctonus are the most destructive biological agents of North and Central American conifer forests. Many species attack standing trees but several attack recently felled or windthrown trees and when populations build, subsequent generations attack and kill standing trees. During outbreaks, several species attack and kill healthy, vigorous trees. All species of Dendroctonus are monogamous (Wood S.L. 1982). Dendroctonus brevicomis LeConte, Western Pine Beetle Distribution Western pine beetle is indigenous to western North America. In the USA, it occurs in Arizona, California, southwestern Colorado, Idaho, western Montana, Nevada, New Mexico, Oregon,

western Texas, Washington and Utah. In Canada, it is found in southern British Columbia. In Mexico, it occurs in the states of Chihuahua, Coahuila, Durango, Nuevo Leon and Zacatecas.

Hosts Ponderosa pine, Pinus ponderosa, is the primary host in Canada and the USA. In California, Coulter pine, P. coulteri, is also attacked. In Mexico, P. arizonica, P. durangensis and P. estevezii are hosts in addition to ponderosa pine.

Importance Western pine beetle is a major pest of pine forests. During outbreaks, group killing of trees is common in dense, overstocked stands of pure, evenaged, young sawtimber sized trees and among dense clumps of pine in mixed conifer forests. During outbreaks, a million or more trees may be killed annually. Tree killing depletes timber supplies, affects stocking levels, disrupts forest management and increases fire danger.

Life History This species has several generations/ year. The number of generations is complicated by

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Fig. 9.3 Distribution of the two old-world species of Dendroctonus: D. armandi and D. micans (based on data from Critchfield & Little 1966, Bevan & King 1983, Wood & Bright 1992, Bright & Skidmore 2002, Kimoto & Duthie-Holt 2006).

climatic variability, re-emergence of parent adults to produce a second or third brood and by overlapping generations. In the northern parts of its range and at high elevations, one complete and a partial second generation can be expected. In southern California and Arizona there are three and sometimes four generations/ year. Adult flight and attack can occur as early as March and as late as November, with the number of adults peaking at different times with respect to elevation. Females initiate attacks and release minute amounts of pheromones, which attract males and other females, and cause a mass attack. During outbreaks, groups of trees are attacked. All life stages occur beneath or in the bark of infested trees, except for a brief period when adults fly to attack new trees. Adults become active when subcortical temperatures reach 7.2–10 C. During attacks, which may last 3 weeks, each female lays about 60 eggs individually in niches cut into the sides of winding S-shaped galleries (Fig. 9.4). Some parent females may emerge and re-attack elsewhere in the same tree or in neighboring trees. Eggs hatch after 1–2 weeks. Larvae feed first in the phloem, where they

construct a short gallery. They then mine into the inner bark, where most development takes place. After completing four instars, they transform into pupae and later adults. Brood adults feed in bark, and spores of blue stain fungi introduced by attacking adults collect in their mycangia to be inoculated into trees they attack. In any given location, elevation is a determining factor where western pine beetle attacks are most abundant. In the USA, attacks occur in forests between elevations of 600 and 2230 m. In north central Arizona, attacks tend to be most abundant at elevations between 1600 and 2230 m. In Canada, infestations usually occur below 300 m and in Mexico above 2400 m.

Description of Stages Adults range from 2.0 to 4.7 mm in length and are about 2.2 times as long as they are wide.

Pest Management Silvicultural tactics, designed to maintain tree and stand vigor, are most appropriate for

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Table 9.1 Distribution and hosts of bark beetles of the genus Dendroctonus (Coleoptera: Curculionidae: Scolytinae). Species

Distribution

Hosts

D. adjunctus Blandford Round headed pine beetle D. approximatus Dietz Larger Mexican pine beetle D. armandi Tsai & Li D. brevicomis LeConte Western pine beetle D. frontalis Zimmerman Southern pine beetle D. jeffreyi Hopkins Jeffrey pine beetle D. mexicanus Hopkins Smaller Mexican pine beetle D. micans Kugelann D murrayanae Hopkins

Guatemala, Mexico, southwestern USA

Pinus spp.

Guatemala, Honduras, Mexico, southwestern USA

Pinus spp.

Central China Southern Canada, northern Mexico, western USA

Pinus armandii Pinus coulteri, P, ponderosa

Belize, Guatemala, Honduras, Mexico, Nicaragua, southern USA Mexico: Baja California USA: California, western Nevada Guatemala, Honduras, Mexico, USA (southeast Arizona) Asia, Europe (including the British Isles) Central and western Canada and the USA

Pinus spp. P. jeffreyi Pinus spp. Picea spp. Pinus banksiana, P. contorta, P. strobus Pinus leiophylla, P. oocarpa Pinus spp.

D. parallelocollis Chapuis D. ponderosae Hopkins Mountain pine beetle D. punctatus LeConte

Guatemala, Honduras, Mexico Southern Canada, western USA

D. pseudotsugae Hopkins Douglas-fir beetle D. rhizophagus Thomas & Bright

Western Canada, northern Mexico, western USA Mexico: Chihuaha, Durango

D. rufipennis (Kirby) Spruce beetle D. simplex LeConte Larch beetle D. terebrans (Olivier) Black turpentine beetle D. valens LeConte Red turpentine Beetle D. vitei Wood

Canada, northeastern and western USA

Picea glauca, P. rubens, P. sitchensis Pseudotsuga menziesii, P. macrocarpa Pinus durangensis, P. engelmannii Picea spp.

Canada, northern USA

Larix laricina

Eastern and southeastern USA

Pinus spp.

Canada, China (introduced), Guatemala, Mexico, northern and western USA Guatemala

Pinus spp.

Canada, USA

Pinus pseudostrobus, P. tenuifolia

Sources: Wood S.L. 1982, Wood & Bright 1992, Bright & Skidmore 2002, Furniss & Johnson 2002, Kimoto & Duthie-Holt 2006.  Important tree killing insect in its natural range.  Important tree killing pest in introduced range.

long-term management. These include sanitation cutting where mature trees with dead tops and branches, short, sparse and chlorotic foliage and dwarf mistletoe, Arceuthobium spp., infections are harvested. Other tactics include thinning and avoidance of mechanical injury to residual trees during timber harvesting. High-value trees near homes or in developed recreation sites can be sprayed in spring–early summer to prevent beetle attacks (DeMars & Roettgering 1982, Wood S.L.

1982, Cibri an Tovar et al. 1995, Fettig et al. 2004, Williams et al. 2008). Dendroctonus frontalis Zimmerman, Southern Pine Beetle Distribution Southern pine beetle is indigenous to southern USA from Ohio, Pennsylvania and West Virginia south to Florida, west to east Texas and portions of

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Fig. 9.4 Characteristic winding S-shaped galleries of western pine beetle, Dendroctonus brevicomis.

Arizona and New Mexico. It also occurs throughout Mexico and Central America, as far south as Nicaragua (Fig. 9.5). Hosts This bark beetle can attack and kill all species of Pinus within its natural range. In southeastern USA, P. echinata, P. rigida, P. serotina, P. taeda and P. virginiana are the principal hosts. In Arizona and New Mexico, P. engelmanni, P. leiophylla var. chihuahua and P. ponderosa are attacked. Hosts in Mexico and Central America include P. ayacahuite, P. arizonica, P. caribaea var. hondurensis, P. durangensis, P. maximinoi, P. oocarpa, P. pringlei, P. tecumani and P. teocote. Importance Southern pine beetle is one of the most destructive bark beetle pests of pine forests in North and Central America. Trees are killed in groups ranging from five to several thousand trees (Plate 41). In the USA, outbreaks have occurred in portions of Alabama, Georgia, Kentucky, Louisiana, Mississippi, North and South Carolina, Tennessee, east Texas, Virginia and West Virginia. Honduras has a history of outbreaks

beginning in the early to mid-1960s. From 1962 to 1965, more than 2 million ha of pine forests were affected. Another outbreak occurred in 1982 in naturally regenerated stands that developed after the 1960s outbreak. A regional outbreak in Central America, including Belize, Guatemala, Honduras and northern Nicaragua, occurred from 2000 to 2003. Infestations were almost exclusively in young, dense, pine forests ranging from 18 to 25 years in age with basal areas exceeding 35 m2/ha. These forests were stressed from overcrowding, recent fires and drought.

Life History Southern pine beetle is a multiple generation species and the number of generations varies with location and climate. In the northern part of its range, for example Virginia or the southern Appalachian Mountains, three generations are typical. In the Piedmont region of Alabama, the Carolinas and Georgia, four to five generations may occur. Further south, there may be six to seven generations. All life stages overwinter in the cambium or inner bark. Adults emerge in spring, when temperatures rise. They mass

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Fig. 9.5 Distribution of the southern pine beetle in southern USA, Mexico and Central America (redrawn from Clark & Nowak 2009).

attack susceptible trees and both sexes are attracted to pheromones produced by attacking females. Once they gain entrance to the cambium, mating occurs and females construct winding S-shaped egg galleries and deposit eggs singly on either side of the gallery. Galleries can be differentiated from those caused by western pine beetle, D. brevicomis, because pupal cells are visible in the inner bark whereas in the case of western pine beetle, they are hidden in the inner bark (Fig. 9.6). A blue stain fungus, Ophiostoma minus (¼ Ceratocystis minor), and an unidentified basiodiomycete are associated with attacking beetles. Larvae hatch within a few days and begin feeding in the cambium. When larvae mature, they construct cells in the inner bark, pupate and develop into adults. Description of Stages Adults are short-legged, stout beetles, range from 2.0 to 3.2 mm long and are about 2.3 times as long as they are wide. The front of the head has a distinct notch and the hind end is smooth and rounded. Mature adults are dark brown to black and newly emerged “callow” adults are soft bodied and amber colored but harden and darken quickly.

Pest Management Prevention includes thinning to reduce stand density, removal of damaged and weakened trees, and harvesting trees before they reach maturity. Direct control tactics involve removal of infested trees via commercial timber sales, rapid processing of logs and destruction of bark, cut and leave, or piling and burning of infested trees. Cut and leave consists of felling all trees with fresh attacks or brood plus a buffer strip of adjacent uninfested trees. This reduces beetle survival within infested trees, disrupts pheromone production and prevents infestation spread (Moser 1975, Thatcher et al. 1980, Thatcher & Barry 1982, Wood S.L. 1982, Paine & Stephen 1987, Cibri an Tovar et al. 1995, Billings et al. 2004).

Dendroctonus micans (Kugelann), Great Spruce Bark Beetle Distribution The origin of D. micans is believed to be Asian boreal conifer forests. Over the past century, it has extended its range into Europe. This has been at least partially aided by increased trade in timber products, especially unprocessed logs. It was first detected

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Fig. 9.6 Galleries of southern pine beetle, Dendroctonus frontalis. Note that pupal cells are visible in the inner bark.

in the UK in 1982 and is believed to have been introduced 10 years previously (see Fig. 9.3).

Hosts Primary hosts are species of Picea including P. abies, P. asperata, P. jezoensis, P. obovata, P. omorika and P. orientalis. It also attacks several North American species of Picea introduced into Europe, including P. breweriana, P. engelmannii, P glauca, P. mariana, P. pungens and P. sitchensis. Attacks also occur on species of Abies, Larix and Pinus.

Importance D. micans is different from the more aggressive Dendroctonus species in that it usually attacks its hosts in low numbers and kills bark in patches. Successive attacks over 5–8 years may be necessary to kill a tree, except during outbreaks. Within most of its natural range, D. micans occurs at low levels and causes little tree mortality. However, outbreaks do occasionally occur. For example, as D. micans extended its range westward into Europe (France and the UK) and southwestern Asia (Republic of Georgia and Turkey) during the late 1900s, outbreaks occurred on more than 200,000 ha of spruce forests. In some cases, older trees were preferentially attacked, while in other

instances all age classes of trees were attacked. D. micans normally colonizes only green standing trees and attacks trees stressed by logging damage, frost, snow, wind, lightning, poor soil nutrition and drought.

Life History The time required to complete a generation varies depending on local conditions and ranges from 10 to 18 months in the UK, from12 to15 months in Turkey and Russia and from 2 to 3 years in Nordic countries. New adults mate under the bark before they emerge. Mating often occurs among siblings. Sex ratios are highly female biased. The typical sex ration is 1 : 10 males : females but can be as high as 1 : 45. After mating, some females remain beneath the bark and simply initiate new galleries nearby. Others emerge and attack elsewhere on the same tree, while others fly to new host trees. Adult flight occurs throughout summer. Mated females construct individual egg galleries in living trees from April to November, depending on local conditions. Egg galleries are constructed primarily in the inner bark (phloem) and females lay from 100 to 150 eggs in a cluster. Larvae feed in a communal, fanlike gallery and pack frass behind themselves. Under laboratory conditions, larvae complete development in about 2 months and pupation in about 1 week. Larvae

Bark and ambrosia beetles and adults overwinter. In spring, brood adults feed and mate under bark for 6–7 weeks. D. micans does not appear to have an aggregation pheromone.

Description of Stages Adults are 6–9 mm long, dark-brown and cylindrical. Legs and antennae are yellow-brown. Morphologically, D. micans is difficult to distinguish from the North American species D. punctatus and they have been suspected of being conspecific. However, it has been established that they are distinct species.

Pest Management Direct control includes sanitation felling and insecticide applications to infested portions of trees. Classic biological control using the predaceous beetle Rhizophagus grandis, which is specific to D. micans, has been used in France, Republic of Georgia and the UK (Bevan & King 1983, Evans et al. 1984, Gregorie 1988, Furniss 1996, Fielding & Evans 1997, Kegley et al. 1997).

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Life History One generation/year is typical but 2 years may be required at high elevations. In portions of California, there are two generations/year. Adults emerge from trees attacked the previous year in late May–early June and may fly until September. Females initiate attack, produce an attractant pheromone and are joined by males. After mating, females construct a vertical egg gallery 10–122 cm long, packed with boring dust (Fig. 9.7) and introduce blue stain fungi into the tree. Eggs are laid singly in niches along both sides of the gallery. They hatch in 10–14 days and larvae feed in the phloem in galleries at right angles to the egg galleries. When mature, larvae construct oval pupation cells. By late summer, most of the current year’s brood have become adults that overwinter under the bark.

Dendroctonus ponderosae Hopkins, Mountain Pine Beetle Distribution Mountain pine beetle occurs in western North America from Alberta and northern British Columbia, Canada south through western USA and northern Mexico.

Hosts All pines, Pinus spp., within its range are attacked. Primary hosts are P. albicaulis, P. contorta, P. flexilis, P. lambertiana, P. monticola and P. ponderosa. Scotch pine, P. sylvestris, and several other pines exotic to western North America are also attacked.

Importance Mountain pine beetle is the most destructive insect pest of pine forests in western North America. Outbreaks have occurred every year since records were kept. Factors that lead to outbreaks vary with the host. Even-aged forests of P. contorta are most susceptible when they reach age 60 years and contain large numbers of trees over 20 cm in diameter with a thick phloem. P. ponderosa forests that are overstocked, have a high basal area and reduced increment are most susceptible to outbreaks.

Fig. 9.7 Egg and larval galleries of mountain pine beetle, Dendroctonus ponderosae.

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Fig. 9.8 Adult mountain pine beetle, Dendroctonus ponderosae.

Description of Stages Adults range from 3.5 to 6.6 mm long. The body is 2.2 times as long as wide. Body color of mature adults is black (Fig. 9.8).

Pest Management Cultural, mechanical and chemical tactics are available to manage mountain pine beetle. However, during outbreaks these tactics have only limited value. Cultural controls vary depending on host tree. Pinus contorta, which typically occurs as even-aged fire originated forests, can be managed by creating a mosaic of age classes over the landscape via small clearcuts, which are easily regenerated by natural seeding. A mosaic of stands of different age classes will have some stands susceptible to outbreaks and others that are too small in diameter to support beetle broods. Susceptibility of P. ponderosa forests to beetle attack can be reduced by thinning. Cutting and burning of infested trees during winter and wrapping infested logs in plastic to create high temperatures are effective against localized infestations. Several chemicals are available to

spray boles of individual, high-value trees near homesites or developed recreation areas to prevent attack. Sprays should be applied in mid- to late May and trees should be treated to a top diameter of about 12 cm (Sartwell & Dolph 1976, Wood S.L. 1982, Amman et al. 1990, Furniss & Johnson 2002). Dendroctonus pseudotsugae, Hopkins, Douglas-fir Beetle Distribution Douglas-fir beetle is found throughout western North America from Alberta and British Columbia, Canada, south into Chihuahua and Durango states of northern Mexico. Hosts The primary host is Douglas-fir, Pseudotsuga menziesii. Big cone Douglas-fir, P. macrocarpa, is a host in southern California and P. flahaulti is attacked in northern Mexico. It also attacks western larch, Larix occidentalis, and can produce broods in windthrow or freshly cut logs but not in standing trees.

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Importance Douglas-fir beetle is the most damaging bark beetle pest of mature Douglas-fir forests and outbreaks have killed millions of cubic meters of Douglas-fir timber. This insect can build up in windthrown material following severe storms, reach epidemic levels and then attack and kill standing trees. It also can build up in standing trees during droughts or following defoliator outbreaks. Life History There is one generation/year. Brood adults and some larvae overwinter. Adults emerge, fly and attack new material from April to early June, depending on local conditions. Those individuals that overwintered as larvae emerge later and adults that emerged early may make a second attack in late June–early July. After mating, females construct a single egg gallery parallel to the grain. Gallery length ranges from 20 to 25 cm. They are packed with frass and may be somewhat longer in windthrown trees. Eggs are deposited alternately along opposite sides of the gallery and hatch in 1–3 weeks. Newly hatched larvae mine in galleries more or less perpendicular to the egg gallery. When feeding is completed, larvae construct a pupal cell at the end of their gallery and pupate (Fig. 9.9). Trees may be infested at varying lengths but usually not higher than a top diameter of 15–20 cm. Description of Stages Adults are stout, cylindrical beetles 4–6 mm long. The head and thorax are black and the elytra are red-brown but may darken with age. Pest Management The pheromone complex has been identified and has been used with some success to manipulate populations. Frontalin and seudenol are attractants and, in combination with volatile components of Douglas-fir resin, can be used to concentrate low populations. Methylcyclohexanone (MCH) disrupts attraction and has been deployed in areas of fresh windthrow to prevent or reduce attacks. Management of Douglas-fir stands, including rapid removal and processing of windthrow, timely harvesting of mature trees and thinning to maintain tree vigor and reduce moisture stress is the most effective long-term tactic for managing this insect (Furniss & Orr 1978, Wood S.L. 1982, Cibrian Tovar et al. 1995, Furniss & Johnson 2002).

Fig. 9.9 Egg and larval galleries of Douglas-fir beetle, Dendroctonus pseudotsugae.

Dendroctonus rufipennis (Kirby), Spruce Beetle (Plate 42) Distribution Spruce beetle is found across the boreal forests of Canada, west to Alaska, and south into the northeastern and Rocky Mountain regions of the USA.

Hosts All species of Picea are hosts. In northeastern USA and adjoining Canada, outbreaks have occurred in red spruce, P. rubens. In the Rocky Mountain region, high-elevation forests of Engelmann spruce, P. engelmannii, are subject to attack and in Alaska, outbreaks have occurred in forests of white spruce, Picea glauca.

Importance This insect is considered the most damaging pest of mature spruce forests in North America.

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Outbreaks have occurred in Arizona, Colorado, Idaho, Maine, Montana, New York, Utah and Wyoming, USA and British Columbia, Canada. In Alaska, an outbreak between 1979 and 1999 spread over 400,000 ha and killed an estimated 30 million trees/year during its peak. In the 1990s, outbreaks in Utah infested over 50,000 ha and killed more than 3 million trees. Lowlevel populations typically exist in fresh windthrow and outbreaks in standing trees are often the result of population increases following high wind events.

Life History Spruce beetle may complete a generation in 1 year on warm sites at low elevations or take up to 3 years in cool locations. A 2-year life cycle is most common. Adults are active from May to October but most attacks occur in early summer. Some brood adults may re-emerge and attack additional trees later in summer. Females initiate attacks and bore into bark and phloem. After mating, they construct egg galleries that range from 6 to 13 cm long. Eggs are laid along alternate sides of the gallery in rows of 4–14 eggs/cm of gallery. Most hatch by August. Larvae bore outward from the egg gallery and feed communally for the first two instars. Third and fourth instars construct individual feeding galleries. Larvae predominate during the first winter, although parent adults and eggs may also be present. During a 2-year life cycle, most larvae pupate 1 year after attack. Pupation occurs in cells at the end of larval galleries and lasts 10–15 days. During the second winter of a 2-year cycle, some adults in standing trees overwinter in their pupal cells, but most emerge, move to the base of the tree and bore into the bark near the root collar. This reduces predation by woodpeckers and winter mortality from cold temperatures. In windthrown trees, most adults overwinter in pupal cells.

Description of Stages Adults are dark brown to black with reddish-brown or black wing covers, approximately 6 mm long and 3 mm wide.

Pest Management Tactics for spruce beetle management include: (i) harvesting of infested and susceptible trees to encourage regeneration of the young; (ii) vigorous forest, salvage logging of windthrown spruce to prevent beetle attack; (iii) use of trap logs to absorb attacking beetles and prevent attacks in standing

trees; (iv) exposure of infested logging residues to direct solar radiation to kill larvae; (v) use of aggregating and anti-aggregating pheromones; and (vi) preventative spraying of high-value trees with chemical insecticides. Most spruce forests in western North America occur at high elevations or are far from roads. Moreover, many of these forests are in designated Wilderness Areas on public lands. Therefore, pest management tactics cannot be applied, resulting in widespread tree mortality (Wood S.L. 1982, Weiss et al. 1985, Holsten et al. 1999, Furniss & Johnson 2002, Ciesla & Mason 2005). Dendroctonus valens LeConte, Red Turpentine Beetle Distribution Red turpentine beetle is indigenous to North America and occurs across Canada, northern and western USA, Mexico, Guatemala and Honduras. This insect was introduced into China, probably during the mid-1990s, via wood packaging material. An outbreak developed in Shanxi Province in 1999 and spread to Hebei, Henan and Shaanxi Provinces. Hosts In North America, hosts include Abies concolor and species of Picea and Pinus. In China, hosts are Pinus tabulaeformis and, occasionally, P. armandii. Importance Within its native range, this bark beetle is considered a secondary invader, which attacks trees with mechanical injury due to lighting or logging or attacked by more aggressive bark beetles. It also attacks and breeds in freshly cut stumps. In China, it has become a destructive forest pest and has killed more than 6 million pines. Life History Depending on location, red turpentine beetle may undergo one or two overlapping generations/ year, except in northernmost locations where more than 1 year may be required. In the northern parts of its range, adults are active from May to October and further south activity may occur throughout the year. Attacks are confined to the lower portion of the bole, usually less than 2 m. On vigorous trees, attacks are indicated by large reddish-yellow pitch tubes. On recently dead trees or stumps, attacks are indicated by the presence of granular frass mixed with dry resin. After successful entry into the cambium and mating, females construct

Bark and ambrosia beetles a vertical egg gallery and deposit clusters of eggs on one side of the gallery. Larvae feed in a communal gallery in the phloem and cambium and produce a large cavity filled with frass. They are active for a minimum of 2 months. Pupation occurs in cells formed in the frass or in short tunnels adjacent to the cavity.

Description of Stages Red turpentine beetle is the largest member of the genus Dendroctonus. Adults are 5.3–8.3 mm long, about 2.1 times as long as they are wide and red-brown in color.

Pest Management Within its natural range in North America, this bark beetle is of little consequence and pest management is not needed. Efforts to manage red turpentine beetle in China include restricting movement of infested pines, population monitoring via trapping and classic biological control.

Related Species Black turpentine beetle, D. terebrans (Olivier) and D. rhizophagus Thomas & Bright, are closely related species. Black turpentine beetle occurs in southeastern USA where it attacks southern yellow pines. Adults are black and life history and habits are similar to red turpentine beetle. D. rhizophagus is found in Chihuahua and Durango, Mexico, where it attacks and breeds in roots of pine seedlings. Adults are similar to red turpentine beetle in overall appearance (Wood S. L. 1982, Cibrian Tovar et al. 1995, Furniss & Johnson 2002, Liu et al. 2006). Phloeosinus Phloeosinus is a genus of monogamous bark beetles that infest conifers of the family Cupressaceae. About 62 species are known worldwide and are found distributed throughout the northern hemisphere. Twenty-seven species and two subspecies are known from North and Central America and 35 species are known from Africa, Asia, Australia and Europe. Several species are occasional pests (Wood S.L. 1982). Phloeosinus armatus Reitter (Plates 43 & 44) Distribution P. armatus is native to the Mediterranean portions of Europe, the Near East and northern Africa. It was detected in California, USA in 1989.

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Hosts Hosts include species of Cupressus, Juniperus and Thuja. In the Mediterranean region, the primary host is Cupressus sempervirens.

Importance P. armatus is a secondary pest and attacks and breeds in stems and large branches of trees suffering from drought, fire, root damage, infection by the canker causing fungus Seiridium cardinalis or plantations that have been established on poor soils. It also damages shoots of host trees during adult feeding, which makes ornamental trees unattractive and reduces tree height and diameter growth. It is a vector of S. cardinalis during adult feeding.

Life History Studies in Israel indicate that this species can have three to four generations/year. Early studies suggested that this insect, as well as the related species, P. bicolor, was bigamous. This is based largely on the presence of a pair of egg galleries radiating from a nuptial chamber. More detailed studies indicate that both P. armatus and P. bicolor are monogamous.

Description of Stages Adults are relatively large beetles, 4–4.5 mm long with a dark red-brown body color, black head and light red-brown legs (Grüne 1979, Bright & Skidmore 2002, Baruch et al. 2005, Haack 2006).

Phloeosinus bicolor (Brulle) (¼ P. aubei Perris) Distribution P. bicolor has a wide distribution and is found from central and southern Europe east to China and south into northern and eastern Africa and the Near East. The author has collected this bark beetle in central Kenya.

Hosts This species feeds on and breeds in species of Chamaecyparis, Cupressus, Juniperus, Platycladus orientalis, Sabina chinensis and Thuja spp.

Importance Attacks are usually confined to broken branches or fresh cut logs. However, it can attack live trees. For example, during the early 1990s it damaged trees in portions of Hungary following several

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consecutive years of drought. A report from Tunisia indicates that it can be a pest of forest and windbreak plantations of Cupressus sempervirens. It is also a vector of the fungus, Seiridium cardinale, which is spread during adult feeding. Life History In central Europe, there are two generations/year and in Tunisia, there are two and a partial third generation/year. In Tunisia, peak adult flight is from March to mid-April with two lesser peaks in May–June and mid-August–mid-October. Adults of the first generation emerge in April and the second generation in June. They feed on shoots of host trees prior to mating. Adults are probably monogamous, despite reports that suggest they are polygamous. Egg galleries are parallel to the wood grain, 4–8 cm long and sometimes forked with two distinct egg galleries. Females may re-emerge to attack additional host material. Description of Stages Adults are 1.4–2.5 mm long, brown in color (Grüne 1979, Mendel 1984, Bright &

Skidmore 2002, Baruch et al. 2005, Belhabib et al. 2007). Scolytus The genus Scolytus consists of about 100 species, which are found across the northern hemisphere and in South America. Individual species may invade either broadleaf trees or conifers. All temperate species are monogamous but several tropical species are bigamous (Wood S.L. 1982). Several breed in the cambium of living trees and at least two Eurasian species are vectors of Dutch elm disease, caused by the fungi Ophiostoma ulmi and O. novo-ulmi (Table 9.2). Scolytus multistriatus (Marsham), Smaller European Elm Bark Beetle Distribution Smaller European elm bark beetle is native to Europe and northern Asia. It was introduced into North America during the early 1900s, probably via elm veneer logs imported from Europe. The insect is

Table 9.2 Representative species of Scolytus (Coleoptera: Curculionidae: Scolytinae): their distribution and hosts. Species

Distribution

Hosts

S. intricatus (Ratzburg)

Europe, northern Africa, northern Asia, Near East British Columbia, Canada northwestern USA Belarus, northern China, Mongolia, Russia

Quercus dalechampii, Q. petraea, Q. robur, other broadleaf trees Larix lyallii, L. occidentalis

S. laricis Blackman Western larch beetle S. morawitzi (Semenov) S. multistriatus (Marsham) Smaller European elm bark beetle S. mundus Wood S. ratzeburgi Janson S. rugulosus (Mu¨ller) Shothole borer S. scolytus (Fabricius) Large elm bark beetle S. schevyrewi Semenov Banded elm bark beetle S. unispinosus LeConte Douglas-fir engraver S. ventralis LeConte Fir engraver

Europe and northern Asia (indigenous). North America, Canada, Mexico, USA, South America: Argentina, Chile (introduced) Mexico Europe, Japan, Mongolia, central and eastern Russia Asia, Europe, northern Africa (indigenous). Australia, North and South America (introduced) Europe

Larix gmelinii, L. kamtschatica, L. sibirica, L. sukaczerii Ulmus spp.

Abies religiosa Betula, Ulmus Malus spp., Prunus spp., Pyrus spp. Ulmus spp.

Asia: China, Korea, Mongolia, Central Asia (indigenous) North America (introduced) Western North America

Prunus, Salix, Ulmus and other broadleaf trees

Western North America

Abies concolor, A. grandis, A. magnifica

Pseudotsuga menziesii

n Tovar et al. 1995, Furniss & Johnson 2002, Kimoto & Duthie-Holt 2006, Wood 2007. Sources: Wood S.L. 1982, Bright & Wood 1992, Cibria

Bark and ambrosia beetles now distributed throughout North America, from British Columbia to Nova Scotia, Canada and from northeastern USA south to Florida and west to California. Infestations have also been reported from Argentina, Chile and Mexico. Hosts All species of Ulmus and Zelkova serrata, a tree native to Japan, are hosts. Importance This insect is a vector of the fungi Ophiostoma ulmi and O. novo ulmi, which cause Dutch elm disease. This disease, which is native to Asia, has caused devastating losses to elms across much of Europe and North America.

Life History This bark beetle can complete two to three generations/year and larvae overwinter in pupal chambers in the bark. Pupation occurs with the onset of warm spring weather and adults emerge in late March– early June, about the time elm foliage is fully expanded. Adults fly directly to weakened or dying elms to breed in the inner bark or to healthy elms, where they feed in branch crotches. Adults, carrying spores of the Dutch elm disease fungus, spread the disease when they feed. Breeding attacks occur in trees weakened by drought, disease or mechanical injury. Elm logs or firewood are also attacked. Females initiate attacks and release an aggregating pheromone that attracts both males and females. After mating, females construct an egg gallery parallel to the wood grain and eggs are deposited individually in niches on either side of the gallery. Larvae feed in individual galleries, generally perpendicular to egg galleries. When fully grown, larvae construct cells in the bark and pupate. During spring and summer, a generation may be completed in 30–40 days.

Description of Stages Adults are 1.9–3.1 mm long and distinctly two-toned with red-brown elytra and a dark brown-black body. Males have a bright yellow brush of hairs on the front of the head. The underside of the posterior end of the body is concave, with a stout spine emerging from the margin of the second abdominal sternite.

Pest Management Overall strategy for managing smaller elm bark beetle and Dutch elm disease is to limit

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supply of breeding material. Elm firewood, broken limbs, stressed trees and trees infected by Dutch elm disease are eliminated by chipping or burning. Severely stressed or dying trees can be injected with herbicides that cause bark to dry and render them unsuitable for breeding. Beetles can also be mass-trapped on sticky traps baited with aggregating pheromones (Wood S.L. 1982, 2007, Drooz 1985). Scolytus schevyrewi Semenov, Banded Elm Bark Beetle Distribution Scolytus schevyrewi is indigenous to China (Heilongjiang, Hebei, Henan, Shaanxi, Ningxia and Xinjiang Provinces), Kazakhstan, Korea, southern Kyrgyzstan, Mongolia, Russia, Tajikistan, Turkmenistan and Uzbekistan. It was detected in Colorado and Utah, USA, in 2003 and is now found throughout much of central and western North America. Museum collections suggest that it had been present in some areas of the USA for at least 10 years.

Hosts In Asia, primary hosts are species of Ulmus, including U. carpinifolia, U. davidiana var. japonica, U. laevis, U. macrocarpa, U. propinqua and U. pumila. Other reported Asian hosts include Caragana korshinskii, Elaeagnus spp., Malus pumila, Prunus armeniaca var. ansu, P. padus, P. persica, P. pseudocerasus, P. salicina, P. yedoensis and Salix spp., including S. babylonica. In the USA, U. pumila and at least two indigenous elms, U. americana and U. thomasi, have been attacked.

Importance S. schevyrewi usually attacks weakened or stressed trees, although it can attack vigorous trees. Young trees tend to be more resistant to attack. Occasional outbreaks can occur that result in widespread tree mortality. In the Karamay region of Xinjiang Province, China, S. schevyrewi is a major pest of elm and has damaged trees in both urban and rural settings. In the USA, elms have been killed in urban settings. Its role as a vector of Dutch elm disease is still undetermined but its life history suggests that it is a potential vector.

Life History In China, S. schevyrewi completes two to three overlapping generations a year, depending on location. Overwintering occurs as mature larvae inside

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pupal chambers or as adults under the bark. Adults emerge in late April–early May. Larvae of the first generation develop from May to June, and become adults by early July. By late August, most secondgeneration larvae construct pupal chambers and overwinter. However, some may continue development and complete a third generation if temperatures are favorable. New adults remain in their pupal chambers for 2–5 days before boring an exit hole through the bark. Sex ratio is slightly female biased (females : males ¼ 1.0 : 0.9). New adults walk along the bark surface after emergence before initiating flight. Adults are most active during warm, sunny weather and feeding occurs on bark at the crotches of tender twigs. Following adult feeding, females attack host trees by constructing individual entrance holes through the bark. Mating occurs on the bark surface and both males and females can mate several times. Each female constructs a single egg gallery in the cambium, parallel to the grain. Egg niches are closely arranged on each side of the gallery and

sealed with a mixture of sawdust and adhesive secretions. Egg galleries usually contain about 60 eggs (range ¼ 23–123 eggs) (Fig. 9.10). Eggs hatch in the order in which they were laid and larvae construct individual galleries. Initially, they are more or less perpendicular to the egg gallery but later they turn upward or downward. Some larval galleries meander or cross each other. Larvae have five instars. When feeding is completed, mature larvae construct pupal chambers in the outer bark at the end of their galleries.

Description of Stages Adults range from 3.2 to 4.2 mm long. Body color is red-brown with a black head. The frons is slightly protruding with striations running toward the clypeus in females and with yellow, inwardly curved frontal hairs on the peripheral edges in males. The elytra are red-brown to black-brown and a dark transverse band may occur on the elytra of some adults.

Fig. 9.10 Egg and larval galleries of banded elm bark beetle, Scolytus schevyrewi (photo by J. Negrón, USDA Forest Service).

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Pest management To date, pest management has been rapid removal and destruction of infested trees (Houping Liu & Haack 2004, Negrón et al. 2005). Scolytus ventralis LeConte, fir Engraver Distribution Fir engraver, Scolytus ventralis, is found throughout much of western North America, from southern British Columbia, Canada, south to Baja California Norte, Mexico and east to western Montana, Colorado and New Mexico, USA. Hosts Primary hosts are firs: Abies concolor, A. grandis and A. magnifica. Trees occasionally attacked include A. lasiocarpa, Pseudotsuga menziesii and Tsuga mertensiana.

Importance Fir engraver is the most important bark beetle of Abies in western North America. Attacks can cause top kill, branch dieback or death of the entire tree. Outbreaks cause extensive tree mortality, for example from 1977 to 1978, an outbreak killed about 1.2 million trees in northern California. Outbreaks often develop following below normal moisture or insect defoliation. Fir engraver is often associated with round headed fir borer, Tetropium abietis, or flat headed fir borer, Melanophila drummondii. During outbreaks, however, it is responsible for most of the tree mortality. Life History Adults emerge in summer and fly in search of host material, i.e. either stressed trees, freshly cut logs or windthrow. They can fly from June to September but most are active between July and August. Females enter the tree first, construct a nuptial chamber and are followed by a male. After mating, females construct a transverse egg gallery, perpendicular to the main stem, about 10–30 cm long, and lay between 100 and 300 eggs singly in niches on either side of the gallery (Fig. 9.11). Within several days, a yellow-brown discoloration due to the fungus Trichosporum symbioticum appears. Eggs hatch within 9–14 days and larvae construct narrow feeding galleries perpendicular to the egg gallery. Larvae require 40–380 days to complete feeding. Pupation occurs in cells at the end of the larval galleries and lasts from 7 to 14 days. Adults remain under the bark for an additional 2 weeks, then emerge.

Fig. 9.11 Egg and larval galleries of fir engraver beetle, Scolytus ventralis.

Description of Stages Adults are shiny dark brownblack beetles, about 4 mm long. When viewed from the side, they have an incurved posterior with a small central bump that is more pronounced in males.

Pest Management Losses can be reduced via removal and utilization of infested trees, removal of fresh cut logs and/or windthrow before brood emergence, and removal of diseased or stressed trees from forests.

Related Species S. mundus Wood occurs in central Mexico where it attacks Abies religiosa. Gallery patterns are similar to those of S. ventralis (Ferrell 1996, Cibri an Tovar et al. 1995, Furniss & Johnson 2002).

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Ips Ips consists of about 60 species indigenous to boreal and temperate conifer forests of Eurasia and North America. Twenty-five species are indigenous to north and central America, were they attack Picea and Pinus. The others are indigenous to Eurasia. Several are important forest pests, second only to Dendroctonus (Table 9.3). Life history and habits of species are similar. Most have multiple generations/year. Adults are polygamous. Males initiate attacks, construct a nuptial chamber in the cambium layer and attract two to seven females. After mating, females construct longitudinal egg

galleries and deposits eggs in niches along each side of the gallery. Multiple egg galleries radiating from a common nuptial chamber often give egg galleries a Y- or X-shaped pattern (Fig. 9.12). Young larvae feed individually perpendicular to egg galleries. Larval galleries increase in width as larvae grow. Pupation occurs in round chambers constructed at ends of larval galleries. Adults are characteristic of bark beetles of the tribe Ipinae. The head is covered by a thoracic shield and not visible when viewed dorsally. The abdominal declivity is concave with each side bearing from three to six spines (Wood S.L. 1982).

Table 9.3 Representative species of Ips (Coleoptera: Curculionidae: Scolytinae): their distribution and hosts. Species

Distribution

Major hosts

I. acuminatus (Gyllenhal) I. avulsus (Eichhoff)

Eurasia USA: southeastern states

3 4

I calligraphus (Germar) Six-spined engraver I. cembrae (Heer) I. confusus (LeConte)

North and Central America, Jamaica Europe North America: southwestern USA, northern Mexico North America (indigenous) Australia (introduced) Asia: China, Kazakhstan, Kyrgyzstan, Russia, Tajikistan and Turkey

Pinus Pinus echinata, P. elliottii, P. palustris, P. taeda Pinus Larix, other conifers Pinus edulis, P. monophylla

4 5

Pinus

5

Larix sibirica, Picea schrenkiana, Pinus sylvestris, P. nigra ssp. pallasiana Picea pungens

4

4

Pinus

5

Picea, Pinus

4

Larix, other conifers

4

Pinus, other conifers

6

Picea, Pinus

5

I. grandicollis (Germar) I. hauseri Reitter Mountain Kyrgyz engraver

I. hunteri Swaine Blue spruce ips I. lecontei Swaine

I. pini (Say) Pine engraver I. subelongatus Motschulsky I. sexdentatus (Boerner)

I. typographus (Linnaeus) Larger European spruce beetle

North America: USA: Colorado, Utah Southwestern USA south through Mexico, Guatemala and Honduras North America: transcontinental distribution Asia: Russia, northern China, northern Mongolia Asia: Turkey across Russia to China and south to Thailand. Europe Eurasia

Number of spines on each side of declivity

6

n Tovar et al. 1995, Bright & Sources: Furniss & Carolin 1977, Gru¨ne 1979, Wood S.L. 1982, Drooz 1985, Abgrall & Soutrenon 1991, Cibria Skidmore 2002, Kimoto & Duthie-Holt 2006, McMillin & DeGomez 2008, FAO 2009b.  Indicates species capable of causing top kill.

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(exotic) and P. merkusii and are hosts in Thailand. Other conifer hosts include Abies nordmanniana (¼ A. bornuelleriana), Larix decidua, Picea obovata and P. orientalis.

Importance This insect usually attacks weakened or windthrown trees. When populations build up in this material, they can attack relatively healthy trees. In some instances, I. acuminatus can kill large numbers of trees. Attacks often occur first in the upper crown causing top kill and subsequent generations attack the lower bole (Plate 45). Life History This beetle has one to two generations/ year. Adult flight occurs in the northern parts of its range or at high altitudes from May to June. In the southern parts of its range or at low altitudes adults fly from April to August. Description of Stages Adults are about 2.2–3.5 mm long and dark red-brown in color. Each side of the abdominal declivity has three spines (Grüne 1979, Vongkalung 1990, Abgrall & Soutrenon 1991, Wood & Bright 1992, Bright & Skidmore 2002). Ips calligraphus (Germar), Six-Spined Engraver Beetle

Fig. 9.12 X-shaped gallery pattern of the six-spined engraver, Ips calligraphus, showing nuptial chamber and four egg galleries, each made by a different female.

Ips acuminatus (Gyllenhal) Distribution I. acuminatus occurs across Eurasia. In Asia it is found from Turkey across Russia to China, Japan, Korea, Mongolia, Syria, Taiwan and Thailand. In Europe it occurs from Spain north to Finland, Norway and Sweden and east to Latvia, Romania and the Republic of Georgia. Hosts Pinus spp. are the predominant hosts. In Europe and the Near East, Pinus cembra, P. mugo, P. nigra and P. sylvestris are attacked. In China, Korea and Mongolia, P. armandii, P. koraiensis, P. sylvestris var. mongolica and P. tabulaeformis are attacked. P. caribaea

Distribution This species is native to north and central America and parts of the Caribbean Basin. It is found in Quebec, Canada and throughout much of the USA. It is also widely distributed in Mexico and throughout the natural range of pine forests in Belize, Guatemala, Honduras and Nicaragua. In the Caribbean, it has been collected in the Bahama Islands, Dominican Republic and Jamaica.

Hosts Species of Pinus, including P. caribaea, P. echinata, P. elliottii, P. michoacana, P. montezumae, P. occidentalis, P. oocarpa, P. ponderosa, P. pseudostrobus, P. resinosa, P. rigida, P. strobus and P. taeda are hosts.

Importance I. calligraphus prefers to invade fresh windthrow, freshly cut stumps and large limbs of recently felled trees. It can also attack living pines stressed by drought or invaded by more aggressive bark

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beetles. Attacks on living trees usually occur in the lower portions of the bole where diameters exceed 15 cm.

Life History The number of generations varies depending on local climatic conditions. In southeastern USA, a generation can be completed in as little as 25 days and there may be as many as six generations/year. Males are joined by three to five females and egg galleries radiate longitudinally from a central nuptial chamber. Larval galleries are often long and transverse.

Description of Stages Adults are 3.8–5.9 mm long and about 2.7 times as long as they are wide. Color of mature adults is red-brown. Each side of the elytral declivity is armed with six spines.

Related Species I. sexdentatus (Boerner) is found across Eurasia where it attacks several species of Larix, Picea and Pinus. This species also bears six spines on either side of the elytral declivity. It is also regarded as a secondary invader often found in association with other bark beetles (Wood S.L. 1982, Drooz 1985, FAO 2009b).

Ips grandicollis (Germar), Southern Pine Engraver Distribution This engraver beetle is widely distributed in eastern North America from Manitoba and southern Quebec, Canada, to Florida and east Texas, USA and south through Mexico to the Bahamas, Cuba, Dominican Republic, Honduras and Jamaica. It was introduced into South Australia around 1943, probably via crating or dunnage and was discovered in Western Australia in 1952. It has since spread to New South Wales, Queensland and Victoria.

Importance Within its natural range, I. grandicollis confines its attacks to standing pines stressed by drought, mechanical injury or other factors. It also attacks freshly cut logs or windthrow. In southeastern USA, I. grandicollis is often associated with other bark beetles including Dendroctonus frontalis, D. terebrans, I. avulsus and I. calligraphus. In Australia, it also attacks stressed, recently felled and windthrown trees but tends to be somewhat more aggressive than in its natural range.

Life History The number of generations/year depends on location and local climate. In southeastern USA, there may be up to six generations/year and a generation can be completed in 25–30 days.

Description of Stages Adults are 2.6–4.6 mm long and red-brown. Each side of the elytral declivity is armed with five spines.

Pest Management In Australia, I. grandicollis has been the target of a classic biological control program using natural enemies imported from the USA. One parasitoid, Roptrocerus xylophagorum, has been established. Three other natural enemies, a parasitoid, Dendrosoter sulcatus, and two predaceous beetles, Thanasimus dubius and Temnochila virescens, have been reared and released but their establishment is doubtful (Wood S.L. 1982, Drooz 1985, Morgan 1989, Lawson & Morgan 1992).

Ips pini (Say), pine engraver (Plate 46) Distribution Pine engraver is a North American species with a transcontinental distribution and is one of the most common pine bark beetles.

Hosts This species can breed in almost any species of Pinus and also infests species of Picea. Hosts Ips grandicollis breeds in species of Pinus, including P. banksiana, P. caribaea, P. durangensis, P. echinata, P. montezumae, P. oocarpa, P. palustris, P. ponderosa, P. pseudostrobus, P. resinosa, P. rigida, P. strobiformis, P. sylvestris, P. taeda, P. tenuifolia and P. virginiana. In Australia it attacks plantations of P. radiata and other introduced pines.

Importance Pine engraver populations build in windthrow, freshly cut logs or logging residues and subsequent generations can attack standing trees. Attacks in standing trees often occur during dry weather, begin in the upper crown and cause top kill.

Bark and ambrosia beetles The lower portions of the bole may become infested by subsequent generations of pine engraver or by species of Dendroctonus (e.g. D. brevicomis, D. ponderosae).

Life History Depending on local conditions, there may be one to five generations/year. Parent adults often re-emerge and make attacks in two to three trees, which causes a confusing overlapping of broods. During late summer, large numbers of beetles may mine under the bark without producing brood. Winter is spent almost exclusively as adults under the bark or in the litter. Description of Stages Adults are red-brown and 3.5–4.2 mm long. Each side of the elytral declivity has four spines. Pest Management Cultural tactics include prompt disposal of logging residues and utilization of windthrown trees. Thinning of overstocked pine stands will reduce hazard of attacks by this insect (Furniss & Carolin 1977, Drooz 1985).

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lasts about 15–17 days. After making initial attacks and laying eggs, some mated females will attack additional host material during late June–early July. Mature adults overwinter in the litter and larvae, pupae and callow adults overwinter underneath the bark of host material. I. subelongatus may continue to attack the same tree over several years. Description of Stages Adults are 4–6 mm long, brown to black in color and have four equally spaced spines on the elytral declivity. The third spine is the largest. The surface of the elytral declivity is covered with long hairs.

Related Species I. cembrae (Heer) is distributed throughout the Larix forests of the Alps and Carpathians in central Europe but has spread to plantations in the Netherlands, Scotland and other countries. Its damage, life history and morphological characteristics are similar to I. subelongatus (Grüne 1979, EPPO 2005b, FAO 2009b). Ips typographus (Linnaeus), Larger European Spruce Bark Beetle (Plates 47 & 48)

Ips subelongatus Motschulsky Distribution This species is indigenous to European Russia and much of the boreal conifer forests of northern Asia including Siberia, Transbaikalia and the Russian Far East, China, North and South Korea and northern Mongolia. It has been introduced into Finland. Hosts Primary hosts are species of Larix, including L. gmelinii, L. leptolepis and L. sibirica. It may occasionally breed in other conifers including Abies spp., Picea spp., Pinus koraiensis, P. sibirica and P. sylvestris. Importance I. subelongatus is considered one of the most damaging pests of larch across Russia and other Asian countries. Attacks often occur in trees previously defoliated by Dendrolimus sibiricus and/or attacked by wood borers such as Xylotrechus altaicus, other insects or in forests damaged by fire. Life History Adult flight usually occurs from midMay to late June in the southern parts of its range and

Distribution This engraver beetle is widely distributed across Eurasia. It occurs over most of Europe and has been introduced into the British Isles. In Asia, it occurs in China, Japan, Korea, Turkey and Asian portions of Russia.

Hosts I. typographus attacks species of Picea. Known hosts are P. abies, P. jezoensis, P. obovata and P. orientalis. Other members of the Pinaceae are also attacked, including species of Abies, Larix and Pinus. According to one report, preferred hosts of Ips typographus in the Caucasus region are pines. P. jezoensis, native to Asia, is attacked by the subspecies, I. typographus japonicus Niijima. The North American P. sitchensis, widely planted in the British Isles, is also attacked but thus far has not been subject to major outbreaks.

Importance This species is a major tree killer of Eurasian spruce forests. Adults attack fresh windthrow or logs stored in the forest for prolonged periods

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Forest entomology: a global perspective complete two generations/year further south. In the north, adults emerge from July to October, depending on time of brood establishment, microclimate and weather. Further south there are two peak flights: May–June for the overwintering adults and July–August for the summer generation. The second generation may emerge in November, but more typically, adults hibernate in the brood tree or forest litter and emerge the following spring. I. typographus overwinters as adults, usually in the duff near the tree where they developed. A few individuals remain beneath the bark during winter, especially in the southern part of its range.

Description of Stages Adults average 4–5.5 mm long and are dark brown in color. Each side of the abdominal declivity has four spines.

Fig. 9.13 Standing Norway spruce attacked by the larger European spruce bark beetle, Ips typographus, following a buildup of beetle populations in windthrow (Bavarian National Park, Germany).

(Fig. 9.13). Subsequent generations attack standing trees. Outbreaks in central and northern Europe have resulted in trees being killed over large areas with losses totaling several million cubic meters of wood. A 7-year outbreak following World War II killed 30 million m3 of spruce in Germany. Some outbreaks in German and Norwegian forests have lasted for 30–50 years. In Norway, an outbreak in the 1970s, which killed 5 million m3 of spruce, led to a substantial reduction of the country’s gross national product.

Life History The number of generations/year depends on temperature. In the northern parts of its range, I. typographus has one generation/year. It can

Pest Management Outbreaks can be prevented through reduction of the amount of host material available to the insect. Cultural tactics include prompt salvage or debarking of windthrown spruces and debarking of logs stored in the forest for extended periods. Direct control includes use of attractant or repellent pheromones to either trap out beetles or reduce attacks on suitable host material (Browne 1968, Grüne 1979, Schwerdtfeger 1981, Krivolutskaya 1983, Forsse & Solbreck 1985, Duelli et al. 1986, Christiansen & Bakke 1988, Eidmann 1992, Wood & Bright 1992, Lozzia 1993, Pfeffer & Skuhravy 1995, Yamaoka et al. 1997, Bright & Skidmore 2002). Orthotomicus Orthotomicus is a small genus of bark beetles closely related to Ips and found in Eurasia and North America. Adults resemble Ips but the spines on the abdominal declivity are less distinct. Twelve species are known worldwide (Wood S.L. 1982).

Orthotomicus erosus (Wollaston), Mediterranean Pine Engraver Distribution O. erosus is widely distributed across northern Africa, Asia and Mediterranean Europe. It has been introduced and become established in California, USA, Chile, South Africa and Swaziland.

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Hosts Primary hosts are pines: Pinus spp., including P. brutia, P. canariensis, P. nigra, P. pinaster, P. pinea, P. sylvestris and P. mugo ssp. uncinata (Europe); and P. armandii, P. kesiya, P. massoniana, P. tabulaeformis and P. yunnanensis (Asia). North and Central American pines planted in areas where this insect is native, or has become established, and have become hosts include P. caribaea, P. coulteri, P. echinata, P. patula, P. radiata and P. strobus. Occasionally, maturing beetles feed in Abies, Cedrus, Picea and Pseudotsuga menziesii. However, it does not breed in hosts other than pines. Importance O. erosus is usually secondary and infests recently fallen trees, broken branches, slash and standing trees that have been wounded or are under stress. During periods of prolonged dry weather, a relatively common occurrence in its natural range, it will attack standing trees. Life History Mediterranean pine engraver completes from two to seven generations/year, depending on temperature. Two generations/year are common in France, Morocco and Turkey. In Israel, it can complete three to five generations/year. Adults overwinter and aggregate beneath the bark of host trees. This species is polygamous. Males bore through the bark to the phloem–cambium layer where they construct a nuptial chamber and are joined by one to three females. After mating, females constructs individual egg galleries radiating from the nuptial chamber and parallel to the grain of the wood. Females lay 26–75 eggs in niches along the sides of the galleries. Larvae feed at right angles to the parent gallery and pass through three instars during their development. Pupation occurs in cells at the end of each gallery (Fig. 9.14). Brood adults feed prior to reaching sexual maturity. This occurs under the bark of the host tree or in another suitable host tree, sometimes of a different species.

Fig. 9.14 Egg and larval galleries of Mediterranean pine engraver beetle, Orthotomicus erosus (northern Cyprus).

prevent population increases. In South Africa, where O. erosus was introduced, a parasitic wasp, Dendrosoter caenopachoides, was introduced (Grüne 1979, Mendel & Halperin 1982, Mendel 1983, Tribe & Kfir 2001, Haack 2004).

Curculionidae (Subfamily Scolytinae – Ambrosia Beetles) Description of Stages Adults are 3–3.8 mm long and red-brown. The head is covered by the thoracic shield and not visible when viewed dorsally. Declivity is concave, with each side armed by four small spines. The second spine from the top is more conspicuous.

Pest Management Prompt treatment of logging slash, windthrow and other potential habitat will

Gnathotrupes Ambrosia beetles of the genus Gnathotrupes invade broadleaf trees and are found in Mexico, Central and South America. They are well represented in Argentina and Chile where they attack species of Nothofagus. About 15 species are known from this area. They are small beetles, 1.3–3.6 mm long, and brown. When

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viewed from above, the head is concealed by the prothorax and they have a concave declivity. Relatively little was known about this group of ambrosia beetles until about 2002 when several species were associated with dieback and mortality of Nothofagus spp. in southern Chile (11th Region). Affected trees include N. betuloides, N. dombeyi and N. pumilio. Attacks occur in groups of trees with chlorotic foliage, branch dieback and tree mortality. At least four species are involved. Gnathotrupes barbifer Schedl, G. nanus (Eichhoff), G. vafer Schedl and G. velatus Schedl. It is not clear if the fungi associated with this group of ambrosia beetles are pathogenic or if the beetles are a contributing factor in a decline event. Females construct galleries and larval cradles while males guard the entrance holes and keep the galleries free of frass (Wood 2007, Aguayo Silva 2008). Trypodendron Trypodendron is a small genus of monogamous ambrosia beetles found in Eurasia and North America, north of Mexico. About 12 species are known. Females initiate attacks and carve an entrance tunnel in a host tree prior to arrival of the male. The tunnel penetrates the bark and continues into the sapwood where it may branch several times. Larvae are reared in cradles arranged in a single series above and below the parent gallery. Cradles are enlarged by larvae as they grow and also serve as pupal chambers (Figs 9.15 & 9.16, Wood S.L. 1982). Trypodendron lineatum (Olivier), Striped Ambrosia Beetle Distribution This ambrosia beetle is found throughout conifer forests of northern Asia, Europe and North America. Hosts Virtually all species of Abies, Larix, Picea, Pinus, Pseudotsuga, Thuja and Tsuga are hosts. It has occasionally been reported from species of Alnus, Betula, Juniperus and Malus.

Importance Infestations cause pinholes with dark stain in the sapwood and reduce lumber quality. T. lineatum is considered the most damaging ambrosia beetle in western North America, especially in coastal

Fig. 9.15 Galleries of an undetermined species of Trypodendron.

British Columbia, Canada. Populations build in windthrow, trees killed by fire or bark beetles, logging residues and logs stored for extended periods. Logs cut in autumn and early winter are most susceptible to attack.

Life History In forests of the Pacific Coast of North America one generation/year is typical but a small portion of adults re-emerge and establish a second brood. Adults become active and attack host material beginning in March, peak in May and continue until August. Development time from egg to adult requires 6–8 weeks. Brood adults emerge from July to September and overwinter in the duff and litter.

Description of Stages Adults are dark brown-black with two yellow-brown longitudinal bands on each elytron. Females are 3.0–3.5 mm long; males are 2.7–3.2 mm long.

Pest Management Spraying logs with chemical insecticides can prevent attacks but has caused

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Fig. 9.16 Individual larval cradles characteristic of ambrosia beetles of the genus Trypodendron.

environmental problems. Prompt processing of logs prior to, and during, beetle flight reduces attacks. Mass trapping of adults in timber storage and processing areas was developed after its aggregation pheromone was synthesized and became commercially available during the 1980s (Bletchley & White 1962, Furniss & Carolin 1977, Wood S.L. 1982, Lindgren & Borden 1983).

usually of a simple branching pattern that may join galleries constructed by other individuals (Fig. 9.17). Eggs are usually placed in clusters at or near the end of branch tunnels. Larvae feed on ambrosia fungi, pupate in the galleries and emerge through entrance holes made by the parent adults (Wood & Bright 1992, Wood 2007) (Table 9.4).

Xyleborus ferrugineus (Fabricius) Xyleborus Xyleborus is the largest and most diverse genus of the subfamily Scolytinae. Several hundred species have been described from all of the tropical and subtropical areas of the world and some species extend their ranges into temperate forest regions. In the tropics, members of this genus can cause significant damage through the destruction of sapwood of recently harvested logs, both in the forest and at sawmill sites. They typically have a wide host range and several species have been introduced and become established. They are polygamous. Males are often reduced in size, flightless and may not leave the parent gallery. Galleries are

Distribution This species occurs from southern USA, south through much of the Caribbean, Mexico, Central and South America south to northern Argentina. It has been introduced into portions of tropical Africa, southern India, Sri Lanka and Micronesia.

Hosts A large number of broadleaf trees are reported hosts, including Carya illinoiensis, Coffea spp., Couma macrocarpa, Eschweilera corrugata, Eucalyptus spp., Lonchocarpus margaretensis, Melicoccus bijugatum, Pithecellobium pinnatum, Protium spp., Sacoglottis procera, Swietenia macrophylla and Theobroma cacao.

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Fig. 9.17 Galleries of an undetermined species of Xyleborus.

Table 9.4 Representative species of Xyleborus (Coleoptera: Curculionidae: Scolytinae): their distribution and hosts. Species

Distribution

Hosts

X. affinis Eichhoff

Tropical America (native) Tropical Africa, southern India, Sri Lanka, Australia, Micronesia and Japan (introduced) Southern USA south through the Caribbean Basin, Mexico, Central and South America south to northern Argentina (native) Tropical Africa, southern India, Sri Lanka and Micronesia (introduced)

Several hundred broadleaf trees

X. ferrugineus (Fabricius)

X. glabratus Eichhoff, Redbay ambrosia beetle X. perforans Wollaston

X. similis Ferrari

India, Japan, Myanmar and Taiwan (native) Southeastern USA (introduced) Cosmopolitan in tropical regions including Africa, Asia, Australia, Caribbean Basin and South America Asia and the Pacific Islands, Mauritius, Tanzania

Sources: Browne 1968, Bright & Skidmore 2002, Wood 2007.

Carya illinoiensis, Coffea spp. Couma macrocarpa, Eschweilera corrugata, Eucalyptus spp., Lonchocarpus margaretensis, Melicoccus bijugatum, Pithecellobium pinnatum, Protium spp., Sacoglottis procera, Swietenia macrophylla and Theobroma cacao Lindera latifolia, Lithocarpus edulis, Litsea elongata, Phoebe lanceolata, Shorea robusta More than 100 host species recorded

Numerous hosts, including tapped and injured Hevea brasiliensis

Bark and ambrosia beetles Importance X. ferrugineus aggressively attacks recently felled logs in forests, log decks and sawmill sites and causes complete destruction of sapwood. It is considered one of the most destructive ambrosia beetles of harvested timber in South America. In Mexico, it attacks pecan plantations and in Brazil it is reportedly one of several ambrosia beetles that attack Eucalyptus plantations. It commonly attacks small-diameter stems as well as large logs but is rare in undisturbed natural forests. It is a vector of the pathogenic fungus, Ceratocystis fimbriata, which causes a wilt disease of cacao.

Life History Newly emerged mated or unmated females fly to seek a new host, usually in early evening. They construct multi-branched tunnels that penetrate deep into the sapwood. Tunnels rarely enter the heartwood. Surface tunnels in the cambium, are common in wet habitats but rare in dry habitats.

Description of Stages Males are 1.6–1.9 mm long and females are 2.0–3.3 mm long. When mature, they are dark red-brown in color. The elytral declivity is steep.

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plants of the family Lauraceae, including avocado, Persea americana, redbay, P. borbonia and sassafras, Sassafras albidum.

Importance This species is the vector of a highly pathogenic fungus, Raffaelea lauricola, which causes a wilt disease and mortality of host plants in southeastern USA. In some locations redbay mortality has exceeded 90%. The palamedes swallowtail butterfly, Papilio palamedes Drury, is dependent upon redbay as a larval host.

Life History Little is known about the life history and habits of this insect. For most species of Xyleborus, males are rare and do not fly. Only females attack host material and construct galleries and brood cradles. If mating occurs, it takes place before adults exit the host plant and is usually between siblings or mother and son. Males are haploid and females are diploid. An unmated female produces only male offspring, with which she later mates to produce females.

Related Species X. affinis Eichhoff is also native to the neotropics and has been introduced to tropical Africa, southern India, Sri Lanka, Australia, Micronesia and Japan. It has a wide host range and is similar in habits to X. ferrugineus (Browne 1968, Flechtmann et al. 2000, Aguilar-Perez et al. 2007, Wood 2007, Wagner et al. 2008).

Description of Stages Adults are small beetles, 2.0 mm long, slender and brown-black in color. The declivity is steep and convex, especially on the posterior portion (Wood & Bright 1992, Rabaglia 2005, Mayfield & Thomas 2009).

Xyleborus glabratus Eichhoff, Redbay Ambrosia Beetle

Megaplatypus (¼ platypus sulcatus) mutatus (Chapuis)

Distribution X. glabratus is indigenous to India, Japan, Myanmar and Taiwan. It was introduced into southeastern USA in about 2002, most likely via solid wood packing material. It was first collected in Georgia, USA, and has since been found in South Carolina and Florida.

Distribution M. mutatus is native to tropical and subtropical areas of South America and reported from Argentina, Bolivia, Brazil, French Guyana, Paraguay, Peru, Uruguay and Venezuela. It has extended its range into temperate regions of the continent, as far south as Neuquen, Argentina. It was detected in Italy near Caserta (Campania) in 2000.

Hosts This insect infests broadleaf trees and shrubs. Known host trees in its natural range include Lindera latifolia, Lithocarpus edulis, Litsea elongata, Phoebe lanceolata and Shorea robusta. In southeastern USA, it infests

Curculionidae (Subfamily Platyponinae – Ambrosia Beetles)

Hosts M. mutatus has a wide host range. It can invade many temperate broadleaf trees including

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species of Acer, Caesalpinia echinata, Citrus, Eucalyptus, Fraxinus, Laurus nobilis, Magnolia grandiflora, Malus, Platanus, Populus, Prunus persica, Persea americana, Pyrus communis, Quercus, Robinia pseudoacacia, Salix, Tilia and Ulmus.

Importance Adults bore into living trees up to about 4 m above ground level. Trees of larger diameters are preferred. They construct long sinuous galleries lined with black fungus mycelium of the ambrosia fungus, Raffaelea santoro. Attacks degrade lumber and cause structural damage. Infested trees may break during high winds. This insect has caused extensive damage to Populus and Salix in Argentina. It has also been reported as a pest of Brazilwood, Caesalpinia echinata, plantations in Brazil. This species poses a threat to poplar plantations worldwide.

Life History Precise data on flight distances are unavailable. However, dispersal is believed to be not more than 100 m for a few individuals when there is a large infestation. After emergence, the adults must find a suitable host within 5 days. Males initiate attacks, construct a short nuptial gallery and attract one or more females. After mating, females construct galleries and cradles for brood and inoculate them with the ambrosia fungus.

Description of Stages Adults are long, thin cylindrical beetles. Body color is dark red-brown with lighter colored antennae and red-brown legs (Casaubon et al. 2006, Girardi et al. 2006, Alfaro et al. 2007, FAO 2007b, EPPO 2007). Myoplatypus (¼ platypus) flavicornis (Fabricius) Distribution This species occurs in eastern USA, from New Jersey south to Florida and west to Texas and Mexico. Hosts Hosts are species of Pinus. It also is occasionally found in broadleaf trees.

Importance The lower boles of dead and dying pines, especially those recently killed by bark beetles

such as Dendroctonus frontalis, are invaded by this insect. Stumps and logs cut during summer are also attacked. In southeastern USA, this species is so abundant that few dying pines, stumps or logs escape attack, which is indicated by copious amounts of fine white boring dust near the base of host trees (Fig. 9.18). Adult galleries and cradles and discoloration associated with the ambrosia fungus causes loss of wood quality.

Life History Galleries are initiated by males and each male is joined by a single female. Apparently pheromones are produced and simultaneous attacks follow. Mated pairs tunnel into the sapwood and introduce ambrosia fungi into the galleries, which may branch extensively and extend into the heartwood. Larvae move freely inside the parental tunnels and excavate individual pupal cells off the main tunnels. Adults emerge through the original entry hole. Normally, only one generation is produced in a tree.

Description of Stages Adults are red-brown and about 5 mm long. The front of the head is flat and clothed with moderately long hairs (Drooz 1985). Platypus Platypus is a large genus of ambrosia beetles. Over 500 species are recognized. They occur in both temperate and tropical forests and damaging species are known from Africa, Asia, Australia, Europe and North and Central America (Browne 1968, Bright & Skidmore 2002). Platypus cylindrus (Fabricius), Oak Pinhole Borer Distribution Oak pinhole borer is found throughout much of Europe from southern England south to the Mediterranean region. It is also found in North Africa (Algeria, Morocco) and the Near East (Iran). Hosts Oaks, Quercus spp., are preferred hosts but other members of the family Fagaceae, including Castanea sativa and Fagus sylvatica, may be attacked. Other reported hosts are Eucalyptus spp. and Populus caspica.

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Fig. 9.18 Heavy boring dust at base of a loblolly pine, Pinus taeda, indicative of attack by the ambrosia beetle Myoplatypus flavicornis (east Texas, USA, photo by R. F. Billings, Texas Forest Service, courtesy of www.forestryimages.org).

Importance P. cylindrus invades severely stressed or recently dead trees. In Algeria, Morocco and Portugal, it is considered a contributing factor in the decline of cork oak, Quercus suber. In the UK, its status has changed from a rare species to a pest. As of 1987, it was listed in the British Red Data Book as rare. However, after a severe storm in 1987, there was an abundance of breeding material and numbers increased dramatically in the early 1990s with concurrent damage. Numbers have continued to be high, probably because of an abundance of oaks affected by dieback and decline.

Description of Stages Adults are 6–8 mm long, cylindrical and dark brown-black in color.

Pest Management Management of infestations includes timely removal and processing of logs to prevent attack and application of chemical sprays to logs in wood yards (Browne 1968, Bright & Skidmore 2002, Henriques et al. 2008, Tilbury 2009).

Platypus granulosus Browne Life History A generation usually takes 2 years to complete although some individuals may complete development in 1 year. Adults are most active between July and mid-September when males bore into logs and stumps of host trees. They are attracted to odors of fermenting sap. Males initiate attacks, bore into the wood for a short distance and then each is joined by a single female. After mating, females construct galleries in the wood and males keep them free of frass. Females lay eggs in cradles constructed in the wood, which they inoculate with ambrosia fungi. Eggs hatch after 2–6 weeks and larvae pass through four or five instars as they feed on ambrosia fungi. The later instars have large mandibles, which they use to extend the galleries.

Distribution P. granulosis beetle is indigenous to Australia and occurs on the island of Tasmania.

Hosts Primary host is Nothofagus cunninghamii. Other hosts include Anodopetalum biglandulosum, Antherosperma moschatum, Eucalyptus spp., Eucryphia lucida, Phyllocladus aspleniifolius and Pinus radiata.

Importance Nothofagus cunninghamii apparently is killed by a fungus associated with P. granulosus, but not the ambrosia fungus that larvae use for food. Other

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hosts are not killed but galleries and discolored wood reduce value of lumber cut from infested trees. Trees with mechanical injury or growing adjacent to recently disturbed areas, such as roads or sites where timber has been harvested, are most susceptible to attack. Recently cut logs and green lumber are also attacked. Life History One year is required to complete a generation. Adults emerge from infested trees and logs in summer and males initiate attacks on suitable host material. After boring about 10 mm into the wood, they await arrival of a female. When females arrive, they move from entrance hole to entrance hole until they locate an unmated male. Mating occurs at the entrance hole. Females then elongate the chamber made by males and bore to the heartwood/sapwood boundary. Boring dust is expelled from the entrance hole by the males. Females lay eggs either singly or in small groups in the tunnel system. When larvae are about 4 mm long, they stop feeding, construct small chambers in the wood and pupate. Description of Stages Adults are about 4 mm long, brown in color and cylindrical (Elliot and deLittle n.d.). Platypus quercivorus Murayama Distribution P. quercivorus is widely distributed in Asia, including temperate, subtropical and tropical portions of India, Indonesia (Java), Japan, Papua New Guinea and Taiwan. In Japan, it is found from the island of Ishigaki Shima north to Honshu. Hosts This insect has a wide host range and invades many broadleaf trees including species of Fagaceae. Castanopsis cuspidata, Pasania glabra, P. (¼ Lithocarpus) edulis, Prunus spp., Quercus acuta, Q. acutissima, Q. gilva, Q. glauca, Q. mongolica (¼ Quercus crispula), Quercus mongolica var. grosseserratus, Q. myrsinifolia (shirakashi), Q. phillyraeoides, Q. salicina, Q. serrata and Q. sessilifolia. Other hosts are Cryptomeria japonica, Ilex chinensis and Lindera erythrocarpa. Importance Beginning in the early 1980s, extensive mortality of oak forests has occurred in western Japan. This condition, referred to as “Japanese oak disease,” is attributed to the fungus Raffaelea quercivora, which is a fungal associate of P. quercivorus. Oak mortality at the

rate of more than 200,000 trees/year has been observed on the west coast of Honshu. Oaks susceptible to the disease are the deciduous species Quercus serrata and Q. mongolica. Other trees of the family Fagaceae present in the area, for example Q. acuta, Q. acutissima, Q. phillyraeoides and Castanopsis cuspidata var. sieboldii, apparently are not affected. It is believed that oaks resistant or tolerant to Raffaelea quercivora co-evolved under a stable relationship between the tree, fungus and beetle during a long evolutionary process. Q. mongolica may not have been part of this co-evolution. This is supported by the fact that P. quercivorus has a low preference for Q. mongolica but exhibits the highest reproductive success in this species. Therefore, P. quercivorus could spread more rapidly in stands with a high component of Q. mongolica. The present epidemic of P. quercivorus may be due to a warmer climate that began in the late 1980s. This allowed it to extend its range to more northerly latitudes and higher altitudes where Q. mongolica occurs. Life History Males initiate attacks on the boles of host trees and excavate galleries for mating from June to October. Initial entry holes bored by males trigger a mass attack. Attacks generally occur near the ground level. A single female joins the male and, after mating, constructs a gallery, which is kept clean by the male. During gallery construction, females inoculate the gallery surface with fungus spores. Adult females deposit eggs at the terminal parts of tunnels 2–3 weeks after the beginning of gallery construction. Eggs are deposited in individual niches. An average of 50–60 larvae develop in a single gallery system but the number can be as high as 160. Larvae feed on the ambrosia fungus that develops on the walls of the galleries. Pupation occurs in the larval galleries. Most brood adults leave their maternal galleries in September and October but some remain until spring. In other cases larvae reach the instar V by late November and overwinter in pupal chambers. Pupation begins the following May, and adults emerge from June to July through entry holes made by parent adults. Description of Stages Adults are red-brown to dark brown with a cylindrical, elongated body that averages 5 mm long. They have a concave declivity armed with spines. The front (prothoracic) legs are adapted for excavation (Wood & Bright 1992, Bright & Skidmore 2002, Kamata et al. 2002, Kubono & Ito 2002, Kobayashii et al. 2003).

Chapter 10

Large Cambium and Wood Boring Insects

INTRODUCTION Many insects, in addition to the bark and ambrosia beetles discussed in the preceding chapter, feed on and breed in the woody tissue of trees. Most wood boring insects are secondary and attack trees weakened by drought, fire, insect defoliation, mechanical injury or other stress. Others attack and bore into the wood of fresh cut logs. Wood borers can reduce quality of lumber sawn from infested logs and damage structural integrity of infested trees, making them more susceptible to wind damage. Several more aggressive species attack live trees and repeated attacks can cause tree death. Still others are vectors of pathogenic fungi or nematodes. Some wood boring insects have long life cycles and require 2 or more years to complete a generation. This allows them to be transported to new locations via international trade in wood products. Once established some species have become extremely damaging. Increased world trade and human travel over the past 20–30 years has accelerated the rate of introduction and establishment of wood boring insects.

This chapter addresses insects of the orders Coleoptera, Lepidoptera and Hymenoptera that utilize woody tissue of tree boles for food and breeding sites. Insects that attack smaller stems and branches are reviewed in Chapter 13 and those that are primarily pests of wood in use are reviewed in Chapter 15.

