Forest Pest Insects

in North America:
A Photographic
Guide

Roy G. Van Driesche

Joseph H. LaForest
Charles T. Bargeron
Richard C. Reardon
Megan Herlihy

FHTET-2012-02
September 2013

Center for Invasive

and
Species
Ecosystem Health
Warnell School of College of Agricultural
Forestry and Natural Resources and Environmental Sciences
Forest Pest Insects in North America:
A Photographic Guide
Forest Pest Insects
in North America:
A Photographic
Guide
Forest Pest Insects
in North America:
A Photographic
Guide
Roy G. Van Driesche1
Joseph H. LaForest2
Charles T. Bargeron2
Richard C. Reardon3
Megan Herlihy1

1
University of Massachusetts, Environmental Conservation/Entomology
2
University of Georgia
3
USDA Forest Service, Forest Health Technology Enterprise Team

Center for Invasive

and
Species
Ecosystem Health
Warnell School of College of Agricultural
Forestry and Natural Resources and Environmental Sciences

FHTET-2012-02
September 2013
Cover Image Credits
Ponderosa pine stand; Chris Schnepf, University of Idaho, Bugwood.org
Male pine butterfly, Neophasia menapia; Terry Spivey, USDA Forest Service,
Bugwood.org

For additional copies of this publication, contact:

Richard C. Reardon
USDA Forest Service
180 Canfield Avenue
Morgantown, WV 26505
304-285-1566
[email protected]

An electronic version of this publication is available online at:

http://www.forestpests.org/book/

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Preface
The information in this publication is intended to help foresters, urban
landscaping employees, or others working with trees recognize some of
the common native and nonnative pest insects affecting trees in North
America, and understand their life cycles and how they damage trees.
Recognition is based on photographs of the pest in various lifestages
rather than on technical keys, and the user’s identification of a pest
may not be accurate if there are many similar species in a group. The
information was drawn from books, websites, factsheets, and some
original literature. This publication is not a guide for specialists. In
many groups, such as the bark beetles and aphids, confirmation of
species identity requires attention to details not visible in photos with
comparisons to other similar species and use of keys. Information on
important biological control agents of each species is provided when
available. Sources for further information (websites and articles) are
given at the bottom of each species’ page; however, an exhaustive
review of the literature was beyond the scope of this project. In most
cases, pages on individual species were reviewed by experts with direct
knowledge of the species. While any residual mistakes remain mine
(Roy Van Driesche), I am deeply indebted to the many people who
greatly improved pages on particular insects with their comments,
photos, and edits. I would also like to thank Wendy Harding for
design and layout of this printed version.
Contents

1. Pear Thrips, Taeniothrips inconsequens (Uzel) (Thysanoptera: Thripidae) . . . . . . . . . . . . . . . . . . . 1

2. Introduced Basswood Thrips, Thrips calcaratus Uzel (Thysanoptera: Thripidae) . . . . . . . . . . . . . . . 3
3. Redbanded Thrips, Selenothrips rubrocinctus (Giard) (Thysanoptera: Thripidae) . . . . . . . . . . . . . . 6
4. Slash Pine Flower Thrips, Gnophothrips fuscus (Morgan) (Thysanoptera: Phlaeothripidae) . . . . . . 9
5. Northern Walkingstick, Diapheromera femorata (Say) (Phasmatodea: Heteronemiidae) . . . . . . . . 11
6. Oak Lace Bug, Corythucha arcuata (Say) (Hemiptera: Tingidae) . . . . . . . . . . . . . . . . . . . . . . . . . . 15
7. Sycamore Lace Bug, Corythucha ciliata (Say) (Hemiptera: Tingidae) . . . . . . . . . . . . . . . . . . . . . . 18
8. Shield-backed Pine Seed Bug, Tetyra bipunctata (Herrich-Schäffer)
(Hemiptera: Scutellaridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
9. Leaffooted Pine Seed Bug, Leptoglossus corculus (Say) (Hemiptera: Coreidae) . . . . . . . . . . . . . . 25
10. Royal Palm Bug, Xylastodoris luteolus Barber (Hemiptera: Thaumastocoridae) . . . . . . . . . . . . . 27
11. Saratoga Spittlebug, Aphrophora saratogensis (Fitch) (Hemiptera: Cercopidae) . . . . . . . . . . . . . 29
12. Pine Spittlebug, Aphrophora parallela (Say) (Hemiptera: Cercopidae) . . . . . . . . . . . . . . . . . . . . . 32
13. Thorn Bug, Umbonia crassicornis (Amyot and Serville) (Hemiptera: Membracidae) . . . . . . . . . . 34
14. Periodical Cicada, Magicicada septendecim L. (Hemiptera: Cicadidae) . . . . . . . . . . . . . . . . . . . . 36
15. Eastern Spruce Gall Adelgid, Adelges abietis (L.) (Hemiptera: Adelgidae) . . . . . . . . . . . . . . . . . . 41
16. Balsam Woolly Adelgid, Adelges piceae (Ratzeburg) (Hemiptera: Adelgidae) . . . . . . . . . . . . . . . 44
17. Hemlock Woolly Adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae) . . . . . . . . . . . . . . . . . 48
18. Cooley Spruce Gall Adelgid, Adelges cooleyi (Gillette) (Hemiptera: Adelgidae) . . . . . . . . . . . . . . 53
19. Pine Bark Adelgid, Pineus strobi (Hartig) (Hemiptera: Adelgidae) . . . . . . . . . . . . . . . . . . . . . . . . 56
20. Pine Leaf Adelgid, Pineus pinifoliae (Fitch) (Hemiptera Adelgidae) . . . . . . . . . . . . . . . . . . . . . . . 59
21. White Pine Aphid, Cinara strobi (Fitch) (Hemiptera: Aphidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
22. Woolly Elm Aphid, Eriosoma americanum (Riley) (Hemiptera: Aphididae; Eriosomatinae) . . . . . 64
23. Woolly Beech Aphid, Phyllaphis fagi (L.) (Hemiptera: Aphidae) . . . . . . . . . . . . . . . . . . . . . . . . . . 66
24. Tuliptree Aphid, Illinoia liriodendri (Monell) (Hemiptera: Aphididae) . . . . . . . . . . . . . . . . . . . . . . . 68
25. Norway Maple Aphid, Periphyllus lyropictus (Kessler) (Hemiptera: Aphidae) . . . . . . . . . . . . . . . . 70
26. Linden Aphid, Eucallipterus tiliae (L.) (Hemiptera: Aphididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
27. Spruce Aphid, Elatobium abietinum (Walker) (Hemiptera: Aphidae) . . . . . . . . . . . . . . . . . . . . . . . 76
28. Beech Scale, Cryptococcus fagisuga Lindinger (Hemiptera: Eriococcidae) . . . . . . . . . . . . . . . . . 79
29. Gillette’s Eriococcin, Eriokermes gillettei (Tinseley)
(Hemiptera: Kermesidae, formerly Eriococcidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
30. Pine Bast Scale, Matsucoccus matsumurae (Kuwana) (Hemiptera: Matsucoccidae) . . . . . . . . . . 86
31. Tuliptree Scale, Toumeyella liriodendri (Gmelin) (Hemiptera: Coccidae) . . . . . . . . . . . . . . . . . . . 88
32. Pine Tortoise Scale, Toumeyella parvicornis (Cockerell) (Hemiptera: Coccidae) . . . . . . . . . . . . . 91
33. Tessellated Scale, Eucalymnatus tessellatus (Signoret) (Hemiptera: Coccidae) . . . . . . . . . . . . . 95
Contents (continued)

34. Elongate Hemlock Scale, Fiorinia externa Ferris (Hemiptera: Diaspididae) . . . . . . . . . . . . . . . . . 97

35. Oystershell Scale, Lepidosaphes ulmi (L.) (Hemiptera: Diaspididae) . . . . . . . . . . . . . . . . . . . . . 101
36. San José Scale, Diaspidiotus perniciosus (Comstock) (Hemiptera: Diaspididae) . . . . . . . . . . . 105
37. Willow Scale, Diaspidiotus gigas (Thiem and Gerneck) (Hemiptera: Diaspididae) . . . . . . . . . . . 110
38. Camphor Scale, Pseudaonidia duplex (Cockerell) (Hemiptera: Diaspididae) . . . . . . . . . . . . . . . 114
39. Coconut Scale, Aspidiotus destructor Signoret (Hemiptera: Diaspididae) . . . . . . . . . . . . . . . . . 117
40. Florida Red Scale, Chrysomphalus aonidum (L.) (Hemiptera: Diaspididae) . . . . . . . . . . . . . . . . 120
41. Magnolia White Scale, Pseudaulacaspis cockerelli (Cooley) (Hemiptera: Diaspididae) . . . . . . . 122
42. White Peach Scale, Pseudaulacaspis pentagona (Targioni-Tozzetti)
(Hemiptera: Diaspididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
43. Citrus Whitefly, Dialeurodes citri (Ashmead) (Hemiptera: Aleyrodidae) . . . . . . . . . . . . . . . . . . . 126
44. The Mealybug Oracella acuta (Lobdell) (Hemiptera: Pseudococcidae) . . . . . . . . . . . . . . . . . . . 130
45. Cottonwood Leaf Beetle, Chrysomela scripta Fabricius (Coleoptera: Chrysomelidae) . . . . . . . . 133
46. Imported Willow Leaf Beetle, Plagiodera versicolora (Laicharting)
(Coleoptera: Chrysomelidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
47. Elm Leaf Beetle, Xanthogaleruca luteola (Müller) (Coleoptera: Chrysomelidae) . . . . . . . . . . . . 139
48. Larger Elm Leaf Beetle, Monocesta coryli (Say) (Coleoptera: Chrysomelidae) . . . . . . . . . . . . . 144
49. Giant Palm Weevil, Rhynchophorus cruentatus (F.) (Coleoptera: Curculionidae) . . . . . . . . . . . . 147
50. Red Palm Weevil, Rhynchophorus ferrugineus (Olivier) (Coleoptera: Dryophthoridae) . . . . . . . 150
51. Pales Weevil, Hylobius pales (Herbst) (Coleoptera: Curculionidae) . . . . . . . . . . . . . . . . . . . . . . 154
52. Pine Root Collar Weevil, Hylobius radicis (Buchanan) (Coleoptera: Curculionidae) . . . . . . . . . . 157
53. Pitch-eating Weevil, Pachylobius picivorus (Germar) (Coleoptera: Curculionidae) . . . . . . . . . . . 160
54. White Pine Weevil, Pissodes strobi (Peck) (Coleoptera: Curculionidae) . . . . . . . . . . . . . . . . . . . 163
55. Eastern Pine Weevil, Pissodes nemorensis Germar (Coleoptera: Curculionidae) . . . . . . . . . . . 167
56. Whitefringed Beetles (Naupactus spp.) (Coleoptera: Curculionidae) . . . . . . . . . . . . . . . . . . . . . 170
57. Yellow-poplar Weevil, Odontopus calceatus (Say) (Coleoptera: Curculionidae) . . . . . . . . . . . . . 174
58. Native Elm Bark Beetle, Hylurgopinus rufipes (Eichhoff)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
59. Smaller European Elm Bark Beetle, Scolytus multistriatus (Marsham)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
60. Spruce Beetle, Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae, Scolytinae) . . . . . 186
61. Black Turpentine Beetle, Dendroctonus terebrans (Olivier)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
62. Eastern Larch Beetle, Dendroctonus simplex (LeConte)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Contents (continued)

63. Southern Pine Beetle, Dendroctonus frontalis (Zimmermann)

(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
64. Western Pine Beetle, Dendroctonus brevicomis LeConte
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
65. Jeffrey Pine Beetle, Dendroctonus jeffreyi Hopkins
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
66. Mountain Pine Beetle, Dendroctonus ponderosae Hopkins
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
67. Douglas-fir Beetle, Dendroctonus pseudotsugae Hopkins
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
68. Engraver Beetle, Ips typographus (L.) (Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . 224
69. Pine Engraver Beetle, Ips pini (Say) (Coleoptera: Curculionidae [formerly Scolytidae]) . . . . . . . 229
70. Southern Pine Engraver, Ips grandicollis (Eichhoff)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
71. Six-spined Ips, Ips calligraphus (Germar) (Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . 238
72. Small Southern Pine Engraver, Ips avulsus Eichhoff
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
73. Silver Fir Beetle, Pseudohylesinus sericeus (Mannerheim)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
74. Fir Engraver, Scolytus ventralis LeConte (Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . 250
75. Columbian Timber Beetle, Corthylus columbianus Hopkins
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
76. Ambrosia Beetles, Xyleborus spp. (Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . 257
77. Redbay Ambrosia Beetle, Xyleborus glabratus Eichhoff
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
78. Walnut Twig Beetle, Pityophthorus juglandis Blackman
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
79. Striped Ambrosia Beetle, Trypodendron lineatum (Olivier)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
80. Black Twig Borer, Xylosandrus compactus (Eichhoff)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
81. Larger Pine Shoot Beetle, Tomicus piniperda (L.)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
82. White Pine Cone Beetle, Conophthorus coniperda (Schwarz)
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290
83. Lodgepole Cone Beetle, Conophthorus ponderosae Hopkins
(Coleoptera: Curculionidae, Scolytinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
85. May and June Beetles, Phyllophaga spp. (Coleoptera: Scarabaeidae) . . . . . . . . . . . . . . . . . . . 300
Contents (continued)

86. Black Vine Weevil, Otiorhynchus sulcatus (Fabricius) (Coleoptera: Curculionidae) . . . . . . . . . . 306
87. Strawberry Root Weevil, Otiorhynchus ovatus (L.) (Coleoptera: Curculionidae) . . . . . . . . . . . . . 310
88. Japanese Beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae) . . . . . . . . . . . . . . . . . 313
89. Twolined Chestnut Borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae) . . . . . . . . . . . . 317
90. Bronze Birch Borer, Agrilus anxius Gory (Coleoptera: Buprestidae) . . . . . . . . . . . . . . . . . . . . . . 321
91. Emerald Ash Borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) . . . . . . . . . . . . . . 326
92. Goldspotted Oak Borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae) . . . . . . . . . . 335
93. Soapberry Borer, Agrilus prionurus Chevrolat (Coleoptera: Buprestidae) . . . . . . . . . . . . . . . . . . 340
94. Hickory Spiral Borer, Agrilus torquatus LeConte (Coleoptera: Buprestidae) . . . . . . . . . . . . . . . . 344
95. Flatheaded Appletree Borer, Chrysobothris femorata (Olivier) (Coleoptera: Buprestidae) . . . . . 347
96. Red Oak Borer, Enaphalodes rufulus (Haldeman) (Coleoptera: Cerambycidae) . . . . . . . . . . . . 350
97. Black Fir Sawyer Beetle, Monochamus urussovii (Fischer von Waldheim)
(Coleoptera: Cerambycidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355
98. Japanese Pine Sawyer, Monochamus alternatus Hope (Coleoptera: Cerambycidae) . . . . . . . . 358
99. Asian Longhorned Beetle, Anoplophora glabripennis Motschulsky
(Coleoptera: Cerambycidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365
100. Poplar Borer, Saperda calcarata Say (Coleoptera: Cerambycidae) . . . . . . . . . . . . . . . . . . . . . 373
102. Cottonwood Borer, Plectrodera scalator (Fabricius) (Coleoptera: Cerambycidae) . . . . . . . . . . 377
103. Banded Hickory Borer, Knulliana cincta (Drury) (Coleoptera: Cerambycidae) . . . . . . . . . . . . . 381
104. Locust Borer, Megacyllene robiniae (Forster) (Coleoptera: Cerambycidae) . . . . . . . . . . . . . . . 383
105. White Oak Borer, Goes tigrinus (De Geer) (Coleoptera: Cerambycidae) . . . . . . . . . . . . . . . . . 387
106. Whitespotted Sawyer, Monochamus scutellatus (Say) (Coleoptera: Cerambycidae) . . . . . . . . 390
107. Bagworm, Thyridopteryx ephemeraeformis (Haworth) (Lepidoptera: Psychidae) . . . . . . . . . . . 393
108. Birch Casebearer, Coleophora serratella (L.) (Lepidoptera: Coleophoridae) . . . . . . . . . . . . . . 398
109. Larch Casebearer, Coleophora laricella (Hübner) (Lepidoptera: Coleophoridae) . . . . . . . . . . . 400
110. Pecan Cigar Casebearer, Coleophora laticornella (Clemens)
(Lepidoptera: Coleophoridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405
111. Oak Skeletonizer, Bucculatrix ainsliella Murtfeldt (Lepidoptera: Bucculatricidae) . . . . . . . . . . . 407
112. Palm Leaf Skeletonizer, Homaledra sabalella (Chambers) (Lepidoptera: Coleophoridae) . . . . 411
113. Mimosa Webworm, Homadaula anisocentra Meyrick (Lepidoptera: Galacticidae) . . . . . . . . . . 413
114. Lodgepole Needleminer, Coleotechnites milleri (Busck) (Lepidoptera: Gelechiidae) . . . . . . . . 416
115. Banded Ash Clearwing, Podosesia aureocincta Purrington and Nielson,
and Ash/Lilac Clearwing, Podosesia syringae (Harris) (Lepidoptera: Sesiidae) . . . . . . . . . . . . 419
116. Red Oak Clearwing Borer, Paranthrene simulans (Grote) (Lepidoptera: Sesiidae) . . . . . . . . . 423
117. Persimmon Borer, Sannina uroceriformis Walker (Lepidoptera: Sesiidae) . . . . . . . . . . . . . . . . 425
Contents (continued)

118. Poplar Clearwing Borer, Paranthrene dollii (Neumoegen) (Lepidoptera: Sesiidae) . . . . . . . . . . 428
119. Carpenterworm, Prionoxystus robiniae (Peck.) (Lepidoptera: Cossidae) . . . . . . . . . . . . . . . . . 431
120. Pecan Carpenterworm, Cossula magnifica (Stecker) (Lepidoptera: Cossidae) . . . . . . . . . . . . 436
121. Spruce Budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) . . . . . . . . . 439
122. Jack Pine Budworm, Choristoneura pinus pinus Freeman (Lepidoptera: Tortricidae) . . . . . . . . 445
123. Large Aspen Tortrix, Choristoneura conflictana (Walker) (Lepidoptera: Tortricidae) . . . . . . . . . 451
124. Sugar Pine Tortrix, Choristoneura lambertiana Busck (Lepidoptera: Tortricidae) . . . . . . . . . . . 456
125. Western Spruce Budworm, Choristoneura occidentalis Freeman
(Lepidoptera: Tortricidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 460
126. Spruce Bud Moth, Zeiraphera canadensis Mutuura and Freeman
(Lepidoptera: Tortricidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 466
127. Cottonwood Twig Borer, Gypsonoma haimbachiana (Kearfott) (Lepidoptera: Tortricidae) . . . . 469
129. Oak Leafroller, Archips semiferanus (Walker) (Lepidoptera: Tortricidae) . . . . . . . . . . . . . . . . . 473
130. Longleaf Pine Seedworm, Cydia ingens (Heinrich) (Lepidoptera: Tortricidae) . . . . . . . . . . . . . 475
131. European Pine Shoot Moth, Rhyacionia buoliana (Denis and Schiffermiiller)
(Lepidoptera: Tortricidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
132. Nantucket Pine Tip Moth, Rhyacionia frustrana (Comstock) (Lepidoptera: Tortricidae) . . . . . . 482
133. Pitch Pine Tip Moth, Rhyacionia rigidana (Fernald) (Lepidoptera: Tortricidae) . . . . . . . . . . . . . 487
134. Southwestern Pine Tip Moth, Rhyacionia neomexicana (Dyar) (Lepidoptera: Tortricidae) . . . . 489
135. Eastern Pine Shoot Borer, Eucosma gloriola Heinrich (Lepidoptera: Tortricidae) . . . . . . . . . . . 491
136. Lodgepole Cone Moth, Eucosma rescissoriana Heinrich (Lepidoptera: Tortricidae) . . . . . . . . . 494
137. Western Pine Shoot Borer, Eucosma sonomana Kearfott (Lepidoptera: Tortricidae) . . . . . . . . 497
138. Douglas-fir Cone Moth, Barbara colfaxiana (Kearfott) (Lepidoptera: Tortricidae) . . . . . . . . . . . 500
139. Pine Webworm, Pococera robustella (Zeller) (Lepidoptera: Pyralidae) . . . . . . . . . . . . . . . . . . . 503
140. Zimmerman Pine Moth, Dioryctria zimmermani (Grote) (Lepidoptera: Pyralidae) . . . . . . . . . . 506
141. Fall Cankerworm, Alsophila pometaria (Harris) (Lepidoptera: Geometridae) . . . . . . . . . . . . . . 509
142. Spring Cankerworm, Paleacrita vernata (Peck) (Lepidoptera: Geometridae) . . . . . . . . . . . . . . 513
143. Eastern Hemlock Looper, Lambdina fiscellaria fiscellaria Guenée
(Lepidoptera: Geometridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 516
144. Western Hemlock Looper, Lambdina fiscellaria lugubrosa (Hulst)
(Lepidoptera: Geometridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519
145. Greenstriped Forest Looper, Melanolophia imitata (Walker) (Lepidoptera: Geometridae) . . . . 522
146. Saddleback Looper, Ectropis crepuscularia (Denis and Schiffermüller)
(Lepidoptera: Geometridae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 524
147. Spearmarked Black Moth, Rheumaptera hastata (L.) (Lepidoptera: Geometridae) . . . . . . . . . 526
148. Elm Spanworm, Ennomos subsignarius (Hübner) (Lepidoptera: Geometridae) . . . . . . . . . . . . 529
Contents (continued)

149. Winter Moth, Operophtera brumata (L.) (Lepidoptera: Geometridae) . . . . . . . . . . . . . . . . . . . . 533

150. Bruce Spanworm, Operophtera bruceata (Hulst) (Lepidoptera: Geometridae) . . . . . . . . . . . . . 536
151. Forest Tent Caterpillar, Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae) . . . . . . . . 538
152. Eastern Tent Caterpillar, Malacosoma americanum (F.) (Lepidoptera: Lasiocampidae) . . . . . . 543
153. Siberian Moth, Dendrolimus sibiricus TschetvericKov (Lepidoptera: Lasiocampidae) . . . . . . . 549
154. Pinkstriped Oakworm, Anisota virginiensis (Drury) (Lepidoptera: Saturniidae) . . . . . . . . . . . . . 553
155. Orangestriped Oakworm, Anisota senatoria (J. E. Smith) (Lepidoptera: Saturniidae) . . . . . . . 557
156. Pandora Moth, Coloradia pandora Blake (Lepidoptera: Saturniidae) . . . . . . . . . . . . . . . . . . . . 560
157. European Gypsy Moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . 563
158. Asian Pink Moth, Lymantria mathura Moore (Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . . 569
159. Nun Moth, Lymantria monacha Moore (Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . . . . . . 573
160. Browntail Moth, Euproctis chrysorrhoea L. (Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . . . 578
161. Pine Tussock Moth, Dasychira pinicola (Dyar) (Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . 583
162. Whitemarked Tussock Moth, Orgyia leucostigma (J. E. Smith)
(Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 586
163. Douglas-fir Tussock Moth, Orgyia pseudotsugata (McDunnough)
(Lepidoptera: Lymantriidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 590
164. Poplar Tentmaker, Clostera inclusa (Hübner) (Lepidoptera: Notodontidae) . . . . . . . . . . . . . . . 594
165. Saddled Prominent, Heterocampa guttivitta (Walker) (Lepidoptera: Notodontidae) . . . . . . . . . 597
166. Walnut Caterpillar, Datana integerrima Grote and Robinson (Lepidoptera: Notodontidae) . . . . 600
167. Variable Oak Leaf Caterpillar, Lochmaeus manteo (Doubleday)
(Lepidoptera: Notodontidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 605
168. Fall Webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae) . . . . . . . . . . . . . . . . . . . . . . 609
169. Arborvitae Leafminer, Argyresthia thuiella (Packard) (Lepidoptera: Argresthiidae) . . . . . . . . . . 614
170. Pine Needle Sheathminer, Zelleria haimbachi Busck (Lepidoptera: Yponomeutidae) . . . . . . . 618
171. Pine Butterfly, Neophasia menapia (Felder and Felder) (Lepidoptera: Pieridae) . . . . . . . . . . . 621
172. Balsam Gall Midge, Paradiplosis tumifex Gagné (Diptera: Cecidomyiidae) . . . . . . . . . . . . . . . 625
173. Poplar Leafcurl Midge, Prodiplosis morrisi Gagné (Diptera: Cecidomyiidae) . . . . . . . . . . . . . . 628
174. Douglas-fir Needle Midge, Contarinia pseudotsugae Condrashoff
(Diptera: Cecidomyiidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 630
175. Pine False Webworm, Acantholyda erythrocephala (L.) (Hymenoptera: Pamphiliidae) . . . . . . 633
176. Redheaded Pine Sawfly, Neodiprion lecontei (Fitch) (Hymenoptera: Diprionidae) . . . . . . . . . . 637
177. Swaine Jack Pine Sawfly, Neodiprion swainei (Middleton) (Hymenoptera: Diprionidae) . . . . . 641
178. European Pine Sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera: Diprionidae) . . . . . . . . . 644
179. Virginia Pine Sawfly, Neodiprion pratti pratti (Dyar) (Hymenoptera: Diprionidae) . . . . . . . . . . . 648
180. Hemlock Sawfly, Neodiprion tsugae Middleton (Hymenoptera: Diprionidae) . . . . . . . . . . . . . . 651
Contents (continued)

181. Introduced Pine Sawfly, Diprion similis (Hartig) (Hymenoptera: Diprionidae) . . . . . . . . . . . . . . 655
182. European Spruce Sawfly, Gilpinia hercyniae (Hartig) (Hymenoptera: Diprionidae) . . . . . . . . . 659
183. Larch Sawfly, Pristiphora erichsonii (Hartig) (Hymenoptera: Tenthridinidae) . . . . . . . . . . . . . . 662
184. Mountain Ash Sawfly, Pristiphora geniculata (Hartig) (Hymenoptera: Tenthridinidae) . . . . . . . 666
185. Yellowheaded Spruce Sawfly, Pikonema alaskensis (Rohwer)
(Hymenoptera: Tenthredinidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 669
186. Birch Leafminer, Fenusa pumila (Leach) (Hymenoptera: Tenthridinidae) . . . . . . . . . . . . . . . . . 673
187. Ambermarked Birch Leafminer, Profenusa thomsoni (Konow)
(Hymenoptera: Tenthridinidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 678
188. Sirex Woodwasp, Sirex noctilio (Fabricius) (Hymenoptera: Siricidae) . . . . . . . . . . . . . . . . . . . 682
189. Asian Chestnut Gall Wasp, Drycosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) . . . . 689
190. Spruce Spider Mite, Oligonychus ununguis (Jacobi) (Acari: Tetranychidae) . . . . . . . . . . . . . . . 692
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697
A Photographic Guide 1

1. Pear Thrips, Taeniothrips inconsequences (Uzel) (Thysanoptera:

Thripidae)

Orientation to Pest Pear thrips, Taeniothrips inconsequences (Uzel), is an invasive species

in North America from Europe, first being recorded in the USA in
California in 1900. Pear thrips have rasping/sucking mouthparts and use
them to feed in early spring on buds and young leaves of hardwood trees.
Potential for damage to sugar maples (Acer sacchrum Marshall) has been
the greatest concern. Feeding injuries to embryonic or developing leaves
result in mature leaves being deformed or tattered. Damage may spike in
some years (e.g., the late 1980s) when thrips emerge earlier with respect
to timing of bud break of key species, but such events are weather-
dependent and do not occur regularly. In general the damage from this
species is light.

Hosts Commonly Attacked Sugar maples (A. sacchrum Marshall) and other hardwoods.

Distribution No map available. The species is found in California, the northeast USA
and southern Canada.

Images of Pear Thrips

Figure 1. Adult pear thrips, Taeniothrips inconsequences. (Left: Pennsylvania

Department of Conservation and Natural Resources - Forestry Archive,
Bugwood.org; right: Pennsylvania Bureau of Forestry)

Figure 2. Larva of pear thrips. (Pennsylvania Department of Conservation and

Natural Resources - Forestry Archive, Bugwood.org)
2 Forest Pest Insects in North America:

Images of Pear Thrips

(continued)

Figure 3. Leaf deformities on sugar maple due to feeding of pear thrips on buds.
(Left: Ronald S. Kelley, Vermont Department of Forests, Parks and Recreation,
Bugwood.org; right: Margaret Miller-Weeks, USDA Forest Service)

Important Biological Control No specialized natural enemies of this thrips are known and the minor
Agents Related to this and irregular damage caused by this pest suggests that it is best left
Pest Species unmanaged or controlled only on limited acreage with pesticides if need
be in years of unusually high damage.

Web Links for Information http://ento.psu.edu/extension/factsheets/pear-thrips-PA; bulletin from

on Pear Thrips Pennsylvania State University on biology and control.
http://ecommons.cornell.edu/bitstream/1813/14346/2/Pear%20Thrips.
pdf; bulletin from Cornell University on biology and control.
http://www.na.fs.fed.us/spfo/pubs/pest_al/pt/pt.htm; bulletin of the
United States Department of Agriculture.
http://www.onlinegardener.com/pests/Pear%20thrips.pdf; bulletin of
Bartlett Tree Company on biology and control.
http://www.dec.ny.gov/docs/lands_forests_pdf/thrips.pdf; article from
“New York Forest Owner” 1993 discussing impact of pear thrips on
forests.

Articles None
A Photographic Guide 3

2. Introduced Basswood Thrips, Thrips calcaratus Uzel (Thysanoptera:

Thripidae)

Orientation to Pest Introduced basswood thrips, Thrips calcaratus Uzel, is a European

species invasive in eastern North America. Basswood thrips have
rasping/sucking mouthparts and in early spring feed on buds, especially
of American basswood (Tilia americana L.). Such bud-feeding causes
mature leaves to be deformed or tattered and defoliation may occur.
Repeated defoliation of American basswood reduces tree growth
and increases rates of dieback from other causes. The species is of
importance in northern Wisconsin, where outbreaks of damage occur.

Hosts Commonly Attacked In North America, this species feeds on American basswood
(T. americana).

Distribution The species is found in New England and Quebec, westward through
Ontario, Pennsylvania, New York, and the Great Lake States.

Images of Introduced
Basswood Thrips

Figure 1. Adult of introduced basswood thrips, Thrips calcaratus. (Kenneth Raffa,

University of Wisconsin, Bugwood.org)

Figure 2. Larva of introduced basswood thrips. (Kenneth Raffa, University of

Wisconsin, Bugwood.org)
4 Forest Pest Insects in North America:

Images of Introduced
Basswood Thrips (continued)

Figure 3. Typical damage to basswood buds from thrips feeding (here by another
thrips, Neohydratothrips tiliae [Hood]). (Steven Katovich, USDA Forest Service,
Bugwood.org)

Figure 4. Defoliation of American basswood by introduced basswood thrips.

(Steven Katovich, USDA Forest Service, Bugwood.org)

Important Biological Control No specialized natural enemies of this thrips are known.
Agents Related to this
Pest Species

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/howtos/ht_bassthrips/ht_bassthrips.

on Introduced Basswood htm; USDA Bulletin on biology of introduced basswood thrips.
Thrips
A Photographic Guide 5

Articles Raffa, K.F. and D.J. Hall. 1988. Thrips calcaratus Uzel (Thysanoptera:
Thripidae), a new pest of basswood trees in the Great Lakes region.
Canadian Journal of Forest Research 19: 1662-1663.
Raffa, K.F. 1991. Biology and impact of Thrips calcaratus Uzel in the
Great Lakes region. In: Parker, B.L., M. Skinner, and T. Lewis (eds.).
Towards Understanding Thysanoptera. USDA Forest Service General
Technical Report NE-147: 317-324.
Werner, S.M., M.A. Albers, T. Cryderman, D. Diminic, R. Heyd, B.
Hrašovic, S. Kobro, S. Larsson, R. Mech, P. Niemela, M. Rousi, K.F.
Raffa, K. Scanlon, and S. Weber. 2006. Is the outbreak status of Thrips
calcaratus Uzel in North America due to altered host relationships?
Forest Ecology and Management 225: 200-206.
6 Forest Pest Insects in North America:

3. Redbanded Thrips, Selenothrips rubrocinctus (Giard) (Thysanoptera:

Thripidae)

Orientation to Pest Redbanded thrips, Selenothrips rubocinctus (Giard), is an invasive

polyphagous tropical thrips of Asian origin found in Hawaii, Florida,
and Puerto Rico. A sucking/rasping insect, its damage is similar in some
regards to that of mite damage on foliage, consisting of a mixture of
discolored areas and the dark areas that result from the liquid deposited
where eggs are inserted in plant tissues. Affected leaves may be distorted
or may drop from plant. Dense thrips populations may defoliate plants.
Thrips also produce honeydew, which leads to growth of sooty molds.

Hosts Commonly Attacked Hosts of redbanded thrips include various fruits (cashew, grape, mango,
avocado, guava), ornamentals, and shade trees.

Distribution Redbanded thrips is found in Hawaii, Florida, and Puerto Rico.

Images of
Redbanded Thrips

Figure 1. Adult of redbanded thrips, Selenothrips rubocinctus. (Lyle Buss,

University of Florida, Bugwood.org)

Figure 2. Pupae of redbanded thrips. (Lyle Buss, University of Florida,

Bugwood.org)
A Photographic Guide 7

Images of Redbanded
Thrips (continued)

Figure 3. Damage by redbanded thrips. (James L. Castner, University of Florida)

Figure 4. Damage of
redbanded thrips on
mango. (Scot Nelson,
University of Hawaii at
Manoa, Bugwood.org)

Important Biological Control Goetheana shakespearei Girault is a self-introduced eulophid parasitoid

Agents Related to this of redbanded thrips found in Florida since 1992, originally detected
Pest Species in a greenhouse. The same parasitoid under the name Dasyscapus
parvipennis was shipped from Trinidad to Puerto Rico in the 1930s and
established. In Australia, a weaver ant is cited as of importance, while in
South Africa, a species of predatory bug (Orius thripoborus [Hesse]) has
been noted as a predator of this thrips.
8 Forest Pest Insects in North America:

Web Links for Information http://edis.ifas.ufl.edu/pdffiles/IN/IN25600.pdf; University of Florida

on Redbanded Thrips factsheet.
https://transact.nt.gov.au/ebiz/dbird/TechPublications.nsf/
56FBE080115393C669256EFF004B540D/$file/752.pdf; an Australian
bulletin on redbanded thrips.

Articles Dennill, G.B. 1992. Orius thripoborus (Anthocoridae), a potential

biocontrol agent of Heliothrips haemorrhoidalis and Selenothrips
rubrocinctus (Thripidae) on avocado fruits in the eastern Transvaal.
Journal of the Entomological Society of Southern Africa 55(2): 255-258.
Bennett, F.D., H.A. Glenn, and R.M. Baranowski. 1993. Goetheana
shakespearei (Hymenoptera: Eulophidae) an immigrant parasitoid of
thrips in Florida and Guadeloupe. Florida Entomologist 76: 395-397.
Peng, R.K. and K. Christian. 2004 The weaver ant, Oecophylla
smaragdina (Hymenoptera: Formicidae), an effective biological control
agent of the red-banded thrips, Selenothrips rubrocinctus (Thysanoptera:
Thripidae) in mango crops in the Northern Territory of Australia.
International Journal of Pest Management 50(2): 107-114.
A Photographic Guide 9

4. Slash Pine Flower Thrips, Gnophothrips fuscus (Morgan) (Thysanoptera:

Phlaeothripidae)

Orientation to Pest The biology of slash pine flower thrips, Gnophothrips fuscus (Morgan),
is poorly known. It is assumed to be widespread throughout the eastern
United States but has only been of concern in seed orchards of slash pine
(Pinus elliottii var. elliottii Engelm.), where feeding on female cones
destroys about a quarter of the potential seed set.

Hosts Commonly Attacked The principal hosts are slash (P. elliottii var. elliottii), jack (P. banksiana
Lamb.), and loblolly (P. taeda L.) pines.

Distribution Slash pine flower thrips is found in the southeastern United States, from
Florida to east Texas, north to Ohio and Maryland.

Figure 1. U.S. distribution of slash

pine flower thrips, Gnophothrips
fuscus. (USDA Forest Service, “Seed
and Cone Insects of Southern Pines”)

Images of Slash Pine

Flower Thrips

Figure 2. Slash pine flower thrips. (István Mikó and the NCSU Insect Museum)
10 Forest Pest Insects in North America:

Images of Slash Pine

Flower Thrips (continued)

Figure 3. Young female slash pine cone showing damage from slash pine flower
thrips. (Bernard H. Ebel, USDA Forest Service, Bugwood.org)

Figure 4. Damage of slash pine

flower thrips as seen in mature cones.
(USDA Forest Service, “Seed and
Cone Insects of Southern Pines”)

Important Biological Control Nothing is known about specialized natural enemies of this species.
Agents Related to this
Pest Species

Web Links for Information http://216.166.86.145/pubs/seed-and-cone/page7.html; USDA Forest

on Slash Pine Flower Thrips Service bulletin “Seed and cone insects of southern pines.”

Articles Fatzinger, C.W. and W.N. Dixon. 1991. Development of sampling

methods for the slash pine flower thrips, Gnophothrips fuscus (Morgan)
(Thysanoptera: Phlaeothripidae). In: Parker, B.L., M. Skinner, and T.
Lewis (eds.). Towards Understanding Thysanoptera. USDA Forest
Service General Technical Report NE-147: 149-161.
A Photographic Guide 11

5. Northern Walkingstick, Diapheromera femorata (Say) (Phasmatodea:

Heteronemiidae)

Orientation to Pest The northern walkingstick, Diapheromera femorata (Say), is the only
walkingstick of economic importance in the USA. While young nymphs
feed on shrubs of various species, older nymphs and adults feed on
leaves of a wide variety of hardwood trees. Populations at times are
dense enough to completely defoliate affected trees. Eggs fall from
trees and when populations are dense, sounds of falling eggs are readily
noticed. Overwintering occurs in the egg stage.

Hosts Commonly Attacked Species fed on by the northern walkingstick include black (Quercus
velutina Lamb.) and red (Q. rubra L.) oaks, American basswood (Tilia
americana L.), American elm (Ulmus americana L.), black locust
(Robinia pseudoacacia L.), cherry (Prunus spp.), and other hardwoods.

Distribution This walkingstick is found in southern Canada and most of the eastern
United States, west to Texas and the Great Plains.

Images of
Northern Walkingstick

Figure 1. Adult of the northern walkingstick, Diapheromera femorata. (Kenneth

Raffa, University of Wisconsin, Bugwood.org)
12 Forest Pest Insects in North America:

Images of Northern Figure 2. Mating

pair of the northern
Walkingstick (continued) walkingstick; small
male mounted on large
female. (Kenneth Raffa,
University of Wisconsin,
Bugwood.org)

Figure 3. Seed-like eggs of the northern walkingstick being examined by an ant.

(Photo copyright Alex Wild/alexanderwild.com)
A Photographic Guide 13

Images of Northern
Walkingstick (continued)

Figure 4. Oak foliage showing feeding of the walkingstick. (Both photosl: James
Solomon, USDA Forest Service, Bugwood.org)

Figure 5. Forest in Mena, Arizona partially defoliated by the walkingstick. (James

Solomon, USDA Forest Service, Bugwood.org)
14 Forest Pest Insects in North America:

Important Biological Control The parasitoid Mesitiopterus kahlii (Ashmead) (Hymenoptera:

Agents Related to this Chrysididae) attacks the D. femorata. Also, birds can feed heavily on
Pest Species walkingsticks when their population densities are high.

Web Links for Information http://wiki.bugwood.org/Archive:South/Walkingstick

on Northern Walkingstick http://www.na.fs.fed.us/spfo/pubs/fidls/walkingstick/walkingstick.htm;
Forest Insect & Disease Leaflet 82, U.S. Department of Agriculture,
Forest Service.

Articles Ignoffo, C.M., D.L. Hostetter, and W.H. Kearby. 1973. Susceptibility of
walkingstick, orangestriped oakworm, and variable oakleaf caterpillar to
Bacillus thuringiensis var. alesti. Environmental Entomology 2: 807-809.
Giese, R.L. and K.H. Knauer. 1977. Ecology of the walkingstick. Forest
Science 23: 45-63.
A Photographic Guide 15

6. Oak Lace Bug, Corythucha arcuata (Say) (Hemiptera: Tingidae)

Orientation to Pest Oak lace bug, Corythucha arcuata (Say), is a minor pest of various oaks
in its native range in North America. The insect may overwinter as either
adults or eggs. Adults have lacey wings held flat over the body with an
ornate pronotum. Nymphs are mostly black with spines. Feeding begins
in early spring at leaf expansion. Eggs are laid on the underside of leaves.
Nymphs and adults occur in colonies on foliage where they feed by
sucking sap from plant tissues. Damage is similar to the yellow stippling
produced by spider mite feeding, but without webbing. Black fecal spots
are also present on foliage infested by lace bugs. Dense populations can
cause premature leaf drop. There are multiple generations per year.

Hosts Commonly Attacked The main hosts of this species are white (Quercus alba L.), bur (Q.
macrocarpa Michx.), and chestnut oaks (Q. prinus L.).

Distribution This bug occurs from Alabama and the Carolinas north to southern
Canada.

Images of Oak Lace Bug

Figure 1. Adult of oak lace bug, Corythucha arcuata. (Joseph Berger,

Bugwood.org)
16 Forest Pest Insects in North America:

Images of Oak Lace Bug

(continued)

Figure 2. Nymphs of oak lace bug on underside of oak leaf. (Jim Baker, North
Carolina State University, Bugwood.org)

Figure 3. Oak foliage showing damage (yellow stippling) on the upper leaf
surface from feeding of oak lace bug. (Jim Baker, North Carolina State University,
Bugwood.org)

Important Biological Control Little is known of the specific natural enemies of the oak lace bug. One
Agents Related to this generalist predator associated with the species is the mirid Deraeocoris
Pest Species nebulosus (Uhler).

Web Links for Information http://www.fs.fed.us/r8/foresthealth/pubs/oakpests/p30.html

on Oak Lace Bug
A Photographic Guide 17

Articles Horn, K.F., C.G. Wright, and M.H. Farrier. 1979. The lace bugs
(Hemiptera: Tingidae) of North Carolina and their hosts. North Carolina
Agricultural Experiment Station Technical Bulletin No. 257: 22 p.
Wheeler, A.G., Jr., B.R. Stinner, and T.J. Henry. 1975. Biology and
nymphal stages of Deraeocoris nebulosus (Hemiptera: Miridae), a
predator of arthropod pests on ornamentals. Annals of the Entomological
Society of America 68: 1063-1068.
18 Forest Pest Insects in North America:

7. Sycamore Lace Bug, Corythucha ciliata (Say) (Hemiptera: Tingidae)

Orientation to Pest Sycamore lace bug, Corythucha ciliata (Say), is a minor pest of
American sycamore (Planatus occidentalis L.) in its native range in
North America but is very important in Europe as an invasive species
on urban plane trees. It has also invaded Australia, China, Turkey, and
Chile. Adults overwinter under the bark flakes of sycamores. Feeding
activity begins in early spring at leaf expansion of host plants. Eggs are
laid on the underside of leaves. Nymphs and adults occur in colonies
on the underside of foliage where they feed by sucking sap from plant
cells. Feeding by low densities of lace bugs results in yellow stippling,
starting along the mid-veins, similar in appearance to the damage caused
by spider mites, but without webbing. Black fecal spots are also present
on the undersides of leaves infested by lace bugs. Dense populations can
cause a white or bronzed appearance to the leaves and premature leaf
drop. In the northeast United States, there are two to four generations
per year.

Hosts Commonly Attacked The principal host of sycamore lace bug is American sycamore
(P. occidentalis), but it occasionally feeds on ash (Fraxinus), hickory
(Carya), and mulberry (Morus).

Distribution This bug is native throughout the eastern USA and southern Canada. It
is also invasive in Europe, Turkey, Australia, China, and Chile. Spread
is likely due to tendency of adults to secrete themselves into crevices
for overwintering, allowing them to be easily moved in crates or cargo
between countries.

Images of
Sycamore Lace Bug

Figure 1. Adult of sycamore lace bug, Corythucha ciliate. (Louis-Michel

Nageleisen, Département de la Santé des Forêts, Bugwood.org)
A Photographic Guide 19

Images of Sycamore
Lace Bug (continued)

Figure 2. Adults of sycamore lace bug feed in groups on the undersides of

leaves (note the black fecal spots). (Jim Baker, North Carolina State University,
Bugwood.org)

Figure 3. Nymphs of sycamore lace bug. (James Solomon, USDA Forest Service,
Bugwood.org)

Figure 4. Size of mature nymphs of sycamore lace bug (black, on left) as

compared to adults. (Steven Katovich, USDA Forest Service, Bugwood.org)
20 Forest Pest Insects in North America:

Images of Sycamore
Lace Bug (continued)

Figure 5. Close up showing the black fecal spots deposited by sycamore lace bug.
(Bruce W. Kauffman, Tennessee Department of Agriculture, Bugwood.org)

Figure 6. Yellow stippling and bronzing of foliage caused by sycamore lace bug.
(Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 7. Sycamore trees bronzed by feeding of sycamore lace bug.

(Terry S. Price, Georgia Forestry Commission, Bugwood.org)
A Photographic Guide 21

Important Biological Control Little is known of the specific natural enemies of the sycamore lace bug.
Agents Related to this The generalist predatory mirid Deraeocoris nebulosus (Uhler) is known
Pest Species to feed on sycamore lace bugs.

Web Links for Information http://entnemdept.ufl.edu/creatures/trees/sycamore_lace_bug.htm; fact

on Sycamore Lace Bug sheet of the University of Florida.

Articles Halbert, S.E. and J.R. Meeker. 1998. The Sycamore Lace Bug,
Corythucha ciliata (Say) (Hemiptera: Tingidae). Entomology Circular
No.387. Florida Dept. Agric. & Consumer Services, Division of Plant
Industry. (Available at http://www.fl-dof.com/publications/fh_pdfs/
Sycamore%20Lace%20Bug.pdf).
Wheeler, A.G., Jr., B.R. Stinner, and T.J. Henry. 1975. Biology and
nymphal stages of Deraeocoris nebulosus (Hemiptera: Miridae), a
predator of arthropod pests on ornamentals. Annals of the Entomological
Society of America 68: 1063-1068.
Horn, K.F., C.G. Wright, and M.H. Farrier. 1979. The lace bugs
(Hemiptera: Tingidae) of North Carolina and their hosts. North Carolina
Agricultural Experiment Station Technical Bulletin No. 257: 22 p.
22 Forest Pest Insects in North America:

8. Shield-backed Pine Seed Bug, Tetyra bipunctata (Herrich-Schäffer)

(Hemiptera: Scutellaridae)

Orientation to Pest The shield-backed pine seed bug, Tetyra bipunctata (Herrich-Schäffer),
feeds on seeds in cones of various pines and thus causes losses in seed
orchards. There is one generation a year, and nymphs and adults feed in
groups. Most damage to seeds is done in late summer or fall and is not
visible, even after the seeds have been extracted from cones.

Hosts Commonly Attacked This insect feeds on various pines, including loblolly (Pinus taeda L.),
slash (Pinus elliottii Engelm.), shortleaf (Pinus echinata Mill.), Virginia
(Pinus virginiana Mill.), eastern white (Pinus strobus L.), red (Pinus
resinosa Sol. ex Aiton), jack (Pinus banksiana Lamb.), and sand (Pinus
clausa [Chapm. ex Engelm.] Sarg.) pines.

Distribution The shield-backed pine seed bug occurs from Virginia north to Ontario
and Quebec and west to Minnesota and Oklahoma.

Images of the Figure 1. Adult shield-

backed pine seed bug,
Shield-backed Pine Seed Bug Tetyra bipunctata.
(John Maxwell, Marlton, NJ,
USA, Bugguide.net)

Figure 2. Eggs of shield-

backed pine seed bug.
(Texas Forest Service
Archive, Texas Forest
Service, Bugwood.org)
A Photographic Guide 23

Images of the Shield-backed

Pine Seed Bug (continued)

Figure 3. Nymph of shield-backed pine seed bug. (USDA Forest Service -

Northeastern Area Archive, USDA Forest Service, Bugwood.org)

Figure 4. Shield-backed pine seed bug adults feeding on cones. (Larry R. Barber,
USDA Forest Service, Bugwood.org)

Important Biological Control Specific natural enemies of this species are unknown.
Agents Related to this
Pest Species

Web Links for Information http://www.forestpests.org/nursery/seedandcone.html; fact sheet “Seed

on Shield-backed and Cone Insects” by Larry R. Barber, Entomologist, Region 8, USDA
Pine Seed Bug Forest Service, Asheville, North Carolina.
24 Forest Pest Insects in North America:

Articles on Shield-backed DeBarr, G.L. 1979. Importance of the seedbugs Leptoglossus corculus
Pine Seed Bug (Say) (Hemiptera: Coreidae) and Tetyra bipunctata (H.-S.) (Hemiptera:
Pentatomidae) and their control in southern pine seed orchards.
Proceedings: a symposium on flowering and seed development in trees.
Starkville, Mississippi: Southern Forest Experiment Station: 330-341.

Gilbert, B.L., S.J. Barras, and D.M. Norris. 1967. Bionomics of Tetyra
bipunctata (Hemiptera: Pentatomidae: Scutellerinae) as associated with
Pinus banksiana in Wisconsin. Annals of the Entomological Society of
America 60: 698-701.

Goyer, R.A. and V.G. Williams.1981. The effects of feeding by

Leptoglossus corculus (Say) and Tetyra bipunctata (Herrich and
Schaffer) on loblolly pine (Pinus taeda L.) conelets. Journal of Georgia
Entomological Society 16: 16-21.

Turgeon, J.J., P. de Groot, and J.D. Sweeney. 2005. Insects of seed cones
in eastern Canada; field guide. Ontario Ministry of Natural Resources/
Forestry Canada, Toronto: 127 p.
A Photographic Guide 25

9. Leaffooted Pine Seed Bug, Leptoglossus corculus (Say) (Hemiptera:

Coreidae)

Orientation to Pest The leaffooted pine seed bug, Leptoglossus corculus (Say), is a sucking
insect that damages cones in pine seed orchards in the southern United
States. Both adults and nymphs fed on seeds in cones of various pines,
greatly reducing production of viable seed. There are several generations
per year in the southern United States.

Hosts Commonly Attacked The leaffooted pine seed bug commonly attacks cones of loblolly
(Pinus taeda L.), slash (P. elliottii Engelm.), shortleaf (P. echinata Mill.),
Virginia (P. virginiana Mill.), eastern white (P. strobus L.), pitch
(P. rigida Mill.) and Table Mountain (P. pungens Lamb.) pines, as well
as cones of some spruce (Picea).

Distribution The leaffooted pine seed bug occurs throughout the eastern United
States.

Images of
Leaffooted Pine Seed Bug

Figure 1. Adult leaffooted pine seed bug, Leptoglossus corculus. See the flat,
expanded lower section of hind legs. (R. Scott Cameron, Advanced Forest
Protection, Inc., Bugwood.org)

Figure 2. String of eggs of leaffooted pine seed bug. (Lacy L. Hyche, Auburn
University, Bugwood.org)
26 Forest Pest Insects in North America:

Images of Leaffooted
Pine Seed Bug (continued)

Figure 3. Nymph of leaffooted pine seed bug. (Tim Tigner, Virginia Department of
Forestry, Bugwood.org)

Figure 4. Leaffooted pine seed bug adult feeding on immature cone of pine.
(Larry R. Barber, USDA Forest Service, Bugwood.org)

Important Biological Control The parasitoid Ooencyrtus leptoglossi Yoshimoto is known to attack the
Agents Related to this eggs of this species.
Pest Species

Web Links for Information http://www.forestpests.org/nursery/seedandcone.html; fact sheet “Seed

on Leaffooted Pine Seed Bug and Cone Insects” by Larry R. Barber, Entomologist, Region 8, USDA
Forest Service, Asheville, North Carolina.

Articles DeBarr, G.L. 1979. Importance of the seedbugs Leptoglossus corculus

(Say) (Hemiptera: Coreidae) and Tetyra bipunctata (H.-S.) (Hemiptera:
Pentatomidae) and their control in southern pine seed orchards.
Proceedings: a symposium on flowering and seed development in trees.
Starkville, Mississippi. Southern Forest Experiment Station: 330-341.
A Photographic Guide 27

10. Royal Palm Bug, Xylastodoris luteolus Barber (Hemiptera:

Thaumastocoridae)

Orientation to Pest Royal palm bug, Xylastodoris luteolus Barber, is a sucking bug that feeds
on immature leaves of royal palm (Roystonea regia [Kunth] O.F. Cook).
If densities are high, this feeding may cause browning of affected foliage.

Hosts Commonly Attacked This sucking bug feeds on royal palm (R. regia), sometimes called
Florida royal palm or Cuban royal palm.

Distribution This insect’s native range is Cuba, but it also occurs in southern Florida.

Images of
Royal Palm Bug

Figure 1. Adults of royal palm bug, Xylastodoris luteolus. (Lyle Buss, University of
Florida, Bugwood.org)

Figure 2. Close view of damage to Florida royal palms by the royal palm bug.
(Doug Caldwell, University of Florida, Bugwood.org)
28 Forest Pest Insects in North America:

Images of Royal Figure 3. Damage to Florida

royal palm frond by the royal
Palm Bug (continued) palm bug. (Doug Caldwell,
University of Florida,
Bugwood.org)

Figure 4. Whole tree view of damage to Florida royal palm frond by the
royal palm bug. (Both photos: Dr. A. D. Ali, Davey Tree Expert Company,
Bugwood.org)

Important Biological Control Specialized natural enemies of this species are unknown.
Agents Related to this
Pest Species

Web Links for Information http://entnemdept.ufl.edu/creatures/orn/palms/royal_palm_bug.htm

on Royal Palm Bug http://edis.ifas.ufl.edu/in254; fact sheet of University of Florida IFAS
Extension.

Articles Weissling, T.J., F.W. Howard, and A.W. Meerow. Royal palm bug,
Xylastodoris luteolus Barber (Insecta: Hemiptera: Thaumastocoridae).
EENY-097 University of Florida IFAS Extension. (Available at http://
edis.ifas.ufl.edu/in254).
Ali, A.D. and D. Caldwell. 2010. Royal palm bug Xylastodoris luteolus
(Hemiptera: Thaumastocoridae) control with soil-applied systemics.
Florida Entomologist 93: 294-297.
A Photographic Guide 29

11. Saratoga Spittlebug, Aphrophora saratogensis (Fitch) (Hemiptera:

Cercopidae)

Orientation to Pest Saratoga spittlebug, Aphrophora saratogensis (Fitch), is a native insect

in North America whose adults damage certain pines, especially red
pine (Pinus resinosa Sol. ex Aiton), usually when grown in plantations.
Damage also occurs on jack (P. banksiana Lamb.) and Scots pines
(P. sylvestris L.). Adult feeding damages pines because toxics injected
into the plant kill tissues, causing dead pockets of xylem and phloem
tissue. Extensive feeding kills branches, stunts and deforms shoots,
and may sometimes kill trees. This species is generally considered
the most serious sap-feeding pest of red pine plantations. In contrast,
nymphs are not pests because they feed on different plants. Young
nymphs feed on understory plants such as brambles (species of Rubus),
orange hawkweed (Pilosella aurantiaca [L.] F.W. Schultz and Schultz-
Bip), pearly everlasting (Anaphalis nubigena DC.), and asters (Aster),
while older nymphs are found on sweetfern (Comptonia peregrine [L.]
J.M. Coulter) and young willows (Salix). Saratoga spittlebug has one
generation each year. On red pine, eggs are laid under the outer scales of
buds in the upper branches. Selection criteria have been developed that
use information on nymphal host plant abundance to pick sites for new
red pine plantations that will have low risk of damage from Saratoga
spittlebug. A site-risk diagram has been developed for this purpose (see
figure in USDA Forest Service leaflet at http://www.na.fs.fed.us/spfo/
pubs/fidls/saratoga/saratoga.htm).

Hosts Commonly Attacked Saratoga spittlebug is known as a pest of some pines, especially red pine
(P. resinosa) grown in plantations.

Distribution This insect occurs where ever its hosts grow, from Maine to Minnesota in
the United States and in the southern portions of the adjacent Canadian
Provinces. It is of particular importance in Michigan and Wisconsin.

Images of
Saratoga Spittlebug

Figure 1a. Adults of Saratoga spittlebug, Aphrophora saratogensis.