COLEOPTERA (BEETLES) Buprestidae (Flat Headed Wood Borers) Agrilus Agrilus is a large genus of over 1000 described species. They occur throughout the world’s forests and rangelands. Some bore in stems of woody or herbaceous plants, for example, red necked cane borer, A. ruficollis (Fabricius), bores in stems of both cultivated and wild raspberries, Sorbus spp. Several species indigenous to Texas, USA, and Mexico, including A. andersoni Hespenheide, A. howdenorum Hespenheide and A. turnbowi

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Table 10.1 Distribution and principle hosts of some important forest species of Agrilus (Coleoptera: Buprestidae). Species

Distribution

Principal hosts

A. angelicus Horn A. anxius Gory Bronze birch borer

USA: California Canada, northern USA

Quercus spp. Betula alleghaniensis, B. papyrifera

A. bilineatus (Weber) Twolined chestnut borer A. coxalis Waterhouse Goldenspotted oak borer

Eastern Canada and USA

Quercus spp.

Guatemala, Mexico USA: Arizona (recently detected in southern California) Eastern Canada and USA North America

Quercus spp.

Cyprus Asia: China, Japan, Korea, Mongolia, Russian Far East, Taiwan North America (introduced) Europe, North Africa, Near East, Russia (east to Siberia) Europe, including British Isles

Quercus alnifolia Fraxinus spp.

A. granulatus (Say) A. liragus Barter & Brown Bronze poplar borer A. roscidus Kienwetter A. planipennis Fairmare Emerald ash borer A. pannonicus Piller & Mitterpacker A. viridis Linnaeus

Populus spp. Populus spp.

Castanea, Fagus, Quercus Alnus glutinosa, Betula spp. Fagus sylvatica, Populus tremula, Quercus spp.

Sources: Browne 1968, Furniss & Carolin 1977, Drooz 1985, Turner & Hawkeswood 1996, Coleman & Seybold 2008.

Nelson, infest stems of mistletoes, Phoradendron spp. At least one species, the St John’s wort root borer, A. hyperici (Creutzer), is considered beneficial and is used to help control St John’s wort, a noxious weed introduced into western North America. Larvae of most Agrilus spp. feed in the cambium and xylem of broadleaf trees (Table 10.1). Adults feed on foliage of host trees before laying eggs. Eggs are laid in bark crevices. Larvae feed in the inner bark and wood and construct winding galleries in the cambium (Fig. 10.1). When mature, they construct pupal cells in the wood or outer bark. Adults emerge through a characteristic D-shaped exit hole in the bark. Infestations can kill all, or a portion of, trees. Adults have cylindrical to flattened bodies with tapered elytra and range in length from 2 to 75 mm. Many are brightly colored in iridescent, metallic hues. Larvae are long and slender, are white to yellow white, legless, with flattened or oval thoracic and abdominal segments and a pair of pincer-like appendages on the last abdominal segment (Bellamy & Houston 2002, Hespenheide 2008).

Agrilus anxius Gory, Bronze Birch Borer Distribution Bronze birch borer is a North American species and widely distributed in Canada and the USA.

Hosts Hosts are birches, especially Betula alleghaniensis and B. papyrifera.

Importance Weakened trees are preferred for attack but under suitable conditions it can attack live trees and cause extensive tree mortality. Between 1930 and 1950, a condition known as “birch dieback” occurred over large areas of eastern Canada and northeastern USA. The dieback was believed to have been incited by a prolonged warming trend that resulted in increased soil temperatures over a 10–20 year period and stressed birch forests. Bronze birch borer attacked and killed many of the stressed birches. This insect also attacks and kills trees stressed by drought or poor soils and is a pest of ornamental birches in urban settings.

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Agrilus bilineatus (Weber), Twolined Chestnut Borer Distribution This species is also native to North America, from the Maritime Provinces of eastern Canada west to the Rocky Mountains and south to Florida and Texas, USA.

Hosts Primary hosts are oaks, Quercus spp. It was also a pest of American chestnut, Castanea dentata, before chestnut blight fungus, Cryphonectria parasitica, was introduced and virtually eliminated this tree from eastern North American forests.

Importance A. bilineatus attacks and kills oaks stressed by drought, suppression or insect defoliation. It has been implicated as a contributing factor to North American oak decline. Tree death may occur following the first year of attack but is typically the result of attacks over 2–3 years.

Fig. 10.1 Winding galleries of Agrilus, probably A. liragus, on quaking aspen, Populus tremuloides (Roosevelt National Forest, Colorado, USA).

Life History In the northern parts of its range, 2 years are required to complete a generation. Further south, there is one generation/year. Adults are active from May or early June to August. They feed on birch foliage and lay eggs in small groups beneath loose curls of bark or in bark crevices. Young larvae feed in the cambium, construct winding galleries across the wood grain and overwinter in the xylem.

Description of Stages Adults are 6–12 mm long, deep green to bronze in color with copper-colored areas on the pronotum. The front of the head is green on males and copper-bronze on females (Hepting 1963, Drooz 1985).

Life History There is one generation/year. Adults are active from April to August and feed on foliage prior to egg deposition. Females lay eggs in clusters in bark cracks and crevices. Larvae hatch in 1–2 weeks and feed in the cambium, where they construct galleries. When full grown, they burrow into the outer bark, construct pupal chambers and overwinter. Pupation occurs the following spring.

Description of Stages Adults are slender, black with two golden stripes along the elytra. Length ranges from 5 to 13 mm (Haack & Acciavatti 1992). Agrilus coxalis Waterhouse, Goldenspotted Oak Borer Distribution This species has been known from Arizona, USA, Mexico and northern Guatemala for many years. In 2004, it was detected near San Diego, California, USA and in 2008 infestations occurred in oak forests of southern California.

Hosts In California, known hosts are Quercus agrifolia, Q. chrysolepis and Q. kelloggii.

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Importance This insect is part of an accelerated episode of oak decline in southern California. Life History Larvae feed in the cambium on the main stem and in larger branches. They kill strips of phloem and cambium and cause dieback and tree death.

Description of Stages Adults are 10 mm long, have a dark green body and elytra. The elytra are adorned with six golden-yellow spots (Coleman & Seybold 2008).

Agrilus pannonicus Pillar and Mitterpacker (¼ A. biguttatus Fabricius) Distribution A. pannonicus is a wide ranging species found throughout Europe, northern Africa, portions of the Near East and Russia as far east as Siberia.

Hosts Hosts are oaks, including Quercus cerris, Q. ilex, Q. petraea, Q. pubescens, Q. robur and Q. suber. Infestations occasionally occur on European beech, Fagus sylvatica, and chestnut, Castanea sativa. There are records of attacks in poplars, Populus spp., but these are considered doubtful. Infestations on North American red oak, Quercus rubra, planted within the geographic range of A. pannonicus are rare.

Importance This insect is a secondary invader of stressed and weakened trees and attacks trees defoliated by insects, such as Tortix viridana, damaged by frost or stressed by warm, dry summers. Infestations result in extensive tree mortality and it is considered a contributing factor in European oak decline. Over the past 20 years, attacks have increased over portions of Europe. In England, for example, this species was regarded as endangered as recently as 1987 but is now a common tree killer in oak woodlands and parks.

Life History A. pannonicus can have one generation/ year but a 2-year cycle is more common. In northern Germany, the larvae hibernate over two winters. Adult flight occurs from May to July. After feeding on oak foliage, females deposit clusters of five to six eggs in bark crevasses. The south-facing sides of large oaks are

preferred for oviposition. Larvae have five instars, feed in the cambium in longitudinal, winding galleries and overwinter inside the bark. When feeding is completed, larvae may have excavated a gallery up to 155 cm long. Pupae develop in the bark, in chambers 10.4–14.4 mm long and 3.0–4.5 mm wide. Adults remain under the bark for about 2 weeks in late spring–early summer and then emerge.

Description of Stages Adults are attractive, slender beetles, 9–12 mm long and metallic green, blue or gold. The posterior third of the elytra have two distinct white marks on their interior edge (Hartman & Blank 1992, Gibbs & Grieg 1997, Gibbs 1999, Moraal & Hilszczanski 2000).

Agrilus planipennis Fairmare, Emerald Ash Borer Distribution This borer is native to China, Korea, Japan, Mongolia, the Russian Far East and Taiwan. In 2002, adults were reared from dead and dying ash in southeastern Michigan, USA. Since its initial discovery in North America, infestations have been detected in Ontario and Quebec, Canada, and several states in the USA including Ohio (2003), northern Indiana (2004), northern Illinois and Maryland (2006), western Pennsylvania and West Virginia (2007), Missouri, Virginia and Wisconsin (2008), Minnesota and New York (2009) and Iowa and Tennessee (2010). Initial introduction into North America is believed to have been via infested wood, probably wooden crating or pallets containing strips of bark imported from Asia. Since its establishment in North America, long-distance overland spread has been facilitated by interstate shipment of fuel wood and nursery stock.

Hosts Species of ash, Fraxinus spp., are the primary hosts of this insect. In its native range, Manchurian walnut, Juglans mandshuria var. sieboldiana David, or Japanese elm, Ulmus davidiana var. japonica, and Japanese wingnut, Pterocarya rhoifolia, are hosts in its natural range. In North America, all indigenous species of ash are either attacked or considered potential hosts.

Importance A. planipennis attacks and kill live trees. In China, attacks occur on trees at the forest edge but

Large cambium and wood boring insects during outbreaks entire stands are killed. Attacks on individual trees may occur over 2–3 years and cause branch dieback, top kill and, eventually, tree death. Since its discovery in North America, this insect has killed tens of millions of ash trees in southeastern Michigan alone. Its presence poses a threat to all species of ash across the North American continent.

Life History Emerald ash borer typically has one generation/year but could require 2 years in colder climates. Adults emerge in May and are active until late June when they feed on foliage. After mating, females lay 65–90 eggs during their life time. They are deposited individually on bark surface or crevices on the main stem or branches. Eggs hatch in 7–10 days. Young larvae enter the cambium and construct S-shaped feeding galleries that become progressively wider as larvae grow. Galleries are packed with fine frass. Fullgrown larvae overwinter and pupate in late April–early May. Adults remain in pupal cells for 1–2 weeks and emerge. They can fly for at least 1 km in search of suitable hosts.

Description of Stages Adults are larger than any of the Agrilus species native to North America and range from 7.5 to 13 mm long. They are slender, elongate and gold-green overall, with darker, emerald green elytra (Plate 49). Mature larvae range from 26 to 32 mm in length, are cream colored and flattened with a brown hard capsule (Plate 50).

Pest Management Attempts have been made, with limited success, to prevent movement of infested nursery stock, logs or fuel wood from spreading out of infested areas via quarantines. In places where the insect is established, infested trees are removed and destroyed. Introduction and establishment of this extremely aggressive Agrilus into North America has presented a series of research and regulatory challenges and, despite all efforts, the insect continues to spread rapidly (Haack et al. 2002, McCullough & Roberts 2002, Cappeart et al. 2005). Agrilus spp. in Africa At least 10 species of Agrilus are known from east Africa and several occurrences of damage have been reported.

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During the 1950s, Agrilus near grandis Gory and Laporte attacked about 40% of the trees in a small plantation of Acacia mollissima in Kenya. Despite copious resin flow on the stems and branches, trees survived. During the 1990s, following a drought in the Sudan, an unidentified species of Agrilus, along with several other Buprestidae, attacked gum Arabic trees, Acacia senegal. The attacks resulted in reduced gum Arabic production, an important non-wood forest product used as a thickener for paints (Schabel 2006). Buprestis Buprestis consists of about 160 species that are among the most colorful beetles of the family Buprestidae (Plate 10). Species occur in Asia, northern Africa, Europe and North America. Most breed in the wood of dead trees and are of minor economic consequence. Some have long life cycles and larvae can remain in dead wood for many years. At least one species, B. aurulenta, the golden buprestid, is known to be a pest of wood in use (see Chapter 15).

Buprestis apricans Herbst, Turpentine Borer Distribution Turpentine borer occurs along the Atlantic and Gulf coastal plains of southeastern USA.

Hosts Hosts are pines, Pinus spp., including the southern yellow pines, P. echinata, P. elliotii, P. palustris, P. rigida and P. taeda.

Importance This insect was once considered the most destructive insect in the turpentine orchards of southeastern USA. Trees attacked by this beetle were weakened and became subject to wind breakage. Lumber value was destroyed and resin production was reduced. In North America, the labor-intensive method of harvesting turpentine by hand has been replaced by other technologies, therefore this insect is no longer considered a pest.

Life History Adults emerge in February or March and feed for a short time on pine needles. Females deposit eggs on exposed wood, especially at the edge of turpentine faces and fire scars. The larvae tunnel in

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the sapwood and construct oval, twisting tunnels filled with resinous frass. Mature larvae construct cells in which they pupate. About 3 years are spent in the larval stage.

Epistomentis pictus Gory

Description of Stages Adults are gray-bronze with a green metallic luster and are about 25 mm long. Their overall shape is elliptical and somewhat flattened. Each elytron bears eight rows of large punctures (Drooz 1985).

Hosts Hosts are species of southern beeches, Nothofagus spp., including N. alpina, N. dombeyi, N. obliqua and N. pumilio.

Buprestis novemmaculata Linnaeus Distribution This species occurs in north Africa, Europe and Asian portions of Russia east to Siberia and Kazakhstan. It is considered rare or endangered in Austria, Germany, and Finland and has been introduced and become established in Chile and New Zealand.

Hosts Pines, Pinus spp., and other conifers are hosts. In Chile, it attacks Pinus radiata.

Importance Limbs, logging residues, stumps or logs that have been dead for at least 1 year are infested. It is not known to cause tree mortality. There is some evidence that severely stressed or fire-damaged pines may be attacked. In Chile, it behaves in much the same manner as it does in its natural range and is not considered a pest.

Life History Life history and habits of B. novemmaculata are poorly understood. Early-instar larvae feed on the inner phloem, while later instars bore into and overwinter in the heartwood. In Europe, 2–4 years are required to complete a generation.

Description of Stages Adults are flattened and metallic brown-black. The area between the eyes and the edges of the pronotum are marked with yelloworange bands. Elytra are deeply grooved and marked with four pairs of yellow-orange spots. Overall length is 13–20 mm. Males are smaller than females (Duran 1963, Francke-Grosman 1963, Billings & Holsten 1969a,b, Eglitis & Holsten 1972, Gara et al. 1980).

Distribution E. pictus is the single representative of a South American genus of Buprestidae and is known from central Chile and Neuquen Province, Argentina.

Importance This is a relatively common insect. During summer, adults are often seen on bark of freshly cut wood, especially wood exposed to direct sunlight. Attacks usually occur in severely stressed, dying or freshly cut trees but during dry periods, vigorous trees can be attacked. Life History Larvae construct galleries in the inner bark and, occasionally, in the sapwood.

Description of Stages Adults are 20–25 mm long. Legs, antennae and body are glossy black. The prothorax has three yellow longitudinal bands and elytra are metallic yellow to orange or bronze and edged with black (Fig. 10.2, Aguayo Silva et al. 2008). Melanophila Species of Melanophila occur across North America and Eurasia. Some species attack and kill trees weakened by drought, air pollution, smog, fire or other injury and are often associated with bark beetles and other wood borers. Some have the ability to detect infrared radiation and attack trees almost immediately after they are fire scorched. Adults are known to bite firefighters and lay eggs on scorched trees while they are still smoldering (Furniss & Carolin 1977, Evans 2005). Melanophila acuminata DeGeer, Black Fire Beetle Distribution Black fire beetle has a holarctic distribution across North America, northern Africa and Eurasia.

Hosts Hosts are conifers including species of Abies, Picea and Pinus.

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Importance Trees weakened by drought or insects, such as hemlock woolly adelgid, Adelges tsugae, are attacked and killed.

Life History A generation is completed in 1 year in dead trees and logs but more than 1 year may be required in living trees. Adults are active from late spring to summer and deposit eggs in groups deep in bark crevasses of weakened, recently dead and dying trees. Larvae construct winding, frass-filled galleries. Later, they construct pupal cells in the outer bark and overwinter.

Description of Stages Adults are black, with a metallic sheen and about 10 mm long. Each elytron has three orange or yellow spots (Drooz 1985, Reardon & Onken 2004).

Melanophila guttulata Gebler, Larch Buprestid Distribution Larch buprestid occurs from the eastern half of European Russia to the Russian Far East. Specimens have been intercepted in Finland in pine imported from Russia. Fig. 10.2 Epistomentis pictus adult on recently cut Nothofagus dombeyi fuel wood (near Lonquimay, IX Region, Chile).

Importance Black fire beetle is a common, but not particularly damaging, species. Life History This species is attracted to, and breeds in, fire-scorched conifers using an infrared heat sensing system. Description of Stages Adults are about 10 mm long and blue-black in color (Drooz 1985, Evans 2005). Melanophila fulvoguttata (Harris), Hemlock Borer Distribution Hemlock borer is found throughout eastern Canada and the USA. Hosts Its favorite host is eastern hemlock, Tsuga canadensis, but it will also attack Larix laricina, Picea spp. and Pinus strobus.

Hosts Primary hosts are Larix spp., including L. gmelinii and L. sibirica. Species of Abies, Picea and Pinus are also attacked.

Importance This insect is one of the most important pests of larch across the Russian boreal forests. It attacks trees weakened by defoliation, fire, or recently cut logs. Attacks occur along the entre length of the bole from root collar to upper crown and may take place over several years on the same tree, ultimately causing tree death.

Life History One to two years are required to complete a generation. Larvae feed in the cambium and wood. In more vigorous trees, they feed mainly in the bark and cambium, whereas in stressed trees, they tend to feed in the wood at a depth of 0.5–1.5 mm. Galleries are winding and frass filled. When larvae are mature, galleries may be up to 15 mm wide.

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Description of Stages Mature larvae are pale yellow to white, 16–18 mm long, with black mandibles. Adults have an elongated bronzed black body, 7–12 mm long. The elytra have three pairs of yellow spots (Vorontsov 1995).

Phaenops gentilis (LeConte), attacks pines, Pinus jeffreyi, P. lambertiana and P. ponderosa in western USA. Adults are iridescent blue-green and lack markings on the elytra. P. californica also invades stressed pines in western USA (Furniss & Carolin 1977, Kolk & Starzyk 1996, Evans et al. 2004).

Phaenops cyanea (Fabricius) Distribution North Africa.

This borer is indigenous to Europe and

Cerambycidae (Round Headed Wood Borers) Anoplophora

Hosts Hosts are pines, Pinus spp., especially P. sylvestris. This insect is found less frequently in Abies alba, Larix decidua and Picea abies. Importance Phaenops cyanea is considered a serious pest of pines, especially during warm, dry years. Trees with thick bark, exposed to direct sunlight along forest edges, are preferred. This insect was first reported as a pest in Germany from 1969 to 1971, when it was involved in a decline of P. sylvestris. The primary cause of the decline was root disease caused by an abnormally high water table. It is also a pest in Poland, especially in older stands weakened by other biotic agents including pollution, fire or moisture deficit. In Finland, Norway and Sweden, seed trees exposed after cutting are attacked during hot, dry summers. Life History There is one generation/year. Adults feed on foliage of host trees prior to mating. Females lay single eggs under bark scales. Newly hatched larvae feed in the inner bark and cambium. They construct pupal cells, overwinter and adults emerge in spring and early summer through oval exit holes. Description of Stages Adults are 8–11 mm long with an oval, flattened body and are dark metallic blue to green in color. Related Species Several species of Phaenops invade conifers in North America. These include the flat headed fir borer, P. drummondii (Kirby), which attacks weakened and stresses conifers (Abies, Larix, Picea, Pseudotsuga menziesii and Tsuga heterophylla) in much of the western provinces of Canada and western USA. Adults are bronze-black and usually have three small yellow spots on the elytra. Flat headed pine borer,

Anoplophora consists of 36 known species distributed across Asia. Adults are striking, with colorful patterns on the elytra and other body parts. These insects are well known in Asian culture and have colorful common names such as “sky beetle” and “starry night sky beetle.” They are a popular subject for wood and ivory carvings. Several species are important pests of broadleaf trees and two have been introduced into North America and Europe (Lingafelter & Hoebeke 2002). Anoplophora glabripennis (Motschulsky), Asian Longhorn Beetle Distribution This species is widely distributed in China, north to the area of Beijing, west through Gansu Province and south into Sichuan Province. It also occurs in North and South Korea. This longhorn beetle has been an insect of international concern since 1996 when infestations were discovered in Brooklyn, New York, USA. Later that year, another infestation was detected in nearby Amityville, New York. The initial method of introduction is believed to have been infested wooden crating with subsequent spread via transport of infested fuel wood. Infestations have since been detected in several other locations in Europe and North America (Table 10.2).

Hosts This insect breeds in temperate broadleaf trees and has a wide host range. In China it prefers species of Acer, Populus and Salix. Other hosts include Melia azedarach, Morus spp., Prunus spp., Pyrus spp., Robinia pseudoacacia and Ulmus spp.

Importance Asian longhorn beetle can invade either vigorous or stressed trees and recently cut logs. Since the 1970s, it has been a major pest of hybrid poplar

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Table 10.2 Worldwide introductions of Asian longhorn beetle, Anoplophora planipennis (Coleoptera: Cerambycidae), 1996–2009. Year

Country

Location

1996 1998 2001 2002 2003

USA USA Austria USA Canada France France Germany USA USA USA Italy USA USA

Brooklyn and Amityville (New York) Chicago (Illinois) Braunau Jersey City (New Jersey) Toronto, Woodbridge (Ontario) Gien (Loiret) Sainte Anne sur Brivet (Loire) Neukirchen Am Inn (Bavaria) Cartaret (New Jersey) Sacramento (California) Linden (New Jersey) Corbetta (Milano) Staten Island and Prall Islands (New York) Worcester (Massachusetts)

2004

2005 2006 2007 2008 Source: Smith et al. 2009.

plantations in northern China, where thousands of trees have been killed. Larvae first attack and kill branches and, within a few years, the entire tree dies. Introduced populations have, to date, been confined to shade and ornamental trees. A major concern in northeastern USA and Canada is its potential for spread into native broadleaf forests with a high component of Acer saccharum and other suitable hosts.

Life History In China, there is normally one generation/year although some individuals may require 2 years. Adults fly from April or May through October with peak activity in July. Newly emerged adults feed on bark of twigs and then mate on trunks and branches. Females chew through the bark to the cambium and usually lay one egg per oviposition site. Most females lay from 25 to 40 eggs, which hatch within 1–2 weeks. Larvae feed first in the cambium and later in the wood, tunneling upward for 10–30 cm through both sapwood and heartwood. They transform to pupae and then adults inside larval galleries in early summer. The new generation of adults exits through 6–18 mm diameter holes that they chew through the bark.

Description of Stages Adults are 25–30 mm long, shiny black with irregular white spots. Antennae are

long with alternating black, white and blue bands (Fig 10.3).

Pest Management In China, infested trees are removed and destroyed. In addition, infested trees are climbed by forest workers during beetle flight and adults are collected and killed by dropping them in a jar of kerosene. In North America, established infestations are treated by removal and destruction of infested and high-risk trees and establishment of quarantine areas to limit infestation spread. As of December 2007, nearly 70,000 infested and high-risk trees had been cut and destroyed, all in urban areas (Li & Wu 1993, Schmutzenhofer et al. 1996, Haack et al. 1997, Canadian Food Inspection Agency/Canadian Forest Service 1998, Lingafelter & Hoebeke 2002, New Jersey Department of Environmental Protection 2002, Tomiczek 2002, Smith et al. 2009).

Anoplophora chinensis (Forster), White Spotted Citrus Longhorned Beetle (¼ A. malasiaca Thomson) Distribution A. chinensis is related to Asian longhorned beetle and occurs primarily in China, Japan and Korea. A few specimens have been reported from

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Fig. 10.3 Asian longhorn beetle, Anoplophora glabripennis, adult (photo by Larry Barber, USDA Forest Service, courtesy of www. forestryimages.org).

Indonesia, Malaysia, the Philippines, Taiwan and Vietnam. This insect has also become a world traveler and has been intercepted at ports of entry. Pathways of introduction include infested nursery stock, especially bonsai. In 2000, an infestation was detected in northern Italy, in a nursery located between the provinces of Milan and Varese. The insect was found in 13 villages, all relatively close to the first point of detection, covering an area of approximately 60 km2. In 2001, a localized infestation was detected near Tukwila, Washington, USA.

Hosts A. chinensis can infest trees important in agriculture, arboriculture and forestry. More than 100 plants have been reported as hosts including Broussonetia papyrifera, Cajanus cajan, Camellia sinensis, Carya illinoensis, Casuarina equisetifolia, C. stricta, Citrus spp., Cryptomeria japonica, Ficus spp., Fortunella

margarita, Hibiscus spp., Juglans regia, Litchi sinensis, Malus pumila, M. spectabilis, Melia azedarach, Morus alba, Populus spp., Platanus spp., Prunus spp., Pyrus communis, Salix spp. and Ziziphus jujuba.

Importance Boring by larvae can kill trees. Moreover, heavily mined branches and stems commonly break, especially during strong winds. This insect is transported via live plant materials or wood in use.

Life History A. chinensis has one generation/year although some individuals may require 2 years to complete development, especially in the northern part of its range. Adults emerge from April to August and are most common in May–July. They are strong fliers but females, heavily laden with eggs, prefer to oviposit either on trees from which they emerged or within

Large cambium and wood boring insects close proximity. Adults feed on the bark, leaves, and leaf petioles of host trees. Females deposit eggs in T-shaped oviposition holes cut in the bark on the lower trunk or exposed roots of host trees. Larvae initially feed in the sapwood and third instar larvae bore deep into the woody tissue. They are present throughout the year. Pupation and adult development takes place in the wood.

Description of Stages Adults are attractive beetles, 25–40 mm long, shiny black in color with a series of white markings on the thorax and elytra. The antennae are marked with alternating black and white segments and are at least as long as the body. The ventral surface is covered with white or blue pubescence.

Pest Management In places where A. chinensis has been introduced and become established, infested trees or trees suspected of being infested have been removed and destroyed (Colombo & Limonta 2001, Washington State Department of Agriculture 2001, Lingafelter & Hoebeke 2002).

Holopterus chilensis Blanchard Distribution This cerambycid is found in temperate forests of Argentina and Chile.

Hosts H. chilensis is known to attack two trees of the genus Nothofagus: N. dombeyi and N. obliqua.

Importance In central and southern Chile, larvae bore in vigorous Nothofagus obliqua. It is rarely found in N. dombeyi. Boring does not kill trees but causes loss of quality and structural integrity of lumber. In a study conducted in the X Region of Chile (Valdivia), between 38% and 46% of N. obliqua examined in three subregions, Coast range, Central Valley and Andean Cordillera, had been attacked.

Life History Relatively little is known about the biology of H. chilensis. This insect attacks the base of the bole of living trees or freshly cut logs. Adults are present throughout the growing season and fresh cut

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logs or trees are subject to attack throughout this period. Larvae bore deep into the wood near the base of the tree, sometimes working their way into the root system.

Description of Stages Adults have a slender body. Males have antennae about 1.2 times the length of the body and females have antennae roughly equal to their body length. Body color is medium red-brown, roughly the color of cured tobacco leaves. Total body length averages 4 cm for males and 5 cm for females (Cameron & Peña 1982, Baldini et al. 1994, Aguilar et al. 1998). Monochamus Monochamus consists of about 150 species distributed across Asia, Africa, Europe and North America (Tables 10.3 & 10.4). Larvae are known as “sawyers” or “sawyer beetles” because they make a loud noise when feeding. Hosts of most boreal and temperate species of Monochamus include conifers of the genera Abies, Larix, Picea, Pinus and Pseudotsuga Several African species breed in broadleaf trees. Most are secondary and breed in trees stressed or recently killed by bark beetles, lightning, disease, climatic- or site-related factors. Upon emergence, adults feed on branches, twigs and foliage of both host and non-host trees. Species that bore deep into the wood reduce lumber quality. If fuelwood is stored inside homes, chewing sounds associated with feeding larvae can be annoying and emerging adults have been known to frighten people. Larvae bore in the cambium and some species bore deep into the wood and cause significant loss of wood quality and structural integrity (Fig. 10.4). Sawyer beetles are also vectors of the pinewood nematode, Bursaphelenchus xylophilus. This nematode is native to North America and causes pine wilt disease in places where it has been introduced. Pine wood nematode has caused extensive mortality of pines in China, Japan, Korea, and, more recently, Portugal. Monochamus larvae become infested while feeding in nematode-infested trees. The nematodes are then spread by adults when they feed on twigs and branches of pines. Nematode “dauerlarvae” emerge from spiracles of adult beetles during feeding, drop on to twigs and penetrate the woody tissue via feeding wounds. Adults are striking longhorn beetles. Overall length ranges from 18 to 30 mm. Males tend to be smaller and

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Table 10.3 Representative species of Monochamus (Coleoptera: Cerambycidae) indigenous to Asia, Europe and Africa. Species

Distribution

Principal hosts

M. alternatus (Hope) Japanese pine sawyer M. bimaculatus Gahan

China, Japan, Korea, Laos, Taiwan, Vietnam China

M. galloprovincialis (Oliver) M. grandis Waterhouse M. griseoplagiatus Thomson

Eurasia and northern Africa Japan, Russia (Kuril Islands) Africa: Uganda

M. guttatus Blessing

Northeast China, Korea, Russian Far East Northeastern China, Mongolia, North Korea, Russia (east of the Urals) Southern Africa

Preliminary Pinus but also found on Abies, Larix and Picea Cinnamomum camphora, Ficus spp., Mallotus philippensis Picea, Pinus Abies, Picea, Tsuga Celtis, Coffea, Cynometra, Khaya, Lachnophylis, Maesopsis eminii Acer, Alnus, Betula, Carpinus, Corylus, Juglans, Quercus, Salix, Ulmus Larix, Pinus sibirica

M. impluviatus Motsch M. leuconotus (Pascoe) White coffee stem borer M. nitens (Bat.) M. saltuarius (Gebler) M. sartor (Fabricius) M. sutor (Linnaeus) M. urossovi (Fisher) Fir sawyer beetle

Japan, eastern Russia Eurasia Eurasia Eurasia Northern China, Finland, northern Japan, Mongolia, North Korea, Russia

Coffea arabica and other broadleaf trees Abies, Larix, Picea Abies, Larix, Picea, Pinus Abies, Picea, Pinus Picea, Pinus Primarily Abies but also on Larix, Picea and Pinus

Sources: Kobayashi et al. 1984, Cherepanov 1991b, Baranchikov 1997, Hao Zheng et al. 2004, Schabel 2006, Waller et al. 2007.

Table 10.4 Representative species of Monochamus (Coleoptera: Cerambycidae) indigenous to North America. Species

Distribution

Principal hosts

M. carolinensis (Oliver) M. clamator (LeConte) M. marmorator Kirby Balsam fir sawyer M. mutator LeConte Spotted pine sawyer M. notatus Drury Northeastern pine sawyer M. obtusus Casey

Eastern USA Western North America, Mexico Eastern Canada and USA south to North Carolina Southern Canada, northern USA

Pinus Pinus, Pseudotsuga menziesii Abies balsamea, A. fraseri

M. scutellatus oregonensis (Say) Oregon fir sawyer M. scutellatus scutellatus (Say) White spotted sawyer M. titillator (Fabricius) Southern pine sawyer

Canada, northern USA

Pinus Abies, Picea, Pinus monticola, P. strobus, Abies

Canada: British Columbia USA: California, Oregon, Washington Western North America

Abies, Larix, Picea, Pinus, Pseudotsuga

North America

Abies, Larix, Picea, Pinus

Bahamas, eastern USA

Pinus

n Tovar et al. 1995, Linsley & Chemsak 1997. Sources: Safranyik & Raske 1970, Furniss & Carolin 1977, Drooz 1985, Cibria

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Fig. 10.4 Larvae of the sawyer beetle, Monochamus galloprovincialis, in Pinus pinaster (Peninsula de Trois, Portugal).

more slender than females and have antennae 2–2.5 times as long as the body. Females have antennae slightly longer than their body length. Body color ranges from red-brown to shiny black. The prothorax and elytra may have lighter markings. Some species have distinct spines on each side of the thorax. Eggs are white, elongated, slightly curved and rounded at both ends. The dimensions are 3 mm long by 1 mm wide. The chorion is a matte silver color, covered by small cells. Larvae of all species are white, legless grubs with an amber head capsule, conspicuous dark-colored mouthparts and distinct abdominal segments. When mature, larvae can be up to 60 mm long. Management of Monochamus should emphasize prevention. This includes not storing recently felled logs in forested areas during adult flight, prompt salvage and utilization of windthrow and/or trees killed by bark beetles, lightning or disease and storage of logs under a water spray at sawmills (Browne 1968, Furniss & Carolin 1977, Kobayashi et al. 1984, Drooz 1985, Speight & Wainhouse 1989, Cherepanov 1991, Mota et al. 1999, USDA APHIS 1999, Evans et al. 2004).

Monochamus alternatus (Hope), Japanese Pine Sawyer Distribution Japanese pine sawyer is indigenous to China, Japan, Korea, Laos and Taiwan.

Hosts Hosts are several Asian pines including Pinus densiflora and P. massoniana. Species of Abies, Picea and Larix may also be attacked.

Importance This insect is the primary vector of pinewood nematode in China, Japan and Korea. Adults are known to carry an average of 18,000 nematodes.

Life History M. alternatus has one generation/year and larvae overwinter in galleries in the wood of infested trees. Adults begin emergence in mid-April with peak activity in May. They feed on branches of pines and if infected by pine wood nematode they transmit the organism. Weakened trees, such as those defoliated by pine caterpillar, Dendrolimus punctatus, newly felled trees or trees dying from pine wilt disease are preferred for oviposition. Early-instar larvae feed in the cambium and instars III–V tunnel into the heartwood. Pupation occurs in late March.

Description of Stages Adults range in length from 15 to 28 mm and from 4.5 to 9.5 mm in width. The color of the body and elytra is orange to dark brown. The basal portion of the first to third antennal segment has gray hairs. The elytra have five longitudinal bands of black and gray rectangular spots (Plate 51, Kobayashi et al. 1984, USDA APHIS 1999).

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Monochamus scutellatus (Say), White Spotted Sawyer Distribution M. scutellatus is a complex of two subspecies. M. scutellatus scutellatus, the white spotted sawyer, is found across most of North America, where it is often the most frequently encountered species of Monochamus. M. scutellatus oregonensis LeConte, the Oregon fir sawyer, is slightly larger and occurs from California north to Alaska and east to Alberta, Canada, and Montana, USA.

Hosts Hosts are species of Abies, Picea, Pinus and Pseudotsuga.

Importance Larvae bore deep into the wood and degrade fire-scorched, injured, dying and recently felled trees. They also damage logs stored in the forest for prolonged periods.

Life History A generation can take 1–2 years to complete with 2 years required in colder climates. Adults emerge April–June. They feed on conifer foliage and twigs, until mating occurs. Females deposit eggs into slits they have chewed into the bark. Larvae emerge, feed in the cambium and wood and overwinter. They pupate near the bark surface and adults emerge in either 1 or 2 years.

host trees and renders them susceptible to breeding attacks. Trees weakened by insect defoliation or fire, recently dead trees and windthrow are also suitable breeding sites. When populations reach high levels, apparently healthy trees are attacked and killed. During the late 1950s, this insect, in association with M. sutor, killed 2 million m3 of timber in the Tomsk Oblast of western Siberia, resulted in the collapse of forest enterprises in the region and delayed the construction of a railroad. From 1971 to 1976, an outbreak damaged 300,000 ha of fir forests in the Krasnoyark Kray in central Siberia.

Life History M. urossovi requires 2 years to complete a generation. Adults fly from late May or early June through late September. Adults feed on branches of conifers, birch and other broadleaf trees. Adults introduce a blue stain fungus, Leptographium sibiricum, into trees during oviposition. Larvae feed in the phloem and sapwood during the first year and overwinter as instar II larvae. During the second year, larvae bore deeper into the wood and construct pupal cells near the surface where they spent their first winter. Pupation and adult emergence occurs during the following spring.

Monochamus urossovi (Fischer), Fir Sawyer

Description of Stages Adults are 18–37 mm long, with antennae 2–2.5 times the body length in males but only slightly longer than the body in females. The pronotum is as long as it is wide. The body and elytra are dark brown to black. The head and pronotum have sparse white or yellowish pubescence. Legs, elytra and antennae are all black with a very slight brass tinge. The elytra of female adults also have spots of white-gray hairs (Isaev et al. 1988, Cherepanov 1991, USDA Forest Service 1991, Baranchikov 1997, Jakobs et al. 2000, Evans et al. 2004).

Distribution M. urossovi occurs from Finland, east across Russia to the Far East.

Megacyllene

Description of Stages Adults are 18–27 mm long, shiny black with a distinct white spot on the scutellum (Plate 52, Furniss & Carolin 1977, Drooz, 1985).

Hosts This sawyer prefers to breed in Abies sibirica, but also attacks species of Larix, Picea and Pinus.

Importance This sawyer is one of the most destructive pests of fir forests in northern Asia. Adult feeding kills the distal portions of stems, reduces foliage area of

Megacyllene consists of about 50 nearctic and neotropical species. Adults are colorful with yellow, orange or black patterns on the thorax and elytra. Several are important pests. One species, M. mellyi (Chevrolat), native to South America, has shown promise as a biological control agent of groundsel, Baccharis spp., in Australia (Table 10.5, McFayden 1983, Parsons & Cuthbertson 1992, Monne & Hovore 2005).

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Table 10.5 Representative species of Megacyllene (Coleoptera: Cerambycidae): their distribution and hosts. Species

Distribution

Hosts

M. antennata (White) M. caryae (Gahan) Painted hickory borer

North America, southwestern USA North America, eastern USA, northern Mexico

Prosopis Carya spp., also Celtis, Fraxinus, Gleditsia triacanthos, Juglans, Morus, Quercus

M. mellyi (Chevrolat)

South America: Argentina, Bolivia, Brazil, Paraguay and Uruguay North America, southern Canada, USA

Baccharis spp.

M. robiniae (Forster) Locust borer

Robinia pseudoacacia, R. neomexicana

Sources: Furniss & Carolin 1977, Drooz 1985, Di Iorio 1995.

Megacyllene robiniae (Forster), Locust Borer Distribution Locust borer is found across the USA and adjoining portions of Canada. Its original range coincided with its host tree, which occurs in the Appalachian Mountains from Pennsylvania to Georgia and in the Ozark Mountains of Arkansas and Missouri. Black locust grows readily on poor sites. Its widespread use to reclaim land damaged by farming and strip mining, as a shade tree and in reforestation has dispersed the insect with its host. It now occurs from eastern Canada south to the Gulf States and west to Washington, Colorado and Arizona, USA.

and overwinter. During the following spring, they bore into the woody part of the tree and cause sap to ooze around small holes. By mid-July, most larvae have transformed into pupae. Mature beetles emerge through openings made by the larvae.

Description of Stages Adults are slender and about 20 mm long with red legs and black antennae. Bright yellow bands encircle their dark black body. A W-shaped band extends across the elytra (Plate 53, Galford 1984, Solomon 1995). Phoracantha

Hosts Black locust, Robinia pseudoacacia, is the most common host. New Mexico locust, R. neomexicana, a tree endemic to southwestern USA, is also attacked (Author’s observation).

Importance Larvae bore in boles and branches of live trees and make them susceptible to wind breakage. Damage from boring and subsequent breakage often results in deformed trees or clumps of sprout growth. Trees growing on poor sites, such as reclaimed strip mines, are especially susceptible to attack.

Life History There is one generation/year. Adults appear in late August–September and feed on flowers of goldenrod, Solidago spp. Females lay eggs under bark scales or in callus tissue around pruning wounds, in cracks in the bark and in other hiding places. Eggs hatch about 1 week later and small, white larvae bore into the inner bark. Larvae construct hibernation cells

Phoracantha consists of about 40 species associated primarily with Eucalyptus in Australia. The genus is well represented in southern Australia and some species occur in the north and in Papua New Guinea. Several are pests of eucalypts within their natural ranges. Others have survived overseas shipments of logs and subsequently become established in eucalypt plantations in Africa, Europe, North and South America. Adults are large, often more than 25 mm long and colorful (Wang 1995, Wang et al. 1999). Phoracantha (¼ Tryphocaria) acanthocera (Macleay), bull’s-eye Borer Distribution This insect occurs in both eastern and western Australia.

Hosts Both broadleaf and coniferous trees are hosts, including Agathis brownii, Araucaria cunninghamii,

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Corymbia calophylla, Eucalyptus diversicolor, E. ficifolia, E. gomphocephala, E. grandis, E. maculata, E. paniculata, E. propinqua, E. redunca, E. resinifera, E. saligna and E. triantha.

Importance This species breeds in living trees. Trees over 30 cm in diameter are preferred. A study in Western Australia indicates that attacks are more prevalent on poor sites and in large trees.

Life History Two years are required to complete a generation and adults emerge in mid-summer. Larvae bore in the sapwood. This borer is considered not capable of surviving long shipments in logs and is therefore at relatively low risk of introduction into other parts of the world.

Description of Stages Adults are 15–29 mm long, body and appendages are red-brown. The elytra are redbrown with yellow markings (Browne 1968, Wang et al. 1999, Farr et al. 2000).

Phoracantha semipunctata (Fabricius), Eucalyptus Longhorn Borer Distribution Eucalyptus longhorn borer is native to Australia. It has been introduced into virtually all Eucalyptus growing regions of the world (Table 10.6).

Hosts

This insect breeds in many species of Eucalyptus.

Importance In Australia it is considered a minor pest that colonizes stressed or down trees. In places where it has been introduced it attacks moisture stressed, polesized and larger trees, causes tree mortality, loss of structural integrity and quality of timber.

Life History Number of generations/year depends on climate. In Portugal and Spain, there is usually one complete generation and a partial second. In warmer climates, average time from egg to adult is 3 months with a range of 2–15 months. Adults are nocturnal and rest under litter, loose bark or bark crevasses during daylight. They feed on flowers and breed in stressed trees or freshly cut logs. Mated females can lay up to 300 eggs in clusters. Eggs hatch within several days and larvae penetrate the bark and begin feeding in the cambium. Mature larvae enter sapwood to pupate.

Description of Stages Males are 15–29 mm and females 19–29 mm long. The head and pronotum are dark red-brown. Males have antennae almost twice as long as the body and females have antennae only slightly longer than the body. The elytra are mostly dark with a zigzag band of yellow or cream about midway and densely punctated.

Pest Management Classic biological control has met with some success where this insect has been introduced. Biological control agents for Phoracantha species include the Australian parasitic wasps Avetianella longoi, Callibracon limbatus, Jarra maculipennis, J. phoracantha; and Helcostizus rufiscutum and Syngaster

Table 10.6 Global distribution of Phoracantha recurva and P. semipunctata (Coleoptera: Cerambycidae). Region

P. recurva

P. semipunctata

Africa

Malawi, Morocco, South Africa, Tunisia, Zambia

Europe Near East North America South America Oceana

Greece, Spain

Algeria, Egypt, Ethiopia, Lesotho, Malawi, Mauritius, Morocco, Mozambique, South Africa, Swaziland, Tunisia, Zambia, Zimbabwe France, Italy, Portugal, Spain Cyprus, Israel, Lebanon, Turkey USA (California) Argentina, Brazil. Chile, Peru, Uruguay Australia, New Zealand, Papua New Guinea

USA: (California) Argentina, Brazil, Chile, Uruguay Australia, New Zealand, Papua New Guinea

 Native. Sources: Smith et al. 1992, FAO 2009b.

Large cambium and wood boring insects lepidus from California (Browne 1968, Duffy 1963, Pook & Forrester 1984, Wang 1995, Paine et al. 2000, Evans et al. 2004).

Phoracantha recurva Newman, Yellow Phoracantha Borer Distribution P. recurva is one of the most commonly occurring species of Phoracantha in Australia. Like P. semipunctata, it is a world traveler and now is established in many parts of the world where eucalypts are grown, including Mediterranean Europe, Africa, South America and California, USA (Table 10.6). In California, it has displaced P. semipunctata in areas where the two species occur, because P. recurva adults emerge several months earlier.

Hosts

Many species of Eucalyptus are hosts.

Importance In Australia, dead and dying trees are infested and populations seem to be held at non-damaging levels by natural enemies. In places where it has been introduced it attacks moisture stressed pole-sized and larger trees, causes tree mortality, loss of structural integrity and quality of lumber.

Life History Studies in California indicate that P. recurva has one generation and a partial second generation/year. Adults emerge from infested material from February to October.

Description of Stages Males range in length from 15 to 29 mm and females from 19 to 29 mm. The head and pronotum are dark red-brown. Males have antennae almost twice as long as the body and females have antennae slightly longer than the body. The elytra are light colored with dark bands at the hind end and are densely punctated (Plate 54).

Pest Management As in the case of P. semipunctata, classic biological control has met with some success in places where this insect has been introduced (Browne 1968, Wang 1995, Bybee et al. 2004).

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Saperda Saperda consists of about 133 species distributed across temperate and boreal forests of Eurasia and North America (Table 10.7). Larvae of most species bore in boles and stems of broadleaf trees. Some species produce galls in stems (see Chapter 12). A few attack living trees and others bore in dying and recently killed trees. Adults feed on leaves and tender bark. Several are pests and poplars and willows are the most common hosts. Saperda calcarata Say, Poplar Borer Distribution Poplar borer is found throughout North America.

Hosts Host trees are poplars, Populus spp., and occasionally willows, Salix spp. In southern USA, eastern cottonwood, P. deltoides, is the primary host. Further north, aspens, P. grandidentata and P. tremuloides, are attacked.

Importance Larvae bore in wood of living trees (Fig. 10.5). Trees with mechanical injury, including former attacks, are preferred. Infested trees have swollen scars on the bole and larger than normal branches. Each larva bores an exit hole through which frass is expelled and sap exudes. Wet areas around the holes blacken and have a varnished appearance. Small trees can be killed by girdling. Large trees are seldom killed but galleries weaken trees and make them subject to wind breakage. Holes made by larvae allow decay fungi to enter and further degrade lumber. This insect is a pest of plantations in southeastern USA and of natural forests and ornamental and shade trees throughout its range.

Life History Two years are normally required to complete a generation in the southern part of its range and 3–5 years in more northerly latitudes. Adults are active during summer and feed on bark of young twigs before laying eggs in small slits in the bark. Eggs are usually deposited on the middle third of the bole. Young larvae bore in the inner bark and later the sapwood where they overwinter. In spring they bore into the heartwood and feed until mature.

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Table 10.7 Representative species of Saperda (Coleoptera: Cerambycidae) indigenous to Eurasia and North America. Species

Distribution

Principal hosts

S. alberti Plavilstshikov

Northeast China, Japan, Korea, northern Mongolia, eastern Russia Northeast China, Japan, North Korea, eastern Russia Canada, USA

Populus, Salix

S. balsamifera Motschulski S. calcarata Say Poplar borer S. candida Fabricius Round headed apple tree borer S. carcharias (Linnaeus) Large poplar longhorn beetle S. inornata Say S. interrupta Gebler S. octomaculata Blessig S. perforata (Pallas) S. populnea (Linnaeus) Small poplar borer S. scalaris (Linnaeus) S. similis Laicharting S. tridentata Olivier Elm borer

Salix Populus, Salix

Eastern Canada and USA

Amelanchier, Crataegus, Malus, Sorbus

Europe, northeast China, Korea, Russia

Populus, Salix

Canada, northern USA Northeast China, Japan, Korea, Russia (Siberia) Northeast China, Japan, Korea, eastern Russia Europe, Russia (Siberia) Eurasia

Populus, Salix Abies, Picea, other conifers Ulmus Populus, Salix Populus, Salix

Europe, China, Japan, Korea, Russia Europe, Near East, northeast China, Korea, northern Mongolia, Russia Southeastern Canada, eastern USA

Betula Salix Ulmus

 Produces stem galls, see Chapter 12. Sources: Drooz 1985, Cherepanov 1991b, Schmutzenhofer et al. 1996, Kimoto & Duthie-Holt 2006.

Description of Stages Adults are 20–28 mm long, gray black or reddish brown in color and densely covered with fine gray and yellow hairs. There are yellowish stripes on the thorax and orange-yellow markings on the elytra. Mature larvae are about 30 mm long (Drooz 1985, Solomon 1995).

Saperda carcharias (Linnaeus), Large Poplar Longhorn Beetle Distribution This species is found over most of Europe, east across Russia to northeastern China and Korea.

Hosts Primary hosts are poplars, Populus spp. Birch, Betula spp., and willow, Salix spp., are also attacked.

Importance Larvae feed in inner bark, sapwood and heartwood. They attack the base of young trees and

occasionally large stems and branches. Larvae bore deep into the wood and heavy infestations can cause tree mortality or predispose trees to other pests.

Life History Two years are normally required to complete a generation. Adults are active from June to early October. They feed on leaves and young shoots and later deposit eggs on host trees.

Description of Stages Adults are 20–30 mm long, yellow-brown, gray or gray-yellow in color and have acutely pointed elytra. The head and pronotum have numerous small black punctures, and the elytra have larger punctures. Each elytron has a faint, indistinct, light transverse band below the mid-point. Antennae are yellow or gray with black apices and extend beyond the elytral apex in males, and do not reach the apex in females (Abgrall & Soutrenon 1991, Kimoto & DuthieHolt 2006).

Large cambium and wood boring insects

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Fig. 10.5 Galleries of the poplar borer, Saperda calcarata, in wood of quaking aspen, Populus tremuloides.

Tetropium Tetropium consists of about 30 species that infest conifers in Asia, Europe and North America. Adults have compound eyes divided into upper and lower lobes, giving the impression they have four eyes instead of two. Most confine their attacks to weakened, recently dead trees or fresh cut logs. However, several have caused severe losses within their natural ranges, usually in association with other bark beetles and woodborers. Larvae can survive long ocean voyages in logs and wood products and are considered a high risk for introduction and establishment into new habitats. One species, T. fuscum, has been introduced into eastern Canada, where it is causing damage to spruce forests in the vicinity of Halifax, Nova Scotia (Table 10.8, Knull 1946, EPPO 2005c, Canadian Food Inspection Agency 2006).

Hosts Conifers of the genera Abies, Larix, Picea and Pinus are hosts. Picea spp. is preferred in Europe whereas pines, Pinus spp., are preferred in Siberia. Importance This insect prefers freshly felled trees, stumps or trees weakened by drought, insects, fungi or air pollution for breeding sites. However, it can attack healthy trees. During the 1990s, it caused tree mortality and loss of lumber quality over some 180,000 ha of spruce forests in Romania and 225,000 m3 of timber resources were affected.

Tetropium castaneum (Linnaeus)

Life History Females lay eggs either singly or in clusters of up to 10 in bark crevices or under bark scales. Early-instar larvae feed on the inner bark and outer sapwood. They construct wide, irregular galleries filled with coarse, granular frass. Mature larvae bore 2–5 cm into the sapwood and form pupal chambers plugged with coarse frass. Adults emerge via oval exit holes, roughly 5 mm in diameter.

Distribution T. castaneum is found over much of Europe, across Russia and east to China, Japan, Korea and Mongolia.

Description of Stages Adults are flat, 8–18 mm long, with antennae that are half the body length.

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Table 10.8 Representative species of Tetropium (Coleoptera: Cerambycidae) indigenous to Eurasia and North America. Species

Hosts

Distribution

T. abietis Fall (round headed fir borer) T. castaneum (Linnaeus)

Abies

North America USA: California, Oregon, Washington Europe Asia: China, Japan, Korea, Mongolia

T. cinnamopterum Kirby Eastern larch borer T. fuscum (Fabricius) Brown spruce longhorn beetle T. gabrieli Weiss Larch longhorn beetle) T. gracilicorne Reitter

Abies, Larix, Picea (preferred in Europe), Pinus (preferred in Siberia) Picea glauca (preferred in western North America) Abies, Larix, Pinus Picea Larix Abies, Larix, Picea, Pinus

T. staudingeri Pic Seven-river spruce borer T. parvulum Casey Northern spruce borer

Picea schrenkiana

T. velutinum LeConte Western larch borer

Abies, Larix occidentalis, Picea, Pinus, Pseudotsuga menziesii, Tsuga heterophylla

Picea engelmannii, P. glauca

North America: transcontinental distribution Central Europe Eastern Canada (introduced) Europe Asia: China, Japan, Kazakhstan, Mongolia, Russia (Siberia, Transbaikalia, Far East). Asia: China, Kazakhstan, Kyrgyzstan, Uzbekistan North America Canada: western provinces USA: Alaska North America Canada: British Columbia USA: California, Montana, Utah

Sources: Furniss & Carolin 1977, Evans et al. 2004, EPPO 2005c, Canadian Food Inspection Agency 2006.

They typically have a black body, brown elytra, and either brown or red antennae and legs. The shiny pronotum is rarely punctured. The elytra are covered with fine hairs and are densely punctured (Evans et al. 2004, Canadian Food Inspection Agency 2006).

Tetropium fuscum Fabricius, Brown Spruce Longhorn Beetle Distribution A Eurasian species, T. fuscum, occurs throughout Europe, as far north as Lapland. It is also reported from Japan and Turkey but is reported as “uncommon” in western Siberia. It was introduced into eastern Canada at Halifax, Nova Scotia around 1990, where it became established. Hosts Within its natural range, its favorite hosts are Abies alba, Picea abies, P. pungens P. sitchensis and Pinus sylvestris. Species of Larix are occasionally attacked. In Canada, the favorite host is Picea rubens but infestations also occur in P. abies, P. glauca and P. mariana.

Importance T. fuscum infests dead and dying trees within its native range. In Canada, it has caused extensive mortality of Picea rubens.

Life History One or two years may be required to complete a generation. Adults are active from June to August. They do not feed. Females deposit eggs singly into bark crevices of suitable host trees. Larvae hatch within 10–14 days and feed under the bark and sapwood, making irregular galleries up to 2 cm wide. The galleries are filled first with brown shredded bark and later with white shredded wood. Larvae molt four times and in autumn bore tunnels in the wood to a depth of 2–5 cm, where they pupate. Pupation occurs from early May through late June. Adults emerge through oval hole, about 7 mm in diameter. Description of Stages Adults are black to dark brown, with a flattened body that varies in length between 8 and 17 mm. The elytra range from brown to red or yellow-brown or straw yellow and bear two to

Large cambium and wood boring insects three distinct longitudinal stripes. A broad white to beige pubescent band is present at the base of the elytra. Short gray-yellow, densely packed hairs cover the first quarter of the elytra. The antennae are red-brown and legs are dark brown. A deep groove is found on the head between the antennae. Fine short hairs cover the body and variations in color and size are common (Juutinen 1955, Cherepanov 1988, Smith & Humble 2000, Smith & Hurley 2000). Tetropium gracilicorne Reitter Distribution T. gracilicorne is an Asian species found across China, northern Japan, Kazakhstan, northern Mongolia and Asian Russia.

Hosts

Hosts are species of Abies, Larix, Picea and Pinus.

Importance This species is an important pest of conifers in its natural range. It attacks trees stressed by insect defoliation, primarily outbreaks of Siberian silk moth, Dendrolimus sibiricus, fire, disease and wind. It is often associated with other bark beetles and woodborers.

Hosts

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The only recorded host is Picea schrenkiana.

Importance T. staudingeri can attack stressed or healthy trees of various ages and subsequent generations may damage the same trees for several consecutive years, ultimately causing tree death. It prefers mature trees and, even in cases where trees are not killed, infestations cause significant loss of tree vigor and wood quality. T. staudingeri occurs in mountain forests, which are important for soil protection against erosion. It is often associated with other bark beetles and woodborers. Life History Mass flight occurs from May to June in the southern portions of its range and at low elevations, and in early July in northern parts of its range and at high altitudes. Eggs are laid in bark cracks of stressed, dying and recently cut trees and stumps or on the lower sides of fresh cut logs. Larvae make large irregular feeding galleries in the sapwood and overwinter for either one or two seasons (EPPO 2005c). Curculionidae (Weevils) Rhyephenes, Spider Weevils

Life History Adults are active from early June to late July. Females lay eggs in clusters of three to five in cracks in the bark of susceptible trees. Young larvae construct irregular feeding galleries in the phloem and later bore into sapwood. In August, larvae may reach a depth of 4–5 cm, where they overwinter. Some larvae overwinter under the bark of thick barked trees. Pupation occurs from May to June.

Description of Stages Adults are flattened, about 10 mm long with a black body. The elytra are light brown and the antenna and legs are red brown. Some individuals may be entirely black (EPPO 2005c, Canadian Food Inspection Agency 2006).

Rhyephenes is a genus of seven species, all of which are found in Chile and neighboring areas of Argentina. Adults are hard-shelled insects with long legs that make them resemble a spider. Rhyephenes mallei (Gay & Solier), Rhyephenes humeralis (Gu erin) Distribution R. maillei occurs in central and southern Chile (V Region south to the X Region). R. humeralis is also found from Chile’s V Region south to the X Region and in Chubut, Neuquen and Santa Cruz Provinces, Argentina.

Tetropium staudingeri Pic, Seven-River Spruce Borer

Hosts Indigenous host plants for both R. maillei and R. humeralis include Cryptocarya alba, Nothofagus dombeyi and Quillaja saponaria. R. maillei also attacks Pinus radiata.

Distribution Seven-river spruce borer is another Asian species found in northwestern China, Kazakhstan, Kyrgyzstan and Uzbekistan.

Importance These insects have habits similar to secondary bark beetles and breed in trees stressed by

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competition, water or nutrient deficiencies, mechanical injury or fire. Recently harvested trees or logging residues may also become infested. They are of minor economic importance and are not known to cause tree mortality. In Chile, breeding attacks have been observed in young P. radiata with heavy foliar damage caused by the pathogen, Phytophthora pinifolia (Author’s observation). Life history Life histories of Rhyephenes spp. are not well known. Both adults and larvae are present throughout the year. Females bore small holes in the bark and deposit eggs inside. The larvae construct galleries between the bark and wood surface. Pupation occurs in chip cocoons beneath the bark. Adults emerge by carving exit holes through the bark and are often seen on the foliage of host trees. They have also been observed on logging residues of P. radiata.

Description of stages Body color of adults is blueblack. R. humeralis has distinct white markings on the outer apical portions of the elytra. Body length ranges from 7 to 15 mm. Larvae are legless grubs, yellow-white in color with a light brown head capsule. Mature larvae are 13–15 mm long (Morrone 1996, Aguayo Silva et al. 2008).

LEPIDOPTERA

Life History Two to four years are required to complete a generation. Adults are nocturnal and active in mid-summer. Females lay eggs in cracks and crevices in the bark of host trees or wounds near the base of the tree. Larvae first burrow irregularly between the bark and wood and later bore tunnels of about 10–15 mm in diameter in the wood. Larvae wander about in their tunnels and eject large amounts of foul smelling frass. Pupation may occur within the tunnel, near the entrance, in a silken cocoon, wood fragments and frass or at the base of the tree in an earthen cocoon.

Description of Stages Adults are gray-brown moths with a wing span of about 90 mm and an abdomen with alternating rings of pale gray and brown. Larvae are 70–80 mm long when full grown, are purple above and yellow below, with a black head and thoracic shield (Browne 1968, Faccioli et al. 1993, Hajek & Bauer 2007).

Alcterogystia (¼ Cossus) cadambae (Moore), Teak Carpenterworm Distribution This species occurs in India in Karnatika, Kerala, Nadu and Tamil States.

Hosts Major host is teak, Tectona grandis. Other hosts are Diospyros melanoxylon, Grewia tiliaefolia and Terminalia bellerica.

Cossidae Cossus cossus Linnaeus, Goat Moth Distribution Cossus cossus is a complex of several subspecies widely distributed across Asia, North Africa and Europe.

Hosts Larvae bore in bark and stems of broadleaf trees, including species of Fagus, Fraxinus, Malus, Populus, Pyrus, Quercus, Salix and Ulmus.

Importance Larval boring weakens branches and stems and can lead to tree death. This insect is an important pest of fruit orchards in Italy. In Gansu Province, China, it is a pest of Salix spp.

Importance This insect is a pest of teak plantations. Larvae bore in branches and boles and cause dieback and deterioration of timber. Damage tends to be more severe in plantations in close proximity to areas of human habitation. These tend to be more subject to human-caused mechanical injury, which provides favorable sites for egg laying.

Life History Generations are continuous and overlapping. Adults are active throughout the year but there are two peaks of activity; May–June and September–October. Females lay eggs on the bark of the bole or branches of host trees. They prefer to lay eggs near sites of mechanical injury or previous infestation. Larvae hatch in about 20 days and feeding causes

Large cambium and wood boring insects girdling of branches and dieback. Duration of larval stage is slightly over 200 days. Pupation occurs in the soil at a depth of 3–4 cm and adults emerge about 11 days later (Mathew et al. 1990, Sudheendrakumar 1994).

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brown scales. Females are larger than males and have a wingspan of about 76 mm. Eggs are olive-brown and oblong to ovoid in shape. Mature larvae are 50–76 mm long, greenish-white and nearly devoid of hairs. The head capsule and legs are shiny brown (Hay & Morris 1970, Solomon 1995).

Prionoxystus robinae (Peck), Carpenterworm Distribution Carpenterworm occurs in southeastern Canada and most of the USA.

Hosts Many broadleaf trees are hosts. Oaks, Quercus spp., especially those of the red oak group, are preferred in eastern and southern USA. In the central states, green ash, Fraxinus pennsylvanica, is favored. Further west, in the Rocky Mountains, poplars, Populus spp., are most frequently attacked and in California, coast live oak, Quercus agrifolia and elms, Ulmus spp., are favorite hosts.

Importance Larvae bore in wood and leave oval, oblong or irregular holes surrounded by stains caused by fungi, which reduces quality and structural integrity of lumber.

Life History Depending on location, between 2 and 4 years are required to complete a generation. Adults are active from late May to mid-July. Life span of an individual moth is about 1 week. Females with a full complement of eggs are unable to fly. After mating, a female can lay 200–1000 eggs. Eggs are usually laid in clusters of two to six eggs each in bark crevices, under vines, lichen or near fresh wounds or wound scars on the boles and larger branches of host trees. Larvae hatch, bore through the bark and into the woody tissue where they construct feeding galleries that ultimately penetrate the heartwood. Completed larval galleries are 15–23 cm in length. Larvae keep their galleries free of frass and push it out of the original entrance hole. Mature larvae spin a loose cocoon in their gallery and pupate. Adults emerge through the larval entrance hole.

Description of Stages Adults are gray, stout-bodied moths. Body and wings are mottled with gray and

Xyleutes ceramica Walker, Teak Beehole Borer Distribution This borer is indigenous to southeastern Asia and the Pacific Islands from Myanmar east through Indonesia, Thailand, Papua New Guinea, the Philippines and the Solomon Islands.

Hosts Hosts include teak, Tectona grandis and Gmelina arborea.

Importance This insect is an important pest of teak plantations. Larvae bore into the wood and cause loss of timber quality and value. Infestation levels of 87–100% of trees in teak plantations have been reported with the level of infestation increasing with age. Another report indicates that 65 individual attacks in trees 60 years old are considered average. A study in Malaysia reports that the annual rate of attack is relatively low (average < 1%/year) but cumulative attacks over time can be up to 65%.

Life History One to two years are required to complete a generation. Light trap studies in Thailand established that adults are active from late February to late March, with a peak in early March. Adults are short lived and may produce up to 50,000 eggs, which are laid in rows in bark crevices. Eggs hatch in 10–20 days and larvae are dispersed on silken threads by wind. Larvae that land on a suitable host spin a protective silken web and chew into the bark and outer sapwood, then inward and upward forming a tunnel that may reach a length of 25 cm. Feeding occurs primarily on young bark that develops on the wound at the entrance to the tunnel. Pupation occurs in a silken cocoon at the upper end of the tunnel. Two to three weeks later, pupae wriggle their way to entrance holes and thrust their bodies partially out of the hole to facilitate adult emergence.

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Description of Stages Adults are large, narrowwinged brown moths with a wingspan ranging from 80 to160 mm. Mature larvae are 60–70 mm long and transversely banded with pink and white bands (Browne 1968, Supachote Eungwijarnpanya et al. 1994, Nair 2001, Ramsden et al. 2002, Gohtoh et al. 2003).

Zeuzera pyrina Linnaeus, Leopard Moth Distribution Leopard moth is indigenous to a large portion of Eurasia and has been introduced into northeastern USA. Infestations were first reported in New Jersey in 1882 and the insect is now known to occur from Massachusetts south to eastern Pennsylvania.

Description of Stages Adults are black and white spotted moths. Wings are semi-transparent, white with black spots. Wingspan of females ranges from 67 to 75 mm. Females are larger and more heavily bodied than males. The thorax is white with six large black spots and one small spot. The abdomen is white with dark cross-bands. Eggs are oval, salmon to orangeyellow in color and about 1.5 mm long. Mature larvae are about 50 mm long, pale yellow with a pink hue. The head, thoracic shield and anal plate are brown-black. The body is sparsely hairy with large, prominent tubercles on each segment (Browne 1968, Solomon 1995).

Sesiidae (Clearwing Moths) Podesesia syringae Harris, Lilac/Ash Borer

Hosts Virtually all broadleaf trees and shrubs are susceptible to attack. In Europe, leopard moth is a pest of fruit trees. It also attacks ornamental Aesculus hippocastanum and species of Castanea, Fraxinus, Juglans, Quercus and Ulmus. In New York, USA, Acer spp. and Ulmus spp. are preferred hosts.

Distribution This species is widely distributed across North America.

Importance Larvae bore in the cambium and wood of branches and boles. Trees of any size are susceptible to attack. Branches are often girdled and killed, resulting in dieback and wind breakage. Attacks on the bole cause degradation of lumber and create infection sites for decay fungi.

Importance This borer is considered a pest throughout its range. Infestation rates of 50% are common in shelterbelts of the Great Plains region of the USA. In the south, trees intended for wood products are degraded and reduced in value. Shade and ornamental trees may be scarred, seriously weakened or killed.

Life History Two years are required to complete a generation. Adults are active from May to September and live only long enough to mate and lay eggs. Females are heavily laden with eggs, seldom fly and lay eggs near where they emerged. Females deposit between 400 and 800 eggs, singly or in small clusters in bark crevices or beneath bark plates. Larvae hatch in about 10 days and bore into the wood, often entering at the nearest bud, twig or branch crotch. They bore into the pith of small stems and the heartwood of larger branches or boles of host trees and construct tunnels up to 12 mm in diameter and 5–15 cm long. Larvae can migrate to larger branches if needed. They spend two winters in the wood and pupate in small chambers near the bark. Pupation takes 4–6 weeks. Pupal cases remain in the exit hole after moths emerge.

Life History There is one generation/year. Adults emerge as early as February in Florida and as late as July further north. Pupal cases are usually present near the exit hole (Plate 55). Eggs are deposited singly or in small clusters in bark crevices and hatch in about 11 days. Young larvae mine in the phloem and cambium and later excavate tunnels 7.5–13.0 cm long in the wood.

Hosts Larvae are borers in boles of lilac, Syringa vulgaris, and ash, Fraxinus spp.

Description of Stages Wings and body are brownblack and legs are marked with black, orange and yellow. Wingspan is about 25 mm. Adults occur in two distinct color morphs, a black morph with a dark brown abdomen and a yellow morph with a light brown abdomen surrounded by yellow bands. These morphs

Large cambium and wood boring insects are geographically distinct. The black morph occurs in eastern North America whereas the yellow morph occurs in western North America. A wide hybridization zone exists in the Midwest and prairie regions. Larvae are white, except for an amber-colored head and thoracic shield, and range from 25 to 34 mm long at maturity.

Pest Management Natural enemies, wound prevention, brood tree removal, burlap trunk wraps and insecticide applications help reduce damage. Pheromone traps to monitor moth flights are helpful to time insecticide applications (Eichlin & Duckworth 1988, Solomon et al. 1993).

HYMENOPTERA

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has occurred in windbreak and shelterbelt plantings. Attacks are typically so heavy that a single poplar can produce 2000 brood adults. Therefore the wood is impossible to use for lumber or other wood products. Moreover, the rate of decay of infested wood is accelerated because of the action of symbiotic fungi associated with this insect. Another impact has been loss of poplar windbreak plantings around agricultural crops and fruit orchards. This exposes orchards to high winds and results in reduced crop yields.

Description of Stages Adult males are black, including antennae and legs. Wings are amber and darker than wings of females. Females are larger with a dark head and thorax. The abdomen has bands of alternating black and amber with a long ovipositor on the last abdominal segment.

Siricidae (Wood Wasps) Tremex fuscicornis (Fabricius) Distribution This species is native to temperate Asia and Europe. In December 1996, a local outbreak of T. fuscicornis was detected between Tamworth and Sydney, New South Wales. The infestation may have been present for 10–15 years prior to its discovery. Infestations were detected in central Chile in February 2000 (V Region and the Region Metropolitana), and was probably established at least 2 years prior to its discovery.

Hosts Hosts in its native range include species of Acer, Alnus, Betula, Carpinus, Celtis, Fagus, Juglans, Populus, Prunus, Quercus, Robinia pseudoacacia, Salix, Ulmus and Zelkova. In Chile, it has been reported from Acer negundo, Populus alba, P. deltoides, P. nigra and Robinia pseudoacacia.

Importance This insect is not a pest in its native range. In Chile, weakened or damaged trees or those that have recently been cut are preferred. However, apparently vigorous trees of some hosts, such as Acer negundo, are subject to attack. On vigorous trees, first attacks occur on branches. This causes dieback and weakening. Brood adults emerging from branches infest the bole. All trees attacked are killed. Extensive damage

Pest Management Direct control methods are not available for this insect. In Chile, an integrated approach using a combination of cutting and destroying infested trees plus introduction of natural enemies has been developed. Low-level populations can be detected by establishment of trap trees injected with a weak herbicide, which attract adults. Two parasitoids, Ibalia leucospoides and Megarhyssa percellus (Hymenoptera: Ichneumonidae) are under evaluation for biological control. M. percellus apparently arrived in Chile concurrently with T. fuscicornis.

Related Species Pigeon tremex, T. columba (Linnaeus), is a North American species found throughout eastern Canada and in the USA as far west as Colorado, Utah and Wyoming. It also invades broadleaf trees. Although this wood wasp is the most abundant in North America, it is not damaging. Adults lay eggs on dead or weakened trees. The associated wood decay fungus is Daedalia unicolor (Smith 1978, Drooz 1985, Baldini U. 2002, CSIRO 2002, Smith & Schiff 2002). Sirex Sirex is a large genus of conifer infesting wood wasps represented in forests of Eurasia and North America. In their natural ranges, they are not considered pests because they confine attacks to weakened and dying trees.

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Fig. 10.6 Worldwide distribution of the wood wasp, Sirex noctilio (Hymenoptera: Siricidae) (redrawn and updated from Ciesla 2003a).

Sirex noctilio Fabricius, Sirex Woodwasp Distribution Sirex woodwasp is native to north Africa, the Azores, Europe, the Near East, Mongolia and Siberia. This insect is easily transported to new locations via shipments of logs, lumber or wooden pallets and, once introduced into areas where suitable host trees occur, it becomes established. S. noctilio has been introduced and become a pest of conifer plantations in several countries of the southern hemisphere where plantations of North American conifers are established. It has also become established in northeastern USA (Fig 10.6, Table 10.9).

Hosts Hosts are conifers of the genus Pinus spp. and, occasionally, Abies, Larix and Picea.

Importance In its native range, S. noctilio confines its attacks to stressed and dying pines and is not a pest. Its associated symbiotic fungus is Amylostereum areolatum, which, in combination with mucus injected into trees

by female wasps during egg laying, is toxic to many species of pines native to North America. In places in the southern hemisphere where S. noctilio has become established, live conifers have been attacked and killed, for example, during an outbreak in Australia between 1987 and 1989, more than 5 million Pinus radiata were killed at a value of $US10–12 million. In Brazil, as of 1998, some 250,000 ha of loblolly, P. taeda, and slash pine, P. elliottii, plantations were known to be infested in the three southernmost states of Brazil (Paran a, Rio Grande do Sul and Santa Catarina) and the insect is spreading. When live trees are attacked, the first indication of infestation is appearance of white resin droplets and oviposition scars on the bark. Foliage of infested pines wilts and turns from green to yellow to reddish brown. A pine plantation infested by S. noctilio typically has a scattering of dead and dying trees (see Plate 6).

Life History S. noctilio can complete a generation in as little as 10 months but may require up to 2 years in cooler climates. Males emerge from infested trees before

Large cambium and wood boring insects

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Table 10.9 Worldwide introduction and establishment of the woodwasp, Sirex noctilio (Hymenoptera: Siricidae). Year

Country

Primary hosts

Early 1900s 1952 1961 1980

New Zealand Australia: Tasmania Australia: mainland Uruguay

1985 1988 1993 1994 2001 2002 2004

Argentina: Entre Rios Province Brazil Argentina: Rio Negro Province South Africa Chile USA: Indiana USA: New York

Pinus radiata P. radiata P. radiata P. elliottii, P. taeda, P. pinaster P. elliottii, P. taeda P. elliottii, P. taeda P. banksiana, P. contorta, P jeffreyi, P. ponderosa P. radiata P. radiata No host reported P. resinosa, P. strobus, P. sylvestris

Sources: Ciesla 2003a, USDA APHIS 2009.

females and may outnumber females by a ratio of 20 : 1. Unmated females lay eggs that produce only males. Mated females produce offspring of both sexes. Adult life span can be up to 12 days but a female that has deposited all of her eggs may live only 3–4 days. Females deposit from 20 to 500 eggs. They drill their ovipositors through the bark and into the outer sapwood and deposit eggs, mucus toxic to some trees and the fungus, A. areolatum. Larvae hatch as early as 9 days but may remain dormant for several months, especially in cooler climates. They feed in the wood and construct large galleries. Pupation takes in the galleries 16–21 days later (Plate 56) and when the female emerges from the pupal case she takes up spores of the symbiotic fungus and stores them in a special organ in her abdomen. Adults bore out of infested trees and leave a round exit hole (Fig. 10.7). They are strong fliers, capable of traveling several kilometers in search of host trees.