(Pennsylvania Department of Conservation and Natural Resources - Forestry
Archive, Bugwood.org)
30 Forest Pest Insects in North America:

Images of Saratoga Figure 1b. Adult Saratoga

spittlebug, Aphrophora
Spittlebug (continued) saratogensis. (Tom Murray)

Figure 2. Eggs of Saratoga spittlebug (dark bodies in woolly mass).

(James B. Hanson, USDA Forest Service, Bugwood.org)

Figure 3. Spittle masses of Saratoga spittlebugs at base of stems of sweetfern,

the favorite host of older nymphs. (Linda Haugen, USDA Forest Service,
Bugwood.org)
A Photographic Guide 31

Images of Saratoga
Spittlebug (continued)

Figure 4. Saratoga spittlebug feeding damage to red pine (Both photos: USDA
Forest Service - Northeastern Area Archive, USDA Forest Service, Bugwood.org)

Important Biological Control Eggs of Saratoga spittlebug are attacked by two parasitoids, Ooctonus
Agents Related to this aphrophorae Milliron and Tumidiscapus cercopiphagus Milliron, but
Pest Species attack rates are low (<5 percent). A fly in the family Pipunculidae,
Verrallia virginica Banks, may be a significant source of mortality
(causing up to 50 percent mortality).

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/saratoga/saratoga.htm; a USDA

on Saratoga Spittlebug Forest Service pest leaflet.
http://www.maine.gov/doc/mfs/saratoga.htm; Maine fact sheet.

Articles Wilson, L.F. 1971. Risk-rating Saratoga spittlebug damage by abundance

of alternate-host plants. Research Note NC-110, North Central Forest
Experiment Station, Forest Service, United States Department of
Agriculture: 4 p.
Linnane, J.P. and E.A. Osgood. 1977. Verrallia virginica (Diptera:
Pipunculidae) reared from the Saratoga spittlebug in Maine. Proceedings
of the Entomological Society of Washington 79(4): 622-623.
32 Forest Pest Insects in North America:

12. Pine Spittlebug, Aphrophora parallela (Say) (Hemiptera: Cercopidae)

Orientation to Pest Pine spittlebug, Aphrophora parallela (Say), is a native insect in North
America that is found in most of the eastern United States and adjacent
southern Canada. Adults and nymphs develop on the same plant species.
The most favored host is Scots pine (Pinus sylvestris L.). Eggs are
deposited under bark and are the overwintering stage. Nymphs initially
feed on twigs, but later move to the trunk, forming protective spittle
masses each time they move. Adults return to twigs to feed. There is
a single generation per year. Feeding of this spittlebug promotes the
spread of a pathogenic fungus, Sphaeropsis sapinea (synonyms S. ellisii
and Diplodia pinea), which invades weakened trees through spittlebug
feeding punctures. This fungus is responsible for a considerable part of
the dead branches associated with this insect.

Hosts Commonly Attacked Pine spittlebugs seem to prefer Scots pine but are also found on six
other pines, three species of spruce (Picea), balsam fir (Abies balsamea
[L.] Miller), larch (Larix), and eastern hemlock (Tsuga canadensis (L.)
Carrière).

Distribution Pine spittlebug is found throughout the eastern United States and
adjacent parts of southern Canada.

Images of Pine
Spittlebug

Figure 1. Adult of pine spittlebug, Aphrophora parallela. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Figure 2. Spittle mass pushed aside to show nymph of pine spittlebug.

(Lacy L. Hyche, Auburn University, Bugwood.org)
A Photographic Guide 33

Important Biological Control The fungus Entomophora aphrophorae Rostrup at times causes high
Agents Related to this mortality to nymphs.
Pest Species

Web Links for Information http://wiki.bugwood.org/Aphrophora_parallela; short fact sheet by

on Pine Spittlebug BugwoodWiki.
http://www.entomology.umn.edu/cues/Web/199Spittlebugs.pdf; fact
sheet with a comparison among spittlebug species.
http://ipm.illinois.edu/diseases/rpds/625.pdf; report on Sphaeropsis blight
or Diplodia tipblight of pines.
http://www.maine.gov/doc/mfs/pinespit.htm; Maine government fact
sheet.

Articles Wilson, L.F. and D.G. Mosher. 1981. How to identify and control pine
spittlebug [Aphrophora parallela]. North Central Forest Experiment
Station, USDA Forest Service, St. Paul, Minnesota: 6 p.
34 Forest Pest Insects in North America:

13. Thorn Bug, Umbonia crassicornis (Amyot and Serville) (Hemiptera:

Membracidae)

Orientation to Pest The thorn bug Umbonia crassicornis (Amyot and Serville) is a sucking
bug that is an occasional pest of ornamentals and fruit trees in southern
Florida. Colonies of these insects may be very visible on the twigs of
their hosts. Feeding may lead to defoliation or twig death in some species
of plants. Also, thorn bugs produce honeydew, which leads to growth of
sooty mold. Eggs are laid in bark of tender twigs and, most interestingly
for insects, females show maternal care by actively tending their
offspring. Young nymphs have three horns instead of the one seen on the
adults. Colonies of insects are chemically defended against predators.

Hosts Commonly Attacked Among plants that suffer important injury when infested with thorn bugs
are species of Hibiscus, powder-puff (Calliandra), woman’s tongue
tree (Albizzia lebbek [L.] Benth.), and species of Acacia. Other hosts
suitable for reproduction include wild tamarind (Lysiloma bahamensis),
tamarind (Tamarindus indica L.), Casuarina sp., Crotalaria sp., rayado
bundleflower (Desmanthus virgatus Benth.), bottle brush (Callistemon
sp.), Jerusalem thorn (Parkinsonia aculeate L.), and dwarf date palm
(Phoenix roebelenii O’Brien).

Distribution This thorn bug is subtropical or tropical and found in Mexico, Central
and South America. In the United States, it is found only in southern and
central Florida.

Images of Thorn Bug

Figure 1. Adult of the thorn bug Umbonia crassicornis. (Lyle Buss, University of
Florida, Bugwood.org)
A Photographic Guide 35

Images of Thorn Bug

(continued)

Figure 2. Adults of this thorn bug are variable in terms of the shape and size of
their horn. (Lyle Buss, University of Florida, Bugwood.org)

Important Biological Control No information is available on specialized natural enemies of this

Agents Related to this species.
Pest Species

Web Links for Information http://edis.ifas.ufl.edu/pdffiles/IN/IN33200.pdf; fact sheet of the

on Thorn Bug University of Florida.

Articles Wood, T.K. 1976. Alarm behavior of brooding female Umbonia

crassicornis (Homoptera: Membracidae). Annals of the Entomological
Society of America 69: 340-344.
36 Forest Pest Insects in North America:

14. Periodical Cicada, Magicicada septendecim L. (Hemiptera: Cicadidae)

Orientation to Pest The periodical cicada, Magicicada septendecim L., is the largest of
periodically emerging cicada species (adult, 40 mm in length). Females
are black on top and males are black with a pattern of 4 or 5 orange-
brown abdominal segments when viewed from below. The female has a
stout ovipositor that is used to insert eggs into the bark of small branches
and twigs. Eggs are laid in batches of 25-30, and many batches may
be laid in the same twig. After eggs hatch, nymphs drop to the ground
and burrow into the soil, where they feed on the plant’s roots. Nymphs
require 17 years to complete their development. Emergence of last instar
nymphs is synchronous, producing mass swarms of adults at 17-year
intervals. Broods (populations in different areas) are not necessarily in
synchrony. Effect of nymphal feeding on roots has not been quantified.
The only significant injury caused by this insect is the dieback of small
branches (“flagging”) from the cuts made for oviposition. While this
injury can be highly visible it is not harmful to larger trees and shrubs
since it only occurs every 17 years and large plants can tolerate this level
of damage. On smaller trees, however, extensive oviposition may be
harmful. Homeowners who need to protect small trees and shrubs from
damage can do so by covering plants with netting to exclude cicadas,
which should be left in place until most cicadas have died off (about
four weeks from their first appearance). Periodical cicadas are often
too numerous to make applications of pesticides practical. The mass
emergences of this insect are much appreciated as a wildlife phenomenon
and this species is best viewed as an impressive act of nature rather than
as a pest.

Hosts Commonly Attacked This cicada lays its eggs in more than 70 species of trees or other
plants, but the most affected trees are oaks (Quercus), hickory (Carya),
honeylocust (Gleditsia triacanthos L.), dogwood (Cornus), apple (Malus
domestica Borkh.), and peach (Prunus persica [L.] Batsch.). But many
other species may also be attacked at times.

Distribution Magicicada septendecim is widely distributed in the eastern United

States.
A Photographic Guide 37

Images of the
Periodical Cicada

Figure 1. Adult of the periodical cicada, Magicicada septendecim, a 17-yr cicada.

(John H. Ghent, USDA Forest Service, Bugwood.org)

Figure 2. Oviposition scars made by the periodical cicada in a branch.

(John H. Ghent, USDA Forest Service, Bugwood.org)

Figure 3. Nymph of periodical cicada, the below-ground life stage. (Bill Gallup,
Bugwood.org)
38 Forest Pest Insects in North America:

Images of the Periodical

Cicada (continued)

Figure 4. Holes in soil made by emerging last instar nymphs of the periodical
cicada. (Jim Occi, BugPics, Bugwood.org)

Figure 5. Adult of a periodical cicada as it begins to molt from the last nymphal
instar after its emergence from the soil. (John H. Ghent, USDA Forest Service,
Bugwood.org)

Figure 6. Cast skins of periodical cicada nymphs produced during the insect’s
mass emergence. (Jim Occi, BugPics, Bugwood.org)
A Photographic Guide 39

Images of the Periodical Figure 7. Flagging of

branches due to oviposition
Cicada (continued) wounds of periodical cicada.
(John H. Ghent,
USDA Forest Service,
Bugwood.org)

Figure 8. Close up of branch flagging from oviposition wounds of periodical

cicada. (Jim Occi, BugPics, Bugwood.org)

Important Biological Control Almost everything seems to eat periodical cicadas including birds,
Agents Related to this squirrels, snakes, spiders, etc. The massive populations of periodical
Pest Species cicadas, however, are able to satiate such predators and they appear to
have little effect in reducing cicada densities.
40 Forest Pest Insects in North America:

Web Links for Information http://insects.ummz.lsa.umich.edu/fauna/michigan_cicadas/periodical/

on the Periodical Cicada index.html; discusses the various species of periodical cicadas.
http://www.magicicada.org/about/species_pages/m_sdecim.php; has
sound files of cicada songs.

Articles Miller, F.D. 1997. Effects and control of periodical cicada Magicicada
septendecim and Magicicada cassini oviposition injury on urban forest
trees. Journal of Arboriculture 23(6): 225-232.
Miller, F. and W. Crowley. 1998. Effects of periodical cicada
ovipositional injury on woody plants. Journal of Arboriculture 24(5):
248-253.
A Photographic Guide 41

15. Eastern Spruce Gall Adelgid, Adelges abietis (L.) (Hemiptera: Adelgidae)

Orientation to Pest Eastern spruce gall adelgid, Adelges abietis (L.), is a pest of the
introduced Norway spruce (Picea abies [L.]) and of various native
North American spruces, especially white spruce (P. glauca [Moench]).
It is an invasive species believed to be of European origin. Damage is
caused by induction of galls on new growth. Adults, which are wingless,
deposit their eggs near buds in spring and newly developing needles
are stimulated by developing nymphs to become enlarged, fusing into
a pineapple-shaped gall, within which nymphs mature. In summer
and early fall, the chambers of the gall open and nymphs emerge and
transform themselves into winged adults, which lay eggs. These hatch
and nymphs overwinter at the base of needles and buds. In spring, when
the gall is newly formed, it is light green in color, but after it matures and
insects emerge, it dries and turns brown, remaining on the branch as a
sign of earlier infestation. Damage results from twig deaths induced by
galls, which makes the tree misshapen. Heavily infested trees may die.

Hosts Commonly Attacked This adelgid feeds on black (P. mariana (Mill.) Britton, Sterns &
Poggenburg), Norway (P. abies), and white (P. glauca) spruces.

Distribution Eastern spruce gall adelgid occurs widely in the eastern United States
from Maine to Tennessee and Illinois, as well as in northern Minnesota,
Wisconsin, and Michigan, and adjacent parts of Canada. Localized
infestations also occur in parts of Montana, Wyoming, and South Dakota.
A map of this distribution is available at http://www.fs.fed.us/nrs/tools/
afpe/maps/ESGA.pdf.

Images of Eastern
Spruce Gall Adelgid

Figure 1. Close view of living galls of eastern spruce gall adelgid, Adelges abietis.
(E. Bradford Walker, Vermont Department of Forests, Parks and Recreation,
Bugwood.org)
42 Forest Pest Insects in North America:

Images of Eastern Spruce Figure 2. Living galls of

eastern spruce gall adelgid
Gall Adelgid (continued) cut in cross section and
showing chambers where
insects feed. (Thérèse
Arcand - Natural Resources
Canada, Canadian Forest
Service)

Figure 3. Dead galls of eastern spruce gall adelgid. (E. Bradford Walker, Vermont
Department of Forests, Parks and Recreation, Bugwood.org)

Figure 4. Openings in gall of eastern spruce gall adelgid after insect emergence.
(Stanislaw Kinelski, Bugwood.org)
A Photographic Guide 43

Images of Eastern Spruce Figure 5. Adult of eastern

spruce gall adelgid,
Gall Adelgid (continued) Adelges abietis. (Thérèse
Arcand - Natural Resources
Canada, Canadian Forest
Service)

Important Biological Control Because damage from this species is not severe, natural enemies have
Agents Related to this not been imported from the pest’s native range into North America for
Pest Species control. Predacious flies have been observed feeding on this adelgid in
Europe (the syphid Cnemodon sp.) and, in Turkey, Leucopis species have
been recorded feeding on related adelgids, but in general little is known
about the natural enemies of this species.

Web Links for Information on http://www.ravageursexotiques.gc.ca/ins_details_eng.asp?pestType=ins&

Eastern Spruce Gall Adelgid lang=en&geID=5306; website of Environment Canada on invasive forest
insects.
http://ohioline.osu.edu/hyg-fact/2000/2550.html; fact sheet of Ohio State
University, includes information on control.

Articles Flaherty, L., D. Quiring, and J.A. Kershaw, Jr. 2010. Post-gall induction
performance of Adelges abietis (L.) (Homoptera: Adelgidae) is
influenced by clone, shoot length, and density of colonising gallicolae.
Ecological Entomology 35: 9-15.
44 Forest Pest Insects in North America:

16. Balsam Woolly Adelgid, Adelges piceae (Ratzeburg) (Hemiptera:

Adelgidae)

Orientation to Pest Balsam woolly adelgid, Adelges piceae (Ratzeburg), attacks only true
firs (Abies). It is native to Eurasia and is a serious invasive pest in
both eastern and western North America. Balsam woolly adelgids are
small, <1 mm in size, and covered with white, woolly wax. Adults
and nymphs feed on both the bole and twigs and inject hormones with
their saliva that causes gouty twigs and growth of compression wood
on the bole. These deformities disrupt water conduction and reduce
tree growth and market value. Heavy infestations eventually kill the
tree. Extensive damage to Fraser fir (Abies fraseri [Pursh] Poir.) in the
southern Appalachian Mountains caused severe ecological damage to
unique mountain-top habitats created by stands of this species. Damage
to balsam (A. balsamea (L.) Mill.) and western firs has caused significant
economic losses due to a combination of elevated rates of tree mortality
and reduced growth. Various predators imported and released during the
1950s from the European Alps failed to bring the pest under biological
control. However, it is possible that the Alps themselves were an invaded
area and that the true native range is the Caucasus Mountains. Thus, this
region has potential as a source of effective natural enemies for this pest.

Hosts Commonly Attacked The host plants of balsam woolly adelgid are Fraser fir (A. fraseri),
balsam fir (A. balsamea), subalpine fir (A. lasiocarpa [Hooker] Nuttall),
Pacific silver fir (Abies amabilis Douglas ex J. Forbes), and grand fir
(A. grandis [Douglas ex D. Don] Lindley).

Distribution Balasm woolly adelgid is found in New York, New England, the
Canadian Maritimes, British Columbia, the Pacific northwestern U.S.
states, and limited areas of the southern Appalachian Mountains and the
central coast of California.

Figure 1. Distribution of
balsam woolly adelgid,
Adelges piceae, in North
America. (USDA Forest
Service, Forest Insect
& Disease Leaflet 118
revised May 2006)
A Photographic Guide 45

Images of Balsam
Woolly Adelgid

Figure 2. Bole of fir tree infested with Figure 3. Close view of balsam woolly
balsam woolly adelgid. (USDA Forest adelgids on bark of fir tree. (Ladd
Service - Ashville Archive, USDA Forest Livingston, Idaho Department of Lands,
Service, Bugwood.org) Bugwood.org)

Figure 4. High magnification of crawler (first instar) of balsam woolly adelgid.

(USDA Forest Service - Ashville Archive, USDA Forest Service, Bugwood.org)
46 Forest Pest Insects in North America:

Images of Balsam
Woolly Adelgid

Figure 5. Goutiness of Fraser fir twigs due to feeding by balsam woolly adelgid.
(William M. Ciesla, Forest Health Management International, Bugwood.org)

Figure 6. Fraser fir trees killed by balsam woolly adelgid. (Robert L. Anderson,
USDA Forest Service, Bugwood.org)

Important Biological Control More than 23 species of natural enemies were imported and released
Agents Related to this into the United States against this pest. Six species established, but
Pest Species remain rare: Aphidecta obliterata (L.) (Coleop.: Coccinellidae),
Scymnus impexus (Mulsant) (Coleop.: Coccinellidae), Laricobius
erichsonii (Rosenhauer) (Coleop.: Derodontidae), Aphidoletes thompsoni
Möhn (Diptera: Cecidomyiidae), Cremifania nigrocellulata Czerny
(Diptera: Chamaemyiidae), and Leucopis obscura (Hal.) (Diptera:
Chamaemyiidae).
A Photographic Guide 47

Web Links for Information http://www.na.fs.fed.us/pubs/fidls/bwa.pdf; U.S. Department of

on Balsam Woolly Adelgid Agriculture, Forest Service Forest Insect & Disease Leaflet 118 (Revised
May 2006).
http://pubs.ext.vt.edu/3006/3006-1452/3006-1452.pdf; Virginia
Cooperative Extension bulleting on balsam woolly adelgid by Scott
Salom and Eric Day.
http://www.ces.ncsu.edu/fletcher/programs/xmas/ctnotes/ctn020.html;
North Carolina State University Cooperative Extension (a short bulletin
discussing how to manage this pest in Christmas trees).

Articles Harris, J.W.E. and A.F. Dawson. 1979. Predator release program for
balsam woolly aphid, Adelges piceae (Homoptera: Adelgidae), in British
Columbia, 1960-1969. Journal of the Entomological Society of British
Columbia 76: 21-26.
Schooley, H.O. and L. Oldford. 1981. An annotated bibliography of the
balsam woolly aphid (Adelges piceae [Ratzeburg]). Information Report
N-X-196, Canadian Forestry Service: 97 p.
Schooley, H.O., J.W.E. Harris, and B. Pendrel. 1984. Adelges piceae
(Ratz.) balsam woolly adelgid (Homoptera: Adelgidae). In: Kelleher, J.S.
and M.A. Hulme (eds.). Biological Control Programmes against Insects
and Weeds in Canada, 1969-1980. Commonwealth Agricultural Bureaux,
Farnham Royal, England: 229-234.
McManamay, R.H., L.M. Resler, J.B. Campbell, and R.A. McManamay.
2011. Assessing the impacts of balsam woolly adelgid (Adelges piceae
Ratz.) and anthropogenic disturbance on the stand structure and mortality
of Fraser fir [Abies fraseri (Pursh) Poir.] in the Black Mountains, North
Carolina. Castanea 76(1): 1-19.
48 Forest Pest Insects in North America:

17. Hemlock Woolly Adelgid, Adelges tsugae Annand (Hemiptera:

Adelgidae)

Orientation to Pest Hemlock woolly adelgid, Adelges tsugae Annand, is an invasive insect
in eastern North America. DNA evidence suggests that the invasive
eastern U.S. population came from Japan and not the western United
States, where the species is also present. Hemlock woolly adelgids are
small in size and to the naked eye only their woolly coverings are easily
visible. The insect has two generations per year and growth occurs from
fall through late spring. Insects in summer are inactive and scarcely
visible at the bases of needles as black dots. Woolly masses (the sign
allowing the species to be recognized) develop in October and are
present thereafter through June of the following year. In the western
United States, A. tsugae feeds on western hemlock (Tsuga heterophylla
[Raf.] Sarg.) but is believed to be native and is not a pest. In the eastern
USA, hemlock woolly adelgid is killing eastern (T. canadensis [L.]
Carrière) and Carolina (T. caroliniana Engelm.) hemlocks in large
numbers from Connecticut south along the Appalachian Mountains.
From Massachusetts north, or at high elevations, tree mortality has been
restrained by higher rates of mortality of adelgids in winter due to low
temperatures. A biological control program is in progress against this
pest, based on specialized predatory beetles that feed only on adelgids,
collected in western North America (Laricobius nigrinus Fender) or
China/Japan (species of Laricobius and various Scymnus ladybird
beetles). To date, releases have not demonstrated any reductions in
adelgid densities from predators. Populations of L. nigrinus have become
well established and abundant in some areas of western North Carolina.

Hosts Commonly Attacked In North America, this species feed only on eastern hemlock
(T. canadensis), Carolina hemlock (T. caroliniana), and western hemlock
(T. heterophylla).

Distribution The hemlock woolly adelgid is found as a native species in the western
United States in the Pacific northwestern states (Oregon, Washington,
Idaho, and Montana) and as an invasive species from Japan in the eastern
United States, from southern Maine to northern Georgia.

Figure 1. Distribution of hemlock

woolly adelgid, Adelges tsugae,
in the eastern United States in 2009.
(USDA Forest Service, Northeastern
Area, Forest Health Protection -
Distribution Maps)
A Photographic Guide 49

Images of
Hemlock Woolly Adelgid

Figure 2. Hemlock woolly adelgid on hemlock. (Michael Montgomery, USDA

Forest Service, Bugwood.org)

Figure 3. Individuals of hemlock woolly adelgid covered with white wool (see
crawler emerging, arrow). (Pennsylvania Department of Conservation and
Natural Resources - Forestry Archive, Bugwood.org)

Figure 4. High
magnification of the
aphid-like body of an adult
hemlock woolly adelgid
after removal of the wool.
(Michael Montgomery,
USDA Forest Service,
Bugwood.org)
50 Forest Pest Insects in North America:

Images of Hemlock
Woolly Adelgid (continued)

Figure 5. Eggs of hemlock woolly adelgid (wool pulled aside for viewing).
(Lorraine Graney, Bartlett Tree Experts, Bugwood.org)

Figure 6. Damage from hemlock woolly adelgid is seen as needle loss, leading to
branch death. (Chris Evans, River to River CWMA, Bugwood.org)
Figure 7. Dead or dying hemlock in
Joyce Kilmer area, North Carolina due
to hemlock woolly adelgid feeding.
(Jason VanDriesche, Bugwood.org)
A Photographic Guide 51

Images of Hemlock
Woolly Adelgid (continued)

Figure 8. Laricobius nigrinus, which feeds only on adelgids, is the most promising
agent released so far against hemlock woolly adelgid. (Ashley Lamb, Virginia
Polytechnic Institute and State University, Bugwood.org)

Important Biological Control There is an on going classical biological control project against this
Agents Related to this pest. As of 2010, the principal species introduced against this pest have
Pest Species been the following four species: Sasajiscymnus tsugae (Sasaji and
McClure) (Coleop.: Coccinellidae), Scymnus sinuanodulus Yu and Yao
(Coleop.: Coccinellidae), Scymnus ningshanensis Yu and Yao (Coleop.:
Coccinellidae), and Laricobius nigrinus Fender (Coleop.: Derodontidae).

Web Links for Information http://na.fs.fed.us/fhp/hwa/biology/biological_control.shtm; information

on Hemlock Woolly Adelgid on hemlock woolly adelgid’s biology, impacts, distribution, and control
(chemical and biological).
http://www.na.fs.fed.us/spfo/pubs/pest_al/hemlock/hwa05.htm; a brief
printable summary of life history and control;
http://www.dcnr.state.pa.us/forestry/woollyadelgid/index.aspx; contains
an annual life cycle diagram.

Articles Zilahi-Balogh, G.M.G., L.T. Kok, and S.M. Salom. 2002. Host
specificity of Laricobius nigrinus Fender (Coleoptera: Derodontidae), a
potential biological control agent of the hemlock woolly adelgid, Adelges
tsugae Annand (Homoptera: Adelgidae). Biological Control 24: 192-198.
Zilahi-Balogh, G.M.G., S.M. Salom, and L.T. Kok. 2003. Development
and reproductive biology of Laricobius nigrinus, a potential biological
control agent of Adelges tsugae. BioControl 48: 293-306.
Mausel, D.L., S.M. Salom, L.T. Kok, and G.A. Davis. 2010.
Establishment of the hemlock woolly adelgid predator, Laricobius
nigrinus (Coleoptera: Derodontidae), in the eastern United States.
Environmental Entomology 39: 440-448.
52 Forest Pest Insects in North America:

Articles on Krapfl, K.J., E.J. Holzmueller, and M.A. Jenkins. 2011. Early impacts of
Hemlock Woolly Adelgid hemlock woolly adelgid in Tsuga canadensis forest communities of the
(continued) southern Appalachian Mountains. Journal of the Torrey Botanical Society
138: 93-106.
Montgomery, M.E., S. Shiyake, N.P. Havill, and R.A.B. Leschen, 2011.
A new species of Laricobius (Coleoptera: Derodontidae) from Japan
with phylogeny and a key for native and introduced congeners in North
America. Annals of the Entomological Society of America 104: 389-401.
A Photographic Guide 53

18. Cooley Spruce Gall Adelgid, Adelges cooleyi (Gillette) (Hemiptera:

Adelgidae)

Orientation to Pest Cooley spruce gall adelgid, Adelges cooleyi (Gillette), is native to
the Rocky Mountain region of North America, where it galls various
species of spruce (Picea). It is also found in Europe, where it was
accidentally introduced through movement of North American conifers
for use in plantations, and where it is a pest in plantations of Douglas-fir
(Pseudotsuga menziesii [Mirbel] Franco). In North America, damage
occurs principally on landscape trees and in plantations. Damage is
not important in native forests. Galls are formed on the apical portion
of shoot, in contrast to eastern spruce gall adelgid (Adelges abietis
[L.]) galls that are found on the basal portion of the twig. Damage is
merely aesthetic, but may be economically significant in Christmas tree
plantations.

Hosts Commonly Attacked The most important host is blue spruce (Picea pungens Engelm.),
but other species attacked include white spruce (P. glauca [Moench]
Voss), Sitka spruce (P. sitchensis [Bong.] Carr.), Engelmann spruce
(P. engelmannii Parry ex Engelm.), and as an alternate host, Douglas-fir
(P. menziesii)

Distribution This species occurs from coast to coast in the northern United States and
throughout the range of white spruce in Canada. Populations occur in the
eastern United States where blue spruce is planted as an ornamental.

Images of Cooley Spruce

Gall Adelgid

Figure 1. Appearance of the large (2-3 cm long) galls caused by Cooley spruce
gall adelgid, Adelges cooleyi. (Whitney Cranshaw, Colorado State University,
Bugwood.org)
54 Forest Pest Insects in North America:

Images of Cooley Spruce

Gall Adelgid (continued)

Figure 2. Close view of living galls of Cooley spruce gall adelgid. (Whitney
Cranshaw, Colorado State University, Bugwood.org)

Figure 3. Living galls of Cooley spruce gall adelgid cut in cross section and
showing chambers where insects feed. (Chris Schnepf, University of Idaho,
Bugwood.org)

Figure 4. Overwintering adults of Cooley spruce gall adelgid. (Whitney Cranshaw,

Colorado State University, Bugwood.org)
A Photographic Guide 55

Images of Cooley Spruce

Gall Adelgid (continued)

Figure 5. Appearance of Cooley spruce Figure 6. Heavy infestation of Cooley

gall adelgids on its alternate host, spruce gall adelgid on Christmas trees.
Douglas-fir. (Ladd Livingston, Idaho Such trees are not saleable. (Eric R.
Department of Lands, Bugwood.org) Day, Virginia Polytechnic Institute and
State University, Bugwood.org)

Important Biological Control Little to nothing is known of the natural enemies of this species.
Agents Related to this Investigations in its native range in the Rocky Mountains of North
Pest Species America may be useful as a source of natural enemies for importation to
Europe, where this species is an invasive pest.

Web Links for Information on http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=5307; website

Cooley Spruce Gall Adelgid of Environment Canada on invasive forest insects.
http://ohioline.osu.edu/hyg-fact/2000/2551.html; fact sheet of Ohio State
University that includes information on control.

Articles Lasota, J.A. and D.J. Shetlar. 1986. Assessing seasonal and spatial
abundance of Adelges cooleyi (Gillette) (Homoptera: Adelgidae) by
various sampling techniques. Environmental Entomology 15: 254-257.
Cranshaw, W.S. 1989. Patterns of gall formation by the Cooley spruce
gall adelgid on Colorado blue spruce. Journal of Arboriculture 15(11):
277-280.
56 Forest Pest Insects in North America:

19. Pine Bark Adelgid, Pineus strobi (Hartig) (Hemiptera: Adelgidae)

Orientation to Pest Pine bark adelgid, Pineus strobi (Hartig), populations are at times
dense and strikingly visible; however, this native adelgid is generally
only a minor pest of eastern white pine (Pinus strobus L.). Even dense
populations do little harm if the tree is otherwise healthy. In plantations
and on Christmas trees, pine bark adelgid infestations may reduce growth
and induce excessive branching, reducing the value of Christmas trees.
The species may have up to five generations per year.

Hosts Commonly Attacked The host of this species is eastern white pine (P. strobus).

Distribution This adelgid is found widely in North America throughout the range of
eastern white pine.

Images of
Pine Bark Adelgid

Figure 1. An infestation of pine bark Figure 2. Closer view of pine trunk

adelgid, Pineus strobi, on the trunk infested with pine bark adelgid.
of eastern white pine. (Petr Kapitola, (Steven Katovich, USDA Forest
State Phytosanitary Administration, Service, Bugwood.org)
Bugwood.org)
A Photographic Guide 57

Images of Pine Bark

Adelgid (continued)

Figure 3. Pine bark adelgids infesting pine branch. (Petr Kapitola, State
Phytosanitary Administration, Bugwood.org)

Figure 4.Excess bud formation from pine bark adelgid feeding can cause shoot
proliferation in tree top. Note also the lady beetle feeding on the adelgids. (Eric
R. Day, Virginia Polytechnic Institute and State University, Bugwood.org)

Figure 5. Laricobius rubidus

(Coleop.: Derodontidae) is an
important predator of pine bark
adelgid in the eastern United States.
(Jim McClarin, BugGuide.net)
58 Forest Pest Insects in North America:

Important Biological Control Because damage from this species is generally not important in North
Agents Related to this America, natural enemies of this species have not been well studied.
Pest Species However, some predators of this adelgid, have been noted in the eastern
United States, including the derodontid beetle Laricobius rubidus
LeConte and chamaemyiid fly Leucopis pinicola Mall.

Web Links for Information http://pubs.ext.vt.edu/2907/2907-1402/2907-1402.pdf; Virginia Tech

on Pine Bark Adelgid extension leaflet updated by Scott Salom.

Articles Sluss, T.P. and B.A. Foote. 1973. Biology and immature stages
of Leucopis pinicola and Chamaemyia polystigma (Diptera:
Chamaemyiidae). The Canadian Entomologist 105: 1443-1452.
A Photographic Guide 59

20. Pine Leaf Adelgid, Pineus pinifoliae (Fitch) (Hemiptera Adelgidae)

Orientation to Pest Pine leaf adelgid, Pineus pinifoliae (Fitch), is native to North America
and feeds alternatively on spruce (Picea) and pine (Pinus). On spruce,
galls are produced. On pine, nymphs feeds openly on terminal foliage.
In the eastern United States, red (Picea rubens Sargent) and black (P.
mariana [Miller] Britton, Sterns and Poggenburg) spruce are the initial
hosts, with adults produced there, migrating to white pine (Pinus strobus
L.). In the western United States, the initial hosts are Engelmann spruce
(Picea engelmannii Parry ex Engelm.), black spruce, blue spruce (P.
pungens Engelm.), and white spruce (P. glauca [Moench] Voss), and the
pine host is western white pine (Pinus monticola Douglas ex D. Don).
Two years are required to complete this cycle among tree hosts. Pine leaf
adelgid can be an important pest in white pine plantations on young trees.
Heavy infestations can reduce growth of pine or even kill trees. Galls on
spruce do not damage the health of the tree.

Hosts Commonly Attacked This species feeds on red (Picea rubens), black (P. mariana), Engelmann
(P. engelmannii), blue (P. pungens), and white (P. glauca) spruce as
primary hosts and eastern white (Pinus strobus) and western white
(P. monticola) pines as alternate hosts.

Distribution This adelgid is found widely in North America wherever the ranges of its
primary and alternative hosts overlap.

Images of
Pine Leaf Adelgid

Figure 1. Close view of pine leaf adelgid, Pineus pinifoliae, on a spruce host.
(Whitney Cranshaw, Colorado State University, Bugwood.org)
60 Forest Pest Insects in North America:

Images of Pine Leaf

Adelgid (continued)

Figure 2. Fresh gall of pine leaf adelgid on a spruce host. (Whitney Cranshaw,
Colorado State University, Bugwood.org)

Figure 3. Old gall of pine leaf adelgid on a spruce host, after adelgid emergence.
(Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 4. Adults of pine leaf adelgid on a pine host. (E. Bradford Walker, Vermont
Department of Forests, Parks and Recreation, Bugwood.org)
A Photographic Guide 61

Images of Pine Leaf

Adelgid (continued)

Figure 5. Flagging tip on pine, a symptom of feeding of pine leaf adelgid.

(Ronald S. Kelley, Vermont Department of Forests, Parks and Recreation,
Bugwood.org)

Important Biological Control No information is available on the natural enemies of this adelgid. They
Agents Related to this are likely to be similar to those recorded for other Pineus and Adelges
Pest Species species in North America.

Web Links for Information http://www.forestpests.org/vermont/pineleafadelgid.html; fact sheet of

on Pine Leaf Adelgid the state of Vermont of biology and control.
http://www.maine.gov/doc/mfs/pineleaf.htm; fact sheet of the state of
Maine on biology and control.

Articles Dimond, J.B. 1974. Sequential surveys for the pine leaf chermid, Pineus
pinifoliae. Technical Bulletin No. 68, Life Sciences and Agriculture
Experiment Station, University of Maine; Orono, Maine, USA: 15 p.
62 Forest Pest Insects in North America:

21. White Pine Aphid, Cinara strobi (Fitch) (Hemiptera: Aphidae)

Orientation to Pest White pine aphid, Cinara strobi (Fitch), is a native insect in North
America that feeds on white pine (Pinus strobus L.). It is easily
recognized by its large size (3-4 mm), slightly dull metallic grey color
with a white stripe on middle of the thorax and spots of white wax on
each side of the abdomen. Eggs are laid in the fall, end-to-end along the
needle and hatch in the spring. White pine aphid feeds on branches and
upper trunks. Because it forms large colonies and is large, it produces
copious amounts of honeydew. It is usually attended by ants. Other than
the honeydew being a nuisance and fostering sooty mold, direct damage
to the tree itself is rare; but occasionally dense aphid populations damage
young pines or individual branches of large trees.

Hosts Commonly Attacked This aphid is only found on white pine (P. strobus).

Distribution White pine aphid is found in the United States from New England to
the Great Lakes states, south to the Carolinas in association with eastern
white pine.

Images of
White Pine Aphid

Figure 1. Close view of white pine aphid, Cinara strobi. (Jim Baker, North
Carolina State University, Bugwood.org)
A Photographic Guide 63

Images of White Pine Figure 2. Colony of white

pine aphid. (William H.
Aphid (continued) Hoffard, USDA Forest
Service, Bugwood.org)

Figure 3. Eggs of white pine aphid on pine needle. (Lacy L. Hyche, Auburn
University, Bugwood.org)

Important Biological Control The braconids Pauesia bicolor (Ashmead) and Pausia xanthothera
Agents Related to this (Smith) are known as parasitoids of this and other aphids in the genus
Pest Species Cinara.

Web Links for Information http://wiki.bugwood.org/Cinara_strobi

on White Pine Aphid

Articles None
64 Forest Pest Insects in North America:

22. Woolly Elm Aphid, Eriosoma americanum (Riley) (Hemiptera: Aphididae;

Eriosomatinae)

Orientation to Pest Woolly elm aphid, Eriosoma americanum (Riley), is a native insect
in North America whose primary hosts are American elm (Ulmus
americana L.) and secondary, summer hosts are serviceberry (called
saskatoon in Canada) and other species of Amelanchier spp. This aphid
flies to elm in the fall where it mates and lays overwintering eggs. These
hatch in the spring and nymphs feed on the edges of young elm leaves,
causing them to roll inward and form a gall over the colony. In early
summer, winged-forms migrate to Almelanchier species where they form
colonies on the roots. Two other species of Eriosoma may also be found
on elm, E. lanigerum (Hausmann) that causing terminal shoots to form
rosettes and E. rileri Thomas, which forms woolly clusters on limbs and
trunks. This aphid is a minor pest on Amelanchier alnifolia (Nutt.), which
is a fruit crop in Manitoba, Canada.

Hosts Commonly Attacked American elm (U. americana) is the primary host, and species of
serviceberry (Amelanchier) are the alternative hosts.

Distribution Woolly elm aphid is widely distributed throughout the eastern United
States and Canada in the range of elm and serviceberry species.

Images of Woolly Figure 1. Winged adult

of woolly elm aphid,
Elm Aphid Eriosoma americanum.
(Government of
Alberta, Agriculture and
Rural Development,
agriculture.alberta.ca)

Figure 2. Elm leaf galled by woolly elm aphid. (Steven Katovich, USDA Forest
Service, Bugwood.org)
A Photographic Guide 65

Images of Woolly Figure 3. Stem of

serviceberry infested by
Elm Aphid (continued) woolly elm aphid. (Whitney
Cranshaw, Colorado State
University, Bugwood.org)

Important Biological Control Given this aphid’s native status and limited damage, little is known
Agents Related to this about its natural enemies. In North America, some predators have been
Pest Species observed feeding on it, including the mirid Saileria irrorata Henry and
the coccinellid Scymnus brulleri Mulsant.

Web Links for Information http://www.gov.mb.ca/agriculture/crops/insects/fad92s00.html; gives

on Woolly Elm Aphid details of biology and effect on saskatoon (A. alnifolia),which is grown
for its fruit in Manitoba, Canada.
http://wiki.bugwood.org/Eriosoma_americanum; entry in Bugwood Wiki.

Articles Miranpuri, G.S. and G.G. Khachatourians. 1996. Bionomics and

fungal control of woolly elm aphid, Eriosoma americanum (Riley)
(Eriosomatidae: Homoptera) on Saskatoon berry, Amelanchier alinifolia.
Journal of Insect Science 9: 33-37.
Fry, K.M. 2001. Eriosoma americanum (Riley), woolly elm aphid
(Homoptera: Pemphigidae). Mason, P.G. and J.T. Huber (eds.).
Biological Control Programmes in Canada, 1981-2000. CABI
Publishing, Wallingford, UK: 120-123.
66 Forest Pest Insects in North America:

23. Woolly Beech Aphid, Phyllaphis fagi (L.) (Hemiptera: Aphidae)

Orientation to Pest Woolly beech aphid, Phyllaphis fagi (L.), is an invasive aphid in North
America, likely of European origin. It feeds on all species of beech,
but high densities are usually only found on European beech (Fagus
sylvatica L.), which is widely used in landscape plantings. The aphid
is usually not a problem in the United States on the American beech,
F. grandifolia Ehrh., growing in forests or low stress sites. This woolly
aphid, which feeds on the undersides of leaves, can be confused with
the beech blight aphid (Grylloprociphilus imbricator [Fitch]), which
begins feeding on the leaves and then moves onto stems. The woolly
beech aphid eggs hatch in the spring and there are about ten generations
per year of apterous and winged forms, which occur together. Huge
populations can occur year after year without causing visible harm to the
tree. The massive amounts of honeydew produced by large colonies can
be a nuisance.

Hosts Commonly Attacked This aphid is found most often on European beech (F. sylvatica L.)
planted in urban or residential landscapes.

Distribution This aphid is widely distributed in both eastern and western North
America, where ever beech is grown.

Images of Woolly
Beech Aphid

Figure 1. Colony of woolly beech aphid, Phyllaphis fagi, on undersurface of

leaves of European beech. (Haruta Ovidiu, University of Oradea, Bugwood.org)
A Photographic Guide 67

Images of Woolly
Beech Aphid (continued)

Figure 2. Close up of woolly beech aphid showing nymphs and older stages
with wool. (Louis-Michel Nageleisen, Département de la Santé des Forêts,
Bugwood.org)

Figure 3. Leaf rolling on European beech, a symptom of damage by woolly

beech aphid. (Louis-Michel Nageleisen, Département de la Santé des Forêts,
Bugwood.org)

Important Biological Control The braconid parasitoids Praon flavinode (Haliday) and Trioxys
Agents Related to this phyllaphidis Mackauer are known to attack this species in Europe, but
Pest Species are not reported from North America.

Web Links for Information http://www.plante-doktor.dk/boegebladluseng.htm; Danish fact sheet on

on Woolly Beech Aphid control of this species.

Articles Iversen, T. and S. Harding. 2007. Biological and other alternative control
methods against the woolly beech aphid, Phyllaphis fagi L., on beech,
Fagus sylvatica, seedlings in forest nurseries. Journal of Pest Science 80:
159-166.
68 Forest Pest Insects in North America:

24. Tuliptree Aphid, Illinoia liriodendri (Monell) (Hemiptera: Aphididae)

Orientation to Pest Tuliptree aphid, Illinoia liriodendri (Monell), is a relatively large aphid
(1.7-2.0 mm), spindle-shaped, pale green and lightly dusted with wax,
that is native to the eastern United States. It occurs east of the Mississippi
River, except northern New England, wherever its hosts, the tuliptree
(Liriodendron tulipifera L.) and some species of magnolia (Magnolia),
are endemic. It is also invasive in many areas where tuliptrees have been
planted as ornamentals outside their native range, including California
and other parts of the western USA, Europe (France, Germany, Italy,
Slovenia, UK), Japan, and Canada (Ontario, British Columbia). This
aphid is of little or no concern in its native range, but in parts of its
invaded range, high aphid densities occur that produce unsightly coatings
of honeydew and sooty mold.

Hosts Commonly Attacked This aphid’s hosts are the tuliptree (L. tulipifera L.) and some species of
magnolia (Magnolia).

Distribution Tuliptree aphid is found throughout the eastern USA and in urban areas
of California and other western states where tuliptrees have been planted.

Image of
Tuliptree Aphid

Figure 1. Adult and nymphs of tuliptree aphid, Illinoia liriodendra.

(Robert L. Anderson, USDA Forest Service, Bugwood.org)
A Photographic Guide 69

Important Biological Control Parasitoids of this aphid in its native range (the eastern United States)
Agents Related to this include a species of Praon, Ephedrus incompletus Provancher, Aphidius
Pest Species polygonaphis (Fitch), A. nigripes Ashmead, and A. rosae Haliday. Three
parasitoids—a Praon species, E. incompletus, and A. polygonaphis—
were imported from the eastern United Statees and released in California,
where A. polgonaphis established but did not completely control the
aphid population.

Web Links for Information http://meta.arsia.toscana.it/meta/meta?id_cms_doc=1&id_news=201; an

on Tuliptree Aphid Italian site covering this species as a pest of ornamentals.

Articles Zuparko, R.L. and D.L. Dahlsten. 1993. Survey of the parasitoids of
the tuliptree aphid, Illinoia liriodendri (Hom: Aphididae), in northern
California. Entomophaga 38: 31-40.
70 Forest Pest Insects in North America:

25. Norway Maple Aphid, Periphyllus lyropictus (Kessler) (Hemiptera:

Aphidae)

Orientation to Pest Norway maple aphid, Periphyllus lyropictus (Kessler), is a European

species that is invasive in the United States, where it occurs on Norway
maple (Acer platanoides L.). This aphid is widespread in eastern North
America, wherever Norway maple is planted. Given that Norway maple
is now considered an invasive forest pest in many areas, this aphid’s
damage to the tree might be considered beneficial. Large populations
can cause leaves to become brown or wrinkled and cause premature
leaf drop. However, the primary concern is usually the large amount
of honeydew it can produce, which is a nuisance to sidewalks and cars
beneath the trees. Both winged and wingless forms occur and the latter
form has a yellow-green body with a “lyre”-shaped brownish marking on
the abdomen.

Hosts Commonly Attacked Its only host in North America is the imported species Norway maple
(A. platanoides).

Distribution This aphid is found widely in eastern North America, wherever Norway
maple has been planted.

Images of
Norway Maple Aphid

Figure 1. Colony of Norway maple aphid, Periphyllus lyropictus.

(Michael Montgomery, USDA Forest Service, Bugwood.org)
A Photographic Guide 71

Images of Norway
Maple Aphid (continued)

Figure 2. Sooty mold and cast skins of Norway maple aphid. (Pennsylvania
Department of Conservation and Natural Resources - Forestry Archive,
Bugwood.org)

Important Biological Control In its native range in Europe, three species of parasitoids have been
Agents Related to this recorded that are believed to be specialized on this genus of aphid:
Pest Species Aphidius setiger Mackauer, Trioxys falcatus Mackauer, and Praon
silvestre Starý. Of these, only A. setiger is known to be present in North
America.

Web Links for Information None

on Norway Maple Aphid

Articles Starý, P. 1972. Host life-cycle and adaptation of parasites of Periphyllus

aphids (Homoptera, Chaitophoridae; Hymenoptera, Aphidiidae). Acta
Entomologica Bohemoslovaca 69(2): 89-96.
72 Forest Pest Insects in North America:

26. Linden Aphid, Eucallipterus tiliae (L.) (Hemiptera: Aphididae)

Orientation to Pest Linden aphid, Eucallipterus tiliae (L.), is invasive in the United States.
It is native to Eurasia, but now occurs worldwide, wherever species of
native or introduced linden trees (Tilia) occur. The winged adults are
easily recognized by the black stripe along the body and the cloudy-
black at the wing edge. Trees planted along streets and by parking areas
are frequently attacked, and the honeydew produced by the aphid makes
walkways and cars parked beneath them sticky. Sooty mold growing
on honeydew blackens the leaves. Progeny are produced continuously
during the growing season. It is attacked by several genera of parasitoids
as well as lady beetles and other generalist predators.

Hosts Commonly Attacked Basswood, also called American linden, (Tilia americana L.) and its
variety, white basswood (T. americana var. heterophylla) are North
American trees that are fed on by this aphid. However, damage to these
plants is rarely serious because they are less susceptible to this aphid
than are the introduced species of Tilia commonly planted in urban
landscapes. Damage to such introduced species is the main focus of
concern with this aphid.

Distribution This aphid is found widely in eastern North America on basswood and on
exotic species of Tilia in landscape plantings. California, especially, has
problems with this aphid on exotic Tilia and has introduced parasitoids
for its biological control.

Images of Linden Aphid Figure 1. Winged adult linden

aphid, Eucallipterus tiliae.
(Bob Gaia)
A Photographic Guide 73

Images of Linden Aphid Figure 2. Nymph of linden

aphid. (Bob Gaia)
(continued)

Figure 3. Group of winged adult linden aphids on underside of Tilia leaf.

(Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 4. Honeydew and sooty mold on linden, typical signs of linden aphid.
(Whitney Cranshaw, Colorado State University, Bugwood.org)
74 Forest Pest Insects in North America:

Images of Linden Aphid Figure 5. Dark discoloration

on cement caused by sooty
(continued) mold growing on honeydew
dripping from linden tree
infested by linden aphid.
(Whitney Cranshaw,
Colorado State University,
Bugwood.org)

Important Biological Control In California, where this is an invasive aphid on exotic street trees, a
Agents Related to this species of European parasitoid, Trioxys curvicaudus Mackauer, was
Pest Species introduced and anecdotal information suggests this introduction lowered
density of aphids on street lindens. In Europe, parasitoids seem not
to be the primary regulator of linden aphid numbers, and generalist
predators and density-dependent feedback from the tree are believed
to be important. This aphid and seven related species occur in northern
China and the Russian Far East; hence, this area may be a good source
of natural enemies. There are ten parasitoid species in the superfamily
Ichneumonoidea and nine in the Chalcidoidea recorded from aphids in
the genus Eucallipterus.

Web Links for Information None found that provide significant information.
on Linden Aphid
A Photographic Guide 75

Articles Dixon, A.F.G. and N.D. Barlow. 1979. Population regulation in the lime
aphid. Zoological Journal of the Linnean Society 67: 255-237.
Zuparko, R. 1983. Biological control of Eucallipterus tiliae [Hom.:
Aphididae] in San Jose, California, through establishment of Trioxys
curvicaudus [Hym.: Aphidiidae]. Entomophaga 28: 325-330.
Hajek, A.E. 1986. New North American records of European parasitoids
(Hymenoptera) of the linden aphid, Eucallipterus tiliae (Aphidoidea:
Drepanosiphidae). Journal of the New York Entomological Society 94:
443-446.
Blackman, R.L. and V.F. Eastop. 1994. Aphids on the World’s Trees: An
Identification and Information Guide. CAB International, Oxfordshire,
United Kingdom.
Zuparko, R.L. and D.L. Dahlsten. 1996. New potential for classical
biological control of Eucallipterus tiliae (Homoptera: Drepanosiphidae.
Biological Control 6: 407-408.
Foottit, R.G., S.E. Halbert, G.L. Miller, E. Maw, and L.M. Russell. 2006.
Adventive aphids (Hemiptera: Aphididae) of America North of Mexico.
Proceedings of the Entomological Society of Washington 108(3): 583-610.
76 Forest Pest Insects in North America:

27. Spruce Aphid, Elatobium abietinum (Walker) (Hemiptera: Aphidae)

Orientation to Pest Spruce aphid, Elatobium abietinum (Walker) (known as “green spruce
aphid” in some places), is native to part of Europe, where it feeds on
Norway spruce (Picea abies [L.] H. Karst). It is seldom a pest on this
species in Europe, causing only infrequent, very minor local outbreaks.
However, it is a major pest in northwestern Europe in plantations of
North American Sitka spruce (P. sitchensis [Bong.] Carr. Outside of
Europe, spruce aphid is invasive in many areas with maritime climates
and also the montane forests of the southwestern United States. In
North America, spruce aphid occurs in natural forests on various spruce
species, mostly in coastal areas of the Pacific northwestern United States
and British Columbia. It also occurs in Arizona, where spruce aphid
outbreaks damage Engelmann spruce (P. engelmannii Parry ex Engelm.),
killing 10 to 24 percent of infested trees, depending on the severity
of injury. In maritime areas, spruce aphid is parthenogenic (females
only), but in its native range and in Arizona the life cycle includes both
parthenogenic and sexual forms.

Hosts Commonly Attacked Spruce aphid feeds on virtually all spruce, including Norway (P. abies),
Sitka (P. sitchensis), Englemann (P. engelmannii), and Colorado blue
spruce (P. pungens Englmann).

Distribution In addition, to its native range in Europe, this aphid has invaded parts of
Russia, Australia, New Zealand, Chile, and North America. It is found in
British Columbia, Ontario, Quebec, California, North Carolina, Oregon,
Arizona, Utah, and Washington.

Images of Spruce Aphid

Figure 1. Spruce aphid, Elatobium abietinum. (Donald Owen, California

Department of Forestry and Fire Protection, Bugwood.org)
A Photographic Guide 77

Images of Spruce Aphid

(continued)

Figure 2. Damage to Sitka spruce from spruce aphid. (Elizabeth Willhite, USDA
Forest Service, Bugwood.org)

Figure 3. Stand of Sitka spruce in Alaska showing damage of spruce aphid

(areas in brown). (Andris Eglitis, USDA Forest Service, Bugwood.org)

Important Biological Control Generalist predators of this species have been noted, but parasitoids
Agents Related to this have not been studied. Pathenogenic fungi, especially species of
Pest Species Entomophthorales, are important natural enemies of spruce aphid.

Web Links for Information http://www.fs.fed.us/r3/resources/health/field-guide/sap/spruce.shtml

on Spruce Aphid http://apps.rhs.org.uk/advicesearch/Profile.aspx?pid=520; a control
factsheet from the UK Royal Horticultural Society.
http://www.oregon.gov/ODF/privateforests/docs/fh/SpruceAphid.
pdf?ga=t
www.science.ulster.ac.uk/lnsrg/elatobium_abietinum.html; University
of Ulster, Northern Ireland. Includes photos of some natural enemies of
spruce aphid.
78 Forest Pest Insects in North America:

Articles on Spruce Aphid Straw, N.A., J.E.L. Timms, and S.R. Leather. 2009. Variation in the
abundance of invertebrate predators of the green spruce aphid Elatobium
abietinum (Walker) (Homoptera: Aphididae) along an altitudinal transect
Elatobium abietinum. Forest Ecology and Management 258: 1-10.
Lynch, A.M. 2004. Fate and characteristics of Picea damaged by
Elatobium abietinum (Walker) (Homoptera: Aphididae) in the White
Mountains of Arizona. Western North American Naturalist 64: 7-17.
A Photographic Guide 79

28. Beech Scale, Cryptococcus fagisuga Lindinger (Hemiptera:

Eriococcidae)

Orientation to Pest Beech scale, Cryptococcus fagisuga Lindinger, is invasive in the

United States and very likely also invasive in Europe. Its native range is
believed, based on studies of mitochondrial DNA, to be in western Asia
(Turkey and the Caucasus Mountains) or southeastern Europe, where
its host is oriental beech (Fagus orientalis Lipsky). It is a moderately
damaging pest in Europe on European beech (F. sylvatica L.) and a
devastating pest in North America of American beech (F. grandifolia
Ehrh.). The insect infests the trunk of beech, inducing cracking which
allows invasion by fungal pathogens (genus Neonectria), which kill the
trunk. Roots survive and trees regenerate, but the disease cycle is too
frequent to permit most trees to reach large size. Trunks are deformed
by cracking and the development of cankers. Visible signs are the white
woolly coverings of scales and the red fruiting bodies of the Neonectria
fungus. About 1 percent of American beech are naturally immune from
attack and remain healthy. Following the accidental introduction of the
scale to the Halifax Arboretum in the late 1800s, an epidemic of beech
bark disease has spread south and west, and now covers a substantial
part of the range of American beech. This insect is apparently completely
parthenogenetic; no males have ever been found.

Hosts Commonly Attacked Oriental beech (F. orientalis) is believed to be the native host. European
beech (F. sylvatica) and American beech (F. grandifolia) are hosts
acquired in invaded areas.

Distribution In North America, beech scale occurs from Nova Scotia to central
Pennsylvania, with infested pockets in West Virginia, North Carolina,
and Michigan, but continues to spread. In Eurasia, the scale is found
from the United Kingdom to Iran. It is found throughout Europe;
however, this area is no longer believed to be the native range. Of
areas studied so far, mitochondrial haplotype diversity is greatest in the
Caucasus Mountains and that area may be part of the original range of
this insect. Surveys are underway in the Caucasus Mountains of Georgia
for parasitoids of this scale.

Figure 1. Distribution of beech

scale, Cryptococcus fagisuga, and
its associated invasive pathogen
in North America in 2005, following
its introduction in about 1890 in
Halifax, Nova Scotia. (USDA Forest
Service, Northeastern Area, Forest
Health Protection: Beech Bark
Disease)
80 Forest Pest Insects in North America:

Images of Beech Scale

Figure 2. Close up of beech scales under their woolly coverings (left) and
with wool removed, showing orange scale bodies (right). (Both photos: Chris
Malumphy, The Food and Environment Research Agency, Bugwood.org)

Figure 3. First instar nymph (crawler) of beech scale. (Joseph O’Brien, USDA
Forest Service, Bugwood.org)

Figure 4. View of trunk of

American beech infested with
beech scale (light colored
material). (Joseph O’Brien, USDA
Forest Service, Bugwood.org)
A Photographic Guide 81

Images of Beech Scale

(continued)

Figure 5. Beech scale on bark of severely damaged American beech (left) (note
also lumpy cankers on trunk caused by beech bark disease); close up of colonies
of beech scale (right). (Left: Joseph O’Brien, USDA Forest Service, Bugwood.org;
right: Chris Malumphy, The Food and Environment Research Agency,
Bugwood.org)

Figure 6. Beech trees in forest killed by beech bark disease. (Joseph O’Brien,
USDA Forest Service, Bugwood.org)

Figure 7. Cracked bark of tree with beech bark disease (left) compared to smooth
grey bark of healthy American beech (right). (Both photos: Joseph O’Brien, USDA
Forest Service, Bugwood.org)
82 Forest Pest Insects in North America:

Images of Beech Scale Figure 8. Bark cracking and

growth of epiphytes (here,
(continued) moss) on trunk are other
signs of beech bark disease.
(Milan Zubrik, Forest
Research Institute - Slovakia,
Bugwood.org)

Figure 9. Dark weeping spots are also a sign of beech bark disease. (Joseph
O’Brien, USDA Forest Service, Bugwood.org)

Figure 10. Discoloration under bark due to Neonectria cankers. (Joseph O’Brien,
USDA Forest Service, Bugwood.org)
A Photographic Guide 83

Images of Beech Scale Figure 11. Orange fruiting

bodies of Neonectria on
(continued) beech bark. (Andrej Kunca,
National Forest Centre -
Slovakia, Bugwood.org)

Important Biological Control No parasitoids of this scale are known from either North America or
Agents Related to this Europe. Studies in both regions found only generalist predators attacking
Pest Species the scale. Currently efforts to find parasitoids are underway in the
Caucasus Mountains, the region now believed to be at least part of the
native range of the scale.