Description of Stages Adults are shiny dark blue or black. Males are smaller than females and the middle abdominal segments are orange. Legs are yellow-red with black tarsi. Hind legs of males are black. The antennae of both sexes are black (Plates 57 & 58).

Pest Management S. noctilio infestations in pine plantations in the southern hemisphere have been managed using a combination of tactics. These include: (i) use of trap trees injected with a weak herbicide to

Fig. 10.7 Round exit hole on loblolly pine, characteristic of emerging adults of the wood wasp, Sirex noctilio (Santa Catarina State, Brazil).

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attract low level populations and ensure early detection; (ii) aerial surveys to detect tree mortality; (iii) thinning to maintain stand vigor and reduce susceptibility to attack; (iv) injection of infested trees with the parasitic nematode, Deladenus siricidicola, which attacks the ovaries of female wood wasps and renders them sterile; and (v) introduction of several parasitic wasps including Ibalia leucospoides, Megarhyssa nortoni, Rhyssa hoferi, R. persuasoria and Schlettererius cinctipes (Zondag 1969, Haugen et al. 1990, Iede et al. 1998, Ciesla 2003a, Haugen & Hoebeke 2005, USDA APHIS 2009). Sirex juvencus Linnaeus Distribution This woodwasp occurs throughout conifer forests of the northern hemisphere. Several subspecies are recognized. S. juvencus juvencus is a European form. S. juvencus atricornis is reported from China, Japan and Mongolia and S. juvencus californicus is known from western USA. An undetermined subspecies occurs over much of Canada and the northeastern and midwestern USA. Packaging material infested with this species has been intercepted in Australia and New Zealand but infestations are not believed established.

Hosts Conifers of the family Pinaceae are hosts. In Asia and Europe, S. juvencus has been reported from Abies alba, A. borisii-regis, Abies cilicia, Abies nordmanniana (¼ A. bornmuelleriana), Abies sachalinensis, Larix decidua, Picea abies, P. jezoensis, P. orientalis and P. sitchensis (native to North America). Pinus nigra and Pinus radiata (native to North America) are hosts in southern Europe and the Near East. A. balsamea, A. lasiocarpa, Larix spp., Picea spp. and Pinus spp. are North American hosts.

Importance This woodwasp confines its attacks to weakened and dead trees or logs and is not considered a pest. The subspecies S. juvencus californicus attacks smog-weakened Pinus ponderosa in California.

Life History Adults begin emergence in July with peak emergence in September. They live an average of 6.9 days. Females insert their saw-tipped ovipositor into the wood and oviposition takes about 8 minutes. Eggs, mucus produced by females and fungal spores are injected into wood during oviposition. Two to three

additional drills may be made at the same site, or the wasp may seek another location. The drills without eggs tend to receive a larger complement of the mucus/ fungus mixture. The fungus associated with the European subspecies is Amylostereum areolatum, the same species associated with S. noctilio. A Canadian study indicates that the fungus Amylostereum (¼Stereum challetii is associated with S. juvencus in Canada. After hatching, larvae construct galleries in the wood. They develop slowly and up to 2 years may be required to complete larval development. After pupation, adults emerge from trees and leave a circular emergence hole. Since this species has a long larval period, adults can emerge from sawn lumber. Natural enemies of S. juvencus include the parasitoids Ibalia sp., Ibalia drewseni, I. leucospoides, I. ruficollis, Rhyssa hoferi, R. jozana and R. persuasoria (Stillwell 1966, Kirk 1975, Furniss & Carolin 1977, Kanamitsu 1978, Spradbery & Kirk 1978, Drooz 1985, Smith & Schiff 2002, Talbot 1977).

Urocerus The genus Urocerus contains the largest of the woodwasps. Adult females range from 27 to 35 mm in length. Species are indigenous to Asia, North Africa, Europe and North America. Urocerus gigas (Linnaeus) Distribution U. gigas is a complex of four subspecies found across conifer forests of Eurasia, north Africa and North America. U. gigas gigas (Linnaeus) occurs throughout Europe, northern Africa (Algeria and probably Morocco and Tunisia) and across Russia to Siberia and the Kamchatka Peninsula. It was probably introduced into the British Isles where it is now established. U. gigas orientalis Maa occurs in China, Japan, Korea and the Russian Far East and Kamchatka Peninsula. U. gigas tibetanus Maa occurs in Xizang Province, China (Tibet). U. gigas flavicornis is a North American subspecies reported from Canada and western USA U. gigas gigas has been introduced into Argentina, where it was discovered in the Provinces of Chebut, Rio Negro and Neuquen in 1993. It has also been introduced into Chile, where it is probably distributed throughout the area of North American conifer plantations.

Large cambium and wood boring insects Hosts Subspecies of the U. gigas complex infest conifers including Abies, Larix, Picea, Pinus and Pseudotsuga. Several dubious host records include species of Cedrus, Chamaecyparis, Fraxinus, Populus and Salix. In Argentina and Chile, where U. gigas gigas has been introduced, it attacks Pinus radiata. Importance In their natural ranges, all subspecies confine attacks to logs and severely weakened trees and are not considered pests. In Chile, U. gigas gigas has thus far behaved as it does in its natural range and has not been damaging. Life History Overall length of a generation is about 3 years. Adult flight is from mid-summer until autumn. Females lay eggs individually in holes pierced with their ovipositor in the wood of host trees. Individual females

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live for 2–4 weeks and can lay up to 350 eggs. Larvae hatch in about 4 weeks and bore into wood at a right angle to the egg tunnel. Later they penetrate deep into the wood. Eventually, they return to a point near the wood surface to pupate and emerge as adults. The symbiotic fungus associated with Urocerus gigas gigas is Amylostereum chailletii.

Description of Stages Adults are robust wasps, cylindrical, up to 35 mm in length. Males are smaller than females. Wings are transparent and amber colored. Abdomen is black with horizontal bands of yellow brown in the basal and posterior segments. The female’s ovipositor is up to 20 mm long (Browne 1968, Furniss & Carolin 1977, Smith 1978, USDA Forest Service 1993, Viitasaari 1984, Smith & Schiff 2002, Klasmer n.d.).

Chapter 11

Sucking Insects

INTRODUCTION Insects with mouthparts equipped for piercing plant tissue and sucking juices are included in the order Hemiptera. This diverse group contains many plant pests. All undergo simple metamorphosis (egg–nymph– adult) but tend to have complex life cycles. Some have alternate hosts and others have both sexual and asexual reproductive stages with winged and wingless forms. Many species occur in colonies and feeding causes wilting and drying of affected plant parts. This leads to growth loss, branch dieback and, in extreme cases, tree death. Aphids, psyllids, plant hoppers and other sucking insects are vectors of plant viruses, microplasmalike organisms (MLOs) or other pathogens (Markham 1988). Many sucking insects also secrete honeydew, which consists of plant sap fed upon by the insects and later excreted. When infestations are heavy, honeydew is produced in large enough quantities to “rain” from infested plants and cause objects such as cars parked under infested trees to become sticky. Ants feed on honeydew and some species have developed a symbiotic relationship with aphids and other sucking

insects. They protect colonies from natural enemies and move individuals to neighboring plants to start new colonies. Sucking insects tend to be small and difficult to detect before plant damage occurs. Therefore, they are easily transported to new locations when live plant materials are moved and, in the absence of natural enemies, have become invasive. Foliage and stem feeding sucking insects are described in this chapter. Those that produce galls are reviewed in Chapter 12 and species that feed on reproductive structures of trees and other woody plants are discussed in Chapter 14.

Psyllidae, Jumping Plant Lice Heteropsylla cubana Crawford, Leucaena Psyllid (Plate 59) Distribution Leucaena psyllid is native to Mexico, Central and northern South America. In 1983, an outbreak was discovered in Florida, USA and a year

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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later infestations were discovered in Hawaii. Between 1985 and 1992, the insect spread across the Pacific Islands, Asia and eastern Africa. In 1985, infestations were detected on Christmas Island, Cook Island, Fiji, Guam, New Caledonia, Okinawa, Papua New Guinea, the Philippines, Saipan, Solomon Islands, Tahiti and Tonga. The following year it was discovered in Cambodia, southern China, Hong Kong, Indonesia, Laos, Malaysia, southern Myanmar, Singapore and Vietnam. Infestations were found in the Andaman Islands, Bangladesh, upper Myanmar, Nepal and Sri Lanka in 1987. By 1991, it had spread to India, Mauritius and Reunion Island and in 1992 it had reached eastern Africa, including Burundi, Kenya, Tanzania and Uganda. Modes of dispersal and spread are not fully understood but are believed to have been in part via monsoon winds and in part via aircraft (Fig. 11.1).

Hosts Host plants are species of Leucaena (Family Leguminosae), including L. leucocephala, a fast-growing tree widely planted in tropical climates as an agroforestry species. This tree has many uses, including forage

for livestock, fuel wood and as a shade tree for agricultural crops such as cocoa, coffee and vanilla.

Importance Nymphs and adults feed on terminal shoots and suck plant juices. Young flowers may also become infested but less frequently than shoots. Within its natural range, the insect is of no concern and little was known about it until it began to appear in other locations. Feeding causes desiccation of shoots and growth loss. The insects also produce honeydew on which sooty mold develops. Heavy infestations can cause tree mortality. Rapid spread of leucaena psyllid across the Pacific into Asia and Africa had severe socioeconomic impacts in countries where leucaena was planted as a fast growing multi-purpose tree.

Life History Leucaena psyllid has several overlapping generations/year. Time from egg to adult ranges from 10 to 20 days. All life stages can be found at the same time. Early-instar nymphs feed in colonies on new shoots but later instars feed in a solitary manner. It has

Fig. 11.1 Spread of leucaena psyllid, Heteropsylla cubana, across the Pacific Islands, Asia, Indian Ocean and eastern Africa 1983–92.

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a seasonal abundance and populations peak during cooler temperatures. During warm weather, life stages may become hard to detect. In some locations, referred to as “ecological refuges”, populations remain at high levels throughout the year.

in crevices between axillary buds and leaf petioles. Nymphs feed on shoots and congregate at bases of leaflets. They envelop themselves in large quantities of white waxy secretions. All life stages are found together on shoots.

Description of Stages Nymphs and adults are small, maximum length of adults is about 2 mm. All life stages are orange.

Description of Stages Eggs are pale yellow or cream and spindle shaped. Early-instar nymphs are also pale yellow but become grey and brown as they grow. They produce white, woolly filaments and masses of powdery, waxy material on the leaves when they feed. Adults are winged and resemble miniature cicadas. They are about 3 mm long, brown with orange patches on the thorax but may appear white if covered with waxy secretions.

Pest Management Infestations are managed via planting of leucaena varieties resistant to, or tolerant of, psyllid feeding and classic biological control. Biological control agents include a predaceous beetle, Curinus coeruleus, and parasitoids Phyllaephagus vaseeni and Tamarixia leucaena. Chemical control is not practical, except for protection of nurseries and young seedlings (Banpot Napompeth & MacDicken 1990, Banpot Napompeth 1994). Ctenarytaina eucalypti (Maskell), Blue Gum Psyllid Distribution This insect is native to Australia (Australian Capital Territory, New South Wales, South Australia, Tasmania, Victoria and Western Australia) and has been introduced and established in many countries where eucalypts are grown, including Argentina, Brazil, Chile, Ireland, Mexico, Sri Lanka, the UK and USA (California).

Hosts Host plants are Eucalyptus globulus and E. pulverulenta.

Importance Nymphs and adults feed on sap of juvenile shoots and leaves on young trees. They are not often seen on older trees and have not been reported on mature foliage. If numbers are high, new shoots become distorted. In California and Ireland, blue gum psyllid has threatened plantings of E. pulverulenta, which is grown for foliage for cut flower arrangements. Life History This insect may have up to five generations/year and the period from egg to adult takes 18–22 days. Adults lay eggs on young, growing shoots of seedlings and small trees. Eggs may also be deposited

Pest Management Classic biological control using the parasitoid Psyllaephagus pilosus has been successful where this psyllid has been introduced. Related Species C. spatulata is also native to Australia and has been introduced into Mediterranean Europe, USA (California), South America (Brazil, Uruguay) and New Zealand. It feeds on several species of Eucalyptus (Phillips 1992, Dahlsten et al. 2002, Purvis et al. 2002, Queiroz Santana & Burkhardt 2007). Glycaspsis brimblecombei Moore, Red Gum Lerp Psyllid (Fig. 11.2) Distribution This psyllid is native to Australia (New South Wales, Northern Territory, Queensland, South Australia). In 1998, it was detected in California, USA and has since appeared in other countries where eucalypts are planted, including Brazil, Chile, Mauritius, Mexico, Portugal and Spain. Hosts G. brimblecombei prefers Eucalyptus camaldulensis but also attacks E. blakelyi, E. brassiana, E. bridgesiana, E. dealbata, E. diversicolor, E. globulus, E. lehmannii, E. mannifera, E. nicholii, E. nitens, E. rudis, E. sideroxylon and E. tereticornis. Importance Infested leaves are covered with small white cones (lerp), honeydew and black sooty molds and feeding causes premature leaf drop. Heavily

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Forest entomology: a global perspective Pest Management Biological control with the parasitoid Psyllaephagus bliteus has shown promise in several countries where this psyllid has been introduced (Dahlsten et al. 2002, Sookar et al. 2003, Queiroz Santana & Burkhardt 2007, Valente & Hodkinson 2009).

Aphididae, Aphids Cinara, Giant Conifer Aphids

Fig. 11.2 Red gum lerp psyllid, Glycaspsis brimblecombei, on foliage of Eucalyptus camaldulensis (VI Region, Chile).

infested leaves are noticeable because spots of white lerp encrust the surfaces. Falling leaves foul the ground beneath infested trees, vehicles parked under trees, and swimming pools. Premature leaf fall beneath trees and on rooftops of buildings poses a fire hazard. Defoliation weaken trees, causes branch dieback and tree death. In southern California, thousands of mature E. camaldulensis have been killed.

Life History In Australia, there are two to four generations/year. Nymphs construct a white conical cover of sugar or lerp, and feed concealed under the shelter. Adults live and hide on undersides of leaves.

Description of Stages Adults are about 3 mm long, pale green with areas of orange and yellow.

Cinara consists of about 200 known species, 150 of which are indigenous to North America, 30 to Europe and the Mediterranean region, and 20 to Asia. They are of minor consequence in their native habitats, although populations occasionally reach high levels. Several have been introduced and established in conifer plantations in the southern hemisphere where they have become damaging. Hosts are conifers of the families Cupressaceae and Pinaceae (Table 11.1). Giant conifer aphids feed in colonies on stems, produce copious quantities of honeydew and are often tended by ants. They have complex life cycles. Both males and females have winged (alate) and wingless (apterous) forms and females that reproduce either sexually or by parthenogenesis. Most have several generations/year. Eggs hatch in spring and produce an asexual or parthenogenic form prevalent throughout the growing season. Parthenogenic females give birth to live young. In temperate climates, a sexual stage appears in autumn and lays eggs, which overwinter. When introduced into warmer climates, parthenogenetic reproduction occurs throughout the year. Adults are 2.6–4.1 mm long, usually dark brown in color with dusty surfaces and waxy areas. Species are difficult to separate and the literature on this group is confusing. Therefore, a taxonomist who specializes in this group should make species determinations, especially when populations are detected in new locations (Blackman & Eastop 1994). Cinara cupressivora Watson & Voegtlin, Cypress Aphid (Plates 60–62) Distribution This species is believed to be indigenous to the Near East, from eastern Greece to south of the Caspian Sea. It has been introduced and become established in portions of Africa, Europe and South America.

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Table 11.1 Representative species of giant conifer aphids, Cinara spp., (Hemiptera: Aphididae): their distribution and hosts. Species

Distribution

Hosts

C. atlantica (Wilson)

Eastern North America, Cuba, Jamaica Southeastern USA, introduced into South Africa

Pinus

C. cronartii Tissot and Pepper

C. cupressi (Buckton)

North America

C. cupressivora Watson and Voegtlin Cypress aphid

Near East. Introduced into Africa and South America

C. maritimae (Dufour)

Mediterranean Region and Near East Introduced into Argentina, Brazil and Chile Europe, Japan, Siberia Eastern Canada, USA Western North America

C. pini (Linnaeus) C. pinivora (Wilson) C. ponderosae Williams C. pseudotsugae Wilson C. strobi (Fitch) C. thujafilina (Del Guercio)

Western North America Eastern North America Asia

On lesions caused by fusiform rust, Cronartium fusiforme on Pinus elliottii, P. serotina and P. taeda Cupressus, other Cupressaceae Cupressaceae. Austrocedrus chilensis, Callitris, Cupressus, Juniperus, Tetraclinis articulata, Thuja, Widdringtonia Pinus

Remarks

Damaging to P. taeda in South Africa

Vector of the fungus Seiridium cardinale Major pest of Cupressus in eastern and southern Africa

Pinus sylvestris Pinus Pinus ponderosa Pseudotsuga menziesii Pinus strobus Cupressaceae. Callitris, Chamaecyparis, Cupressus, Juniperus, Thuja, Widdringtonia

Source: Blackman & Eastop 1994.

Hosts Within its native habitat, C. cupressivora infests Cupressus sempervirens. In places where it has been introduced, it feeds on a many plants of the family Cupressaceae, including Austrocedrus chilensis, Callitris spp., Cupressus spp., Juniperus spp., Tetraclinis articulata, Thuja spp. and Widdringtonia spp.

Importance Life stages feed in colonies on stems. This insect is a pest of Cupressus lusitanica and other Cupressaceae in places where it has been introduced. In eastern and southern Africa, infestations were detected in Malawi in 1986, where it was initially identified as C. cupressi (Buckton). From Malawi, the insect spread into neighboring Burundi, Rwanda, Tanzania and Uganda. In 1990, infestations were discovered in Kenya, Zambia and Zimbabwe and in 2003 the insect

was discovered in Ethiopia. In eastern and southern Africa, populations build to high levels just prior to the end of the rainy seasons. During the dry seasons, foliage of infested trees dries and turns red-brown. Initially, infested trees have some degree of recovery during the subsequent rainy season and produce new foliage. However, stress caused by a combination of aphid feeding and dry seasons is cumulative and trees die. In Kenya, the native Juniperus procera is also infested but has sustained relatively little damage. In 2003–4, infestations were discovered in Chile, where both natural forests of Austrocedrus chilensis and plantations of Cupressus macrocarpa were damaged (Author’s observations).

Life History Because of the confused taxonomy prior to 1999, when this species was described, knowledge of

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its life history and habits is based on studies conducted on populations that may, or may not, be of this species. In Africa, only parthenogenic forms are known.

Pest Management In Africa, classic biological control with the parasitoid Pauesia juniperinum has met with some success. In Kenya, extensive plantations of C. lusitanica, which once comprised 46% of the country’s forest plantation resource, have been replaced by species of Eucalyptus (Mustafa 1987, Ciesla 1991, Ciesla et al. 1995, Watson et al. 1999, FAO n.d. (b)). Elatobium The genus Elatobium consists of five species. Three attack conifers, one occurs on Salix and another attacks Trochodendron. One species is an important pest of spruce, Picea spp.

Elatobium abietinum (Walker), Green Spruce Aphid Distribution Green spruce aphid is native to continental Europe. It has been introduced and become established in Australia (Tasmania), Chile, the Falkland Islands, Iceland, New Zealand, western North America (British Columbia, Canada; Arizona, New Mexico, Oregon and Washington, USA) and the UK.

Hosts Species of spruce, Picea spp., are either actual or potential hosts. True firs, Abies spp. are also subject to attack, but less commonly so than spruce.

Importance Life stages feed on lower surfaces of needles. Feeding causes discoloration, loss of old needles and growth reduction. Damage can be especially severe to Picea sitchensis, native to North America and widely planted in the UK. Severe damage has also occurred on both P. engelmannii and P. pungens in portions of southwestern USA.

Life History In its native range in continental Europe, the life cycle is holocyclic. Parthenogenic populations increase in autumn and spring, with both

winged male and female forms. Sexual forms appear in autumn that produce an overwintering egg stage. Introduced populations in the UK lack a sexual stage and parthenogenic reproduction continues throughout the year. Males have been detected in populations in Arizona but occurrence of sexual reproduction is yet to be determined.

Description of Stages Nymphs are wingless, oval, green and approximately 1 mm long. Adults may be winged or wingless, about 2 mm long, but otherwise similar in body shape and color to nymphs. They have a yellow-green head and dull red eyes. Eggs are oval and yellow-red to brown or black (Speight & Wainhouse 1989, Blackman & Eastop 1994, Halldórsson et al. 2003, Lynch 2009). Phloeomyzus passerinii Signoret, Woolly Poplar Aphid Distribution Woolly poplar aphid is found over much of Europe and the Near East. It is also known from two locations in North America, an unspecified site in the Atlantic region of Canada and one in Maine, USA, where it was probably introduced.

Hosts Poplars, Populus spp., are hosts. The preferred host is P. nigra and its hybrids, including P.  euroamericana. P. alba is resistant to attack.

Importance Stems and branches are infested. Smooth-barked trees are preferred. Heavy infestations can cause growth loss, branch dieback and tree mortality. This insect is a pest of fast-growing hybrid poplar plantations, especially in Mediterranean Europe, including portions of France, Greece, Italy and Spain. In northern Italy, infestations have caused mortality of P.  euroamericana plantations. This aphid is also of concern in the Near East and is reported as a pest in Iran, Israel and Turkey.

Life History Sexual forms have been reported in the UK but most typically only asexual forms are found. This insect typically undergoes a succession of generations of alate and apterous female adults that produce live young. Each female can produce more than 1000

Sucking insects offspring and there may be as many as 10–12 generations/year. Adults produce copious quantities of flocculent white woolly wax and form dense colonies on trees of almost any age, preferably in shaded portions of stands or on the north-facing sides of stems.

Description of Stages Adults and nymphs are small, yellow-green and hidden under a dense covering of woolly wax secreted by the adults.

Pest Management Infestations can be treated with contact insecticides. Considerable work has been done to identify strains or clones of fast-growing poplars resistant to attack (Browne 1968, Blackman & Eastop 1994, Allegro & Cagelli 1996, Karahroodi et al. 2006, Bankhead-Dronnet et al. 2008).

Adelgidae Adelges Adelges is a small genus of conifer infesting aphid-like insects. Species occur throughout conifer forests of Eurasia and North America. They have complex life cycles and some have alternate hosts with a sexual form that produces galls on spruce, Picea spp. These are reviewed in Chapter 12. Two species that infest branches or boles of “secondary” hosts have been introduced into new locations where they have caused severe damage. Adelges tsugae (Annand), Hemlock Woolly Adelgid (Plates 63 & 64) Distribution This adelgid is native to Asia, including southwestern China, India, Japan, Nepal and Taiwan. It was discovered in western USA and Canada during the 1920s and initially was believed to be introduced. Infestations were first reported in eastern USA in 1951, near Richmond, Virginia. By 2005, infestations were established over portions of 16 eastern states from Maine to Georgia. A study based on analyses on mitochondrial DNA, concludes that populations of hemlock woolly adelgid in eastern North America were the result of an introduction from low-elevation forests in Japan. Populations in western North America may be either indigenous or introduced.

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Hosts In its native range, both hemlock, Tsuga spp., and spruce, Picea spp., are hosts. Hemlock hosts in Asia include T. chinensis in China, T. diversifolia and T. sieboldii in Japan, and T. dumosa in the Himalayas. Spruce hosts are Picea jezoensis hondoensis and Picea polita. Western hemlock, Tsuga heterophylla, and mountain hemlock, T. mertensiana, are hosts in western North America. Eastern hemlock, T. canadensis and Carolina hemlock, T. caroliniensis, are attacked in eastern USA. North American spruces apparently are unsuitable hosts for this adelgid.

Importance In Asia and western USA, hemlock woolly adelgid causes little or no damage. Feeding on eastern and Carolina hemlocks causes needles to desiccate and buds to stop growing. Within a few months of heavy infestation, trees appear gray-green, needles drop, and little or no new foliage is produced. Foliage loss and dieback of major limbs is visible in 2–4 years. An infested hemlock may survive for several years, but foliage is usually sparse at branch tips and in the upper crown. Weakened trees succumb to disease and attacks from other insects, such as hemlock borer, Melanophila fulvoguttata.

Life History This adelgid has a complex life cycle. In North America and Japan there are three generations/ year: sistens, which overwinter; progrediens, which remain on hemlock; and sexuparae, which migrate to spruce. In May–June, “winged” sexuparae and the “wingless” progrediens develop simultaneously. In June, the sexuparae females fly to a spruce host, on which they lay eggs. These “unfertilized” eggs hatch into sexuales which, in North America, fail to develop. The wingless female progrediens deposit eggs on hemlock, which hatch in June and July. First-instar nymphs, or crawlers, look for a place to feed. These eventually attach themselves to the bases of needles where they feed briefly before becoming inactive. This period of inactivity lasts until mid-October when feeding resumes and nymphs mature during autumn and winter. They develop a waxy overcoat as they mature into adults. The sisten generation matures into adults in February after four nymphal instars. During March– May, adult sistens produce a single ovisac containing up to 300 eggs which develop into sexuparae and progrediens adults about 4 weeks after eggs hatch. The number of eggs that become sexuparae increases with

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adelgid density. This could be the result of declining host nutrition.

Description of Stages Adults are less than 1.5 mm long and vary from dark red-brown to purple-black in color. When they mature, they are covered with woollike wax filaments (ovisacs). Ovisacs are present from late autumn to early summer on the underside of the outermost branch tips of hemlocks.

Pest Management Populations established in highelevation forests or in northeastern USA are subject to high levels of overwintering mortality. Cultural, regulatory, chemical and biological tactics have been used to reduce this insect’s rate of spread and protect individual trees. Moving bird feeders away from hemlock trees and removing isolated infested trees from a woodlot helps prevent further infestations. Quarantines help prevent movement of infested materials into un-infested areas. Chemicals, including horticultural oils and insecticidal soaps, are effective when trees are saturated. This ensures that the insecticide comes in to contact with all of the population. Systemic insecticides have proven effective when applied to the soil around the base of trees or injected into the stem. Chemical control is limited to individual tree treatments in readily accessible, non-environmentally sensitive areas and is not practical in forests. The most promising tactic for hemlock woolly adelgid management is classic biological control. As is the case with other Adelgidae, there are no known parasitoids of this insect. Several predaceous beetles, including Scymnus ningshanensis, Scymnus sinuanodulus and Sasajiscymnus tsugae, and Laricobius nigrinus and fungi have been introduced from Asia or western North America with some success (McClure & Cheah 1999, Reardon & Onken 2004, Shields & Cheah 2005, USDA Forest Service 2005, Havill et al. 2006, Canadian Food Inspection Agency 2008).

Adelges piceae (Ratzeburg), Balsam Woolly Adelgid (Plates 65 & 66) Distribution Balsam woolly adelgid is native to central Europe. It was introduced into southeastern Canada and northeastern USA during the early 1900s. It appeared on the North American Pacific Coast

around 1929 and in the southern Appalachian Mountains in the 1950s. Infestations spread into northern Idaho during the 1980s.

Hosts All species of true fir, Abies spp., are hosts. In Europe, Abies alba is the primary host. In North America, A. balsamea and A. fraseri in the east and A. amabilis, A. grandis and A. lasiocarpa in the west are hosts.

Importance In Europe, this insect causes little or no damage although trees support heavy populations. Asian species of Abies are intermediate in sensitivity to attack; some are damaged, others are not. In North America, balsam woolly adelgid has become an important pest of fir forests. Two forms of damage may occur. Infestations on twigs and branches cause “gouting,” the development of swellings at the buds and branch nodes. This causes a crown decline, branch dieback and top kill. The other is bole attack, where infestations cause a red growth of sapwood known as “rotzholz” or redwood. Species sensitive to attack can die in 2–3 years due to bole infestations. A. fraseri, a species endemic to the southern Appalachian Mountains, and A. lasiocarpa, of western North America, are especially sensitive to stem attack.

Life History Balsam woolly adelgid has two to four generations/year, depending on location and elevation. Two generations are most common. Most overwinter as sessile nymphs (neosistens) and develop into adults by late June. They produce a waxy covering that appears as white, woolly spots about the size of a pin-head on the bole, limbs, and buds of host trees. In North America, only females occur. They lay eggs under the woolly masses for about 6 weeks. Eggs hatch within a few days and early-instar nymph, known as “crawlers” move about for several weeks. Crawlers are easily moved via air currents and this is how populations disperse. Crawlers transform into a sessile neosistens stage, which lasts from 2 to 8 weeks. Secondgeneration adults are abundant by late September– early October and egg-laying continues until about mid-November. A partial third generation occurs in the southern Appalachian Mountains, and a partial fourth generation occurs in low-elevation forests of Oregon and Washington.

Sucking insects Description of Stages Adults are dark purple, roughly spherical in shape, less than 1 mm long, and almost invisible to the naked eye. They are typically wingless although a rare winged form (progrediens) has been seen in Europe and eastern Canada. Eggs are produced under the adults and are amber in color. Nymphs are orange with black eyes. All life stages are best seen with a hand lens.

Pest Management Classic biological control was attempted after infestations were discovered in North America. Balsam woolly adelgid has no known parasitoids but several predators. They include the beetles Aphidecta obliterata, Laricobius erichsonii and Pullus impexus, and flies Aphidoletes thompsoni and Leucopis obscura. Several have become established but none has had a significant impact on populations. Ground application of chemical insecticides is effective but expensive and only practical in high value areas such as Christmas tree plantings (Ragenovich & Mitchell 2006). Pineus, Pine Woolly Adelgids Pineus consists of about 21 species distributed throughout conifer forests of the northern hemisphere. Hosts

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are species of Picea and Pinus. Some are holocyclic, have both sexual and asexual stages and alternate between a sexual, gall forming stage on Picea and an asexual form on Pinus (see Chapter 12). Other species are autocyclic and have asexual forms on either Picea or Pinus (Table 11.2). Several have been introduced and become damaging. All species are similar in appearance, difficult to separate and life stages live under a covering of white wool. Some were originally described from places where they were introduced and their natural ranges are not well understood (Blackman & Eastop 1994, Havill & Foottit 2007). Pineus boerneri Annand (¼P. laevis (Maskell)) Distribution This species, originally described from Pinus radiata in California, USA, is believed to be Eurasian in origin where it has also been reported as P. laevis (Maskell) and P. pini Maquart. Much of the literature on this species appears under these former names. This species has been introduced into Africa, Australia, North America (northeastern USA and Hawaii), South America and New Zealand. It was first reported in Africa from both Kenya and Zimbabwe in 1968 and spread rapidly across pine plantations throughout the continent. Initial introduction into

Table 11.2 Representative anholocyclic, non-gall making species of Pineus (Hemiptera. Adelgidae), their distribution and hosts. Species

Distribution

Hosts

P. abietinus Underwood & Balch

Western North America

P. boerneri Annand

Eurasia? Africa, Australia, North America (northeastern USA and Hawaii), South America and New Zealand (introduced) China North America Western Himalayas Europe North America (introduced) Europe (indigenous) China

Abies amabalis, A. grandis, A. lasiocarpa Pinus

P. P. P. P.

cladogenous Fang & Sun coloradensis (Gillette) ghanii Yaseen & Ghani pineoides (Cholodkovsky)

P. pini (Macquardt) P. piniyunnanensis Zhang, Zhong & Zhang 1992 P. simmondsi Yaseen & Ghani P. strobi (Hartig) P. wallichianae Yaseen & Ghani

Western Himalayas North America (indigenous) Europe (introduced) Western Himalayas

Pinus Pinus Pinus Picea Pinus

Pinus Pinus monticola, P. strobus, P. peuce Pinus

Sources: Browne 1968, Furniss & Carolin 1977, Drooz 1985, Blackman & Eastop 2004, Havill & Foottit 2007.

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Africa was probably via P. taeda scion material imported from Australia.

Hosts This insect infests various pines, Pinus spp., including P. halepensis from Mediterranean Europe and P. caribaea and P. radiata from North and Central America. In Africa, P. elliottii, P. kesiya and P. radiata are attacked and P. patula and P. taeda are only slightly susceptible. P. resinosa is attacked in eastern North America.

Damage Nymphs and adults suck plant juices from needles and stems and cause deformity, growth loss, branch dieback and occasional tree mortality (Fig. 11.3). Excreted honeydew is a medium for sooty mold.

Life History Life history studies conducted in Africa indicate that reproduction occurs entirely by parthenogenesis. Both winged and wingless adults occur but wingless forms are more common. Aphids feed on the base of needles and on tender young bark. In Kenya and Zimbabwe, this insect is most abundant during dry seasons.

Pest Management Classic biological control using a predator of the genus Leucopis has been attempted in Chile, New Zealand and the USA (Hawaii) (McClure 1985, Murphy et al. 1991, Diekmann et al. 2002, Blackman & Eastop 1994, Schabel 2006).

Pineus strobi (Hartig), Pine Bark Adelgid Distribution This adelgid is found throughout North America and has been introduced into Europe, including the UK.

Hosts Primary host is eastern white pine, Pinus strobus, both in North America and continental Europe. Western white pine, Pinus monticola, is infested in western North America. In the British Isles, it is also found on P. peuce.

Importance Large patches of white cottony material may cover the bole and branches giving them a whitewashed appearance (Fig. 11.4). Although mature eastern white pines may become unsightly when heavily infested, they are not damaged. However, young trees and nursery stock may suffer damage. Needles turn yellow on heavily infested young trees and feeding may cause stunting or death.

Fig. 11.3 Pine woolly aphid, Pineus boerneri, infestation on pine stem and resultant deformity (western Uganda).

Life History In spring, females lay eggs that produce both winged and wingless females. Wingless forms remain on the host tree and reproduce several times. Some of the winged forms may fly to spruce instead of white pine, where they settle on the needles, lay eggs and die because the larvae cannot survive on spruce. Up to five generations have been observed in north central USA (Drooz 1985, Blackman & Eastop 2004).

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into motile nymphs (crawlers) that may feed on the foliage or shoots. In late summer, the nymphs seek a permanent feeding site and molt into a sessile, legless form that overwinters. Adult females are red-brown, wingless and range in size from 1.6 to 4.7 mm long. Body shape is oval or pear shaped and coarsely wrinkled. Males are smaller and winged. They have a prominent brush of long, waxy filaments at the end of the abdomen. Eggs are amber and oval in shape. First-instar nymphs somewhat resemble female adults but are smaller and average 0.4 mm in length. The posterior of the nymphs have two long sensory hairs. Second-instar nymphs are elliptical, legless and lack antennae (Bean & Godwin 1971, Foldi 2001). Matsucoccus acalyptus Herbert, Piñon Needle Scale Distribution Piñon needle scale is indigenous to southwestern USA. Hosts Hosts are piñon pines, Pinus edulis and P. monophylla.

Fig. 11.4 Infestation of pine bark aphid, Pineus strobi, on eastern white pine, Pinus strobus (Fort Collins, Colorado, USA).

Margarodidae, Pine Bast Scales Matsucoccus Matsucoccus consists of about 34 species that infest conifers of the genus Pinus; in North American 19 species are known, there are 16 in the USA, two in Mexico and one in the Dominican Republic. The remainder occur in Eurasia (Table 11.3). The taxonomy of several species has been unclear. Several are important pests. All species have similar life cycles. They may have one or two generations/year and winter is spent as a sessile nymph. Adults are active in early spring–summer. Females are wingless but motile and lack mouthparts. Males are winged. After mating, females deposit eggs in an ovisac that remains attached to the abdomen. Eggs hatch

Importance Heavy feeding during outbreaks can kill small trees. Successive years of feeding on large trees results in loss of older foliage and makes them susceptible to attack by the bark beetle, Ips confusus. Outbreaks have caused damaged trees in several national parks including Grand Canyon National Park in Arizona and Mesa Verde National Park in Colorado (Furniss & Carolin 1977). Matsucoccus feytaudi Ducasse, Maritime Pine Bast Scale Distribution This scale is native to Morocco, Portugal and Spain and but has spread into portions of France (including Corsica) and northern Italy. Hosts

This scale has a single host, Pinus pinaster.

Importance Little or no damage is caused in Atlantic coastal forests. When introduced into the maritime Alps region of southeastern France it destroyed nearly all of the pine forests over an area of approximately 120,000 ha. This may have been due to climatic differences and the genetic susceptibility of local provenances of P. pinaster. M. feytaudi has also damaged

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Table 11.3 Representative species of Matsucoccus (Hemiptera: Margarodidae): their distribution and hosts. Species

Distribution

Hosts

M. acalyptus Herbert Pin˜on needle scale M. feytaudi Ducasse Maritime pine bast scale M. josephi Bodenheimer & Harpaz Israeli pine bast scale M. matsumurae (Kuwana) Japanese pine bast scale (¼ M. resinosae Bean & Godwin, red pine scale, ¼ M. thunbergiana Miller & Park M. pini Green

USA: southwestern states

Pinus edulis, P. monophylla

Morocco, Portugal, Spain (native). France, Italy (introduced) Israel

Pinus pinaster

Japan (indigenous), North America, China, South Korea (introduced)

Central, northern and eastern Europe, including the UK

Pinus brutia, P. eldarica, P. halepensis Pinus spp. (seven species of the resinosae group)

Pinus mugo, P. sylvestris

Sources: Furniss & Carolin 1977, McClure 1983, 1985, 1990, Kolk & Starzyk 1996, Foldi 2001, Booth & Gullan 2006, Liphschitz & Mendel 2006.

pines in the Liguria region of northern Italy and on the French island of Corsica. Attacks occur on the bark and feeding causes yellowing of the foliage. Weakened trees are susceptible to attack by secondary insects such as the bark beetle Tomicus destruens and the weevil Pissodes notatus (Abgrall & Soutrenon 1991, Anderson 2005, CAB International 2001).

Matsucoccus matsumurae (Kuwana), Japanese Pine Bast Scale (¼M. resinosae Bean & Godwin, Red Pine Scale ¼ M. thunbergianae Miller & Park) Distribution This scale is native to Japan and has been introduced into portions of China, Korea and eastern North America. It may have been introduced into New York City in 1939 during the World Fair on exotic pines and is now found in portions of eastern Canada and northeastern USA.

Hosts Seven species of Pinus of the “resinosae” group are known hosts. P. densiflora, P. luchuensis and P. thunbergii in Japan, P. insularis, P. massoniana, P. tabulaeformis and P. taiwanensis in China and Taiwan, and P. resinosa in eastern North America. Importance M. matsumurae is relatively innocuous on its native host pines in Japan. However, it has caused extensive damage to P. massoniana and P. tabulaeformis

in China and the entity formerly known as M. resinosae has killed entire plantations of P. resinosa in portions of northeastern USA and southeastern Canada (Bean & Godwin 1971, McClure 1983, 1985, 1990, Booth & Gullan 2006). Matsucoccus pini Green Distribution This scale occurs over much of Europe, including the British Isles.

Hosts Primary host is Pinus sylvestris but other pines native to Europe, such as P. mugo, may also become infested.

Importance M. pini is often associated with forests affected by air pollution from industrial sources and is one of the primary agents associated with decline and mortality of pines. Heavy infestations can cause tree mortality (Kolk & Starzyk 1996).

Pseudococcidae, Mealybugs Maconellicoccus hirsutus (Green), Pink Hibiscus Mealybug Distribution This species is believed native to southern Asia and has spread to central and northern Africa,

Sucking insects northern Australia, India, Pakistan and the Pacific Islands. In 1994, it was detected for the first time in the western hemisphere on the Caribbean island of Grenada. In has since spread to Guyana and Venezuela in South America and across much of the Caribbean, including the islands of Anguilla, Antiqua, British Virgin Islands, St Eustatius, St Kitts, Nevis, St Lucia, St Vincent, Trinidad and Tobago. Infestations have also been detected in Mexico (1999) and the USA, including Hawaii (1983), California (1999), Florida (2002), Louisiana (2006) and Texas (2007). Hosts In addition to species of Hibiscus, this mealybug feeds on over 76 families and 200 genera of plants, including species of importance in agriculture, horticulture and forestry. Preferred hosts are plants in the families Fabaceae, Malvaceae and Moraceae. Hosts of importance in forestry include Hibiscus elatus and Tectona grandis.

Importance Feeding occurs in colonies and, if left undisturbed, they will expand into large masses of white, waxy deposits on branches, fruit, foliage and, in extreme cases, entire plants. During feeding, a toxic saliva is injected into the plant that causes leaf curling, shoot distortion, stunting and, occasionally, death. Leaves show a characteristic curling similar to that caused by some plant viruses. Heavily infested plants have shortened internodes leading to rosetting and a “bunchy top” appearance. Black sooty mold may develop due to honeydew secretions. Infestations in Grenada have killed trees. Prior to initiation of a classic biological control program, crop losses were estimated at US$ 3.5 million.

Life History Pink hibiscus mealybug can complete a generation in 23–30 days and, under optimum conditions, can have up to 15 generations/year. Reproduction occurs by parthenogenesis if males are absent. In temperate climates, overwintering may occur in the soil or on host plants, either as eggs or adults. In tropical climates, they are active throughout the year. Females deposit up to 650 eggs in an egg sac of white wax, usually in clusters on twigs, branches and bark of host plants. Eggs hatch into motile nymphs (crawlers), which disperse over the plant, especially toward tender growing portions. Female nymphs have four instars and males have three. The final male instar is inactive and

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remains in a cocoon of mealy wax. Eggs and nymphs are easily dispersed by winds. Eggs are covered with sticky wax, which easily attaches to animals or humans for dispersal.

Description of Stages Adults of both sexes are about 3 mm long. Females are pink and have a white, waxy covering. They are wingless and ovoid in shape. Males have a pair of wings and two long waxy tails and can fly. Eggs are 0.3–0.4 mm long, orange when first deposited but turn pink with age. Nymphs resemble adults but are smaller.

Pest Management Classic biological control using imported predators and parasitoids has met with success in some areas and is considered the best long-term tactic for management of this insect. About 21 parasitoids and 41 predators are known to attack this insect worldwide, including the ladybird beetle Cryptolaemus montrouzieri and the egg parasitoid Anagyrus kamali (USDA APHIS 1999, EPPO 2005d). Oracella acuta (Lobdell), Loblolly Pine Scale (Fig. 11.5) Distribution This insect is indigenous to southeastern USA and was introduced into Guangdong Province, China in 1988.

Hosts Pinus echinata, P. palustris, P. taeda and P. virginiana are hosts within its natural range. In China, P. elliottii is the main host but P. massoniana, an indigenous species, is also infested.

Importance This insect is usually of minor importance in its native habitat but can appear in large numbers in seed orchards following use of chemical insecticides for treatment of other insects. Following its introduction into China on pine scion material collected in the USA and grafted on to rootstocks in 1988, the insect spread rapidly. By June 1995, over 212,500 of plantations were infested. Heavy infestations cause premature loss of needles, reduced shoot growth, shorter needles and bud mortality. Height growth can be reduced by 25–30%. Infestations can also occur on cones and cause deformity.

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Forest entomology: a global perspective Eriococcidae Cryptococcus fagisuga Lindinger, Beech Scale Distribution Native to Europe, beech scale was introduced into North America around 1890 near Halifax, Nova Scotia, Canada. It was discovered in the USA in 1929 in Boston, Massachusetts and spread rapidly throughout eastern Canada and northeastern USA. Infestations have spread south and west to West Virginia (1981), Virginia (mid-1980s), North Carolina and Tennessee (1994) and Michigan (2000).

Hosts Hosts are beeches, Fagus spp. In Europe, the host tree is F. sylvatica and in North America, F. grandifolia is infested.

Fig. 11.5 Branch of slash pine, Pinus elliottii, infested by loblolly pine scale, Oracella acuta (Guangdong Province, China).

Life History In its natural range, O. acuta, has four to five generations/year. It has at least that many generations in China. Motile nymphs or crawlers overwinter under resin cells. Nymphs and adults feed on buds and expanding shoots of pines. They produce cells of white resin used for a protective cover. Honeydew exudations provide a medium for sooty mold. Sexual reproduction can occur but most reproduction is by parthenogenesis and most adults are females. Dispersal is by movement of crawlers via air currents.

Description of Stages Males are winged and females lack wings. Eggs are pale orange. Nymphs are pale rose in color (Clarke et al. 1990, Sun et al. 1996, Diekmann et al. 2002).

Importance Beech scale infests the boles, which causes drying and cracking of the bark. Trees are subsequently invaded by fungi of the genus Nectria, which cause decline and death. Reports from Europe as early as 1849, describe extensive death of beech forests. Beech scale, readily visible on trees, was considered the cause of death until 1914, when it was learned that a fungus, then identified as N. ditissima, infected trees infested by the scale. In North America, the primary invading fungus is N. coccinea var. faginata. A second species, N. galligena, is occasionally involved. Since its introduction into North America, this scale/ fungus complex has caused extensive decline and mortality of F. grandifolia, a component of northern hardwood forests.

Life History Beech scale has one generation/year. Overwintering occurs as instar II nymphs. Adults appear in spring and egg deposition begins in mid summer. All adults are females and reproduce asexually. Eggs are deposited in strings of four to eight eggs attached to the bark. Eggs hatch in late summer and continue hatching until early winter. They hatch into crawlers with well-developed legs and antennae. Some migrate to cracks in the bark, where they settle and feed. Others are washed to the ground by rains and die. Still others are carried to neighboring trees by air currents. They molt into instar II nymphs, which lack legs and are covered with wax.

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Description of Stages Adults are yellow, elliptical soft-bodied scales, 0.5–1.0 mm long. They have redbrown eyes, rudimentary antennae and legs and numerous glands that secrete white woolly wax. Nymphs are yellow-white and about 0.3 mm long.

Pest Management Direct control in forested areas is impractical but infested ornamental trees can be treated with insecticides. Cold winter temperatures (35oC) can kill scales not protected by snow. There are indications that individual trees may be resistant to infestation (Houston & O’Brien 1983, Drooz 1985, Ciesla & Mason 2005).

Gossyparia spuria (Modeer), European Elm Scale (Fig. 11.6) Distribution This insect is native to Europe and introduced and established in North America. It is widely distributed across the USA and portions of Canada.

Hosts

All species of elms, Ulmus spp., are hosts.

Importance European elm scale infests leaves and branches of elms and is found in both forests and urban settings. It is common on ornamental elms and heavy infestations can cause branch dieback. Honeydew produced by the scales promotes growth of sooty mold fungi. Heavily infested trees are especially visible in winter when branches appear black with profuse mold growth.

Life History This scale has one generation/year and instar II nymphs overwinter in bark crevices. In some areas, males are not produced and females reproduce asexually. When males occur, male nymphs develop into adults during January–March and are most abundant in early spring. Females appear when elm seeds develop in spring. They move to branches where mating occurs and females develop waxy sacs. Egg deposition begins when leaves develop and continues into late summer. One female may lay up to 400 eggs. Eggs hatch into motile nymphs (crawlers) within an hour of deposition and move to undersides of leaves to feed near the primary veins. Later, they

Fig. 11.6 Colony of European elm scale, Gossyparia spuria, on a branch of American elm, Ulmus americana (Fort Collins, Colorado, USA).

migrate back to the branches, molt into the second instar and overwinter.

Description of Stages Female adults are brown or green-brown soon after they molt but turn gray with age. They are covered with a waxy sac that leaves only the top center of the scale exposed. Body fluids of the females are orange. Males are winged and may have either long or short wings. Instar I nymphs are yellow and instar II nymphs are red brown.

Pest Management Crawlers are susceptible to applications of either horticultural oils or chemical insecticides to the foliage. Instar II nymphs are less susceptible to foliar applications. Soil treatments using

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the systemic insecticide imidacloprid have proven effective for control of this scale (Drooz 1985, Colorado State University 2004). Kermesidae, Kermes Scales Allokermes kingii (Cockerell), Northern Red Oak Kermes Distribution This species is native to North America and is found throughout the much of eastern and southwestern USA.

Hosts Many species of oak, Quercus spp., are hosts and Q. rubra and Q. velutina are favored.

Female adults are covered with a protective shell that is an integral part of their bodies. Secretion of wax from pores creates a very convex shell. Color is pale yellowbrown, marbled with a slightly dark red tint and with small black spots covering the entire surface. Antennae and legs have six and five segments, respectively. Pest Management Pruning of infested branches and removal of scales by hand and destruction of infested plant material will reduce infestations on urban trees (Turner & Buss 2004). Kermococcus vermilis Donkin (¼ Kermes vermilis, K. ilicis, Coccus ilicis) Distribution This scale occurs in Mediterranean Europe and the Near East.

Importance Feeding causes branch dieback, reduced growth and sooty mold, which grows on honeydew secreted by the scales. Tree death may occur as a result of heavy infestations. This insect is a pest of ornamental trees and of little or no consequence in forests. Life History This scale has one generation/year in Virginia and two in Florida. In Florida, first-generation nymphs (crawlers) appear in late May. After hatching, they remain under parental brood chambers until conditions favor their dispersal. Females migrate to large branches and males to bark crevices. Crawlers molt to a second instar by mid-July and instar II females migrate to tree wounds or new growth, often near leaf petioles. They become sessile and secrete a hard waxy covering over their bodies. Instar II males migrate further down on the tree stems, become sessile and secrete a white, felt-like waxy pupal case. Females molt to a third instar, and become adults from late August to mid-December. Eggs are deposited in brood chambers from early September to mid-December. Each female can lay an average of 3000 eggs. The female’s abdomen shrinks after eggs are laid. Dead females may remain on the host plant for a year or more after the next brood emergences. Second-generation crawlers appear in mid-September, molt into instar II by mid-October and overwinter. By late April, they become mature adults and deposit eggs until mid-June. Description of Stages Adult females are about 5 mm long, 4.3 mm wide and about 3.5 mm high.

Hosts Two species of Mediterranean oaks, Quercus coccifera and Quercus ilex, are hosts.

Importance This insect was once a highly valued species because female adults were a source of red dye to color wool and silk before other red dye sources were available. The pigment is kermesic acid and the color descriptors crimson (English), carmosine (French), karmir (Armenian), kimiz (Persian) and kirmina (Sanskrit) all refer to the pigment derived from this insect. At one time this scale was reared commercially on oak trees in southern France, Spain and other portions of the Mediterranean. The scales were harvested before dawn by women carrying lanterns and picking the insects with fingernails that were kept long for this purpose (Ciesla 2002a).

Kerriidae Kerria (¼ Laccifer) lacca (Kerr), Lac Insect Distribution Lac insect is an Asian species found in China, India, Myanmar, Pakistan and Thailand.

Hosts The lac insect feeds on stems of over 100 species of broadleaf trees, especially those that produce gums, latex, mucilage, resins or have high tannin content.

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Importance This insect is known for secretion of lac, a scarlet substance used to dye wool and silk, as a cosmetic, a medicinal drug and as shellac to coat candies and give fresh fruits and vegetables a glossy finish. It has been cultured in India and, to a lesser extent, Myanmar and Thailand and is considered one of the most valuable of all plant parasitic insects. Major tree species used for lac culture include Acacia nilotica, Butea monosperma, Ficus elastica, Ficus spp., Ougeinia dalbergioides, Schleichera oleosa, S. trijuga, Shorea talura and Ziziphus mauritania.

Life History In India, there are two, rarely three generations/year. Females live within a cell formed of lac, which has two respiratory pores and an anal pore kept open by long wax filaments. Males are short lived and fertilize females via the anal pores. Females also have the capacity to produce eggs without fertilization. They produce several hundred eggs in their cells and die soon thereafter. Instar I nymphs are motile, swarm out of the mother’s cell and migrate to young succulent shoots, where they settle and feed in dense colonies. As they feed, they secrete lac, which first appears as a shiny coating over the bodies and later spreads and thickens to form a crusty surface that covers the entire colony.

Description of Stages Female adults lack wings, eyes and legs. They are sac-like and about 1.5 mm long. Male adults have both winged and wingless forms. They are red or, rarely, yellow with eyes and legs but no functional mouthparts.

Related Species Species of the genera Afrotachardina, Austrotacharidia, Laccifer, Metatachardia, Tachardina and Tachordiella also produce lac but of inferior quality and are used only locally (Browne 1968).

Diaspididae, Armored Scales Aonidiella orientalis (Newstead), Oriental Scale (Fig. 11.7) Distribution This scale is widely distributed in tropical regions of the eastern hemisphere, including portions of Africa, Southeast Asia and the Indian

Fig. 11.7 Infestation of oriental scale, Aonidiella orientalis, on foliage of neem, Azadirachta indica (Kano, Nigeria).

subcontinent. The insect may have originated in Asia and then spread into Africa.

Hosts Oriental scale has a wide host range including species of Citrus and a number of tropical forest species. Aegle marmelos, Anacardium occidentale, Azadirachta indica, Butea monosperma, Cassia fistula, Chloroxylon swietenia, Dalbergia sissoo, Melia azedarach, Santalum album, Schleichera trijuga, Swietenia mahogani, Syzygium cuminii, Tamarindus indica and Ziziphus mauritania.

Importance Scales infest leaf petioles, foliage, fruit and stems. During heavy infestations, foliage becomes discolored and branch dieback occurs. This causes

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growth loss and death of young trees. This scale is considered an important pest of citrus and in India is a pest of Dalbergia sissoo plantations where periodic outbreaks have caused severe damage. Infestations also interfere with the culture of the beneficial lac insect, Kerria (¼ Laccifer) lacca, in India (see section in this chapter). In Nigeria, oriental scale is a pest of neem, Azadirachta indica, where it causes widespread damage to ornamental trees across the northern part of the country. Damaged foliage turns brown and dies.

Description of Stages Adults are flattened, circular or slightly oblong and 1.6 mm in diameter. They vary in color from yellow-light or red brown (Browne 1968, Boa 1995). Lepidosaphes ulmi (Linnaeus), Oystershell Scale (Figs 11.8 & 11.9) Distribution This scale is probably of European origin and introduced into North America. It is now present throughout the USA and much of Canada.

Life History Oriental scale has multiple overlapping generations and egg deposition is continuous throughout the year.

Hosts Many broadleaf trees and shrubs are hosts including species of Acer, Betula, Buxus sempervirens,

Fig. 11.8 Oystershell scale, Lepidosaphes ulmi, on stem of quaking aspen (Fort Collins, Colorado, USA).

Fig. 11.9 Close up of colony of oystershell scales showing distinctive oyster shape of the scales (Fort Collins, Colorado, USA).

Sucking insects Fagus, Fraxinus, Malus, Populus, Prunus, Pyrus, Salix, Syringa vulgaris and Ulmus. Trees with smooth bark are preferred. Importance Oystershell scale colonizes boles and large branches. This is a common pest of ornamental and shade trees but of little or no concern in forests. Feeding causes cracks in the bark surface and branch dieback. Heavy infestations that persist for long periods can kill trees. Life History Number of generations varies with host and location. In Maryland, for example, infestations on Acer and Syringa vulgaris have two generations/year whereas populations on Populus and Salix in the same area have one generation/year. The overwintering stage is usually eggs, clustered in groups of 50–100, under the cover of the adult scale. Crawlers emerge in May and move to feeding sites on twigs and branches. Adult females are present in July. By late July, they have formed a scale covering under which eggs are laid.

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Description of Stages The waxy cover of mature female adults is about 2.5 mm long, gray-brown, and noticeably convex, resembling miniature oyster shells. The cover of the scale may vary according to host. In Maryland, the cover of populations on Populus is red-brown with two transverse yellow bands. On Acer and Syringa vulgaris, populations have a dark brown cover. On other species, the covers are gray or graybrown. White eggs are found beneath the cover of the female. The crawler stage is pale yellow and less than 1 mm long. Adult males have a single pair of wings. When observed closely, adult males are often misidentified as parasitoids as they walk over infested twigs.

Pest Management Management tactics for oystershell scale include pruning and destruction of infested twigs and branches. The most vulnerable life stage of the insect to insecticides is the crawler, which is active from May to June. They can be managed with applications of horticultural oils or chemical insecticides (Drooz 1985).

Fig. 11.10 Black pine leaf scale, Nuculaspis californica, on needle of ponderosa pine, Pinus ponderosa (Spokane, Washington, USA).

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Nuculaspis californica Coleman, Black Pine Leaf Scale (Fig. 11.10) Distribution Black pine leaf scale is native to North America and widely distributed in southeastern Canada and eastern USA, the Rocky Mountains and the Pacific coast.

Hosts This species attacks several pines, including Pinus echinata, P. rigida in eastern North America, P. jeffreyi, P. lambertiana, P. ponderosa and P. sabiniana in the west, and P. cembroides in Mexico. Abies concolor and Pseudotsuga menziesii are also infested. Importance Needles of host trees are infested and feeding causes spots of yellow discoloration. Low-level populations of about one scale/5 cm of needle cause little or no damage. Heavy infestations can cause premature needle loss, thin crowns and growth loss. Weakened trees are subject to attack by bark beetles. Heavy populations are often associated with conditions that disrupt activities of this scale’s natural enemies. Deposits of dust from roads, excavations or emissions from industrial plants are known to cause outbreaks. Repeated use of chemical insecticides around orchards or mosquito habitats can cause outbreaks in surrounding pine forests if sprays kill the scale’s natural enemies.

Life History This scale has one generation/year in the northern parts of its range and two generations/ year further south. In northern locations males emerge and mate with females, which are immobile, in early–mid-June. Eggs hatch into crawlers between mid-July and early August. Crawlers select permanent feeding sites 2–3 weeks later and develop into adults to spend the winter. In southerly locations, the first generation of crawlers is present from midMay to mid-June and second generation crawlers are present during early–mid-August. During warm autumns and winters, some scales may start a third generation.

Description of Stages The cover of the adult female is oval, convex and black with light gray margins and a yellow central area. The male scale cover is similar to the female but is smaller and more elongate. Eggs are light yellow.

Pest Management Direct control of infestations in forested areas is not practical but individual ornamental trees can be treated effectively with contact sprays timed to treat the crawlers or application of systemic insecticides (Edmunds 1973, Drooz 1985, Ferrell 1986).

Chapter 12

Gall Insects

INTRODUCTION Plant galls are tumor-like growths of tissue produced by host plants in response to chemical and/or mechanical stimuli caused by invading organisms. These stimuli cause accelerated production of plant growth hormones. Agents that cause plant galls include bacteria, fungi, mites, nematodes, parasitic plants (e.g. mistletoes), viruses and insects. Virtually all plants are susceptible to organisms that produce galls. In most cases, galls are more conspicuous than the organisms that cause them and many are characteristic enough to identify the agent that caused the gall, often to species (Russo 2007). Estimates of the worldwide number of gall producing insects range from 21,000 to 211,000 species, many of which are yet to be described. Their distribution pattern suggests that the greatest species richness occurs in warm temperate regions with sclerophyllous vegetation (Mediterranean climates). It is likely that tropical rain forests may harbor a greater number of gall forming insects because of their plant diversity. However, relatively little is presently known about the insect

fauna of tropical rain forests (Espirito-Santo & Wilson 2007). Many insects produce galls on trees. Orders of insects that contain gall producers include the Thysanoptera (thrips), Hemiptera (aphids), Coleoptera (beetles), Lepidoptera (moths and butterflies), Diptera (flies) and Hymenoptera (bees and wasps). Often insect galls are more of a cosmetic concern or curiosity than damaging. Therefore, gall insects tend to be more of a concern on ornamental and shade trees than in forests. However, several gall making insects can cause branch dieback and tree mortality in forests and are considered pests.

HEMIPTERA Psyllidae (Jumping Plant Lice) Pachypsylla celtidismamma Riley, Hackberry Nipple Gall Distribution This insect is native to North America and is found wherever its host trees occur.

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Hosts Hosts are hackberries, Celtis sp., especially Celtis occidentalis.

Importance Heavy infestations result in leaves covered with galls, which cause leaf curl and premature leaf drop. Heavy infestations are unsightly but cause little or no permanent injury.

Life History This species has one generation/year and adults overwinter in bark crevices. They mate in spring and females deposit eggs on the undersides of expanding leaves. Nymphs hatch about 10 days later and begin feeding, which causes leaf tissue to expand into a pouch around the nymph. They go through several instars and become adults in September, when they tend to congregate on window screens, front doors and siding of homes.

Description of Galls and Life Stages Galls appear as small swellings of yellow to yellow-green tissue on leaves or petioles (Fig. 12.1). Adults are about 3 mm long and resemble tiny cicadas.

Pest Management Infestations can be prevented by application of a contact insecticide to foliage of hackberry trees shortly after leaves develop in spring (Drooz 1985, Colorado State University 2004).

Fig. 12.1 Foliage of hackberry, Celtis occidentalis, with galls of hackberry nipple gall, Pachypsylla celtidismamma (Fort Collins, Colorado, USA).

Phytolyma spp.

used as a substitute for teak, Tectona grandis, for furniture, boats and construction.

Psyllids of the genus Phytolyma consist of a complex of at least three species: P. fusca Walker, P. lata Walker and P. tuberculata (Alibert).

Distribution Species of Phytolyma are found in tropical Africa from Sierra Leone east to Tanzania. All three occur in Ghana.

Hosts Hosts are species of Milicia. P. lata is most commonly found on M. regia and P. fusca occurs more frequently on M. excelsia. Lumber cut from species of Milicia is of significant commercial value and commands high prices internationally. The lumber is often

Importance Nymphs produce galls on stems, shoots and leaves. When plant tissues become heavily infested, decay sets in and branch dieback occurs. Trees are rarely killed by early attacks. They may sprout auxiliary shoots, which are subsequently attacked. Sustained attacks cause deformity, growth loss and may eventually kill trees. Nursery seedlings and young, vigorous plantations suffer the greatest damage and in some cases 100% nursery and plantation failure has occurred.

Life History Eggs are deposited in rows or, less frequently, are scattered on buds, shoots or leaves and hatch about 8 days later. Instar I nymphs crawl over

Gall insects

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the plant surface, penetrate plant tissue and break down epidermal cells, which causes a fermentation of the leaf parenchyma and gall formation. Young leaves are preferred. Galls are formed within 1–2 days and completely enclose the nymph. Nymphs feed inside the gall tissue and undergo five instars over 2–3 weeks. When feeding is completed, galls become turgid, split and release adults. Period of development from egg to adult varies between 22 and 45 days, depending on species and location. Galls and adults are present throughout the year but tend to be less abundant from January to March and in September.

Description of Galls Leaf galls are globular to ovate in shape and occur most frequently on leaf midribs. Galls on stems or shoots tend to be somewhat oblong. Clusters of galls on young leaves or shoots may coalesce and become a mass of gall tissue (Fig. 12.2).

Pest Management Applications of systemic insecticides, either to the soil or applied directly to the galls, provide effective protection of trees. Other tactics include establishment of mixed species plantations or planting Milicia under a cover of other trees. Selection of genetic strains of Milicia resistant to attack also offers promise (Wagner et al. 2008).

Aphididae Mordwilkoja vagabunda (Walsh), Poplar Vagabond Gall Distribution This gall insect is a North American species, widely distributed across much of Canada and the USA. Populations of this insect have also been detected in Turkey.

Fig. 12.2 Galls on foliage of Milicia sp. caused by a gall forming psyllid of the genus Phytolyma (Ghana, photo by S. Sky Stephens, Colorado State Forest Service).

elongate into a branch. Heavy infestations affect entire trees. Infestations are often restricted to a few trees.

Life History Aphids feed on growing shoots of host trees in spring and form the gall. They vacate galls in mid-summer but return in fall to deposit eggs, which overwinter inside the galls.

Hosts Species of Populus are hosts. In southern USA, P. deltoides is a common hosts and further north, P. tremuloides is attacked. In Turkey, infestations have been detected on P. nigra. Alternate hosts include species of loosestrife, Lythrum spp.

Description of Gall Galls are large and convoluted, green or brown in color and 5–7 cm long. They occur at the tips of shoots.

Importance Colonies of feeding nymphs cause new shoots to become distorted and produce galls rather than

Pest Management If galls should become unsightly on ornamental trees, infestations can be treated with

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a spring application of an insecticide effective against aphids or they can be removed by hand clipping (Ostry et al. 1988, Colorado State University 2004, Toper Kagin & Yildiz 2010).

Adelges cooleyi (Gillette), Cooley Spruce Gall Adelgid (Plate 67) Distribution Cooley spruce gall aphid is native to western North America and has been introduced and become established in eastern North America and Europe.

Adelgidae Hosts Primary hosts are species of Picea. Alternate host is Pseudotsuga menziesii.

Adelges Of the over 35 species of Adelges recognized as of 2007, 17 are known to produce galls on species of Picea. They are found throughout the natural ranges of spruces in the northern hemisphere (Table 12.1, Havill & Foottit 2007).

Importance Galls are of little consequence under forest conditions. However, on seedlings and saplings in nurseries or ornamental trees, the insect can become

Table 12.1 Representative species of gall making species of Adelges (Hemiptera: Adelgidae), on Picea: their distribution and secondary hosts. Species

Distribution

Secondary host

A. abietis (Linnaeus) Eastern spruce gall aphid A. cooleyi (Gillette) Cooley spruce gall aphid A. glandulae (Zhang) A. isedakii Eichhorn A. japonicus Monzou A. knucheli (Schneider-Orelli & Schneider A. lariciatus (Patch)

Europe (indigenous) North America (introduced) Western North America (indigenous) Eastern North America, Europe (introduced) China Japan Japan Western Himalayas Eastern North America west to Alberta and Saskatchewan, Canada Europe Estonia, Finland, Russia (indigenous) Kyrgyzstan (introduced) Italy Caucasus (indigenous), now widely distributed across Europe Australia (Tasmania), New Zealand (introduced) Europe, China, Japan Caucasus Europe Japan Asia North America (introduced) Europe

None

A. laricis Vallot A. lapponicus Kholodkorsii A. nebrodensis (Binazzi & Covassi) A. nordmanniana (Eckstein)

A. pectinata (Cholodkovsky) A. prelli (Grossman) A. tardoides (Cholodkovsky) A. torii (Eichhorn) A. tsugae (Annand) Hemlock woolly adelgid A. viridis (Ratzeburg) 

Pseudotsuga menziesii (foliage) Abies Larix Larix kaempferi (foliage) Abies Larix laricina, L. lyallii (buds and cones) Larix decidua (foliage) None Abies Abies nordmanniana (foliage)

Abies Abies Larix Larix Tsuga spp. (shoots) Larix decidua (foliage)

See Chapter 11 for summary of this species. Populations established in North America fail to develop or produce galls on indigenous species of Picea. Sources: Furniss & Carolin 1977, Drooz 1985, Kolk & Staryk 1996, EPPO 2003, Havill & Foottit 2007, Canadian Food Inspection Agency 2008, Sano et al. 2008. 

Gall insects a pest because branch tips are killed, causing stunting and deformity. Moreover, large numbers of galls on ornamental trees are unsightly.

Life History Cooley spruce gall aphid is holocyclic and follows the example given in Chapter 6 (see Fig. 6.3). This insect requires two hosts, i.e. spruce and Douglas-fir, and goes through five generations which take 2 years to complete. In late summer–autumn, winged sexuparae fly from Douglas-fir to spruce, where they lay eggs and die. Their roof-like wings shelter the eggs, which develop into males and females of the sexuales generation. Sexuales feed near the body of their dead mothers and when mature, disperse toward the center ofthe tree to mate and lay eggs. Females deposit one egg, which hatches into a wingless fundatrix. The fundatrix nymph crawls to, or near, a bud and overwinters. Their feeding initiates gall formation. In spring, fundatrix adults deposit a large mass of eggs that hatch into gallicolae nymphs and migrate to the developing gall. They feed inside the gall and when development is complete, winged gallicolae

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adults either migrate to Douglas-fir to initiate the exulis and sexupara generations (Fig. 12.3) or remain on spruce to initiate anholocyclic fundatrix and gallicola generations.

Description of Galls and Life Stages Galls resemble miniature pineapples or small cones, occur on branch tips and may persist for years. They range from 12 to 75 mm in length and are light green to dark purple, later changing to brown after adults emerge. Most life stages are small and covered with a white waxy coating.

Pest Management High-value ornamental trees can be sprayed with either a horticultural oil or contact insecticide in late autumn to kill overwintering stages or in spring, at bud burst before galls form. Galls can be pruned from infested ornamental spruce (Furniss & Carolin 1977, Drooz 1985, Munson 2006, Havill & Foottit 2007).

Fig. 12.3 Woolly aphid generation of Cooley spruce gall aphid, Adelges cooleyi, on foliage of its secondary host, Douglas-fir, Pseudotsuga menziesii (Gallatin National Forest, Montana, USA).

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Table 12.2 Gall forming species of Pineus (Hemiptera: Adelgidae): their distribution and primary and secondary hosts. Species

Distribution

Primary host

Secondary host

P. armandicola Zhang, Zhong & Zhang P. cembrae (Cholodkovsky) P. floccus (Patch) Red spruce adelgid P. orientalis (Dreyfus) P. pinifoliae (Fitch) Pine leaf chermes

China Europe, China, Japan Eastern North America

Picea likiangensis Picea spp. Picea rubens P. mariana Picea orientalis Picea spp.

Pinus armandii Pinus koraiensis, Pinus spp. Pinus strobus

Caucasus Mountains North America

Pinus Pinus monticola, P. strobus

 lu 2003, Havill & Foottit 2007. Sources: Drooz 1985, Kaygin & Canak¸ ¸ ciog

Pineus Of the 29 species of Pineus recognized as of 2007, at least five species are holocyclic and have sexual generations that produce galls on spruce, Picea spp. (Table 12.2, Havill & Foottit 2007).

indicate that there is a threat of high populations on pine the following year.

Description of Gall Galls are small and conical in shape. They are green or purple when first formed and later turn red-brown.

Pineus pinifoliae (Fitch), Pine Leaf Chermes Distribution P. pinifoliae is a North America species and is found over most of the range of its hosts.

Hosts The sexual form attacks spruce, Picea spp., and asexual forms attack pines of the white or soft pine group, including western white pine, Pinus monticola, and eastern white pine, P. strobus.

Pest Management On ornamental spruce, galls can be hand pruned before mid-June and burned to reduce cosmetic damage to ornamental trees (Drooz 1985).

COLEOPTERA Cerambycidae Saperda populnea (Linnaeus), Small Poplar Borer

Importance Galls are formed on spruce, which can be unsightly on ornamental trees but of little consequence in forests.

Life History Four and sometimes five generations are part of the 2-year life cycle of this adelgid. On spruce, insects are hidden under bud scales or in galls. On Pinus strobus they are more readily seen. In June of the first year, winged adults fly from spruce to white pine, where they lay eggs on old needles and die. Wingless nymphs emerge, crawl to the axes of newly expanding shoots, insert their mouthparts and begin feeding. They mature in May of the second year and fly back to spruce to lay eggs. Eggs on spruce hatch in June and nymphs feed on needles. When mature, they mate and the eggs produce the insects that hibernate under spruce bud scales and later form galls. Heavy gall production on spruce may

Distribution This insect is indigenous to Eurasia from western Europe to China. It has been introduced and become established in much of western North America from British Columbia, Canada, south to Arizona, California and New Mexico, USA.

Hosts Hosts are species of poplar, Populus spp., and willow, Salix spp.

Importance Larvae bore into twigs and cause galls at the point of infestation. Small poplar borer is one of the most injurious and widespread pests of poplars and, less frequently, willows in parts of Europe, China and Russia. This insect is reported as a pest of Populus nigra plantations and also of natural forests of P. tremula in Turkey.

Gall insects Life History In the UK, 2 years are usually required to complete a generation but some individuals can complete a generation in 1 year while others may need 3 years. Adults are active from May to July and lay eggs on the bark surface of host trees.

Description of Galls and Life Stages Galls are globose and appear on stems and branches. Adults are 10–15 mm long, have slim, gray to black, heavily punctated bodies and appendages. The dorsal surface of the head and thorax is rimmed with pale orange pubescence and the elytra are marked with spots of pale orange pubescence.

Related Species S. inornata Say and S. concolor LeConte are North American gall making species of Saperda. Larvae bore in wood of young poplars and produce globose galls on stems and branches. Damage is worst to nursery whips and 1–3- year-old trees in plantations. Affected branches occasionally break off or die above the gall. Most trees, however, overgrow the galls and injured trees tend to recover height growth within 2–3 years. Females lay eggs in spring in niches cut in the bole and branches. A single larva usually develops in each niche and bores under the bark, causing a gall to form around the injured area. Larva bore into the center of the gall in late summer, where they pupate and overwinter (Browne 1968, Furniss & Carolin 1977, Ostry et al. 1988, Abgrall & Soutrenon 1991, Solomon  zbek et al. 2009). 1995, Schmutzenhofer et al. 1996, O

HYMENOPTERA Tenthredinidae Sawflies of the genera Euura, Phyllocolpa and Pontania of family Tenthredinidae produce galls on willows and poplars. Euura spp. produce closed galls on buds, petioles and stems of willows. Phyllocolpa spp. produce open galls by deforming leaf margins of host trees. Pontania spp. produce closed galls on Salix (Smith 1968).

Euura spp. Distribution Euura is a large genus of gall making sawflies found ithroughout the northern hemisphere.

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Hosts Species of willow, Salix spp. are hosts. Most species of Euura confine their attacks to a single species of Salix and some species may even show preference for certain clones. E. shibayanagii Togahsi, for example, a Japanese species, is found only on S. japonica. An exception is E. mucronata (Hartig), a species with a holarctic distribution that produces galls on stems of at least 30 species of willows. Some specialists believe, however, that E. mucronata may be a complex of several closely related species.

Importance Larvae form galls on stems, twigs, petioles or buds of willows. Members of the subgenus Gemmura produce bud galls and members of the subgenus Euura produce galls on petioles or stems. Galls can be abundant in willow stands but they do not appear to cause significant injury. E. mucronata prefers to attack plants with long shoots.

Life History These sawflies typically have one generation/year. Adults appear in spring and mating and oviposition occurs within 1–2 hours after emergence. Larvae feed on plant tissue, become enveloped in galls and spend the entire growing season feeding. Sometimes more than one larva will occur in a gall. Some species overwinter in the galls as prepupae and others drop to the ground and overwinter in cocoons in the soil near the base of the plant.