Web Links for Information http://www.ecostudies.org/people_sci_lovett_beech_bark_disease.html;

on Beech Scale ecosystem effects of beech bark disease.
http://nrs.fs.fed.us/disturbance/invasive_species/bbd_resistant_beech/;
research on resistance in American beech to beech bark disease.
http://threatsummary.forestthreats.org/threats/threatSummaryViewer.
cfm?threatID=117
http://en.wikipedia.org/wiki/Beech_bark_disease
http://www.sel.barc.usda.gov/catalogs/eriococc/Cryptococcusfagisuga.
htm; ScaleNet catalog, giving a complete list of all recorded natural
enemies, hosts, and distribution records, and an index to all other
information published on this insect.
84 Forest Pest Insects in North America:

Articles on Beech Scale Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A Review and Recommendations.
FHTET-96-19. USDA Forest Service, Morgantown, West Viriginia,
USA: 20-22.
Forrester, J.A., G.G. McGee, and M.J. Mitchell. 2003. Effects of beech
bark disease on aboveground biomass and species composition in a
mature northern hardwood forest, 1985 to 2000. Journal of the Torrey
Botanical Society 130(2): 70-78.
Gwiazdowski, R.A., R.G. Van Driesche, A. Desnoyers, S. Lyon, San-an
Wu, N. Kamatad and B.B. Normark. 2006. Possible geographic origin
of beech scale, Cryptococcus fagisuga (Hemiptera: Eriococcidae), an
invasive pest in North America. Biological Control 39: 9-18.
A Photographic Guide 85

29. Gillette’s Eriococcin, Eriokermes gillettei (Tinseley) (Hemiptera:

Kermesidae, formerly Eriococcidae)

Orientation to Pest Gillette’s eriococcin, Eriokermes gillettei (Tinseley) (syn. is Eriococcus

gillettei), is a native North American scale. This species is found on
native junipers (Juniperus) but is not damaging. Occasionally higher
densities develop on junipers planted as ornamentals. Adult females are
yellowish before forming an ovisac but then turn brownish purple. In
Maryland, there is one generation per year and the scale overwinters in
the egg stage.

Hosts Commonly Attacked This scale is restricted to feeding on species of juniper (Juniperus).

Distribution This scale is known from Florida, Maryland, Virginia, and several
western states, but likely has a larger U.S. distribution, including most of
the southeastern United States.

Image of Figure 1. Gillette’s eriococin,

Eriokermes gillettei, on
Gillette’s Eriococcin juniper. (Raymond Gill,
California Department
of Food and Agriculture,
Bugwood.org)

Important Biological Control Parasitoids of this scale have not been reported but its characteristic
Agents Related to this low density in natural habitats suggests it may be kept under control by
Pest Species specialized parasitoids.

Web Links for Information http://www.sel.barc.usda.gov/catalogs/kermesid/Eriokermesgillettei.htm;

on Gillette’s Eriococcin ScaleNet catalog, giving a complete list of all recorded natural enemies,
hosts, and distribution records, and an index to all other information.

Articles Miller, D.R. and G.L. Miller. 1993. Description of a new genus of scale
insect with a discussion of relationships among families related to the
Kermesidae (Homoptera: Coccoidea). Systematic Entomology 18:
237-251. (This article introduced the current name of the species.)
86 Forest Pest Insects in North America:

30. Pine Bast Scale, Matsucoccus matsumurae (Kuwana) (Hemiptera:

Matsucoccidae)

Orientation to Pest Pine bast scale, Matsucoccus matsumurae (Kuwana), (formerly referred
to as red pine scale, Matsucoccus resinosae, in the United States) is a
Japanese scale that is invasive in the United States. Along with six Asian
species of pines, it attacks red pine (Pinus resinosa Sol. ex Aiton), its
only native North American host. It occurs in plantations of red pine
planted south of their native range in Connecticut, New York, New
Jersey, and Pennsylvania. The infested area is expanding slowly.

Hosts Commonly Attacked Pine bast scale attacks several Asian pines, but its only North American
native host is red pine (P. resinosa).

Distribution This scale is native to Japan and is invasive in China, South Korea,
Sweden, and eastern North America.

Image of Figure 1. Pine bast scale,

Matsucoccus matsumurae.
Pine Bast Scale (USDA APHIS PPQ CPHST)

Important Biological Control Natural enemies associated with pine bast scale in North America are
Agents Related to this principally generalist predators (coccinellids, anthocorid bugs, and
Pest Species cecidomyiid flies). In China, predation on eggs and young nymphs by the
anthocorid Elatophilus nipponensis Hiura is a critical factor affecting the
pest. Also in both China and Japan, the generalist predatory coccinellid
Harmonia axyridis Pallas is believed to suppress this scale’s density. This
coccinellid in now found as an invasive species throughout the range of
red pine scale in the northeast United States but definitive studies on its
impact on this scale are lacking.

Web Links for Information http://www.sel.barc.usda.gov/catalogs/matsucoc/

on Pine Bast Scale Matsucoccusmatsumurae.htm; ScaleNet catalog, giving a complete list of
all recorded natural enemies, hosts, and distribution records, and an index
to all other information published on this insect.
A Photographic Guide 87

Articles Cheng, H.Y. and W.J. Ming. 1979. Population dynamics and
biological control of Matsucoccus matsumurae Kuwana (Homoptera:
Margarodidae). Acta Entomologica Sinica 22: 149-155.
McClure, M.S. 1986. Role of predators in regulation of endemic
populations of Matsucoccus matsumurae (Homoptera: Margarodidae) in
Japan. Environmental Entomology 15: 976-983.
Ming, W.J., Q.J. Ge, and H.Y. Zheng. 1983. Studies of some major
predaceous natural enemies of Matsucoccus matsumurae (Kuwana).
Journal of Nanjing Technological College of Forest Products 3: 19-29.
88 Forest Pest Insects in North America:

31. Tuliptree Scale, Toumeyella liriodendri (Gmelin) (Hemiptera: Coccidae)

Orientation to Pest Tuliptree scale, Toumeyella liriodendri (Gmelin), is native to the eastern
United States, where it is widely distributed. It also occurs in urban areas
in California, where it is invasive. It mainly feeds on twigs of tuliptree
(Liriodendron tulipifera L.) and magnolia (Magnolia). Heavy infestations
are common and these can (1) kill seedlings, (2) destroy leaders or
branches leading to bushy form, and (3) remove nutrients and lower tree
vigor. The scale has one generation per year except in the southern part
of its range and it generally overwinters as young nymphs on twigs.

Hosts Commonly Attacked This scale feeds on tuliptree (L. tulipifera) and species of magnolia
(Magnolia), as well as hosts in eight other plant families.

Distribution This scale is found from New York south to Florida and west to the
Mississippi River (see Burns and Donley [1970] for a distribution map),
and has invaded urban areas of California.

Images of
Tuliptree Scale

Figure 1. Group of mature female tuliptree scales, Toumeyella liriodendri, on twig

of host. (A. Steven Munson, USDA Forest Service, Bugwood.org)
A Photographic Guide 89

Images of Tuliptree Figure 2. Mature female of

tuliptree scale surrounded
Scale (continued) by crawlers. (Gerald J.
Lenhard, Louiana State Univ,
Bugwood.org)

Figure 3. Overwintering second instar nymphs of tuliptree scale on twig.

(Lacy L. Hyche, Auburn University, Bugwood.org)

Figure 4. Empty cast skins of male “pupae” of tuliptree scale. (Lacy L. Hyche,
Auburn University, Bugwood.org)
90 Forest Pest Insects in North America:

Important Biological Control Many species of parasitoids and predators are reported attacking tuliptree
Agents Related to this scale in the United States, but at the same time, the scale is reported as
Pest Species frequently being abundant and damaging. Its ability to thrive in spite of
its many enemies may be because it is very frequently tended by ants.

Web Links for Information http://www.fs.fed.us/r6/nr/fid/fidls/fidl-92.pdf; USDA Forest Service

on Tuliptree Scale leaflet #92 with photos of damage to tree form.
http://ento.psu.edu/extension/factsheets/tuliptree-scale; fact sheet from
Pennsylvania State University; discusses control.
http://www.ca.uky.edu/entomology/entfacts/ef435.asp; fact sheet from
University of Kentucky; discusses control.
http://cfextension.ifas.ufl.edu/agriculture/nursery_production/documents/
Magnolia.pdf; University of Florida; discusses control on magnolia.
http://www.sel.barc.usda.gov/catalogs/coccidae/Toumeyellaliriodendri.
htm; ScaleNet catalog, giving a complete list of all recorded natural
enemies, hosts, and distribution records, and an index to all other
information published on this insect.

Articles Burns, D.P. and D.E. Donley. 1970. Biology of the tuliptree scale,
Toumeyella liriodendri (Homoptera: Coccidae). Annals of the
Entomological Society of America 63: 228-235.
Simpson, J.D. and P.L. Lambdin. 1983. Life history of the tuliptree
scale, Toumeyella liriodendri (Gmelin), on yellow-poplar in Tennessee.
Tennessee Farm and Home Science 125: 2-5.
A Photographic Guide 91

32. Pine Tortoise Scale, Toumeyella parvicornis (Cockerell) (Hemiptera:

Coccidae)

Orientation to Pest The pine tortoise scale, Toumeyella parvicornis (Cockerell), is native
to the United States and is found widely over the northeastern, north
central, and southeastern portions of the country. It is also invasive in
Puerto Rico and the Turks and Caicos Islands in the Caribbean. It has
one generation per year in the northern parts of its range, and four or
more generations in the southern portions. The first instars (crawlers)
are the mobile dispersal stage. Crawlers eventually settle either on the
shoot or on the needles. The choice of settling site may be host-specific.
Settled females on shoots are brown with dark markings and have
the hemispherical tortoise shape for which they are named. Females
developing on the needles are elongate and initially are light green with
darker green stripes, eventually turning brownish. The males develop on
the same plant part as the females. Adult males are winged and smaller
than the females. The scales excrete honeydew, which provides a growth
medium for sooty mold. Heavy infestations may result in tree mortality.

Hosts Commonly Attacked Most pines within its range can be infested by this scale but important
hosts include Scotch (Pinus sylvestris), Jack (P. banksiana), Virginia
(P. virginiana), loblolly (P. taeda), and slash (P. elliottii) pines.

Distribution This scale is found from eastern New York, south to Florida and west
through the Dakotas, Colorado, and Texas wherever its hosts are found.
It is also known to occur in Manitoba and Ontario, Canada, and has been
introduced into parts of the Caribbean.

Images of
Pine Tortoise Scale

Figure 1. Pine tortoise scales, Toumeyella parvicornis; note the tortoise shell
shape and color pattern. (Lacy L. Hyche, Auburn University, Bugwood.org)
92 Forest Pest Insects in North America:

Images of Pine Tortoise

Scale (continued)

Figure 2. Sooty mold growing on honeydew excreted by pine tortoise scales

on pine needles. (R. Scott Cameron, Advanced Forest Protection, Inc.,
Bugwood.org)

Figure 3. Adult (shoot form) of the female pine tortoise scale with emerging
crawlers (left) and the needle form female with settled crawlers (right).
(Left: Jill O’Donnell, MSU Extension, Bugwood.org; right: Albert (Bud) Mayfield,
USDA Forest Service, Bugwood.org)

Figure 4. Settled crawlers of pine tortoise scale. (Lacy L. Hyche, Auburn

University, Bugwood.org)
A Photographic Guide 93

Images of Pine Tortoise

Scale (continued)

Figure 5. Young nymphs of pine tortoise scale. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Figure 6. Damage to Virginia

pine from pine tortoise scale.
(Lacy L. Hyche, Auburn
University, Bugwood.org)

Figure 7. Ladybird beetles feeding on pine tortoise scales. (Lacy L. Hyche,

Auburn University, Bugwood.org)
94 Forest Pest Insects in North America:

Important Biological Control Many species of parasitoids and predators are reported attacking pine
Agents Related to this tortoise scale in the United States, and in most settings, natural enemies
Pest Species are sufficient to maintain the scale at low densities. Damaging scale
populations develop either when ants tend scale colonies (see Wilkinson
and Chellman, 1979) or where dust or insecticides reduce natural enemy
numbers in seed orchards or Christmas tree farms (see Clarke et al.,
1992).

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/pine_tort_scale/pine_tort_scale.

on Pine Tortoise Scale htm; USDA FS leaflet, with information on biology, life cycle and
control of this scale.
http://www.plantpath.cornell.edu/trees/PTortSc.html; fact sheet from
Cornell University.
http://www.entomology.umn.edu/cues/Web/186PineTortoiseScale.pdf;
fact sheet from University of Minnesota.

Articles Wilkinson, R.C. and C.W. Chellman. 1979. Toumeyella scale, red
imported fire ant, reduce slash pine growth. Florida Entomologist 62:
71-72.
Clarke, S.R., J.F. Negron, and G.L. DeBarr. 1992. Effects of four
pyrethroids on scale insect (Homoptera) populations and their natural
enemies in loblolly and shortleaf pine seed orchards. Journal of
Economic Entomology 85: 1246-1252.
A Photographic Guide 95

33. Tessellated Scale, Eucalymnatus tessellatus (Signoret) (Hemiptera:

Coccidae)

Orientation to Pest Tessellated scale, Eucalymnatus tessellatus (Signoret), occurs as an

invasive species in the southern United States and parts of Africa,
Australia, Asia and Europe. It is believed to be native to South America.
The adult female is oval or pear-shaped and is reddish to dark brown in
color. There are one or two generations per year. All scales are female
and give birth directly to crawlers through egg retention.

Hosts Commonly Attacked Tessellated scale has an extremely broad host range, attacking more
than 50 families of plants, including many monocots and dicots used as
ornamentals or as fruit trees. Common host plants include palms, figs,
and mango (Mangifera indica L.).

Distribution In the United States, this scale is found outdoors in various southern
states, particularly Florida. In northern areas, the scale is found in
greenhouses.

Image of Figure 1. Closer view of a

single adult of tessellated
Tesselated Scale scale, Eucalymnatus
tessellatus. (United States
National Collection of Scale
Insects Photographs Archive,
USDA Agricultural Research
Service, Bugwood.org)

Important Biological Control The parasitic wasp Metaphycus stanleyi Compere is a natural enemy of
Agents Related to this tessellated scale.
Pest Species

Web Links for Information http://edis.ifas.ufl.edu/pdffiles/IN/IN82600.pdf; fact sheet of the

on Tesselated Scale University of Florida.
http://www.entomology.umn.edu/cues/inter/inmine/Scalet.html; fact
sheet from University of Minnesota.
http://www.sel.barc.usda.gov/catalogs/coccidae/Eucalymnatustessellatus.
htm; ScaleNet catalog, giving a complete list of all recorded natural
enemies, hosts, and distribution records, and an index to all other
information published on this insect.
96 Forest Pest Insects in North America:

Article on Tesselated Scale Ray, C.H. and M.L. Williams. 1981. Redescription and lectotype
designation of the tessellated scale, Eucalymnatus tessellatus (Signoret)
(Homoptera: Coccidae). Proceedings of the Entomological Society of
Washington 83: 230-244. (Provides descriptions of three immature stages
and adult female, a key for the separation of instars, a list of hosts, and
information on distribution.).
A Photographic Guide 97

34. Elongate Hemlock Scale, Fiorinia externa Ferris (Hemiptera:

Diaspididae)

Orientation to Pest Elongate hemlock scale, Fiorinia externa Ferris, an invasive scale in
the USA, is native to Japan and China that is a pest of eastern hemlock
(Tsuga canadensis [L.] Carrière) in the eastern United States. The
scale has long-lived adult females that reproduce slowly over much of
the year, leading to a population with highly overlapping life stages.
In New England there is one generation per year but there are two in
North Carolina. Damage can be serious in the eastern United States on
hemlock, with densities one- to three-thousand-fold higher than in Japan,
causing chlorosis, needle drop, and tree mortality.

Hosts Commonly Attacked This scale attacks principally eastern hemlock (T. canadensis), but
sometimes species of spruce or fir (Abies), Douglas-fir (Pseudotsuga
menziesii [Mirbel] Franco), or yew (Taxus) are infested.

Distribution Elongate hemlock scale is native to Japan and China but is invasive in
North America from New Hampshire to Georgia and west to Michigan,
within range of eastern hemlock. This scale is also invasive in Europe
(UK, France).

Figure 1. North American distribution of elongate hemlock scale, Fiorinia externa.

(USDA Forest Service - AFPE Online Mapping)
98 Forest Pest Insects in North America:

Images of Elongate
Hemlock Scale

Figure 2. Heavy infestation of elongate hemlock scale on eastern hemlock,

showing many white males and some brown adult females. (Eric R. Day, Virginia
Polytechnic Institute and State University, Bugwood.org)

Figure 3. Adult female

elongate hemlock scales
(brown). (Kristopher Abell,
University of Massachusetts,
Bugwood.org)

Figure 4. Eggs (yellow) of

elongate hemlock scale
inside opened scale cover of
adult female. (Pennsylvania
Department of Conservation
and Natural Resources -
Forestry Archive,
Bugwood.org)

Figure 5. Settled first

instar nymphs (“crawlers”,
see light colored, upper
right quarter of photo) of
elongate hemlock scale.
(Rich Cowles, Connecticut
Agricultural Experiment
Station)
A Photographic Guide 99

Images of Elongate
Hemlock Scale (continued)

Figure 6. Second instar nymph (mid sized, yellow stages) of elongate hemlock
scale; also showing brown adult females and white males. (Pennsylvania
Department of Conservation and Natural Resources - Forestry Archive,
Bugwood.org)

Figure 7. Adult female elongate hemlock scales (left) with parasitoid emergence
holes; adult Encarsia citrina (right), an often abundant but generally ineffective
polyphagous parasitoid found attacking elongate hemlock scale in both North
America and Japan. (Both photos, Kristopher Abell, University of Massachusetts,
Bugwood.org)

Figure 8. Cybolephalus nr.

nipponicus Endrody-Younga,
an introduced predatory
beetle that sometimes preys
on elongate hemlock scale.
(Pennsylvania Department
of Conservation and Natural
Resources - Forestry
Archive, Bugwood.org)
100 Forest Pest Insects in North America:

Important Biological Control Studies by Kristopher Abell (Ph.D. dissertation, Entomology, University
Agents Related to this of Massachusetts, 2010) have shown that Encarsia citrina (Crawford),
Pest Species a polyphagous aphelinid parasitoid commonly found attacking elongate
hemlock scale in both eastern North America and Japan, is only one of
many Encarsia, Aphytis and other parasitoid species that attack the scale
in Japan, which, except for E. citrina, are not present in North America.
In addition, this same research has confirmed that E. citrina and the
susceptible life stage of the scale are asynchronous throughout North
America, even where two complete generations of the scale occur as
Japan. The lack of synchrony was attributed to overlapping generations
and delayed senescence of the adult female scale. Importations of species
of parasitoids from Japan has potential to reduce densities of elongate
hemlock scale in North America.

Web Links for Information http://ento.psu.edu/extension/factsheets/elongate-hemlock-scale; fact

on Elongate Hemlock Scale sheet from Pennsylvania State University.
http://www.na.fs.fed.us/spfo/pubs/pest_al/ehscale/ehscale.htm; fact sheet
from USDA Forest Service, Northeastern Area.
http://www.hgic.umd.edu/_media/documents/hg81.pdf; fact sheet from
Maryland Cooperative Extension Service.
http://www.fcla.edu/FlaEnt/fe89p527.pdf; article on results of predator
survey in North Carolina and Tennessee, USA.

Articles McClure, M.S. and M.B. Fe. 1977. Fiorinia externa and Tsugaspidiotus
tsugae (Homoptera: Diaspididae): distribution, abundance, and new
hosts of two destructive scale insects of eastern hemlock in Connecticut.
Environmental Entomology 6: 807-811.
Preisser, E.L., J.S. Elkinton, and K. Abell. 2008. Evolution of increased
cold tolerance during range expansion of the elongate hemlock
scale Fiorinia externa Ferris (Hemiptera: Diaspididae). Ecological
Entomology 33: 709-715.
Abell, K. 2010. Population dynamics and biological control of elongate
hemlock scale, Fiorinia externa. Ph.D. dissertation, Entomology,
University of Massachusetts, Amherst, Massachusetts.
A Photographic Guide 101

35. Oystershell Scale, Lepidosaphes ulmi (L.) (Hemiptera: Diaspididae)

Orientation to Pest Oystershell scale, Lepidosaphes ulmi (L.), is a widespread species

of uncertain origin, now found in almost every temperate area in the
world. It is known to be invasive in North America, Australia, New
Zealand, South Africa, and Japan. It is also found, potentially as a native
species, in North Africa and throughout temperate regions of Europe
and Asia. It is one of the most polyphagous insects known, reported
from 72 different families of plants. Oystershell scale is found mostly
on branches and trunks of broadleaf shrubs and trees, in both natural
habitats and planted landscapes. The coloration of the adult female varies
with the host species. The life history of the species also varies with
host, with more generations per year on some hosts and fewer on others.
The species overwinters as eggs on all host species. Dense populations
may kill branches, whole trees, or even forest stands. This variable
cosmopolitan species may in reality be a complex of several biotypes
that are being lumped together (see Gharib, 1978). Some populations are
parthenogenetic, consisting entirely of females.

Hosts Commonly Attacked Oystershell scale is a polyphagous species that attacks many broad leaf
woody species, including apple (Malus), ash (Fraxinus), beech (Fagus),
birch (Betula), elm (Ulmus), maple (Acer), poplar (Populus), species of
Prunus, and willow (Salix).

Distribution This invasive species is found throughout the United States and southern
Canada.

Images of Figure 1. Oystershell scale,

Lepidosaphes ulmi, is named
Oystershell Scale for the shape of the scale
covering, resembling an
oyster shell. (United States
National Collection of Scale
Insects Photographs Archive,
USDA Agricultural Research
Service, Bugwood.org)
102 Forest Pest Insects in North America:

Images of Oystershell Figure 2. View of dense

colony of oystershell scales.
Scale (continued) (United States National
Collection of Scale Insects
Photographs Archive, USDA
Agricultural Research
Service, Bugwood.org)

Figure 3. Adult oystershell scale inverted to reveal masses of white eggs,

the overwintering stage. (Whitney Cranshaw, Colorado State University,
Bugwood.org)

Figure 4. One female oystershell scale surrounded by recently settled crawlers.

(Whitney Cranshaw, Colorado State University, Bugwood.org)
A Photographic Guide 103

Images of Oystershell
Scale (continued)

Figure 5. Dense population of oystershell scale on aspen in Colorado. (Whitney

Cranshaw, Colorado State University, Bugwood.org)

Figure 6. Willows in natural habitat killed by oystershell scale. (USDA Forest

Service - Ogden Archive, USDA Forest Service, Bugwood.org)
104 Forest Pest Insects in North America:

Important Biological Control Despite a large literature on this species, relatively little is known
Agents Related to this about its potential for classical biological control because the species
Pest Species is principally of concern in apple orchards, where the perception is that
no scales can be tolerated. But in a forestry or shade tree context, that
concern should not apply. Two natural enemies are frequently mentioned
in connection with this scale, the predatory mite Hemisarcoptes malus
(Shimer) and the aphelid parasitoid Aphytis mytilaspidis (Le Baron).
While both of these occur in North America, neither were deliberately
introduced. Indeed, little to no work has been done to determine the
native range of this scale (likely Central Asia or the Russian Far East)
and then determine which species of natural enemies in that area are
most effective against low density scale populations. Also, this species
of scale may in reality be a complex of species or races and this should
be sorted out with DNA methods to show which races are of concern in
North America and where are they found in the native range. Finally,
even some natural enemies like A. mytilaspidis, which is present in the
United State, might be improved by a new introduction of the species
that is better adapted to cold areas, where the existing race does not do
well but where much of the damage from oystershell scale occurs.

Web Links for Information http://www.ext.colostate.edu/pubs/insect/05513.html; fact sheet from

on Oystershell Scale Colorado State University.
http://ento.psu.edu/extension/factsheets/oystershell-scale; fact sheet from
Pennsylvania State University.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Lepidosaphesulmi.htm;
ScaleNet catalog, giving a complete list of all recorded natural enemies,
hosts, and distribution records, and an index to all other information
published on this insect.

Articles Gharib, B. 1978. On two new distinct biological races of Lepidosaphes

ulmi L. (Homoptera, Coccoidea, Diaspidinae). Comptes Rendus
Hebdomadaires des Seances de l’Academie des Sciences, D 286(18):
1313-1314. (In French).
A Photographic Guide 105

36. San José Scale, Diaspidiotus perniciosus (Comstock) (Hemiptera:

Diaspididae)

Orientation to Pest San José scale, Diaspidiotus perniciosus (Comstock), is native to Asia
(northern China, Korea, the Russian Far East) and is invasive in North
America and in other temperate and subtropical areas throughout the
world. It is found throughout the United States and Canada. It is a highly
polyphagous species, reported from 43 different families of plants,
but is economically most troublesome on members of the rose family.
The scale has multiple generations per year in many areas and, if not
suppressed by parasitoids, can be very damaging, killing heavily infested
trees. It is not a pest in native forests, but is common on ornamental,
shade, and fruit trees.

Hosts Commonly Attacked Common hosts include many rosaceous plants, especially apple (Malus),
plums and cherries (Prunus), peaches (P. persica (L.) Batsch), almonds
(P. dulcis [Mill.] D.A.Webb), pear (Pyrus), and Pyracantha.

Distribution This invasive species, while native to northeastern Asia, is found

throughout North America, and more generally around the world in
temperate and subtropical regions.

Images of
San José Scale

Figure 1. Adults (left) of San José scale, Quadraspidiotus perniciosus; females

(right) with scale cover removed to show scale body. (Left: United States National
Collection of Scale Insects Photographs Archive, USDA Agricultural Research
Service, Bugwood.org; right: Washington State University)
106 Forest Pest Insects in North America:

Images of San José

Scale (continued)

Figure 2. Crawler of San José scale. (Washington State University)

Figure 3. Nymphs of San

José scale settled around
two adult females. (United
States National Collection of
Scale Insects Photographs
Archive, USDA Agricultural
Research Service,
Bugwood.org)

Figure 4. Adult male of San José scale (left); drawing of male (right).
(Left: Washington State University; right: FCIT - http://etc.usf.edu/clipart)
A Photographic Guide 107

Images of San José Figure 5. View of twig

infested by San José scale.
Scale (continued) (United States National
Collection of Scale Insects
Photographs Archive, USDA
Agricultural Research
Service, Bugwood.org)

Figure 6. Damage on apples from San José scale. Note both scales and red
discoloration around scales. (Central Science Laboratory, Harpenden Archive,
British Crown, Bugwood.org)

Figure 7. Encarsia perniciosi, an important parasitoid of San José scale.

(HYPPZ)
108 Forest Pest Insects in North America:

Images of San José Figure 8. Twice stabbed

ladybird, Chilocorus orbus
Scale (continued) Drea, a predator of San José
scale. (Jack Kelly Clark,
University of California
- Statewide IPM Program,
Bugwood.org)

Important Biological Control As San José scale moved around the world into various apple growing
Agents Related to this regions, it came in contact with local parasitoids, some of which adopted
Pest Species this scale as a new host. Also, several of the Asian parasitoids associated
with this scale in its native range moved naturally with the scale to new
regions (on apple breeding stock). Worldwide, the list of parasitoids is
long and complex (see Van Driesche et al., 1996, pp. 28-31, see under
web links for URL). Of these many parasitoids, the most important
in North America are two North American species, Aphytis diaspidis
(Howard) and A. mytilaspidis (Le Baron) and two Chinese parasitoids,
Encarsia perniciosi (Tower) and Coccophagoides kuwanae (Silvestri).
In much of the United States, this scale is now generally held at
acceptable levels by some combination of these parasitoids, unless they
are absent naturally or have been suppressed by local use of pesticides.