Description of Gall and Life Stages Gall characteristics vary according to species. Bud galls tend to have a swollen appearance. Stem galls vary from narrow to bulbous to elongate. Some tend to be white and others are dark green, red or red-brown. Adults range in size from 3 to 8 mm and are black or a combination of black and amber in color. Larvae are yellow to light green in color with black eyes and a brown head capsule.

Pest Management Since galls appear to have little or no effect on tree health, control methods have not been developed (Smith 1968, Togashi 1980, Drooz 1985, Price et al. 1987, Nyman 2002, Colorado State University 2004).

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Pontania proxima (Lepeltier), Willow Red Gall Sawfly Distribution This sawfly has a wide distribution. Records of its occurrence include Australia (Tasmania), Europe, India and North America.

Hosts

Hosts are willow, Salix spp.

Importance Galls are produced on leaves. The galls are conspicuous but appear to have little effect on tree health. They are more of a curiosity than a pest.

Life History In most locations there are two generations/year. Adults emerge and deposit eggs in spring on young expanding leaves. Females also deposit a chemical into the leaf tissue that stimulates gall formation. Larvae feed within the galls and when fully grown, drop to the ground and spin cocoons. Some individuals pupate and later merge as adults to produce a second generation. Others remain dormant until the following season.

Description of Stages Galls are round, red and often occur in clusters. Adults are small black wasps.

Pest Management Developing larvae are often heavily parasitized. Other insects may also utilize galls for their development and kill the sawfly larvae. No pest management methods have been developed for this insect (Smith 1968, Naumann et al. 2002, Colorado State University 2004).

Eulophidae

Hosts Host plants are species of Erythrina. Approximately 110 species are known worldwide, mostly from the tropics. Common names are tiger’s claw, Indian coral and wiliwili-haole. They are used as ornamentals, living fences and as nitrogen fixing plants in agroforestry. Species of Erythrina have showy red flowers, which have special meanings to indigenous cultures. In some cultures the red flowers signal the arrival of spring and serve as a working calendar. In others it signals the time to plant sweet potatoes. In coastal cultures, occurrence of Erythrina blossoms provides a sign that it is time to catch flying fish.

Importance Larvae induce gall formation on leaflets and petioles of host plants. As infestations progress, leaves curl and appear deformed while petioles and shoots become swollen. Heavily galled leaves and stems result in growth loss and reduced vigor.

Life History This insect has multiple generations and period of development from egg to adult is about 20 days. Sex ratio of adults is 7 : 1 males: females. Females carry an average of 320 eggs. Eggs are inserted into young leaf and stem tissue. A gall forms as larvae develop in plant tissue. Pupation occurs inside the gall and adult wasps cut exit holes through the gall and emerge.

Description of Galls and Stages Tiny green galls occur in large numbers on the foliage and leaf petioles of infested trees. They are globular in shape and thick walled. Affected leaves are curled and deformed. Females are 1.45–1.6 mm long, dark brown with yellow markings. The head is yellow and antennae are pale brown. Males are smaller, 1.0–1.15 mm long and color of body, head and antennae is white to pale yellow as opposed to yellow in female.

Quadrastichus erythrinae Kim, Erythrina Gall Wasp Distribution This gall wasp is believed indigenous to Africa but its origin is uncertain. It was described in 2004 from specimens collected in Mauritius, Reunion and Singapore. Since its original description, infestations have been detected in China, Florida and Hawaii, USA, India, the Philippines and Taiwan.

Pest Management Attempts to eradicate localized infestations have been unsuccessful. Removal of Erythrina trees growing near ports has been suggested as a means to prevent establishment of this insect. Pruning of gall infested branches has met with some success. Systemic insecticides provide effective short-term control of infestations. Exploration for natural enemies to

Gall insects be used in classic biological control programs is underway (ISSG 2006, Wiley & Skelley 2006). Leptocybe invasa Fisher & LaSalle, Blue Gum Chalcid Distribution This species was described from the Near East in 2004. It is believed to have originated in Australia but has not yet been collected there. It is presently known from Africa (Algeria, Kenya, Morocco, Tanzania and Uganda), Asia (Thailand, Vietnam), Europe (France, Italy, Portugal and Spain), the Near East (Iran, Israel, Jordan, Syria and Turkey) and USA (Florida).

Hosts Known hosts are Eucalyptus spp., including E. botryoides, E. bridgesiana, E. camaldulensis, E. globulus, E. grandis, E. gunnii, E. robusta, E. saligna, E. tereticornis and E. viminalis.

Importance L. invasa produces galls on midribs, petioles and stems of new shoots. Heavy infestations cause deformed leaves and shoots, resulting in growth loss. Damage to young plantations and nursery seedlings has been reported but tree mortality has not been observed.

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Pest Management No pest management tactics are currently available for this insect. Classic biological control is being investigated (Mendel et al. 2004, EPPO 2008, FAO 2009b). Ophelimus Ophelimus is an Australian genus, which contains over 50 described species and others, which are yet to be described. In addition, there are uncertainties regarding the taxonomy, biology, distribution, host range and natural enemies of this genus. Species have been assumed to be gall inducers and this is probably the largest group of gall inducing eulophids on Eucalyptus. The biology is known for only a few species and there is some indication that at least some species are not true gall inducers but are associated with galls in other ways, possibly as parasitoids (La Salle et al. 2009). Ophelimus eucalypti (Gahan), Eucalyptus Gall Wasp Distribution This gall wasp is undoubtedly native to Australia but, ironically, the insect has not been recorded there. It was described from near Wellington, New Zealand in 1922. In 1987, infestations were discovered in eucalypt plantations on the North Island and it has since spread throughout the North Island and portions of the South Island. It has also been introduced into the Mediterranean region of Europe: Greece (2002), Italy (2000), Spain (2003); also into the Near East (Iran, Israel) and Africa (Kenya, Morocco, Uganda).

Life History In Iran, Israel and Turkey, two to three overlapping generations occur per year. Female adults insert eggs in the epidermis of young leaves on both sides of the midrib, in the petioles and twigs within 1–2 weeks of bud break. Larvae develop inside of the galls and adults emerge, leaving round exit holes. With the exception of one record of males in Turkey, reproduction is by parthenogenesis. Average time of development time from egg to adult is 132.6 days at room temperature.

Hosts Several eucalypts are hosts. In New Zealand, Eucalyptus botryoides, E. deanei, E. grandis and E. saligna are attacked. E. camaldulensis and E. globulus are also reported as hosts.

Description of Galls and Life Stages Galls are green at first and later become pink. Surface of the galls is glossy. After wasps emerge, galls turn light brown on foliage and red on stems and loose their glossy surface. Mean length of a gall containing a single wasp is 2.1 mm. Adults (females) are 1.1–1.4 mm long with black bodies and lighter colored antennae and legs.

Importance On E. globulus, galls are produced on branches and midribs. In New Zealand, galls are produced on leaves of E. botyroides and E. saligna and the insect may be a different biotype. The heavily galled leaves abscise from the tree, and cause widespread defoliation, growth loss and loss of vigor. To date, populations in the Mediterranean region have not been damaging.

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Life History In New Zealand, this insect is believed to have two overlapping generations/year. Adults are active in August and December–January. They live for about 10 days. Females deposit a single egg in the side of the leaf and galls develop in rows. One female can deposit about 350 eggs. Female larvae produce a circular protruding gall and male larvae produce a pit gall. About 3 months are required for larval development. Galls are green when the larvae are young and turn brown as the larvae mature.

Description of Galls and Life Stages Female larvae of O. eucalypti induce circular, protruding galls on the leaves of E. botryoides and E. saligna, whereas the males induce pit galls on the same species. Adults are 1.3–1.7 mm long and have a black and yellow body, gray antennae and hyaline wings (Withers et al. 2000, Raman & Withers 2003, EPPO 2006).

in some manner. Two species of Selitrichodes are parasitoids of the blue gum chalcid, Leptocybe invasa. S. globulus is the first member of this genus known to induce galls. Little is presently known about its life history and habits.

Description of Galls and Stages Galls are brown and occur on branches and, occasionally, on leaves. They consist of multiple chambers in which larvae and pupae develop. In heavy infestations, they occur continually with up to 20 galls/5 mm of branch. Adults are small wasps. Females are 0.95–1.5 mm long and males are 0.85 mm long. The color of head, thorax and abdomen is dark brown with a few pales yellow area. Legs and antennae are pale yellow with brown tips. Eyes are large and red-brown in color (La Salle et al. 2009).

Cynipidae Selitrichodes globulus La Salle & Gates, Blue Gum Gall Wasp Distribution Blue gum gall wasp was discovered in Los Angeles County, California, USA in November 2008. This species is believed to be Australian in origin but is yet to be found in what is thought to be its native range.

Hosts Blue gum, Eucalyptus globulus, is the only reported host.

Importance This insect forms numerous small galls on branches. Galls are unsightly and heavy infestations lead to tree decline. Heavily infested branches dry and crack and could serve as sites for invasion of pathogenic fungi. This insect has the potential to spread into other parts of the world where blue gum is widely planted in forest plantations. In California, blue gum is widely planted as an ornamental but has become naturalized in some areas where it is considered invasive. This gall wasp has the potential to serve as a biological control agent in those areas where the tree is invasive.

Life History Selitrichodes consists of 12 described species and is believed to contain other, as yet undescribed, species. Most appear to be associated with galls

Amphibolips confluenta (Harris), Oak-Apple Gall Distribution Several species of wasps of the family Cynipidae produce large robust galls on oaks and are known as oak apples. Amphibolips confluenta is one of the more commonly occurring North American species and is found wherever its host plants occur naturally or have been planted.

Hosts Hosts are species of Quercus. Species of the red oak group, such as Q. coccinea, Q. rubra and Q. velutina, are preferred.

Importance Oak apples are one of the most common galls of oak in North America. They cause no permanent injury to their host trees. Heavy infestations on ornamental trees may be unsightly but this gall is more of a curiosity than a pest.

Life History Adult wasps hatch from galls in June and July. Males and females mate and then drop to the ground. Female wasps burrow in the soil at the base of host trees and inject eggs into the roots. Larvae hatch and feed on the roots for over 1 year before becoming pupae. Only wingless female wasps hatch from the

Gall insects subterranean pupae. They crawl up trunks of host trees in early spring and inject an egg into the midrib of a new leaf. Larvae are small and round. As they grow, they cause a chemical reaction inside the leaf that forms a gall around the larvae. As larvae feed and grow, galls increase in size.

Description of Gall and Life Stages Galls are 12–50 mm in diameter and filled with a fibrous mass (Plate 68). They are produced on the midribs or petioles of leaves. Galls formed during spring are green, but become light brown when dry with a thin, papery shell. Adults are small, dark wasps with an oval, compressed abdomen. Larvae are small and globe shaped.

Pest Management Natural enemies are usually sufficient to keep populations under control. If galls are abundant and unsightly on ornamental trees, they can be hand picked and destroyed.

Related Species Andricus californicus (Bassett) occurs on Quercus garryana in the Siskiyou Mountains of southern Oregon and northern California. Callirhytis quercuspomiformis (Bassett) forms large apple-like galls with a gnarled surface on California live oak, Q. agrifolia. Another species, Trichoteras vaccinifoliae (Ashmead) forms conspicuous galls on canyon live oak, Quercus chrysolepis. Both species are found in California, USA. In Europe, oak apples are caused by Biorhiza pallida (Olivier) (McCracken & Egbert 1922, Drooz 1985, Solomon et al. 1999).

Cynips Cynips is a large genus of gall wasps. Many species form galls on foliage and stems of Quercus. They have alternating sexual and asexual generations. Both adults and the galls produced are morphologically distinct. Early classifications of the genus Cynips often recognized these generations as separate species. Adults of the sexual generation range in size from 2.2 to 2.7 mm and females are slightly larger than males. Body color is brown to black and legs tend to be yellow to dark yellow. The antennae are long and have 14 segments. Adults of the asexual generation tend to be somewhat

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larger and range in length from 2.2 to 4.0 mm. Body color ranges from orange to dark brown with a black underside. Legs are chestnut-brown. Cynips gallae tinctoria, Aleppo Gall Distribution Aleppo gall, also known as Turkey gall, Levant gall, gall-nut, gall of commerce and ink marble, is found in eastern Europe and the Near East, including portions of Greece, Hungary, Iraq and Turkey.

Hosts Hosts are species of Quercus, including Q. aegilops, Q. infectoria, Q. pedunculata and others.

Importance This insect produces galls on stems and branches. Although galls may be unsightly on ornamental trees, they have been of commercial importance since the time of ancient Greece and have been used for a variety of purposes. The primary use of Aleppo gall was in the manufacture of ink known as iron gallotannate ink or iron-gall ink. Aleppo gall has high tannic acid content and also contains gallic acid. Gallic acid produced permanent inks that did not fade. When durable ink was required, as is in the case of legal documents, local laws often required that the records be made of inks produced from Aleppo gall. Ninth and tenth century monks used ink made from the galls of this insect. As late as the 20th century, inks purchased by the Treasury of the United States, Bank of England, the German Chancellery and the Danish Government were required to be made from acids extracted from the Aleppo gall. A permanent black dye, used to dye leather, has also been produced from Aleppo galls. In 1914, US$ 17,000 worth of Aleppo gall nuts were imported from Baghdad to dye sealskins. Women also used this dye to color their hair black.

Description of Galls Galls are spherical, hard and brittle, dark blue-green or olive green, nearly spherical in shape and 12–18 mm in diameter.

Related Species Galls produced by C. insana Westwood, on Quercus infectoria, were a source of a dye known as “Turkey red” that was used in Basra, Iraq.

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These galls are known as the mad apple of Sodum, Dead Sea fruit or the Mecca or Bissorah gall (Fagan 1918, Felt 1940, Grieve 1971, Berenbaum 1995, Ciesla 2002a). Cynips quercusfolii Linnaeus, Oak Bud Gall Wasp, Cherry Gall

Disholcaspis quercusmamma (Walsh), Rough Oak Bullet Gall Wasp (Plate 69) Distribution This gall wasp occurs in central USA as far west as Colorado.

Hosts Host trees are swamp white oak, Quercus bicolor and burr oak, Q. macrocarpa.

Distribution This insect is widely distributed across Asia, the Near East and Europe.

Hosts Species of oaks, Quercus spp., are hosts. In the UK, this insect is often abundant on Q. petraea and Q. robur.

Importance Galls are formed on buds and leaves of host plants. Damage is largely cosmetic in nature and it is regarded as more of a curiosity than a pest.

Life History This insect has two generations/year, i.e. a sexual generation and an asexual generation. In spring, larvae of the sexual generation form galls on dormant buds of host trees. In late summer, the asexual generation forms galls on veins of the lower surfaces of expanded leaves.

Description of Galls Galls of the sexual stage are small, ovoid, dark red to purple when first formed and later turn black. Their surfaces are covered with small, pointed scales. Galls of the asexual generation have an average diameter of 17 mm and spherical in shape. They are yellow or green, marked with pink or red when first formed and turn brown when mature. The common name “oak bud gall wasp” refers to the sexual generation and “cherry gall” to the asexual generation.

Related Species Two other species of Cynips, C. divisa Hartig and C. longiventris Hartig, are also found on species of Quercus across Eurasian oak forests. Both have alternating sexual and asexual generations. The asexual generation of C. divisa produces round galls, about 8 mm in diameter, pale yellow with red-brown markings and known as the red pea gall (Browne 1968).

Importance Round, woody galls are produced on twigs and branches of host trees. Galls remain on branches for several years after the insects have completed their development and are most conspicuous during winter when foliage is not present. In some cases, branches of host trees are covered with galls, which are unsightly.

Life History Life history of this insect is not well understood. There may be more than one generation/ year. Adults are active in fall and deposit eggs on twigs and branches. Eggs hatch in spring and larval feeding stimulates gall production. There is one larva/gall. Description of Gall and Life Stages Galls first appear as small green eruptions or bumps. Their color changes to red and later dark brown as insects mature. Completely formed galls are rounded with a point at the apex and 8–15 mm long. Wasps are 2–3 mm long, black to brown and ant-like in appearance. Larvae are white and legless and the head is indistinct.

Related Species D. cinerosa Bassett produces galls on Quercus fusiforme and Q. virginiana in southeastern Louisiana, Oklahoma and Texas, USA. This insect has two generations/year, one sexual and the other asexual. Galls of the spring–early summer sexual generation are relatively inconspicuous. The late summer galls produced by the asexual generation are large, range in size from 3 to 25 mm and are known as mealy oak galls (Morgan & Frankie 1982, Eckberg & Cranshaw 1995). Dryocosmus kuriphilus Yamamatsu, Chestnut Gall Wasp (Plate 70) Distribution Chestnut gall wasp is native to China. It was introduced into Japan in the early 1940s and

Gall insects Korea in the early 1960s. In 1974, infestations of this species were detected in Georgia, USA and it has since spread into adjoining states. This insect has also been detected in northwestern Italy, where it is spreading rapidly.

Hosts The vegetative buds of species of Castanea that produce three nuts per burr are attacked by this gall wasp. In China, C. mollissima is attacked and in Korea and Japan, C. crenata and C. crenata  C. mollissima are hosts. In the USA, C. mollissima and the native C. dentata are attacked and in northwestern Italy the host is C. sativa. Gall formation has not been detected on North American species of Castanea that produce one nut per burr, the chinkapins, such as C. pumila and C. alnifolia.

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Pest Management The parasitoid complex of this species is well known. In Japan, a classic biological control program, which involved release of the parasitoid Torymus sinensis, resulted in its establishment and spread. Ten years after release, gall formation in chestnut buds was significantly reduced. The same species has been released in northwestern Italy for biological control of this insect (Payne 1978, Moriya et al. 1989, Wang et al. 2001, Quacchia et al. 2008).

DIPTERA Cecidomyiidae Dasineura

Importance Galls on vegetative buds disrupt shoot growth, suppress shoot elongation and reportedly reduce fruiting. Observations in its native habitat in China on Castanea mollissima by the author, however, indicate that most galls are produced in the inner crowns of host trees, whereas chestnuts are produced on the outermost branches, i.e. those exposed to direct sunlight. These observations suggest that, at least where the insect is native, its impact on chestnut production is questionable.

Life History Chestnut gall wasp has one generation/ year. Overwintering takes place as instar I larvae inside buds of host trees. In spring, when buds normally begin to break, they develop into a rose-colored gall about 8–15 mm in diameter. Larvae feed inside the gall for 20–30 days and pupate. Adults, all of which are females, emerge in late May–early June. They deposit from three to five eggs in a cluster inside buds. More than one adult may deposit eggs in the same bud and some buds are known to contain 10–25 eggs. Larvae hatch within about 40 days, usually in late July, and larval development is slow throughout autumn and winter.

Description of Stages Adults are black wasps about 3 mm long. Eggs are oval, milky white and 0.1–0.2 mm long. Larvae are 2.5 mm long when mature and milky white in color. Pupae are black and 2.5 mm long.

Dasineura is a large genus of gall midges with at least 300 known species. Larvae produce galls on many plants of importance in agriculture, forestry and horticulture.

Dasineura spp., Larch Gall Midges Distribution Larch gall midges are a complex of four species found across Europe and Asia that have similar habits and cause similar damage (Table 12.3).

Hosts Hosts are species of larch, Larix spp., found within the respective ranges of each species of larch gall midge (Table 12.3).

Importance Galls are produced in buds of host trees and kill buds. As a result, trees are unable to produce flower buds and seed yields are either significantly reduced or non-existent. Trees are not killed by infestations and new twigs are formed on gall bearing branches, which are subject to attack in subsequent years. Repeated heavy infestations can cause reductions in both radial and height growth. In Siberia, damage was first noted during the late 1960s when seed orchards were established. In some instances, up to 90% of buds were killed in seed orchards by D. rozhkovi. In Poland infestation levels of D. kellneri vary between 4.4% and 20.9%.

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Table 12.3 Species of larch bud moths, Dasineura spp. (Diptera: Cecidomyiidae): their distribution and hosts. Species

Distribution

Hosts

D. kellneri Henschel D. rozhkovi Mam. et. Nik.

Central Europe Russia, southern and eastern Siberia and northeastern Mongolia Russia, southern Siberia Japan

Larix decidua Larix czekanowski, L. gmelinii L. sibirica

D. verae Skuhrava D. nipponica Inouye

Larix gmelinii, L. sibirica L. kaempferi

Source: Baranchikov 2006.

Life History All species have one generation/year. Adults are active from April to May, depending on temperature, which coincides with the development of larch needles. Up to 60% of the adults may hatch in 1 day. Studies on the life history of D. rozhkovi in Siberia (reported as D. laricis) indicate that there are approximately twice as many females as males. Males live from 1 to 3 days and females live somewhat longer. Females have an average of 60–85 eggs. They deposit eggs singly between needles and bract scales on the buds of spur shoots of larch. Larvae hatch within 6–9 days and crawl between the needles to the next year’s buds and bore inside the bud. Larval feeding causes a gall to form. Galls become visible in June and a single larva feeds inside each gall. In late August, instar IV larvae leave the gall and weave thick white cocoons between external gall scales. Larvae overwinter in the cocoon and pupate in late May of the following year. Adults emerge 5–7 days later. Description of Gall and Life Stages Galls are 5–12 mm  4–8 mm, light brown and shaped like a miniature artichoke. Instar IV larvae are pale orange and are 3–3.2 mm long. Pest Management Bud mortality can be reduced through application of contact and systemic insecticides. Timing is critical if contact insecticides are used because of the cryptic nature of the insect and must be applied during the first week of adult emergence. Direct control in seed orchards must be done at least once every 3 years to keep populations at a tolerable level (Isaev et al. 1988, Baranchikov 2006, Skrzypczynska 2007).

Oligotrophus betheli Felt, Juniper Tip Midge Distribution This midge is a North America species found in central and western USA and British Columbia, Canada.

Hosts Species of Juniperus are infested, including J. occidentalis, J. osteosperma, J. scopulorum and J. virginiana. Infestations are especially common on J. osteosperma in Colorado and Utah, USA. Importance Apical conical galls are formed on the tips of host plants. Infestations are of little consequence in forested areas but may be unsightly on ornamental trees.

Description of Gall Galls occur at branch tips and vaguely resemble a flower or small cone. They are blue green when first formed in spring/early summer and turn red-brown toward the end of the growing season (Fig. 12.4).

Related Species O. juniperinus (Linnaeus), O. panteli (Kieffer) and O. gemmarum (Rübsammen) produce galls on Juniperus in Europe and O. nezu (Kikuti) causes similar damage in Japan. O. apicis Appleby & Neiswander damages tips of Juniperus in central USA (Appleby & Neiswanger 1965, Furniss & Carolin 1977, Harris et al. 2006).

Oligotrophus

Pinyonia edulicola (Gagne), Piñon Spindle Gall Midge

Members of this genus infest both broadleaf trees and conifers throughout the northern hemisphere. Several species form galls on branch tips of Juniperus spp.

Distribution Piñon spindle gall midge occurs in Colorado and possibly other states in southwestern USA.

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Description of Gall and Life Stages Galls are spindle shaped and develop at the base of the needles (Fig. 12.5). The portion of needle affected by the gall may become yellow or red. Adults are small, delicate flies, about 2.5 mm long, with an orange abdomen. Larvae are small, orange maggots Plate 71, (Furniss & Carolin 1977, Colorado State University 2004). Rhabdophaga strobiloides Osten Sacken, Willow Pine Cone Gall Distribution This species is native to North America and is found throughout much of the continent.

Hosts

Species of willow, Salix spp., are hosts.

Importance Larvae feed on terminal shoots and prevent stems from elongating. Feeding produces a compact gall. The galls cause no permanent injury but may be unsightly on ornamental trees.

Life History There is one generation/year. Adults emerge in late April–early May and deposit eggs singly on the terminal buds of willow just as the buds begin to swell. Feeding by the larva causes buds to swell as it develops. Larvae overwinter in the gall and pupate in early spring. Fig. 12.4 Gall on branch of Utah juniper, Juniperus osteosperma, caused by the juniper tip midge, Oligotrophus betheli (Colorado National Monument, Colorado, USA).

Hosts Host is piñon pine, Pinus edulis. Galls can also form on bristlecone pine, P. aristata, but insects do not complete development.

Description of Gall Galls are rosettes of tightly clustered leaves that resemble a pine cone (Fig. 12.6). They are green at first and later turn brown and may persist on trees for several seasons after the larva has completed development. Galls can be up to 25 mm in diameter.

Importance Galls are produced at the base of current year’s needles. Infested needles turn brown and drop prematurely. Damage is usually of a minor, cosmetic nature.

Related Species The European rosette gall midge, R. rosaria (Loew), produces a similar gall on Salix alba and other willows in Europe (Fig. 12.7, Colorado State University 2004, Koll ar 2007).

Life History This insect has one generation/year and overwinters as larvae in the galls. Each gall contains 5–40 larvae. Adults emerge in late June–early July.

Taxodiomyia cupressiananassa (Osten Sacken), Cypress Twig Gall Midge Distribution Cypress twig gall midge occurs in southern and central USA and has been reported from

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Fig. 12.5 Open gall of piñon spindle gall midge Pinyonia edulicola showing orange larvae (Fort Collins, CO, USA).

Alabama, Florida, Illinois, Indiana, Louisiana and Tennessee.

Hosts Bald cypress, Taxodium distichum, is the only known host.

Importance This insect produces galls on branch tips of host trees. There are some indications that certain trees are more susceptible to attack than others. Galls cause little or no damage but are unsightly on ornamental trees. In addition, heavily infested branches tend to droop under the weight of the galls.

Life History Cypress twig gall midge has two generations/year (Fig. 12.8). Adults emerge beginning in mid-May and emergence can occur over an extended period. However, about 95% of the adults emerge over a period of 3 weeks. Mating usually occurs on the same day as emergence and females deposit eggs on the

developing foliage of host trees. They deposit an average of 120 eggs during their 1–2 day life span and young larvae induce gall formation. Galls first appear as small pink swellings of the branchlets at the feeding site and increase in size rapidly. Larvae gradually move toward the longitudinal axis of the gall and construct a small chamber. Each gall contains an average of 16 larvae (range 1–30) and larger galls tend to produce a greater number of larvae. Adults of the first generation emerge from late July to mid-September and second generation larvae feed until mid-September. In late summer, galls containing larvae fall from infested trees. Larvae overwinter inside galls and pupate in April.

Description of Gall and Life Stages Galls are oval and occur at the terminal portion of branchlets. Several needles of host trees protrude through the gall surface. When mature, they resemble tiny pineapples (Fig. 12.9). Color is initially pink and changes to light green or white because of a covering of fine powdery material on the gall surface. When adults emerge, galls

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Fig. 12.6 Willow pine cone gall caused by the midge, Rhabdophaga strobiloides (Roosevelt National Forest, Colorado, USA).

Fig. 12.7 Rosette galls on willow caused by the midge, Rhabdophaga rosaria (Weltenberg, Bavaria, Germany).

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Fig. 12.8 Seasonal history of cypress twig gall midge, Taxodiomyia cupressiananassa (redrawn from Chen & Appleby 1984).

turn a copper-brown hue. Galls are about 20–30 mm long, 20 cm wide. Adults are small flies with clear wings covered with short setae. The thorax and appendages of males are tan and the abdomen is tan to orange in color. Female abdomens are orange-red. Males average 1.57 mm in length and females are 2.17 mm long. Larvae are 0.6 mm long and 0.09 mm wide when first hatched and light orange in color. As they grow, they change body color to orange red. They are about 1.5 mm long and 0.07 mm wide when mature.

Thecodiplosis japonensis Uchida et Inouye, Pine Needle Gall Midge

Pest Management Infestations on ornamental trees can be treated by removal and destruction of galls before adults emerge. This reduces the number of galls produced during the following season (Chen & Appleby 1984, Gomez & Mizell 2009).

Importance Larvae produce galls at the base of needles and infested needles dry and become brown. Feeding damage causes growth loss and tree mortality. Outbreaks have occurred in the coastal regions of Japan and in Korea. T. japonensis is considered one of the most

Distribution This gall midge is native to Japan and was discovered in South Korea in 1929. The insect is now widespread over much of South Korea, including the island of Jeju. It is also present in North Korea.

Hosts Two pines, Pinus densiflora and P. thunbergii, are hosts.

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soon after emergence and deposit eggs on surfaces of the cavity between a pair of developing needles in a fascicle. Eggs hatch within a week after they are deposited and larvae move to the base of needles where they form galls. An individual gall can contain from 1 to 18 larvae. Larvae feed inside the galls and pass through three instars. Mature instar III larvae drop from the galls to the soil in late fall, crawl into the litter, spin cocoons and overwinter. Pupation takes place in April and adults emerge beginning in late May.

Description of Gall and Life Stages Spindleshaped galls are formed at the base of infested needles. Larvae are pale yellow and range from 1.81 to 2.25 mm long. Adults are small midges. The size of larvae, pupae and adults are larger in galls that produce a small number of individuals. Adults from galls with small numbers of individuals tend to produce more viable eggs.

Pest Management Severely damaged and dead trees have been harvested. Two parasitoids, Inostemma matsutama and I. seoulis, build up in heavily infested areas and after several years become significant natural control factors. Fig. 12.9 Galls on bald cypress, Taxodium distichum, caused by the cypress twig gall midge, Taxodiomyia cupressiananassa (New Iberia, Louisiana, USA).

destructive insect pests of pines within both its natural and introduced range, especially in Korea where more than half of forest area is composed of host pines. In South Korea, infestations tend to be most severe at the leading edge of the zone of infestation. They peak 6–7 years after initial establishment, then decline. After 12 years the proportion of infested foliage stabilizes at a low level. Infestations in South Korea peaked in 1961 when some 410,000 ha of pine forests were infested. In 1992, 212,000 ha of South Korea’s pine forests suffered damage.

Life History This midge has one generation/year. Adults emerge from late May to late July with a peak in mid-June. Adults live for about 1 day. Females mate

Related Species T. brachyptera Schw€ ag is a European species that forms galls on Pinus sylvestris but is not nearly as damaging as T. japonensis (Sone & Takeda 1983, Jeon et al. 1993, Lee & Lee 1993, Sone 1993).

Argomyzidae Hexomyza (¼ Melanagromyza) schineri (Giraud), Poplar Twig Gall Fly Distribution This insect has a circumpolar distribution and occurs across much of the northern hemisphere.

Hosts Poplars, Populus spp., and occasionally willows, Salix spp., are attacked. In North America, P. tremuloides is a favorite host.

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Fig. 12.10 Galls on quaking aspen, Populus tremuloides, caused by poplar twig gall fly, Hexomyza schineri (Fort Collins, Colorado, USA).

Importance Heavy infestations give host plants a gnarled, knobby appearance and can be unsightly on ornamental trees, especially during leaf-off periods. They also cause streaks or pith flecks in wood harvested from infested trees. Beginning in the 1980s, this insect reached epidemic proportions on ornamental trees in the vicinity of Denver, Colorado, USA and other communities along the eastern slopes of the Rocky Mountains.

Life History There is one generation/year. However, some individuals may have a second generation during warm years. Overwintering occurs inside the gall as mature larvae. Pupation takes place in late winter– early spring within the gall and later most pupae drop to the ground. Adults emerge as new shoots develop on host trees. During daytime, they may be seen resting on

leaves. After mating, females move to developing shoots where they deposit eggs inside the shoots. Larvae hatch and produce galls in response to their feeding.

Description of Gall and Life Stages Galls are spherical and occur on new shoots. They remain in place and continue to grow as the plant grows (Fig. 12.10). Adults are stout bodied, shiny dark flies about 4 mm long.

Pest Management Control of this insect on ornamental trees is difficult. Use of chemical insecticides has met with little success and infestations are often too heavy to allow hand pruning of galled branches to be practical (Browne 1968, Colorado State University 2004).

Chapter 13

Tip, Shoot and Regeneration Insects

INTRODUCTION Many insects, especially representatives of the orders Coleoptera and Lepidoptera, invade tips and shoots of trees and woody plants. Feeding by larvae kills the growing tips of affected trees and causes deformity, multiple branching and growth loss. Tip and shoot injury often occurs on young trees growing either in plantations or in natural stands. Other insects, especially weevils, strip bark from seedlings and kill them. This chapter reviews insects that damage tips and shoots of trees and those that damage seedlings.

COLEOPTERA (BEETLES) Bostrichidae (Branch and Twig Borers) Apate Bostrichid beetles of the genus Apate occur primarily in the tropics but can also be found in some temperate forests. They breed in wood of dead and fallen trees and

reduce the sapwood to a fine dust. Adults bore in branches of live trees. Some species are pests of young trees and ornamentals.

Apate monachus Fabricius Distribution This twig beetle occurs throughout Africa, including Madagascar, and the Mediterranean region. In has been introduced into some islands of the Caribbean and Brazil.

Hosts Many temperate and tropical broadleaf trees are hosts, including Acer orientalis, Azadirachta indica, Dalbergia sissoo, Eucalyptus spp., Terminalia ivoriensis and others.

Importance Larvae develop in felled trees and timbers and reduce the sapwood to a fine dust. Adults tunnel in boles of young trees and small stems and branches of large trees. Feeding by adults either kills

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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branches or makes them susceptible to wind breakage. Young trees in plantations are especially susceptible to damage.

Importance Larvae bore in branches and construct galleries. Infested branches are either killed outright or break during high winds and heavy rains. The impact of this longhorn beetle is usually greater in plantations than in natural forests and on trees that are stressed.

Life History Adults construct short tunnels into the wood and deposit eggs. Larvae feed parallel to the grain and construct tunnels that may be up to 40 cm long and 1.3 cm wide. They consume all of the available sapwood and may enter the heartwood. Tunnels are tightly packed with fine frass. Pupations occur in an oval chamber in the sapwood. Adults may remain in their pupal chambers for 1–2 weeks before they bore a circular exit hole, emerge and feed in stems of living trees.

Life History Adults emerge and feed on branches of I. paraguariensis. Branches between 30 and 40 mm in diameter are preferred for adult feeding. Mating occurs shortly after emergence and females deposit eggs on branches 15–20 mm in diameter. Larvae bore in the branches, pupate and emerge.

Description of Stages Adults are elongated, somewhat cylindrical, dark brown beetles. The head is bent downward and barely visible when viewed from above.

Description of Stages Adults are 26 mm long and stout. The body color is variable but the typical pattern is white to gray-white. Legs are dark gray to black and the thorax and elytra have gray-black markings. The antennae have alternating dark and light segments (Fig. 13.1).

Pest Management To reduce damage to young trees, removal of dead wood, which can be used as breeding sites, from areas scheduled for tree planting is recommended.

Related Species A. terebrans Pallas occurs in subSaharan Africa and in the Near East and has also been introduced into portions of the Caribbean Basin and Brazil. Its habits are similar to A. monachus. A. indistincta Murray occurs in eastern and southern Africa where it breeds in the stems and branches of dead fallen trees of many species. Adults also injure young living trees by tunnelling into small stems and branches to feed (Browne 1968, Schabel 1996, Wagner et al. 2008).

Cerambycidae Hedypathes betulinus (Klug) Distribution This twig borer is found in northern Argentina, southern Brazil and Paraguay.

Hosts Ilex paraguariensis, the foliage of which is the source of yerba mate, a herbal tea and locally important non-wood forest product, is the only host.

Pest Management Direct control in yerba mate plantations usually involves collection and destruction of adults. Work is underway to develop a pathogenic nematode, Steinernema carpocapsae, for control of this borer (Guedes et al. 2000, FAO 2007c, Alves et al. 2009).

Curculionidae (Weevils or Snout Beetles) Hylobius Weevils of the genus Hylobius are indigenous to Eurasia and North America. Seven species are found in North America and four in Europe, some of which extend their ranges into Asia. At least four species occur in Japan. Most are associated with conifers of the family Pinaceae and several are important pests of young plantations. Breeding occurs in fresh stumps and roots of recently killed pines and other conifers, which causes no damage. Emerging adults feed on bark of young conifer seedlings, girdle and kill them. Several species are known to be vectors of fungi of the genus Leptographium, some of which are pathogenic (Table 13.1, Viiri 2004). A notable exception to the association of Hylobius weevils with conifers is H. transversovittatus Goeze. This

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Fig. 13.1 Adult Hedypathes betulinus on branch of yerba mate, Ilex paraguariensis (near Colombo, Parana State, Brazil). Table 13.1 Representative species of Hylobius (Coleoptera: Curculionidae): their distribution and hosts. Species

Distribution

Hosts

H. abietis Linnaeus, Large pine weevil H. albosparsus Boheman White spotted weevil H. angustus Faust

Eurasia

Picea, Pinus, Pseudotsuga menziesii

Asia including China, Japan, Korea, Mongolia and Russia Northern India and Pakistan

All Pinaceae. Larix and Pinus are preferred Cedrus deodara, Picea smithiana, Pinus griffithii Pinus, occasionally other Pinaceae, Thuja

H. pales (Herbst) Pales weevil H. piceus (DeGeer) H. radicis Buchanan Pine root collar weevil

H. rhizophagus Millers, Benjamin & Warner Pine root tip weevil H. transversovittatus Goeze

Eastern North America Central and northern Europe, Siberia, northern North America Southern Canada, Newfoundland west to Manitoba USA, northeastern states, west to Minnesota and south to Virginia North central USA and adjoining Canada

Larix, Pseudotsuga menziesii, occasionally Pinus Pinus

Europe, introduced into Canada and USA for biological control of purple loosestrife

Lythrum salicaria

Pinus banksiana, other Pinus spp.

Sources: Browne 1968, Drooz 1985, EPPO 2000, Kolk & Starzyk 1996, McAvoy et al. 2002.

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weevil is native to Europe and feeds on the foliage of purple loosestrife, Lythrum salicaria, considered a noxious weed in North America. This species has been introduced into Canada and the USA for biological control of this plant (McAvoy et al. 2002). Hylobius abietis Linnaeus, Large Pine Weevil Distribution Large pine weevil occurs across Eurasia. In Europe it is found from Spain and France east to European Russia. It was introduced into the British Isles during the early part of the 20th century. In Asia, it occurs across Asian Russia and east into China.

Hosts Adults feed on seedlings of pine, Pinus spp., spruce, Picea spp., and Douglas-fir, Pseudotsuga menziesii. Root systems of recently dead conifers and fresh cut stumps, especially pines, are used as breeding sites.

Importance Adults strip bark from young seedlings and kill them. This insect is of considerable importance in reforestation areas across Eurasia. In northern and

western Europe, it is considered the single most important reforestation pest.

Life History Development from egg to adult varies according to altitude and latitude. In the UK, there are two generations/year but further north there may be only a single generation. Adults live up to 4 years and deposit eggs in late spring–early summer on exposed roots and root collars of dying conifers or on stumps of the previous year’s timber harvesting operations. Larvae bore under the bark, gradually widening their galleries and larvae of the second generation or adults overwinter. On roots with thin bark, galleries penetrate into the woody tissue. The following spring, they construct cells at the end of the larval galleries and pupate. Brood adults emerge in early summer and feed on the bark of conifer shoots and seedlings. They overwinter in the litter, mate and lay eggs the following spring (Fig. 13.2).

Description of Stages Adults are 7–14 mm long and the head has a long snout characteristic of weevils.

Fig. 13.2 Life history of the weevil, Hylobius abietis (Coleoptera. Curculionidae) (redrawn from Abgrall & Soutrenon 1991).

Tip, shoot and regeneration insects The body color is dark brown. Yellow-gold scales form three transverse, irregular, slightly curved stripes on the elytra. These markings tend to fade on older specimens.

Pest Management Reforestation of harvested conifer forests should be postponed for 2–3 years to avoid serious damage to seedlings. Chemical control involves dipping seedling in an insecticide or spraying seedlings prior to planting. Other tactics include digging ditches with vertical sides around young seedlings for trapping and destruction of adults or attracting adults with fresh pieces of conifer bark or pit traps packed with fresh conifer branches sprayed with insecticide (Browne 1968, Heritage et al. 1989, Abgrall & Soutrenon 1991, Day et al. 2004). Hylobius pales (Herbst), Pales Weevil Distribution Pales weevil is essentially the North American counterpart of H. abietis and is found throughout North America east of the Great Plains and north to Ontario, Canada.

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are March–May and July–August. After a brief period of feeding, adults fly to recently cut, damaged or recently dead pines. They mate and females lay eggs in the roots. Larvae feed downward in long galleries and pupate in cells in the outer sapwood. Pupation and adult emergence may take place in late summer or autumn, or larvae may overwinter and pupate the following spring.

Description of Stages Adults are large weevils, 7–12 mm long, dark red-brown with tufts of yellowwhite hairs on the elytra and thorax.

Pest Management Cultural control of pales weevil involves delay of planting of cut over areas for 1 year in the south and 1–2 years in the north, or until weevils have completed their life cycle and left the area. Seedlings are dipped in a solution containing a contact insecticide prior to planting or there is application of a spray after planting (Nord et al. 1984, Drooz 1985). Pissodes

Hosts This weevil breeds in all species of pine that occur within its natural range. Favorite hosts are Pinus echinata, P. rigida, P. strobus and P. taeda. It has also been reported on species of Abies, Larix, Picea, Pseudotsuga, Thuja and Tsuga.

Importance Adults feed on bark of young seedlings. Pales weevil is the most serious pest of pine seedlings on recently cut over sites, reforested areas or Christmas tree plantations. Mortality of seedlings in plantations of 30–60% is not uncommon and mortality rates of over 90% have been reported.

Life History Life history and habits vary according to location. In northern locations, there is usually one generation/year but further south there may be a second generation. In the north, adults overwinter in the litter and larvae overwinter in the roots of freshly cut stumps. In the south, adults may be active throughout the winter but do not reproduce. Adults emerge from March to June. They are nocturnal and feed on young conifer seedlings. Peak periods of adult activity

Pissodes consists of 47 known species widely distributed across Eurasia and North and Central America. Twenty-nine species occur in the western hemisphere and 18 in Eurasia (Table 13.2). Adults are brown, covered with scales that form patterns of white or yellow spots on the elytra and range in length from 8 to10 mm. Eggs are oval, yellow-white and about 0.43 mm long. Larvae are crescent or C-shaped legless grubs with light brown heads. Pupae are white and turn dark brown before they develop into adults. All are associated with trees of the family Pinaceae. They bore in the cambium and sometimes in the pith of host trees. One species, P. validirostris (Sahlberg), feeds on cones of Pinus (see Chapter 14). Most species attack weakened, dying or recently dead trees. Several are damaging pests and tend to damage young trees (Langor et al. 1999, Xiuxia Lu et al. 2007).

Pissodes strobi Peck, White Pine Weevil Distribution This insect is native to most of Canada and the northern portions of the USA, south to northern Georgia and the central Rocky Mountains.

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Table 13.2 Representative species of Pissodes (Coleoptera: Curculionidae): their distribution and hosts. Species

Distribution

Hosts

P. approximatus Hopkins Northern pine weevil

Eastern North America, west to Manitoba, Canada, and Minnesota, USA and south to North Carolina, USA Europe, Asian Russia

Picea, Pinus

Pinus, rarely Larix and Picea

Central and northern Europe, Siberia Southeastern USA

Picea Cedrus, Pinus

Northern Africa, western Asia, Europe

Pinus, also Larix decidua and Picea abies Abies, Picea Pinus

P. castaneus (De Geer) (¼ P. notatus Fabricius) P. hercyniae (Herbst) P. nemorensis Germar Deodar weevil P. notatus Fabricius Banded pine weevil P. piceae (Illiger) P. pini (Linnaeus) P. radiatae Hopkins Monterrey pine weevil P. strobi Peck White pine weevil P. terminalis Hopping Lodgepole terminal weevil P. validrostris (C.R. Sahlberg) P. yunnanensis (Langor & Zhank) Yunnan pine weevil

Europe, Asian Russia China, Japan, Kazakhstan, Korea, Russia, Europe California, USA, introduced into Uruguay-initially misidentified as P. castaneus Canada, northern and western USA Northern and western Canada, western USA China, Russia, Europe Southwestern China

Pinus attenuata, P.contorta, P. muricata, P. radiata Picea, Pinus Pinus banksiana, P. contorta Pinus spp. (cones) Pinus yunnanensis

e 2007, Xiuxia Lu Sources: Browne 1968, Furniss & Carolin 1977, Drooz 1985, Langor et al. 1998, Kimoto & Duthie-Holt 2006, Alfaro & Lavalle et al. 2007.

Hosts Hosts and host preference varies with location. In eastern North America this weevil is most damaging to Pinus strobus but also occurs on Picea abies, P. glauca, P. mariana, P. pungens, P. rubens, Pinus banksiana, P. resinosa and other species. In western North America, species of Picea, including P. engelmannii, P. glauca and P. sitchensis, are the preferred hosts.

Importance White pine weevil attacks, breeds in and kills leaders of host trees. Attacks cause stem deformation, reduction in height growth and increased susceptibility to decay fungi (Fig. 13.3). Tree mortality due to attacks can occur but the most significant damage is stem deformation. Deformation is usually more severe on trees growing in the open than on trees under a forest canopy. White pine weevil is considered the most economically important native insect pest of pine and spruce regeneration in Canada. It is also a major pest in portions of the USA. It is believed that widespread

establishment of pure plantations of Pinus strobus and later Picea abies in eastern North America may have led to increases in damage by this insect over the last century.

Life History White pine weevil has one generation/ year and adults overwinter in the litter beneath host trees (Fig. 13.5). In early spring, adults emerge, walk to nearby host trees, crawl up the bole or fly on warm, sunny days. They feed, mate and lay eggs in the bark of the previous year’s leader. Eggs are laid from late April to mid-July, with a peak in May and early June, and hatch in about 10 days. Larvae feed in the cambium and phloem, moving downwards, which kills the leader. Pupation occurs in chip cocoons in the leader and brood adults emerge from August to September (Fig. 13.4). Adults drop to the litter to overwinter. In coastal British Columbia, Canada, a small portion of larvae and pupae overwinter in the leader and emerge as adults the following spring.

Tip, shoot and regeneration insects

Fig. 13.3 Dead leader and multiple branching on Engelmann spruce, Picea engelmannii, caused by white pine weevil, Pissodes strobi (Waterton Lakes National Park, Alberta, Canada).

Description of Stages Adults are small brown weevils, 4–6 mm long. Elytra and body are covered with irregular patches of brown and white scales. A large white patch and a brown patch are found near the apex of each elytron. Pest Management Damage can be reduced by maintaining high tree density in open-grown plantations, planting host trees under shade or nurse crops and pruning and destruction of infested leaders in small plantations (Drooz 1985, Alfaro & Lavallee 2007). Pissodes yunnanensis Langor & Zhack, Yunnan Pine Weevil Distribution ern China.

This weevil is indigenous to southwest-

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Fig. 13.4 Chip cocoon of white pine weevil, Pissodes strobi (Roosevelt National Forest, Colorado, USA).

Hosts

The only known host is Pinus yunnanensis.

Importance P. yunnanensis attacks the upper bole on current or 1-year-old growth and, occasionally, upper lateral branches of trees less than 20 years old. It prefers to attack 8–10-year-old trees. Attacks often kill the leader and cause stem forking and crooking. After 2–3 years of consecutive attacks, trees may die. This insect is an important pest of young P. yunnanensis plantations. The incidence of attack has increased due to extensive establishment of large areas of pine plantations in southwestern China.

Life History There is one generation/year. Adults emerge from infested stems from April to mid-May.

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Fig. 13.5 Life histories of two species of Pissodes, P. strobi (North America) and P. yunnanensis (China) (based on data from Drooz 1985, Zhang et al. 2004, Alfaro & Lavallee 2007).

Eggs appear in June, instar I larvae in early July and instar IV larvae in March of the following year. Early-instar larvae feed in the phloem. Instar III larvae move into the sapwood or pith where they overwinter. Instar IV larvae excavate pupal chambers or chip cocoons in the outer sapwood or pith. Pupation occurs from late March to early May (Fig. 13.5).

Description of Stages Adults are 6–6.5 mm long, with a brown body color. They are slightly darker on the apical two thirds of the snout (Langor et al. 1999, Zhang et al. 2004).

Curculionidae (subfamily Scolytinae) Several bark beetles are of concern because they damage regeneration by adult feeding rather than because of breeding attacks. Others breed in and can kill branches of host trees. Hylastes ater (Paykull), Black Pine Beetle Distribution H. ater is a Eurasian bark beetle, widely distributed over much of Europe, Asia and the Near East including China, Cyprus, Japan, Korea, Turkey and

Asian Russia. It has been introduced into Australia, Chile, New Zealand and South Africa.

Hosts Primary hosts are pines. In Europe, Pinus nigra, P. pinaster and P. sylvestris are hosts. North American species of Pinus that are known hosts include P. ponderosa and P. radiata. Other hosts include species of Abies, Larix, Picea, Pseudotsuga menziesii, Araucaria cunninghamii and Thuja.

Importance Breeding attacks are confined to freshly cut stumps, logging residues and logs in direct contact with the ground and are not damaging. Adults feed at the base of seedlings, which can lead to mortality (Plate 72). Vigorous seedlings may survive adult feeding but wounds are encrusted with resin. In New Zealand, H. ater is associated with two fungi known to cause root disease, Leptographium truncatum and L. procerum.

Life History There are one to three generations/year andadultsandlarvaecanbepresentthroughouttheyear. Adults are monogamous and breeding attacks consist of short entry tunnels that lead to an oblique nuptial chamber where mating occurs. Eggs are deposited singly in galleries 80–130 mm long, constructed parallel to the

Tip, shoot and regeneration insects wood grain. Each female lays about 100 eggs in frasspacked niches. Larvae construct feeding galleries at right angles to the egg galleries and later take on a random pattern. They pupate in cells at the end of larval galleries and emerge to feed on young conifer seedlings. Description of Stages Adults are 4.5–5.2 mm long, red-brown when newly developed and turn dark brown or black when mature. Antennae are red-brown. The pronotum is as wide as the elytra and the frons is marked with dense punctures. The elytra are coarsely punctate–striate.

Pest Management Tactics used to prevent or suppress damage caused by Hylobius spp. are also applicable to management of H. ater (Clark 1932, Browne 1968, Milligan 1978, Grüne 1979, Neumann 1987, Ciesla 1988, MacKenzie 1992). Hylurgus ligniperda (Fabricius), Golden Haired Bark Beetle Distribution H. ligniperda occurs throughout Europe, including Great Britain and the Mediterranean

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Basin, the Caucasus Mountains and western Siberia. It is also found in the Azores (Portugal), Canary Islands (Spain), Madeira Islands (Portugal) and Morocco. It is established in Australia, Brazil, Chile, Japan, New Zealand, South Africa, Sri Lanka, Uruguay and the USA (New York and California).

Hosts This beetle attacks species of Pinus. Within its natural range it attacks P. canariensis, P. halepensis, P. brutia, P. nigra, P. nigra ssp. pallasiana, P. pinaster, P. pinea and P. sylvestris. In places where it has been introduced, P. elliottii, P. montezumae, P. patula, P. radiata and P. strobus are hosts.

Importance H. ligniperda breeds in fresh stumps or slash from recently felled trees and buried logs or portions of logs in contact with the soil. Occasionally, the base and roots of weakened or wounded standing trees are attacked. In Chile, H. ligniperda has been found in standing trees during periods of extended drought. When adults feed, they strip the bark from seedlings and kill them (Fig. 13.6). In Chile, seedlings that have malformed roots caused by poor planting (J-rooting) or natural pine regeneration with bark injury caused

Fig. 13.6 Pinus radiata seedling girdled by adult feeding of the beetle Hylurgus ligniperda. (VIII Region, Chile).

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by other insects or small mammals are more likely to be selected for adult feeding than vigorous seedlings.

Life History H. ligniperda has at least two generations/year and is monogamous. Females initiate attacks, bore through the bark and construct a small nuptial chamber. A male joins her and mating occurs. After mating, females construct a long winding egg gallery and lay eggs individually in niches. A female can lay up to 500 eggs. Larvae feed beneath the bark and pupate once they mature. Adults emerge and fly to new hosts at various times of the year.

Description of Stages Adults are 2 mm wide, 6 mm long, dark brown to black and covered with golden-red hairs, which give this insect its common name. The hairs are most conspicuous on the posterior slopes of the elytra.

Pest Management Tactics used to prevent or suppress damage caused by Hylobius spp. and Hylastes ater are also applicable to management of this species (Browne 1968, Ciesla 1988, Wood & Bright 1992).

examination by a taxonomist specializing in this group for species identification (Furniss & Carolin 1977, Wood, S.L. 1982). Pityophthorus spp., Piñon Twig Beetles Distribution The piñon twig beetles consist of a complex of at least 12 species and are found throughout the ranges of piñon pines in southwestern USA and northern Mexico and are usually referred to as Pityophthorus spp. (Table 13.3). Hosts Hosts include all of the piñon pines, principally Pinus cembroides, P. edulis and P. monophylla. Some species also infest other species of pines.

Importance These beetles attack the tips of branches and cause dieback. Tip dieback is a common sight, especially on piñon pines in urban areas where damage is more noticeable. During periods of below normal precipitation, infestations tend to move further down the branch and kill larger branches. Several piñon twig beetles, along with the piñon ips, Ips confusus, caused extensive damage to piñon forests throughout southwestern USA between 2002 and 2005.

Pityophthorus, Twig Beetles The genus Pityophthorus consists of about 300 described species. Over 200 species are known from North and Central America, 50 from South America and 50 from Asia, Africa and Europe. They are small beetles, 1.7–2.2 mm long and most are of little or no socioeconomic importance. Breeding occurs under a variety of situations. Some species breed in dead trees, often after invasion by more aggressive bark beetles such as Dendroctonus, Ips or Scolytus. Others breed in unhealthy seedlings, shaded branches, injured tops, logging residues and broken branches. Some species are of concern because they invade and kill shoots of conifers or broadleaf trees. At least one species has become a vector of a pathogenic fungus. Adults are polygamous and gallery systems consist of a nuptial chamber from which radiate several egg galleries each occupied by a female. Eggs are deposited singly along the egg gallery. Larvae feed in the cambium and pupate at the end of the larval mines. Most species have at least two generations/year. Most species of Pityophthorus are similar in appearance and require

Life History Piñon twig beetles have two or more generations/year depending on local climate. Adults emerge in spring and fly to a suitable host. Males initiate attacks and release aggregation pheromones that attract multiple females. They construct a nuptial chamber and mate with three to five females. After mating, females construct galleries that encompass the branch. Eggs are deposited in niches along sides of galleries. Larvae feed in the cambium, pupate and emerge. Winter is passed as adults that tunnel into the pith of twigs of host trees.

Description of Stages Adults are small cylindrical beetles about 1.5–2 mm long and dark red-brown. A specialist in bark beetle taxonomy is needed to separate the species involved.

Pest Management Pruning and destruction of infestedbranchesisan effective tacticforlow-levelpopulations. However, during heavy infestations this tactic will

Tip, shoot and regeneration insects

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Table 13.3 Species of Pityophthorus (Coleoptera: Curculionidae: Scolytinae) known to infest piñon pines in Mexico and southwestern USA: their distribution and hosts. Species

Distribution

Hosts

P. barberi Blackman

Mexico: Durango, Nuevo Leon USA: Colorado, New Mexico USA: Arizona, California, Colorado, Utah Mexico: Durango USA: Arizona, Colorado, New Mexico Canada: British Columbia Mexico: Baja California, Chihuahua, Hidalgo, Nuevo Leon USA: throughout the western states Mexico: Durango USA: Arizona, Colorado, Nevada, New Mexico, South Dakota USA: California, Colorado Mexico: Chihuahua USA: Arizona, New Mexico Mexico: Baja California, Chihuahua, Durango USA: Arizona, California, Nevada, New Mexico, Utah USA: California, New Mexico Mexico: Coahuila, Durango, Hidalgo, Nuevo Leon, Quertaco, Tlaxcala Canada: Alberta, British Columbia Mexico: Baja, California, Coahuila, Durango, Nuevo Leon USA: western states including Alaska USA: New Mexico

Pinus cembroides, P. edulis

P. blandus Blackman P. brevis Blackman P. confertus Swaine

P. deletus LeConte

P. keeni (Blackman) P. lecontei Bright P. modicus Blackman

P. punctifrons Bright P. schwartzi Blackman P. tuberculatus Eichhoff

P. woodi Bright

Pinus edulis, P. monophylla Pinus ayachuite, P. edulis, P. ponderosa, P. strobiformis Pinus spp., including P. cembroides and P. edulis

Pinus cembroides, P. edulis, P. monophylla. Less common in P. flexilis, P. ponderosa and P. strobiformis Pinus edulis, P. monophylla Pinus cembroides, P. edulis Pinus cembroides, P. edulis, P. monophylla, P. sabiniana

Pinus edulis, P. monophylla, P. quadrifolia Pinus cembroides, P. edulis, P. greggii, P. lumholtzii, P. montezumae, P. teocote Pinus spp., including P. cembroides, P. edulis, P. monophylla, Picea spp. (rare)

P. edulis

Source: Wood, S.L. 1982.

result in removal of most of the crown and affected trees cannot be saved. Contact insecticides, applied in early spring (March–April) can prevent attacks in high-value urban trees (Wood, S.L. 1982, Sandoval n.d.). Pityophthorus boycei Swaine Distribution This twig beetle is widely distributed in western North America from British Columbia, Canada, south to California and east to South Dakota and Colorado, USA. Hosts Three species of pines are reported hosts: Pinus aristata, P. contorta and P. ponderosa.

Importance This is a little known species but it is capable of infesting twigs and shoots of apparently

healthy, vigorous pines. Infested twigs are killed and subject to wind breakage. Infestations can, at times, be heavy; for example, in 2009 infestations were so heavy in P. aristata near the summit of Thirty-Nine Mile Mountain, an isolated peak west of Colorado Springs, Colorado, USA, that damage was aerially visible. Approximately 70–80% of the branch tips had been killed in the heaviest area of infestation, giving the affected area a brown discoloration similar to that caused by defoliating insects (Plate 73, Author’s observation). Infestations and resultant damage were also observed during 2008 and 2009 on the Eldorado National Forest, California where up to 50% of twigs were affected.1

1

Personal communication, Joel Egan and Martin MacKenzie, USDA Forest Service, Sonora, California.

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Life History Little is known about the life history and habits of this species. Adults construct tunnels in the pith of new shoots, where they deposit eggs. Larvae enlarge and extend the tunnels and emerge as adults before the needles turn brown. Adults overwinter in pine shoots (Plate 74).

Description of Stages and dark brown.

Boulder, Colorado Springs and Denver, Colorado, USA. The walnut twig beetle/thousand cankers disease complex poses a threat to native forests of J. nigra in eastern and central USA, the wood of which is valued for furniture, gunstocks and other quality wood products. This concern became a reality when infestations were detected in eastern Tennessee in 2010.

Adults are 1.9–2.9 mm long

Pest Management No pest management tactics are available for this insect (Wood, S.L. 1982). Pityophthorus juglandis Blackman, Walnut Twig Beetle Distribution Prior to about 2001, walnut twig beetle occurred in Arizona, a small area of southern California and New Mexico, USA, and northern Mexico. This insect has expanded its range northward and is now known to occur in central California and portions of Idaho, Oregon, Utah and Washington. In 2010, infestations were discovered in Tennessee.

Hosts Arizona walnut, Juglans major, is the host within its traditional range. In its expanded range, it attacks several species of Juglans, including California walnut, J. californica, and black walnut, J. nigra.

Importance Historically, walnut twig beetle has been of little or no consequence. It causes dieback on twigs and smaller branches of stressed J. major. In its expanded range it has become associated with several new hosts. Attacks on these hosts involve branches of larger diameter resulting in crown dieback. In addition, this twig beetle has developed an association with the fungus Geosmithia morbida. As beetles bore in the twigs, the fungus is introduced. In some species, especially J. nigra, this leads to development of cankers under the bark, a condition known as “thousand cankers disease.” Cankers restrict flow of nutrients and eventually kill the tree, sometimes within a year. A second fungus, Fusarium solani, has also been recovered from trees attacked by this beetle but its role as a pathogen is less clear. This insect–fungus association has killed many ornamental J. nigra in cities such as

Life History Two or more generations/year may occur and winter is spent as adults. Males initiate attacks in early May and introduce the Geosmithia fungus, which grows in advance of the beetle. They construct a nuptial chamber and attract and mate with several females. After mating, females construct egg galleries that radiate from the nuptial chamber and lay eggs. Larvae develop just under the bark and then enter the bark to pupate. A generation can be completed in less than 2 months. Description of Stages Adults are 1.5–1.9 mm long and yellow-brown in color. Pest Management No effective pest management tactics are available for this insect (Wood, S.L. 1982, NAPPO 2008, Tisserat et al. 2009, Colorado State University n.d.). Tomicus Bark beetles of the genus Tomicus are called “pine shoot beetles” because the adults tunnel in and kill branch tips. Damage can be significant because they kill branches and cause crown deformity. In addition, needles on affected branches die and turn red, which is unsightly on ornamental and Christmas trees. Some species breed in recently killed trees and others breed in, and kill, weakened or stressed trees. Seven species are known. All occur in Asia and/or Europe and are considered among the more damaging insects of Eurasian pine forests (Table 13.4). One species, T. piniperda Linnaeus, has been introduced into North America (Haack & Kucera 1993, Kirkendall et al. 2008).

Tomicus destruens (Wollaston) Distribution This insect is indigenous to the Mediterranean region of north Africa, Europe (southern

Tip, shoot and regeneration insects

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Table 13.4 Species of Tomicus (Coleoptera: Curculionidae: Scolytinae): their distribution and hosts. Species

Distribution

Hosts

T. brevipilosus (Eggers)

Asia: China, India (Assam), Japan, Korea

T. destruens (Wollaston)

Mediterranean Europe and the Near East

T. minor (Hartig) T. pilifer (Spessivtsew) T. piniperda Linnaeus

Eurasia Asian Russia, northern China North Africa, Asia, Europe (indigenous), North America (introduced) Russia. Siberia, Far East, Sakhalin Island

Pinus koraiensis, P. insularis, P. parvifolia, P. yunnanensis P. brutia, P. canariensis, P. halapensis, P. pinaster and P. pinea Picea, Pinus and other conifers Pinus armandii, P. koraiensis, P. tabulaeformis Picea, Pinus and other conifers

T. puellus (Reitter) T. yunnanensis Kirkendall & Faccoli Yunnan shoot borer

Yunnan Province, China

Abies holophylla, A. nephrolepis, Picea ajanensis, P. jezoensis, Pinus koraiensis Pinus yunnanensis

Source: Browne 1968, Wood & Bright 1992, Haack & Kucera 1993, Kirkendall et al. 2008.

France, Italy, Portugal and Spain) and the Near East (Cyprus, Israel and Turkey).

Hosts Hosts include Pinus brutia, P. canariensis, P. halepensis, P. pinea, and P. pinaster.

Importance T. destruens causes two types of damage. Adults feed in tender young shoots and cause tip dieback and growth loss. Breeding attacks in boles of living trees cause tree mortality. T. destruens can be an aggressive tree killer and attacks trees stressed by drought, defoliation, root disease, poor site and related factors. In Italy, pines growing in coastal areas where the water table is high and occasionally exposed to salt water are vulnerable to attack by root disease fungi and ultimately T. destruens. In northern Cyprus, trees in excess of about 30 cm diameter breast height (dbh) are subject to top kill and often have attacks in the larger branches of live trees (Fig. 13.7).

Life History The life cycle of this bark beetle is variable, depending on local conditions. Studies in northern and central Italy indicate that there are two generations/year. In northern Italy, overwintering takes place as adults in shoots or stumps of host trees. Adults emerge, fly and attack trees between March and April. In central Italy, all life stages may overwinter, adult flight begins in February and new host material is attacked from March to April. First-generation adults

mature in both locations in early summer and begin feeding in pine shoots. Attacks by the second generation begin in August in both locations. In northern Italy, flight and attacks are completed by onset of winter while in southern Italy, beetle flight and attacks may occur into the winter months.

Description of Stages Adults are cylindrical with a smooth and rounded elytral declivity and are dark brown. The head is visible when viewed from above and length is 3.5–4.8 mm (Abgrall & Soutrenon 1991, Wood & Bright 1992, Nanni & Tiberi 1997, Kohlmayr et al. 2002, Ciesla 2004). Tomicus piniperda Linnaeus Distribution This species is indigenous to North Africa and across Asia and Europe. It has been introduced into North America, where it was first detected in 1992 and is now established over much of northeastern and north central USA and adjoining Canada. Hosts In Europe, Pinus mugo, P. nigra, P. radiata and P. sylvestris are attacked. In Asia, T. piniperda is reported from Pinus armandii, P. koraiensis and P. massoniana. In North America, P. sylvestris is the preferred host but P. banksiana, P. nigra, P. resinosa and P. strobus are also attacked. Other conifer hosts include various spruces, Picea abies, P. obovata and P. smithiana, Larix decidua and Pseudotsuga menziesii.

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Fig. 13.7 Galleries and brood of the bark beetle, Tomicus destruens, in branch of Pinus brutia (northern Cyprus).

Importance The most severe damage caused by T. piniperda is destruction of shoots by adults during adult feeding (Plate 75). When shoot feeding is severe, tree height and diameter growth is reduced. Christmas trees attacked by this insect are unsightly because of dead shoots. Trees with heavy feeding injury tend to have narrow crowns due to adult feeding. Breeding attacks in stumps, weakened trees, fresh cut logs or logging residues cause little damage. Life History T. piniperda has one generation/year. Adults become active during the first warm days of spring and colonize recently cut stumps, logs or the trunks of severely weakened trees. They use volatile host terpines, such as a-pinene, to locate suitable breeding site and do not produce aggregation pheromones. Females initiate gallery construction and one male joins each female. After mating, females construct vertical egg galleries 10–25 cm long in the cambium. They lay eggs singly in niches cut into each side of the egg gallery. After hatching, larvae construct horizontal feeding galleries 4–9 cm in length. Pupation occurs in cells at the end of the larval galleries. Brood adults tunnel through the outer bark and create exit holes about 2 mm in diameter. They fly to

crowns of live, healthy pines of all ages but prefer the tallest trees. Adults feed in the upper half of the crown from May to October in the temperate portions its range. During maturation feeding, each adult may damage one or two shoots. They usually enter 1-year-old or current year’s shoots and tunnel into the center of the shoot and bore outwards, hollowing out 2–3 cm of the shoot. After several weeks, adults emerge and enter other shoots. Infested shoots generally bend near the point where the beetles entered, turn yellow to red, eventually break off and fall to the ground. In warmer parts of its range, adults overwinter in the shoots. In colder climates adults exit shoots after the first frost and enter the thick bark at the base of pines to spend the winter.

Description of Stages Overall appearance of adults is similar to that of T. destruens and a specialist is needed to separate the two species. Pest Management In Europe, where harvested logs are often left in the forest for extended periods, breeding attacks are prevented by debarking of logs, exposure of logs to solar radiation, or application of chemical insecticides (Browne 1968, Grüne 1979, Abgrall &

Tip, shoot and regeneration insects Soutrenon 1991, Haack & Kucera 1993, Bright & Skidmore 2002, Kohlmayr et al. 2002). Tomicus yunnanensis Kirkendall & Faccoli, Yunnan Shoot Borer Distribution This insect has to date been found only in Yunnan Province, China.

Hosts Pinus yunnanensis is the primary host. Shoot feeding can also occur in P. armandii, P. densata and P. kesiya var. langbianensis.

Importance This is an unusually aggressive species. Adults aggregate in large numbers to feed on shoots of Pinus yunnanensis and subsequently attack boles of the same trees for breeding. Outbreaks have damaged over 200,000 ha of pine forests.

Life History This species, described in 2008, was once thought to be an Asian form of T. piniperda. Like T. piniperda, T. yunnanensis has one generation/year. Breeding attacks tend to occur in mid- and lower boles of trees whereas T. piniperda prefers to attack the lower bole. Adults emerge in early spring and fly toward crowns of pines to feed on shoots. They aggregate in large numbers and damage four to five shoots per adult as compared to one to two shoots for T. piniperda. Following shoot feeding, adults mass attack the boles of the same trees. The primary blue-stain fungus associated with this species is Leptographium yunnanese.

Description of Stages Adults are dark red-brown beetles, 4.3–5.4 mm long (Kirkendall et al. 2008).

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Some species are leaf rollers, others bore in shoots and tips and still others invade flowers, fruits and seeds. Several are important forest pests. Species that invade shoots and stems are reviewed in this chapter and species that invade cones of conifers in Chapter 14. Eucosma sonomana Kearfott, Western Pine Shoot Borer Distribution Western pine shoot borer is found in western North America including British Columbia, Canada and Arizona, California, Oregon, Colorado and Montana, USA and throughout much of Mexico.

Hosts The principal host in Canada and the USA is Pinus ponderosa. P. contorta, P. jeffreyi and Picea engelmannii are also infested. In Mexico, several pines are hosts.

Importance Larvae bore into the center of terminal shoots. Infestations cause reductions in height growth of up to 25% each year the terminal shoot is attacked. Attacked terminals can lose dominance, which leads to multiple leaders and deformity. Lateral shoots that are attacked are usually killed. This insect is considered an important pest of pine plantations.

Life History Western pine shoot borer has one generation/year. Adults emerge in spring and are active for 2–3 months. Females deposit eggs on elongating shoots. Terminal shoots that are robust, and presumably more suitable for larval development, are preferred for egg laying. Larvae hatch within several days, bore into shoots and mine the pith. They complete development by mid-summer. There is usually one larva per infested shoot. When feeding is completed, larvae chew circular exit holes in the lower part of the shoot and drop to the ground. Winter is passed as pupae in cocoons in the litter underneath infested trees.

Tortricidae (Subfamily Olethreutinae) Eucosma Eucosma is a large genus of moths distributed throughout the northern hemisphere and into the subtropical and tropical regions of Asia. They have a wide rage of habits.

Description of Stages Adults are small moths with copper-red forewings marked with two bright gray transverse bands. They have a wingspan of 16–22 mm. Larvae are about 15 mm long when mature. Body color is pale pink-red with a dark head capsule.

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Pest Management Damage caused by this insect in pine plantations can be reduced by applications of either contact or systemic insecticides or use of pheromones for either mating disruption or to trap and kill male moths.

Related Species E. gloriola Heinrich, the eastern pine shoot borer, is found in eastern Canada and northeastern and north central USA and bores in the shoots of Pinus banksiana, P. resinosa, P. sylvestris and other pines. It is also considered a pest of young plantations (Furniss & Carolin 1977, Cibrian Tovar et al. 1995, Randall 2005). Rhyacionia Rhyacionia consists of several species of pine infesting shoot borers. Most are native to North America and several are considered pests of young pine stands of both natural origin and plantations. One Eurasian species has been introduced into both North and South America and has caused extensive damage to both young plantations and ornamental trees (Table 13.5). Rhyacionia buoliana (Dennis and Schiffermüller), European Pine Shoot Moth Distribution European pine shoot moth is native to North Africa (Algeria, Morocco and Tunisia) and throughout Eurasian conifer forests. Two subspecies

are recognized in Europe: Rhyacionia buoliana buoliana occurs in the north and R. buoliana thurificana in the Mediterranean region. Other subspecies occur eastward from Europe to Japan. It was first detected in the USA in 1914; it spread rapidly and by 1970 was established across the northeastern states and adjacent Canada, with limited occurrences in British Columbia, Oregon and Washington and. It appeared in Argentina in 1939 and is now also present in Chile and Uruguay.

Hosts This insect prefers three-needled pines and has been reported from 30 or more species. In its native habitat, Pinus halepensis, P. mugo, P. nigra, P. pinaster, P. pinea and P. sylvestris are known hosts. In France, pines most frequently attacked are P. nigra, P. sylvestris, P. ponderosa (introduced) and P. radiata (introduced). Pines rarely attacked are P. halapensis, P. pinaster and P. pinea. In places where it has been introduced into North America, P. contorta, P. mugo, P. muricata, P. ponderosa, P. radiata, P. resinosa and P. taeda are hosts. In Argentina and Chile, P. radiata is the primary host and P. taeda is the primary host in Uruguay.

Importance Feeding by larvae causes infested shoots to form a crook, which causes deformity and adversely affects the form of trees grown for lumber (Fig. 13.8). It is also damaging in nurseries and Christmas tree plantations, where infestations can render trees unsightly and unmarketable. Within its natural range, European pine shoot moth is most aggressive in central Europe

Table 13.5 Distribution of representative species of pine infesting Rhyacionia (Lepidoptera: Tortricidae). Species

Distribution

R. buoliana (Dennis and Schiffermu¨ller) European pine shoot moth R.. bushnelli (Busck) Western pine tip moth R. frustrana (Comstock) Nantucket pine tip moth R.. neomexicana (Dyar) Southwestern pine tip moth R.. rigidana (Fernald) pitch pine tip moth R.. subtropica Miller Subtropical pine tip moth R.. zozana (Kearfott)

Eurasia (indigenous), North and South America (introduced) Central USA and introduced into several western states Eastern USA from Massachusetts south to Florida, and west to Missouri, Oklahoma, Texas. Introduced into California. Central and southwestern USA Eastern USA, Maine and New York, south to Florida and west to Missouri and Texas Southeastern USA, range corresponds to range of Pinus elliottii California, Oregon, Washington

Sources: Yates 1968, Furniss & Carolin 1977, Powell & Miller 1978, Yates et al. 1981, Drooz 1985, Abgrall & Soutrenon 1991.

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south central Chile, this insect completes only one generation/year. Adults usually mate within the first day after emergence and females are active from late spring to early summer, usually at dusk. Eggs are deposited at the bases of needles and buds. Larvae hatch, mine in needles and then bore into buds. Infested buds are killed by midsummer. Instar III larvae cease to feed in late summer and remain quiescent in the tunnels over winter. They resume activity in early spring when they feed in new buds and elongating shoots. Pupation occurs inside feeding tunnels in spring and adults emerge 1–2 weeks later.

Description of Stages Adults have orange forewings marked with several irregular silver lines and hind wings are gray. Wingspan is approximately 20 mm. Eggs are disk shaped and about the diameter of a pine needle. They are yellow when first laid and later turn orange and brown. Mature larvae are about 15 cm long. Bodies are light brown with black prothoracic and anal plates. Pupae are red-brown and 10 mm long.

Fig. 13.8 Branch crook on Pinus radiata caused by European pine shoot moth, Rhyacionia buoliana, Argentina (photo by Paula Klasmer, Instituto Nacional de Tecnologia Agropecuaria, Argentina).

(Hungary, Romania and Slovakia) and portions of Ireland and the Netherlands. In Chile, it has become a major pest of P. radiata plantations. In North America, it can be damaging to young pine plantations and ornamental pines.

Life History R. buoliana buoliana has one generation/ year, as do populations in the USA. R. buoliana thurificana has two generations/year. It has been suggested that populations in Argentina may have more than one generation/year but this remains to be confirmed. Some confusion exists concerning R. buoliana varieties and generations/year, and it is uncertain at this time which variety exists in Chile. However, in Regions IX and X of

Pest Management Several pest management approaches are used to reduce losses with varying degrees of success. Chemical control can be effective at three stages in its life cycle: migrating larvae in early spring, adults in early summer and newly hatched larvae in early summer. Timing of spray applications directed against adults and newly hatched larvae can be done via use of pheromone traps. This tactic can effectively protect nurseries, Christmas tree plantings and ornamental trees but is impractical over large areas of forest plantations. In Europe, interplanting of pine plantations with lupines, Lupinus spp., has reduced incidence of attack and resultant damage, presumably because lupine blossoms provide a food source for parasitoids. Mating disruption, using pheromones, has also proven effective. In places where European pine shoot moth has been introduced, classic biological control has been used with reasonable success. In Chile, for example, a consortium of forest companies has established a laboratory for mass rearing and release of the parasitoid, Orgilus obscurator (Miller 1967, Abgral & Soutrenon 1991, L angstrüm et al. 2004, Controladora de Plagas Forestales 1997, Cerda et al. 1985).

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Rhyacionia frustrana (Comstock) (Nantucket Pine Tip Moth) Distribution Nantucket pine tip moth is widely distributed over eastern USA from Massachusetts south to Florida, west to Missouri, Oklahoma, Texas and California. The population in California is most likely an introduction and has been traced to a shipment of infested seedlings from Georgia in 1967. It also occurs in the Caribbean region and Central America, including Cuba, the Dominican Republic, Guatemala, Honduras, Jamaica, Mexico (Oaxaca) and Nicaragua.

Hosts At least 20 species of pines are indigenous to eastern USA, the Caribbean and Central America. In the southern and southeast states, hosts are Pinus echinata and P. rigida, P. sylvestris, P. taeda and P. virginiana are favored in the northeastern, New England and middle Atlantic states. P. echinata is preferred in the central states and in California the host is P. radiata.

Importance Larvae bore in shoots, buds and conelets, cause growth loss, stem deformity and destruction of conelets. Most damage occurs during the first 5 years in the life of a host pine. This tip moth is considered an important pest of young pine plantations, natural stands of young pines, Christmas tree plantings, ornamental pines and pine seed orchards in eastern USA.

Life History Nantucket pine tip moth can have up to five generations/year, depending on local climatic conditions. It overwinters as pupae inside damaged shoots, cones or buds. Adults emerge in the early spring, sometimes as early as February. After mating, females deposit eggs on new shoots, conelets or last year’s shoots of pines. In cool weather (late winter or early spring), eggs may take 30 days to hatch, but can hatch in as early as 5 days during hot weather. Young larvae may feed on the outside of new shoots for a few days and then bore into shoots, conelets and buds. Larvae feed for about 3–4 weeks and pupate in the damaged tissues.

Description of Stages Female adults are larger than males. Head, body and appendages are covered with gray scales. The wings are mottled rusty-red and forewings are marked with dark basal patch bordered by

a lighter cross band that is narrower than the basal patch. Wingspan of forewings of males is 4.0–7.0 mm and of females is 4.0–7.5 mm. Eggs are slightly convex, about 0.8 mm in diameter, opaque white when first laid and later turn yellow-medium green at maturation. Young larvae are cream colored with a black head. As the larvae grow, body color becomes light orangebrown. They are about 9 mm long when mature. Pupae are light-dark brown and 4.6–7.5 mm long.

Pest Management Preventative tactics include establishment of pine plantations on suitable sites to reduce stress and encourage good growth. Rapid crown closure of pine plantations should be encouraged. Growth of flowering plants in the understory may favor natural enemies. Infested shoots and conelets can be removed by hand pruning if infestation levels are low. Insecticides may be applied at the time of adult flight and egg deposition. The attractant pheromone for this insect has been identified and synthesized and timing of insecticide applications can be determined by deployment of pheromone-baited traps.

Related Species Two related species of Rhyacionia often infest the same trees as the Nantucket pine tip moth. These are the pitch pine tip moth, R. rigidana (Fernald), and the subtropical pine tip moth, R. subtropica Miller. The pitch pine tip moth is more prevalent and difficult to distinguish from Nantucket pine tip moth. The range of the subtropical pine moth is restricted to Florida and southern Georgia, Mississippi and South Carolina, USA. Adults and pupae of three species of Rhyacionia with overlapping host ranges, R. frustrana, R. rigidana and R. subtropica (Miller), are difficult to separate but keys to their identification are available (Yates 1969, Powell & Miller 1978, Yates et al. 1981).

Pyralidae Hypsipyla, Mahogany Shoot Borers Hypsipyla consists of 11 known species of pyralid moths. All occur in the tropics. Four species are found in the western hemisphere and seven in the eastern hemisphere. Two are important pests of trees of the family Meliaceae, including species of the genera Cedrela, Khaya, Swietenia and Toona. Larvae feed in shoots,

Tip, shoot and regeneration insects flowers and fruits of host trees. Shoot feeding causes severe damage in young plantations of mahoganies and severely limits ability to grow trees in plantations. The remaining nine species have limited distributions, are not important pests and little is known of their biology (Griffiths 2001, Horak 2001). Hypsipyla grandella (Zeller) Distribution This mahogany shoot borer is a neotropical species and is found from Florida, USA and southern Mexico south to Peru and Brazil and throughout the Caribbean Islands (Fig. 13.9).

Hosts Cedrela mexicana, C. odorata, Khaya senegalensis, Swietenia macrophylla and S. mahogani are hosts of this borer. Importance Thisinsectisa major pest of youngtreesin tropical forest plantations and stem attacks cause growth loss and deformity. Trees aged 2 years and older are susceptible to attack. Attacks in fruits result in premature shedding but is of less importance than stem attacks.

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Life History Life history and habits of H. grandella and its old world counterpart, H. robusta are similar. One to two months are generally required to complete a generation but this may extend to 5 months if larvae enter diapause. Females mate once and deposit between 200 and 450 eggs over five to eight nights on green shoots of host trees. Adults are strong fliers and can locate suitable trees over considerable distances. Mortality of eggs and early-instar larvae is high but even low populations cause significant damage and have an adverse effect on tree form. Larvae bore longitudinally in the center of the shoot and later pupate inside feeding galleries or in soil or plant material at the base of the tree. Habits of larvae that feed in flowers and fruit of host trees are less well known. In areas of low temperature or seasonal drought, stem feeding larvae enter a diapause. All larvae that feed on fruits enter diapause.

Description of Stages Adults are brown to graybrown with a wingspan of 23–45 mm. Forewings are gray to brown with rust red on the lower portion and white scales with black spots toward the wing tips. Wing veins are distinctively overlaid with black. Eggs

Fig. 13.9 Worldwide distribution of the mahogany shoot borers, Hypsipyla grandella and H. robusta (Lepidoptera. Pyralidae) (redrawn from Griffiths 2001).

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are oval, flattened and 0.5–1.0 mm long and wide. They are white when first deposited and, if fertilized, develop red and white bands within 24 hours.

Pest Management Silvicultural tactics for both H. grandella and H. robusta include establishment of mixed orenrichmentplantings,varyingtreedensity,provisionof shade, promoting vigorous tree growth in nurseries and plantations, and selection of resistant or tolerant host trees. At present, there no single reliable, cost-effective, environmentally sound chemical insecticide available to control these insects. Chemical control is most applicable in nurseries or as part of an integrated program to achieve temporary population reduction over limited areas (Browne 1968, Griffiths 2001, Horak 2001).

Hypsipyla robusta (Moore) Distribution H. robusta is indigenous to the old world tropics. It is found in the western Africa from Senegal east to Nigeria and in eastern Africa in Mozambique, Tanzania and the island of Madagascar. In Asia, it occurs from the Indian continent south and east through the mainland of Southeast Asia, Indonesia, the Philippines, Papua New Guinea and the east coast of Australia (Fig. 13.9).

Hosts Hosts are species of Meliaceae including Cedrela mexicana, C. odorata, Chloroxylon swietenia, Chukrasia tabularis, Entandrophragma angolense, E. utile, Khaya anthotheca, K. grandiflora, K. ivorensis, K. nyasica, K. senegalensis, Lovoa trichilioides, Swietenia macrophylla, S. mahogani, Toona australis, T. ciliata and T. serrata.

Importance Larvae are shoot borers but also attack flowers and fruit. Primary damage is deformity and multiple branching caused by shoot infestation. Trees can be attacked as early as age 2–3 years. Infestations tend to be heaviest on young vigorous trees growing in full sunlight. In Sri Lanka, infestations are found at altitudes as high as 1700 m and have destroyed plantations of Toona serrata. In Ghana and Nigeria, infestations have limited the ability to establish plantations of Khaya and Lovoa and in Ghana plantation survival rates as low as 9% are reported. This insect is considered a major pest of Swietenia plantations in Malaysia.

Life History Life history and habits are similar to H. grandella. Number of generations varies with location. In portions of India, 6 months may be required to complete a generation. Winter is usually spent as instar IV larva. In equatorial regions there is a complex of overlapping generations and development from egg to adult can be completed in as little as 2 months. One larva may infest more than one shoot or fruit during development. Pupation occurs in a cocoon constructed in the larval tunnel or other sheltered place. Description of Stages Adults are small with a wingspan of 30–50 mm. Instar I–III larvae are light brown to red and instar IV larvae are blue-green. The head, pronotal and anal plates are black. Each side of the body has five rows of black spots. Pest Management Available tactics to reduce damage by this insect in plantations of Meliaceae are similar to those given for H. grandella (Browne 1968, Atuahene 1996, Griffiths 2001, Horak 2001, Wagner et al. 2008). Dioryctria Dioryctria is a large genus of conifer infesting insects and contains many species of economic importance. Larvae bore into buds, cones, shoots and stems of host trees and are found throughout the boreal, temperate and subtropical conifer forests of the northern hemispheres. Larvae of some species have variable habits and invade shoots, stems and/or cones. Species that are primarily shoot and stem borers are discussed in this chapter (Table 13.6) and those that infest cones are reviewed in Chapter 14. Dioryctria sylvestrella Ratzburg, Maritime Pine Borer Distribution Maritime pine borer is a Eurasian species widely distributed in Europe from Portugal and France north to Belgium, the Netherlands and the UK and east across Asia to Thailand and Vietnam.

Hosts This insect attacks several species of Pinus, including Pinus halapensis, P. nigra, P. pinaster and P. pinea.

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Table 13.6 Representative stem and shoot infesting species of Dioryctria (Lepidoptera. Pyralidae), their distribution and hosts. Species

Distribution

Hosts

D. assamensis Muutura D. horneana (Dyar) D. raoi Muutura D. rubella Hampson Tusam pitch moth D. sylvestrella Ratzburg Maritime pine borer D. zimmermani Grote Zimmerman pine moth

India Cuba India Asia: Indonesia, Philippine Islands

Pinus Pinus Pinus Pinus

Eurasia, east to Thailand and Vietnam Canada and northern USA

Pinus caribaea, P. halapensis, P. kesiya, P. merkusii, P. nigra, P. pinaster, P. pinea Pinus, Pseudotsuga menziessi

kesiya caribaea roxburghii, Pinus spp. caribaea, P. kesiya, P. merkusii

Sources: Browne 1968, Mutuura 1971, Drooz 1985, Abgrall & Soutrenon 1991, FAO 2007a, Nair 2001, 2007.

Importance Maritime pine borer is considered an important pest of young pine plantations. Larvae bore in stems of young trees and cause a copious resin flow, which weakens trees and increases susceptibility to attack by other agents. Resin flow, caused by pruning of live branches, increases susceptibility to attack. Life History Throughout most of its range, there is one generation/year. However, in the Mediterranean region, two generations/year are common. In areas where there is a single generation, adult flight occurs from late July to August. Eggs are present from late July to early September. Beginning in early August, eggs hatch and young larvae bore into the stems of host trees and construct galleries. Larvae overwinter and pupation occurs from June to late August. Description of Stages Adults have a wingspan of 28–35 mm. The forewings have a dark brown background with light gray bands. There is a red-brown spot on the lower portion of the wing toward the body. The hind wings are light gray-brown. Larvae are yellowwhite with a red-brown head capsule and thoracic shield. Each abdominal segment bears dark spots at bases of the setae (Abgrall & Souternon 1991, Jactel et al. 1996, Nair 2007). Dioryctria zimmermani Grote, Zimmerman Pine Moth Distribution Zimmerman pine moth is found throughout much of northern USA and adjoining portions of Canada.

Hosts Hosts are various species of Pinus. Douglas-fir, Pseudotsuga menziesii, is also subject to attack. Importance Larvae feed in buds, cambium and outer xylem tissue of limbs and larger branches of young pines. Infestations can cause stem deformity, branch dieback and reduced height growth. Individual trees may endure repeated attacks. Damage has been especially severe in Christmas tree plantations in portions of Canada and north central USA.

Life History This insect has one generation/year. Adults are active from mid-July to August. They deposit eggs at the edge of wounds, on masses of resin, bark crevices or terminal buds. Eggs hatch within 8–10 days and young larvae enter bark recesses, spin hibernacula and overwinter without feeding. Feeding begins in May and June, first in the bark and later in the cambium of new terminal or lateral branches.

Description of Stages Adults have gray forewings blended with red-brown and marked with transverse, zigzag white and black lines. The hind wings are yellowwhite. Mature larvae range from off-white, pink or pale green with prominent small black setal bases on each thoracic and abdominal segment. Length is about 18 mm (Drooz 1985).

Chapter 14

Insects of Tree Reproductive Structures

INTRODUCTION Reproductive structure of trees, including pollen, flowers, nuts and cones, are food sources and breeding sites for many insects. Species that damage tree fruits, such as the codling moth, Cydia pomonella, are important agricultural pests. Those that damage fruiting structures of forest trees cause reduction in seed available for tree nurseries that produce seedlings for reforestation and afforestation. In addition, these insects can affect supplies of chestnuts, acorns or nuts that are important non-wood forest products. These insects have become subjects of increased interest since seed orchards were established to produce seed and tree seedlings with desirable genetic traits, such as superior form, faster growth rates or resistance to disease, for forest plantations (Turgeon et al. 1994). Several orders, including the Coleoptera, Diptera, Hemiptera, Hymenoptera and Lepidoptera, include pests of acorns, nuts, fruits, cones and seeds of forest trees. Some have the ability to feed in a variety of habitats,

including buds, foliage, stems and shoots, in addition to tree reproductive structures. Western spruce budworm, Choristoneura occidentalis, for example, an insect indigenous to western North America and profiled in Chapter 7, is a major forest defoliator. Larvae can also bore into cones and destroy seeds (Dewey 1970). Insects that damage reproductive structures of trees are profiled in this chapter.

HEMIPTERA Lygaeidae (Seed Bugs) Orsillus depressus (Dallas), O. maculatus Fieber, Cypress Seed Bugs Distribution O. depressus is found throughout most of Europe, including the UK, northern Africa and Asia east to Tajikistan. It is more common in the western Mediterranean region and is the dominant seed bug on the Iberian Peninsula (Portugal

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and Spain). O. maculatus also occurs throughout the Mediterranean region but is more abundant in the eastern portions.

of O. maculatus (Rouault et al. 2005, 2007, Bouaziz & Roques 2006). Coreidae (Leaf-Footed Bugs)

Hosts O. depressus feeds on cones and seeds of Cupressus, Juniperus and Pinus. It has also adapted to cones and seeds of the North American Chamaecyparis lawsoniana, which is commonly planted as an ornamental in the UK and other parts of Europe. O. maculatus is closely associated with the Mediterranean cypress, Cupressus sempervirens, and may also invade cones of other Cupressus or, rarely, Chamaecyparis lawsoniana.

Importance Nymphs and adults of both species feed on the seeds of host trees, which results in reduced yields. O. depressus also feeds on foliage.

Life History Life history of the two species is believed to be similar and has been studied in detail for O. maculatus. Females lay eggs on cones and they are deposited in all seasons except winter on the inner scales of partially open cones. Females also deposit eggs in exit holes cut through cone scale by emerging adults of the seed chalcid, Megastigmus wachtli. This rather unique habit of egg deposition is believed to be an adaptive strategy to avoid attacks by egg parasitoids of the genus Telenomus. There are five nymphal instars and all are present in the field from May to November. Adults are also present throughout much of the year but are most abundant in September, when cones mature and most of eggs are deposited. Nymphs and adults seek shelter inside cones of host trees to overwinter.

Description of Stages Life stages of both species are similar in appearance. Adults are about 16 mm long and brown with red highlights. Adults of O. maculatus tend to be darker in color with more distinct red highlights. O. maculatus usually has a dark pronotal spot whereas O. depressus has a vertical black line from the top to bottom of the pronotum. Eggs are light yellow when first laid and later turn dark yellow with red punctuations. Nymphs are pink-white when first hatched or after each molt and turn dark brown. Instar I and II nymphs of O. depressus are slightly longer than those

Leptoglossus Leptoglossus consists of more than 40 known species. All except one are indigenous to the western hemisphere. They are characterized by flattened, dilated hind tibiae that range in shape from lanceolate to broadly scalloped. Several are damaging to agricultural crops including tree fruits, cucurbits, tomatoes and cacao (Mitchell 2000). Species of importance in forestry suck juices from conifer seeds and damage seed crops. Leptoglossus occidentalis Heidemann, Western Conifer Seed Bug Distribution This seed bug is distributed from Alberta and British Columbia, Canada south to Mexico. Its range has been expanding and is now found as far east as western New York and Pennsylvania, USA. It has also been introduced and become established in Italy and Slovenia.

Hosts Seeds of many conifers are hosts including Pinus monticola, P. ponderosa other species of Pinus, Pseudotsuga menziesii and Calcocedrus decurrens.

Importance Both adults and nymphs feed on conifer seeds by piercing through cone scales into developing seeds. When feeding occurs before the seed coat hardens, contents of the seed are completely removed and the seed coat collapses. Heavy infestations cause losses of seed crops. Losses of up to 41% have been documented for Pseudotsuga menziesii and up to 26% for Pinus monticola. Adults also feed on developing male flowers in spring and this causes them be stunted and deformed. In addition, adults often enter homes in autumn to seek a warm overwintering site and they can become household pests. Life History There is one generation/year. Adults overwinter and often invade homes in autumn. In spring, eggs are deposited in rows of four to five on

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needles of host trees. Eggs hatch about 10 days later. Both adults and nymphs feed on developing cones or seeds.

Rhopalidae (Scentless Plant Bugs)

Description of Stages Adults are 15–18 mm long, 4–6 mm wide and red-brown to dark gray in color. There is a faint white zigzag stripe across the mid-point of its upper surface (Plate 76). Nymphs are orange and brown and become red-brown to brown as they develop and grow. Eggs are barrel shaped.

Distribution Boxelder bugs are found across North America. B. trivittata is an eastern species found as far west as Alberta, Canada and Arizona, Utah and Montana, USA. B. rubrolineata is found from British Columbia, Canada, south to California and east to Texas, USA.

Pest Management Heavy infestations in seed orchards are treated with ground applications of contact insecticides.

Hosts Primary host is boxelder, Acer negundo. Other species of Acer and Fraxinus are also hosts. Fruit trees and some agricultural crops are also used as food sources.

Related Species L. corculus (Say), the southern pine seed bug, is found in southeastern USA. Nymphs and adults feed on all species of Pinus indigenous to the region. Instar II nymphs of this species feed on ovules in conelets and cause conelet abortion. Later instars and adults feed on seeds (Furniss & Carolin 1977, Hedlin et al. 1980, Roques et al. 2006, Klass 2008).

Importance Nymphs insert stylets into seeds of host plants and suck the juices (Fig. 14.1). Adults feed on leaves, flowers and seeds. Until seeds develop, they live on the ground and feed on low vegetation and old seeds. These insects rarely cause severe damage in terms of seed loss. However, they are considered a nuisance because in autumn they may enter buildings

Boisea trivittata (Say), Boxelder Bug, Boisea rubrolineata (Barber), Western Boxelder Bug

Fig. 14.1 Nymphs of boxelder bug, Boisea trivittata, on seeds of boxelder, Acer negundo (Fort Collins, Colorado, USA).

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to overwinter. Occurrence of female (seed bearing) box elder near homes increases the risk of adults and nymphs entering homes.

Life History Boxelder bugs may have one or two generations/year, depending on location. Eggs are deposited in crevices in the bark, foliage or seeds of host trees. Nymphs feed on plant juices and molt five times. In autumn, large swarms of boxelder bug nymphs and adults may be seen. Adults overwinter.

Description of Stages Both species are virtually identical in appearance. Adults are about 12 mm long, dark brown or black with narrow red lines on the upper surface of the body. The abdomen under the wings is bright red. Nymphs are similar but lack wings (Plate 9).

Pest Management Removal of female boxelder trees from homesites eliminates the risk of boxelder bugs from entering homes to overwinter. Risk of entry into

homes can be reduced by repairing windows or door screens that are torn, screening of crawl space openings, vents and louvers, caulking of cracks around windows, doors, vents, light fixtures, pipes, conduits and air conditioners, and attaching weather-stripping to door bottoms (Furniss & Carolin 1977, Anton 1993).

COLEOPTERA Curculionidae (Weevils) Curculio Curculio is a large genus of weevils found across much of the northern hemisphere. Larvae feed in fruits and nuts of several genera of temperate broadleaf trees and can cause severe damage to nut crops (Table 14.1). They have long snouts or rostrums adapted for boring. Snouts of female adults may be twice as long as their bodies. They deposit eggs in holes bored in the nuts of host plants. Larvae feed in nuts and destroy the contents. When feeding is completed, they pupate in the soil.

Table 14.1 Representative species of Curculio (Coleoptera: Curculionidae) that damage acorns and nuts of forest trees, their distribution and hosts. Species

Distribution

Hosts

C. caryae Horn Pecan weevil C. caryatrypes (Boheman) Large chestnut weevil C. davidi Farmaire C. elephas Gyllenhall Chestnut weevil C. fulvus Chittenden C. glandium Marsham C. neocorylus Gibson Hazelnut weevil C. nucum Linnaeus C. occidentalis (Casey) Filbert weevil C. proboscideus Fabricius C. sayi Gyllenhall Small chestnut weevil C. sikkimensis (Heller) Chestnut curculio C. sulcatulus (Casey)

Eastern North America

Carya

Eastern USA

Castanea dentata

China Europe, North Africa, Near East

Castanea mollissima, Quercus Castanea sativa, Quercus

Southeastern USA Europe Eastern North America

Quercus virginiana Corylus, Quercus petraea, Q. robur Corylus

Europe British Columbia, Canada, western USA Eastern North America Eastern USA

Corylus, Quercus petraea, Q. robur Corylus californica, Lithocarpus densiflorus, Quercus Quercus Castanea dentata

China, Japan

Castanea

Eastern North America

Quercus

Sources: Browne 1968, Drooz 1985, Furniss & Carolin 1977, Shen 1993, Menu et al. 1995, Hill 2002, Venette et al. 2003, Pelsue et al. 2003, Yan Shanchun et al. 1993.

Insects of tree reproductive structures

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Curculio elephas Gyllenhal, Chestnut Weevil

Mechoris cumulatus (Vos)

Distribution Chestnut weevil is distributed over much of Europe, northern Africa and the Near East.

Distribution This weevil is found in China and, possibly, Japan and Korea.

Hosts Fruits of trees of the family Fagaceae, including Castanea sativa, C. vesca, Quercus robur and Q. suber are hosts.

Hosts Hosts are Chinese chestnut, Castanea mollissima and oaks, Quercus spp.

Importance Curculio elephas is an important pest of Castanea and Quercus in its native range. It is considered one of the most damaging pests of nuts of Castanea sativa, which are an important non-wood forest product in southern Europe. Adult feeding at the base of young nuts can cause up to 20% premature nut drop and combined larval and adult feeding can cause up to 90% crop loss. Feeding on acorns of Quercus may not impede germination but nutrient reserves required for seeding survival are severely depleted.

Importance This insect is one of several pests of chestnut orchards in China. Adults feed on branches containing chestnuts burrs, causing them to break and kill the developing chestnut. Breeding attacks occur in drying chestnut burrs on broken branches (Fig. 14.2).

Life History In southern France, adults emerge from late August to October. Adults feed on fruits of host trees for about 1 week. Females then bore holes in developing acorns or chestnuts and deposit eggs, usually one egg per fruit. Larvae feed inside the nuts and pass through four instars. When feeding is completed, larvae drop to the soil to overwinter. Some larvae may remain in the soil for an extended diapause that can last 2–3 years. Pupation usually occurs in July–August. Description of Stages Adults are 6–9 mm long, yellow-gray in color and covered with thick pubescence. The rostrum is slender and strongly curved. The rostrum of females is as long as the body. Larvae are 15 mm long when mature. The body color is white with a brown head. Pest Management Populations can be reduced by cultural and sanitation practices. In home plantings, nuts should be gathered daily as soon as they fall and stored so that emerging weevil larvae cannot enter the soil to re-infest acorns the following year. If all newly emerged larvae are destroyed for 3–4 consecutive years, weevil populations can be reduced to tolerable levels. Three to four insecticide applications, beginning in early August and repeated at 10-day intervals, will provide effective control of adults (Menu & Debouzie 1995, Venette et al. 2003).

Fig. 14.2 Stem containing developing burrs of Chinese chestnut, Castanea mollissima, cut by an adult weevil, Mechoris cumulates (Henan Province, China).

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Life History There is one generation/year. Adults are active in spring and summer. They feed on branches of host trees and later lay eggs on chestnut burrs. Larvae develop inside the chestnuts. When feeding is completed, larvae emerge and burrow into the soil to overwinter.

Description of Stages Adults are blue-black shiny weevils with 10 lines of small, light-colored spots on the elytra (Fig. 14.3). Larvae are white, legless grubs and have a light brown head capsule (Wang Gaoping et al. 2001). Pissodes validirostris Gyllenhal, Pine Cone Weevil Distribution This insect is found across pine forests of Eurasia from the UK and Spain, across Russia and east to China.

Hosts Virtually all species of Pinus indigenous to Eurasia, with the exception of Swiss stone pine,

P. cembra, are hosts. Scotch pine, P. sylvestris, is a favorite host and in Spain, P. pinea is attacked. P. contorta, native to North America and planted in portions of northern Europe, is also a host.

Importance Larvae feed in pine cones, which is an unusual habit for weevils of the genus Pissodes (see Chapter 13). A report from Inner Mongolia, China indicates reductions of 30–50% in Scotch pine seed yields and, in some cases, losses as high as 80%. In Scotland, larvae have been observed tunneling near the bases of leading shoots of P. sylvestris. This species has been suggested as a possible candidate for biological control of P. pinaster, which is considered invasive in South Africa. It is prolific and destroys all of the seeds in cones that are attacked. However, one drawback of its use as a biological control agent is that it would also attack cones of more desired pines.

Life History Females deposit eggs in cones. Larvae develop and feed inside cones and destroy the seed. Adults superficially scour the bark of host pines but

Fig. 14.3 Adult weevil, Mechoris cumulatus (Henan Province, China).

Insects of tree reproductive structures have no harmful effects. In Sweden, more adults emerge from cones of the introduced lodgepole pine, P. contorta, than from cones of its native host, P. sylvestris.

Description of Stages Adults appear much the same as other species of this genus (see Chapter 13) and would require a specialist to identify them to species.

Pest Management In China, a species of Scambus is an effective parasitoid. The percentage of larval parasitism is commonly 20–30% and sometimes as high as 45% or more, which significantly regulates the weevil population (Browne 1968, Wang Zhiying et al. 1995, Yue Shukui et al. 1997, Moran et al. 2000, Durmont & Roques 2001, Lindeloew & Bjoerkman 2001).

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Curculionidae, Subfamily Scolytinae Conophthorus (Cone Beetles) Conophthorus consists of 13 species found in North America from southern Canada through Mexico (Table 14.2, Wood S.L. 1982). All but one species, C. banksianae, infest and destroy cones of Pinus. All are monogamous. Several can cause severe damage to seed crops. Life history and habits of most species are similar. All have one generation/year and winter is spent in cones on the ground. Adults emerge in spring and fly to the tops of pines. Females bore into the petiole at the base of the cone, leaving a pitch tube at the point of attack (Fig. 14.4). She then bores toward the cone tip and is joined by a male. After mating, females construct a gallery along the entire length of the cone and lay eggs. Larvae feed in the seeds and cone tissue and pupate in

Table 14.2 Species of Pinus infesting cone beetles, Conophthorus (Coleoptera: Curculionidae: Scolytinae): their distribution and hosts. Species

Distribution

Hosts

C. apachecae Hopkins

Mexico: Chihuahua, Durango USA: Arizona Canada: Ontario USA. Michigan, Wisconsin Mexico: central Mexico Canada: Nova Scotia – Ontario USA: Maine to Minnesota and south to North Carolina USA: Missouri Mexico: countrywide USA: Arizona, Colorado, New Mexico, Texas, Utah Mexico: Hidalgo, Puebla, Veracruz n Mexico: Michoaca Mexico: Baja California Norte USA: California, Nevada, Utah Canada: British Columbia Mexico: countrywide USA: western states USA: California

Pinus engelmannii

C. banksianae (McPherson) Jack pine tip beetle C. conicolens Wood C. coniperda (Schwartz) White pine cone beetle C. echinatae Wood C. edulis Hopkins (¼ C. cembroides Wood) C. mexicanus Wood C. michoacanae Wood C. monophyllae Hopkins C. ponderosae Hopkins

C. radiatae (Hopkins) Monterrey pine cone beetle C. resinosae Hopkins Red pine cone beetle C. teocotum Wood

P. banksiana P. douglasiana, P. leiophylla P. strobus

P. echinata P. cembroides, P. discolor, P. edulis P. patula P. michoacana P. monophylla Pinus spp.

P. radiata

USA: northeastern states

P. resinosa

n Mexico: Michoaca

P. teocote

 Bores in terminal portions of twigs. n Tovar et al. 1995, Bright & Skidmore 2002. Sources: Wood S.L. 1982, Drooz 1985, Cibria

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Fig. 14.4 Conelet of bristlecone pine, Pinus aristata, with a pitch tube indicate of attack by a cone beetle of the genus Conophthorus (Pike National Forest, Colorado, USA).

cells at the end of tunnels. In late summer–early autumn, infested cones die and fall from the tree. Some adults emerge from cones and attack first-year cones but most remain in the cones until the following spring. Conophthorus coniperda (Schwartz), White Pine Cone Beetle Distribution White pine cone beetle is found in southern Canada from Nova Scotia west to Ontario and in eastern USA from Maine west to Minnesota and south to western North Carolina.

Hosts

have been destroyed in northeastern USA. Most damage occurs when second-year cones are attacked. Description of Stages Adults are smaller than most species of Conophthorus; their length is 2.4–3.0 mm and color is dark brown-black (Drooz 1977, Wood S.L. 1982). Conophthorus ponderosae Hopkins Distribution This insect ranges from southern British Columbia, Canada, throughout western USA and most of Mexico.

The host is eastern white pine, Pinus strobus.

Importance This insect attacks first- and secondyear cones, shoots and, occasionally buds and male flowers. It is one of the most damaging pests of eastern white pine seed. During some years, entire seed crops

Hosts This cone beetle attacks many species of Pinus, including P. aristata, P. arizonica, P. ayacahuite, P. contorta, P. cooperi, P. douglasiana, P. durangensis, P. flexilis, P. hartwegii, P. jeffreyi, P. lambertiana, P. leiophylla, P. montezumae, P. monticola, P. pringlei, P. pseudostrobus, P. rudis, P. strobiformis and P. washoensis.

Insects of tree reproductive structures

303

Importance C. ponderosae is an important pest of pine seed throughout its range. In Mexico it is considered the most important pest of pine seed where it damages from 40% to 87% of cone crops of P. hartwegii, P. montezumae and P. rudis.

via seed shipments. Several North American species, the most noteworthy of which is the Douglas-fir seed chalcid, M. spermatrophus, have been introduced into other locations (Hedlin et al. 1980, Roques & Skrzypczy nska 2003).

Description of Stages Adults vary in color from brown to black and range in size from 2.5 to 4.2 mm long with a robust body (Furniss & Carolin 1977, Wood 1982, Cibrian Tovar et al. 1995).

Megastigmus bipunctatus (Swederus) Distribution This chalcid is distributed from Algeria, Morocco, across Europe east to western Siberia, Kazakhstan and Uzbekistan.

HYMENOPTERA Hosts

Hosts are Juniperus communis and J. thurifera.

Torymidae Megastigmus, Seed Chalcids Megastigmus is a large genus of about 126 described species distributed throughout the world. They have a diversity of habits. Some species produce galls in host plants, others parasitize gall insects and some are facultative parasitoids that use both the insect and the gall tissue as food. Some 59 species of Megastigmus invade seeds of trees and shrubs of the families Cupressaceae, Pinaceae, Rosaceae and Taxodiaceae. Conifer seed infesting species of Megastigmus are found throughout the northern hemisphere (Table 14.3). Most species are specialized in their method of attack and feeding habits. Larval development takes place within a single seed and some larvae enter an extended diapause that may last several years. When infestations are active, there is no external evidence of damage on infested seeds and there is little or no difference in weight between infested and normal seeds. Presence of larvae in seeds is detected by radiography. Adults emerge by cutting a round exit hole in the seed coat. This provides the only external evidence that the seed has been infested. Twenty-one species of Megastigmus are recognized in Europe, North Africa and Asia Minor. They are known from 29 native conifers of genera Abies, Cedrus, Cupressus, Juniperus, Larix and Picea and 43 of the exotic conifers introduced to Europe. Some species attack seeds of plants of the family Rosaceae (Amelanchier, Rosa, Sorbus) and Anacardiaceae (Pistacia, Schinus). In North America, 11 species are known to infest seeds of conifers of the family Pinaceae. Seed infesting species of Megastigmus lend themselves to introduction and establishment in new locations

Importance Larvae feed in and destroy seeds of junipers. Studies in Spain indicate that this chalcid can attack up to one third of the cones per tree.

Description of Stages Adults range in length from 1.8 to 3.2 mm. The body color is brown, olive green or gray and black and antennae are gray-brown.

Related Species M. bipunctatus is one of about nine known species of Megastigmus that infest and destroy seeds of Juniperus in Africa, Asia and Europe. Others include M. certus, M. pingii and M. somaliensis. Additional, as yet undescribed, species may also exist. For example, in 1993 and 1997 the author observed emergence holes characteristic of Megastigmus on seeds of Juniperus polycarpos var. seravschanica in the Ziarat Forest, Balochistan Province, Pakistan (Plate 77). Adults recovered from this material were identified as Megastigmus sp. In addition, the author has observed similar exit holes in Juniperus osteosperma in western Colorado and Utah, USA but has yet to recover adults (Garcıa 1997, Roques & Skrzypczy nska 2003).

Megastigmus spermatrophus Wachtl, Douglas-Fir Seed Chalcid Distribution This seed chalcid is indigenous to western North America from eastern Alberta and British Columbia south to southern California and Arizona, USA, and northern Mexico. It has been introduced into Europe and New Zealand.

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Table 14.3 Representative conifer seed infesting species of Megastigmus (Hymenoptera: Torymidae): their distribution and hosts. Species

Distribution

Hosts

M. albifrons Walker Ponderosa pine seed chalcid

North America: southern British Columbia, Canada, south into northern Mexico North America: Canada, USA (Appalachian Mountains, Rocky Mountains, northern Pacific coast) Introduced and established in eastern, central and western Europe North Africa: Algeria, Morocco Europe east to western Siberia, Russia, Kazakhstan and Uzbekistan Kygyzstan India

Pinus ponderosa

M. atedius Walker

ska M. atlanticus Roques & Skrzypczn M. bipunctatus (Swederus) M. certus Nikol’skaya M. cupressi Matur Cypress seed fly M. grandiosus Yoshimoto Montezuma pine seed chalcid M. milleri (Milliron)

€ rster) M. pictus (Fo M. pingii Roques & Sun M. pinsapinis Hoffmeyer M. pinus Parfitt Fir seed chalcid

M. schimitscheki Novitsky M. somaliensis Hussey M. spermatrophus Wachtl Douglas-fir seed chalcid

M. wachtli Seitner

Mexico British Columbia, Canada south to California, USA Introduced and established in western Europe Eurasia, UK to western China China Western Mediterranean, Algeria, Italy, Morocco, Spain British Columbia, Canada, south to California, USA Introduced and established over most of Europe Southeastern Europe, Near East Eastern Africa North America: southern British Columbia and Alberta, Canada, south to California, Arizona, USA and northern Mexico. Introduced and established in Europe and New Zealand Mediterranean Europe and Africa, Near East

Picea spp., Pinus strobus

Cupressus spp. Juniperus communis, J. thurifera Juniperus sabina, J. semiglobosa Cupressus torulosa Pinus hartwegii, P. montezumae, P. rudis Abies

Larix Juniperus pingii Abies, Cedrus Abies, Pinus aristata

Cedrus brevifolia Juniperus procera Pseudotsuga menziesii, P. macrocarpa

Cupressus sempervirens, Cupressus spp.

ska 2003, Bouaziz & Roques 2006. Sources: Browne 1968, Hedlin et al. 1980, Rasplus et al. 2000, Roques & Skrzypczn

Hosts Douglas-fir, Pseudotsuga menziesii, and big cone Douglas-fir, P. macrocarpa are hosts.

Importance This species is a major seed destroying pest of Douglas-fir and in North America damages from 2% to 20% of the seed crop. Because infested seeds are difficult to detect, they have been shipped to other parts

of the world where Douglas-fir is planted. In Europe, it is considered a major pest of Douglas-fir seed orchards and plantations where up to 100% of the seed crops have been destroyed are reported.

Life History Adults emerge from seeds between midApril and late May depending on location and feed on

Insects of tree reproductive structures aphid honeydew and flower nectar. Sex ratio is usually 1 : 1 but unmated females can produce eggs that develop into males. Females insert their ovipositors through young cone scales and deposit a single egg into a developing seed. A female can lay about 150 eggs. Larvae hatch in 3–5 days, pass through five instars and mature in 6–8 weeks. Each larva remains in the seed and consumes the entire contents. Mature larvae overwinter in the seeds and pupate in early spring. About 20% of the larvae remain in an extended diapause for 1 year or longer. Description of Stages Females are small wasps, about 3.4 mm long, with a yellow body and red eyes. Larvae are grub-like, legless and with poorly defined body parts. Pest Management Exposure of 2-year-old Douglasfir seed to temperatures of 45oC for 27 hours and 1-year-old seed to 45oC for 33 hours effectively kills larvae inside of stored seeds with no adverse effect on germination or seedling growth (Ruth & Hedlin 1974, Hedlin et al. 1980, Roques & Skrzypcz nska 2003, Roques et al. 2006, Mailleux et al. 2008). Megastigmus wachtli Seitner Distribution The original natural range of this seed chalcid probably coincided with its primary host, Cupressus sempervirens, which is from northern Iran west to the Greek Island of Crete. Today it is found throughout Mediterranean Europe, the Near East and northern Africa where its host tree has been planted. Hosts Seeds of the Mediterranean cypress, C. sempervirens, are the primary host. Seeds of several species of Cupressus indigenous to North America and established in the Mediterranean region, including C. abramsiana, C. arizonica, C. bakeri and C. goveniana are occasional hosts. Importance Studies in Algeria indicate that this chalcid is responsible for 30–40% seed loss. It is often part of a complex of seed and cones insects that affect Cupressus in the Mediterranean region including Pseudococcyx tessulatana (Staudinger), Orsillus depressus and

305

O. maculatus, which can collectively cause over 95% seed loss.

Life History In Algeria, adults emerge over a 6-week period between early September and mid-October. Males emerge before females. Adult life span is about 30 days. Only older females can lay eggs, which are deposited in scales of first-, second- and third-year cones. Unfertilized females can lay viable eggs but all progeny are males. Sex ration varies from year to year. A larva remains inside a single seed and consumes the entire contents of the seed during its development.

Description of Stages Adults are 3.0–4.5 mm long with a dirty yellow body color (Rasplus et al. 2000, Bouaziz & Rouques 2006).

LEPIDOPTERA Tortricidae Cydia Cydia (¼ Laspeyresia) is a large and economically important genus and is represented throughout much of the world. Several species, such as the codling moth, C. pomonella, are important pests of agricultural crops. Others invade the fruiting structures of trees important in forestry. Still others, such as C. succedana, which feeds on the seeds of gorse, Ulex europaeus, a noxious weed introduced into many temperate regions of the world, are of value as biological control agents. Life stages of species of Cydia are similar in appearance and difficult to identify to species. They are small moths with a wingspan of 10–20 mm. The forewings are usually dark brown or metallic gray with distinct silver cross-bars or bands. The one exception is C. anaranjada, which has orange forewings with white bands. Larvae are 10–15 mm long when mature and white-cream in color (Table 14.4, Hedlin et al. 1980, Hill & Gourlay 2002). Cydia illutana dahuricolana Kuznetsov, Dahurian Larch Seed Moth Distribution The Dahurian larch seed moth is indigenous to northern China, Japan, Mongolia and Asian

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Table 14.4 Representative species of Cydia (Lepidoptera: Tortricidae) that damage tree seeds, their distribution and hosts. Species

Distribution

Hosts

C. anaranjada (Miller) Slash pine seed worm C. caryana (Fitch) Hickory shuckworm C. ethelinda (Meyrick) C. fagiglandana (Zeller) Beech seed worm €ffer C. illutana illutana Herrich-Scha C. illutana dahuricolana Kuznetsov Dahurian larch seed moth C. ingens (Heinrich) Longleaf pine seed worm C. latisigna Miller C. montezuma Miller

USA: Florida, southern Alabama, Georgia, Louisiana, Mississippi Southern Canada and eastern USA

Pinus elliotii, P. palustris, P. taeda

Northern India and Pakistan Europe

Picea smithiana Fagus sylvatica

Europe: Austria, Finland, France, UK Asia: China, Japan, Mongolia, Russia USA: southeastern states

Abies, Larix, Picea Larix gmelinii, L. sibirica, Picea obovata, other conifers

C. piperana (Kearfott) Ponderosa pine seed worm C. splendana (Hu¨bner) Acorn moth C. strobilella (Linnaeus) Spruce seed moth ) C. toreuta (Grote Eastern pine seed worm

Western Canada and USA, northern Mexico Europe, Near East

Mexico Mexico

Carya spp.

Pinus elliotii, P. palustris, P. taeda Pinus engelmannii, P. michoacana Pinus hartwegii, P. montezumae, P. pseudostrobus, P. rudis Pinus jeffreyi, P. ponderosa Castanea sativa, Quercus

Eurasia, North America

Picea

Eastern North America

Pinus banksiana, P. contorta, P. echinata, P. resinosa, P. taeda, P. virginiana

Sources: Hedlin et al. 1980, Drooz 1985, EPPO 2002c, Alford 2007.

Russia, including southern and eastern Siberia, Transbaikalia and the Far East. Hosts This seed worm infests seeds and cones of Larix gmelinii (¼ L. dahurica) and L. sibirica, species of Picea, especially P. obovata, Abies and other conifers. Importance Losses of 20–30% of seed production of Larix sibirica are typical. Infestations in cones of Picea can exceed 70%. Losses of seed production in Abies spp. can be as high as 60%. This insect is often associated with other insects in cones of Larix spp., which can collectively destroy 80–95% of the seed crop.

Life History There is one generation/year and adult flight usually occurs between May and late July with peak activity in June. Adult flight lasts for 2–3 weeks. Females deposit one to three eggs on developing

conelets and eggs hatch within 6–10 days. Early-instar larvae enter the cone and initially feed on seeds. Later instar larvae may feed on other parts of the cone. Damaged portions of the cone secrete resin and are unable to open and distribute seeds. Typically on)e, but occasionally two to three, larvae develop in a cone. Feeding is usually completed by August–September, when larvae emerge from infested cones, drop to the forest floor and overwinter in white cocoons. Pupation takes place in the overwintering site during the following spring (Fig. 14.5).

Pest Management Direct control with applications of chemical and biological insecticides is done in Russia and other countries when infestation levels are high.

Related Subspecies Cydia illutana illutana HerrichSch€ affer occurs in portions of Europe, including Austria,

Insects of tree reproductive structures

307

Fig. 14.5 Life histories of three species of cone and seed infesting insects of the genus Cydia (Lepidoptera: Tortricidae) (based on data from Hedlin et al. 1980, Drooz 1985, EPPO 2002c, Alford 2007).

Finland and European Russia, but populations tend to be low as is damage to seed crops. However, this subspecies is expanding its range and is now present France and the UK (EPPO 2002c.)

developing nuts. They mature in autumn and drop to the ground to overwinter in cocoons. Pupation occurs in June–July, although some larvae may diapause over a second winter (Fig. 14.5, Alford 2007).

Cydia splendana (Hübner), Acorn Moth

Cydia toreuta (Grot e), Eastern Pine Seed Worm

Distribution C. splendana is widely distributed across Europe and portions of the Near East.

Distribution This seed worm is widely distributed across eastern North America from New Brunswick, Ontario and Quebec, Canada, south to Florida and eastern Texas, USA.

Hosts Nuts and acorns of Castanea sativa, Juglans spp. and Quercus spp. are hosts.

Importance Larvae bore into and destroy all, or a portion of, the nuts of host trees. Heavy infestations can result in heavy losses of nut crops.

Hosts Most hard or yellow pines, Pinus, within its natural range are hosts, including Pinus banksiana, P. echinata, P. resinosa, P. taeda and P. virginiana. P. contorta, indigenous to western North America, is also reported as a host.

Life History There is one generation/year and adults are active from July to September. Eggs are deposited singly on leaves of host trees close to a developing nut. Eggs hatch in 10–12 days and larvae bore into the

Importance Larvae bore into the upper part of second-year cones and feed on seeds as they spiral around cone scales and may destroy a significant portion of the seed crop. Damaged seeds tend to adhere

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Forest entomology: a global perspective

to the cone scales. Infestations appear to be heaviest on open-grown pines with branches to the ground or on trees in open stands.

Life History This species has one generation/year and winter is passed as mature larvae in the woody axis of cones. Pupation occurs in spring and adults emerge 1–2 weeks later. Females lay eggs on the base of spines, the surface of the scales or the stalk of second-year cones. Young larvae bore between cone scales, enter a seed and consume the entire contents, leaving the seed filled with frass. They construct a silk-lined tunnel to the next seed. As the cone matures, larvae burrow from the last seed damaged to the cone axis to spend the winter (Fig. 14.5, Plate 78). Related Species Other species of Cydia that damage pine seed in North America include C. anaranjada (Miller) and C. ingens (Heinrich) in southeastern USA and C. piperana (Kearfott) in western Canada, the USA and northern Mexico (Table 14.4). Their life histories are similar to C. toreuta (Hedlin et al. 1980, Drooz 1985). Eucosma Larvae of several species of Eucosma bore in and destroy cones of conifers and can cause significant losses of seed crops (Table 14.5).

Eucosma cocana Kearfott, Shortleaf Pine Coneworm Distribution This species is found over much of eastern USA, from Connecticut south to Florida and west to eastern Texas. Hosts Several species of hard pines within its range, including shortleaf pine, Pinus echinata, loblolly pine, P. taeda, and possibly pitch pine, P. rigida, and Virginia pine, P. virginiana, are hosts. Importance Larvae bore in cones and destroy the contents. Larvae are able to move from cone to cone and all cones on a branch may be killed. This species is especially damaging to cones of P. echinata and, in some locations, can destroy 20% of the cone crop. It is a minor pest of P. taeda. Life History There is one generation/year. Adults emerge and are active in late April and May. Eggs are deposited in small overlapping clusters tucked between scales leaves of cone stalks. Young larvae bore into cones and feed gregariously. As they develop, they consume much of the cone’s interior and then move to neighboring cones and enter near the cone base. When feeding is completed, mature larvae drop to the ground and pupate in the soil.

Table 14.5 Representative species of cone infesting Eucosma (Lepidoptera: Tortricidae): their distribution and hosts. Species

Distribution

Hosts

E. bobana Kearfott Pin˜on coneworm E. cocana Kearfott Shortleaf pine coneworm E. impropria Meyrick

Southwestern USA, northern Mexico

Pinus cembroides, P. edulis, P. monophylla, P. pinceana Pinus echinata, P. taeda

E. monitorana Heinrich Red pine cone borer E. ponderosa Powell Western pine coneworm E. rescissoriana Heinrich Lodgepole cone borer E. tocullionana Heinrich White pine cone borer

Eastern and southern USA Asia: northeastern China, Mongolia, Russia Europe: Czech Republic, Poland, Russia, Slavic Republic (introduced) Southern Canada, northeastern and north central USA Southern British Columbia, Canada, western USA, northern Mexico Western North America from southern Alberta, Canada south to California, Idaho, Oregon and Washington Southern Canada, northeastern and north central USA

n Tovar et al. 1995, EPPO 2002c Sources: Hedlin et al. 1980, Cibria

Larix spp., L. gmelinii, L. sibirica

Pinus resinosa, P. virginiana Pinus jeffreyi, P. ponderosa Pinus contorta, P. monticola Pinus strobus, Tsuga canadensis

Insects of tree reproductive structures Description of Stages Adults have a wingspan of 18–22 mm. Forewings are tan and red-brown with variable gray patterns. Hind wings are dark gray. Larvae are 10–15 mm long when mature with a purple to pink body color and a prominent dark patch on the dorsum of the last abdominal segment (Hedlin et al. 1980).

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Pyralidae Dichocrocis (¼ Conogethes) punctiferalis Guen ee, Yellow Peach Borer Distribution This insect is widely distributed throughout Asia from India east to China, Japan, Korea and Papua New Guinea. It also occurs in Australia.

Pseudococcyx tessulatana (Staudinger) Distribution This species is found over much of the Mediterranean region, including southern Europe, northern Africa and the Near East.

Hosts Cones and seeds of trees of the family Cupressaceae, including species of Cupressus, Tetraclinis articulata and Thuja orientalis, are hosts.

Importance Larvae feed in cones and destroy seeds. In Italy, it is considered the major insect pest of Cupressus sempervirens cones in seed orchards established to produce progeny resistant to a canker disease caused by the fungus Seiridium cardinale.

Life History In Spain, there are up to three overlapping generations/year. Pupae overwinter and emerge as adults in spring as the new conelets develop. Adults mate and females deposit eggs singly on the surface of the developing conelets. Larvae hatch, feed for a short period on the conelet’s exterior and then bore into the cone and feed on the interior, including the seeds. Infested cones do not open in the normal fashion, which allows larvae to continue feeding and construct pupal chambers.

Description of Stages Adults are 12–15 mm long and gray. Forewings are rose colored with a series of black, regular transverse bands. Eggs are circular and 1.2 mm in diameter. Mature larvae are dull rose with a dark head and pronotal shield. Pupae are 7–8 mm long and red-brown.

Pest Management In Italy, sleeve cages, designed to prevent females from depositing eggs on conelets and insecticide applications, have been used to reduce seed losses (Templado 1976, Cantini & Battisti 2001, Bouaziz & Roques 2006).

Hosts Yellow peach borer attacks the fruits and seeds of a wide range of temperate and tropical plants of agricultural importance. It also feeds on at least one species of Pinus, P. massoniana, and is a pest of burrs of Castanea mollissima.

Importance Larvae bore into and tunnel in stems and fruits of host plants. Infested fruits and shoots have deposits of frass and webbing near where larvae enter. In Castanea, larvae typically enter the fruit near the point of attachment to the stem (Fig. 14.6).

Life History Yellow peach borer is a multi-generation species. In central China, there are three to four generations/year. Overwintering occurs as mature larvae in bark crevasses or in fruits in storage. Adults first appear in May and begin to lay eggs. During the growing season, duration from egg to adult is about 6 weeks.

Description of Stages Adults are colorful orangeyellow moths with a wing span of 25 cm and conspicuous black spots on the wings and body. Mature larvae are about 25 mm long, with a grey-green body color with pink highlights (Astridge & Elder 1999).

Dioryctria Larvae of many species of Dioryctria feed in cones of conifers throughout the northern hemisphere and several are important cone and seed pests. In North America, at least 16 species are known to infest conifer cones and additional, lesser known species are found in the pines forests of the Caribbean Basin and Central America. Other cone infesting species are found in

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Forest entomology: a global perspective Importance Larvae are principally cone borers but can also invade shoots of conifers. It is considered the dominant cone borer in Eurasia and causes significant loss of cone crops throughout its range. A report from India indicates that about 30% of the cones of Pinus wallichiana are infested by this insect. All seeds within infested cones are destroyed.

Life History There is one generation/year. Adult flight occurs in mid- or late summer and eggs are deposited on young cones or shoots. Larvae bore into the tissue of host trees and feed until autumn. They eject large quantities of resin mixed with frass from infested cones or shoots. Larvae overwinter in cones or shoots and pupate in a papery, silken cocoon until May. Some larvae may emerge from cones that have fallen to the ground and pupate in the soil.

Description of Stages Adults are gray and white with a wingspan of about 25 mm. Forewings are smooth and contain a pattern of dark and light gray scales with diffuse white cross-bands and flecking. Hind wings are pale gray and unmarked. Larvae are either red or green, have a brown head and thoracic shield and are about 18 mm long when mature.

Fig. 14.6 Burr of Chinese chestnut, Castanea mollissima, with boring dust indicative of attack by yellow peach borer, Dichocrocis punctiferalis (Henan Province, China).

Asia and Europe (Table 14.6, Bradley 1969, Hedlin et al. 1980, Nair 2007). Dioryctria abietella Dennis & Schiffermueller Distribution This species is widely distributed across Eurasia from the UK east to northern India, Pakistan and Thailand.

Hosts Many trees of the family Pinaceae are reported hosts. These include Abies alba, A. pindrow, A. spectabilis, Cedrus deodara, Picea abies, P. smithiana, Pinus gerardiana, P. griffithii, P. mugo, P. roxburghii, P. sylvestris, P. wallichiana and Pseudotsuga menziesii.

Related Species D. abietivorella (Grote), the fir coneworm, is widely distributed across the conifer forests of Canada, northeastern and western USA and northern Mexico where it infests species of Abies, Larix, Picea, Pinus and Pseudotsuga menziesii. This species has, at times, been considered synonymous with D. abietella (Browne 1968, Hedlin et al. 1980, Nair 2007).

Dioryctria amatella (Hulst), Southern Pine Coneworm Distribution Southern pine coneworm is found in southeastern USA from southern New Jersey and Maryland south to Florida and west to eastern Texas.

Hosts Primary hosts are the southern yellow pines including Pinus echinata, P. elliottii, P. palustris, P. taeda and P. virginiana. Bald cypress, Taxodium distichum, is also known to be attacked.

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Table 14.6 Representative species of cone infesting species of Dioryctria (Lepidoptera: Pyralidae): their distribution and hosts. Species

Distribution

Hosts

D. abietella Dennis & Schiffermueller ) D. abietivorella (Grote Fir coneworm D. amatella (Hulst) Southern pine coneworm

Eurasia from the UK east to northern India, Pakistan and Thailand Canada, northeastern and western USA, northern Mexico Southeastern USA

Abies, Cedrus, Picea, Pinus

) D. auranticella (Grote Ponderosa pine coneworm D. castanea Bradley D. clarioralis (Walker) Blister coneworm D. disclusa Heinrich Webbing coneworm D. erythropasa (Dyar)

Southern British Columbia, Canada, western USA, northern Mexico Northeastern India Eastern and southern USA, Cuba

D. merkeli Matuura & Monroe Loblolly pine coneworm D. mutatella Fuchs D. pinicolella Amsel D. pseudotsugella Monroe

Southeastern USA

D. pygmaella Ragonot Bald cypress coneworm D. reniculelloides Matuura & Monroe Spruce coneworm D. rossi Munroe

Eastern Canada and the USA Arizona, USA, Belize, Guatemala, Honduras, Mexico, Nicaragua

Europe Mexico Alberta, British Columbia, Canada, western USA, northern Mexico Southeastern USA Canada, northeastern, north central and eastern USA, northern Mexico British Columbia, Canada, western USA, northern Mexico

Abies, Larix, Picea, Pinus, Pseudotsuga menziesii Pinus echinata, P. elliottii, P. palustris, P. taeda, P. virginiana, Taxodium distichum Pinus attenuata, P. ponderosa Pinus kesiya Pinus caribaea, P. echinata, P. elliottii, P. palustris, P. taeda Picea glauca, Pinus Pinus caribaea, P. douglasiana, P. lawsonii, P.leiophylla, P. maximinoi, P. michoacana, P. oocarpa P. echinata, P. elliottii, P. palustris, P. taeda, P. virginiana Pinus sylvestris Abies religiosa, Pinus Pseudotsuga menziesii Taxodium distichum Abies, Larix laricina, Picea, Pinus contorta, Pseudotsuga menziessii, Tsuga heterophylla Pinus durangensis, P. ponderosa

n Tovar et al. 1995, Hedlin et al. 1980, Nair 2007. Sources: Bradley 1969, Cibria

Importance Larvae bore in flowers, shoots, conelets infected by southern cone rust caused by the fungus Cronartium strobilinum, stems and branches infected by fusiform rust caused by the fungus C. fusiforme, first-year conelets and second-year cones of host trees (Plate 79). Infestations in cones cause significant loss of seed crops. This species is considered the most damaging of the coneworms in southeastern USA, especially in seed orchards. Life History This insect has multiple generations. The autumn generation of adults deposits eggs on buds or fusiform rust galls. Eggs hatch and young larvae overwinter. The following spring, they bore into rust galls, buds or migrate to male and female flowers

to feed. Later they may bore into developing conelets. Subsequent overlapping generations invade rust infected conelets or developing cones.

Description of Stages Adults have forewings with a gray or dark brown ground color shaded with black and marked with sharply contrasting white patches and cross-bands. Hind wings are light brown-gray. Wingspan is about 30 mm. Mature larvae are redpurple-brown above and paler, often with green highlights below and about 20 mm long when mature.

Pest Management Infestations of this species and associated seed and cone insects of southern pine

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seed orchards are controlled with carefully timed applications of chemical sprays. Timing of sprays is done with traps baited with pheromones (Hedlin et al. 1980, Drooz 1985).

DIPTERA Cecidomyiidae, Midges Contarinia Contarinia is a large genus of midges with a wide range of habits. Many species form galls on plants of importance in agriculture and horticulture and others feed on foliage of trees and shrubs. Two North American species attack cones. Contarinia oregonensis Foote, Douglas-Fir Cone Gall Midge Distribution This midge occurs in western North America from central British Columbia and southwestern Alberta south through western USA into northern Mexico.

Hosts Cones of Douglas-fir, Pseudotsuga menziesii, are infested by this insect.

Importance This species is considered a major pest of Douglas-fir seed. It is found throughout the range of its host tree but tends to be more abundant in the Pacific coast region than in the more

arid regions of the interior west. During periods of heavy infestation, up to 100% of the seed crop may be destroyed.

Life History There is one generation/year and adults emerge in early spring just as the flowers of Douglas-fir open and are ready for pollination. Eggs are deposited near the young ovules and become intermixed with pollen grains. Eggs hatch in May and early June, about 2–3 weeks after they are laid. Larvae bore into scales of the developing cones and spend the summer. Each larva occupies a separate cell where a small swelling or gall forms. Larvae pass through three instars and during instar III, they assume a U shape inside the gall. When cones mature in autumn, the larvae drop to the litter and form cocoons, often in the remnants of old male flowers. Most larvae pupate in late winter/early spring. A portion of the larvae enter an extended diapause for 1 or more years.

Description of Stages Adults are tiny midges, dark in color and 3–4 mm long. Mature larvae are pink to orange in color and about 2.8 mm long. Pupae are orange when first formed, later become dark brown and are encased in a delicate oblong cocoon.

Related Species Larvae of the Douglas-fir cone scale midge, C. washingtoniensis Johnson, feed in cone scales of Douglas-fir but do not cause direct damage to seeds and are not considered a major pest (Hedlin et al. 1980).

Chapter 15

Insects of Wood in Use

INTRODUCTION Wooden structures, furniture, posts, poles and other wood products are subject to attack by several groups of insects. They can be extremely destructive and cause millions of US$ worth of damage. Major groups of insects that damage wood products include termites (order Isoptera), wood boring beetles (order Coleoptera) and bees, wasps and ants (order Hymenoptera).

ISOPTERA (TERMITES) Termites are among the most common of wood infesting insects. Like the wood boring beetles, termites are unable to digest cellulose and have developed symbiotic relationships with protozoa and bacteria that can digest this material. Termites are social insects with highly developed, complex societies composed of castes that perform specific functions for the colony. There are two broad classes of termites: dampwood or subterranean termites, and drywood or powder post termites. Dampwood termites can infest either living

trees or wood in use but require an external moisture source. Those that forage above ground for food protect themselves with shelter tubes, sometimes called mud tubes. These are made from particles of soil or wood and bits of debris held together by salivary secretions. The tubes may be thinly constructed or large and thick walled to accommodate many termites moving vertically between the soil and food source (Fig. 15.1). Dry wood and powder post termites, on the other hand, are able to survive on moisture available in the wood and do not require external moisture sources. Termites are both beneficial and highly destructive. In forests, they are generally beneficial because they help break down woody tissue of dead trees and recycle nutrients. However, some species attack, severely damage and even kill trees. When humans began to build wooden structures, termites found an additional food source and became pests. Termites readily invade buildings, posts, poles and furniture. They bore into wood, construct extensive tunnels and weaken its structural integrity. Termites cause millions of US$ of damage to wooden structures every year and extermination of

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Fig. 15.1 Subterranean termite shelter tubes on bark of Acacia nilotica along the Blue Nile in the Sudan.

termite colonies from homes and other buildings has become an important business. Termites also produce large quantities of methane, a greenhouse gas (Zimmerman et al. 1982, Triplehorn & Johnson 2005). Termites are most abundant in the tropics. In some locations, their numbers may range from 2000 to 4000 individuals/m2 of land area and in extreme cases there may be 10,000 individuals/m2 of land area. Some tropical species construct large and elaborate nests, which may be several meters high.

Termite Castes In a typical termite society, there are at least three castes: reproductives, workers and soldiers. The reproductive caste consists of a king and queen, usually one pair per colony, which develop into fully winged adults. They may be white or dark in color, depending on

species. The king is generally small but the queen can develop an enlarged abdomen, which increases her egg laying capacity. In some tropical species, queens can be up to 110 mm long as compared with 10–20 mm for the king. Once mature, they leave their colony of origin in a swarming flight, shed their wings and, as pairs, seek a nesting site to establish a new colony. In the early stages of colony establishment, reproductives feed their immatures and tend to the nest. The first nymphs produced develop into worker and soldier castes and workers ultimately assume the role of caring for immatures. Workers are 3–4 mm long, white, sterile and blind. They are typically the most numerous members of a colony and may work 24 hours per day for their entire lives. They gather food, construct tunnels and repair and enlarge the nest. The activity of workers is what causes termites to be pests. Soldiers have orange or amber colored heads, which are heavily sclerotized. Their bodies are white. They protect the colony against natural enemies. Soldiers have strong mandibles, which they use to crush enemies, especially ants. Some species have pointed snouts, which eject white, sticky latex to trap enemies. In some species, there may also be a secondary reproductive caste. These are wingless and replace the reproductives should they die (Triplehorn & Johnson 2005).

Termite Pest Management Subterranean termite infestations in wooden structures are best treated by professional exterminators. Two overall tactics are available: use of soil insecticides (termiticides) or baits. Treating soil with a termiticide creates a chemical barrier beneath the structure that termites will either avoid or die should they come in contact with it. If a building is constructed on a concrete slab, the termiticide is applied before the slab is poured. When infestations occur after a structure is built, termiticides can be applied by rodding, drilling or trenching. Rodding involves injecting a termiticide into the soil at intervals around the perimeter of the structure with a specially designed injection tool. Drilling involves making holes through concrete slabs, walkways, patios, floors, etc. to treat the soil beneath the structure. Trenching involves digging a shallow

Insects of wood in use trench around the base of the structure, applying the termiticide to the trench and backfill and then refilling the trench. Baits were developed during the early 1990s and provide an alternative to termiticides. Bait stations are placed around the structure to be protected. The bait is either a paper or cardboard-like material or textured cellulose impregnated with an insecticide that slowly kills the termites. Termites that feed on the bait receive a dose of active ingredient that does not kill them immediately but gives them sufficient time to feed to other members of the colony. Eventually, the colony is severely reduced in size or killed outright (Cabrera et al. 2009). Several tactics are available for treatment of drywood termites. Fumigation or tenting is a highly technical procedure that involves covering the infested structure with a gas-tight tarpaulin and releasing a fumigant inside the enclosure. The fumigant is then aerated from the structure after a set exposure time. Heat or excessive cold can also be used to treat drywood termite in a portion of a building such as an attic, porch or bedroom. Hot air is applied using high-output propane heaters. Excessive cold is used primarily for treating wall voids or similar small enclosures in a structure. Liquid nitrogen is pumped into the voids until the temperature falls to a level lethal to the termites. Local or “spot” treatments include wood injection and surface applications, microwave energy, electrocution, and wood replacement (Scheffran & Nan-Yau Su 2005).

Termites as Invasives Termites have been moved long distances by human transport of wood products, crating and dunnage. Characteristics of termites that favor their introduction and establishment in new locations include: . . .

colonies reach maturity and produce dispersal flights in ship’s, containers, and/or wood products; distributions are disjunct and sympatric with human habitation; infestations are associated with human structures.

However, introduction of old world species into the western hemisphere has been relatively unusual. To date, only five old world species have become established in the western hemisphere (Scheffrahn et al. 2004).

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Mastotermitidae Mastotermes darwiniensis Frogatt, Darwin, Great Northern or Giant Termite Distribution This termite is found in tropical regions of Australia, generally north of the Tropic of Capricorn in northern Western Australia, the Northern Territory, northern Queensland and the coastal islands.

Hosts Great northern termite has a wide host range that includes wood in use, living trees and some agricultural crops such as sugar cane.

Importance This termite is the largest in size of all of the world’s termites and the most destructive subterranean termite in northern Australia. It can invade wooden buildings, bridges, posts, poles and other plant and animal products. It also girdles and kills living trees and damages field crops such as sugar cane.

Life History The great northern termite is a subterranean non-mound builder that can also nest in boles and stems of trees and stumps. Most subterranean galleries occur in the first 30 cm of soil but can be found as deep as 70 cm. Following colonization flights and mating, queen reproductives lay eggs in masses of about 20 eggs. A colony can consist of up to 100,000 individuals. In areas of abundant food sources, colonies can subdivide into units that are independent of the main colony (Hadlington 1996).

Kalotermitidae Cryptotermes brevis (Walker), West Indian Drywood Termite Distribution West Indian drywood termite is presently the most widespread drywood termite in the subtropics and tropics. It is found throughout much of Africa, Australia, the Pacific Islands, Central and South America, the Caribbean islands and portions of North America. In Europe, it is reported from Spain and in Asia from Hong Kong. This species was first described from Jamaica in 1853. However, it is not believed to be indigenous to this region and its origin is unknown.

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Infestations were likely spread from port to port by ships carrying wood products. In the USA, it is found in the Hawaiian Islands and the coastal regions of the southeastern states. It was probably introduced into the USA at Key West, Florida, prior to 1919.

Termopsidae, Dampwood Termites Porotermes adamsoni (Froggatt)

Hosts Wood of both conifers and broadleaf species can serve as host material.

Distribution This termite is indigenous to the eastern coastal regions of Australia, from southern Queensland to South Australia, inland to the Australian Capital Territory and south to the island of Tasmania. It has been introduced into New Zealand.

Importance C. brevis is an extremely destructive termite and accounts for losses of about US$ 120 million annually in the USA and untold damage worldwide. It readily attacks small objects such as wooden furniture, headboards, cabinets and picture frames.

Hosts Species of Eucalyptus, including Eucalyptus amygdalina, E. baxteri, E. globulus, E. obliqua, E. pauciflora, E. regnans, E. sieberi, E. tenuiramis and E. viminalis are hosts. Colonies have also been found in the wood of Pinus radiata.

Life History Each year, a proportion of nymphs in a colony develop into winged reproductives that leave the colony in a series of dispersal flights. Flights occur over several weeks and this is the only time members of the colony leave the confines of their galleries. Dispersal flights are usually the first sign of an infestation. Winged reproductives fly between dusk and dawn from April through June and are attracted to lights. Smaller flights may also occur in autumn. After a brief flight, they shed their wings, segregate into male/female pairs and seek a suitable site for excavation of a nuptial chamber. Preferred sites are defects in wood such as cracks, crevices, knots or nail holes. When the chamber is large enough to accommodate the pair, its opening is sealed with an intestinal secretion. During the first 6 months, the first eggs develop into workers. Soldiers first appear during the second or third year of colony development. A colony matures in about 5 years at which time it produces its first crop of winged reproductives. Colonies can live over 10 years and contain over a 1000 individuals. Several colonies may live in close proximity and share common galleries.

Importance Porotermes adamsoni is considered an important pest in New South Wales, Tasmania and Victoria, Australia. Trees in forests and urban settings, tree stumps, poles and houses are subject to attack. This termite often builds nests in living trees and attacks are initiated in wood where there is some decay. Once established, colonies can invade sound wood of living trees. This termite also invades wood in use, especially when timbers are in direct contact with damp soil. Infested timbers often have partially eaten shells of wood remaining.

Related Species C. dudleyi Banks is native to Indonesia and Malaysia and has become established in Brazil, Colombia, Costa Rica, the Lesser Antilles and Trinidad and Tobago. C. havilandi (Sj€ostedt) is native to Africa and is established in Brazil, the Guyanas and the Caribbean islands (CAB International 1980, Scheffrahn & Nan-Yau Su 2005).

Life History P. adamsoni does not form large central colonies. Instead they live in small independent groups in the wood. While some colonies are headed by primary reproductives, many are headed by neotenics or secondary reproductives, suggesting that primary reproductives have a short life span. Colonies usually contain a few thousand individuals. They do not have a subterranean gallery system and generally have no contact with the soil. Colonies produce pipes or tunnels made of an earthen material known as “mudgut.” Pest Management Control of infestations in poles and homes involves removal of nests and spraying with chemical insecticides. Related Species Two related species with somewhat similar habits are P. quadricollis (Rambur), native to portions of Argentina and Chile, and P. planiceps

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(Sjostedt), native to the Western Cape region of South Africa (Browne 1968, Ross 2005, Elliott & deLittle n.d.).

than fire or floods combined. It causes more than 70% of the termite damage to buildings in Australia.

Rhinotermitidae

Life History A colony may consist of well over 1 million individuals and can establish nests in broken trees and stumps, behind retaining walls, under stacks of firewood and in timbers of fence posts. Colonies can construct subterranean feeding tunnels or galleries from the central nest to other food sources including live trees. Cases have been documented of feeding galleries extending from six to 10 trees, sometimes of different species (Browne 1968, Wood 1978, Hadlington & Staunton 2007).

Coptotermes Subterranean termites of the genus Coptotermes are found throughout tropical and subtropical regions of the world (Table 15.1). They invade wood products and also attack and kill live trees. Some species are extremely destructive and a few have been introduced into new locations.

Coptotermes formosanus (Shiraki), Formosan Subterranean Termite

Coptotermes acinaciformis Froggart Distribution This damp wood termite is widely distributed in Australia and has been introduced into Fiji and New Zealand.

Hosts Hosts includes species of Eucalyptus and Pinus radiata.

Distribution This termite is believed native to China, Japan and Taiwan. Over the past century it has become established in South Africa, the Hawaiian Islands and southeastern USA where it is now known to occur in Alabama, Georgia, Florida, Louisiana, Mississippi, South Carolina, Tennessee and Texas.

Importance This species is the most widely distributed and destructive timber pest in Australia. Recent surveys suggest that it causes more damage to homes

Hosts Formosan subterranean termite has a wide host range and is known to infest over 50 species of plants as well as wood in use.

Table 15.1 Representative species of Coptotermes (Isoptera: Rhinotermitidae) and their distribution. Species

Native distribution

Introduced locations

Remarks

C. acinaciformis Froggart

Australia

Fiji, New Zealand

C. formosanus (Shiraki) Formosan subterranean termite C. gestroi (Wasmann) (¼ C. havilandi Holmgren) C. lacteus Beerburrum

Asia: China, Japan, Taiwan

South Africa, USA

Asia: Indonesia, Malaysia, Thailand Australia

Brazil, Caribbean Basin, USA (Florida)

Considered Australia’s most damaging termite. Invades both live trees and wood in use Considered one of the world’s most destructive termites. Invades both live trees and wood in use Invades both buildings and live trees

C. sjostedti Holmgren

Africa

Caribbean Basin (Guadeloupe)

C. testaceus Linnaeus

Caribbean Islands, South America

Sources: Browne 1968, Scheffrahn et al. 2004, Jenkins et al. 2007.