Web Links for Information http://www.fs.fed.us/foresthealth/technology/pdfs/FHTET_96_19.pdf;

on San José Scale Van Driesche et al. 1996; see pages 28-31.
http://www.virginiafruit.ento.vt.edu/SJS.html; fact sheet from Virginia
Tech.
http://www.inra.fr/hyppz/RAVAGEUR/6quaper.htm; good photographs.
http://jenny.tfrec.wsu.edu/opm/displaySpecies.php?pn=490; Washington
State University—lengthy, high quality fact sheet; excellent photo gallery
on species.
http://www.nysipm.cornell.edu/factsheets/treefruit/pests/sjs/sjs.asp; fact
sheet from Cornell University.
http://www.ipm.ucdavis.edu/PMG/r4301911.html; University of
California at Davis fact sheet covering life history and use of pesticides
for control on fruit.
http://www.canr.msu.edu/vanburen/sanjosefs.html; fact sheet from
Michigan State University.
http://www.caf.wvu.edu/kearneysville/pest_month/insectfocusapril98.
htm; fact sheet of University of California.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Diaspidiotusperniciosus.
htm; ScaleNet catalog, giving a complete list of all recorded natural
enemies, hosts, and distribution records, and an index to all other
information published on this insect.
A Photographic Guide 109

Articles CAB (Commonwealth Agricultural Bureaux). 1986. International

Institute of Entomology, Distribution maps of pests, No. 7.
Commonwealth Agricultural Bureaux, United Kingdom. [Map of world
distribution of San José scale].
110 Forest Pest Insects in North America:

37. Willow Scale, Diaspidiotus gigas (Thiem and Gerneck) (Hemiptera:

Diaspididae)

Orientation to Pest Willow scale, Diaspidiotus gigas (Thiem and Gerneck), is an invasive
scale found in North America. It is also widely distributed across the
northern temperate zone in the Old World, from Algeria and western
Europe to Siberia and China. It has one generation per year in the
northern United States and overwinters as partly grown nymphs or young
adults. It feeds on branches of species of willow (Salix) and poplar
(Populus) and is found on both shade trees and in natural vegetation.

Hosts Commonly Attacked Willow scale feeds on species of willow (Salix) and poplar (Populus).

Distribution This scale is found across the United States, with specimens recorded
from both western parts of the country (Oregon, Washington, Idaho,
Montana, Colorado, Utah, and Wyoming), the midwest (Wisconsin,
Ohio) and the northeast (New York, Pennsylvania, Rhode Island). It is
also found in eastern Canada (Nova Scotia, New Brunswick, Ontario).

Images of Willow Scale

Figure 1. Mixed stages of willow scale, Diaspidiotus gigas, on trunk of an aspen

tree. (Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 2. Close up view

of mixed stages of willow
scale on aspen. (Whitney
Cranshaw, Colorado State
University, Bugwood.org)
A Photographic Guide 111

Images of Willow Scale

(continued)

Figure 3. Crawlers of willow scale escaping from female scale. (Bob Hammon,
Bugwood.org)

Figure 4. Overwintering stage (2nd instar nymphs) of willow scale on bark.

(Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 5. Two views of damage from willow scale to aspen bark; left view of
whole trunk showing “bubbling” or pitting effect on bark (middle of photo) and
on right, close up of damage. (Both photos: Whitney Cranshaw, Colorado State
University, Bugwood.org)
112 Forest Pest Insects in North America:

Images of Willow Scale Figure 6. Aspen trees in

Aspen, Colorado, that are
(continued) heavily infested with willow
scale. (Bob Hammon,
Bugwood.org)

Figure 7. Chilocorus
kuwanae, a predator of
willow scale in its native
range in China. (Tom Murray,
Bugwood.org)
A Photographic Guide 113

Important Biological Control The aphelinid parasitoids Pteroptrix longgiclava (Girault) and Encarsia
Agents Related to this gigas (Tshumakova) attack willow scale in China and are attracted to
Pest Species volatiles from the scale, suggesting these may be specialized species of
value in biological control. Other parasitoids known to attack this scale
in Europe or other regions include Aphytis mytilaspidis (Le Baron),
Pteroptrix dimidiatus (Westwood), and Azotus matritensis Merc. Aphytis
mytilaspidis, however, is a widely distributed polyphagous species that
attacks many diaspidids. Predators known to feed on this scale include
Chilocorus kuwanae Silvestri (an Asian ladybird already introduced
into North America for control of another invasive scale, Unaspis
euonymi [Comstock]), Chilocorus bipustulatus (L.) and Coccinella
bipunctata ( L.). The native North American species Chilocorus stigma
has been observed feeding on D. gigas in Colorado. Other potential
willow scale predators known from Europe include the predatory bugs
Loricula pselaphiformis Curt. and Loricula elengatula Baerensprung
(Microphysidae) and the anthocorids Ectemnus nigriceps E. Wagner and
Temnostethus longirostris (Horv.).

Web Links for Information http://www.fs.fed.us/r1-r4/spf/fhp/publications/bystate/BFO-PR-08-01_

on Willow Scale woodriver.pdf; photos and description of recent problems on aspen from
willow scale in Idaho.
http://wci.colostate.edu/shtml/AspenScale.shtml; Colorado State
University factsheet.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Diaspidiotusgigas.htm;
ScaleNet catalog, giving a complete list of all recorded natural enemies,
hosts, and distribution records, and an index to all other information
published on this insect.

Articles Lelláková-Dusková, F. 1963. The morphology, metamorphosis, and

lifecycle of the scale insect Quadraspidiotus gigas (Thiem and Gerneck).
Acta Entomologica Musei Nationalis Prague 5: 611-648.
Nakahara, S. 1982. A Checklist of the Armored Scales of the Coterminous
United States. Plant Protection and Quarantine Animal and Plant Health
Inspection Service, United States Department of Agriculture. 110 p.
Chi, D.F., S.C. Yan, X.H. Zhao, and Z.H. Wen. 2003 The taxis of chalcid
parasitoids to their host and analysis of the volatile[s] from fixed first
instar nymph[s] of Quadraspidiotus gigas (Thiem & Gerneck). Journal
of Northeast Forestry University 31(2): 20-22.
Chi, D.F., R.O. Rubio, S.C. Yan, D.D. Zhang, and Z.H. Wen. 2002.
Foraging behavior of parasitoid chalcid to the essential oil from bark of
Populus pseudo-simonii × P. nigra and Quadraspidiotus gigas. Journal
of Forestry Research 13(4): 255-259.
Ma, L., C.D. Li, J.Q. Liu, Y. Sun, X.L. Wang, Y.J. Ji, W.S. Du, L.
Yang, G.L. Su, and B.H. Zhao. 1997. Predatory function of Chilocorus
kawanae Silvestri on Quadraspidiotus gigas (Thiem et Gerneck).
Journal of Northeast Forestry University 25(2): 64-67.
114 Forest Pest Insects in North America:

38. Camphor Scale, Pseudaonidia duplex (Cockerell) (Hemiptera:

Diaspididae)

Orientation to Pest Camphor scale (also called camellia scale), Pseudaonidia duplex
(Cockerell), is an invasive Asian scale found in the southern United
States. The scale infests leaves, twigs and fruit of various plants, but
especially the camphor tree (Cinnamomum camphora [L.] Sieb.). In Asia,
it is a pest of tea plantations. In Louisiana, the scale has three generations
per year and overwinters primarily as mated adult females.

Hosts Commonly Attacked This scale feeds on over 200 different host plants in Louisiana alone,
including the camphor-tree (C. camphora) and some species of citrus
(Citrus). In Asia it is a pest of tea (Camellia sinensis [L.] Kuntze).

Distribution The United States distribution of this scale reaches from Florida and
Georgia, west to Texas.

Images of Camphor Scale

Figure 1. Camphor scale, Pseudaonidia duplex, adults; adult and one nymph on
fruit (left) and adults and several nymphs on foliage (right). (Both photos: Lyle
Buss, University of Florida, Bugwood.org)

Figure 2. Camphor scales on Virginia creeper (Parthenocissus quinquefolia [L.]

Planch.) leaf. (Lyle Buss, University of Florida, Bugwood.org)
A Photographic Guide 115

Images of Camphor Scale

(continued)

Figure 3. Camphor scales on wax-myrtle (Myrica spp.), with scale covers

turned over to reveal scale bodies (orange). (Lyle Buss, University of Florida,
Bugwood.org)

Figure 4. Camphor scales on wax-myrtle twig. (Lyle Buss, University of Florida,

Bugwood.org)

Important Biological Control In Taiwan in tea plantations, this scale is attacked by the parasitoid
Agents Related to this Neochrysocharis sp., which can cause 42-61 percent mortality
Pest Species (Shiao, 1978).

Web Links for Information http://www.doacs.state.fl.us/pi/enpp/ento/entcirc/ent028.pdf; a Florida

on Camphor Scale entomology circular that gives a host list.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Pseudaonidiaduplex.htm;
ScaleNet catalog, giving a complete list of all recorded natural enemies,
hosts, and distribution records, and an index to all other information
published on this insect.
116 Forest Pest Insects in North America:

Articles on Camphor Scale Shiao, S.N. 1977. Bionomics of the camellia scale, Pseudaonidia duplex
(Cockerell), in northern parts of Taiwan. I. Life history and mortality
factors. Plant Protection Bulletin, Taiwan 19(2): 65-77.
Shiao, S.N. 1978. Bionomics of the camellia scale, Pseudaonidia duplex
(Cockerell) (Homoptera: Diaspididae) in northern parts of Taiwan. II.
Natural enemies. Plant Protection Bulletin, Taiwan 20(3): 210-223.
A Photographic Guide 117

39. Coconut Scale, Aspidiotus destructor Signoret (Hemiptera: Diaspididae)

Orientation to Pest Coconut scale, Aspidiotus destructor Signoret, has an extremely broad
host range, feeding on over 60 different families of plants. As a pest of
coconut (Cocos nucifera L.), it normally feeds on the undersurfaces of
fronds, but when infestations are very dense, even fruits may be infested.
Palms may be killed by heavy infestations, but in most cases, predators
suppress the scale to less destructive levels.

Hosts Commonly Attacked Coconut scale is a common pest of coconut (C. nucifera) and banana
(Musa), and also infests avocado (Persea americana Miller), bird of
paradise (Strelitzia), breadfruit (Artocarpus altilis [Parkinson] Fosberg),
cassava (Manihot esculenta Crantz), cotton (Gossypium), ginger
(Zingiber officinale Roscoe), guava (Psidium), mango (Mangifera indica
L.), mock orange (Philadelphus), mountain apple (Syzygium malaccense
[L.] Merr. and L.M. Perry), oil palm (Elaeis), papaya (Carica papaya
L.), pandanus (Pandanus), frangiapani (Plumeria), rubber trees (Hevea
brasiliensis Müll. Arg.), sugarcane (Saccharum), and tea (Camellia
sinensis [L.] Kuntze).

Distribution At a world level, this scale is widely distributed in tropical areas,

including Puerto Rico. The mainland United States distribution of this
scale includes Florida, Georgia, North Carolina, and Virginia.

Images of Coconut Scale

Figure 1. Several adults of coconut Figure 2. Crawlers of coconut scale

scale, Aspidiotus destructor, interspersed among large adults.
on a palm frond. (United States (United States National Collection of
National Collection of Scale Insects Scale Insects Photographs Archive,
Photographs Archive, USDA USDA Agricultural Research Service,
Agricultural Research Service, Bugwood.org)
Bugwood.org)
118 Forest Pest Insects in North America:

Images of Coconut Scale

(continued)

Figure 3. Colonies of various density of coconut scale on palm fronds.

(Both photos: United States National Collection of Scale Insects Photographs
Archive, USDA Agricultural Research Service, Bugwood.org)

Figure 4. Discolored streaks

in palm fronds are a sign
of coconut scale damage.
(United States National
Collection of Scale Insects
Photographs Archive, USDA
Agricultural Research
Service, Bugwood.org)
A Photographic Guide 119

Images of Coconut Scale Figure 5. Coconuts heavily

infested with coconut scale.
(continued) (Infonet B. Loehr)

Figure 6. Encarsia citrina

is a polyphagous aphelinid
that attacks many diaspidid
scales, including coconut
scale. (Kristopher Abell,
University of Massachusetts,
Bugwood.org)

Important Biological Control A great many natural enemies of coconut scale have been reported (see
Agents Related to this summary in Waterhouse and Norris, 1987). See also discussion in Frank
Pest Species and Foltz (1996) for specific natural enemies present in Florida and
Puerto Rico (see articles below for URL). Among the most important of
these are the ladybird beetle Cryptognatha nodiceps Marshall and the
parasitoid Encarsia citrina (Crawford).

Web Links for Information http://www.spc.int/pps/pest_of_the_month_-_january_2005.htm;

on Coconut Scale discusses biology and control of coconut scale with reference to islands
of the Pacific.
http://www.extento.hawaii.edu/kbase/crop/type/a_destru.htm; fact sheet
from the Hawaiian Extension Service.
http://www.ctahr.hawaii.edu/adap/Publications/ADAP_pubs/2000-5.pdf;
fact sheet of ADAP (Agricultural Development in American Pacific).
http://www.fs.fed.us/foresthealth/technology/pdfs/FHTET_96_20.PDF;
Frank and Foltz (1996); FHTET Bulletin of USDA Forest Service, see
page 13.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Aspidiotusdestructor.htm;
ScaleNet catalog, giving a complete list of all recorded natural enemies,
hosts, and distribution records, and an index to all other information
published on this insect (over 470 references).

Article Waterhouse, D.F. and K.R. Norris. 1987. Biological Control: Pacific
Prospects. ACIAR, Inkata Press, Melbourne, Australia: 62-71.
120 Forest Pest Insects in North America:

40. Florida Red Scale, Chrysomphalus aonidum (L.) (Hemiptera:

Diaspididae)

Orientation to Pest Florida red scale, Chrysomphalus aonidum (L.), is an Asian scale
invasive in the United States. It is one of the most polyphagous insects
known, recorded from 70 families of plants. It is most often reported on
agricultural or ornamental plants but occurs in natural vegetation as well.
The scale feeds on foliage, producing yellow areas. Heavy infestations
may cause foliage to drop.

Hosts Commonly Attacked Citrus (Citrus), holly (Ilex), and palms are the more commonly affected
groups of plants.

Distribution The U.S. distribution of this scale is from North Carolina, south and west
to Texas, as well as Hawaii. It is common in greenhouses throughout the
country. It is also common in most tropical and subtropical parts of the
world, including Puerto Rico.

Images of
Florida Red Scale

Figure 1. Florida red scale, Chrysomphalus aonidum, on dragontree (Dracaena

marginata Lam.) (left) and on citrus leaf (right). (Left: United States National
Collection of Scale Insects Photographs Archive, USDA Agricultural Research
Service, Bugwood.org; right: Pedro Torrent Chocarro, Bugwood.org)

Figure 2. Florida red scale

on an orange. (Pedro Torrent
Chocarro, Bugwood.org)
A Photographic Guide 121

Important Biological Control This scale has been the focus of parasitoid introductions in Texas and
Agents Related to this Florida. The aphelinid Aphytis holoxanthus DeBach from Israel has
Pest Species provided substantial control in both locations.

Web Links for Information http://www.biosecurity.govt.nz/pests-diseases/plants/florida-red-scale/frs-

on Florida Red Scale report.htm; a New Zealand fact sheet.
http://www.sel.barc.usda.gov/catalogs/diaspidi/Chrysomphalusaonidum.
htm; ScaleNet catalog, giving a complete list of all recorded natural
enemies, hosts, and distribution records, and an index to all other
information published on this insect.

Articles Clancy, D.W., A.G. Selhime, and M.H. Muma. 1963. Establishment of
Aphytis holoxanthus as a parasite of Florida red scale in Florida. Journal
of Economic Entomology 56: 603-605.
Steinberg, S., H. Podoler, and D. Rosen. 1986. Biological control of the
Florida red scale, Chrysomphalus aonidum, in Israel by two parasite
species: Current status in the coastal plain. Phytoparasitica 14: 199-204.
122 Forest Pest Insects in North America:

41. Magnolia White Scale, Pseudaulacaspis cockerelli (Cooley) (Hemiptera:

Diaspididae)

Orientation to Pest Magnolia white scale, Pseudaulacaspis cockerelli (Cooley) (known

formerly as the false oleander scale), is an invasive Asian scale found
in the southern United States. It is one of the most polyphagous insects
known, reported from 75 different plant families.

Hosts Commonly Attacked Common host plants include magnolia (Magnolia), bird of paradise
(Strelitzia), oleander (Nerium oleander L.), azalea (Azalea), plumeria
(Plumeria), mango (Mangifera indica L.), kukui (Aleurites moluccana
[L.] Willd.), cycads, and palms.

Distribution This scale is likely native to southern Asia, but is invasive in many parts
of the world, including Hawaii and Puerto Rico. In North America, it
is invasive in the southern United States (Florida to Maryland, west to
Texas and Kansas).

Images of Magnolia
White Scale

Figure 1. Adults of magnolia white scale, Pseudaulacaspis cockerelli, on areca

palm (Areca sp.) (left); close up of magnolia white scale adults (right). (Left:
United States National Collection of Scale Insects Photographs Archive, USDA
Agricultural Research Service, Bugwood.org; right: Gillian W. Watson, California
Department of Food & Agriculture)
A Photographic Guide 123

Images of Magnolia
White Scale (continued)

Figure 2. Parasitoid emergence hole in cover of white magnolia scale.

(Charles Olsen, USDA APHIS PPQ, Bugwood.org)

Important Biological Control Some specialized parasitoids of this scale are known from its native
Agents Related to this south Asian range, but none have been used for biological control of this
Pest Species scale in North America.

Web Links for Information http://edis.ifas.ufl.edu/pdffiles/IN/IN30600.pdf; fact sheet of the

on Magnolia White Scale University of Florida.
http://www.extento.hawaii.edu/kbase/crop/Type/p_cocker.htm; fact sheet
of the Hawaiian Cooperative Extension Service on biology and control.
http://www.sel.barc.usda.gov/catalogs/diaspidi/
Pseudaulacaspiscockerelli.htm; ScaleNet catalog, giving a complete list
of all recorded natural enemies, hosts, and distribution records, and an
index to all other information published on this insect.

Articles Rosen, D. and P. DeBach. 1986. Three new species of Aphytis

(Hymenoptera: Aphelinidae), parasites of Pseudaulacaspis spp.
(Homoptera: Diaspididae) in India and Australia. Entomophaga 31:
139-151.
124 Forest Pest Insects in North America:

42. White Peach Scale, Pseudaulacaspis pentagona (Targioni-Tozzetti)

(Hemiptera: Diaspididae)

Orientation to Pest White peach scale, Pseudaulacaspis pentagona (Targioni-Tozzetti), is an

invasive scale found in the southeast United States, likely native to Japan
or China. It has one of the broadest host ranges reported for any insect,
attacking 85 families of plants, including more than 115 genera in Florida
alone. The scale has four generations per year in Florida and three in the
northern parts of its U.S. range.

Hosts Commonly Attacked A polyphagous species, white peach scale is common on peach (Prunus
persica [L.] Batsch), mulberry (Morus), persimmon (Diospyros
virginiana L.), catalpa (Catalpa), hibiscus (Hibiscus), and privet
(Ligustrum).

Distribution In the United States, white peach scale is found from Florida west to
Texas and north to Maryland and Tennessee. It also occurs in Puerto Rico
and Hawaii.

Images of White
Peach Scale

Figure 1. Close up of adults of white peach scale, Pseudaulacaspis pentagona.

(Left: Pennsylvania Department of Conservation and Natural Resources -
Forestry Archive, Bugwood.org; right: Jeffrey W. Lotz, Florida Department of
Agriculture and Consumer Services, Bugwood.org)

Figure 2. Adult female of white peach scale with scale cover removed, showing
eggs. (Lyle Buss, University of Florida, Bugwood.org)
A Photographic Guide 125

Images of White
Peach Scale (continued)

Figure 3. Dense bark infestations of white peach scale. (Both photos:

Eric R. Day, Virginia Polytechnic Institute and State University, Bugwood.org)

Important Biological Control Two species of Encarsia (E. berlesei Howard and E. diaspidicola
Agents Related to this Silvestri) are important parasitoids of white peach scale.
Pest Species

Web Links for Information http://entnemdept.ufl.edu/creatures/orn/scales/white_peach_scale.htm;

on White Peach Scale fact sheet of the University of Florida.
http://www.sel.barc.usda.gov/catalogs/diaspidi/
Pseudaulacaspispentagona.htm; ScaleNet catalog, giving a complete list
of all recorded natural enemies, hosts, and distribution records, and an
index to all other information published on this insect.

Articles Neumann, G., P.A. Follett, R.G. Hollingsworth, and J.H. de León.
2010. High host specificity in Encarsia diaspidicola (Hymenoptera:
Aphelinidae), a biological control candidate against the white peach scale
in Hawaii. Biological Control 54: 107-113.
León, J.H. de, G. Neumann, P.A. Follett, and R.G. Hollingsworth.
2010. Molecular markers discriminate closely related species Encarsia
diaspidicola and Encarsia berlesei (Hymenoptera: Aphelinidae):
biocontrol candidate agents for white peach scale in Hawaii. Journal of
Economic Entomology 103: 908-916.
Kreiter, P. and A. Panis. 1998. Inventory of the natural enemies of
the white peach scale Pseudaulacaspis pentagona (Targioni-Tozzetti,
1886) in the world (Homoptera, Diaspididae). Bulletin de la Société
Entomologique de France 103: 263-271.
Hanks, L.M. and R.F. Denno. 1993. The white peach scale,
Pseudaulacaspis pentagona (Targioni-Tozzetti) (Hemiptera:
Diaspididae): life history in Maryland, host plants, and natural enemies.
Proceedings of the Entomological Society of Washington 95: 79-88.
126 Forest Pest Insects in North America:

43. Citrus Whitefly, Dialeurodes citri (Ashmead) (Hemiptera: Aleyrodidae)

Orientation to Pest Citrus whitefly, Dialeurodes citri (Ashmead), is a polyphagous invasive

whitefly of Asian origin that was once a major pest of citrus in the United
States, as well as infesting a range of other plants. Like all whiteflies, this
species has sessile nymphs that are affixed to plant surfaces, where they
feed by removing plant sap, debilitating the host. Citrus whitefly secretes
honeydew, leading to growth of sooty molds on infested plants.

Hosts Commonly Attacked Citrus whitefly attacks many species, most of which are themselves
introduced crops or ornamentals. These include citrus (Citrus), allamanda
(Allamanda), banana shrub (Michelia figo [Lour.] Spreng), cape jasmine
(Gardenia jasminoides J. Ellis), chinaberry (Melia azedarach L.), laurel
cherry (Prunus laurocerasus L.), crape myrtle (Lagerstroemia spp.),
Ficus macrophylla Desf. ex Pers., gardenia (Gardenia jasminoides J.
Ellis), myrtle (Myrtus spp.), mock olive (Notelaea), and pear (Pyrus).
Among the native North American plants attached are osage orange
(Maclura pomifera [Raf.] Schneid.), water oak (Quercus nigra L.),
persimmon (Diospyros), and green ash (Fraxinus pennsylvanica
Marshall).

Distribution In North America, citrus whitefly is found in the southeastern United

States, from Florida west to Texas and north to Virginia, and also in
California.

Images of Citrus Whitefly

Figure 1. Adults of citrus whitefly, Dialeurodes citri. (Both photos: Lyle Buss,
University of Florida, Bugwood.org)

Figure 2. Nymph of citrus

whitefly. (Florida Division of
Plant Industry Archive, Florida
Department of Agriculture
and Consumer Services,
Bugwood.org)
A Photographic Guide 127

Images of Citrus Whitefly

(continued)

Figure 3. Cast skin (top) of citrus whitefly. (Florida Division of Plant Industry
Archive, Florida Department of Agriculture and Consumer Services,
Bugwood.org)

Figure 4. Leaf infested with citrus whitefly (white dots dispersed over leaf).
(Jeffrey W. Lotz, Florida Department of Agriculture and Consumer Services,
Bugwood.org)
128 Forest Pest Insects in North America:

Images of Citrus Whitefly Figure 5. Sooty mold

growing on honeydew
(continued) produced by citrus
whitefly. (INRA:
Alain Fraval)

Figure 6. The aphelinid

parasitoid Encarsia
lahorensis, the
key natural enemy
of citrus whitefly.
(Jeffrey W. Lotz,
Florida Department
of Agriculture and
Consumer Services,
Bugwood.org)

Important Biological Control This invasive whitefly has come under virtually complete biological
Agents Related to this control, at least on citrus, due the influence of the introduced parasitoid
Pest Species Encarsia lahorensis Howard.
A Photographic Guide 129

Web Links for Information http://entnemdept.ufl.edu/creatures/citrus/citrus_whitefly.htm; a

on Citrus Whitefly University of Florida fact sheet on biology, hosts, distribution and
control of citrus whitefly.
http://edis.ifas.ufl.edu/pdffiles/IN/IN42700.pdf; a fact sheet of the
University of Florida, discussing the biology and life history of the
parasitoid Encarsia lahorensis.

Articles Bellows, T.S. and C. Meisenbacher. 2007. Field population biology of

citrus whitefly, Dialeurodes citri (Ashmead) (Heteroptera: Aleyrodidae),
on oranges in California. Population Ecology 49: 127-134.
130 Forest Pest Insects in North America:

44. The Mealybug Oracella acuta (Lobdell) (Hemiptera: Pseudococcidae)

Orientation to Pest The mealybug Oracella acuta (Lobdell) is native to the southeastern
United States. It has up to five generations per year in the southern part
of the range. First generation crawlers are the mobile dispersal stage.
Once female crawlers finish dispersing, they either settle on the shoot
or occasionally between the needles near the fascicle. Females secrete
a characteristic white resin cell that covers their body. The tips of new
shoots are the preferred settling site, though the entire shoot may be
colonized when populations are high. The resin cells, shoots, and needles
may become covered with black, sooty mold growing on honeydew
produced by the mealybug. Males develop on the shoot within a covering
called a test. First generation males are apterous, with subsequent
generation males being alate. Infestations rarely cause tree mortality,
but they may severely retard growth. Oracella acuta is a secondary pest,
usually becoming a problem following insecticide applications targeting
other insects.

Hosts Commonly Attacked Hosts of this mealybug include loblolly (Pinus taeda L.), slash (P.
elliottii Engelm.), Virginia (P. virginiana Miller), shortleaf (P. echinata
Miller), and longleaf (P. palustris Miller) pine.

Distribution In North America, this mealybug occurs from Florida north to Virginia
and west to Texas. It is also invasive in China, where it has become a
major pest in slash pine plantations.

Images of the Mealybug Figure 1. A colony of the

mealybug Oracella acuta
Oracella acuta on slash pine. (William
M. Ciesla, Forest Health
Management International,
Bugwood.org)
A Photographic Guide 131

Images of the Mealybug Figure 2. Close up of

the mealybug Oracella
Oracella acuta (continued) acuta, showing an
adult female and her
eggs. (Steve Clarke,
USDA Forest Service,
Bugwood.org)

Figure 3. A resin cell,

formed by feeding of
the mealybug Oracella
acuta. (Steve Clarke,
USDA Forest Service,
Bugwood.org)
132 Forest Pest Insects in North America:

Important Biological Control Three native parasitoids, Zarhopalus debarri Sun, Acerophagus coccois
Agents Related to this E. Smith, and Allotropa oracellae Masner, help regulate this mealybug’s
Pest Species population size in the southeast United States. All three parasitoids were
imported to China and released in heavily infested slash pine plantations,
but failed to suppress the pest. One hypothesis about this failure is that it
is due to the lack of mealybugs available as hosts in the summer months
in China.