Builds mounds up to 2 m tall. Damages wood in use in coastal regions Detected in a mangrove forest in Guadeloupe. Damage potential questionable Known to attack and kill Hevea brasiliensis trees after they have been weakened by foliage diseases

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Importance Formosan termite is one of the world’s most destructive termites. It has been referred to as a “super termite” because it can build large colonies that cover an area of roughly 100 m2, infests a wide range of structures, including boats and high-rise condominiums, and consumes wood at a rapid rate. This termite is aggressive both in its feeding habits and foraging tenacity. One established, colonies can cause severe structural damage to wooden homes in as little as 2 years. Although it prefers to establish colonies that can maintain contact with the ground to provide moisture, ground contact is not necessary. Formosan termites have become a major problem in the city of New Orleans, Louisiana, USA. They are believed to infest about 30% of the historic live oaks, Quercus virginiana, in the city as well as many residences in the famous French Quarter. New Orleans is a perfect habitat for Formosan termite infestations because of its high relative humidity and closely spaced older homes. This termite causes about US$ 300 million dollars worth of damage annually in New Orleans alone. Infestations of this termite in the levees that protect the city from flooding may have been, at least in part, responsible for their failure during Hurricane Katrina in 2005.

Life History In southeastern USA, swarms of winged termites (alates) usually occur from April to July on calm, warm and humid evenings. They are large and can consist of tens of thousands of individuals. The swarming termites are attracted to lights and can be found around windows, light fixtures, window sills and other well-lit areas. After landing on the ground, they break off their wings and search for a mate. When a mate is found, male and female (king and queen) search for a crevice in damp ground or wood, hollow out a small chamber and crawl inside. Within a few days, the queen lays eggs and hatching brood is fed by the king and queen. Immatures develop into soldiers and workers. If the king or queen dies or the colony becomes large, secondary reproductives may appear and lay eggs. Formosan termite colonies build large nests that are usually subterranean but can be above ground and not connected to the soil. The nests are built from a material known as carton, which workers make from a mixture of soil, chewed wood, other plant matter, their saliva and feces. Nests can be large with some containing an estimated 8 million termites (Henderson 2008, Cabrera et al. 2009).

Reticulitermes Reticulitermes is a large genus of subterranean termites with species distributed over much of the northern hemisphere (Table 15.2). Most species are damaging to wood products. The taxonomy of this group is still unclear and is being addressed with analysis of mitochondrial DNA and subject to revision. A 2005 study, for example, concluded that the eastern subterranean termite, R. flavipes, of North America and a European species, R. santonensis, are synonymous (Clement et al. 2001, Austin et al. 2005). Reticulitermes flavipes (Kollar), Eastern Subterranean Termite Distribution This species is the most widely distributed termite in North America and occurs from Ontario, Canada through the eastern, central and southern USA, south to Mexico and Guatemala. It apparently has been introduced and become established in Austria, France, Germany, Chile and Uruguay. Infestations have also been detected in Oregon, USA and on Grand Bahama Island in the Caribbean. It is most abundant in regions with warm, humid climates.

Hosts Eastern subterranean termite can invade wood in use of virtually any species.

Importance Eastern subterranean termite is the most damaging indigenous termite in North America. It can invade wooden structures, utility poles, fence posts, furniture, books, paper, plastic and buried utility cables (Figs 15.2 & 15.3). It also occasionally attacks living trees, shrubs and field crops.

Life History Swarms of reproductives typically appear in early spring. After a short flight, they loose their wings and pair off. The king and queen seek a suitable nesting site in moist soil near a source of wood and start a new colony. After mating, the queen lays eggs, which hatch in 80–90 days. Most eggs hatch into worker nymphs. Workers and soldiers mature in about 1 year and reproductives require 2 years to mature (Plate 80). Workers construct earthen tubes on foundations, water pipes, sills and joists to provide a connection from the moist soil to wood, their food

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Table 15.2 Representative species of Reticulitermes (Isoptera: Rhinotermitidae) indigenous to Europe, the Near East and North America and their distributions. Species

Distribution

R. arenicola Goellner

North America: northwestern Indiana, southwestern Michigan and Boston, Massachusetts, USA Europe: Balkan region Europe: southwestern France, northeastern and central Spain Near East: Israel North America: East of the Missouri River from southern Ontario, Canada south to Florida, Mexico, Guatemala, Europe: Austria, France, Germany (introduced) South America: Chile, Uruguay (introduced) Europe: southwestern France, northwestern and southern Spain and Portugal North America: southeastern USA North America: Pacific coast region

ment) R. balkanensis (Cle ment R. banyulensis Cle R. clypeatus Lash R. flavipes Kollar Eastern subterranean termite (¼ R. santonensis Feytuad) ment R. grassei Cle R. hageni Banks R. hesperus Banks Western subterranean termite R. lucifugus Rossi Mediterranean termite R. tibialis Banks Dryland subterranean termite R. virginicus Banks Dark southern subterranean termite

Europe: southeastern France, Italy, Turkey North America: southwestern Great Basin and midwestern regions of the USA North America: southeastern USA

ment et al. 2001, Luchetti et al. 2004, Weimin Ye et al. 2004, Austin et al. 2005. Sources: Cle

Fig. 15.2 Damage to construction lumber by subterranean termites of the genus Reticulitermes.

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Forest entomology: a global perspective system for the nest. Some build open chimneys or vent holes in their mounds, whereas others build completely enclosed mounds that exchange gases through porous thin-walled tunnels.

Macrotermes bellicosus Smeathman Distribution M. bellicosus is widely distributed in tropical Africa, including Ghana, Kenya, Malawi, Nigeria, Tanzania, Uganda and Zambia.

Hosts Trees, agricultural plants and grasses can serve as food for the fungus gardens of this termite. Decaying grasses are a food source in the absence of wood.

Importance Structural lumber is damaged throughout this termite’s range. In addition, it is reported to be a pest of cacao in Ghana and of cloves on the island of Zanzibar. Living trees, including species of Eucalyptus and neem, Azadirachta indica, are damaged in Nigeria. This termite can invade root systems and lower branches of many forest plantation trees and will girdle and kill large trees. The swarming alates reportedly have high nutritional value and are trapped and sold as a food item in markets (Fig. 15.4).

Fig. 15.3 Heavy damage to the interior of a tree bole by subterranean termites of the genus Reticulitermes.

source (Smith & Johnston 1970, Scheffrahn et al. 1999, Scheffrahn et al. 2004, Austin et al. 2005, McKern et al. 2006).

Termitidae Macrotermes Termites of the genus Macrotermes are widely distributed throughout Africa and Southeast Asia. All are subterranean and cultivate fungi that aid in digestion of cellulose. They construct large mounds, the largest non-human-made structures in the world. The mounds enclose a network of tunnels and form a ventilation

Life History M. bellicosus is one of the world’s largest termites and has a complex, highly evolved colonial organization. It is a fungus growing species, which builds irregular, open spire-like mounds that can be up to 9 m tall.

Related Species Other species of Macrotermes indigenous to Africa include M. goliath Sjoetedt, found in the highlands of east Africa and across much of the continent south of the Congo rain forest. M. natalensis Haviland is widely distributed south of the Sahara (Browne 1968, Wagner et al. 2008). Nasutitermes Species of Nasutitermes are found throughout the tropics and subtropics. Many species are arboreal and nest on or above ground level. Twelve species are known to

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Fig. 15.4 Swarming alates of Macrotermes bellicosus have a high nutritional value and are commonly sold in markets in Africa as a food item (Lusaka, Zambia, photo by Maggie Beveridge).

damage buildings. Soldiers are characterized by having pointed heads (Fig. 15.5, Scheffrahn et al. 2002).

Nasutitermes walkeri (Hill) Distribution This species is indigenous to Australia and occurs along the eastern coast of the country. It has been intercepted in New Zealand but is not established.

Hosts trees.

N. walkeri attacks wood of many broadleaf

Importance Decayed and weathered wood is subject to attack, especially if it is in contact with the soil. This termite is especially damaging in humid environments where there is poor ventilation.

Life History Castes include queen, king, soldiers, workers and secondary reproductives. Large arboreal nests are constructed in trees on the main bole, a fork or on larger branches. The arboreal nest is connected to another part of the colony in a subterranean environment, usually in the root system of the same tree. Shelter tubes on the surface of the tree connect the two portions of the colony. Subterranean tunnels radiate from the base of trees to various food sources.

Pest Management Physical removal of the arboreal nest and flooding the subterranean nest in the root system can destroy the colony.

Related Species N. graveolus (Hill) is also indigenous to the east coast of Australia and across the northern part of the country west to Darwin. N. costalis

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Fig. 15.5 Soldiers of termites of the genus Nasutitermes (Guatemala, photo by David Cappaert, courtesy of www. forestryimages. org).

(Holmgren) is widely distributed in the West Indies and infestations were discovered in East Kilbride, Scotland in 2000 and in Florida, USA in 2002. N. nigriceps (Haldeman) is found from Panama north to Mexico and on the islands of Jamaica and Grand Cayman. N. acajutlae (Holmgren) occurs in Puerto Rico, the British and US Virgin Islands, Trinidad and Guyana (Hadlington 1996, Thorne et al. 1996, Thompson & Herbert 1998, Scheffrahn et al. 2002).

are typically subterranean although they may also build nests in trees. Like Macrotermes, some species construct vertical, hollow chimneys made of clay that can rise 2 m above the surface of the ground (Fig. 15.6). The chimney opens into a large subterranean airspace and provides ventilation for the nest (Turner 1994, Aanen & Eggleston 2005).

Odontotermes badius (Haviland), Crater Termite

Odontotermes

Distribution This termite is found over much of subSaharan Africa.

Odontotermes consists of about 78 species distributed across tropical Africa and Asia. They cultivate fungi inside their nests for food. The fungus gardens are continuously provided with plant materials such as grasses, leaves and bits of wood by workers. Their nests

Hosts Many food sources are used to nurture subterranean fungus gardens cultivated by this species for food, including grasses, leaves and wood.

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underground cell. Female reproductives can lay one egg every 2 seconds. When the colony is well established, vertical shafts are built, which open at the start of the rains. A nest may be 2 m in diameter and can contain many fungus gardens.

Pest Management Direct control of this termite involves pouring contact insecticides down each of the vertical shafts during the rainy season.

Related Species O. latericius (Haviland) is abundant in dead wood in eastern and southern Africa and also damages wooden structures. The nests of this species are equipped with clay chimneys (Browne 1968, Schabel 2006, Hall 2008).

COLEOPTERA Bostrichidae, Subfamily Lyctinae (Powder Post Beetles) Lyctus

Fig. 15.6 A termite chimney constructed by an unidentified termite of the family Termitidae (southern Kenya).

Importance This species is considered one of the most damaging termites in eastern and southern Africa. Nests are often found under wooden buildings. Damage can occur to wooden structures, tree roots, young plantation trees and tree nurseries. In Kenya, this termite is also considered a pest of pastures. Unlike other Odontotermes, this species builds inconspicuous nests that may be visible only by the presence of a rise in soil elevation and small soil craters. Nests lack the chimneys and conspicuous ventilation shafts built by other species of Odontotermes.

Life History Colonies are established by reproductives that fly at dusk at the onset of rainy seasons. They segregate by pairs, shed their wings and build an

Beetles of the genus Lyctus are small, red-brown, brown or black and range from 2 to 7.5 mm long. They have a prominent head not covered by the prothorax with short antennae with a two-segmented club. They are cosmopolitan species, found throughout the world and each geographic region has its complex of indigenous and introduced species. Larvae usually bore in wood of ring porous tree such as ash, Fraxinus spp., or oak, Quercus spp. (Fig 15.7). In some parts of the world, powder post beetles are second to termites in their ability to damage wood.

Lyctus brunneus (Stephens), Brown Lyctus Beetle Distribution This powder post beetle is cosmopolitan in its distribution and is reported from Africa, Australia, Europe and North America.

Hosts Sapwood of wood harvested from broadleaf trees, including species of Eucalyptus, Fraxinus, Quercus and Ulmus, is invaded.

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Fig. 15.7 Sapwood of gambel oak, Quercus gambelii, with a heavy infestation of powder post beetle of the genus Lyctus (Dolores County, Colorado, USA).

Importance Infestations can begin prior to the delivery of lumber to construction sites and continue after the lumber is incorporated into the building. This beetle is also a common pest of furniture, rafters and flooring and attacks bamboo furniture. Damage is easily overlooked during early stages of infestation. In later stages, damaged wood may be reduced to a powdered mass inside a thin intact skin of wood. In Africa, it is considered the most destructive of the lycinid beetles.

brown oval spots on either side of the body. They have three pairs of legs and are up to 6 mm long when mature (Drooz 1985, Schabel 2006, Elliot & deLittle n.d.).

Life History Adults emerge from infested wood during summer and deposit eggs on pores of susceptible wood. Eggs hatch 2–3 weeks later and larvae bore into the sapwood. They generally feed parallel to the wood grain and fill their tunnels with fine, tightly packed frass. About 1 year is spent in the larval stage and pupation takes place just below the surface of the infested wood. Pupation lasts from 2 to 4 weeks.

Hosts L. planicollis feeds on seasoned wood of ring porous trees, including species of Carya, Fraxinus and Quercus.

Description of Stages Adults are 4–7 mm long and red-brown to black in color, flattened and elongated. Antennae are clubbed. Larvae are pale cream with light

Lyctus planicollis Leconte, Southern Lyctus Beetle Distribution This insect is native to North America and is most common in southeastern USA.

Importance This powder post beetle readily invades dry, seasoned wood less than 5 years old. Plywood, wooden crating, furniture, tool handles, picture frames and baskets are subject to attack.

Life History One year is usually required to complete a generation but life span can vary from 6 months to

Insects of wood in use 4 years. Development is faster in heated buildings. Larvae cause the most damage and account for most of the variation in length of life cycle. Adults mate shortly after emerging from wood. Females lay eggs in pores, small cracks and crevices on the surface of bare, unfinished wood. They lay from 15 to 50 eggs over a 1week period and eggs hatch in 1–3 weeks. Young larvae bore deeper into the wood. Early instars tend to feed parallel to the wood grain but later instars feed across the grain. They fill their tunnels with a powderlike frass that gives them their common name. Larvae may tunnel all the way to the surface and eat away most of the material but will not break the wood surface. Therefore, infested wood has a paper-like outer shell that is easily broken. Generally, larvae confine their feeding to the sapwood. In warm buildings they can develop in all seasons, but most activity occurs in spring and summer. Larvae bore close to the surface to pupate and pupation lasts from 12 to 30 days. Adults chew to the surface and leave circular exit holes 0.8–1.5 mm in diameter. Description of Stages Adults are red-brown, sometimes black and 4–6 mm long. The body is elongated and slightly flattened. The head is prominent and not covered by the pronotum. Antennae have 11 segments and the last two segments are broadened into a club. The elytra are striated with double rows of small puncture marks between the striations. A sparse, short, yellowish pubescence covers the body. Larvae are yellow-white, C-shaped and about 7.5 mm long with an enlarged spiracle on the eighth abdominal segment. They have well-developed legs. Eggs are translucent, about 1 mm long and cylindrical with rounded ends. Related Species The western lyctus beetle, L. cavicollis LeConte occurs in western North America but has expanded its range. In California, it attacks the wood of introduced eucalyptus trees (Furniss & Carolin 1977, Drooz 1985, Brammer 2003).

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Hosts Favorite hosts are wood of spruce, Picea spp. and pine, Pinus spp. The wood of Douglas-fir, Pseudotsuga menziesii, and western hemlock, Tsuga heterophylla, is also invaded. This species can also invade wood of temperate broadleaf species but is rarely found in wood of tropical broadleaf trees.

Importance Larvae produce a network of galleries in wood and wood products 1–2 mm in diameter. Despite its common name, this insect is more of a pest of structural timber than furniture. However, it can enter homes via infested furniture and then spread to areas of exposed, unfinished wood. It also infests twigs and branches of dead trees. Life History Depending on location, adults emerge from May to July and live from 2 to 4 weeks. They fly during warm weather and retreat into their exit holes to mate. Females lay 20–80 eggs, which they deposit individually between pieces of wood, in old exit holes or pupation chambers. Eggs hatch in 3–4 weeks at 65% relative humidity. Larvae feed for about 2 years at 22–24  C and 70% relative humidity but will live for 3–5 years under drier conditions. When breeding in branches of dead trees, length of a generation is about 1 year. Description of Stages Adults are 2.5–6 mm long, cylindrical and red-brown. Elytra have punctures in longitudinal rows and the last three antennal segments are longer than the others. Mature larvae are about 6 mm long, yellow-white with dark brown mouthparts. Bodies are covered with a scattering of yellow hairs (French 1968, Furniss & Carolin 1977, Drooz 1985.

Pest Management should be replaced.

Infested

structural

lumber

Anobiidae (Death Watch and Spider Beetles) Anobium punctatum (DeGeer), Furniture Beetle

Buprestidae (Flat Headed Wood Borers)

Distribution This beetle is probably of European origin, where it is widely distributed. It also occurs in North America and has been introduced into Australia and New Zealand.

Buprestis aurulenta Linnaeus, Golden Buprestrid Distribution Golden buprestid is found in British Columbia, Canada and western USA.

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Hosts Hosts are conifers, including species of Abies, Picea, Pinus and Pseudotsuga.

Importance In forests, the larvae mine in and around fire scars and mechanical injuries on live trees. This insect also infests wood in buildings, especially in the Pacific northwest (Oregon, Washington). Larvae mine timbers and boards. Infestations may originate in the forest, in lumberyards or in exposed portions of wooden structures.

Life History In the dry wood of buildings, development from egg to adult may be prolonged from 30 to 50 years. In forests, the time required to complete a generation is much shorter but still requires several years.

Description of Stages Adults are attractive beetles, iridescent blue or green in color with the margins of the elytra bordered with copper. They range in size from 12 to 20 mm (Plate 81, Furniss & Carolin 1977).

earlier than those infesting wood in cooler basements. Eggs are laid in fan-shaped clusters in holes or tight crevasses. Stacked lumber and cracks and checks in the wood of houses are especially desirable sites for attack. Young larvae feed near the surface of the wood and older larvae bore deeper. They produce tunnels filled with tightly packed frass. Larvae seldom break through the surface of the wood. Severely damaged timbers, on the brink of collapse, may appear perfectly sound.

Description of Stages Adults are slightly flattened, brown-black beetles and range in length from 8 to 20 mm. The head and thorax are covered with gray hairs. There are two darker spots on the thorax and faint lighter markings on the elytra. Infestations can be treated by replacing infested lumber and fumigating homes.

Pest Management Heavy infestations in buildings require replacement of damaged wood (Becker 1976, Drooz 1985).

HYMENOPTERA Cerambycidae (Round Headed wood Borers) Hylotrupes bajulus Linnaeus, Old-House Borer Distribution This wood borer is native to Europe and has been introduced to other regions of the world, including Asia, Australia and North America.

Hosts Lumber of seasoned conifers, especially pine, Pinus spp., and spruce, Picea spp., are subject to attack.

Importance Contrary to its name, the beetle is usually found in newly constructed wooden buildings. Structural timbers, framing and other wood can be weakened by larval mines.

Life History In the southern portions of its range, 3–5 years may be required to complete a generation and further north an additional 2 years may be required. The life cycle may vary in a single building. For example, adults in warm attics may emerge 2 years

Formicidae Camponotus The genus Camponotus consists of about 1000 species found throughout the world in both temperate and tropical ecosystems. Some species are known as carpenter ants and nest in damp wood whereas others nest in soil. Wood nesting species do not feed on the wood but construct extensive networks of galleries. Nests established in wooden buildings cause structural damage and result in high costs for repair, replacement of damaged wood and control. In North America alone, 23 species of Camponotus invade wood in use and seven are capable of causing significant damage (Robinson 2005). Camponotus pennsylvanicus (DeGeer), Black Carpenter Ant Distribution This ant is widely distributed in eastern North America, from Quebec and Ontario, Canada and in eastern USA west to North Dakota.

Insects of wood in use Hosts Nesting occurs in the wood of many species temperate broadleaf trees and conifers.

Importance Live and dead standing trees, decaying logs and stumps, poles and wooden buildings are used as nesting sites. Homes and other wooden buildings can be invaded by black carpenter ants from nearby nests in forested areas. Entry is gained through openings around the building’s foundation or from tree branches in contact with the building. The most commonly infested parts of wooden buildings are support timbers, porch pillars, sills, girders, joists, studs, window casings and external trim. Galleries are similar in appearance to those produced by termites except that they run across the grain and are free of frass and sawdust (Figs 15.8 & 15.9).

Life History Black carpenter ants are social insects and have worker and reproductive castes. Reproductive males and females engage in nuptial flights from May to late June. Mated females then establish nests in cavities in wood and seal themselves inside. They lay eggs and rear their first broods of workers

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on salivary secretions and are smaller than normal because they are inadequately fed. Subsequent broods are fed by workers and resulting individuals are larger. These workers expand the nest by constructing a network of galleries in the wood. Nests must be established for at least 3 years before a new crop of reproductives appear. These ants feed on live and dead insects, honeydew, sap, juices of ripened fruits and trash. They also feed on a variety of foods prepared in homes including sweets, raw and cooked meat and fruits. Colonies established for several years usually consist of a singe reproductive female, large numbers of winged males and virgin females, and thousands of workers.

Description of Stages Black carpenter ants are among the largest of ants and workers range from 6 to13 mm long. Body color is usually black but some individuals may have red highlights on the body and red legs.

Pest Management Homes that are well built, with concrete foundations and good clearance, are generally

Fig. 15.8 Spruce lumber damaged by black carpenter ants, Camponotus pennsylvanicus.

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Fig. 15.9 Heavy damage to wood by black carpenter ants, Camponotus pennsylvanicus.

safe from invasion by carpenter ants. Removal of woody debris from near homesites, avoiding transport of infested wood into wooden structures and cutting back branches in contact with wooden buildings will prevent establishment of infestations. Infestations in wooden buildings require removal of nesting sites and replacement of damaged wood. Related Species C. modoc Wheeler occurs in the Pacific northwest region of Canada and the USA and is a structural pest. C. ligniperdus (Latrielle) is a European species found from Sweden south to Italy

and Spain and east to western Russia and is considered one of that region’s largest ants. C. compressus Fabricius, native to India and Pakistan, nests in trees and is considered beneficial because it tends the lac insect, Kerria lacca, and protects it from predators. C. saundersi Emery is native to southeastern Asia and is one of several species whose workers have greatly enlarged mandibular glands. They can release their contents and spray a toxic substance to attack natural enemies and are known as “exploding ants” (Browne 1968, Drooz 1985, Robinson 2005).

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Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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Subject and Taxonomic Index A Abies 8, 34, 102, 122, 128, 129, 144, 146, 161, 180, 189, 193, 195, 196, 208, 209, 213, 214, 215, 216, 220, 221, 222, 223, 228, 231, 238, 240, 256, 277, 280, 285, 303, 304, 306, 310, 311, 326 Abies alba 122, 210, 222, 230, 240, 310 Abies amabalis (Pacific silver fir) 122, 240, 241 Abies balsamea 120, 121, 214, 230, 240 Abies borisii-regis (Bulgarian fir) 230 Abies cilicia 122, 230 Abies concolor (white fir) 11, 122, 146, 184, 186, 189, 252 Abies fraseri 214, 240 Abies grandis (grand fir) 122, 146, 186, 189, 240, 241 Abies holophylla (Manchurian fir) 285 Abies lasiocarpa (subalpine fir) 7, 24, 122, 125, 186, 189, 230, 240, 241 Abies magnifica (red fir) 117 Abies nephrolepis 285 Abies nordmanndiana (Nordmann fir) 191, 230, 256 Abies pindrow 310 Abies religiosa (oyamel, sacred fir) 44, 124, 125, 186, 189, 311 Abies sachalinensis (Sakhalin fir) 120, 230 Abies sibirica 216 Abies spectabilis 310 Acacia 114 Acacia mearnsii 137 Acacia mollisima 207

Acacia nilotica 249, 314 Acacia senegal (gum Arabic) 207 Acer (maple) 6, 114, 123, 124, 125, 140, 141, 144, 145, 146, 148, 200, 210, 214, 226, 227, 250, 297 Acer negundo (boxelder) 227, 297 Acer orientalis 273 Acromyrmex 167 Acromyrmex ambiguus 168 Acromyrmex asperus 168 Acromyrmex coronatus 168 Acromyrmex crossispinus 168 Acromyrmex diasi 168 Acromyrmex diseiger 168 Acromyrmex gallardoi 168 Acromyrmex heyeri 168 Acromyrmex hispidus 168 Acromyrmex landolti 168 Acromyrmex laticeps 168 Acromyrmex laticeps nigosetosus 168 Acromyrmex lobicornis 168 Acromyrmex lundi 168 Acromyrmex lrystrix 168 Acromyrmex mesopotamicus 168 Acromyrmex niger 168 Acromyrmex octospinosus 168 Acromyrmex rugosus 168 Acromyrmex subterraneus 168 Acromyrmex pulvercus 168 Acromyrmex silvestris 168 Acromyrmex striatus 168 Acromyrmex versicolor 168 Adelges 102, 239–241, 256–257 Adelges abietis (eastern spruce gall aphid) 256 Adelges cooleyi (Cooley spruce gall adelgid) 256–257

Adelges piceae (balsam woolly adelgid) 36, 240–242 Adelges glandulae 256 Adelges isedakii 256 Adelges japonicus 256 Adelges knucheli 256 Adelges lapponicus 256 Adelges lariciatus 256 Adelges laricis 256 Adelges nebrodensis 256 Adelges nordmanniana 256 Adelges pectinata 256 Adelges piceae 240–241 Adelges prelli 256 Adelges tardiodes 256 Adelges torii 256 Adelges tsugae 36, 76, 95, 209, 239–240 Adelges viridis 256 Adelgidae 102, 239–242, 256–258 Adenium 139 Aegle marmelos 249 Aesclepias (milkweed) 44 Aesculus 6 Aesculus hippocastanum (horse chestnut) 42, 226 Afrotachardina 249 Agathis Agathis brownii 217 Agathis pumila 117 Agrilus 203–207 Agrilus andersoni 203 Agrilus angelicus 204 Agrilus anxius (bronze birch borer) 204–205 Agrilus bilineatus (twolined chestnut borer) 24, 27, 204, 205 Agrilus coxalis (goldenspotted oak borer) 204, 205

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

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356

Subject and Taxonomic Index

Agrilus (Continued ) Agrilus granulatus 203 Agrilus howdenorum 203 Agrilus hyperici 204 Agrilus pannonicus 24, 27 Agrilus liragus (bronze poplar borer) 204 Agrilus near grandis 207 Agrilus pannonicus 204, 205 Agrilus planipennis (emerald ash borer) 36, 42, 204, 206–207 Agrilus roscidus 204 Agrilus ruficollis (red necked cane borer) 203 Agrilus sp. 207 Agrilus turnbowi 203 Agrilus viridis 204 agroforestry 39–40 Agromyzidae (leaf-miner flies) 111, 271–272 Aletergystia cadambae (teak carpenterworm) 224 Allamaranda cathartica 139 Allokermes 103, 248 Allokermes kingii (northern red oak kermes) 248 Alnus (alder) 8, 120, 125, 130, 131, 143, 145, 146, 148, 196, 214 Alnus glutinosa (common alder, European alder) 204 Alsophila pometaria (fall cankerworm) 124, 125 Amalanchier 125, 146, 220, 303 ambrosia beetles 173–174, 196–202 Amphibolips confluenta (oak apple) 262 Amylostereum 230–231 Amylostereum aerolatum 32, 38, 84, 228, 230, 231 Amylosterium chailetti 231 Anacardiaceae 303 Anacardium occidentale (cashew) 249 Anagyrus kamali 245 Anaphes nitens 155 Anastus sp. 128 anholcyclic 102, 257 Anodopetalum biglandosum 201 Andricus californicus 263 Anobiidae (death watch or spider beetles) 104, 325 Anobium punctatum (furniture beetle) 325 Anoplophora 44, 210–213 Anoplophora chinensis (citrus longhorn beetle) 211–213

Anoplophora glabripennis (Asian longhorn beetle) 36, 210–211 Anoxia matutinalis matutinalus 151 Antheraea 43 Antheraea pernyi (tussar moth) 43 Antheraea yamamai 43 Antherosperma moschatum 201 Aonidiella orientalis (oriental scale) 42, 249–250 Apate 273 Apate distincta 274 Apate monachus 273–274 Apate terebrans 274 Aphidecta obliterata 241 Aphididae (aphids) 102, 236, 255 Aphidoletes thompsoni 241 Apis mellifera (honey bee) 42, 107 Araucaria cunninghamii 217, 280 Arbutus 130 Arctiidae (tiger moths) 110, 148 Argidae 107, 158 Armillaria 24, 27 Arthropoda 97 Atta 168 Atta bispherica 169 Atta capiguara 169 Atta cephalodes 169 Atta colombica 169 Atta goiania 169 Atta insularis 169 Atta laevigata 169 Atta mexicana 169 Atta opaciceps 169 Atta robusta 169 Atta saltensis 169 Atta sexdens 169 Atta silvi 169 Atta texana (town ant, Texas leaf cutting ant) 169–171 Atta vollenweiler 169 Austrotacharidia 249 Austrocedrus chilensis 237 Avetianella longoi 218 Azadirachta indica (neem) 41, 42, 88, 249, 250, 273, 320 B Baccharis 216–217 Bacillus thuringiensis (Bt) 55, 81, 82, 89, 135, 141, 148 bark beetles 10, 106, 173–196 Beauvaria bassiana 23, 83, 128

Betula (birch) 8, 120, 123, 124, 125, 130, 131, 135, 141, 143, 144, 145, 146, 148, 153, 165, 186, 196, 204, 214, 220, 227, 250 Betula allegheniensis (yellow birch) 165, 204 Betula ermanii 165 Betula glandulifera 165 Betula maximowicziana 165 Betula papyrifera (paper birch) 165, 204 Betula pendula 165 Betula platyphylla 165 Betula populifolia (gray birch) 165 Betula turkestanica 165 Betulaceae 8 Biorhiza pallida 263 Bischofia javanica 114 Bissorah gall 264 biological control 43 biome 2 Boisea 98, 297 Boisea rubrolineata (western box elder bug) 297 Boisea trivittata (box elder bug) 98, 297 Bostrichidae (twig borers and powder post beetles) 104, 273, 323–325 Bombyx mori (silkworm) 42 Broussonetia papyrifera (paper mulberry) 212 Bupalis piniarius (pine looper) 37, 84, 125 Buprestidae (flat headed wood borers) 104, 105, 203–210, 325–326 Buprestis 207–208 Buprestis apricans (turpentine borer) 207–208 Buprestis aurulenta (golden buprestid) 325–326 Buprestis novemmacula 39, 208 Bursaphelenchus 32 Bursaphelenchus xylophilis (pinewood nematode) 33, 60, 84, 213 Butea monosperma (flame of the forest) 249 Buxus sempervirens (boxwood) 250 C Cactoblastis cactorum 109 Caesalpina echinata (Brazilwood) 200 Cajanus cajan 212

Subject and Taxonomic Index Calcocedrus decurrens (incense cedar) 296 Callibracon limbatus 218 Callirhytis quercuspomiformis 263 Callitris 237 Calomicrus apicalis 152 Calotropis procera (giant milkweed) 139 Camelia sinensis (tea) 212 Cameraria 116 Cameraria cincinnatiella 116 Cameraria hamadryadella 116 Cameraria ohridella (horse-chestnut leaf miner) 42, 114–115 Camponotus 326 Camponotus compressus 328 Camponotus ligniperdus 328 Camponotus modoc 328 Camponotus pennsylvanicus (black carpenter ant) 326–328 Camponotus saundersi 328 Caragana 141 Caragana korshinskii (peashrub) 187 carbon 1 carbon dioxide (CO2) 27, 28 Cargola arena 38 carmosine 249 Carpinus (hornbeam) 123, 135, 144, 214, 227 Carya 6, 124, 125, 148, 217, 298, 306, 324 Carya illinoiensis (pecan) 197–198, 212 Cassia fistula (golden shower tree) 249 Castanea 135, 204, 226, 265, 298, 306, 307 Castanea alnifolia 265 Castanea crenata (Japanese chestnut) 265 Castanea crenata x mollisima 265 Castanea dentata (American chestnut) 205, 265, 298 Castanea mollisima (Chinese chestnut) 265, 298, 299, 309, 310 Castanea pumila 265 Castanea sativa (European chestnut) 140, 206, 265, 298, 299, 307 Castanea vesca (Spanish chestnut) 299 Castanopsis cuspidata 202 Casuarina 212 Casuarina equisitfolia (Australian pine) 212 Casuarina stricta 212

Caucarium 214 Ceanothus 130, 131, 146 Ceanothus velutinus Cecidomyiidae (gall midges) 111, 265–271, 312 Cedrela 37, 290, 291 Cedrela mexicana 291, 292 Cedrela odorata (Spanish cedar) 169, 291, 292 Cedrus 122, 133, 195, 231, 303, 304, 311 Cedrus brevifolia 304 Cedrus deodara 275, 310 Cedrus libani (cedar of Lebanon) 152 Celtis (hackberry) 214, 217, 227, 254 Celtis occidentaltis (eastern hackberry) 254 Cerambycidae (round headed wood borers 105, 106, 210, 258, 326 Cerospatus volupis 155–156 chalcoprax 59 chapparal 8 chemical 74, 76, 87 ecology 17–19 allecochemicals 18 allomones 17, 18 kairomones 17, 18, 84 pheromones 17–18, 84, 85 semiochemicals, 17, 19, 58, 84 synomomes 17, 19 insecticide 73, 76, 89, 92 botanicals 88 carbamates 88 chlorinated hydrocarbons 88 horticultural oils 89 inorganic 87 insect growth regulators 89 insecticidal soaps 89 organophosphates 88 resistance 89 synthetic pyrethroids 88 systemic insecticides 88 pesticide 74, 75 registration 89–90 Chamaecyparis 185, 231, 237, 296 Chamaecyparis lawsonina (Port Orford cedar), 296 Chilecomandia moorei (butterworms) 109 Choristoneura 118–123 Choristoneura biennis (two-year cycle budworm) 119 Choristoneura carana californicum 119

357

Choristoneura carana carana 119 Choristoneura conflicata (large aspen tortrix) 119 Choristoneura diversana 119, 120 Choristoneura fumferana (spruce budworm) 16, 25, 34, 60, 82, 118, 119 Choristoneura lambertiana lambertiana 119 Choristoneura lambertiana ponderosana 119 Choristoneura lambertiana subretiniana 119 Choristoneura lambertiana (uncertain status) 119 Choristoneura murinana (fir budworm) 119, 122 Choristoneura occidentalis (western spruce budworm) 11, 24, 26, 34, 60, 64, 82, 93, 118, 119, 122–123, 293 Choristoneura pinus pinus (jack pine budworm) 119 Choristoneura orea 119 Choristoneura retiniana 119 Cinnamomum camphora (camphor tree) Chloroxylon swietenia (Ceylon satinwood) 249, 292 Chyrptolaemus montrouzieri 245 Chrysomelidae 105, 152 Chrysocharis laricinellae 117 Chrysomima semilteria 38 Chrysontomyia ruforum 19 Chrysophtharta 152 Chrysophtharta agricola 152 Chrysophtharta bimaculata (Tasmanian eucalyptus leaf beetle) 152 Chrysophtharta varicollis 152 Chukrasia tabularis 292 Cinnamomium camphora 214 Cinara (giant conifer aphids) 236 Cinara atlantica 237 Cinara cronartii 237 Cinara cupressi 39, 237 Cinara cupressivora (cypress aphid) 39, 63, 74, 78, 80, 236–238 Cinara maritimae 39, 237 Cinara pini 237 Cinara pinivora 237 Cinara ponderosae 237 Cinara pseudotsugae 237 Cinara thujafilina 237 Cinara strobi 237 Citrus 169, 200, 212, 249

358

Subject and Taxonomic Index

Clania variegata (giant bagworm) 114 climate 19, 22, 24 climate change 27–30 Coccoloba (seagrape) 158 Coccoloba uvifera 44, 158 Coccoloba venosa 158 Coffea (coffee) 214, 197–198 Coffea arabica 214 Coleophora laricella (larch casebearer) 56, 109, 117 Coleophoridae (casebearers) 108, 117 Coleoptera 98, 99, 104, 203, 253, 258, 273, 295, 298, 313, 323 Coleotechnites 117–118 Coleotechnites apictriopunctella 118 Coloetechnites milleri (lodgepole needle miner) 117–118 Coloetechnites ponderosae 118 Coleotechnites piceaella 118 Coleotechnites starki 118 Coleotechnites thujiella 118 Colophospermium mopane (mopane) 42, 136 Coloradia 135–136 Coloradia doris (Black Hills pandora moth) 136 Coloradia pandora (pandora moth) 42, 135–136 Coloradia velda 136 Conophthorus 174, 301–303 Conophthorus apachecae 301 Conophthorus banksianae (jack pine tip weevil) 301 Conophthorus conicolens 301 Conophthorus coniperda (white pine cone beetle) 301, 302 Conophthorus echinatae 301 Conophthorus edulis 301 Conophthorus mexicanus 301 Conophthorus michoacanae 301 Conophthorus monophyllae 301 Conophthorus ponderosae 301–303 Conophthorus radiatae 33, 301 Conophthorus resinosae 301 Conophthorus teocotum 301 Contarinia 312 Contarinia oregoniensis 312 Contarinia washingtoniensis 312 Coptotermes 317–318 Coptotermes acinaciformis 317 Coptotermes formosanus 36, 317–318 Coptotermes gestroi 317 Coptotermes lacteus 317

Coptotermes sjostedi 317 Coptotermes testaceus 317 Coreidae (leaf-footed bugs) 102, 296–297 Cornus florida (flowering dogwood) 61 Corylus 130, 135, 141, 143, 214, 298 Corylus californica 298 Corymbia calophylla 218 Cossidae (carpenter or goat moths) 109, 224–226 Cossus cossus (goat moth) 224 Cotoneaster 141 Couma macrocarpa (leche caspi) 197–198 Crateagus 130, 141, 143, 153, 220 crimson 248 Cronartium 237 Cronartium fusiforme 237, 311 Cronartium strobilinum 311 Cryptocarya alba (peumo) 223 Crytpococcus fagisuga (beech scale) 36, 246 Cryphalus piceae 122 Cryphonectria parasitica 205 Cryptocarya alba 137 Cryptomeria japonica 202, 212 Cyrptolaemus montrouzieri 245 Cryptotermes 315–316 Cryptotermes brevis (West Indian drywood termite) 315–316 Cryptotermes dudleyi 316 Cryptotermes havilandi 316 Ctenarytaina 235 Ctenarytaina eucalypti (blue gum psyllid) 40, 235 Ctenarytainia spatulata 235 Ctenomorphodes tessulata 151 Cunninghamia lanceolata 37 Cupressaceae 185, 236, 237 Cupressus 38, 185, 237, 296, 303, 304, 305 Cupressus abramsiana (Santa Cruz cypress) 305 Cupressus arizonica (Arizona cypress) 305 Cupressus bakeri 305 Cupressus goveniana 305 Cupressus lusitanica 39, 78, 80, 125, 127, 169, 237, 238 Cupressus macrocarpa (Monterrey cypress) 237 Cupressus sempervirens (Mediterranean cypress) 41, 185, 237, 296, 304, 305, 309 Cupressus torulosa 304

Curculio 298–299 Curculio caryae (pecan weevil) 298 Curculio caryatrypes (large chestnut weevil) 298 Curculio davidi 298 Curculio elephas (chestnut weevil) 298 Curculio fulvus 298 Curculio glandium 298 Curculio neocorylus (hazelnut weevil) 298 Curculio nucum 298 Curculio occidentalis (filbert weevil) 298 Curculio proboscidens 298 Curculio sayi (small chestnut weevil) 298 Curculio sikkimensis (chestnut curculio) 298 Curculio sulcatulus 298 Curculionidae 99, 105, 154, 173–175, 223, 274–287, 298–303 Curinus coeruleus 235 Cydia 305–308 Cydia anaranjada (slash pine seedworm) 305, 306, 308 Cydia caryana (hickory shuckworm) 306 Cydia ethelinda 306 Cydia fagilandana 306 Cydia illutana dahuricola (Dahurian larch seed moth) 305–307 Cydia illutiana illutiana 306 Cydia ingens (longleaf pine seedworm) 306, 308 Cydia latisigna 306 Cydia montezuma 306 Cydia piperana (ponderosa pine seed worm) 306, 308 Cydia pomonella 89, 295, 305 Cydia splendana (acorn moth) 306, 307 Cydia strobilella (spruce seed moth) 306 Cydia succediana 305 Cydia toreuta (shortleaf pine coneworm) 306, 307, 308 Cydonia 143 Cynipidae (gall wasps) 108, 262 Cynips 263 Cynips divisa 264 Cynips gallae tinctora (Allepo gall) 43, 263

Subject and Taxonomic Index Cynips insana 263 Cynips longiventris 264 Cynips quercusfolii (oak bud gall wasp, cherry gall) 264 Cynometra 214 D Daedalia unicolor 227 Danaus plexippus (monarch butterfly) 44 dauerlarvae 213 Deladenus siricola 58, 84, 230 Dalbergia sissoo (Indian rosewood) 249, 250, 273 Dasineura (larch gall midges) 265–266 Dasineura kellneri 265, 266 Dasineura rozhkovi 24, 265, 266 Dasineura nipponica 266 Dasineura verae 266 Dasychira grisefacta (western pine tussock moth) 139–140 dauerlarvae 213 Dead Sea fruit 264 declines and diebacks 26–27 Deladenus siricola 230 Dendranthema cineariaefolium 88 Dendroctonus 34, 86, 173, 175–185 Dendroctonus adjunctus (roundheaded pine beetle) 177 Dendroctonus approximatus (larger Mexican pine beetle) 177 Dendroctonus armandi 176, 177 Dendroctonus brevicomis 20, 175–178, 179 Dendroctonus frontalis (southern pine beetle) 19, 20, 25, 29, 34, 61, 70, 77, 79, 177–179, 180, 192, 200 Dendroctonus jeffreyi (Jeffrey pine beetle) 177 Dendroctonus mexicanus (smaller Mexican pine beetle) 177 Dendroctonus micans 176, 177, 179–181 Dendroctonus murrayanae 177 Dendroctonus rhizophagus 177 Dendroctonus rufipennis (spruce beetle) 20, 29, 34, 177, 183–184 Dendroctonus simplex (larch beetle) 177 Dendroctonus parallelocollis 177

Dendroctonus ponderosae (mountain pine beetle) 11, 12, 18, 19, 20, 22, 29, 34, 69, 78, 86, 174, 177, 181–182 Dendroctonus pseudotsugae (Douglas-fir beetle) 10, 20, 34, 86, 177, 182–183 Dendroctonus punctatus 177 Dendroctonus terebrans (black turpentine beetle) 177, 185, 192 Dendroctonus rhizophagus 177, 185 Dendroctonus rufipennis 86, 177, 183–184 Dendroctopnus simplex 177 Dendroctonus valens (red turpentine beetle) 36, 177, 184–185 Dendroctonus vitei 177 Dendrolimus 127, 128 Dendrolimus houi (Yunnan pine caterpillar) 128 Dendrolimus kikuchii 128 Dendrolimus sibiricus (Siberian silk moth) 128–130, 193 Dendrolimus pini 33, 37, 127–128 Dendrolimus punctatus (pine caterpillar) 37, 50, 60, 61, 82, 83, 85, 128–129 Dendrolimus sibiricus (Siberian silk moth) 33, 82, 128–130 Dendrolimus spectabilis 33, 128 Dendrolimus tabulaeformis 128 Dendrosoter 192 Dendrosoter caenopachoides 195 Dendrosoter sulcatus 192 density dependent 19 density independent 19, 22 Diapheromera femorata 149 Diaspididae (armored scales) 104, 249 Dichocrocis punctiferalis (yellow peach borer) 309–310 Didymuria violescens (spurlegged phasmid) 150–151 Diprion pini 19 Diprionidae (conifer sawflies) 107, 158–165 Dirohabda 43, 105, 152 Diorhabda carinata 153 Diorhabda carinulata 153 Diorhabda elongata 153 Diorhabda meridionalis 153 Dirohabda sublineata 153 Diospyros 6, 148 Diospyros melanoxylon (east Indian ebony) 224

359

Dioryctria 62, 292, 309–312 Dioryctria abietella 310, 311 Dioryctria abietivorella 310, 311 Dioryctria amatella 310, 311 Dioryctria assamensis 293 Dioryctria aurantcella 311 Dioyrctria castanea 311 Dioryctria claioralis 311 Dioryctria disclusa 311 Dioryctria erythropasa 311 Dioryctria horneana 293 Dioryctria merkeli 311 Dioryctria mutatella 311 Dioryctria pinicolella 311 Dioryctria pseudotsugella 311 Dioryctria pygmaella 311 Dioryctria raoi 293 Dioryctria reniculloides 311 Dioryctria rossi 311 Dioryctria rubella (Tusam pitch moth) 293 Dioryctria pinicolella 311 Dioryctria sylvestrella (maritime pine borer) 292–293 Dioryctria zimmermani (Zimmerman pine moth) 293 Diptera (flies) 110, 253, 265, 295, 312 Dipterocarpaceae 3 Disholcaspis 264 Disholcaspis cinerosa 264 Disholcaspis quercusmamma (rough oak bullet gall wasp) 264 Dryocoetes confusus (western balsam bark beetle) 24 Dryocosmus kuriphilus (chestnut gall wasp) 264–265 E Early Warning System 60 Echites umbellate 139 Elatobium 238 Elatobium abietinum (green spruce aphid) 238 Elaeagnus 187 Endoxyla leucomolcha 109 Ennomos subsignarius (elm spanworm) 124 Entomophaga maimaiga 83 Entadrophragma 292 Entadrophragma angolense 292 Entadrophragma utile 292 Epirrita autumnata (autumnal moth) 125 Epistomentis pictus 208–209

360

Subject and Taxonomic Index

Erannis 125 Erannis defoliara (mottled umber moth) 125 Erannis jacobsonii (Jacobson’s spanworm) 125 Erannis tiliara (linden looper) 124, 125 Ericoccidae (felt scales) 102, 246 Eriokermes 103 Ernobius punctulatus 33 Erythrina 260 Eschweilera corrugata 197–198 Eucalyptus 8, 36, 38, 107, 147, 154, 155, 197, 200, 201, 218, 238, 261, 273, 316, 317, 320, 323 Eucalyptus amygdalina 316 Eucalyptus baxteri 316 Eucalyptus blakelyi 235 Eucalyptus botryoides 261 Eucalyptus brassiana 235 Eucalyptus bridegsiana 235, 261 Eucalyptus camaldulensis (red gum) 154, 168, 235, 261 Eucalyptus citrodora 147 Eucalyptus cloeziana 147 Eucalyptus dealbata 235 Eucalyptus deanei 261 Eucalyptus delegatensis (alpine ash) 150, 152 Eucalyptus diversicolor 218, 235 Eucalyptus ficifolia 218 Eucalyptus globulus 78, 235, 261, 262, 316 Eucalyptus gomphocephala 218 Eucalyptus grandis 147, 218, 261 Eucalyptus gunnii 261 Eucalyptus lehmannii 235 Eucalyptus maculata 218 Eucalyptus maidenii 154 Eucalyptus mannifera 235 Eucalyptus nesophila 147 Eucalyptus nicholii 235 Eucalyptus nitens 152, 235 Eucalyptus obliqua 316 Eucalyptus paniculata 218 Eucalyptus pauciflora 316 Eucalyptus propinqua 218 Eucalyptus pulveriana 235 Eucalyptus punctata 154 Eucalyptus redunca 218 Eucalyptus regnans 150, 152, 316 Eucalyptus resinifera 218 Eucalyptus robusta (swamp mahogany) 154, 261 Eucalyptus rudis 235

Eucalyptus saligna 218, 261 Eucalyptus sideroxylon (mugga, red ironbark) 235 Eucalyptus sieberti 316 Eucalyptus smithii (gully gum) 154 Eucalyptus tereticornis 235, 261 Eucalyptus tenuiramis 316 Eucalyptus triantha 218 Eucalyptus viminalis 154, 155, 261, 316 Eucalyptus urophylla 147 Eucosma 287–288, 308–309 Eucosma bobana 308 Eucosma cocana (eastern pine coneworm) 308 Eucosma gloriola 288 Eucosma impropria 308 Eucosma monitorana 308 Eucosma ponderosa 308 Eucosma rescissoriana 308 Eucosma sonomana (western pine shoot borer) 87, 287–288 Eucosma tocullionana 308 Eucryphia lucida (leatherwood) 201 Eulophidae 108, 260 Euproctis 140–141 Euproctis chrsorrhoea (browntail moth) 140 Euproctis kargalika (Turkistan browntail moth) 33, 140–141 Eupcroctis similis (yellowtail moth) 140 Euura 259 Euura mucronata 259 Euura shibayangii 259 Evita hylaninaria blandaria 124, 125 F Fabaceae 245 Fagaceae 200, 202 Fagus 6, 120, 123, 125, 135, 153, 224, 227, 246, 251 Fagus grandifolia (American beech) 246 Fagus sylvatica (European beech) 6, 200, 204, 206, 246, 306 feedback 21, 22 negative 21 delayed negative 21 positive 22 rapid negative 21 Fenusa 165 Fenusa dohrni 165 Fenusa pusilla (birch leaf miner) 165 Fenusa ulmi 165

Ficus 212, 214, 249 Ficus elastica 249 foliage feeding insects 113–171 forest 1, 2 boreal or tiaga 8–9, 10 dynamics 9, ecosystems 2 health 73, 74, 95 plantations 34, 295 primary 1 temperate 5–6, 10 Mediterranean forests 8, temperate broadleaf and mixed forests 2, 6 temperate conifer forests 2, 7 tropical/subtropical 2–3, 9–10 mangroves 4 tropical and subtropical conifer forests 2, 4 tropical and subtropical dry broadleaf forests 2, 3 tropical and subtropical moist broadleaf forests 2, 3 urban 40–42 forest entomology as a carrier 44–47 applied forest entomology 46 government agencies 46 private industry 46–47 educational requirements 44–45 international opportunities 47 research 45–46 forest research institutes 45 universities 45 forest insect management 73–95 monitoring 49–72 objectives 49–50 surveillance and reporting 50–51 sampling sample frequency 54 sample location 54 sample size 53–54 sample unit 51–53 Formicidae (ants) 108, 167, 326 Fortunella margarita (kumquat) 212 Fraxinus (ash) 6, 42, 120, 130, 141, 148, 200, 204, 206, 217, 224, 225, 226, 231, 251, 297, 323, 324 Fraxinus excelsior (European ash) 155 Fraxinus pennsylvanica (green ash) 225 frontalin 183

Subject and Taxonomic Index fundatrix 257 fungi 83–84 Fusarium subglutinans f. spp. pini 33 fynbosch 8 G gall insects 253–272 gallicolae 257 Gelechiidae (leaf miners) 109, 117 geographic information systems (GIS) 64, 66 Geometridae (inchworms) 109, 124–127 Geosmithia 28, 284 Geosmithia morbida 284 Gemmura 259 Gilpinia 37, 158–159 Gilpinia abieticola 159 Gilpinia fennica 159 Gilpinia fructetorum 159 Gilpinia hercyniae (European spruce sawfly) 36, 38, 84, 158–159 Gilpinia leksawasdii 159 Gilpinia marshalli 159 Gilpinia pallida 159 Gilpinia pindrowi 159 Gilpinia polytoma 159 Gilpinia ventralis 159 Gilpinia virens 159 Gleditsia tricanthos (honey locust) 153, 217 Glena bisulcata 38, 125–127 global positioning systems (GPS) 66 Glycaspis brimblecombei (red gum lerp) 40, 235 Gmelina arborea 169, 225 Gnathotrupes 195–196 Gnathotrupes barbifer 196 Gnathotrupes nanus 196 Gnathotrupes vafer 196 Gnathotrupes velatus 196 Gonipterus 154 Gonipterus gibberus 154 Gonipterus scutellatus (eucalyptus weevil) 40, 154 Gossyparia spuria (European elm scale) 246 Gracillaridae (leaf blotch miners) 108, 114 Guarda macrocarpa 9 gouting 240 Grewia tiliaefolia 224 Grypocentrus aloipes 165 Gypchek 83

H Hedypathes betulinus 274–275 Helianthocampa 110 Helcostizus rufiscutum 218 Hemiptera (aphids, scales, bugs) 101, 233, 253, 295 Heteronemiidae 100, 149 Heterobasidium annosum 24 Heteropsylla cubana (leucaena psyllid) 40, 79, 233–235 Hevea brasiliensis (rubber tree) 198 Hexomyza schineri (poplar twig gall fly) 271–272 Hibiscus 212, 245 Hibiscus elatus 245 Himatanthus sucuuba 139 holocyclic 102, 257 Holopterus chilensis 213 honey 42 Hylastes ater (black pine beetle) 280–281, 282 Hylamorpha elegans 151 Hylobius 61, 274–277, 281, 282 Hylobius abietis (large pine weevil) 275–277 Hylobius albosparsus (white spotted weevil) 275 Hylobius angustus 275 Hylobius pales (pales weevil) 37, 275, 277 Hylobius piceus 275 Hylobius radicis (pine root collar weevil) 275 Hylobius rhizophagus (pine root tip weevil) 275 Hylobius transversovittatus 274 Hyloptrupes bajulus (old-house borer) 326 Hylurgus ligniperda (golden haired bark beetle) 39, 281–282 Hymenoptera (bees, wasps, ants) 107, 155, 203, 227, 253, 259, 295, 313, 326 Hyphantrea cunea (fall webworm) 36, 148 Hypsipyla 290–292 Hypsipyla grandella 24, 37, 291–292 Hypsipyla robusta 24, 37, 291, 292 I Ibalia 227, 230 Ibalia drewseni 230 Ibalia leucospoides 227, 230 Ibalia ruficollis 230

361

Ilex 202, 274 Ilex chinensis 202 Ilex paraguarensis 274 Imbrasia 42, 136, 137 Imbrasia belina (mopane worm, mopane emperor moth) 42, 136 Imbrasia nicitans 137 Inostemma 271 Inostemma matsutama 271 Inostemma seoulis 271 insects indigenous 33 introduced 33, 34 integrated pest management (IPM) 73–76, 87, 94, 95 action process 74, 75–76 decision process 74–75 strategies prevention 76 suppression 76 tactics 76 application technology 76, 90–94 meteorology 94 modeling behavior of aerial applications 94 spray systems 92 fixed wing aircraft 91 helicopters 91 calibration and characterization 92 ground application equipment 90 biological control 76, 79–84 augmentative 81 classic 80 microbials 80 bacteria 80 fungi 83 nematodes 84 viruses 80 chemical insecticides 76, 87–90 botanicals 88 carbamates 88 chlorinated hydrocarbons 88 horticultural oils 89 inorganic 87–88 insecticidal soaps 89 insecticide resistance 89 insect growth regulators 89 organophosphates 88 synthetic pyrethroids 88 systemic insecticides 88–89 cultural 76, 78 genetic 76, 79

362

Subject and Taxonomic Index

tactics (Continued ) mechanical 76, 79 regulatory 76 semiochemicals 76, 84–87 anti-aggregation 85–87 mass trapping 84–85 mating disruption 87 systems 94 Ips 32, 34, 98, 173, 190–195 Ips acuminatus 20, 34, 190, 191 Ips avulsus 34, 190, 192 Ips calligraphus (six-spined engraver beetle) 20, 190, 191–192 Ips cembrae 190, 193 Ips confusus 34, 190, 243 Ips grandicollis (southern pine engraver) 39, 190, 192 Ips hauseri (mountain Kyrgyz engraver) 190 Ips hunteri (blue spruce ips) 190 Ips latidens 19 Ips lecontei 190 Ips pini (pine engraver) 19, 20, 34, 190, 192–193 Ips sexdentatus 190, 192 Ips subelongatus 190, 193 Ips typographus (larger European spruce beetle) 10, 20, 23, 34, 59, 67, 84, 85, 99, 100, 190, 193–194 Ips typographus japonicus 193 ipslure 59 Isoptera (termites) 100, 313–328 J Jarra 218 Jarra maculipennis 218 Jarra phoracantha 218 Juglans (walnut) 6, 28, 143, 148, 214, 217, 226, 227, 284, 307 Juglans californica 284 Juglans major 284 Juglans mandshuria (Manshurian walnut) 206 Juglans nigra (black walnut) 28, 284 Juglans regia (English walnut, Persian walnut) 212 Juniperus (juniper) 7, 122, 144, 185, 196, 237, 296, 303 Juniperus communis (common juniper) 303, 304 Juniperus occidentalis (western juniper) 114, 267 Juniperus osteosperma (Utah juniper) 267, 303

Juniperus pingii 304 Juniperus polycarpos var. seravscanica 303 Juniperus procera (East African pencil cedar) 4, 237, 303, 304 Juniperus sabina 304 Juniperus scopulorum (Rocky Mountain juniper) 267 Juniperus semiglobosa 304 Juniperus thurifera 304 Juniperus viginiana (eastern red cedar) 267 K Kalotermitidae 101, 315–316 karmina 249 karroo 8 kermesic acid 249 Kermesidae (gall scales) 103, 248 Kermococcus 103, 248 Kermococcus vermilis 248 Kerria lacca (lac insect) 103, 248–249, 250, 328 Kerriidae (lac scales) 103, 248–249 kimiz 249 kirmina 249 Khaya (African mahogany) 37, 290, 292 Khaya anthotheca 292 Khaya grandiflora 292 Khaya ivorensis 292 Khaya nyasica 292 Khaya senegalensis 291, 292 Kerria lacca (lac insect) 43, 248, 328 L Lambdina 125 Lambdina fiscellaria fiscellaria (hemlock looper) 125 Lambdina fiscellaria lugubrosa (western hemlock looper) 125 large cambium and wood boring insects 203–231 Laricobius 240 Laricobius erichsonii 241 Laricobius nigrinus 240 Larix (larch) 8, 102, 109, 117, 128, 129, 144, 146, 180, 190, 192, 193, 196, 209, 213, 214, 215, 216, 221, 222, 223, 228, 230, 231, 256, 265, 275, 277, 280, 303, 304, 306, 308, 310, 311 Larix czekanowsii 266

Larix decidua (European larch) 25, 117, 123, 125, 191, 210, 230, 256, 266, 285 Larix gmelinii 125, 186, 193, 209, 266, 306, 308 Larix kaempferi 256, 266 Larix kamtschatica 186 Larix laricina (eastern larch) 117, 177, 209, 256, 311 Larix leptolepis (Japanese larch) 143, 193 Larix lyalli (subalpine larch) 186, 256 Larix occidentalis (western larch) 117, 122, 124, 146, 186, 222 Larix sibirica (Siberian larch) 125, 143, 186, 190, 193, 209, 266, 306, 308 Larix sukaczerii 186 Lathrolestes nigricollis 165 Lasiocampidae (lappet moths, tent caterpillars) 109, 127–132 Laurus nobilis (sweet bay, laurel) 200 Lespedeza 120 Lepidoptera (moths, skippers, butterflies) 108, 113–148, 203, 224–227, 253, 273, 287, 295, 305 Lepidosaphes ulmi (oystershell scale) 250–251 Leptocybe invasa (blue gum chalcid) 40, 261 Leptoglossus 102, 296 Leptoglossus corculus (southern pine seed bug) 297 Leptoglossus occidentalis (western conifer seed bug) 296 Leptographium 173, 274, 280 Leptographium procerum 280 Leptographium sibiricum 216 Leptographium truncatum 280 Leptographium yunnanesis 286 Leucaena 234 Leucana leucocephala 39, 79, 234 Leucoma salicis (satin moth) 141–142 Leucopis 241, 242 Leucopis obscura 241 Lindera 198 Lindera erythrocarpa 202 Lindera latifolia 198 Liriodendron 6 Liquidambar 6, 148 Liquidambar styraciflua 7

Subject and Taxonomic Index Lindgren funnel trap 59, 85 Litchi sinensis 212 Lithocarpus 198, 298 Lithocarpus densiflorus 298 Lithocarpus edulis 198 Lithrea caustica (litre) 137 Litsea elongata 198 Llamandra cathartica 139 Lonchocarpus margaretensis 197–198 Lophyrotoma interrupta (cattle poisoning sawfly) 157 Lovoa trichiliodes 292 Lupinus 289 Lyctinae 104, 323 Lyctus 323–325 Lyctus brunneus (brown lyctus beetle) 323–324 Lyctus cavicollis (western lyctus beetle) 325 Lyctus planicollis (southern lyctus beetle) 324–325 Lygaeidae (seed bugs) 101, 295 Lymantria 142–145 Lymantria dispar (gypsy moth) 16, 21, 24, 26, 33, 36, 41, 58, 67, 82, 83, 84, 87, 99, 141–144 Lymantria dispar asiatica (Asian gypsy moth) 33, 142 Lymantria dispar dispar (European gypsy moth) 142–143 Lymantria dispar japonica 143 Lymantria fumida 84 Lymantria mathura 143 Lymantria monacha (nun moth) 17, 18, 33, 37, 54, 82, 84, 143–145 Lymantria obfuscata (Indian gypsy moth) 143 Lymantria umbrosa (Hokkiado gypsy moth) 143 Lymantriinae (tussock moths) 16, 110, 139–148 Lythrum 256, 275, 276 Lythrum calicaria (purple loosestrife) 275, 276 M Maesopsis eminnii (umbrella tree) 137 Maconellicoccus hirsutus (pink hibiscus mealybug) 244–245 Macrotermes 320–321 Macrotermes goliath 320 Macrotermes natalensis 320 Mad apple of Sodum 264

Magnolia 6, 200 Magnolia grandiflora (southern magnolia) 200 Malacosoma 79, 130, 131 Malacosoma americanum (eastern tent caterpillar) 131 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma constrictum (Pacific tent caterpillar) 131 Malaocosoma disstria (forest tent caterpillar) 16, 26, 82, 131–132 Malacosoma incurvum (southwestern tent caterpillar) 131 Malacosoma indica (Indian tent caterpillar) 131 Malacosoma nuestria 131 Malacosoma tigris 131 Mallotus philippenis (kamala tree) 214 Malus (apple) 120, 124, 125, 130, 131, 141, 143, 145, 146, 153, 196, 200, 220, 224, 251 Malus pumila 187, 212 Malus spectablis 212 Malvaceae 245 Mangifera indica (mango) 169 maquis 8 mattaral 8 Matsucoccus 243–244 Matsucoccus acalyptus (piñon needle scale) 243–244 Matsucoccus feytaudi (maritime bast scale) 243–244 Matsucoccus josephi (Israeli pine bast scale) 244 Matsucoccus matsumurae 39, 244 Matsucoccus pini 244 mangroves 4–5 Margarodidae (giant coccids or bast scales) 102, 243 Mastotermes darwinensis 101, 315 Mastotermitidae 101, 315 Mecca gall 264 Mechoris cumulatus 299–300 Megacyllene 216–217 Megacyllene antennata 217 Megacyllene caryae (painted hickory borer) 217 Megacyllene mellyi 216 Megacyllene robinae 217 Megarhyssa 230 Megarhyssa nortoni 230 Megarhyssa percellus 230

363

Megastigmus 296, 303–305 Megastigmus albifrons (ponderosa pine seed chalcid) 304 Megastigmus atedius 304 Megastigmus atlanticus 304 Megastigmus bipunctatus 303, 304 Megastignus certus 303, 304 Megastigmus grandiosus (Montezuma pine seed chalcid) 304 Megastigmus milleri 304 Megastigmus pictus 304 Megastigmus pingii 303, 304 Megastigmus pinsapinis 304 Megastigmus pinus (fir seed chalcid) 304 Megastigmus schimitscheki 304 Megastigmus somaliensis 303, 304 Megastigmus spermatrophus (Douglas-fir seed chalcid) 303–305 Megastigmus wachtli 304, 305 Megaplatypus mutatus 199–200 Melanophila 208–210 Melanophila accuminata (black fire beetle) 208–209 Melanophila fulvoguttata (hemlock borer) 209, 239 Melanophila guttulata (larch buprestid) 209–210 Melia azedarach (Chinaberry) 210, 212, 249 Meliaceae 290 Melicoccus bijugatum 197–198 metamorphosis 98–99 complete 98, 99 simple 98, 233 Metatachardia 249 methylcyclohexanone (MCH) 183 Metrosideros polymorpha (ohia) 26, Microplasma-like organisms (MLOs) 233 Mikania 147 Milicia 254–255 Milicia excelsia 254 Milicia regia 254 Monochamus 32, 33, 213–216 Monochamus alternatus (Japanese pine sawyer) 60, 214, 215–216 Monochamus bimaculatus 214 Monochamus caroliniensis 214 Monochamus clamator 214 Monochamus galloprovincialis 214, 215 Monochamus grandis 214 Monochamus griseoplagiatus 214

364

Subject and Taxonomic Index

Monochamus (Continued ) Monochamus guttatus 214 Monochamus impluviatus 214 Monochamus marmorator (balsam fir sawyer) 214 Monochamus mutator 214 Monochamus notatus (northeastern pine sawyer) 214 Monochamus nitens 214 Monochamus obtusus 214 Monochamus saltuarius 214 Monochamus sartor 214 Monochamus scutellatus (white spotted sawyer) 214, 216 Monochamus scutellatus oregoniensis 214, 216 Monochamus scutellatus scutellatus 214, 216 Monochamus sutor 214 Monochamus titillator (southern pine sawyer) 214 Monochamus urossovi (fir sawyer) 214, 216 Moraceae 245 Moran effect 25, 26 Morpho spp. 44 Mordwilkoja vagabunda (poplar vagabond gall) 255–256 Morus 42, 143, 210 Morus alba (white mulberry) 212, 217 Myoplatypus flavicornis 200 Myricaria 153 N Nanokermes 103 Nasutitermes 320–321 Nasutitermes acajutlae 321 Nasutitermes costalis 320–321 Nasutitermes graveolus 320 Nasutitermes nigriceps 321 Nasutitermes walkeri 320–321 Nectria 246 Nectria coccinea var. faginata 246 Nectria ditissima 246 Nectria galligena 246 Nematus oligospilus 167 Neodiprion 160–164 Neodiprion abietis (balsam fir sawfly) 161 Neodiprion autumnalis 160–162 Neodiprion dailingensis 161 Neodiprion edulicolus (piñon sawfly) 161 Neodiprion fulviceps 161

Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion nanulus contortae 156, 161 Neodiprion nanulus nanulus (red pine sawfly) 161 Neodiprion pinetum 161 Neodiprion pratti pratti (Virginia pine sawfly) 161 Neodiprion sertifer (European pine sawfly) 38, 84, 161, 163–164 Neodiprion swainei (Swain’s jack pine sawfly) 57, 161 Neodiprion taedae linearis 160, 161 Neodiprion tsugae (hemlock sawfly) 161 Neodiprion xiangyunicus 161 Neomatus oligospilus 39 Nepytia 125 Nepytia freemani (false hemlock looper) 125 Nepytia janetae 125 Nesodiprion 164 Nesodiprion biremis 37, 164 Nesodiprion japonica 164 Noctuidae (owlet or miller moths) 16, 110, 139–148 Nothofagus (southern beech) 7, 8, 151, 195, 196, 208, 213 Nothofagus alpina (rauli) 156, 208 Nothofagus betuloides 196 Nothofagus dombeyi (coigüe) 137, 196, 208, 213, 223 Nothofagus cunnighamii (myrtle beech) 201 Nothofagus obliqua (roble) 137, 156, 208, 213 Nothofagus pumilio (lenga) 196, 208 Nuculaspis californica (black pine leaf scale) 251–252 Nyssa 131 O Odontata dorsalis (locust leaf miner) 153–154 Odontotermes 322 Odontotermes badius 322–323 Odontotermes latericus 323 Orgilus obscurator 80, 288 Oligotrophus 266–267 Oligotrophus apicis 266 Oligotrophus betheli (juniper felt tip midge) 266–267 Oligotrophus gemmarum 266

Oligotrophus juniperinus 266 Oligotrophus nezu 266 Oligotrophus panteli 266 Olla v-nigrum 80 Operophtera 125 Operophtera bruceata (bruce spanworm) 125 Operophtera brummata (winter moth) 125 Ophelimus 261 Ophelimus eucalypti (eucalyptus gall wasp) 40, 261–262 Olea europea (olive) 155 Ophiostoma 173 Ophiostoma minus 179 Ophiostoma novo ulmi 32, 186, 187 Ophiostoma ulmi 32, 186, 187 Oracella acuta (loblolly pine scale) 23, 245–246 Orgyia 145–147 Orgyia antiqua (rusty tussock moth) 145–146 Orgyia cana 146 Orgyia leucostigma 146 Orgyia mixta 38, 146 Orgyia pseudotsugata (Douglas-fir tussock moth) 11, 15, 21, 24, 25, 34, 82, 84, 146–147 Orgyia thyellina 146 Ormiscodes 137–138 Ormiscodes amphimone 137 Ormiscodes cinnamomea 38, 137–138 Orsillus 295–296, 305 Orsillus depressus (cypress seed bug) 295–296, 305 Orsillus maculatus (cypress seed bug) 295–296, 305 Orthotomicus 194 Orthotomicus erosus (Mediterranean pine engraver) 34, 194 Osmanthus fragrans 147 Ophiostoma 32, 173, 179, 186 Ophiostoma minus 179 Ophiostoma novo ulmi 32, 186 Ophiostoma ulmi 32, 186 Ougeinia dalbergioides 249 Opuntia 109 Oxydia trychiata 38 P Pachylobius picivorus 61 Pachypsylla celtidismamma (hackberry nipple gall) 253–254

Subject and Taxonomic Index Paleacrita vernata (spring cankerworm) 124 Panaxia quadripunctata 44 Panolis flammea (pine beauty moth) 37, 38 Papilio palamedes 199 Paropsis 152 Paropsis atomaria 152 Paraopsis charbdis 152 Pasania 202 Pasania edulis 202 Pasania glabra 202 Pauesia juniperinum 238 Pawlonia 114 Pawlonia tomentosa 114 Perga 156 Perga affinis 156 Perga affinis affinis 156 Perga affinins atrata 156 Pergagrapta bella 157 Pergidae 107, 155 Persea 199 Persea americana (redbay) 199 Persea borbonia 199 Peumos boldos (boldo) 137 Phaenops 210 Phaenops californica 210 Phaenops cyanea 210 Phaenops drummondi (flatheaded fir borer) 210 Phaenops gentilis (flatheaded pine borer) 210 Phasmatoidea (walkingsticks) 100, 149 Phigalea titea 124 Plumeria (frangipani) 139 Plumeria alba 139 Plumeria obtuse 139 Plumera rubra 139 Phllocnistis populiella (aspen leaf miner) 116 Phillyrea latifolia 155 Phloeomyzus passerinii (woolly poplar aphid) 238–239 Phloeosinus 185–186 Phloeosinus armatus 185 Phloeosinus bicolor 185–186 Phoebe lanceolata 198 Phoracantha 217–219 Phoracantha acanthocera (bulls-eye borer) 217–218 Phoracantha recurva (yellow phoracantha borer) 40, 219 Phoracantha semipunctata (eucalyptus longhorn borer) 40, 218–219

Phoradendron 204 Phyllaephagus 235 Phyllaephagus pilosus 235 Phyllaephagus vaseeni 235 Phyllocnistis populiella (aspen leaf miner) 116–117 Phylocladius aspleniifolius 201 Phyllocolpa 259 Phytoloma 254–255 Phytoloma excelsia 254 Phytoloma fusca 254 Phytoloma tuberculata 254 Phytophthora pinifolia 223 Physocarpus 130 Psyllaephagus 235, 236 Psyllaephagus bliteus 236 Psyllaephagus pilosus 235 Psyllaephagus vaseeni 235 Picea (spruce) 8, 102, 114, 128, 129, 144, 145, 146, 159, 161, 177, 180, 183, 190, 192, 193, 195, 196, 209, 213, 214, 215, 216, 220, 221, 222, 223, 228, 230, 231, 238, 256, 258, 275, 276, 277, 278, 280, 283, 285, 303, 304, 306, 310, 311, 325, 326 Picea abies (Norway spruce) 37, 144, 159, 180, 193, 210, 222, 230, 278, 285, 310 Picea ajanensis 285 Picea asperata (dragon spruce) 180 Picea brewerana (Brewer spruce) 180 Picea engelmanni (Engelmann spruce) 122, 125, 139, 180, 183, 222, 238, 278, 279, 287 Picea glauca (white spruce) 10, 120, 122, 139, 159, 177, 180, 183, 222, 278, 311 Picea jezoensis (Jezo spruce) 180, 193, 230, 239, 285 Picea likiangensis 258 Picea mariana (black spruce) 120, 180, 222, 258, 278 Picea obovata (Siberian spruce) 180, 191, 193, 285, 306 Picea omorika (Serbian spruce) 180 Picea orientalis (Caucasian spruce) 180, 191, 193, 230, 258 Picea polita 239 Picea pungens (blue spruce) 146, 180, 190, 222, 238, 278 Picea rubens (red spruce) 120, 177, 183, 222, 258, 278

365

Picea schrenkiana (Schrenk’s spruce) 190, 222, 223 Picea sitchensis (Sitka spruce) 36, 159, 177, 180, 222, 230, 238, 278 Picea smithiana 275, 285, 306, 310 Pimpla 128 Pinaceae 7, 236, 275, 303 Pineus 102, 241–242, 258–259 Pineus abietinus 241 Pineus armandicola 258 Pineus boerneri 36, 38, 79, 241–242 Pineus cembrae 258 Pineus cladogenous 241 Pineus coloradensis 241 Pineus floccus (red spruce adelgid) 258 Pineus ghanii 241 Pineus laevis 241 Pineus orientalis 258 Pineus pineoides 241 Pineus pini 241 Pineus pinifoliae (pine leaf chermes) 258 Pineus pinyunnanensis 241 Pineus simmondsi 241 Pineus strobi (pine bark adlegid) 241–243 Pineus wallichianae 241 Pinus (pine) 7, 8, 36, 38, 102, 114, 125, 128, 129, 133, 146, 161, 163, 164, 177, 180, 181, 184, 190, 191, 192, 193, 195, 196, 200, 207, 208, 209, 213, 214, 216, 221, 222, 223, 228, 230, 231, 237, 241, 242, 244, 258, 275, 276, 277, 278, 280, 281, 283, 285, 293, 296, 297, 301, 307, 309, 311, 317, 325, 326 Pinus albicaulis (whitebark pine) 122, 181 Pinus aristata (Colorado bristlecone pine) 267, 283, 302, 304 Pinus arizonica 161, 164, 175, 177, 302 Pinus armandi 128, 177, 184, 191, 195, 258, 285, 286 Pinus attenuata 278, 311 Pinus ayacahuite 164, 178, 283, 302 Pinus banksiana (jack pine) 57, 161, 177, 192, 229, 275, 278, 285, 288, 301, 306, 307 Pinus brutia 53, 133, 134, 152, 195, 244, 281, 285

366

Subject and Taxonomic Index

Pinus (pine) (Continued ) Pinus canariensis (Canary Island pine) 195, 281, 285 Pinus caribaea 191, 192, 195, 242, 293, 311 Pinus caribaea var. hondurensis 178 Pinus cembra (Swiss stone pine) 123, 161, 191, 300 Pinus cembroides (Mexican piñon) 252, 282, 283, 301, 308 Pinus contorta (lodgepole pine) 7, 12–13, 36, 78, 123, 161, 174, 177, 181, 228, 278, 283, 288, 300, 302, 308, 311 Pinus contorta ssp. contorta 38, 135, 287, Pinus contorta ssp. latifolia 122 Pinus cooperi 302 Pinus coulteri (Coulter pine) 135, 175, 177, 195 Pinus densata (Sikang pine) 286 Pinus densiflora (Japanese red pine) 128, 215, 244, 270 Pinus discolor (border piñon) 301 Pinus douglasiana 301, 302 Pinus durangensis 164, 175, 177, 178, 192, 302, 311 Pinus echinata (shortleaf pine) 161, 169, 178, 190, 191, 192, 195, 207, 245, 252, 277, 290, 301, 306, 307, 308, 310, 311 Pinus edulis (piñon pine) 7, 135, 161, 165, 190, 243, 244, 267, 282, 301, 308 Pinus eldarica 244 Pinus ellioti (slash pine) 37, 38, 128, 161, 190, 207, 229, 237, 242, 281, 282, 283, 306, 310, 311 Pinus engelmanni 161, 164, 177, 178, 301, 306 Pinus estevezii 175 Pinus flexilis (limber pine) 122, 181, 283, 302 Pinus gerardiana 310 Pinus greggi 283 Pinus griffithi 275, 310 Pinus halapensis (Aleppo pine) 133, 151, 242, 244, 281, 285, 288, 292, 293 Pinus hartwegii 302, 303, 304, 306 Pinus insularis 244, 285 Pinus jeffreyi (Jeffrey pine) 135, 177, 229, 252, 287, 302, 306, 308

Pinus kesiya 159, 164, 195, 242, 286, 293, 311 Pinus koraiensis 191, 192, 258, 285 Pinus lambertiana (sugar pine) 135, 181, 252, 302 Pinus lawsonii 311 Pinus leiophylla 164, 177, 178, 301, 311 Pinus luchuensis 244 Pinus lumholtzii 283 Pinus massoniana (Mason pine) 37, 128, 195, 215, 244, 245, 285, 309 Pinus maximinoi 178, 311 Pinus merkusii 4, 128, 164, 191, 293 Pinus michoacana 164, 191, 301, 306, 311 Pinus monophylla (single-leaf piñon) 136, 161, 165, 190, 243, 244, 282, 283, 301, 308 Pinus montezumae 164, 191, 192, 281, 283, 302, 303, 304, 306 Pinus monticola (western white pine) 122, 181, 214, 241, 242, 258, 296, 302, 308 Pinus mugo 191, 244, 285, 288, 310 Pinus mugo ssp. uncinata 195 Pinus muricata 278, 288 Pinus nigra (Austrian pine) 133, 191, 195, 230, 281, 285, 288, 292, 293 Pinus nigra ssp maritima 133, 280 Pinus nigra ssp pallasiana 152, 190, 281 Pinus occidentalis 191 Pinus oocarpa 164, 177, 178, 191, 192, 311 Pinus palustris (longleaf pine) 161, 190, 192, 207, 245, 306, 310, 311 Pinus parvifolia 285 Pinus patula 169, 195, 242, 281, 301 Pinus peuce (Macedonian pine) 241, 242 Pinus pinaster (maritime pine) 133, 195, 215, 229, 243, 244, 280, 281, 288, 292, 293 Pinus pinea (stone pine) 133, 151, 195, 281, 288, 292, 293, 300 Pinus pinceana 308

Pinus ponderosa 7, 11, 78, 122, 135, 139, 160, 161, 165, 175, 177, 178, 181, 191, 192, 230, 237, 251, 252, 280, 283, 287, 288, 296, 304, 306, 308, 311 Pinus pringeli 178, 302 Pinus pseudostrobus 164, 177, 191, 192, 302, 306 Pinus quadrifolia 283 Pinus radiata (Monterrey pine) 33, 36, 38, 78, 80, 133, 137, 146, 164, 195, 201, 208, 223, 229, 230 , 213, 241, 242, 278, 280, 281, 285, 288, 289, 290, 301, 316 Pinus roxburghii 293, 310 Pinus resinosa (red pine) 159, 161, 191, 192, 229, 242, 244, 278, 285, 288, 301, 306, 307, 308 Pinus rigida (pitch pine) 161, 178, 191, 192, 207, 252, 277, 290 Pinus rudis 302, 303, 304, 306 Pinus sabiniana (digger pine) 252, 283 Pinus sibirica (Siberian pine) 193, 214 Pinus serotina (pond pine) 178, 237 Pinus strobiformis 192, 283, 302 Pinus strobus (eastern white pine) 128, 161, 177, 191, 195, 209, 214, 229, 237, 241, 242, 258, 277, 278, 281, 285, 301, 302, 304, 308 Pinus sylvestris (Scotch pine) 19, 37, 38, 123, 125, 127, 128, 133, 144, 159, 161, 163, 190, 191, 192, 193, 195, 210, 222, 228, 237, 242, 244, 280, 281, 285, 288, 290, 300, 310, 311 Pinus sylvestris var. mongolica 191 Pinus tabulaeformis 128, 161, 184, 191, 195, 244, 285 Pinus taeda (loblolly pine) 37, 38, 78, 85, 128, 161, 169, 178, 190, 191, 192, 207, 229, 237, 242, 245, 277, 290, 306, 308, 310, 311 Pinus taiwanensis 244 Pinus tenuifolia 177, 192 Pinus teocote 161, 164, 178, 283, 301 Pinus thunbergii 128, 244, 270 Pinus virginiana 161, 178, 192, 245, 290, 306, 307, 308, 310, 311

Subject and Taxonomic Index Pinus wallichiana 310 Pinus washoensis 302 Pinus yunnanensis 161, 195, 278, 279, 285, 286 Pinyonia edulicola (piñon spindle gall midge) 266–267 Pissodes 277–280 Pissodes approximatus (northern pine weevil) 278 Pissodes castaneus 278 Pissodes hercyniae 278 Pissodes nemorensis (deodar weevil) 278 Pissodes notatus (banded pine weevil) 278 Pissodes piceae 278 Pissodes pini 278 Pissodes radiatae (Monterrey pine weevil) 278 Pissodes strobi (white pine weevil) 24, 37, 277–280 Pissodes terminalis (lodgepole terminal weevil) 278 Pissodes validirostris 277, 278, 300–301 Pissodes yunnanensis 278, 279–280 Pistacia 141, 303 Pithecellobium pinnatum 197–198 Pityokteines curvidens 122 Pityophthorus 282–284 Pityophthorus barberi 283 Pityophthorus blandus 283 Pityophthorus brevis 283 Pityophthorus boycei 283–284 Pityophthorus confertus 283 Pityophthorus deletus 283 Pityophthorus juglandis (walnut twig beetle) 28, 284 Pityophthorus keeni 283 Pityophthorus lecontei 283 Pityophthorus modicus 283 Pityophthorus punctifrons 283 Pityophthorus schwartzi 283 Pityophthorus tuberculatus 283 Pityophthorus woodi 283 Platanus 200, 212 Platanus occidentalis (American sycamore) 114 Platycladius orientalis 185 Platyponinae (ambrosia beetles, pinhole borers) 106, 173–174, 199–202 Platypus 200–202 Platypus cylindrus (oak pinhole borer) 200–201

Platypus granulosus 201–202 Platypus quercivorus 29, 202 Plodia interpunctella 89 Podacanthus wilkinsonnii 151 Podesesia syringae (lilac/ash borer) 62, 227 Pontania 259–260 Pontania proxima (willow red gall sawfly) 260 Population 15–24 abundance 15 cycles 25 density 15 dynamics 15 eruptions 25 processes 16 Populus (poplar, cottonwood) 8, 37, 114, 120, 123, 125, 130, 131, 140, 141, 143, 148, 167, 200, 204, 210, 212, 219, 220, 224, 225, 227, 238, 251, 256, 258, 271 Populus alba 227, 238 Populus caspica 200 Populus deltoides (eastern cottonwood) 219, 227, 256 Populus x euroamericana 238 Populus grandidentata (bigtooth aspen) 219 Populus nigra 227, 238, 256, 258 Populus tremula 204, 258 Populus tremuloides (quaking aspen) 116, 219, 256, 271, 272 Porotermes 316–317 Porotermes adamsoni 316–317 Porotermes planiceps 316 Porotermes quadricollis 316 Prionoxytus robinae (carpenterworm) 225 Prosopis 217 Protium 197–198 Prunus (cherry, plum) 120, 125, 130, 131, 143, 144, 149, 153, 202, 210, 212, 227, 251 Prunus armeniaca 187 Prunus padus 187 Prunus persica 187, 200 Prunus pseudocerasus 187 Prunus salicina 187 Prunus yedoensis 187 Pseudips 33 Pseudococcidae (mealybugs) 103, 244–246, 305 Pseudococcyx tessulatana 309

367

Pseudoperga lewisii (pale brown sawfly) 157 Pseudosphinx tetrio (frangipani hawk moth) 139 Pseudotsuga 102, 196, 214, 216, 231, 277 Pseudotsuga flahaulti 182 Pseudotsuga macrocarpa (bigcone Douglas-fir), 177, 182, 304 Pseudotsuga menziesi (Douglas-fir) 7, 11, 33, 34, 37, 122, 125, 139, 145, 146, 177, 182, 189, 195, 214, 222, 237, 252, 256, 257, 275, 276, 280, 285, 293, 296, 304, 310, 311, 312, 325, 326 Psidium 147 Psidium cattelianum (strawberry guava) 147 Psidium guajava (guava) 147 Psychidae (bagworms) 108, 113 Psyllaephagus pilosus 235 Psyllidae (jumping plant lice) 102, 233, 253 Pterocarya rhoifolia (Japanese wingnut) 206 Pullus impexus 241 Purshia 131 Purshia tridentata (bitterbrush) 130, 146 Pyralidae (snout moths) 109, 290–292, 309–312 Pyrrhalta luteola (elm leaf beetle) 154 Pyrus 141, 143, 200, 210, 212, 224, 251 Pyrus communis 200, 212 Q Quadrastichus erythrinae (erythrina gall wasp) 260–261 Quadraspidiotus perniciosus 89 Quercus (oak) 6, 7, 43, 103, 114, 120, 123, 124, 125, 130, 131, 135, 140, 141, 143, 144, 146, 149, 153, 200, 202, 204, 205, 214, 217, 224, 225, 226, 227, 248, 262, 263, 264, 298, 299, 306, 307, 323, 324 Quercus acuta 202 Quercus acutissima 202 Quercus aegilops 263 Quercus agrifolia 225, 263 Quercus alnifolia 204 Quercus bicolor 264 Quercus cerris 135, 206 Quercus chrysolepis 146, 205, 263

368

Subject and Taxonomic Index

Quercus (oak) (Continued ) Quercus coccifera 248 Quercus coccinea (scarlet oak) 262 Quercus dalechampii 186 Quercus dilatata 131 Quercus fusiforme 264 Quercus gambelii (gambel oak) 103 Quercus garryana (Garry oak) 263 Quercus gilva 202 Quercus glauca 202 Quercus ilex (Holm oak) 206, 248 Quercus incana 131 Quercus infectoria 263 Quercus kellogii (California black oak) 146, 205 Quercus macrocarpa (burr oak) 264 Quercus mongolica 29, 202 Quercus mongolica var. grosseserratus 202 Quercus myrsinifolia 202 Quercus pendiculata 263 Quercus petraea (sessile oak) 135, 186, 206, 264, 298 Quercus phillyraeoides (ubame oak) 202 Quercus pubescens 206 Quercus robur (English oak) 135, 186, 206, 264, 298, 299 Quercus rubra (northern red oak) 206, 248 Quercus salcina 202 Quercus serrata 202 Quercus sessilifolia 202 Quercus suber (cork oak) 206, 299 Quercus velutina (black oak) 248, 262 Quercus virginiana (live oak) 248, 264, 298 Quillaja saponora (soapberry) 223 R Raffaelea Raffaelea lauricola 199 Raffaelea quercivora 29, 202 Raffaelea santoro 200 remote sensing 62–68 aerial forest health surveys 62–66 aerial observer training 66 aerial photography 66–68 automated flight following 66 touch screen computers 66 Reticulitermes 318–320 Reticulitermes arenicola 319 Reticulitermes balkanensis 319

Reticulitermes banyulensis 319 Reticulitermes clypeatus 319 Reticulitermes flavipes (subterranean termite) 318–320 Reticulitermes grassei 319 Reticulitermes hageni 319 Reticulitermes hesperus 319 Reticulitermes lucifugus 319 Reticulitermes santonensis 319 Reticulitermes tibialis 319 Reticulitermes virginicus 319 Rhabdadenia biflora 139 Rhabdophaga 267, 269 Rhabdophaga rosaria 267, 269 Rhabdophaga strobiloides (willow pine cone gall) 267, 269 Rhamnus 120, 146 Rhamnus californica 146 Rhinotermitidae 101, 317–320 Rhododendron 120 Rhopalidae (scentless plant bugs) 102, 297–298 Rhyacionia 24, 288–290 Rhyacionia buoliana 38, 39, 80, 288–290 Rhyacionia buoliana buoliana 288, 289 Rhyacionia buoliana thurificana 288, 289 Rhyacionia bushnelli (western pine tip moth) 288 Rhyacionia frustrana (Nantucket pine tip moth) 37, 288, 290 Rhyacionia neomexicana (southwestern pine tip moth) 288 Rhyacionia rigidana (pitch pine tip moth) 288, 290 Rhyacionia subtropica (subtropical pine tip moth) 288, 290 Rhyacionia zozana 288 Rhizophagus grandis 181 Rhus triloba 130 Rhyephenes 223–224 Rhyephenes mallei 223–224 Rhyephenes humeralis 223–224 Rhyssa 230 Rhyssa hoferi 230 Rhyssa jozana 230 Rhyssa persuasoria 230 Ribies 130, 141 risk/hazard rating 68–72 Robinia (locust) 143 Robinia neomexicana (New Mexico locust) 217

Robinia pseudoacacia (black locust) 114, 153, 200, 210, 217, 227 Roptrocerus xylophagorum 192 Rosa 130, 141, 143, 146, 303 Roscaeae 8, 140, 303 rotzholz 240 Rubus 141 S Sabina chinensis 185 Sacoglottis procera 197–198 Salicaceae 8 Salix (willow) 8, 114, 120, 125, 130, 131, 140, 141, 144, 145, 146, 148, 167, 187, 200, 210, 212, 214, 220, 224, 227, 231, 238, 251, 258, 259, 267, 271 Salix alba 267 Salix babylonica (weeping willow) 187 Salix japonica 259 Santalum album (sandalwood) 249 Saperda 219–221 Saperda alberti 220 Saperda balsamifera 220 Saperda calcarata (poplar borer) 219–221 Saperda candida (roundheaded appletree borer) 220 Saperda carcharias (large poplar longhorn beetle) 220–221 Saperda concolor 220, 259 Saperda interrupta 220 Saperda inorata 220, 259 Saperda octomaculata 220 Saperda perforata 220 Saperda populnea (small poplar borer) 220, 258–259 Saperda scaiarias 220 Saperda similis 220 Saperda tridentata (elm borer) 220 Sarsina violascens 147 Sasajiscymnus tsugae 240 Sassafras albidum (sassafras) 199 Saturniidae (giant silk moths) 110, 135 Scarabaeidae (June or cockchafer beetles) 104, 151 Schinus 303 Schinus latifolius (molle) 137 Schleichera 249 Schleichera oleosa 249 Schleichera trijuga 249 Schlettererius cinctipes 230

Subject and Taxonomic Index Sclerophyllous 8 Scolytinae (bark beetles and ambrosia beetles) 99, 106, 173–199, 280–287, 301–303 Scolytus 173, 186–189 Scolytus intricatus 186 Scolytus laricis (western larch beetle) 186 Scolytus morawitzi 186 Scolytus mundus 186 Scolytus multistriatus (smaller European elm bark beetle) 20, 32, 41, 186–187 Scolytus ratzeburgi 186 Scolytus rugulosus (shothole borer) 186 Scolytus schevyrewi (banded elm bark beetle) 59, 186, 187–189 Scolytus scolytus 21, 32, 186 Scolytus unispinosus (Douglas-fir engraver) 186 Scolytus ventralis (fir engraver beetle) 34, 186, 189 Scymnus 240 Scymnus ningshanensis 240 Scymnus sinuanodulus 240 seed orchards 295 Seiridium cardinalis 185, 186, 237, 309 Seltrichodes globulus (blue gum gall wasp) 262 sequential sampling 56–58 Sequoia sempervirens 8 Sesiidae (clearwing moths) 109, 226–227 Sericocerus 158 Sericocerus krugii 158 Sericoceros mexicanus 44, 158 seudenol 183 sexupara 257 Shorea 249 Shorea robusta (sal) 198, 249 Shorea talura 249 silk 42 Sirex 227–230 Sirex noctilio (sirex woodwasp) 32, 38, 39, 58, 74, 78, 84, 85, 94, 228–230 Sirex juvencus 230 Sirex juvencus atricornis 230 Sirex juvencus californicus 230 Sirex juvencus juvencus 230 Siricidae (woodwasps) 107, 227–230

Solidago spp. (goldenrod) 217 Sorbus (mountain ash) 9, 203, 220 Sphingidae (sphink or hawk moths, hornworms) 110, 138 Steinernema carpocapsae 274 Stereonychus fraxini (ash weevil) 155 sucking insects 233–252 Swietenia 38, 249, 290, 292 Swietenia macrophylla (big leaf mahogany) 169, 197–198, 291, 292 Swietenia mahogany 249, 291, 292 synchronous cohort senescence 27 Syngaster lepidus 218–219 Syringa vulgaris (lilac) 155, 226, 251 Syzygium cuminii 249 T Tachardina 249 Tachordiella 249 Tachinidae 128 Tamarix (salt cedar) 43, 152 Tamarixia leucaenae 235 Tamarindus indica (tamarind) 249 Taxodiomyia cupressianassa (cypress twig gall midge) 267–271 Taxodium distichum (bald cypress) 114, 268, 310, 311 taxonomy 99–100 Tectona grandis (teak) 169, 224, 225, 245 Telenomus 80, 128, 296 Telenomos alsophilae 80 Temnochila virescens 192 Tenthredinidae 107, 165, 259 Terminalia 224, 273 Terminalia bellerica 224 Terminalia ivoriensis 273 termites 313–323 castes 314 invasives 315 pest management 314–315 Termitidae 101, 320–321, 323 Tetraclinis articulata 237 Tetropium 221–223 Tetropium abietis (roundheaded fir borer) 222 Tetropium castaeum 221–222 Tetropium cinnamopterum (eastern larch borer) 222 Tetropium fuscum (brown spruce longhorn beetle) 36, 222–223

369

Tetropium gabrielli (larch longhorn beetle) 222 Tetropium gracilicorne 222–223 Tetropium parvulum (northern spruce borer) 222 Tetropium staudingeri (Seven-river spruce beetle) 222–223 Tetropium velutinum (western larch borer) 222 Thanasimus dubius 19, 61, 192 Thaumetopea 16, 18, 110, 132–135 Thaumetopoea pityocampa (pine processionary caterpillar) 28, 32, 60, 79, 82, 84, 132 Thaumetopoea processionae (oak processionary caterpillar) 28, 132, 135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 17, 32, 53, 60, 79, 82, 132 Thaumetopoeidae (processionary caterpillars) 110, 132–135 Thecodiplosis 70, 270 Thecodiplosis brachyptera 270 Thecodiplosis japonensis (pine needle gall midge) 70, 270 Theobroma cacao (cacao) 197–198 Thermopisdae (dampwood termites) 101, 316–317 Thuja 185, 196, 237, 275, 277, 280, 309 Thuja occidentalis (northern white cedar) 114 Thuja orientalis 309 Thuja plicata 8 Thyridopteryx ephemaraeformis (bagworm) 113–114 thrips 253 Thysanoptera 253 Tilia (basswood, linden) 6, 114, 124, 125, 140, 143, 149, 200 Tip, shoot and regeneration insects 273–293 TM Biocontrol 1 82 Tomicus 284–287 Tomicus brevipilosus 285 Tomicus destruens 34, 284–286 Tomicus minor 285 Tomicus pilifer 285 Tomicus piniperda 21, 284–287 Tomicus puellus 285 Tomicus yunnanensis (Yunnan shoot borer) 285, 287

370

Subject and Taxonomic Index

Toona 290, 292 Toona autralis 292 Toona cilicata 292 Toona serrata 292 Tortricidae (leaf rollers, budworms) 109, 118–124, 287–290, 305–309 Tortrix viridana (green oak tortrix) 33, 123 Torymidae 108, 303–305 Torymus sinensis 265 Traumetocampa 110 tree reproductive structures 295–312 Tremex 227 Tremex columbia 227 Tremex fuscicornis 227 Trichoteras vaccinifoliae 263 Trichosporum symbioticum 189 Trochodendron 238 Trypodendron 196–197 Trypodendron lineatum (striped ambrosia beetle) 21, 85, 196–197 Tsuga (hemlock) 102, 129, 144, 196, 214, 239, 256, 275 Tsuga canadensis (eastern hemlock) 76, 125, 209, 239, 308 Tsuga caroliniensis (Carolina hemlock) 239 Tsuga chinensis 239 Tsuga diversifolia 239 Tsuga dumosa 239 Tsuga mertensiana (mountain hemlock) 122, Tsuga heterophylla (western hemlock) 7, 122, 125, 139, 145, 146, 161, 189, 222, 239, 311, 325

Tsuga mertensiana (mountain hemlock) Tsuga sieboldii 239 Turkey red 263 U Ulex europaeus 305 Ulmus (elm) 114, 120, 124, 125, 141, 146, 148 , 149, 153, 154, 186, 187, 200, 210, 214, 220, 224, 225, 226, 227, 246, 251, 323 Ulmus americana (American elm) 41, 166, 187 Ulmus carpinifolia 187 Ulmus davidana (David elm, Japanese elm) 187, 206 Ulmus laevis 187 Ulmus macrocarpa 187 Ulmus propinqua 187 Ulmus pumila 187 Ulmus thomasi (rock elm) 187 Urocerus 230–231 Urocerus gigas 230 Urocerus gigas flavicornis 230 Urocerus gigas gigas 230, 231 Urocerus gigas orientalis 230 Urocerus gigas tibetatunus 230 Urtica 123 V Vaccinium 123, 145, 146 viruses 81–83 baculoviruses 81 cytoplasmic viruses (CPV) 81 granulosis viruses (GV) 81 nucleopolyhedrosis viruses (NPV) 81, 82 Retroviridae 81

W Widdringtonia 237 wildfire 12 windstorms 10 windthrow 10, 11 wood in use 313–328 X Xyleborus 197–199 Xyleborus affinis 198, 199 Xyleborus ferrugineus 197–199 Xyleborus glabratus (redbay ambrosia beetle) 198–199 Xyleborus perforans 198 Xyleborus similis 198 Xyleutes ceramica (teak beehole borer) 225 Xylotrechus altaicus 193 Y yerba mate 274 Z Zadiprion 164–165 Zadiprion falsus 164–165 Zadiprion rohweri 165 Zadiprion townsendi 165 Zeiraphera 123–124 Zeiraphera diniana (gray larch bud moth 25, 26, 30, 84, 123–124 Zeriraphia improbana 124 Zeiraphera sp. 124 Zelkova 227 Zelkova serrata 187, Zeuzera pyrina (leopard moth) 226 Ziziphus 212, 249 Ziziphus mauritania 249 Ziziphus jujube 212

Host Index A Abies Foliage feeders Choristoneura fumiferana (spruce budworm) 119, 120–122 Choristoneura murinana (fir budworm) 119, 122 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Dendrolimus punctatus (pine caterpillar) 128–129 Lymantria monacha (nun moth) 144–145 Neodiprion abietis (balsam fir sawfly) 161 Orgyia antiqua (rusty tussock moth) 145–146 Bark and ambrosia beetles Dendroctonus micans 177, 180 Ips subelongatus 193 Ips typographus (larger European spruce beetle) 193 Orthotomicus erosus (Mediterranean pine engraver) 195 Scolytus ventralis (fir engraver beetle) 186, 189 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Melanophila acuminata (black fire beetle) 208 Melanophila guttulata (larch buprestid) 209 Monochamus alternatus (Japanese pine sawyer) 214, 215 Monochamus grandis 214 Monochamus nitens 214 Monochamus notatus (northeastern pine sawyer) 214 Monochamus obtusus 214

Monochamus saltuarius 214 Monochamus sartor 214 Monochamus scutellatus (white spotted sawyer) 214, 216 Phaenops drummondi (flatheaded fir borer) 210 Saperda interrupta 220 Sirex noctilio (sirex woodwasp) 228 Sirex juvencus 230 Tetropium abietis (roundheaded fir borer) 222 Tetropium castaeum 221–222 Tetropium cinnamopterum (eastern larch borer) 222 Tetropium gracilicorne 222–223 Tetropium velutinum (western larch borer) 222 Urocerus gigas 231 Sucking insects Adelges glandulae 256 Adelges nebrodensis 256 Adelges pectinata 256 Adelges piceae (balsam woolly adelgid) 240 Adelges prelli 256 Elatobium abietinum (green spruce aphid) 238 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Hylobius pales (pales weevil) 277 Tree reproductive structures Dioryctria abietella 310, 311 Dioryctria abietivorella (fir coneworm) 310, 311 Dioryctria reniculelloides (spruce coneworm) 311 Cydia illutiana illutiana 306 Megastigmus milleri 304 Megastigmus pinsapinis 304

Megastigmus pinus (fir seed chalcid) 304 Wood in use Buprestis aurulenta (golden buprestid) 326 Abies alba Foliage feeders Choristoneura murinana (fir budworm) 119, 122 Wood borers Phaenops cyanea 210 Tetropium fuscum (brown spruce longhorn beetle) 222 Sirex juvencus 230 Sucking insects Adelges piceae (balsam woolly adelgid) 240 Tree reproductive structures Dioryctria abietella 310, 311 Abies amabalis (Pacific silver fir) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Choristoneura orea 119 Sucking insects Adelges piceae (balsam woolly adelgid) 240 Pineus abietinus 241 Abies balsamea Foliage feeders Choristoneura fumiferana (spruce budworm) 119, 120–122 Wood borers Monochamus marmorator (balsam fir sawyer) 214 Sirex juvencus 230 Sucking insects Adelges piceae (balsam woolly adelgid) 240

Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

371

372

Host Index

Abies borisii-regis (Bulgarian fir) Wood borers Sirex juvencus 230 Abies cilicata Foliage feeders Choristoneura murinana (fir budworm) 119, 122 Wood borers Sirex juvencus 230 Abies concolor (white fir) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Choristoneura retiniana 119 Orgyia pseudotsugata (Douglas-fir tussock moth) 146–147 Bark and ambrosia beetles Scolytus ventralis (fir engraver beetle) 186, 189 Sucking insects Nuculaspis californica (black pine leaf scale) 252 Abies fraseri Wood borers Monochamus marmorator (balsam fir sawyer) 214 Sucking insects Adelges piceae (balsam woolly adelgid) 240 Abies grandis (grand fir) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Orgyia pseudotsugata (Douglas-fir tussock moth) 146–147 Bark and ambrosia beetles Scolytus ventralis (fir engraver beetle) 186, 189 Sucking insects Adelges piceae (balsam woolly adelgid) 240 Pineus abietinus 241 Abies holophylla (Manchurian fir) Tip, shoot and regeneration insects Tomicus puellus 285 Abies lasiocarpa (subalpine fir) Foliage feeders Choristoneura biennis (two-year cycle budworm) 119 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Nepytia janetae 125

Bark and ambrosia beetles Scolytus ventralis (fir engraver beetle) 186, 189 Wood borers Sirex juvencus 230 Sucking insects Adelges piceae (balsam woolly adelgid) 240 Pineus abietinus 241 Abies magnifica (red fir) Foliage feeders Coleotechnites milleri (lodgepole needle miner) 117–118 Abies nephrolepis Tip, shoot and regeneration insects Tomicus puellus 285 Abies nordmanniana (Nordmann fir) Bark and ambrosia beetles Ips acuminatus 191 Wood borers Sirex juvencus 230 Gall insects Adelges nordmanniana 256 Abies pindrow Foliage feeders Gilpinia pindrowi 159 Tree reproductive structures Dioryctria abietella 310, 311 Abies religiosa (oyamel, sacred fir) Foliage feeders Evita hyalinaria blandaria 124 Bark and ambrosia beetles Scolytus mundus 186, 189 Tree reproductive structures Dioryctria pinicolella 311 Abies sachalinensis (Sakhalin fir) Wood borers Sirex juvencus 230 Abies sibirica Wood borers Monochamus urossovi (fir sawyer) 216 Abies spectabilis Tree reproductive structures Dioryctria abietella 310, 311 Acacia Foliage feeders Clania variegata (giant bagworm) 114 Acacia mearnsii Foliage feeders Imbrasia nicitans 137 Acacia mollisima Wood borers Agrilus near grandis 207

Acacia nilotica Sucking insects Kerria lacca (lac insect) 248–249 Acacia senegal (gum Arabic) Wood borers Agrilus sp. Acer (maple) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Erannis tiliaria (linden looper) 124, 125 Euproctis chrysorrhoea (browntail moth) 140 Euproctis kargalika (Turkistan browntail moth) 140–141 Hyphantria cunea (fall webworm) 148 Leucoma salicis (satin moth) 141–142 Lymantria monacha (nun moth) 144–145 Malacosoma disstria (forest tent caterpillar) 131–132 Malacosoma nuestria 131–132 Operophtera bruceata (bruce spanworm) 125 Operophtera brumata (winter moth) 125 Orgyia antiqua (rusty tussock moth) 145–146 Thyridopteryx ephemeraeformis (bagworm) 113–114 Tortrix viridana (green oak tortrix) 123 Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Monochamus guttatus 214 Tremex fuscicornis 227 Sucking insects Lepidosaphes ulmi (oystershell scale) Tree reproductive structures Boisea rubrolineata (western boxelder bug) 297 Boisea triivittata (boxelder bug) 297 Acer negundo (boxelder) Wood borers Tremex fuscicornis 227 Tree reproductive structures Boisea rubrolineata (western boxelder bug) 297

Host Index Boisea triivittata (boxelder bug) 297 Acer orientalis Tip, shoot and regeneration insects Apate monachus 273 Adenium Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Aegle marmelos Sucking insects Aonidiella orientalis (oriental scale) 249 Aesculus hippocastanum (horse chestnut) Foliage feeders Cameraria ohridella (horse-chestnut leaf miner) 114–115 Wood borers Zeuzera pyrina (leopard moth) 226 Agathis Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Allamanda cathartica Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Alnus (alder) Foliage feeders Choristoneura diversana 119, 120 Epirrita autumnata (autumnal moth) 125 Fenusa dohrni 166 Hyphantria cunea (fall webworm) 148 Lymantria dispar (gypsy moth) 143 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma disstria (forest tent caterpillar) 131–132 Malacosoma nuestria 131–132 Operophtera bruceata (Bruce spanworm) 125 Orgyia antiqua (rusty tussock moth) 145–146 Bark and ambrosia beetles Tremex fuscicornis 227 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Monochamus guttatus 214

Alnus glutinosa (common alder, European alder) Wood borers Agrilus viridis 204 Amelanchier Foliage feeders Operopthera bruceata (Bruce spanworm) 125 Orgyia cana 146 Wood borers Saperda candida (roundheaded appletree borer) 220 Tree reproductive structures Megastigmus 303 Anacardium occidentale (cashew) Sucking insects Aonidiella orientalis (oriental scale) 249 Anodopetalum biglandosum Bark and ambrosia beetles Platypus granulosus 201 Antherosperma moschatum Bark and ambrosia beetles Platypus granulosus 201 Araucaria cunninghamii Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Austrocedrus chilensis Sucking insects Cinara cupressivora (cypress aphid) 237 Azadirachta indica (neem) Sucking insects Aonidiella orientalis (oriental scale) 249 Tip, shoot and regeneration insects Apate monachus 273 Wood in use Macrotermes bellicosus 320 B Baccharis Wood borers Megacyllene mellyi 217 Betula (birch) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Choristoneura diversana 119, 120 Erannis defoliaria (mottled umber moth) 125

373

Erannis tiliaria (linden looper) 124, 125 Epirrita autumnata (autumnal moth) 125 Euproctis kargalika (Turkistan browntail moth) 140–141 Fenusa pusilla (birch leaf miner) 165–167 Hyphantria cunea (fall webworm) 148 Lymantria dispar (gypsy moth) 143 Lymantria monacha (nun moth) 144–145 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma disstria (forest tent caterpillar) 131–132 Operophtera brumata bruceata (bruce spanworm) 125 Operopthera brummata (winter moth) 125 Orgyia antiqua (rusty tussock moth) 145–146 Thaumetopoea processionaria (oak processionary caterpillar) 135 Tortrix viridana (green oak tortrix) 123 Bark and ambrosia beetles Scolytus ratzeburgi 186 Wood borers Agrilus viridis 204 Monochamus guttatus 214 Tremex fuscicornis 227 Saperda carcharias (large poplar longhorn beetle) 220 Saperda scaiarias 220 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Betula alleghaniensi (yellow birch) Wood borers Agrilus anxius (bronze birch borer) 204–205 Betula ermanii Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Betula glandulifera Foliage feeders Fenusa pusilla (birch leaf miner) 165–167

374

Host Index

Betula maximowicziana Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Betula papyrifera (paper birch) Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Wood borers Agrilus anxius (bronze birch borer) 204–205 Betula pendula Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Betula platyphylla Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Betula populifolia (gray birch) Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Betula turkestanica Foliage feeders Fenusa pusilla (birch leaf miner) 165–167 Bischofia javanica Foliage feeders Clania variegata (giant bagworm) 114 Broussonetia papyrifera (paper mulberry) Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Butea monosperma (flame of the forest) Sucking insects Aonidiella orientalis (oriental scale) 249 Kerria lacca (lac insect) 248–249 Buxus sempervirens (boxwood) Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 C Caesalpinia echinata (Brazilwood) Bark and ambrosia beetles Megaplatypus mutatus 200 Calcocedrus decurrens (incense cedar) Tree reproductive structures Leptoglossus occidentalis (western conifer seed bug) 296 Callitris Sucking insects

Cinara cupressivora (cypress aphid) 237 Carpinus (hornbeam) Foliage feeders Lymantria monacha (nun moth) 144–145 Malacosoma nuestria 131–132 Thaumetopoea processionea (oak processionary caterpillar) 135 Tortrix viridana (green oak tortrix) 123 Wood borers Monochamus guttatus 214 Tremex fuscicornis 227 Cajanus cajan Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Calotropis procera (giant milkweed) Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Callitris Sucking insects Cinara cupressivora (cypress aphid) 237 Cinara thujafilina 237 Camelia sinensis (tea) Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Caragana Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Caragana korshinskii (peashrub) Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Carya Foliage feeders Erannis tiliaria (linden looper) 124, 125 Hyphantria cunea (fall webworm) 148 Wood borers Megacyllene caryae (painted hickory borer) 217 Tree reproductive structures Curculio caryae (pecan weevil) 298 Cydia caryana (hickory shuckworm) 298, 306

Wood in use Lyctus planicollis (southern lyctus beetle) 324 Carya illinoensis (pecan) Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Wood borers Anoplophora chinensis (citrus longhorn borer) 212 Castanea Foliage feeders Malacosoma nuestria 131–132 Thaumetopoea processionaria (oak processionary caterpillar) 135 Wood borers Zeuzera pyrina (leopard moth) 226 Gall insects Dryocosmus kuriphilus (chestnut gall wasp) 265 Tree reproductive structures Curculio sikkimensis (chestnut curculio) 298 Castanea crenata (Japanese chestnut) Gall insects Dryocosmus kuriphilus (chestnut gall wasp) 265 Castanea dentata (American chestnut) Wood borers Agrilus bilineatus (twolined chestnut borer) 205 Gall insects Dryocosmus kuriphilus (chestnut gall wasp) 265 Tree reproductive structures Curculio caryatrypes (large chestnut weevil) 298 Curculio sayi (small chestnut weevil) 298 Castanea mollissima (Chinese chestnut) Gall insects Dryocosmus kuriphilus (chestnut gall wasp) 265 Tree reproductive structures Curculio davidi 298 Dichocrocis punctiferalis (yellow peach borer) 309 Mechoris cumulatus 299 Castanea sativa (European chestnut) Foliage feeders Euproctis chrysorrhoea (browntail moth) 140 Bark and ambrosia beetles Platypus cylindrus (oak pinhole borer) 201

Host Index Gall insects Dryocosmus kuriphilus (chestnut gall wasp) 265 Tree reproductive structures Curculio elephas (chestnut weevil) 298, 299 Cydia splendana (acorn moth) 306, 307 Castanea vesca (Spanish chestnut) Tree reproductive structures Curculio elephas (chestnut weevil) 299 Castanopsis cuspidata Bark and ambrosia beetles Platypus quercivorus 202 Cassia fistula (golden shower tree) Sucking insects Aonidiella orientalis (oriental scale) 249 Casuarina equisitfolia (Australian pine) Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Casuarina stricta Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Caucarium Wood borers Monochamus griseoplagiatus 214 Ceanothus Foliage feeders Malacosoma californicum (western tent caterpillar) 130–131 Ceanothus velutinus Foliage feeders Orgyia cana 146 Cedrela mexicana Tip, shoot and regeneration insects Hypsipyla grandella 291 Hypsipyla robusta 292 Cedrela odorata (Spanish cedar) Foliage feeders Atta cephalotes 169 Tip, shoot and regeneration insects Hypsipyla grandella 291 Hypsipyla robusta 292 Cedrus Foliage feeders Choristoneura murinana (fir budworm) 119, 122 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135

Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Urocerus gigas 231 Tip, shoot and regeneration insects Pissodes nemorensis (deodar weevil) 278 Tree reproductive structures Dioryctria abietella 310, 311 Megastigmus pinsapinis 304 Cedrus brevifolia Tree reproductive structures Megastigmus schimitscheki 304 Cedrus deodara Tip, shoot and regeneration insects Hylobius angustus 275 Tree reproductive structures Dioryctria abietella 310, 311 Cedrus libani (cedar of Lebanon) Foliage feeders Calomicrus apicalis 152 Celtis (hackberry) Wood borers Megacyllene caryae (painted hickory borer) 217 Monochamus griseoplagiatus 214 Tremex fuscicornis 227 Gall insects Pachypsylla celtidismamma (hackberry nipple gall) 254 Celtis occidentalis (eastern hackberry) Gall insects Pachypsylla celtismamma (hackberry nipple gall) 254 Cecropia Foliage feeders Acromyrmex octospinosus 168 Chamaecyparis Bark and ambrosia beetles Phloeosinus bicolor 185 Wood borers Urocerus gigas 231 Sucking insects Cinara thujafilina 237 Chamaecyparis lawsoniana (Port Orford cedar) Insects of tree reproductive structures Orsillus depressus (cypress seed bug) 296 Orsillus maculatus (cypress seed bug) 296

375

Chukrasia tabularis Tip, shoot and regeneration insects Hypsipyla robusta 292 Cinnamomum camphora (camphor tree) Wood borers Monochamus bimaculatus 214 Citrus Foliage feeders Atta cephalotes 169 Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Anoplophora chinensis (citrus longhorn borer) 212 Sucking insects Aonidiella orientalis (Oriental scale) 249 Coccoloba (seagrape) Foliage feeders Sericoceros krugii 158 Sericoceros mexicanus 158 Coccoloba uvifera Foliage feeders Sericoceros mexicanus 158 Coccoloba venosa Foliage feeders Sericoceros mexicanus 158 Coffea (coffee) Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Wood borers Monochamus griseoplagiatus 214 Coffea arabica Wood borers Monochamus leuconotus 214 Colophospermum mopane (mopane) Foliage feeders Imbrasia belina (mopane worm, mopane emperor moth) 136–137 Chloroxylon swietenia (Ceylon satinwood) Sucking insects Aonidiella orientalis (oriental scale) 249 Tip, shoot and regeneration insects Hypsipyla robusta 292 Corylus Foliage feeders Leucoma salicis (satin moth) 141–142 Lymantria dispar (gypsy moth) Malacosoma nuestria 131–132 Thaumetopoea processionea (oak processionary caterpillar) 135

376

Host Index

Corylus (Continued ) Wood borers Monochamus guttatus 214 Tree reproductive structures Curculio glandium 298 Curculio neocorylus (hazelnut weevil) 298 Curculio nucum 298 Corylus californica Tree reproductive structures Curculio occidentalis (filbert weevil) 298 Corymbia calophylla Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Cotoneaster Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Malacosoma nuestria 131–132 Couma macrocarpa (leche caspi) Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Crataegus Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Lymantria dispar (gypsy moth) Malacosoma nuestria 131–132 Wood borers Saperda candida (roundheaded appletree borer) 220 Cynometra Wood borers Monochamus griseoplagiatus 214 Cryptocarya alba (peumo) Foliage feeders Ormiscodes cinnamomea 137–138 Wood borers Rhyephenes mallei 223 Rhyephenes humeralis 223 Cryptomeria japonica Bark and ambrosia beetles Platypus quercivorus 202 Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Cupressus Bark and ambrosia beetles Phloeosinus armatus 185 Phloeosinus bicolor 185 Sucking insects Cinara cupressi 237

Cinara cupressivora (cypress aphid) 237 Cinara thujafilina 237 Tree reproductive structures Megastigmus atlanticus 304 Megastigmus wachtli 304, 305 Orsillus depressus (cypress seed bug) 296 Orsillus maculatus (cypress seed bug) 296 Pseudococcyx tessulatana 309 Cupressus abramsiana (Santa Cruz cypress) Tree reproductive structures Megastigmus wachtli 304, 305 Cupressus arizonica (Arizona cypress) Tree reproductive structures Megastigmus wachtli 304, 305 Cupressus bakeri Tree reproductive structures Megastigmus wachtli 304, 305 Cupressus goveniana Tree reproductive structures Megastigmus wachtli 304, 305 Cupressus lusitanica Foliage feeders Atta cephalotes 169 Glena bisulca 125, 127 Sucking insects Cinara cupressivora (cypress aphid) 237 Cupressus sempervirens (Mediterranean cypress) Bark and ambrosia beetles Phloeosinus armatus 185 Phloeosinus bicolor 185 Sucking insects Cinara cupressivora (cypress aphid) 237 Insects of tree reproductive structures Megastigmus wachtli 304–305 Orsillus maculatus (cypress seed bug) 296 Cupressus torulosa Tree reproductive structures Megastigmus cupressi (cypress seed fly) 304 Cydonia Foliage feeders Lymantria obfuscata (Indian gypsy moth) 143 Cynometra Wood borers Monochamus griseoplagiatus 214

D Dalbergia sissoo (Indian rosewood) Sucking insects Aonidiella orientalis (oriental scale) 249 Tip, shoot and regeneration insects Apate monachus 273 Diospyros Hyphantria cunea (fall webworm) 148 Diospyros melanoxylon (East Indian ebony) Wood borers Alcterogystia cadambae (teak carpenterworm) 224 E Echites umbellata Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Elaeagnus Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 186, 187 Entandrophragma angolense Tip, shoot and regeneration insects Hypsipyla robusta 292 Entandrophragma utile Tip, shoot and regeneration insects Hypsipyla robusta 292 Erythrina Gall insects Quadrastichus erythrinae (erythrina gall wasp) 260 Eschweilera corrugate Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Eucalyptus Foliage feeders Chrysophtharta agricola 152 Chrysophtharta bimaculata (Tasmanian eucalyptus leaf beetle) 152 Chrysophtharta varicollis 152 Didymuria violescens (spurlegged phasmid) 150–151 Gonipterus gibberus 155 Gonipterus scutellatus (eucalyptus weevil) 154 Lophyrotoma interrupta (cattle poisoning sawfly) 157 Paropsis atomaria 152 Paraopsis charbdis 152 Perga affinis 156

Host Index Pergarograpta bella 157 Pseudoperga lewisii (pale brown sawfly) 157 Sarsina violascens 147–148 Bark and ambrosia beetles Megaplatypus mutatus 200 Platypus cylindrus (oak pinhole borer) 201 Platypus granulosus 201 Xyleborus ferrugineus 197–198 Wood borers Phoracantha recurva (yellow phoracantha borer) 219 Phoracantha semipunctata (eucalyptus longhorn borer) 218–219 Gall insects Ctenarytainia spatulata 235 Leptocybe invasa (blue gum chalcid) 261 Ophelimus eucalypti (eucalyptus gall wasp) 261 Tip, shoot and regeneration insects Apate monachus 273 Wood in use Coptotermes acinaciformis 317 Lyctus brunneus (brown lyctus beetle) 323 Lyctus cavicollis (western lyctus beetle) 325 Macrotermes bellicosus 320 Porotermes adamsoni 316 Eucalyptus amygdalina Wood in use Porotermes adamsoni 316 Eucalyptus baxteri Wood in use Porotermes adamsoni 316 Eucalyptus blakelyi Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus botryoides Gall insects Leptocybe invasa (blue gum chalcid) 261 Ophelimus eucalypti (eucalyptus gall wasp) 261 Eucalyptus brassiana Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus bridgesiana Sucking insects

Glycaspis brimblecombei (red gum lerp) 235 Gall insects Leptocybe invasa (blue gum chalcid) 261 Eucalyptus camaldulensis (red gum) Foliage feeders Acromyrmex laticeps nigrosetosus 168 Gonipterus scutellatus (eucalyptus weevil) 154 Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Gall insects Leptocybe invasa (blue gum chalcid) 261 Ophelimus eucalypti (eucalyptus gall wasp) 261 Eucalyptus citriodora Foliage feeders Sarsina violascens 147–148 Eucalyptus cloeziana Foliage feeders Sarsina violascens 147–148 Eucalyptus dealbata Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus deanei Gall insects Ophelimus eucalypti (eucalyptus gall wasp) 261 Eucalyptus delegatensis (alpine ash) Foliage feeders Chrysophtharta bimaculata (Tasmanian eucalyptus leaf beetle) 152 Didymuria violescens (spurlegged phasmid) 150–151 Eucalyptus diversicolor Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus ficifolia Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus globulus (blue gum) Sucking insects Ctenarytaina eucalypti (blue gum psyllid) 235

377

Glycaspis brimblecombei (red gum lerp) 235 Gall insects Leptocybe invasa (blue gum chalcid) 261 Ophelimus eucalypti (eucalyptus gall wasp) 261 Selitrichodes globulus (blue gum gall wasp) 262 Wood in use Porotermes adamsoni 316 Eucalyptus gomphocephala Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus grandis Foliage feeders Sarsina violascens 147–148 Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Gall insects Leptocybe invasa (blue gum chalcid) 261 Ophelimus eucalypti (eucalyptus gall wasp) 261 Eucalyptus gunnii Gall insects Leptocybe invasa (blue gum chalcid) 261 Eucalyptus lehmannii Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus maculata Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus maidenii (maiden’s gum) Foliage feeders Gonipterus scutellatus (eucalyptus weevil) 154 Eucalyptus mannifera Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus nesophila (Melville Island bloodwood) Foliage feeders Sarsina violascens 147–148 Eucalyptus nicholii Sucking insects Glycaspis brimblecombei (red gum lerp) 235

378

Host Index

Eucalyptus nitens Foliage feeders Chrysophtharta bimaculata (Tasmanian eucalyptus leaf beetle) 152 Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus obliqua Wood in use Porotermes adamsoni 316 Eucalyptus paniculata Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus pauciflora Wood in use Porotermes adamsoni 316 Eucalyptus propinqua Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus pulverulenta Sucking insects Ctenarytaina eucalypti (blue gum psyllid) 235 Eucalyptus punctata Foliage feeders Gonipterus scutellatus (eucalyptus weevil) 154 Eucalyptus redunca Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus regnans Foliage feeders Didymuria violescens (spurlegged phasmid) 150–151 Chrysophtharta bimaculata (Tasmanian eucalyptus leaf beetle) 152 Wood in use Porotermes adamsoni 316 Eucalyptus resinifera Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus robusta (swamp mahogany) Foliage feeders Gonipterus scutellatus (eucalyptus weevil) 154 Gall insects Leptocybe invasa (blue gum chalcid) 261

Eucalyptus rudis Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus saligna Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Gall insects Leptocybe invasa (blue gum chalcid) 261 Eucalyptus sideroxylon (mugga, red ironbark) Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Eucalyptus sieberi Wood in use Porotermes adamsoni 316 Eucalyptus smithii (gully gum) Foliage feeders Gonipterus scutellatus (eucalyptus weevil) 154 Eucalyptus tereticornis Sucking insects Glycaspis brimblecombei (red gum lerp) 235 Gall insects Leptocybe invasa (blue gum chalcid) 261 Eucalyptus tenuiramis Wood in use Porotermes adamsoni 316 Eucalyptus triantha Wood borers Phoracantha acanthocera (bulls-eye borer) 217–218 Eucalyptus urophylla Foliage feeders Sarsina violascens 147–148 Eucalyptus viminalis Foliage feeders Gonipterus scutellatus (eucalyptus weevil) 154 Gall insects Leptocybe invasa (blue gum chalcid) 261 Eucryphia lucida (leatherwood) Bark and ambrosia beetles Platypus granulosus 201 F Fagus Foliage feeders Choristoneura diversana 119, 120

Erannis defoliaria (mottled umber moth) 125 Malacosoma nuestria 131–132 Operophtera bruceata (Bruce spanworm) 125 Thaumetopoea processionaria (oak processionary caterpillar) 135 Tortrix viridana (green oak tortrix) 123 Wood borers Cossus cossus (goat moth) 224 Tremex fuscicornis 227 Sucking insects Cryptococcus fagisuga (beech scale) 246 Lepidosaphes ulmi (oystershell scale) 250–251 Fagus grandifolia (American beech) Sucking insects Cryptococcus fagisuga (beech scale) 246 Fagus sylvatica (European beech) Foliage feeders Choristoneura diversana 119, 120 Erannis defoliaria (mottled umber moth) 125 Malacosoma nuestria 131–132 Operophtera bruceata (bruce spanworm) 125 Thaumetopoea processionaria (oak processionary caterpillar) 135 Tortrix viridana (green oak tortrix) 123 Bark and ambrosia beetles Platypus cylindrus (oak pinhole borer) 201 Wood borers Agrilus pannonicus 206 Agrilus viridis 204 Sucking insects Cryptococcus fagisuga (beech scale) 246 Tree reproductive structures Cydia fagiglandana 306 Ficus Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Monochamus bimaculatus 214 Sucking insects Kerria lacca (lac insect) 248–249

Host Index Ficus elastica Sucking insects Kerria lacca (lac insect) 248–249 Fortunella margarita (kumquat) Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Fraxinus (ash) Foliage feeders Choristoneura diversana 119, 120 Hyphantria cunea (fall webworm) 148 Leucoma salicis (satin moth) 141–142 Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Agrilus planipennis (emerald ash borer) 204, 206–207 Cossus cossus (goat moth) 224 Megacyllene caryae (painted hickory borer) 217 Podesesia syringae (lilac/ash borer) 226 Urocerus gigas 231 Zeuzera pyrina (leopard moth) 226 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Tree reproductive structures Boisea rubrolineata (western boxelder bug) 297 Boisea trivittata (boxelder bug) 297 Wood in use Lyctus brunneus (brown lyctus beetle) 323 Lyctus planicollis (southern lyctus beetle) 324 Fraxinus excelsior (European ash) Foliage feeders Stereonychus fraxini (ash weevil) 155 Fraxinus pennsylvanica (green ash) Wood borers Prionoxystus robinae (carpenterworm) 225 G Gleditsia triacanthos (honey locust) Foliage feeders Odontata dorsalis (locust leaf miner) 153–154 Wood borers

Megacyllene caryae (painted hickory borer) 217 Gmelina arborea Foliage feeders Atta cephalotes 169 Wood borers Xyleutes ceramica (teak beehole borer) 225 Grewia tiliaefolia Wood borers Alcterogystia cadambae (teak carpenterworm) 224

H Hevea brasiliensis (rubber tree) Bark and ambrosia beetles Xyleborus similis 198 Hibiscus Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Sucking insects Maconellicoccus hirsutus (pink hibiscus mealybug) 245 Hibiscus elatus Sucking insects Maconellicoccus hirsutus (pink hibiscus mealybug) 245 Himatanthus Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 I Ilex chinensis Bark and ambrosia beetles Platypus quercivorus 202 Ilex paraguariensis Tip, shoot and regeneration insects Hedypathes betulinus 274 J Juglans (walnut) Foliage feeders Lymantria obfuscata (Indian gypsy moth) 143 Hyphantria cunea (fall webworm) 148 Wood borers Megacyllene caryae (painted hickory borer) 217 Monochamus guttatus 214 Tremex fuscicornis 227 Zeuzera pyrina (leopard moth) 227

379

Tip, shoot and regeneration insects Pityophthorus juglandis (walnut twig beetle) 284 Tree reproductive structures Cydia splendana (acorn moth) 307 Juglans californica Tip, shoot and regeneration insects Pityophthorus juglandis (walnut twig beetle) 284 Juglans major Tip, shoot and regeneration insects Pityophthorus juglandis (walnut twig beetle) 284 Juglans mandshuria (Manchurian walnut) Wood borers Agrilus planipennis (emerald ash borer) 206–207 Juglans nigra (black walnut) Tip, shoot and regeneration insects Pityophthorus juglandis (walnut twig beetle) 284 Juglans regia (English walnut, Persian walnut) Wood borers Anoplophora chinensis (citrus longhorn borer) 212 Juniperus (juniper) Foliage feeders Choristoneura murinana (fir budworm) 119, 122 Malacosoma nuestria 131–132 Bark and ambrosia beetles Phloeosinus armatus 185 Trypodendron lineatum (striped ambrosia beetle) 196 Sucking insects Cinara cupressivora (cypress aphid) 237 Cinara thujafilina 237 Gall insects Oligotrophus apicis 266 Oligotrophus betheli (juniper felt tip midge) 266 Oligotrophus gemmarum 266 Oligotrophus juniperinus 266 Oligotrophus nezu 266 Oligotrophus panteli 266 Tree reproductive structures Megastigmus certus 303, 304 Orsillus depressus (cypress seed bug) 296 Juniperus communis (common juniper) Tree reproductive structures Megastigmus bipunctatus 304

380

Host Index

Juniperus occidentalis (western juniper) Gall insects Oligotrophus betheli (juniper felt tip midge) 266 Juniperus osteosperma (Utah juniper) Gall insects Oligotrophus betheli (juniper felt tip midge) 266 Juniperus pingii Tree reproductive structures Megastigmus pingii 303, 304 Juniperus polycarpos Tree reproductive structures Megastigmus bipunctatus 304 Juniperus procera (East African pencil cedar) Sucking insects Cinara cupressivora (cypress aphid) 237 Tree reproductive structures Megastigmus somaliensis 303, 304 Juniperus scopulorum (Rocky Mountain juniper) Gall insects Oligotrophus betheli (juniper felt tip midge) 266 Juniperus thurifera Tree reproductive structures Megastigmus bipunctatus 304 Juniperus virginiana (eastern red cedar) Foliage feeders Thyridopteryx ephemeraeformis (bagworm) 113–114 Gall insects Oligotrophus betheli (juniper felt tip midge) 266 K Khaya (African mahogany) Wood borers Monochamus griseoplagiatus 214 Khaya anthotheca Tip, shoot and regeneration insects Hypsipyla robusta 292 Khaya grandiflora Tip, shoot and regeneration insects Hypsipyla robusta 292 Khaya ivorensis Tip, shoot and regeneration insects Hypsipyla robusta 292 Khaya nyasica Tip, shoot and regeneration insects Hypsipyla robusta 292 Khaya senegalensis Tip, shoot and regeneration insects

Hypsipyla grandella 291 Hypsipyla robusta 292 L Lachnophylis Monochamus griseoplagiatus 214 Larix (larch) Foliage feeders Coleophora laricella (larch casebearer) 117 Dendrolimus sibiricus (Siberian silk moth) 128–130 Lymantria monacha (nun moth) 144–145 Malacosoma nuestria 131–132 Nesodiprion japonica 164 Orgyia antiqua (rusty tussock moth) 145–146 Orgyia thyellina 146 Bark and ambrosia beetles Ips cembrae 190 Ips sexdentatus 192 Ips subelongatus 190,193 Ips typographus (larger European spruce beetle) 190, 193 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Melanophila guttulata (larch buprestid) 209 Monochamus alternatus (Japanese pine sawyer) 214, 215 Monochamus impluviatus 214 Monochamus nitens 214 Monochamus urossovi (fir sawyer) 214, 216 Phaenops drummondi (flat headed fir borer) 210 Tetropium castaneum 221–222 Tetropium cinnamopterum (eastern larch borer) 222 Tetropium fuscum (brown spruce longhorn beetle) 222 Tetropium gabrieli (larch longhorn beetle) 222 Tetropium gracilicorne 222–223 Sirex juvencus 230 Sirex noctilio (sirex woodwasp) 228 Urocerus gigas 231 Sucking insects Adelges isedakii 256 Adelges tardoides 256 Adelges torii 256

Gall insects Dasineura sp. 265–266 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Hylobius albosparsus (white spotted weevil) 275 Hylobius pales (pales weevil) 275, 277 Hylobius piceus 275 Pissodes castaneus 278 Tree reproductive structures Cydia illutana illutana 306 Dioryctria abietivorella 310, 311 Eucosma impropriana 308 Megastigmus pictus 304 Larix czekanowski Gall insects Dasineura rozhkovi 265–266 Larix decidua (European larch) Foliage feeders Bupalus piniaria (pine looper) 125 Coleophora laricella (larch casebearer) 117 Zeiraphera diniana (gray larch bud moth) 123 Bark and ambrosia beetles Ips acuminatus 190, 191 Wood borers Phaenops cyanea 210 Sirex juvencus 230 Sucking insects Adelges laricis 256 Adelges viridis 256 Gall insects Dasineura kellneri 265–266 Tip, shoot and regeneration insects Pissodes notatus (banded pine weevil) 278 Tomicus piniperda 285 Larix gmelinii Foliage feeders Erannis jacobsoni (Jacobson’s spanworm) 125 Bark and ambrosia beetles Scolytus morawitzi 186 Ips subelongatus 193 Wood borers Melanophila guttulata (larch buprestid) 209 Gall insects Dasineura rozhkovi 265–266 Dasineura verae 265–266 Tree reproductive structures

Host Index Cydia illutana dahuricolana (Dahurian larch seed moth) 306 Eucosma impropria 308 Larix kaempferi Sucking insects Adelges japonicus 256 Larix kamtschatica Bark and ambrosia beetles Scolytus morawitzi 186 Gall insects Dasineura nipponica 265–266 Larix laricina (eastern larch) Foliage feeders Coleophora laricella (larch casebearer) 117 Zeiraphera sp. 124 Bark and ambrosia beetles Dendroctonus simplex (larch beetle) 177 Wood borers Melanophila fulvoguttata (hemlock borer) 209 Sucking insects Adelges lariciatus 256 Tree reproductive structures Dioryctria reniculelloides (spruce coneworm) 311 Larix leptolepis (Japanese larch) Foliage feeders Lymantria umbrosa (Hokkiado gypsy moth) 143 Bark and ambrosia beetles Ips subelongatus 193 Larix lyallii (subalpine larch) Bark and ambrosia beetles Scolytus laricis (western larch beetle) 186 Sucking insects Adelges lariciatus 256 Larix occidentalis (western larch) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Coleophora laricella (larch casebearer) 117 Zeiraphera improbana 124 Bark and ambrosia beetles Dendroctonus pseudotsugae (Douglas-fir beetle) 182 Scolytus laricis (western larch beetle) 186 Wood borers

Tetropium velutinum (western larch borer) 222 Larix sibirica (Siberian larch) Foliage feeders Erannis jacobsoni (Jacobson’s spanworm) 125 Lymantria dispar asiatica (Asian gypsy moth) 143 Bark and ambrosia beetles Ips hauseri 190 Scolytus morawitzi 186 Wood borers Melanophila guttulata (larch buprestid) 209 Gall insects Dasineura rozhkovi 265–266 Dasineura verae 265–266 Tree reproductive structures Cydia illutana dahuricolana (Dahurian larch seed moth) 306 Eucosma impropria 308 Larix sukaczerii Bark and ambrosia beetles Scolytus morawitzi 186 Laurus nobilis (sweet bay, laurel) Bark and ambrosia beetles Megaplatypus mutatus 200 Lespedeza Foliage feeders Choristoneura diversana 119, 120 Leucaena Sucking insects Heteropsylla cubana (leucaena psyllid) 234 Leucana leucocephala Sucking insects Heteropsylla cubana (leucaena psyllid) 234 Lindera erythrocarpa Bark and ambrosia beetles Platypus quercivorus 202 Lindera latifolia Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Liquidambar Foliage feeders Hyphantria cunea (fall webworm) 148 Litchi sinensis Wood borers Anoplophora chinensis (citrus longhorn beetle) 212

381

Lithocarpus edulis Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Lithrea caustica (litre) Foliage feeders Ormiscodes cinnamomea 137–138 Litsea elongata Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Llamandra cathartica Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Lonchocarpus margaretensis Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Lovoa trichilioides Tip, shoot and regeneration insects Hypsipyla robusta 292 M Maesopsis eminii (umbrella tree) Foliage feeders Imbrasia nicitans 137 Wood borers Monochamus griseoplagiatus 214 Magnolia grandiflora (southern magnolia) Bark and ambrosia beetles Megaplatypus mutatus 200 Mallotus philippensis (kamala tree) Wood borers Monochamus bimaculatus 214 Malus (apple) Foliage feeders Choristoneura diversana 119, 120 Erannis tiliaria (linden looper) 124, 125 Euproctis kargalika (Turkistan browntail moth) 140–141 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma americanum (eastern tent caterpillar) 131 Malacosoma indica (Indian tent caterpillar) 131 Malacosoma nuestria 131–132 Operopthera brummata (winter moth) 125 Orgyia antiqua (rusty tussock moth) 145–146 Orgyia thyellina 146 Bark and ambrosia beetles Megaplatypus mutatus 200

382

Host Index

Malus (apple) (Continued ) Scolytus rugulosus (shothole borer) 186 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Cossus cossus (goat moth) 224 Saperda candida (round headed appletree borer) 220 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Malus pumila Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 186, 187 Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Malus spectabilis Wood borers Anoplophora chinensis (citrus longhorn beetle) 212 Mangifera indica (mango) Foliage feeders Atta cephalotes 169 Melia azedarach (Chinaberry) Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Anoplophora chinensis (citrus longhorn beetle) 212 Sucking insects Aonidiella orientalis (oriental scale) 249 Melicoccus bijugatum Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Mikania Foliage feeders Sarsina violascens 147–148 Milicia Gall insects Phytoloma sp. 254 Phytoloma tuberculata 254 Milicia excelsia Gall insects Phytoloma fusca 254 Milicia regia Gall insects Phytoloma excelsia 254 Morus (Mulberry) Foliage feeders Lymantria obfuscata (Indian gypsy moth) 143

Malacosoma nuestria 131–132 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Megacyllene caryae (painted hickory borer) 217 Morus alba (white mulberry) Wood borers Anoplophora chinensis (citrus longhorn borer) 212 N Nothofagus (southern beech) Foliage feeders Hylamorpha elegans 151 Ormiscodes cinnamomea 137–138 Bark and ambrosia beetles Gnathotrupes barbifer 196 Gnathotrupes nanus 196 Gnathotrupes vafer 196 Gnathotrupes velatus 196 Wood borers Epistomentis pictus 208 Nothofagus alpina (rauli) Foliage feeders Cerospastus volupis 155 Wood borers Epistomentis pictus 208 Nothofagus cunninghamii (myrtle beech) Bark and ambrosia beetles Platypus granulosus 201 Nothofagus dombeyi (coigüe) Foliage feeders Ormiscodes cinnamomea 137–138 Wood borers Epistomentis pictus 208 Holopterus chilensis 213 Rhyephenes humeralis 223 Rhyephenes mallei 223 Nothofagus obliqua (roble) Foliage feeders Cerospastus volupis 155 Ormiscodes cinnamomea 137–138 Wood borers Epistomentis pictus 208 Holopterus chilensis 213 Nothofagus pumilio (lenga) Wood borers Epistomentis pictus 208 O Olea europea (olive) Foliage feeders Stereonychus fraxini (ash weevil) 155

Osmunthus fragrans Foliage feeders Sarsina violascens 147–148 Ougeinia dalbergioides Sucking insects Kerria lacca (lac insect) 248–249 P Pasania glabra Bark and ambrosia beetles Platypus quercivorus 202 Pawlonia Foliage feeders Clania variegata (giant bagworm) 114 Persia americanum (redbay) Bark and ambrosia beetles Megaplatypus mutatus 200 Xyleborus glabratus (redbay ambrosia beetle) 198–199 Persia borbonia Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Peumus boldus (boldo) Foliage feeders Ormiscodes cinnamomea 137–138 Plumeria (frangipani) Foliage feeders Pseudosphiinx tetrio (frangipani hawk moth) 139 Phillyrea latifolia Foliage feeders Stereonychus fraxini (ash weevil) 155 Phoebe lanceolata Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Phyllocladus aspleniifolius Bark and ambrosia beetles Platypus granulosus 201 Picea (spruce) Foliage feeders Choristoneura murinana (fir budworm) 119, 122 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Coleotechnites piceaella 118 Dendrolimus sibiricus (Siberian silk moth) 128–130 Gilpinia hercyniae (European spruce sawfly) 158–159 Gilpinia polytoma 159

Host Index Lymantria monacha (nun moth) 144–145 Neodiprion abietis (balsam fir sawfly) 161 Orgyia antiqua (rusty tussock moth) 145–146 Thyridopteryx ephemeraeformis (bagworm) 113–114 Bark and ambrosia beetles Dendroctonus micans 177, 180 Dendroctonus rufipennis (spruce beetle) 177, 183 Dendroctonus valens (red turpentine beetle) 184 Ips pini (pine engraver) 190, 192 Ips sexdentatus 190, 192 Ips subelongatus 190, 193 Ips typographus (larger European spruce beetle) 190, 193 Orthotomicus erosus (Mediterranean pine engraver) 195 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Melanophila acuminata (black fire beetle) 208 Melanophila fulvoguttata (hemlock borer) 209 Melanophila guttulata (larch buprestid) 209 Monochamus alternatus (Japanese pine sawyer) 214, 215 Monochamus galloprovincialis 214 Monochamus grandis 214 Monochamus nitens 214 Monochamus notatus (northeastern pine sawyer) 214 Monochamus saltuarius 214 Monochamus sartor 214 Monochamus scutellatus (white spotted sawyer) 214, 216 Monochamus sutor 214 Monochamus urossovi (fir sawyer) 214, 216 Phaenops drummondi (flat headed fir borer) 210 Saperda interrupta 220 Tetropium castaneum 221–222 Tetropium gracilicorne 222–223 Tetropium velutinum (western larch borer) 222 Sirex juvencus 230 Sirex noctilio (sirex woodwasp) 228 Urocerus gigas 231

Sucking insects Elatobium abietinum (green spruce aphid) 238 Gall insects Adelges abietis (eastern spruce gall aphid) 256 Adelges cooleyi (Cooley spruce gall adelgid) 256–257 Adelges glandulae 256 Adelges isedakii 256 Adelges japonicus 256 Adelges knucheli 256 Adelges lariciatus 256 Adelges laricis 256 Adelges lapponicus 256 Adelges nebrodensis 256 Adelges nordmanniana 256 Adelges pectinata 256 Adelges prelli 256 Adelges tardoides 256 Adelges torii 256 Adelges tsugae (hemlock woolly adelgid) 239 Adelges viridis 256 Pineus cembrae 258 Pineus pinifoliae (pine leaf chermes) 258 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Hylobius abietis (large pine weevil) 275–276 Hylobius pales (pales weevil) 275, 277 Pissodes approximatus (northern pine weevil) 278 Pissodes castaneus 278 Pissodes hercyniae 278 Pissodes piceae 278 Pissodes strobi 278 Pityophthorus tuberculatus 283 Tomicus minor 285 Tree reproductive structures Cydia illutana illutana 306 Cydia strobilella (spruce seed moth) 306 Dioryctria abietivorella 310, 311 Dioryctria abietella 310, 311 Dioryctria reniculelloides (spruce coneworm) 311 Megastigmus atedius 304 Wood in use Anobium punctatum (furniture beetle) 325

383

Buprestis aurulenta (golden buprestid) 326 Hylotrupes bajulus (old-house borer) 326 Picea abies (Norway spruce) Bark and ambrosia beetles Dendroctonus micans 180 Ips typographus (larger European spruce beetle) 193 Foliage feeders Gilpinia hercyinae 159–159 Wood borers Phaenops cyanea 210 Tetropium fuscum (brown spruce longhorn beetle) 222 Sirex juvencus 230 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tip, shoot and regeneration insects Pissodes notatus (banded pine weevil) 278 Pissodes strobi (white pine weevil) 278 Tomicus piniperda 285 Tree reproductive structures Dioryctria abietella 310, 311 Picea ajanensis Tip, shoot and regeneration insects Tomicus puellus 285 Picea asperata (dragon spruce) Bark and ambrosia beetles Dendroctonus micans 180 Picea breweriana (Brewer spruce) Bark and ambrosia beetles Dendroctonus micans 180 Picea engelmannii (Engelmann spruce) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Dasychira grisefacta (western pine tussock moth) 139–140 Nepytia janetae 125 Bark and ambrosia beetles Dendroctonus micans 180 Dendroctonus rufipennis (spruce beetle) 183 Wood borers Tetropium parvulum (northern spruce borer) 222 Sucking insects Elatobium abietinum (green spruce aphid) 238 Tip, shoot and regeneration insects

384

Host Index

Picea engelmannii (Engelmann spruce) (Continued ) Eucosma sonomana (western pine shoot borer) 287 Pissodes strobi (white pine weevil) 278 Picea glauca (white spruce) Foliage feeders Choristoneura fumiferana (spruce budworm) 119, 120–122 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Glipinia hercyniae 158–159 Dasychira grisefacta (western pine tussock moth) 139–140 Bark and ambrosia beetles Dendroctonus micans 180 Dendroctonus punctatus 177 Dendroctonus rufipennis (spruce beetle) 183 Wood borers Tetropium cinnamopterum (eastern larch borer) 222 Tetropium fuscum (brown spruce longhorn beetle) 222 Tetropium parvulum (northern spruce borer) 222 Tip, shoot and regeneration insects Pissodes strobi (white pine weevil) 278 Tree reproductive structures Dioryctria disclusa (webbing coneworm) 311 Picea jezoensis (Jezo spruce) Bark and ambrosia beetles Dendroctonus micans 180 Ips typographus (larger European spruce beetle) 193 Wood borers Sirex juvencus 230 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Picea likiangensis Gall insects Pineus armandicola 258 Picea mariana (black spruce) Foliage feeders Choristoneura fumiferana (spruce budworm) 119, 120–122 Bark and ambrosia beetles Dendroctonus micans 180 Wood borers

Tetropium fuscum (brown spruce longhorn beetle) 222 Gall insects Pineus floccus (red spruce adelgid) 258 Tip, shoot and regeneration insects Pissodes strobi (white pine weevil) 278 Picea obovata (Siberian spruce) Bark and ambrosia beetles Dendroctonus micans 180 Ips acuminatus 191 Ips typographus (larger European spruce beetle) 193 Tip, shoot and regeneration insects Tomicus piniperda 285 Tree reproductive structures Cydia illutana dahuricolana (Dahurian larch seed moth) 306 Picea orientalis (Caucasian spruce) Bark and ambrosia beetles Dendroctonus micans 180 Ips acuminatus 191 Ips typographus (larger European spruce beetle) 193 Wood borers Sirex juvencus 230 Gall insects Pineus orientalis 258 Picea omorika (Serbian spruce) Bark and ambrosia beetles Dendroctonus micans 180 Picea polita Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Picea pungens (blue spruce) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Orgyia pseudotsugata (Douglas-fir tussock moth) 146–147 Bark and ambrosia beetles Dendroctonus micans 180 Ips hunteri (blue spruce ips) 190 Wood borers Tetropium fuscum (brown spruce longhorn beetle) 222 Sucking insects Elatobium abietinum (green spruce aphid) 238