Web Links for Information None

on the Mealybug
Oracella acuta

Articles Clarke, S.R., G.L. Debar, and C.W. Berisford. 1990. Life history of
Oracella acuta (Homoptera: Pseudococcidae) in loblolly pine seed
orchards in Georgia. Environmental Entomology 19: 99-103.
Sun, J.-H., S.R. Clarke, G.L. Debarr, and C.W. Berisford. 2004.
Parasitoid complex of the mealybug Oracella acuta (Lobdell)
(Hemiptera: Pseudococcidae) in Georgia, USA. Journal Entomological
Science 39(1): 11-22. (Available at http://www.srs.fs.usda.gov/pubs/ja/
ja_sun005.pdf).
Clarke, S.R., H.-B. Yu, M.-R. Chen, G.L. Debarr, and J.-H. Sun. 2010.
Classical biological control program for the mealybug Oracella acuta
in Guangdong Province, China. Insect Science 17: 129-139. (Available
at http://onlinelibrary.wiley.com/doi/10.1111/j.1744-7917.2009.01292.
x/pdf).
A Photographic Guide 133

45. Cottonwood Leaf Beetle, Chrysomela scripta Fabricius (Coleoptera:

Chrysomelidae)

Orientation to Pest Cottonwood leaf beetle, Chrysomela scripta Fabricius, is a native insect
in North America that feeds on various species of poplar (Populus),
willow (Salix), and alder (Alnus) and is found throughout the United
States. Adults and larvae feed on leaves, and the yellow eggs are
laid in clusters on foliage. Young larvae feed in groups and are leaf
skeletonizers. Young larvae emit chemical exudates for defense against
predators. Older larvae and adults feed individually. Pupation is on
the host plant or nearby materials. Adults overwinter under bark, litter
or forest debris, and there are several generations per year (3-4 in the
midwestern United States and up to 6-8 in the southern United States).
Feeding results in reduced growth and multi-forked tops. Damage is of
greatest concern in cottonwood or willow plantations.

Hosts Commonly Attacked Various species of poplar (Populus) and occasionally willow (Salix) are
attacked by this beetle.

Distribution Cottonwood leaf beetle is found throughout the United States.

Images of Cottonwood
Leaf Beetle

Figure 1. Adults (left) of the cottonwood leaf beetle, Chrysomela scripta, on

leaf, and close up of adult beetle (right). (Left: Andrew J. Boone, South Carolina
Forestry Commission, Bugwood.org; right: Gerald J. Lenhard, Louiana State
University, Bugwood.org)

Figure 2. An egg mass (yellow) of cottonwood leaf beetle on leaf (left); a close
view of an egg mass (right). (Left: James Solomon, USDA Forest Service,
Bugwood.org; right: Lacy L. Hyche, Auburn University, Bugwood.org)
134 Forest Pest Insects in North America:

Images of Cottonwood
Leaf Beetle (continued)

Figure 3. A group of cottonwood leaf beetle larvae feeding on leaf (left); a

close view of a single larva (right). (Left: Lacy L. Hyche, Auburn University,
Bugwood.org; right: Gerald J. Lenhard, Louisiana State University, Bugwood.org)

Figure 4. Dorsal view

of pupa of cottonwood
leaf beetle. (Gerald J.
Lenhard, Louiana State
University, Bugwood.org)

Figure 5. Damage to eastern cottonwood terminal from cottonwood leaf beetle

(left); and close up of damage to leaf from group of feeding larvae (right). (Left:
James Solomon, U.S. Forest Service, Bugwood.org; right: Lacy L. Hyche, Auburn
University, Bugwood.org)

Figure 6. Eggs of
cottonwood leaf beetle
are preyed on by many
generalist predators;
here, by the ladybird
beetle Neoharmonia
venusta (Melsheimer).
(James Solomon,
U.S. Forest Service,
Bugwood.org)
A Photographic Guide 135

Important Biological Control Natural enemies of this beetle include the microsporidian Nosema scripta
Agents Related to this Bauer and Pankratz, the ladybird beetle Coleomegilla maculata (De
Pest Species Geer), and the pteromalid parasitoid Shizonotus latus (Wlk.).

Web Links for Information http://wiki.bugwood.org/Archive:South/Chrysomela_scripta; article in

on Cottonwood Leaf Beetle Bugwood Wiki.
http://www.ento.okstate.edu/ddd/insects/leafbeetle.htm; a fact sheet of
Oklahoma State University compares willow leaf beetle, Chrysomela
knabi with cottonwood leaf beetle, Chrysomela scripta.
http://aces.nmsuledu/pubs/_circulars/circ552.html; provides information
on control.

Articles Bauer, L.S. and H.S. Pankratz. 1993. Nosema scripta n. sp.
(Microsporida: Nosematidae), a microsporidian parasite of
the cottonwood leaf beetle, Chrysomela scripta (Coleoptera:
Chrysomelidae). Journal of Eukaryotic Microbiology 40(2): 135-141.
Andersen, D.C. and S.M. Nelson. 2002. Effects of cottonwood leaf
beetle, Chrysomela scripta (Coleoptera: Chrysomelidae), on survival
and growth of Fremont cottonwood (Populus fremontii) in northwest
Colorado. American Midland Naturalist 147: 189-203.
Tenczar, E.G. and V.A. Krischik. 2006. Management of cottonwood
leaf beetle (Coleoptera: Chrysomelidae) with a novel transplant soak
and biorational insecticides to conserve coccinellid beetles. Journal of
Economic Entomology 99: 102-108.
Coyle, D.R., E.R. Hart, J.D. McMillin, L.C. Rule, and R.B. Hall. 2008.
Effects of repeated cottonwood leaf beetle defoliation on Populus growth
and economic value over an 8-year harvest rotation. Forest Ecology and
Management 255(8/9): 3365-3373.
136 Forest Pest Insects in North America:

46. Imported Willow Leaf Beetle, Plagiodera versicolora (Laicharting)

(Coleoptera: Chrysomelidae)

Orientation to Pest Imported willow leaf beetle, Plagiodera versicolora (Laicharting), is an

introduced insect in North America of European or Asian origin. Adults
overwinter in protected locations near host trees. Adults begin feeding
in early spring at leaf expansion and lay clusters of oval yellow eggs
on leaves. Both larvae and adults are foliage feeders, skeletonizing and
causing shot hole damage, respectively, to the leaves of willows and
poplars in both landscape plantings and natural habitats. There are two
(northern climates) to four (southern climates) generations per year.

Hosts Commonly Attacked Hosts of imported willow leaf beetle include various species of poplar
(Populus) and willow (Salix).

Distribution This insect is found in North America (throughout the eastern United
States and southern Canada, and in parts of Alaska), Central Europe,
Japan, and China. The exact native range is not known.

Images of Imported
Willow Leaf Beetle

Figure 1. Adult of imported willow leaf beetle, Plagiodera versicolora. (Jim Baker,
North Carolina State University, Bugwood.org)

Figure 2. Shot-hole type feeding damage caused by adults of imported willow

leaf beetle. (Lacy L. Hyche, Auburn University, Bugwood.org)
A Photographic Guide 137

Images of Imported
Willow Leaf Beetle
(continued)

Figure 3. Larvae and feeding damage of imported willow leaf beetle. (Paul
Weston, Cornell University, Bugwood.org)

Figure 4. Clump of willow showing branches defoliated by imported willow leaf

beetle. (U.S. Forest Service - Ogden Archive, USDA Forest Service,
Bugwood.org)

Figure 5. The coccinellid Neoharmonia venusta, a predator of imported willow

leaf beetle eggs. (James Solomon, USDA Forest Service, Bugwood.org)
138 Forest Pest Insects in North America:

Important Biological Control In North America, the chalcids Schizonotus rotundivenris (Girault) and
Agents Related to this Schizonotus latus (Walker) parasitize the imported willow leaf beetle.
Pest Species Eggs are eaten by predators, especially the coccinellid Neoharmonia
venusta (Melsheimer).

Web Links for Information on http://wiki.bugwood.org/Archive:Poplar/Plagiodera_versicolora

Imported Willow Leaf Beetle

Articles Wade, M.J. and F. Breden. 1986. Life history of natural populations of
the imported willow leaf beetle, Plagiodera versicolora (Coleoptera:
Chrysomelidae). Annals of the Entomological Society of America
79: 73-79.
A Photographic Guide 139

47. Elm Leaf Beetle, Xanthogaleruca luteola (Müller) (Coleoptera:

Chrysomelidae)

Orientation to Pest Elm leaf beetle, Xanthogaleruca luteola (Müller), is an introduced

insect in North America of European or Asian origin that feeds on elms
(Ulmus). It overwinters as an adult in protected locations. Eggs are laid
in clusters on elm leaves and larvae feed as leaf skeletonizers, while
adults produce shot-hole damage to leaves. Eggs are pointed, rather
than oval in outline. Pupation occurs at the base of infested trees. Two
generations may occur per year.

Hosts Commonly Attacked This beetle feeds on various imported or native elms (Ulmus), and is
important as a pest of American elm (Ulmus americana L.).

Distribution This invasive insect is found throughout the United States and southern
Canada, wherever elms occur naturally or have been planted as
ornamentals.

Images of
Elm Leaf Beetle

Figure 1. Adults of elm leaf beetle, Xanthogaleruca luteola. (Both photos:

Clemson University - USDA Cooperative Extension Slide Series, Bugwood.org)

Figure 2. Cluster of elm leaf beetle eggs. (Gyorgy Csoka, Hungary Forest
Research Institute, Bugwood.org)
140 Forest Pest Insects in North America:

Images of Elm Leaf

Beetle (continued)

Figure 3. Larvae of elm leaf beetle (young larvae, top; older larvae, middle;
close up of mature larva, bottom). (Top: Lacy L. Hyche, Auburn University,
Bugwood.org; middle, bottom: Gyorgy Csoka, Hungary Forest Research Institute,
Bugwood.org)
A Photographic Guide 141

Images of Elm Leaf

Beetle (continued)

Figure 4. Pupae of elm leaf beetle. (Whitney Cranshaw, Colorado State

University, Bugwood.org)

Figure 5. Elm leaves (top) skeletonized by elm leaf beetle larvae, and with shot-
hole damage from adult feeding (below). (Top: James Solomon, USDA Forest
Service, Bugwood.org; bottom: Pennsylvania Department of Conservation and
Natural Resources - Forestry Archive, Bugwood.org)
142 Forest Pest Insects in North America:

Images of Elm Leaf

Beetle (continued)

Figure 6. Oomyzus gallerucae is an important egg parasitoid affecting elm leaf

beetle populations in some parts of the United States. (Bruno Lavoue)

Figure 7. The tachinid Erynniopsis antennata is an introduced larval parasitoid

of elm leaf beetle. (Jack Kelly Clark, University of California - Statewide IPM
Program, Bugwood.org)

Important Biological Control Oomyzus gallerucae (Fonscolombe) (=Tetrastichus gallerucae [Boy])

Agents Related to this is an introduced egg parasitoid of elm leaf beetle that is found in
Pest Species Missouri, where it is of major importance, and in California. The tachinid
Erynniopsis antennata (Rondani) is an introduced larval parasitoid
of elm leaf beetle that is established in California and of moderate
importance there.
A Photographic Guide 143

Web Links for Information http://californiaagriculture.ucanr.org/landingpage.cfm?article=ca.

on Elm Leaf Beetle v052n02p18&fulltext=yes; an IPM plan for control of elm leaf beetle or
street trees in California.
http://www.ext.colostate.edu/pubs/insect/05521.html; fact sheet from
Colorado State University Extension.
http://extension.missouri.edu/publications/DisplayPub.aspx?P=g7356;
fact sheet from University of Missouri Extension.
http://www.hgic.umd.edu/_media/documents/hg113_000.pdf; fact sheet
from Maryland Cooperative Extension Service.

Articles Dreistadt, S.H. and D.L. Dahlsten. 1990. Distribution and abundance of
Erynniopsis antennata (Dipt.: Tachinidae) and Tetrastichus brevistigma
(Hym.: Eulophidae), two introduced elm leaf beetle parasitoids in
northern California. BioControl 35: 527-536.
Dreistadt, S.H. and D.L. Dahlsten. 1991. Establishment and
overwintering of Tetrastichus gallerucae (Hymenoptera: Eulophidae),
an egg parasitoid of the elm leaf beetle (Coleoptera: Chrysomelidae) in
northern California. Environmental Entomology 20: 1711-1719.
Meiners, T. and M. Hilker. 1997. Host location in Oomyzus gallerucae
(Hymenoptera: Eulophidae), an egg parasitoid of the elm leaf beetle
Xanthogaleruca luteola (Coleoptera: Chrysomelidae). Oecologia 112:
87-93.
Puttler, B. and W.C. Bailey. 2003. Establishment of Oomyzus gallerucae
(Hymenoptera: Eulophidae), an egg parasite of the elm leaf beetle
(Coleoptera: Chrysomelidae), in Missouri and adjacent states. Biological
Control 27: 20-24.
144 Forest Pest Insects in North America:

48. Larger Elm Leaf Beetle, Monocesta coryli (Say) (Coleoptera:

Chrysomelidae)

Orientation to Pest The larger elm leaf beetle, Monocesta coryli (Say), is a native insect in
the United States that is of concern in Florida as an occasional pest of
elms (Ulmus). It has a similar biology to the invasive elm leaf beetle,
Xanthogaleruca luteola (Müller), but is not as damaging, in part because
it has only one generation per year. Young larvae feed in groups on elm
leaves but older larvae disperse and feed singly. Larval feeding causes
leaves to become skeletonized, giving trees a brown or defoliated look.
Mature larvae leave trees to pupate in the soil.

Hosts Commonly Attacked The larger elm leaf beetle feeds on various elms (Ulmus).

Distribution This beetle occurs spottily from Florida to Pennsylvania, and west to
Ohio and Kansas.

Images of Large
Elm Leaf Beetle

Figure 1. Views of the adult larger elm beetle, Monocesta coryli. (Both photos:
Clemson University - USDA Cooperative Extension Slide Series, Bugwood.org)

Figure 2. Female of larger elm

leaf beetle with egg mass.
(North Carolina Forest Service
Archive, Bugwood.org)
A Photographic Guide 145

Images of Large
Elm Leaf Beetle (continued)

Figure 3. Young larvae of larger elm leaf beetle (left) feed in groups; close up
of one larva (right). (Both photos: Gerald J. Lenhard, Louiana State University,
Bugwood.org)

Figure 4. Defoliated elms fed on by larger elm leaf beetle. (Both photos: Jim
Baker, North Carolina State University, Bugwood.org)
146 Forest Pest Insects in North America:

Important Biological Control No natural enemies of this species have been well documented, but the
Agents Related to this tachinid Icelia triquetra (Olivier) is reported to attack this species.
Pest Species

Web Links for Information http://entnemdept.ufl.edu/creatures/trees/le_leaf_beetle.htm; fact sheet of

on Larger Elm Leaf Beetle the University of Florida.

Articles Anderson, D. and C.S. Papp. 1961. The larger elm leaf beetle, Monocesta
coryli (Say). Proceedings of the Entomological Society of Washington
63: 203-207.
Kelsheimer, E.G. 1945. Notes on the great elm leaf beetle. Florida
Entomologist 28: 25-27.
Kelsheimer, E.G. 1957. Larger elm leaf beetle (Monocesta coryli).
Cooperative Economic Insect Report, Plant Pest Control Division,
Agricultural Research Service, U.S. Department of Agriculture 7: 650.
Madden, A.H. 1940. Larger elm leaf beetle (Monocesta coryli Say).
Insect Pest Survey Bulletin, Bureau of Entomology and Plant Quarantine,
U.S. Department of Agriculture 20: 408.
A Photographic Guide 147

49. Giant Palm Weevil, Rhynchophorus cruentatus (F.) (Coleoptera:

Curculionidae)

Orientation to Pest Giant palm weevil (also called palmetto weevil), Rhynchophorus
cruentatus (F.), is native to the West Indies, South America, and
coastal areas of the southern United States. This species attacks various
introduced and native palms. The adult is variable in color pattern. Adults
emit aggregation pheromones to concentrate attack on stressed trees.
Eggs are laid in the bases of leaves or in wounds in a dying palm. Larvae
feed primarily in the soft tissue surrounding the apical meristem. Mature
grubs migrate to the periphery of the stem or petioles and prepare a
cocoon from palm fibers. The weevil attacks lightning-struck or stressed
palms, such as ones that have recently been transplanted. Healthy palms
in natural settings are rarely damaged.

Hosts Commonly Attacked The giant palm weevil is closely associated with cabbage palmetto (Sabal
palmetto [Walker] Loddiges ex J.A. et Schultes), which is native to the
southeastern United States. The native saw palmetto (Serrenoa repens
[Bart.] Small) is an alternate host. Infestations have also been found in
several introduced palms such as the Canary Island date palm (Phoenix
canariensis Hortorum ex Chabaud), Phoenix dactylifera L., Pritchardia
sp., Washingtonia sp., royal palms (Roystonea sp.), Latania sp., coconut
palm (Cocos nucifera L.), and Caryota sp.

Distribution This weevil is found in the United States in areas where palms grow,
from Florida to Texas and north along the coast to South Carolina.

Images of
Giant Palm Weevil

Figure 1. Adult of giant palm weevil (also called palmetto weevil), Rynchophorus
cruentatus. The adult is variable in color (right). (Left: Jim Occi, BugPics,
Bugwood.org; right: Doug Caldwell, University of Florida, Bugwood.org)
148 Forest Pest Insects in North America:

Images of Giant
Palm Weevil (continued)

Figure 2. Larvae of giant palm weevil in Bismark palm (Bismarckia nobilis

Hildebrandt and H.Wendl.). (Left: Wikipedia Commons; right: Lyle Buss,
University of Florida, Bugwood.org)

Figure 3. Cocoon of giant palm weevil, inside of which the larva pupates.
(Lyle Buss, University of Florida, Bugwood.org)

Figure 4. Left: Sabal palm (Sabal sp.)

killed by giant palm weevil; bottom:
Canary island date palm, Phoenix
canariensis, killed by giant palm weevil.
(Both photos: Robin M. Giblin-Davis)
A Photographic Guide 149

Important Biological Control Natural enemies of this species are not reported from North America but
Agents Related to this some have been noted in other parts of its range. However, since this
Pest Species weevil is a native species in the United States and since damage occurs
principally in planted palms in landscapes, the use of pesticides is likely
to be an adequate and more efficient method of control.

Web Links for Information http://entnemdept.ufl.edu/creatures/orn/palmetto_weevil.htm; fact sheet

on Giant Palm Weevil of the University of Florida.
http://www.doacs.state.fl.us/pi/pest_alerts/pdf/giantpalmweevils.pdf;
fact sheet “Giant palm weevils of the genus Rhynchophorus (Coleoptera:
Curculionidae) and their threat to Florida palms”; FDACS-Division of
Plant Industry, March 2010.

Articles Giblin-Davis, R.M. and F.W. Howard. 1989. Vulnerability of stressed

palms to attack by Rhynchophorus cruentatus (Fabricius) (Coleoptera:
Curculionidae) and insecticidal control of the pest. Journal of Economic
Entomology 82: 1185-1190.
Giblin-Davis, R.M., A.C. Oehschlager, A. Perez, G. Gries, R. Gries,
T.J.Weissling, C.M. Chinchilla, J.E. Peña, R.H. Hallett, and H.D. Pierce,
Jr. 1996. Chemical and behavioral ecology of palm weevils. Florida
Entomologist 79: 153-167. (Available at http://www.fcla.edu/FlaEnt/
fe79p153.pdf)
150 Forest Pest Insects in North America:

50. Red Palm Weevil, Rhynchophorus ferrugineus (Olivier) (Coleoptera:

Dryophthoridae)

Orientation to Pest The red palm weevil, Rhynchophorus ferrugineus (Olivier), is native
to south Asia and Melanesia, where it is a serious pest of palms. Since
the 1980s, this weevil has spread rapidly throughout many other parts
of the world, reaching the United States (southern California) in 2010.
The adult is 2-5 cm long and usually rusty red, but color variants
are common. Larvae feed by tunneling in the trunks of palm trees,
weakening and eventually killing trees. Red palm weevil is a major
pest of date palms (Phoenix dactylifera L.) and coconut palms (Cocos
nucifera L.). Palms damaged by red palm weevil show (1) foliage
injuries that include straight edges instead of pointed tips, notching of
lateral pinnae, and sometimes a linear series of “windows” across pinnae,
(2) tunnels in the trunk or at the base of fronds, (3) oozing of viscous
fluids from tunnels; (4) frass at entrance of feeding tunnels, (5) empty
pupal cases and the bodies of dead adults around heavily infested palms,
and (6) breaking of the trunk, or toppling of the palm crown. This weevil
is a pest of both cultivation (agricultural or landscaping palms) and
native palms in wildlands.

Hosts Commonly Attacked The red palm weevil is especially damaging economically to date
palms (P. dactylifera) and coconut palms (C. nucifera). Other hosts
that have been recorded include palms in the genera Areca, Arenga,
Borassus, Caryota, Corypha, Livistona, Metroxylon, Oreodoxa, Sabal,
Trachycarpus, and Washingtonia. Of potential concern in the United
States is the risk to native palms in the genera Washingtonia and Sabal.

Distribution Red palm weevil is from south Asia and Melanesia, but since the 1980s,
it has spread to Africa, Mediterranean Europe, the Caribbean, and
southern California (USA).

Images of Red
Palm Weevil

Figure 1. Adult red palm weevil (dark form found in California), Rhynchophorus
ferrugineus. (John Kabashima, UC Cooperative Extension, Bugwood.org)
A Photographic Guide 151

Images of Red
Palm Weevil (continued)

Figure 2. Larva and pupa of red palm weevil (red form of adult, from France).
(Christina Hoddle, University of California - Riverside, Bugwood.org)

Figure 3. Red palm weevil larva in open pupal cell. (Mike Lewis, Center for
Invasive Species Research, Bugwood.org)
152 Forest Pest Insects in North America:

Images of Red
Palm Weevil (continued)

Figure 4. Palm frond with typical damage from red palm weevil. (Mike Lewis,
Center for Invasive Species Research, Bugwood.org)

Figure 5. Landscape palm damaged by invasive red palm weevils.

(Christina Hoddle, University of California - Riverside, Bugwood.org)
A Photographic Guide 153

Images of Red Figure 6. Native

California fan palms,
Palm Weevil (continued) Washingtonia filifera
(Lindl.) H.Wendl., at
Mountain Palm Springs
in Anza Borrego State
Park in southern
California may be hosts
for red palm weevil.
(Bill Sullivan for
ABDNHA, Bugwood.org)

Important Biological Control Biological control of the red palm weevil has so far been focused on
Agents Related to this control methods that can be applied to agricultural palm plantations or
Pest Species landscape palms in urban areas. Should native palms in wildlands be
affected, classical biological control (based on importation of parasitoids
of the weevil from its native range) should be investigated.

Web Links for Information http://www.palms.org/palmsjournal/2002/redweevil.htm; a fact sheet

on Red Palm Weevil about this pest in the Mediterranean region.
http://cisr.ucr.edu/red_palm_weevil.html; University of California fact
sheet with photos of red palm weevil and its damage.
http://www.cdfa.ca.gov/phpps/rpw/index.html; California Department of
Agriculture site with photos of pest.

Articles Murphy, S.T. and B.R. Briscoe. 1999. The red palm weevil as an alien
invasive: biology and the prospects for biological control as a component
of IPM. Biocontrol News and Information 20(1): 35N-46N. (Available at
http://www.iraqi-datepalms.net/uploadedfiles/review%20article.pdf).
Faleiro, J.R. 2006. A review of the issues and management of the red
palm weevil Rhynchophorus ferrugineus (Coleoptera: Rhynchophoridae)
in coconut and date palm during the last one hundred years. International
Journal of Tropical Insect Science 26 (3): 135-154.
154 Forest Pest Insects in North America:

51. Pales Weevil, Hylobius pales (Herbst) (Coleoptera: Curculionidae)

Orientation to Pest The pales weevil, Hylobius pales (Herbst), is native to North America,
where it is the most serious pest of young seedling pines in both cutover
lands undergoing regeneration and in recently planted plantations or
Christmas tree plantations where stumps have not been removed. This
weevil is also the vector of Leptographium procerum, the causal agent of
Leptographium root disease. The species overwinters as adults beneath
litter or as larvae in roots. Eggs are laid in the roots of fresh stumps,
where larvae then create galleries and feed. Pupation occurs in the outer
sapwood. There is usually one generation per year and adults can live up
to 2 years. Adult feeding on young seedlings (up to 1 cm in diameter) is
the most important type of damage because it may cause girdling and tree
death. Adult feeding can also kill tips of branches. In cut over forest land,
damage can be lowered by delaying replanting for 1 or 2 years to allow
adults to emerge from infested stumps and leave the area. In Christmas
tree plantations, damage can be lowered by delaying replanting of cut
areas, treating stumps with pesticides to kill adult weevils, or maintaining
stumps in a living condition to prevent breeding (as in coppice harvesting
of Christmas trees).

Hosts Commonly Attacked The pales weevil affects all pines (Pinus spp.) native to North America
and also many exotic pines. It is also found in spruce (Picea spp.), fir
(Abies spp.), juniper (Juniperus spp.), larch (Larix spp.), hemlock (Tsuga
spp.), northern white cedar (Thuja occidentalis L.), and Douglas fir
(Pseudotsuga menziesii [Mirbel] Franco).

Distribution The pales weevil is found throughout eastern North America, west to the
Great Plains and north to southern Canada.

Images of
Pales Weevil

Figure 1. Adult of the pales weevil, Hylobius pales. (Clemson University - USDA
Cooperative Extension Slide Series, Bugwood.org)
A Photographic Guide 155

Images of Pales Weevil Figure 2. Larvae of

pales weevil, removed
(continued) from galleries.
(USDA Forest Service
publication “Weevils”)

Figure 3. Left, pale weevil larvae develop in roots of stumps left after harvest;
right, bark removed from stump to show the breeding galleries of pales weevil.
(Left: Eric R. Day, Virginia Polytechnic Institute and State University,
Bugwood.org; right: Mary Ann Hansen, Virginia Polytechnic Institute and
State University, Bugwood.org)

Figure 4. Feeding by adult

pales weevils can girdle
and kill seedlings (left) or
branches (right), destroying
forest regeneration or
damaging the shape and
appearance of Christmas
trees. (Left: USDA Forest
Service - Northeastern
Area Archive, USDA Forest
Service; right: Eric R. Day,
Virginia Polytechnic Institute
and State University,
Bugwood.org)
156 Forest Pest Insects in North America:

Important Biological Control The only known specialized natural enemy of this species is the
Agents Related to this euphorine braconid parasitoid Microctonus pachylobi Muesebeck, which
Pest Species develops in the adult weevil.

Web Links for Information http://www.forestpests.org/nursery/weevils.html; fact sheet on weevils of

on Pales Weevil forestry importance.
http://wiki.bugwood.org/Hylobius_pales; fact sheet on biology and
control of pales weevil.
http://pubs.ext.vt.edu/2902/2902-1102/2902-1102.pdf; fact sheet from
Virginia Tech.
http://www.fl-dof.com/publications/fh_pdfs/pine%20reproductive%20we
evils.pdf; short article on pine reproduction weevils.

Articles Zanzot, J.W., G. Matusick, and L.G. Eckhardt. 2010. Ecology of root-
feeding beetles and their associated fungi on longleaf pine in Georgia.
Environmental Entomology 39: 415-423.
Nevill, R.J. and S.A. Alexander. 1992. Transmission of Leptographium
procerum to eastern white pine by Hylobius pales and Pissodes
nemorensis (Coleoptera; Curculionidae). Plant Disease 76: 307-310.
Dixon, W.N. and J.L. Foltz. 1990. Pine reproduction weevils, Hylobius
pales (Herbst) and Pachylobius picivorus (Germar) (Coleoptera:
Curculionidae). Entomology Circular #332, University of Gainesville,
Florida.
Lynch, A.M. 1984. The pales weevil, Hylobius pales (Herbst): a
synthesis of the literature. Journal of the Georgia Entomological Society
19 (3, Suppl. 1).
Corneil, J.A. and L.F. Wilson. 1981. How to identify and control
pales weevil [Hylobius pales] in Christmas tree plantations. St. Paul,
Minnesota: North Central Forest Experiment Station, USDA Forest
Service: 5 p.
A Photographic Guide 157

52. Pine Root Collar Weevil, Hylobius radicis (Buchanan) (Coleoptera:

Curculionidae)

Orientation to Pest The pine root collar weevil, Hylobius radicis (Buchanan), is native
to eastern North America, where it is damaging in Scots pine (Pinus
sylvestris L.) Christmas tree farms and in red pine (Pinus resinosa Sol. ex
Aiton) plantations, but not natural forests. Eggs are laid in the root collar
zone of healthy trees. Larval feeding may kill young trees or badly stress
older trees. That stress can predispose mature trees to lethal attacks by
bark beetles. This weevil is associated with several fungi linked to red
pine decline. Damage may be reduced in a young stands before canopy
closure by pruning away lower branches and removing litter and soil
around the base of trunks to reduce weevil oviposition.