Tip, shoot and regeneration insects Pissodes strobi (white pine weevil) 278 Picea rubens (red spruce) Foliage feeders Choristoneura fumiferana (spruce budworm) 119, 120–122 Bark and ambrosia beetles Dendroctonus punctatus 177 Dendroctonus rufipennis (spruce beetle) 183 Wood borers Tetropium fuscum (brown spruce longhorn beetle) 222 Gall insects Pineus floccus (red spruce adelgid) 258 Tip, shoot and regeneration insects Pissodes strobi (white pine weevil) 278 Picea schrenkiana (Schrenk’s spruce) Bark and ambrosia beetles Ips hauseri 190 Wood borers Tetropium staudingeri (Seven-river spruce borer) 222–223 Picea sitchensis (Sitka spruce) Foliage feeders Choristoneura orea 119 Bark and ambrosia beetles Dendroctonus micans 180 Dendroctonus punctatus 177 Ips typographus (larger European spruce beetle) 193 Wood borers Tetropium fuscum (brown spruce longhorn beetle) 222 Sirex juvencus 230 Sucking insects Elatobium abietinum (green spruce aphid) 238 Tip, shoot and regeneration insects Pissodes strobi (white pine weevil) 278 Picea smithiana Tip, shoot and regeneration insects Hylobius angustus 275 Tomicus piniperda 285 Tree reproductive structures Cydia ethelinda 306 Dioryctria abietella 310, 311 Pithecellobium pinnatum Bark and ambrosia beetles Xyleborus ferrugineus 197–198

Host Index Pinus (pine) Foliage feeders Bupalus piniaria (pine looper) 125 Choristoneura murinana (fir budworm) 119, 122 Clania variegata (giant bagworm) 114 Coloradia pandora (pandora moth) 135–136 Dendrolimus houi (Yunnan pine caterpillar) 128 Dendrolimus kikuchii 128 Dendrolimus pini 127, 128 Dendrolimus sibiricus (Siberian silk moth) 128–130 Lymantria monacha (nun moth) 144–145 Neodiprion fulviceps 161 Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion sertifer (European pine sawfly) 161, 163–164 Nesodiprion biremis 164 Nesodiprion japonica 164 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Thyridopteryx ephemeraeformis (bagworm) 113–114 Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus adjunctus (round headed pine beetle) 177 Dendroctonus approximatus (larger Mexican pine beetle) 177 Dendroctonus frontalis (southern pine beetle) 177–179 Dendroctonus mexicanus (smaller Mexican pine beetle) 177 Dendroctonus micans 177 Dendroctonus ponderosae (mountain pine beetle) 177, 181 Dendroctonus rhizophagus 177 Dendroctonus terebrans (black turpentine beetle) 177 Dendroctonus valens (red turpentine beetle) 177, 184 Ips acuminatus 190, 191 Ips calligraphus (six-spined engraver beetle) 190, 191 Ips grandicollis (southern pine engraver) 190, 192

Ips lecontei 190 Ips pini (pine engraver) 190, 192 Ips sexdentatus 190, 192 Ips typographus (larger European spruce beetle) 190, 193 Myoplatypus flavicornis 200 Orthotomicus erosus (Mediterranean pine engraver) 195 Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Buprestis apricans (turpentine borer) 207 Buprestis novemmaculata 208 Melanophila acuminata (black fire beetle) 208 Melanophila guttulata (larch buprestid) 209 Monochamus alternatus 214, 215 Monochamus caroliniensis 214 Monochamus clamator 214 Monochamus galloprovincialis 214 Monochamus mutator 214 Monochamus nitens 214 Monochamus notatus (northeastern pine sawyer) 214 Monochamus saltuarius 214 Monochamus sartor 214 Monochamus scutellatus (white spotted sawyer) 214, 216 Monochamus sutor 214 Monochamus titillator (southern pine sawyer) 214 Monochamus urossovi (fir sawyer) 214, 216 Phaenops californica 210 Phaenops gentilis (flat headed pine borer) 210 Tetropium castaneum 221–222 Tetropium cinnamopterum (eastern larch borer) 222 Tetropium gracilicorne 222–223 Tetropium velutinum (western larch borer) 222 Sirex juvencus 230 Sirex noctilio (sirex woodwasp) 228 Urocerus gigas 231 Sucking insects Cinara atlantica 237 Cinara maritimae 237 Cinara pinivora 237 Pineus boerneri 241–242 Pineus cembrae 258 Pineus cladogenous 241

385

Pineus coloradensis 241 Pineus ghanii 241 Pineus orientalis 258 Pineus pineoides 241 Pineus pini 241 Pineus pinifoliae 258 Pineus piniyunnanensis 241 Pineus simmondsi 241 Pineus wallichianae 241 Tip, shoot and regeneration insects Dioryctria raoi 293 Dioryctria zimmermani (Zimmerman pine moth) 293 Hylastes ater (black pine beetle) 280 Hylobius abietis (large pine weevil) 275–276 Hylobius albosparsus (white spotted weevil) 275 Hylobius pales (pales weevil) 275, 277 Hylobius piceus 275 Hylobius radicis (pine root collar weevil) 275 Hylobius rhizophagus (pine root tip weevil) 275 Hylurgus ligniperda (golden haired bark beetle) 281 Pissodes approximatus (northern pine weevil) 278 Pissodes nemorensis (deodar weevil) 278 Pissodes notatus (banded pine weevil) 278 Pissodes pini 278 Pissodes terminalis (lodgepole terminal weevil) 278 Pissodes validirostris 278, 300 Rhyacionia buoliana (European pine shoot moth) 288 Rhyacionia bushnelli (western pine tip moth) 288 Rhyacionia frustrana (Nantucket pine tip moth) 288, 290 Rhyacionia neomexicana (southwestern pine tip moth) 288 Rhyacionia rigidana (pitch pine tip moth) 288 Rhyacionia subtropica (subtropical pine tip moth) 288 Rhyacionia zozana 288 Tomicus minor 285 Tree reproductive structures Conophthorus ponderosae 301, 302

386

Host Index

Pinus (pine) (Continued ) Cydia toreuta (shortleaf pine coneworm) 306, 307 Dioryctria abietella 310, 311 Dioryctria abietivorella 310, 311 Dioryctria disclusa (webbing coneworm) 311 Dioryctria pinicolella 311 Dichocrocis punctiferalis (yellow peach borer) 309 Leptoglossus corculus (southern pine seed bug) 297 Leptoglossus occidentalis (western conifer seed bug) 296 Megastigmus atedius 304 Pissodes validirostriss (pine cone weevil) 278, 300 Wood in use Anobium punctatum (furniture beetle) 325 Buprestis aurulenta (golden buprestid) 326 Hylotrupes bajulus (old-house borer) 326 Pinus albicaulis (whitebark pine) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Bark and ambrosia beetles Dendroctonus ponderosae (mountain pine beetle) 181 Insects of wood in use Pinus aristata (Colorado bristlecone pine) Gall insects Pinyonia edulicola (piñon spindle gall midge) 267 Tip, shoot and regeneration insects Pityophthorus boycei 283 Tree reproductive structures Conophthorus ponderosae 302 Megastigmus pinus (fir seed chalcid) 304 Pinus arizonica Foliage feeders Neodiprion autumnalis 160–161 Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus brevicomis (western pine beetle) 175–177 Dendroctonus frontalis (southern pine beetle) 177–179 Tree reproductive structures Conophthorus ponderosae 302

Pinus armandi Foliage feeders Dendrolimus spectabilis 128 Bark and ambrosia beetles Dendroctonus armandi 177 Dendroctonus valens (red turpentine beetle) 184 Ips acuminatus 191 Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Pineus armandicola 258 Tip, shoot and regeneration insects Tomicus pilifer 285 Tomicus piniperda 285 Tomicus yunnanensis (Yunnan shoot borer) 285, 287 Pinus attenuata Tip, shoot and regeneration insects Pissodes radiatae (Monterrey pine weevil) 278 Tree reproductive structures Dioryctria auranticella (ponderosa pine coneworm) 311 Pinus ayacahuite Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Tip, shoot and regeneration insects Pityophthorus brevis 283 Tree reproductive structures Conophthorus ponderosae 302 Pinus banksiana (jack pine) Foliage feeders Choristoneura pinus pinus (jack pine budworm) 119 Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion nanulus nanulus (red pine sawfly) 161 Neodiprion swainei (Swain’s jack pine sawfly) 161 Bark and ambrosia beetles Dendroctonus murrayanae 177 Ips grandicollis (southern pine engraver) 192 Wood borers Sirex noctilio (sirex woodwasp) 228 Tip, shoot and regeneration insects Conophthorus banksianae (jack pine tip weevil) 301 Eucosma gloriola (eastern pine shoot borer) 288 Hylobius rhizophagus (pine root tip weevil) 275

Pissodes strobi (white pine weevil) 278 Pissodes terminalis (lodgepole terminal weevil) 278 Tomicus piniperda 285 Tree reproductive structures Cydia toreuta (eastern pine seedworm) 306, 307 Pinus brutia Foliage feeders Calomicrus apicalis 152 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Matsucoccus josephi (Israeli pine bast scale) 244 Tip, shoot and regeneration insects Hylurgus ligniperda (golden haired bark beetle) 281 Tomicus destruens 285 Pinus canariensis (Canary Island pine) Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Tip, shoot and regeneration insects Hylurgus ligniperda (golden haired bark beetle) 281 Tomicus destruens 285 Pinus caribaea Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Ips acuminatus 191 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Pineus boerneri 242 Tip, shoot and regeneration insects Dioryctria horneana 293 Dioryctria rubella (Tusam pitch moth) 293

Host Index Tree reproductive structures Dioryctria clarioralis (blister coneworm) 311 Dioryctria erythropasa 311 Pinus cembra (Swiss stone pine) Foliage feeders Zeiraphera diniana (gray larch bud moth) 123 Neodiprion pinetum 161 Bark and ambrosia beetles Ips acuminatus 191 Pinus cembroides (Mexican piñon) Sucking insects Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Pityophthorus barberi 283 Pityophthorus confertus 283 Pityophthorus deletus 283 Pityophthorus lecontei 283 Pityophthorus modicus 283 Pityophthorus schwartzi 283 Pityophthorus tuberculatus 283 Tree reproductive structures Conophthorus edulis 301 Eucosma bobana (piñon coneworm) 308 Pinus contorta (lodgepole pine) Foliage feeders Choristoneura lambertiana (uncertain status) 119 Choristoneura lambertiana ponderoasana 119 Choristoneura lambertiana subretiniana 119 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Coleotechnites milleri (lodgepole needle miner) 117–118 Coleotechnites starki 118 Coloradia pandora (pandora moth) 135–136 Neodiprion nanulus contortae 161 Zeiraphera diniana (gray larch bud moth) 123 Bark and ambrosia beetles Dendroctonus murrayanae 177 Dendroctonus ponderosa (mountain pine beetle) Wood borers Sirex noctilio (sirex woodwasp) 228

Tip, shoot and regeneration insects Eucosma sonomana (western pine shoot borer) 287 Pissodes radiatae (Monterrey pine weevil) 278 Pissodes terminalis (lodgepole terminal weevil) 278 Pityophthorus boycei 283 Rhyacionia buoliana (European pine shoot moth) 288 Tree reproductive structures Conophthorus ponderosae 302 Cydia toreuta (eastern pine seedworm) 306, 307 Dioryctria reniculelloides (spruce coneworm) 311 Eucosma recissoriana (lodgepole cone borer) 308 Pissodes validirostris (pine cone weevil) 300 Pinus cooperi Tree reproductive structures Conophthorus ponderosae 302 Pinus coulteri (Coulter pine) Foliage feeders Coloradia pandora (pandora moth) 135–136 Bark and ambrosia beetles Dendroctonus brevicomis (western pine beetle) 175–177 Orthotomicus erosus (Mediterranean pine engraver) 195 Pinus densata (Sikang pine) Tip, shoot and regeneration insects Tomicus yunnanensis (Yunnan shoot borer) 285, 287 Pinus densiflora (Japanese red pine) Foliage feeders Dendrolimus spectabilis 128 Wood borers Monochamus alternatus (Japanese pine sawyer) 215 Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Gall insects Thecodiplosis japonensis (pine needle gall midge) 270 Pinus discolor (border piñon) Tree reproductive structures Conophthorus edulis 301 Pinus douglasiana Tree reproductive structures Conophthorus conicolens 301

387

Conophthorus ponderosae 302 Dioryctria erythropasa 311 Pinus durangensis Foliage feeders Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus brevicomis (western pine beetle) 175–177 Dendroctonus frontalis (southern pine beetle) 177–179 Dendroctonus rhizophagus 177 Ips grandicollis (southern pine engraver) 192 Tree reproductive structures Conophthorus ponderosae 302 Dioryctria rossi 311 Pinus echinata (shortleaf pine) Foliage feeders Atta texana (town ant, Texas leaf cutting ant) 169–171 Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion pinetum 161 Neodiprion pratti pratti (Virginia pine sawfly) 161 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Ips avulsus 190 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Buprestis apricans (turpentine borer) 207 Sucking insects Nuculaspis californica (black pine leaf scale) 252 Oracella acuta (loblolly pine scale) 245 Tip, shoot and regeneration insects Hylobius pales (pales weevil) 275, 277 Rhyacionia frustrana (Nantucket pine tip moth) Tree reproductive structures Conophthorus echinatae 301 Cydia toreuta (shortleaf pine seedworm) 306, 307 Dioryctria amatella (southern pine coneworm) 310, 311

388

Host Index

Pinus echinata (Continued ) Dioryctria clarioralis (blister coneworm) 311 Dioryctria merkeli (loblolly pine coneworm) 311 Eucosma cocana (eastern pine coneworm) 308 Pinus edulis (piñon pine) Foliage feeders Coloradia pandora (pandora moth) 135–136 Neodiprion edulicolus (piñon sawfly) 161 Zadiprion rohweri 165 Bark and ambrosia beetles Ips confuses 190 Sucking insects Matsucoccus acalyptus (piñon needle scale) 243–244 Gall insects Pinyonia edulicola (piñon spindle gall midge) 267 Tip, shoot and regeneration insects Pityophthorus blandus 283 Pityophthorus brevis 283 Pityophthorus confertus 283 Pityophthorus deletus 283 Pityophthorus keeni 283 Pityophthorus lecontei 283 Pityophthorus modicus 283 Pityophthorus punctifrons 283 Pityophthorus schwartzi 283 Pityophthorus tuberculatus 283 Pityophthorus woodi 283 Tree reproductive structures Conophthorus edulis 301 Eucosma bobana (piñon coneworm) 308 Pinus eldarica Sucking insects Matsucoccus josephi (Israeli pine bast scale) 244 Pinus ellioti (slash pine) Foliage feeders Dendrolimus punctatus (pine caterpillar) 128–129 Bark and ambrosia beetles Ips avulsus 190 Ips calligraphus (six-spined engraver beetle) 191 Wood borers Sirex noctilio (sirex woodwasp) 228

Sucking insects Cinara cronartii 237 Oracella acuta (loblolly pine scale) 245 Pineus boerneri 242 Tip, shoot and regeneration insects Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia subtropica (subtropical pine tip moth) 288 Tree reproductive structures Cydia anaranjada (slash pine seedworm) 306 Cydia ingens (longleaf pine seedworm) 306 Dioryctria amatella (southern pine coneworm) 310, 311 Dioryctria clarioralis (blister coneworm) 311 Dioryctria merkeli (loblolly pine coneworm) 311 Pinus engelmannii Foliage feeders Neodiprion autumnalis 160–161 Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Dendroctonus rhizophagus 177 Wood borers Buprestis apricans (turpentine borer) 207 Tree reproductive structures Conophthorus apachecae 301 Cydia latisigna 306 Pinus estevezii Bark and ambrosia beetles Dendroctonus brevicomis (western pine beetle) 175–177 Pinus flexilis (limber pine) Foliage feeders Choristoneura lambertiana ponderosana 119 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Bark and ambrosia beetles Dendroctonus ponderosae (mountain pine beetle) 181 Tip, shoot and regeneration insects Pityophthorus deletus 283 Tree reproductive structures Conophthorus ponderosae 302 Pinus gerardiana Tree reproductive structures Dioryctria abietella 310, 311

Pinus greggi Tip, shoot and regeneration insects Pityophthorus schwartzi 283 Pinus griffithi Tip, shoot and regeneration insects Hylobius angustus 275 Tree reproductive structures Dioryctria abietella 310, 311 Pinus halapensis (Aleppo pine) Foliage feeders Anoxia matutinalis matutinalis 151 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Sucking insects Pineus boerneri 242 Matsucoccus josephi (Israeli pine bast scale) 244 Tip, shoot and regeneration insects Dioryctria sylvestrella (maritime pine borer) 292, 293 Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia buoliana (European pine shoot moth) 288 Tomicus destruens 285 Pinus hartwegii Tree reproductive structures Conophthorus ponderosae 302 Cydia montezuma 306 Megastigmus grandiosus (Montezuma pine seed chalcid) 304 Pinus insularis Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Tip, shoot and regeneration insects Tomicus brevipilosus 285 Pinus jeffreyi (Jeffrey pine) Foliage feeders Coloradia pandora (pandora moth) 135–136 Bark and ambrosia beetles Dendroctonus jeffreyi (Jeffrey pine beetle) 177 Wood borers Phaenops gentilis (flat headed pine borer) 210 Sirex noctilio (sirex woodwasp) 228

Host Index Sucking insects Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Eucosma sonomana (western pine shoot borer) 287 Tree reproductive structures Conophthorus ponderosae 302 Cydia piperana (ponderosa pine seed worm) 306 Eucosma ponderosa (western pine coneworm) 308 Pinus kesiya Foliage feeders Gilpinia leksawasdii 159 Gilpinia marshalli 159 Nesodiprion biremis 164 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Pineus boerneri 242 Tip, shoot and regeneration insects Dioryctria assamensis 293 Dioryctria rubella (Tusam pitch moth) 293 Tomicus yunnanensis (Yunnan shoot borer) 285, 287 Tree reproductive structures Dioryctria castanea 311 Pinus koraiensis Bark and ambrosia beetles Ips acuminatus 191 Ips subelongatus 193 Sucking insects Pineus cembrae 258 Tip, shoot and regeneration insects Tomicus brevipilosus 285 Tomicus pilifer 285 Tomicus piniperda 285 Tomicus puellus 285 Pinus lambertiana (sugar pine) Foliage feeders Choristoneura lambertiana lambertiana 119 Coloradia pandora (pandora moth) 135–136 Bark and ambrosia beetles Dendroctonus ponderosae (mountain pine beetle) 181 Wood borers Phaenops gentilis (flat headed pine borer) 210 Sucking insects

Nuculaspis californica (black pine leaf scale) 252 Tree reproductive structures Conophthorus ponderosae 302 Pinus lawsonii Tree reproductive structures Dioryctria erythropasa 311 Pinus leiophylla Foliage feeders Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus parallelocollis 177 Dendroctonus frontalis (southern pine beetle) 177–179 Tip, shoot and regeneration insects Tree reproductive structures Conophthorus conicolens 301 Conophthorus ponderosae 302 Dioryctria erythropasa 311 Pinus luchuensis Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Pinus lumholtzii Tip, shoot and regeneration insects Pityophthorus schwartzi 283 Pinus massoniana (Mason pine) Foliage feeders Dendrolimus punctatus (pine caterpillar) 128–129 Dendrolimus tabulaeformis 128 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Monochamus alternatus (Japanese pine sawyer) 215 Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Oracella acuta (loblolly pine scale) 245 Tip, shoot and regeneration insects Tomicus piniperda 285 Tree reproductive structures Dichocrocis punctiferalis (yellow peach borer) Pinus maximinoi Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Tree reproductive structures Dioryctria erythropasa 311 Pinus merkusii Foliage feeders

389

Dendrolimus punctatus (pine caterpillar) 128–129 Nesodiprion biremis 164 Bark and ambrosia beetles Ips acuminatus 191 Tip, shoot and regeneration insects Dioryctria rubella (Tusam pitch moth) 293 Pinus michoacana Foliage feeders Zadiprion falsus 164–165 Bark and ambrosia beetles Ips calligraphus (six-spined engraver beetle) 191 Tree reproductive structures Conophthorus michoacanae 301 Cydia latisigna 306 Dioryctria erythropasa 311 Pinus monophylla (single-leaf piñon) Foliage feeders Coloradia velda 136 Neodiprion edulicolus (piñon sawfly) 161 Zadiprion rohweri 165 Bark and ambrosia beetles Ips confusus 190 Sucking insects Matsucoccus acalyptus (piñon needle scale) 243–244 Tip, shoot and regeneration insects Pityophthorus blandus 283 Pityophthorus deletus 283 Pityophthorus keeni 283 Pityophthorus modicus 283 Pityophthorus punctifrons 283 Pityophthorus tuberculatus 283 Tree reproductive structures Conophthorus monophyllae 301 Eucosma bobana (piñon coneworm) 308 Pinus montezumae Bark and ambrosia beetles Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Tip, shoot and regeneration insects Pityophthorus schwartzi 283 Tree reproductive structures Conophthorus ponderosae 302 Cydia montezuma 306 Megastigmus grandiosus (Montezuma pine seed chalcid) 304

390

Host Index

Pinus monticola (western white pine) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Bark and ambrosia beetles Dendroctonus ponderosae (mountain pine beetle) 181 Wood borers Monochamus notatus (northeastern pine sawyer) 214 Sucking insects Pineus pinifoliae (pine leaf chermes) 258 Pineus strobi (pine bark adelgid) 242 Tip, shoot and regeneration insects Hylurgus ligniperda (golden haired bark beetle) 281 Tree reproductive structures Conophthorus ponderosae 302 Eucosma recissoriana (lodgepole cone borer) 308 Pinus mugo Bark and ambrosia beetles Ips acuminatus 191 Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Matsucoccus pini 244 Tip, shoot and regeneration insects Rhyacionia buoliana (European pine shoot moth) 288 Tomicus piniperda 285 Tree reproductive structures Dioryctria abietella 310, 311 Pinus muricata Tip, shoot and regeneration insects Pissodes radiatae (Monterrey pine weevil) 278 Rhyacionia buoliana (European pine shoot moth) 288 Pinus nigra (Austrian pine) Foliage feeders Calomicrus apicalis 152 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Bark and ambrosia beetles Ips acuminatus 191

Ips hauseri 190 Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Sirex juvencus 230 Tip, shoot and regeneration insects Dioryctria sylvestrella (maritime pine borer) 292, 293 Hylastes ater (black pine beetle) 280 Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia buoliana (European pine shoot moth) 288 Tomicus piniperda 285 Pinus occidentalis Bark and ambrosia beetles Ips calligraphus (six-spined engraver beetle) 191 Pinus oocarpa Foliage feeders Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Dendroctonus parallelocollis 177 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Tree reproductive structures Dioryctria erythropasa 311 Pinus palustris (longleaf pine) Foliage feeders Neodiprion lecontei (redheaded pine sawfly) 161–162 Bark and ambrosia beetles Ips avulsus 190 Ips grandicollis (southern pine engraver) 192 Wood borers Buprestis apricans (turpentine borer) 207 Sucking insects Oracella acuta (loblolly pine scale) 245 Tree reproductive structures Cydia anaranjada (slash pine seedworm) 306 Cydia ingens (longleaf pine seedworm) 306 Dioryctria amatella (southern pine coneworm) 310, 311 Dioryctria clarioralis (blister coneworm) 311

Dioryctria merkeli (loblolly pine coneworm) 311 Pinus parvifolia Tip, shoot and regeneration insects Tomicus brevipilosus 285 Pinus patula Foliage feeders Atta cephalotes 169 Glena bisulca 125, 127 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Pineus boerneri 242 Tip, shoot and regeneration insects Hylurgus ligniperda (golden haired bark beetle) 281 Tree reproductive structures Conophthorus mexicanus 301 Pinus peuce (Macedonian pine) Sucking insects Pineus strobi (pine bark adelgid) 242 Pinus pinaster (maritime pine) Foliage feeders Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Sirex noctilio (sirex woodwasp) 228 Sucking insects Matsucoccus feytaudi (maritime bast scale) 243–244 Tip, shoot and regeneration insects Dioryctria sylvestrella (maritime pine borer) 292, 293 Hylastes ater (black pine beetle) 280 Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia buoliana (European pine shoot moth) 288 Tomicus destruens 285 Tree reproductive structures Pissodes validirostris (pine cone weevil) 300

Host Index Pinus pinceana Tree reproductive structures Eucosma bobana (piñon coneworm) 308 Pinus pinea (stone pine) Foliage feeders Anoxia matutinalis matutinalis 151 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Tip, shoot and regeneration insects Dioryctria sylvestrella (maritime pine borer) 292, 293 Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia buoliana (European pine shoot moth) 288 Tomicus destruens 285 Tree reproductive structures Pissodes validirostris (pine cone weevil) 278, 300 Pinus ponderosa Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Coleotechnites ponderosae 118 Coloradia doris (Black Hills pandora moth) 136 Coloradia pandora (pandora moth) 135–136 Dasychira grisefacta (western pine tussock moth) 139–140 Neodiprion autumnalis 160–161 Neodiprion fulviceps 161 Neodiprion nanulus contortae 161 Zadiprion townsendi 165 Bark and ambrosia beetles Dendroctonus brevicomis (western pine beetle) 175–177 Dendroctonus ponderosae (mountain pine beetle) 181 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Wood borers Phaenops gentilis (flat headed pine borer) 210

Sirex juvencus californicus 230 Sirex noctilio (sirex woodwasp) 228 Sucking insects Cinara ponderosae 237 Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Eucosma sonomana (western pine shoot borer) 287 Hylastes ater (black pine beetle) 280 Pityophthorus boycei 283 Pityophthorus brevis 283 Pityophthorus deletus 283 Tree reproductive structures Conophthorus ponderosae 302 Cydia piperana (ponderosa pine seed worm) 306 Dioryctria auranticella (ponderosa pine coneworm) 311 Dioryctria rossi 311 Eucosma ponderosa (western pine coneworm) 308 Megastigmus albifrons (ponderosa pine seed chalcid) 304 Pinus pringeli Bark and ambrosia beetles Dendroctonus frontalis 177–179 Tree reproductive structures Conophthorus ponderosae 302 Pinus pseudostrobus Foliage feeders Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus vitei 177 Ips calligraphus (six-spined engraver beetle) 191 Tree reproductive structures Conophthorus ponderosae 302 Cydia montezuma 306 Pinus radiata (Monterrey pine) Foliage feeders Ormiscodes cinnamomea 137–138 Orgyia mixta 146 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Zadiprion falsus 164–165 Bark and ambrosia beetles Ips grandicollis (southern pine engraver) 192

391

Orthotomicus erosus (Mediterranean pine engraver) 195 Platypus granulosus 201 Wood borers Buprestis novemmaculata 208 Rhyephenes mallei 223 Sirex juvencus 230 Sirex noctilio (sirex woodwasp) 228 Urocerus gigas 231 Sucking insects Pineus boerneri 241–242 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Hylurgus ligniperda (golden haired bark beetle) 281 Pissodes radiatae (Monterrey pine weevil) 278 Rhyacionia buoliana (European pine shoot moth) 288 Rhyacionia frustrana (Nantucket pine tip moth) 288, 290 Tomicus piniperda 285 Tree reproductive structures Conophthorus radiatae 301 Wood in use Coptotermes acinaciformis 317 Porotermes adamsoni 316 Pinus resinosa (red pine) Foliage feeders Choristoneura pinus pinus (jack pine budworm) 119 Gilpinia frutetorum 159 Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion pinetum 161 Bark and ambrosia beetles Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Wood borers Sirex noctilio (sirex woodwasp) 228 Sucking insects Pineus boerneri 242 Matsucoccus matsumurae (Japanese pine bast scale) 244 Tip, shoot and regeneration insects Eucosma gloriola (eastern pine shoot borer) 288 Pissodes strobi (white pine weevil) 278

392

Host Index

Pinus resinosa (red pine) (Continued ) Rhyacionia buoliana (European pine shoot moth) 288 Tomicus piniperda 285 Tree reproductive structures Conophthorus resinosae 301 Cydia toreuta (eastern pine seedworm) 306, 307 Eucosma monitorana (red pine cone borer) 308 Pinus rigida (pitch pine) Foliage feeders Neodiprion pinetum 161 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Wood borers Buprestis apricans (turpentine borer) 207 Sucking insects Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Hylobius pales (pales weevil) 275, 277 Rhyacionia frustrana (Nantucket pine tip moth) Tree reproductive structures Eucosma cocana (shortleaf pine coneworm) 308 Pinus roxburghii Tip, shoot and regeneration insects Dioryctria raoi 293 Tree reproductive structures Dioryctria abietella 310, 311 Pinus rudis Tree reproductive structures Conophthorus ponderosae 302 Cydia montezuma 306 Megastigmus grandiosus (Montezuma pine seed chalcid) 304 Pinus sabiniana (digger pine) Sucking insects Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Pityophthorus modicus 283 Pinus sibirica (Siberian pine) Bark and ambrosia beetles Ips subelongatus 193

Wood borers Monochamus impluviatus 214 Pinus serotina (pond pine) Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Sucking insects Cinara cronartii 237 Pinus strobiformis Bark and ambrosia beetles Ips grandicollis (southern pine engraver) 192 Tip, shoot and regeneration insects Pityophthorus deletes 283 Tree reproductive structures Conophthorus ponderosae 302 Pinus strobus (eastern white pine) Foliage feeders Dendrolimus spectabilis 128 Neodiprion pinetum 161 Neodiprion swainei (Swain’s jack pine sawfly) 161 Bark and ambrosia beetles Ips calligraphus (six-spined engraver beetle) 191 Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Melanophila fulvoguttata (hemlock borer) 209 Monochamus notatus (northeastern pine sawyer) 214 Sirex noctilio (sirex woodwasp) 228 Sucking insects Cinara strobi 237 Pineus floccus (red spruce adelgid) 242 Pineus strobi (pine bark adelgid) 242 Gall insects Pineus pinifoliae (pine leaf chermes) Tip, shoot and regeneration insects Hylobius pales (pales weevil) 277 Hylurgus ligniperda (golden haired bark beetle) 281 Pissodes strobi (white pine weevil) 278 Tomicus piniperda 285 Tree reproductive structures Conophthorus coniperda (white pine cone beetle) 301, 302 Eucosma tocullionana (white pine cone borer) 308

Pinus sylvestris (Scotch pine) Foliage feeders Bupalus piniaria (pine looper) 125 Choristoneura lambertiana (uncertain status) 119 Dendrolimus pini 128 Gilpinia fructetorum 159 Gilpinia pallida 159 Gilpinia virens 159 Neodiprion sertifer (European pine sawfly) 161, 163–164 Neodiprion swainei (Swain’s jack pine sawfly) 161 Thaumetopoea pityocampa (pine processionary caterpillar) 133–135 Thaumetopoea wilkinsonii (eastern pine processionary caterpillar) 133–135 Zeiraphera diniana (gray larch bud moth) 123 Bark and ambrosia beetles Dendroctonus ponderosae (mountain pine beetle) 181 Ips acuminatus 191 Ips grandicollis (southern pine engraver) 192 Ips hauseri 190 Ips subelongatus 193 Orthotomicus erosus (Mediterranean pine engraver) 195 Wood borers Phaenops cyanea 210 Tetropium fuscum (brown spruce longhorn beetle) 222 Sirex noctilio (sirex woodwasp) 228 Sucking insects Cinara pini 237 Matsucoccus matsumurae (Japanese pine bast scale) 244 Matsucoccus pini 244 Gall insects Thecodiplosis brachyptera 271 Tip, shoot and regeneration insects Eucosma gloriola (eastern pine shoot borer) 288 Dioryctria abietella 310, 311 Hylastes ater (black pine beetle) 280 Hylurgus ligniperda (golden haired bark beetle) 281 Rhyacionia buoliana (European pine shoot moth) 288

Host Index Rhyacionia frustrana (Nantucket pine tip moth) 288, 290 Tomicus piniperda 285 Tree reproductive structures Dioryctria mutatella 311 Pissodes validirostris (pine cone weevil) 300 Pinus tabulaeformis Foliage feeders Dendrolimus spectabilis 128 Dendrolimus tabulaeformis 128 Neodiprion dailingensis 161 Bark and ambrosia beetles Dendroctonus valens (red turpentine beetle) 184 Orthotomicus erosus (Mediterranean pine engraver) 195 Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Tip, shoot and regeneration insects Tomicus pilifer 285 Pinus taeda (loblolly pine) Foliage feeders Atta texana (town ant, Texas leaf cutting ant) 169–171 Dendrolimus punctatus (pine caterpillar) 128–129 Dendrolimus spectabilis 128 Neodiprion lecontei (redheaded pine sawfly) 161–162 Neodiprion taedae linearis (loblolly pine sawfly) 160, 161 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Ips avulsus 190 Ips calligraphus (six-spined engraver beetle) 191 Ips grandicollis (southern pine engraver) 192 Wood borers Sirex noctilio (sirex woodwasp) 228 Sucking insects Cinara cronartii 237 Oracella acuta (loblolly pine scale) 245 Pineus boerneri 242 Tip, shoot and regeneration insects Hylobius pales (pales weevil) 277 Rhyacionia buoliana (European pine shoot moth) 288 Rhyacionia frustrana (Nantucket pine tip moth) 289

Tree reproductive structures Cydia anaranjada (slash pine seedworm) 306 Cydia ingens (longleaf pine seedworm) 306 Cydia toreuta (eastern pine seedworm) 306, 307 Dioryctria amatella (southern pine coneworm) 310, 311 Dioryctria clarioralis (blister coneworm) 311 Dioryctria merkeli (loblolly pine coneworm) 311 Eucosma cocana (shortleaf pine coneworm) 308 Pinus taiwanensis Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Pinus tecumani Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Pinus tenuifolia Bark and ambrosia beetles Dendroctonus vitei 177 Ips grandicollis (southern pine engraver) 192 Pinus teocote Foliage feeders Neodiprion autumnalis 160–161 Zadiprion falsus 164–165 Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Tip, shoot and regeneration insects Pityophthorus schwartzi 283 Tree reproductive structures Conophthorus teocote 301 Pinus thunbergiana Foliage feeders Dendrolimus spectabilis 128 Dendrolimus tabulaeformis 128 Sucking insects Matsucoccus matsumurae (Japanese pine bast scale) 244 Gall insects Thecodiplosis japonensis (pine needle gall midge) 270 Pinus virginiana (Virginia pine) Foliage feeders Neodiprion pratti pratti (Virginia pine sawfly) 161

393

Bark and ambrosia beetles Dendroctonus frontalis (southern pine beetle) 177–179 Ips grandicollis (southern pine engraver) 192 Sucking insects Oracella acuta (loblolly pine scale) 245 Tip, shoot and regeneration insects Rhyacionia frustrana (Nantucket pine tip moth) 288, 290 Tree reproductive structures Cydia toreuta (eastern pine seedworm) 306, 307 Dioryctria amatella (southern pine coneworm) 310, 311 Dioryctria merkeli (loblolly pine coneworm) 311 Eucosma cocana (shortleaf pine coneworm) 308 Eucosma monitorana (red pine cone borer) 308 Pinus washoensis Tree reproductive structures Conophthorus ponderosae 302 Pinus wallichiana Tree reproductive structures Dioryctria abietella 310, 311 Pinus yunnanensis Bark and ambrosia beetles Orthotomicus erosus (Mediterranean pine engraver) 195 Foliage feeders Neodiprion xiangyunicus 161 Tip, shoot and regeneration insects Pissodes yunnanensis (Yunnan pine weevil) 278, 279 Tomicus brevipilosus 285 Tomicus yunnanensis (Yunnan shoot borer) 285, 287 Pistacia Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Tree reproductive structures Megastigmus 303 Platanus Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Anoplophora chinensis (citrus longhorn beetle) 212

394

Host Index

Platanus occidentalis (American sycamore) Foliage feeders Thyridopteryx ephemeraeformis (bagworm) 113–114 Platycladius armatus Bark and ambrosia beetles Phloeosinus bicolor 185 Plumeria (frangipani) Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Populus (poplar, cottonwood) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Choristoneura diversana 119, 120 Clania variegata (giant bagworm) 114 Euproctis chrysorrhoea (browntail moth) 140 Hyphantria cunea (fall webworm) 148 Leucoma salicis (satin moth) 142–151 Lymantria dispar (gypsy moth) 142–144 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma disstria (forest tent caterpillar) 131–132 Malacosoma incurvum (southwestern tent caterpillar) 131 Malacosoma nuestria 131–132 Operophthera bruceata (Bruce spanworm) 125 Operophthera brumata (winter moth) 125 Tortrix viridana (green oak tortrix) 123 Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Agrilus granulatus 204 Agrilus liragus (bronze poplar borer) 204 Anoplophora chinensis (citrus longhorn beetle) 212 Anoplophora glabripennis (Asian longhorn beetle) 210 Cossus cossus (goat moth) 224

Prionoxystus robinae (carpenterworm) 225 Saperda alberti 220 Saperda calcarata (poplar borer) 219–220 Saperda carcharias (large poplar longhorn beetle) 220 Saperda concolor 259 Saperda inorata 259 Saperda perforata 220 Saperda populnea (small poplar borer) 220, 258 Tremex fuscicornis 227 Urocerus gigas 231 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Phloeomyzus passerinii (woolly poplar aphid) 238 Gall insects Hexomyza schineri (poplar twig gall fly) 271 Mordwilkoja vagabunda (poplar vagabond gall) 255 Saperda concolor 259 Saperda inornata 259 Populus caspica Bark and ambrosia beetles Platypus cylindrus (oak pinhole borer) 201 Populus deltoides (eastern cottonwood) Wood borers Saperda calcarata (poplar borer) 219 Gall insects Mordwilkoja vagabunda (poplar vagabond gall) 255 Populus x euroamericana Sucking insects Phloeomyzus passerinii (woolly poplar aphid) 238 Populus grandidentata (bigtooth aspen) Foliage feeders Choristoneura conflictana (large aspen tortrix) 119 Wood borers Saperda calcarata (poplar borer) 219 Populus nigra Sucking insects Phloeomyzus passerinii (woolly poplar aphid) 238 Gall insects Mordwilkoja vagabunda (poplar vagabond gall) 255

Saperda populnea (small poplar borer) 258 Populus tremula Wood borers Agrilus viridis 205 Gall insects Saperda populnea (small poplar borer) 258 Populus tremuloides (quaking aspen) Foliage feeders Choristoneura conflictana (large aspen tortrix) 119 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma disstria (forest tent caterpillar) 131–132 Phyllocnistis populiella (aspen leaf miner) 116–117 Wood borers Saperda calcarata (poplar borer) 219 Gall insects Hexomyza schineri (poplar twig gall fly) 271 Mordwilkoja vagabunda (poplar vagabond gall) 255 Prosopis Wood borers Megacyllene antennata 217 Protium Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Prunus (cherry, plum) Foliage feeders Choristoneura diversana 119, 120 Diapheromera femorata 149–150 Lymantria monacha (nun moth) 144–145 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma americanum (eastern tent caterpillar) 131 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma disstria (forest tent caterpillar) 131–132 Malacosoma incurvum (southwestern tent caterpillar) 131 Malacosoma nuestria 131–132 Nematus oligospilus 167 Operophthera bruceata (Bruce spanworm) 125 Operophthera brumata (winter moth) 125

Host Index Bark and ambrosia beetles Scolytus schevyrewi 186, 187 Scolytus rugulosus (shothole borer) 186 Platypus quercivorus 202 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Anoplophora chinensis (citrus longhorn borer) 212 Tremex fuscicornis 227 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Prunus armeniaca Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Prunus padus Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Prunus persica Bark and ambrosia beetles Megaplatypus mutatus 200 Scolytus schevyrewi (banded elm bark beetle) 187 Prunus pseudocerasus Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Prunus salicina Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Prunus yedoensis Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Pseudotsuga Bark and ambrosia beetles Trypodendron lineatum (striped ambrosia beetle) 196 Tip, shoot and regeneration insects Hylobius pales (pales weevil) 277 Wood in use Buprestis aurulenta (golden buprestid) 326 Pseudotsuga flahaulti Bark and ambrosia beetles Dendroctonus pseudotsugae (Douglasfir beetle) 182 Pseudotsuga macrocarpa (bigcone Douglas-fir) Foliage feeders

Choristoneura carana carana 119 Bark and ambrosia beetles Dendroctonus pseudotsugae (Douglasfir beetle) 177, 182 Wood borers Phaenops drummondi (flat headed fir borer) 210 Tree reproductive structures Megastigmus spermatrophus (Douglas-fir seed chalcid) 304 Pseudotsuga menziesii (Douglas-fir) Foliage feeders Bupalus piniaria (pine looper) 125 Choristoneura carana californicum 119 Choristoneura occidentalis (western spruce budworm) 119, 122–123 Nepytia freemani (false hemlock looper) 125 Orgyia antiqua (rusty tussock moth) 145–146 Orgyia pseudotsugata (Douglas-fir tussock moth) 146–147 Bark and ambrosia beetles Dendroctonus pseudotsugae (Douglasfir beetle) 177, 182 Orthotomicus erosus (Mediterranean pine engraver) 195 Scolytus ventralis (fir engraver beetle) 186, 189 Scolytus unispinosus (Douglas-fir engraver) 186 Wood borers Monochamus clamator 214 Monochamus scutellatus (white spotted sawyer) 214, 216 Tetropium velutinum (western larch borer) 222 Sucking insects Adelges cooleyi (Cooley spruce gall adelgid) 256 Cinara pseudotsugae 237 Nuculaspis californica (black pine leaf scale) 252 Tip, shoot and regeneration insects Dioryctria zimmermani (Zimmerman pine moth) 293 Hylastes ater (black pine beetle) 280 Hylobius abietis (large pine weevil) 275–276 Hylobius piceus 275 Tomicus piniperda 285

395

Tree reproductive structures Dioryctria abietella 310, 311 Dioryctria abietivorella 310, 311 Dioryctria pseudotsugella 311 Dioryctria reniculelloides (spruce coneworm) 311 Contarinia oregonensis (Douglas-fir cone gall midge) 312 Contarinia washingtoniensis (Douglas-fir cone scale midge) 312 Leptoglossus occidentalis (western conifer seed bug) 296 Megastigmus spermatrophus (Douglas-fir seed chalcid) 304 Wood in use Anobium punctatum (furniture beetle) 325 Psidium cattelianum (strawberry guava) Foliage feeders Sarsina violascens 147–148 Psidium guajava (guava) Foliage feeders Sarsina violascens 147–148 Pterocarya rhoifolia (Japanese wingnut) Wood borers Agrilus planipennis (emerald ash borer) 206–207 Purshia Foliage feeders Malacosoma californicum (western tent caterpillar) 130–131 Purshia tridentata (bitterbrush) Foliage feeders Malacosoma californicum (western tent caterpillar) 130–131 Orgyia cana 146 Pyrus Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma nuestria 131–132 Bark and ambrosia beetles Scolytus rugulosus (shothole borer) 186 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Cossus cossus (goat moth) 224 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251

396

Host Index

Pyrus communis Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Anoplophora chinensis (citrus longhorn borer) 212 Q Quercus (oak) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Choristoneura diversana 119, 120 Diapheromera femorata 149–150 Erannis defoliaria (mottled umber moth) 125 Erannis tiliaria (linden looper) 124, 125 Euproctis chrysorrhoea (browntail moth) 140 Euproctis kargalika (Turkistan browntail moth) 140–141 Lymantria dispar (gypsy moth) 142–144 Lymantria monacha (nun moth) 144–145 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma constrictum (Pacific tent caterpillar) 131 Malacosoma nuestria 131–132 Malacosoma tigris 131 Operophthera bruceata (bruce spanworm) 125 Operophthera brumata (winter moth) 125 Orgyia thyellina 146 Thaumetopoea processionea (oak processionary caterpillar) 135 Thyridopteryx ephemeraeformis (bagworm) 113–114 Bark and ambrosia beetles Megaplatypus mutatus 200 Platypus cylindrus (oak pinhole borer) 201 Wood borers Agrilus angelicus 204 Agrilus bilineatus (twolined chestnut borer) 204, 205 Cossus cossus (goat moth) 224 Megacyllene caryae (painted hickory borer) 217

Monochamus guttatus 214 Prionoxystus robinae (carpenterworm) 225 Tremex fuscicornis 227 Zeuzera pyrina (leopard moth) 227 Sucking insects Allokermes kingii (northern red oak kermes) 249 Gall insects Amphibolips confluenta (oak apple) 262 Biorhiza pallida 263 Cynips gallae tinctoria (Aleppo gall) 263 Cynips divisa 264 Cynips longiventris 264 Tree reproductive structures Curculio elephas 298, 299 Curculio davidi 298 Curculio occidentalis (filbert weevil) 298 Curculio proboscidens 298 Curculio sulcatulus 298 Mechoris cumulatus 299 Wood in use Lyctus brunneus (brown lyctus beetle) 323 Lyctus planicollis (southern lyctus beetle) 324 Quercus acuta Bark and ambrosia beetles Platypus quercivorus 202 Quercus acutissima Bark and ambrosia beetles Platypus quercivorus 202 Quercus aegilops Gall insects Cynips gallae tinctoria (Aleppo Gall) 263 Quercus agrifolia (coast live oak) Wood borers Agrilus coxalis (goldenspotted oak borer) 204, 205 Prionoxystus robinae (carpenterworm) 225 Gall insects Callirhytis quercuspomiformis 263 Quercus alba (white oak) Foliage feeders Cameraria cincinnatiella 116 Cameraria hamadryadella 116 Quercus alnifolia Wood borers Agrilus roscidus 204

Quercus bicolor (swamp white oak) Gall insects Disholcaspis quercusmamma (rough oak bullet gall wasp) 264 Quercus cerris Foliage feeders Thaumetopoea processionea (oak processionary caterpillar) 135 Wood borers Agrilus pannonicus 206 Quercus chrysolepis (canyon live oak) Foliage feeders Orgyia cana 146 Wood borers Agrilus coxalis (goldenspotted oak borer) 204, 205 Gall insects Trichoteras vaccinifoliae 263 Quercus coccifera (kermes oak) Sucking insects Kermococcus vermilis 249 Quercus coccinea (scarlet oak) Gall insects Amphibolips confluenta (oak apple) 262 Quercus dalechampii Bark and ambrosia beetles Scolytus intricatus 186 Quercus dilatata Foliage feeders Malacosoma indica (Indian tent caterpillar) 131 Quercus fusiforme (Texas live oak) Gall insects Disholcaspis cinerosa 264 Quercus garryana (Garry oak) Gall insects Andricus californicus 263 Quercus gilva Bark and ambrosia beetles Platypus quercivorus 202 Quercus glauca Bark and ambrosia beetles Platypus quercivorus 202 Quercus ilex (Holm oak) Wood borers Agrilus pannonicus 206 Sucking insects Kermococcus vermilis 249 Quercus incana Foliage feeders Malacosoma indica (Indian tent caterpillar) 131

Host Index Quercus infectoria Gall insects Cynips gallae tinctoria (Aleppo gall) 263 Cynips incana 263 Quercus kelloggii (California black oak) Foliage feeders Orgyia cana 146 Wood borers Agrilus coxalis (goldenspotted oak borer) 205 Quercus macrocarpa (burr oak) Foliage feeders Cameraria cincinnatiella 116 Cameraria hamadryadella 116 Gall insects Disholcaspis quercusmamma (rough oak bullet gall wasp) 264 Quercus mongolica Bark and ambrosia beetles Platypus quercivorus 202 Quercus myrsinifolia Bark and ambrosia beetles Platypus quercivorus 202 Quercus pendiculata Gall insects Cynips gallae tinctoria (Aleppo gall) 263 Quercus petraea (sessile oak) Foliage feeders Thaumetopoea processionea (oak processionary caterpillar) 135 Bark and ambrosia beetles Scolytus intricatus 186 Wood borers Agrilus pannonicus 206 Gall insects Cynips quercusfolii (oak bud gall wasp, cherry gall) 264 Tree reproductive structures Curculio nucum 298 Quercus phillyraeoides (ubame oak) Bark and ambrosia beetles Platypus quercivorus 202 Quercus pubescens Wood borers Agrilus pannonicus 206 Quercus robur (English oak) Foliage feeders Thaumetopoea processionea (oak processionary caterpillar) 135

Bark and ambrosia beetles Scolytus intricatus 186 Gall insects Cynips quercusfolii (oak bud gall wasp, cherry gall) 264 Tree reproductive structures Curculio elephas (chestnut weevil) 298, 299 Curculio nucum 298 Quercus rubra (northern red oak) Sucking insects Allokermes kingii (northern red oak kermes) 249 Gall insects Amphibolips confluenta (oak apple) 262 Quercus salcina Bark and ambrosia beetles Platypus quercivorus 202 Quercus serrata Bark and ambrosia beetles Platypus quercivorus 202 Quercus sessilifolia Bark and ambrosia beetles Platypus quercivorus 202 Quercus suber (cork oak) Wood borers Agrilus pannonicus 206 Tree reproductive structures Curculio elephas (chestnut weevil) 298–299 Quercus velutina (black oak) Sucking insects Allokermes kingii (northern red oak kermes) 249 Gall insects Amphibolips confluenta (oak apple) 262 Quercus virginiana (live oak) Gall insects Disholcaspis cinerosa 264 Tree reproductive structures Curculio fulvus 298 Quillaja saponora (soapberry) Wood borers Rhyephenes mallei 223 Rhyephenes humeralis 223 R Ribies (currant, gooseberry) Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141

397

Rhabdadenia biflora Foliage feeders Pseudosphinx tetrio (frangipani hawk moth) 139 Rhamnus Foliage feeders Choristoneura diversana 119, 120 Rhamnus californica Foliage feeders Orgyia cana 146 Rhododendron Foliage feeders Choristoneura diversana 119, 120 Robinia (locust) Foliage feeders Lymantria obfuscata (Indian gypsy moth) 143 Robinia neomexicana (New Mexico locust) Wood borers Megacyllene robinae 217 Robinia pseudoacacia (black locust) Foliage feeders Odontata dorsalis (locust leaf miner) 153–154 Thyridopteryx ephemeraeformis (bagworm) 113–114 Bark and ambrosia beetles Megaplatypus mutatus 200 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Megacyllene robinae (locust borer) 217 Tremex fuscicornis 227 Rosa Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Lymantria obfuscata (Indian gypsy moth) 143 Malacosoma nuestria 131–132 Tree reproductive structures Megastigmus 303 Rubus Foliage feeders Euproctis kargalika (Turkistan browntail moth) 140–141 Malacosoma nuestria 131–132 S Sabina chinensis Bark and ambrosia beetles Phloeosinus bicolor 185

398

Host Index

Sacoglottis procera Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Salix (willow) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Choristoneura diversana 119, 120 Euproctis chrysorrhoea (browntail moth) 140 Euproctis kargalika (Turkistan browntail moth) 140–141 Hyphantria cunea (fall webworm) 148 Leucoma salicis (satin moth) 141–142 Lymantria monacha (nun moth) 144–145 Malacosoma californicum (western tent caterpillar) 130–131 Malacosoma incurvum (southwestern tent caterpillar) 131 Malacosoma nuestria 131–132 Nematus oligospilus 167 Operophthera bruceata (bruce spanworm) 125 Operophthera brumata (winter moth) 125 Orgyia antiqua (rusty tussock moth) 145–146 Orgyia cana 146 Thyridopteryx ephemeraeformis (bagworm) 113–114 Bark and ambrosia beetles Megaplatypus mutatus 200 Scolytus schevyrewi (banded elm bark beetle) 187 Wood borers Cossus cossus (goat moth) 224 Monochamus guttatus 214 Saperda alberti 220 Saperda balsamifera 220 Saperda carcharias (large poplar longhorn beetle) 220 Saperda perforata 220 Saperda populnea (small poplar borer) 220, 258 Saperda similis 220 Tremex fuscicornis 227 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Gall insects Euura spp. 259 Euura mucronata 259

Hexomyza schineri (poplar twig gall fly) 271 Rhabdophaga rosaria 267 Rhabdophaga strobiloides (willow pine cone gall) 267 Pontania proxima (willow red gall sawfly) 260 Salix babylonica (weeping willow) Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Salix japonica Gall insects Euura shibayanagii 259 Santalum album (sandalwood) Sucking insects Aonidiella orientalis (oriental scale) 249 Sassafras albidum (sassafras) Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Schinus Tree reproductive structures Megastigmus 303 Schinus latifolius (molle) Foliage feeders Ormiscodes cinnamomea 137–138 Schleichera oleosa Sucking insects Kerria lacca (lac insect) 248–249 Schleichera trijuga Sucking insects Aonidiella orientalis (oriental scale) 249 Schleichera trijuga Sucking insects Kerria lacca (lac insect) 248–249 Shorea robusta (sal) Bark and ambrosia beetles Xyleborus glabratus (redbay ambrosia beetle) 198–199 Shorea talura Sucking insects Kerria lacca (lac insect) 248–249 Sorbus Foliage feeders Malacosoma nuestria 131–132 Wood borers Saperda candida (round headed appletree borer) 220 Tree reproductive structures Megastigmus 303 Swietenia macrophylla (big leaf mahogany)

Foliage feeders Atta cephalotes 169 Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Tip, shoot and regeneration insects Hypsipyla grandella 291 Hypsipyla robusta 292 Swietenia mahogani Sucking insects Aonidiella orientalis (oriental scale) 249 Tip, shoot and regeneration insects Hypsipyla grandella 291 Hypsipyla robusta 292 Syringa vulgaris (lilac) Foliage feeders Malacosoma nuestria 131–132 Stereonychus fraxini (ash weevil) 155 Wood borers Podesesia syringae (lilac/ash borer) 226 Sucking insects Lepidosaphes ulmi (oystershell scale) 250–251 Syzygium cuminii Sucking insects Aonidiella orientalis (oriental scale) 249 T Tamarix (salt cedar) Foliage feeders Diorhabda carinata 153 Diorhabda carinulata 153 Diorhabda elongata 153 Diorhabda meridionalis 153 Diorhabda sublineata 153 Tamarindus indica (tamarind) Sucking insects Aonidiella orientalis (oriental scale) 249 Taxodium distichum (bald cypress) Foliage feeders Coleotechnites apicitripunctella 118 Thyridopteryx ephemeraeformis (bagworm) 113–114 Gall insects Taxodiomyia cupressiananassa (cypress twig gall midge) 267–268 Tree reproductive structures Dioryctria amatella (southern pine coneworm) 310, 311

Host Index Dioryctria pygmaella (baldcypress coneworm) 311 Tectona grandis (teak) Foliage feeders Atta cephalotes 169 Wood borers Alcterogystia cadambae (teak carpenterworm) 224 Xyleutes ceramica (teak beehole borer) 225 Sucking insects Maconellicoccus hirsutus (pink hibiscus mealybug) 245 Terminalia bellerica Wood borers Aletergystia cadambae (teak carpenterworm) 224 Terminalia ivoriensis Tip, shoot and regeneration insects Apate monachus 273 Tetraclinis articulata Sucking insects Cinara cupressivora (cypress aphid) 237 Tree reproductive structures Pseudococcyx tessulatana 309 Theobroma cacao (cacao) Bark and ambrosia beetles Xyleborus ferrugineus 197–198 Thuja Bark and ambrosia beetles Phloeosinus armatus 185 Phloeosinus bicolor 185 Trypodendron lineatum (striped ambrosia beetle) 196 Sucking insects Cinara cupressivora (cypress aphid) 237 Cinara thujafilina 237 Tip, shoot and regeneration insects Hylastes ater (black pine beetle) 280 Hylobius pales (pales weevil) 275, 277 Thuja occidentalis (northern white cedar) Foliage feeders Coleotechnites apicitripunctella 118 Thyridopteryx ephemeraeformiss (bagworm) 113–114 Thuja orientalis Tree reproductive structures Pseudococcyx tessulatana 309 Tilia (basswood, linden) Foliage feeders Erannis tiliaria (linden looper) 124, 125

Euproctis chrysorrhoea (browntail moth) 140 Diapheromera femorata 149–150 Lymantria dispar (gypsy moth) 142–144 Thyridopteryx ephemeraeformis (bagworm) 113–114 Operophthera brumata (winter moth) 125 Bark and ambrosia beetles Megaplatypus mutatus 200 Toona autralis Tip, shoot and regeneration insects Hypsipyla robusta 292 Toona cilicata Tip, shoot and regeneration insects Hypsipyla robusta 292 Toona serrata Tip, shoot and regeneration insects Hypsipyla robusta 292 Tsuga (hemlock) Foliage feeders Dasychira grisefacta (western pine tussock moth) 139–140 Dendrolimus sibiricus (Siberian silk moth) 128–130 Bark and ambrosia beetles Trypodendron lineatum (striped ambrosia beetle) 196 Wood borers Monochamus grandis 214 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tip, shoot and regeneration insects Hylobius pales (pales weevil) 277 Tsuga canadensis (eastern hemlock) Foliage feeders Coleotechnites apicitripunctella 118 Lambdina fiscellaria fiscellaria (hemlock looper) 125 Wood borers Melanophila fulvoguttata (hemlock borer) 209 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tree reproductive structures Eucosma tocullionana (white pine cone borer) 308 Tsuga caroliniensis (Carolina hemlock) Sucking insects Adelges tsugae (hemlock woolly adelgid) 239

399

Tsuga chinensis Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tsuga diversifolia Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tsuga dumosa Sucking insects Adelges tsugae 239 Tsuga mertensiana (mountain hemlock) Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tsuga heterophylla (western hemlock) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Lambdina fiscellaria lugubrosa (western hemlock looper) 125 Neodiprion tsugae (hemlock sawfly) 161 Orgyia antiqua (rusty tussock moth) 145–146 Wood borers Phaenops drummondi (flat headed fir borer) 210 Tetropium velutinum (western larch borer) 222 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 Tree reproductive structures Dioryctria reniculelloides (spruce coneworm) 311 Wood in use Anobium punctatum (furniture beetle) 325 Tsuga mertensiana (mountain hemlock) Foliage feeders Choristoneura occidentalis (western spruce budworm) 119, 122–123 Coleotechnites milleri (lodgepole needle miner) 117–118 Bark and ambrosia beetles Scolytus ventralis 186, 189 Sucking insects Adelges tsugae (hemlock woolly adelgid) 239

400

Host Index

Tsuga sieboldii Sucking insects Adelges tsugae (hemlock woolly adelgid) 239 U Ulmus (elm) Foliage feeders Alsophila pometaria (fall cankerworm) 125 Diapheromera femorata 149–150 Choristoneura diversana 119, 120 Erannis defoliara (mottled umber moth) 125 Erannis tiliaria (linden looper) 124, 125 Euproctis kargalika (Turkistan browntail moth) 140–141 Fenusa ulmi (elm leaf miner) 165–167 Hyphantria cunea (fall webworm) 148 Malacosoma nuestria 131–132 Operopthera brummata (winter moth) 125 Pyrrhalta luteola (elm leaf beetle) 154 Thyridopteryx ephemeraeformis (bagworm) 113–114 Bark and ambrosia beetles Megaplatypus mutatus 200 Scolytus multistriatus (smaller European elm bark beetle) 186, 187 Scolytus ratzeburgi 186 Scolytus scolytus (European elm bark beetle) 186 Scolytus schevyrewi (banded elm bark beetle) 186, 187 Wood borers Anoplophora glabripennis (Asian longhorn beetle) 210 Cossus cossus (goat moth) 224 Monochamus guttatus 214

Prionoxytus robinae (carpenterworm) 225 Saperda octomaculata 220 Saperda tridentata (elm borer) 220 Tremex fuscicornis 227 Zeuzera pyrina (leopard moth) 226 Sucking insects Gossyparia spuria (European elm scale) 246 Lepidosaphes ulmi (oystershell scale) 250–251 Wood in use Lyctus brunneus (brown lyctus beetle) 323 Ulmus americana (American elm) Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus carpinifolia Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus davidana (David elm, Japanese elm) Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Wood borers Agrilus planipennis (emerald ash borer) 206–207 Ulmus laevis Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus macrocarpa Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus propinqua Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus pumila Bark and ambrosia beetles

Scolytus schevyrewi (banded elm bark beetle) 187 Ulmus thomasi (rock elm) Bark and ambrosia beetles Scolytus schevyrewi (banded elm bark beetle) 187 V Vaccinium Foliage feeders Operophthera brumata (winter moth) 125 Orgyia antiqua (rusty tussock moth) 145–146 W Widdringtonia Sucking insects Cinara cupressivora (cypress aphid) 237 Cinara thujafilina 237 Z Zelkova Wood borers Tremex fuscicornis 227 Zelkova serrata Bark and ambrosia beetles Scolytus multistriatus (smaller European elm bark beetle) 186,

addstr 350, "|%2e [PRINT] \n" wdb 350 187 Ziziphus mauritania Sucking insects Aonidiella orientalis (oriental scale) 249 Kerria lacca (lac insect) 248–249 Ziziphus jujube Wood borers Anoplophora chinensis (citrus longhorn beetle) 212

Plate 1 Foliage discoloration of horse chestnut, Aesculus hippocastanum, caused by the leaf miner, Cameraria ohridella, in Garmisch, Germany. This insect, whose origin is unknown, has become invasive and damaging to a highly favored shade tree across much of Europe. Cited on page 42.

Plate 2 An adult monarch butterfly, Danaus plexippus, rests on a plant as it migrates to spend the winter in groves of sacred fir, Abies religiosa, in central Mexico where they are a tourist attraction and subject of curiosity (near Linville Gorge, North Carolina, USA). Cited on page 44. Forest Entomology: A Global Perspective, First Edition. William M. Ciesla. Ó 2011 William M. Ciesla. Published 2011 by Blackwell Publishing Ltd.

Plate 3 Plantations of Cupressus lusitanica damaged by cypress aphid, Cinara cupressivora (near Eldoret, Kenya). Cited on page 62.

Plate 4 Defoliation of sacred fir or oyamel, Abies religiousa by the geometrid, Evita hyalinaria blandaria (San Felipe del Progreso, Mexico State, Mexico). Cited on page 62.

Plate 5 Mortality of lodgepole pine, Pinus contorta, caused by mountain pine beetle, Dendroctonus ponderosa (Rocky Mountain National Park, Colorado, USA). Cited on page 62.

Plate 6 Scattered tree mortality in a loblolly pine, Pinus taeda, plantation caused by the wood wasp, Sirex noctilio (Parana State, Brazil). Cited on page 62.

Plate 7 Color infrared image of a group of pines killed by southern pine beetle, Dendroctonus frontalis. Yellow crowns are dying pines (faders), red-brown crowns are pines with green crowns and bright red crowns are broadleaf trees (Oak Ridge, Tennessee, USA). Cited on page 66.

Plate 8 Color infrared image taken from a NASA ER-2 aircraft of a mixed broadleaf forest that had received an aerial spray to protect it from defoliation by gypsy moth, Lymantria dispar. Gray bands are areas of defoliation due to inadequate coverage by the spray aircraft (western Maryland, USA). Cited on page 68.

Plate 9 A colony of nymphs and adults of the boxelder bug, Boisea trivittata (Homoptera: Rhopalidae) (Fort Collins, Colorado, USA). Cited on page 98.

Plate 10 Adult of the metallic wood borer, Buprestis rufipes (Coleoptera: Buprestidae) (Central Louisiana, USA). Cited on page 104.

Plate 11 Adult longhorn beetle, Adesmus hemisphilus (Coleoptera: Cerambycidae) (Parque Nacional Foz do Igua¸cu, Brazil). Cited on page 105.

Plate 12 Mature larva of the geometrid Meris alticola (Lepidoptera: Geometridae) (North Park, Colorado, USA). Cited on page 109.

Plate 13 Adult hawk moth, Hyles lineata (Lepidoptera: Sphingidae) in flight (Fort Collins, Colorado, USA). Cited on page 110.

Plate 14 Larva of the frangipani sphinx, Pseudosphinx tetrio (Lepidoptera: Sphingidae) (Island of Tobago, Trinidad and Tobago). Cited on page 110.

Plate 15 Mature larva of western spruce budworm, Choristoneura occidentalis (Culebra Range, Colorado, USA). Cited on page 122.

Plate 16 Pupa of western spruce budworm, Choristoneura occidentalis. (Culebra Range, Colorado, USA). Cited on page 122.

Plate 17 Adult western spruce budworm, Choristoneura occidentalis (Wet Mountains, Colorado, USA). Cited on page 122.

Plate 18 Defoliation of Abies concolor by western spruce budworm. (North La Veta Pass, Colorado, USA). Cited on page 122.

Plate 19 Pine caterpillar, Dendrolimus pini, mature larva (Lithuania). Cited on page 128.

Plate 20 Pine caterpillar, Dendrolimus punctatus, mature larva (near Vinh, Vietnam). Cited on page 129.

Plate 21 Colony of the western tent caterpillar, Malacosoma californicum (Culebra Range, Colorado, USA). Cited on page 130.

Plate 22 Western tent caterpillar, female adult and egg mass (Photo by J. E. Dewey, USDA Forest Service). Cited on page 130.

Plate 23 Aerial view of defoliation of Populus tremuloides by western tent caterpillar (Sangre de Cristo Range, Colorado, USA). Cited on page 130.

Plate 24 Colony of mature larvae of the pine processionary caterpillar, Thaumetopoea wilkinsonii (northern Cyprus). Cited on page 134.

Plate 26 Gypsy moth, Lymantria dispar, female adults and egg-masses (Dover, Delaware, USA). Cited on page 142. Plate 25 Gypsy moth, Lymantria dispar, mature larva (near Bucharest, Romania). Cited on page 142.

Plate 27 Heavy defoliation of a mixed oak forest by gypsy moth (South Mountain, Pennsylvania, USA). Cited on page 142.

Plate 28 Adult male Douglas-fir tussock moth, Orgyia pseudotsugata. Cited on page 147.

Plate 29 Mature larva of Douglas-fir tussock moth, Orgyia pseudotsugata (near Aspen Park, Colorado, USA). Cited on page 147.

Plate 30 Larva of fall webworm, Hyphantria cunea, on narrowleaf cottonwood, Populus angustifolia (near Fort Collins, Colorado, USA). Cited on page 148.

Plate 31 Larvae of a diorhabda beetle, Diorhabda sp., feeding on foliage of salt cedar (near Moab, Utah, USA). Cited on page 152.

Plate 32 Heavy defoliation of salt cedar by a diorhabda beetle, Diorhabda sp. (near Moab, Utah, USA). Cited on page 152.

Plate 33 Larva of elm leaf beetle, Pyrrhalta luteola (Taos, New Mexico, USA). Cited on page 154.

Plate 34 Larva of elm leaf beetle, Pyrrhalta luteola (Taos, New Mexico, USA). Cited on page 154.

Plate 35 Adult female and eggs of a sea grape sawfly, Sericoceros mexicanus (Roatan Island, Honduras). Cited on page 158.

Plate 36 Colony of larvae of Sericoceros mexicanus feeding on foliage of sea grape, Coccoloba uvifera (Roatan Island, Honduras). Cited on page 158.

Plate 37 Adult male of a pine sawfly, Neodiprion autumnalis (Elbert County, Colorado, USA). Cited on page 160.

Plate 38 Larval colony of a pine sawfly, Neodiprion autumnalis, feeding on foliage of ponderosa pine, Pinus ponderosa (Photo by Patricia M. Ciesla). Cited on page 160.

Plate 39 Red boring dust indicative of attack by a bark beetle, Phloeosinus bicolor, in Juniperus procera (near Maralal, Kenya). Cited on page 173.

Plate 40 White boring dust indicative of attack by an unidentified species of ambrosia beetle on a recently fallen tree (Jardin Botanico Dr Alfredo Barrera, Puerto Morales, Mexico). Cited on page 174.

Plate 41 Aerial view of tree mortality caused by southern pine beetle, Dendroctonus frontalis (photo by R.F. Billings, Texas Forest Service). Cited on page 178.

Plate 42 Heavy mortality of high-elevation Engelmann spruce, Picea engelmannii, caused by spruce beetle, Dendroctonus rufipennis (Rio Grande National Forest, Colorado, USA). Cited on page 183.

Plate 43 Galleries of the bark beetle, Phloeosinus armatus, in Cupressus sempervirens (northern Cyprus). Cited on page 185.

Plate 44 Dead branch tips on Cupressus sempervirens due to adult feeding by the bark beetle, Phloeosinus armatus (northern Cyprus). Cited on page 185.

Plate 45 Top kill of Pinus sylvestris caused by the engraver beetle, Ips acuminatus (Lower Caucuses, Republic of Georgia). Cited on page 191.

Plate 46 Top kill of Pinus ponderosa caused by the pine engraver beetle, Ips pini (Warm Springs Indian Reservation, Oregon, USA). Cited on page 192.

Plate 47 Mature adult of the larger European spruce bark beetle, Ips typographus (Bavaria, Germany). Cited on page 193.

Plate 48 Egg and larval galleries of the larger European spruce bark beetle, Ips typographus (Bavarian National Park, Germany). Cited on page 193.

Plate 49 Emerald ash borer, Agrilus planipennis, adult (Photo by David Cappaert, courtesy of forestryimages.org). Cited on page 207.

Plate 50 Emerald ash borer, Agrilus planipennis, larva in gallery (Photo by David Cappaert, courtesy of forestryimages.org). Cited on page 207.

Plate 51 Adult Japanese pine sawyer, Monochamus alternatus, Anhui Province, China. Cited on page 215.

Plate 52 Adult northern pine sawyer, Monochamus scutellatus (Graham County, North Carolina, USA). Cited on page 216.

Plate 54 Yellow phoracantha borer, Phoracantha recurva, on eucalyptus (near Rancagua, VI Region, Chile). Cited on page 219. Plate 53 Adult black locust borer, Megacyllene robinae, on goldenrod, Solidago spp. (Linville Gorge, North Carolina, USA). Cited on page 217.

Plate 55 Pupal cases of lilac-ash borer, Podesesia syringae, by exit hole (Fort Collins, Colorado, USA). Cited on page 226.

Plate 56 Pupa of the woodwasp, Sirex noctilio, in larval gallery (Santa Catarina State, Brazil). Cited on page 229.

Plate 57 Female adult of the woodwasp, Sirex noctilio (Santa Catarina State, Brazil). Cited on page 229.

Plate 58 Adult male of the woodwasp, Sirex noctilio (Santa Catarina State, Brazil). Cited on page 229.

Plate 59 Infestation of leucaena psyllid, Heteropsylla cubana, on new shoots of Leucaena leucocephala (Morogoro, Tanzania). Cited on page 233.

Plate 60 Colony of cypress aphids, Cinara cupressivora, on stem of Cupressus lusitanica (Muguga, Kenya). Cited on page 236.

Plate 61 Monterrey cypress, Cupressus macrocarpa, damaged by cypress aphid, near Rancagua (VI Region, Chile). Cited on page 236.

Plate 62 Cypress hedge with damage caused by cypress aphid, Cinara cupressivora (Concha y Toro Winery, near Santiago, Chile). Cited on page 236.

Plate 63 Hemlock woolly adelgid, Adelges tsugae, infestation on branch of eastern hemlock, Tsuga canadensis (Pisgah National Forest, North Carolina, USA). Cited on page 239.

Plate 64 Mortality of large eastern hemlock caused by hemlock woolly adelgid, Adelges tsugae (Joyce Kilmer Memorial Forest, Nantahala National Forest, North Carolina, USA). Cited on page 239.

Plate 65 Gout on branches of Fraser fir, Abies fraseri, caused by balsam woolly adelgid, Adelges piceae (Black Mountains, North Carolina, USA). Cited on page 240.

Plate 66 Tree mortality in high-elevation spruce–fir forest caused by balsam woolly adelgid, Adelges piceae (Great Smoky Mountains National Park, North Carolina, USA). Cited on page 240.

Plate 67 Cooley spruce gall adelgid, Adelges cooleyi, on blue spruce, Picea pungens (Fort Collins, Colorado, USA). Cited on page 256.

Plate 68 Oak apple gall caused by the gall wasp, Amphibolips confluenta, on northern red oak, Quercus rubra. (Photo by Brian Howell, USDA Forest Service). Cited on page 263.

Plate 69 Galls on burr oak, Quercus macrocarpa, caused by rough oak bullet gall wasp, Disholcaspis quercusmamma (Fort Collins, Colorado, USA). Cited on page 264.

Plate 70 Galls on Chinese chestnut, Castanea mollissima, caused by chestnut gall wasp, Dryocosmus kuriphilus (Photo by E. Richard Hoebecke, Cornell University, Ithaca, New York, USA, courtesy of forestryimages.org). Cited on page 264.

Plate 71 Open gall of piñon spindle gall midge Pinyonia edulicola showing orange larvae (Fort Collins, CO, USA). Cited on page 267.

Plate 72 Mortality of Pinus radiata regeneration caused by adult feeding of the bark beetle Hylastes ater (V Region, Chile). Cited on page 280.

Plate 73 Twig mortality on bristlecone pine, Pinus aristata, caused by the twig beetle, Pityophthorus boycei (photo by Brian Howell, USDA Forest Service). Cited on page 283.

Plate 74 Adult twig beetles, Pityophthorus boycei, in shoot of bristlecone pine, Pinus aristata (Pike National Forest, Colorado, USA). Cited on page 284.

Plate 75 Shoot damage on pine caused by adult feeding of the bark beetle, Tomicus piniperda (photo by E. Richard Hoebecke, Cornell University, Ithaca, New York, USA, courtesy of forestryimages.org). Cited on page 286.

Plate 76 Adult western conifer seed bug, Leptoglossus occidentalis, on screen door in autumn (Photo by Patricia M. Ciesla). Cited on page 297.

Plate 77 Exit holes characteristic of seed chalcids of the genus Megastigmus on seeds of Juniperus polycarpos var. seravschanica (Ziarat Forest, Balochistan Province, Pakistan). Cited on page 303.

Plate 78 Larva of a seed worm, Cydia sp., overwintering in the axis of a pine cone (Photo by Larry J. Barber, USDA Forest Service). Cited on page 308.

Plate 79 Larvae, adult and cone damage by southern pine coneworm, Dioryctria amatella (Photo by R.F. Billings, Texas Forest Service). Cited on page 311.

Plate 80 Workers of subterranean termites of the genus Reticulitermes (Photo by David Cappaert, courtesy of forestryimages.org). Cited on page 318.

Plate 81 Adult golden buprestrid, Buprestis aurulenta (Idaho, USA). Cited on page 326.