Hosts Commonly Attacked The most frequently affected species is Scots pine (P. sylvestris),
followed by red pine (P. resinosa), when grown in plantation.

Distribution The pine root collar weevil is found from Newfoundland in Canada,
south to Virginia and west to Minnesota and Manitoba.

Images of Pine
Root Collar Weevil

Figure 1. Adult of the pine root collar weevil, Hylobius radicis. (Jennifer C. Giron
Duque, University of Puerto Rico, Bugwood.org)
158 Forest Pest Insects in North America:

Images of Pine Root

Collar Weevil (continued)

Figure 2. Larva of pine root collar weevil dissected out of wood. (James B.
Hanson, USDA Forest Service, Bugwood.org)

Figure 3. Damage of pine root collar weevil in a mixed red and Scots pine stand.
(Steven Katovich, USDA Forest Service, Bugwood.org)

Important Biological Control Very few natural enemies have been recorded attacking pine collar
Agents Related to this weevil, the main ones being the braconids Bracon radicis Shenefelt and
Pest Species Miller and Microctonus pachylobi Muesebeck.

Web Links for Information on http://wiki.bugwood.org/Archive:Northeast/Hylobius_radicis; fact sheet

Pine Root Collar Weevil from Bugwood Wiki.
A Photographic Guide 159

Articles Wilson, L.F. and I. Millers. 1983. Pine root collar weevil – its ecology
and management. Technical Bulletin No. 1675, United States Department
of Agriculture, Washington, DC: 34 p.
Klepzig, K.D., K.F. Raffa, and E.B. Smalley. 1991. Association of an
insect-fungal complex with red pine decline in Wisconsin. Forest Science
37(4): 1119-1139.
Rieske, L.K. and K.F. Raffa. 1993. Potential use of baited pitfall traps in
monitoring pine root weevil (Hylobius pales, H. radicis, and Pachylobius
picivorus) populations and infestation levels. Journal of Economic
Entomology 86: 475-485.
160 Forest Pest Insects in North America:

53. Pitch-eating Weevil, Pachylobius picivorus (Germar) (Coleoptera:

Curculionidae)

Orientation to Pest The pitch-eating weevil, Pachylobius picivorus (Germar), is native

pest of pines in the eastern United States that is especially important in
southern states. This weevil’s biology and damage are very similar to that
of the pales weevil, Hylobius pales (Herbst). Damage occurs in cut-over
stands of various kinds of pines, where adults feed either on the inner
bark of small stems of the remaining trees or on planted or naturally
regenerated pine seedlings. Seedling death is the most important kind of
damage. Eggs are laid in green roots of recently cut stumps or those of
dead or dying trees. Adults dig through soil to reach roots for oviposition.
Larvae feed in the roots’ cambial layer and eventually pupate there,
after having constructed a cell of wood chips. Harvest creates abundant
stumps, allowing population increase of this insect and focusing damage
on cut-over sites. This weevil is one of several species associated with
fungi believed to be the cause of decline of red pine (Pinus resinosa Sol.
ex Aiton).

Hosts Commonly Attacked This weevil feeds on various pines, but especially shortleaf (P. echinata
Mill.), loblolly (P. taeda L.), and slash pines (P. elliottii Engelm.) in
the southern United States, and Scots pine (P. sylvestris L.), in the
northeastern/northcentral part of the country.

Distribution The pitch-eating weevil is found throughout the eastern United States,
but is most common from Virginia to Florida and west to Oklahoma and
Texas.

Images of
Pitch-eating Weevil

Figure 1. Pitch-eating weevil, Pachylobius picivorus, on a pine stem (left), the

adult’s feeding site; and close up (right). (Left: Terry S. Price, Georgia Forestry
Commission, Bugwood.org; right: Marvin Smith)
A Photographic Guide 161

Images of Pitch-eating Figure 2. Feeding on

pine stem by adults of
Weevil (continued) the pitch-eating weevil.
(Robert L. Anderson,
USDA Forest Service,
Bugwood.org)

Figure 3. Larva of pitch-eating weevil, extracted from a root, the larval feeding
site. (Wayne N. Dixon, Florida Department of Agriculture and Consumer
Services, Bugwood.org)
162 Forest Pest Insects in North America:

Important Biological Control Very few specialized natural enemies have been recorded attacking pitch-
Agents Related to this eating weevil, the main one being Microctonus pachylobi Muesebeck,
Pest Species which attacks the adult stage.

Web Links for Information http://wiki.bugwood.org/Pachylobius_picivorus; Bugwood Wiki fact

on Pitch-eating Weevil sheet.

Articles Walker, A.I. 1975. The biology and habits of the pales weevil, Hylobius
pales (Herbst), and the pitcheating weevil, Pachylobius picivorus
(Germar). Dissertation Abstracts International, B 35(8): 3962.
Dixon, W.N. and J.L. Foltz. 1990. Pine reproduction weevils, Hylobius
pales (Herbst) and Pachylobius picivorus (Germar) (Coleoptera:
Curculionidae). Entomology Circular #332, University of Gainesville,
Florida.
Rieske, L.K. and K.F. Raffa. 1990. Use of a monitoring system to
evaluate pesticide efficacy and residual activity against two pine
root weevils, Hylobius pales and Pachylobius picivorus (Coleoptera:
Curculionidae), in Christmas tree farms. Great Lakes Entomology 23:
189-193.
Rieske, L.K. and K.F. Raffa. 1991. Effect of varying ethanol and
turpentine levels on attraction of two pine root weevil species,
Hylobius pales and Pachylobius picivorus (Coleoptera: Curculionidae).
Environmental Entomology 20: 48-52.
A Photographic Guide 163

54. White Pine Weevil, Pissodes strobi (Peck) (Coleoptera: Curculionidae)

Orientation to Pest White pine weevil, Pissodes strobi (Peck), is native to North America.
This weevil can breed in a wide range of native and introduced pine and
spruce species. Eastern populations attack especially eastern white pine
(Pinus strobus L.) and Norway spruce (Picea abies (L.) H. Karst.). In
the western United States, Engelmann spruce (P. engelmanni Hopkins)
and Sitka spruce (P. sitchensis [Bong] Carr.) are attacked. These western
populations were formerly felt to be separate species and were studied
under the names Pissodes engelmanni Hopkins and Pissodes sitchensis
Hopkins, respectively. Cross breeding experiments, however, found all
populations to be inter-fertile and thus these western populations were
synonymized under the name P. strobi, which is how the name is used
here. The important damage by this species is caused by larvae, which
tunnel in and kill leaders, inducing branching and crooked trunks. Adults
overwinter in the litter and feed on the cambium of the main terminals
of their hosts. Eggs are laid in these terminals and larvae feed as cambial
borers, often then girdling and killing the terminal. Larvae pupate in
chambers made in the woody part of the stem. This species is considered
the most serious pest of eastern white pine regeneration. Losses can be
very high (up to 40 percent) to merchantable timber in heavily infested
stands over the life of a cohort of trees. Attack is recognized by oozing
of pitch from the terminal, followed by wilting and terminal death (and
change of color to reddish brown). Signs of former attacks are seen
in bushy or crooked tree form. Damage to young white pines may be
partially reduced by maintaining a shady over-story of hardwoods,
as weevil oviposition is greatest in full sun. Natural enemies are not
sufficient to prevent significant losses when silvicultural factors present
are favorable to this insect.

Hosts Commonly Attacked A variety of native and introduced pines and spruces, especially
eastern white (Pinus strobus), and jack (P. banksiana Lamb.) pines and
Norway (Picea abies), white (P. glauca [Moench] Voss), Engelmann
(P. engelmanni), and Sitka (P. sitchensis) spruce.

Distribution The white pine weevil is found widely in both the eastern and western
United States and Canada, in the ranges of its principal pine and spruce
hosts.
164 Forest Pest Insects in North America:

Images of White Pine

Weevil

Figure 1. Adult of white pine weevil, Pissodes strobi (left), feeding on terminal;
close up (right) of adult. (Left: A. Steven Munson, USDA Forest Service,
Bugwood.org; right: U.S. Department of Agriculture Forest Service)

Figure 2. Group of white pine weevil larvae feeding in a stem (left), and close up
(right). (Left: Dave Powell, USDA Forest Service, Bugwood.org; right: Joseph
O’Brien, USDA Forest Service, Bugwood.org)
A Photographic Guide 165

Images of White Pine

Weevil (continued)

Figure 3. Leader of spruce (left) killed by white pine weevil; and close up (right)
of pitch oozing from pine leader fed on by adults of white pine weevil. (Left: Troy
Kimoto, Canadian Food Inspection Agency, Bugwood.org; right: E. Bradford
Walker, Vermont Department of Forests, Parks and Recreation, Bugwood.org)

Figure 4. Damage to form of pines from white pine weevil injury. Left, old pine
with multiple crooked trunks; right, bush-like pines with many leaders. (Left:
Steven Katovich, USDA Forest Service, Bugwood.org; right: Joseph O’Brien,
USDA Forest Service, Bugwood.org)
166 Forest Pest Insects in North America:

Important Biological Control Not surprisingly, given that this native insect has a large geographical
Agents Related to this range and feeds on several tree genera, a variety of parasitoids and
Pest Species predators have been found associated with white pine weevil brood,
including the parasitoids Eurytoma pissodes Girault, Microbracon
(now Bracon) pini Muesebeck, Dolichomitus terrebrans nubilipennis
(Viereck), Coeloides pissodes (Ashmead), and Allodorus crassigaster
(Provancher). The principal predator of importance is Lonchaea corticis
Taylor. See Mills and Fischer (1986) for a summary and discussion
of white pine weevil natural enemies. However, none of these species
reduce populations of white pine weevil to non-damaging levels.
Consequently, the concept of introducing new natural enemies from
Pissodes species native to Europe was studied (Kennis and Mills, 1994),
but no species were ever actually introduced.

Web Links for Information http://cfs.nrcan.gc.ca/subsite/weevil/home-accueil; fact sheet of Natural

on White Pine Weevil Resources Canada.
http://www.na.fs.fed.us/spfo/pubs/howtos/ht_white/white.htm; USDA
Forest Service bulletin on how to manage white pine weevil damage.
http://ir.library.oregonstate.edu/xmlui/handle/1957/11007; thesis
providing a hazard rating system for white pine weevil risk to Sitka
spruce in Oregon, USA.

Articles Mills, N.J. and P. Fischer. 1986. The entomophage complex of Pissodes
weevils, with emphasis on the value of P. validirostris as a source of
parasitoids for use in biological control, pp. 297-307. In: Roques, A.
(ed.). Proceedings of the 2nd Conference on the Cone and Seed insects
Working Party, Briancon, France, September 3-5, 1986.
Kenis, M. and N.J. Mills. 1994. Parasitoids of European species of the
genus Pissodes (Col: Curculionidae) and their potential for the biological
control of Pissodes strobi (Peck) in Canada. Biological Control 4(1):
14-21.
Daoust, G. and M.J. Mottet. 2006. Effect of white pine weevil (Pissodes
strobi Peck) on plantations of Norway spruce (Picea abies [L.] Karst.).
Part 1: Productivity and quality of stems. Forestry Chronicle 82(4):
538-549.
A Photographic Guide 167

55. Eastern Pine Weevil, Pissodes nemorensis Germar (Coleoptera:

Curculionidae)

Orientation to Pest Eastern pine weevil, Pissodes nemorensis Germar, is now the name that
is applied to what was formerly known as northern pine weevil (Pissodes
approximatus Hopkins), which is no longer considered a valid species.
Eastern pine weevil is native to North America, where it feeds on cedars
(Cedrus) and pines (Pinus). In the southern United States, adults are
active in fall, winter and spring, and are inactive in summer. Eggs are
laid in the spring in stems or branches. The northern form appears to
commonly breed also in stumps of cut trees and becomes a pest in
plantations as numbers build up. Larvae feed in the cambium and pupate
in the wood in cells make of wood chips. Adults feed on branches and
terminals, which may be killed, leading to bushiness or crooked stems.
Small trees may be killed by larval feeding.

Hosts Commonly Attacked In the northern part of its range, eastern pine weevil breeds in pines,
especially red (Pinus resinosa Sol. ex Aiton) and Scots (P. sylvestris L.)
pines. In the southern United States, both cedars and pines are attacked.
Cedars used as hosts include both native species and exotic cedars
such as deodar cedar (Cedrus deodara [Roxb.] G. Don), Atlas cedar
(C. atlantica [Endl.] Manetti ex Carrière), and cedar of Lebanon
(C. libani A. Rich.). In the southern USA, various North American pines
are used as hosts, especially loblolly (P. taeda L.), shortleaf (P. echinata
Mill.), and longleaf (P. palustris Mill.)

Distribution The southern form of this weevil is found in the southeastern

United States, north to Pennsylvania. The northern form (the old
P. approximatus) is found from the Atlantic coast to Manitoba and
Minnesota and south to North Carolina. The species is not present in the
western United States. It is an invasive pest in some countries, including
South Africa, where it is a pest in plantations of exotic pines.

Images of Eastern
Pine Weevil

Figure 1. Adult of eastern pine weevil, Pissodes nemorensis. (Gerald J. Lenhard,

Louiana State University, Bugwood.org)
168 Forest Pest Insects in North America:

Images of Eastern
Pine Weevil (continued)

Figure 2. Eggs of eastern pine weevil. (Mark Fontaine, University of Florida,

Bugwood.org)

Figure 3. Larvae of eastern pine weevil feeding in gallery (left) and mature larva
(right) ready for pupation. (Left: Rayanne Lehman, Pennsylvania Department
of Agriculture, Bugwood.org; right: Gerald J. Lenhard, Louiana State University,
Bugwood.org)

Figure 4. Pupa of eastern pine weevil in cocoon (left), and cocoons on trunk
with bark removed (right). (Left: Gerald J. Lenhard, Louiana State University,
Bugwood.org; right: Chris Evans, River to River CWMA, Bugwood.org)
A Photographic Guide 169

Important Biological Control The braconid parasitoid Coeloides pissodis (Ashmead) is an important
Agents Related to this natural enemy of eastern pine weevil.
Pest Species
Web Links for Information http://wiki.bugwood.org/Archive:South/Pissodes_nemorensis; Bugwood
on Eastern Pine Weevil Wiki fact sheet on general biology of insect.
http://www.eppo.org/QUARANTINE/insects/Pissodes_nemorensis/
PISONE_ds.pdf; quarantine data sheet on species for Europe; detailed
information on biology and other topics.
http://www.fabinet.up.ac.za/tpcpweb/pamphlets/pissodes.pdf; fact sheet
on weevil as pest in South Africa providing considerable detail on
biology and management.

Articles Gebeyehu, S. and M.J. Wingfield. 2003. Pine weevil Pissodes

nemorensis: threat to South African pine plantations and options for
control. South African Journal of Science 99(11/12): 531-536.
170 Forest Pest Insects in North America:

56. Whitefringed Beetles (Naupactus spp.) (Coleoptera: Curculionidae)

Orientation to Pest Whitefringed beetles, formerly regarded as a separate genus

(Graphognathus), are now an informal “species group” within a much
larger genus (Naupactus). The name whitefringed beetle comes from
the white lines that run the length of the grey-to-brown body, two on
each side, one above and one below the eye. There are four species,
all invasive, in the United States: Naupactus leucoloma (Boheman),
N. peregrinus (Buchanan), N. minor (Buchanan), and N. fecundus
(Buchanan). These beetles are pests of various agricultural crops, but
also injure young pine seedlings when trees are planted into converted
farmland in the southern United States. Adults feed on the foliage of
hundreds of plant species, but damage is of minor importance. Larvae
feed on roots and their damage is more serious, especially on crops or
young pines in plantations and nurseries. Like the adults, the larvae feed
on a diverse set of hosts, although their full host range is more difficult to
document. The U.S. populations of all four species contain only females,
although populations with males are found in the native range in southern
South America. Adults have fused wing covers and cannot fly. Eggs are
laid in masses on objects on or near the soil. Larvae drop into the soil
where they feed on roots and then and pupate in earthen cells where they
overwinter. There is only one generation per year.

Hosts Commonly Attacked Adults feed on nearly 400 species of plants, including many crops.
Larvae feed on roots of some trees, such as peach (Prunus persica [L.]
Batsch), pecan (Carya illinoinensis [Wangenh.] K.Koch), tung (Vernicia
fordii [Hemsl.] Airy Shaw), willow (Salix), and pine (Pinus).

Distribution Originally from southern South America (Argentina, Peru, Chile,

Uruguay), whitefringed beetles are found as invasive species in the
United States from Florida north to Virginia and Illinois and west to
Texas.

Images of
Whitefringed Beetles

Figure 1. Adult of one of the whitefringed beetles, Naupactus peregrinus; see

the enhanced white lines in drawing on right. (Left: Russ Ottens, University
of Georgia, Bugwood.org; right: Department of Entomology and Nematology,
University of Florida)
A Photographic Guide 171

Images of Whitefringed
Beetles (continued)

Figure 2. Larva of a species of whitefringed beetle in root (left) and close up

(right). (Left: Terry S. Price, Georgia Forestry Commission, Bugwood.org; right:
Edward L. Barnard, Florida Department of Agriculture and Consumer Services,
Bugwood.org)

Figure 3. Feeding damage of adult whitefringed beetles (left), and close up of

adult feeding on blueberry leaf (right). (Left: Louis Tedders, USDA Agricultural
Research Service, Bugwood.org; right: Jerry A. Payne, USDA Agricultural
Research Service, Bugwood.org)

Figure 4. Feeding damage of larvae of whitefringed beetles on root of sycamore

tree. (James Solomon, USDA Forest Service, Bugwood.org)
172 Forest Pest Insects in North America:

Images of Whitefringed
Beetles (continued)

Figure 5. “Scalping” is the use of machinery to remove a thin layer of thatch

and top soil before planting a new block of pine seedlings. (Edward L. Barnard,
Florida Department of Agriculture and Consumer Services, Bugwood.org)

Figure 6. Unscalped pine plantation (left), showing loss of seedling establishment

vs. scalped plot (right), where whitefringed beetles have been removed and
pine seedling survival increased. (Both photos: Edward L. Barnard, Florida
Department of Agriculture and Consumer Services, Bugwood.org)

Figure 7. Scalped plot (left of man) vs unscalped plot (right of man) in slash
pine (Pinus elliottii Engelm.) plantation, showing cumulative effect of larvae of
whitefringed beetles on growth of slash pines when beetle are suppressed before
planting via “scalping” (removal of thatch and bit of top soil just before planting)
or not. (Edward L. Barnard, Florida Department of Agriculture and Consumer
Services, Bugwood.org)
A Photographic Guide 173

Important Biological Control No information was located on specialized natural enemies of this group
Agents Related to this of beetles.
Pest Species
Web Links for Information http://entnemdept.ufl.edu/creatures/field/beetles/whitefringed_beetles.
on Whitefringed Beetles htm; University of Florida fact sheet on biology and control.
http://www.eppo.org/QUARANTINE/insects/Naupactus_leucoloma/
GRAGLE_ds.pdf; EPPO datasheet on this quarantine pest (from
Europe’s point of view) containing information on biology and damage.
http://keys.lucidcentral.org/keys/sweetpotato/key/
Sweetpotato%20Diagnotes/media/html/TheProblems/Pest-
Root&StemInsects/WhitefringedBeetles/whitefringed%20beetle%2
0home.htm; factsheet of University of Queensland (Australia) with
information on recognition, biology, ecology, and control.

Articles Anderson W.H. 1938. A key to separate the larvae of the white-fringed
beetle, Naupactus leucoloma Boh., from the larvae of closely related
species. USDA Bureau of Entomology and Plant Quarantine Circular
E-422. 3 p.
Ahmad, R. 1974. Studies on Graphognathus leucoloma (Boh.) Col.:
Curculionidae) and its natural enemies in the central provinces of
Argentina. Technical Bulletin No. 17, Commonwealth Institute of
Biological Control: 19-28.
Harlan, D.P. and J.W. McGuire. 1977. Separation of four species of
whitefringed beetles by measuring the length and width of the eggs
(Coleoptera: Curculionidae). Journal of the Georgia Entomological
Society 12: 125-128.
Lanteri, A.A. and A.E. Marvaldi. 1995. Graphognathus Buchanan a
new synonym of Naupactus Dejean and systematics of the N. leucoloma
species group (Coleoptera: Curculionidae). Coleopterists Bulletin 49(3):
206-228.
174 Forest Pest Insects in North America:

57. Yellow-poplar Weevil, Odontopus calceatus (Say) (Coleoptera:

Curculionidae)

Orientation to Pest Yellow-poplar weevil, Odontopus calceatus (Say), feeds on yellow-

poplar (Liriodendron tulipifera L.), magnolia (Magnolia), and sassafrass
(Sassafras albidum [Nuttall] Nees.). The weevil overwinters as an adult
and upon emergence, adults feed on buds or developing leaves. Eggs
are laid in leaf veins and larvae feed as leafminers. Multiple larvae may
coexist in a single common mined area. Outbreaks in yellow-poplar
have occurred since 1960 in eastern Kentucky, Tennessee, Ohio, West
Virginia, and Virginia.

Hosts Commonly Attacked This weevil feeds on yellow-poplar (L. tulipifera), magnolia (Magnolia),
and sassafras (S. albidum).

Distribution This weevil is found throughout the eastern United States within the
range of its hosts.

Images of
Yellow-poplar Weevil

Figure 1. Adults of yellow-poplar weevil, Odontopus calceatus, on leaf and (inset)

close up. (Lacy L. Hyche, Auburn University, Bugwood.org; inset: Rich Kelly)
A Photographic Guide 175

Images of Yellow-poplar
Weevil (continued)

Figure 2. Oviposition wounds (left) and eggs of yellow-poplar weevil (revealed by

dissection) (right) in leaf vein. (Both photos: Lacy L. Hyche, Auburn University,
Bugwood.org)

Figure 3. Larval mine (left) and larvae (right) (seen by opening mine) of yellow-
poplar weevil. (Both photos: Lacy L. Hyche, Auburn University, Bugwood.org)

Figure 4. Cocoons in leaf (left) and pupa (right) (revealed by opening cocoon)
of yellow-poplar weevil. (Both photos: Lacy L. Hyche, Auburn University,
Bugwood.org)

Important Biological Control Unspecified species of parasitoids have been noted to destroy up to
Agents Related to this 50 percent of larvae in mines.
Pest Species

Web Links for Information http://www.ca.uky.edu/entomology/entfacts/ef414.asp; University of

on Yellow-poplar Weevil Kentucky fact sheet.
http://www.ag.auburn.edu/enpl/bulletins/lmweevil/lmweevil.htm; fact
sheet of Auburn University, with extensive photos.
http://www.fs.fed.us/outernet/r6/nr/fid/fidls/fidl-125.pdf; USDA Forest
Service Pest Leaflet.
176 Forest Pest Insects in North America:

Articles on Burns, D.P. and L.P. Gibson. 1968. The leaf mining of yellow-poplar. The
Yellow-poplar Weevil Canadian Entomologist 100: 421-429.
Hyche, L.L. 1994. The yellow-poplar leaf-mining weevil. A guide to
recognition and habits in Alabama. Bulletin No. 622 of the Alabama
Agricultural Experiment Station, Alabama Agricultural Experiment
Station, Auburn University: 13 p.
A Photographic Guide 177

58. Native Elm Bark Beetle, Hylurgopinus rufipes (Eichhoff) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Native elm bark beetle, Hylurgopinus rufipes (Eichhoff), is a native
North American insect associated with various species of elm and other
trees. Before the introduction of the exotic pathogen causing Dutch
elm disease, this insect was of minor importance. During the historical
period of mass epidemics of Dutch elm disease, this native bark beetle
served as a vector of the pathogen. This process continues in wild lands
where native elms survive and reproduce before succumbing to Dutch
elm disease. Although H. rufipes has been displaced over much of its
range by the introduced smaller European elm bark beetle (Scolytus
multistriatus [Marsham]), H. rufipes remains the dominant vector of
Dutch elm disease in northern areas where S. multistriatus is limited by
low temperatures. Adult beetles emerge in early spring and fly to healthy
elms to feed for a short period in the bark of branches and twigs. After
feeding, beetles fly to dying and recently dead trees for oviposition.
Females construct egg galleries that are oriented horizontally, across the
grain of the wood. Eggs are laid along the egg gallery and larvae feed in
tunnels that run parallel to the grain and perpendicular to the egg gallery.
Larvae pupate in small cells at the ends of the feeding gallery. In some
instances, larvae and newly developed adults will overwinter in these
galleries, with adults emerging the following spring through small round
exit holes. At other times, adults emerge before winter; these adults fly to
healthy elm trees to feed on branches and to chew overwintering galleries
in the bark, generally of the lower stem. If as larvae these adults had
developed in trees dying from Dutch elm disease, they are very likely to
carry spores of the fungus that causes Dutch elm disease. When they visit
healthy elm trees to feed on branches or to create overwintering galleries,
these beetles can transmit the pathogen. There are one to two generations
of this bark beetle per year, depending on location.

Hosts Commonly Attacked Rock (Ulmus thomasii Sarg.) and American elm (U. americana L.) are
the major hosts of this bark beetle, but it also attacks American basswood
(Tilia americana L.), and species of ash (Fraxinus).

Distribution This native bark beetle was formerly common throughout eastern North
America from northern Alabama and Mississippi to southern Canada,
but it has been displaced over most of this area by the introduced smaller
European elm bark beetle. It is, however, still common in colder areas
such as northern New York, New England, northern Minnesota, and
southern Canada, where the smaller European elm bark beetle does not
survive winter temperatures.
178 Forest Pest Insects in North America:

Images of Native
Elm Bark Beetle

Figure 1. Adult native elm bark beetle, Hylurgopinus rufipes. (J.R. Baker and
S.B. Bambara, North Carolina State University, Bugwood.org)

Figure 2. Egg gallery

(horizontal) and larval
galleries (vertical)
of the native elm
bark beetle. (John
A. Williams, USDA
Forest Service,
Bugwood.org)
A Photographic Guide 179

Images of Native Elm

Bark Beetle (continued)

Figure 3. Historical record of the change the new pathogen made in the
importance of this bark beetle. (Darren Blackford, USDA Forest Service,
Bugwood.org)

Figure 4. Two views of American elms dying of Dutch elm disease. (Left:
Linda Haugen, USDA Forest Service, Bugwood.org; right: Roland J. Stipes,
Virginia Polytechnic Institute and State University, Bugwood.org)
180 Forest Pest Insects in North America:

Images of Native Elm

Bark Beetle (continued)

Figure 5. Streaking and discoloration of the vascular tissue in elms infected with
Dutch elm disease. (North Carolina Forest Service Archive, Bugwood.org)

Figure 6. The clerid Enoclerus nigripes, an important predator of the native elm
bark beetle. (Charley Eiseman, Bugwood.org)

Important Biological Control The major predator of this bark beetle in Connecticut is the clerid
Agents Related to this Enoclerus nirgripes Say and the principal parasitoid is Spathius
Pest Species canadensis Ashmead.

Web Links for Information http://www.extension.umn.edu/distribution/horticulture/DG1420.html;

on Native Elm Bark Beetle fact sheet from the University of Minnesota.
http://www.na.fs.fed.us/spfo/pubs/howtos/ht_ded/ht_ded.htm#intro;
recognition and management of Dutch elm disease.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=2850. Natural
Resources Canada fact sheet.
A Photographic Guide 181

Articles Thompson, H.E. and J.G. Matthysse. 1972. Role of the native elm bark
beetle, Hylurgopinus rufipes (Eichh.), in transmission of the Dutch
elm disease pathogen, Ceratocystis ulmi (Buisman) C. Moreau. Search
Agriculture (Entomology) 2(1,V): 16 p.
Gardiner, L.M. 1981. Seasonal activity of the native elm bark beetle,
Hylurgopinus rufipes, in central Ontario (Coleoptera: Scolytidae).
The Canadian Entomologist 113: 341-348.
182 Forest Pest Insects in North America:

59. Smaller European Elm Bark Beetle, Scolytus multistriatus (Marsham)

(Coleoptera: Curculionidae, Scolytinae)

Orientation to Pest Smaller European elm bark beetle, Scolytus multistriatus (Marsham), is
an introduced bark beetle that is one of the two main vectors of Dutch
elm disease (along with the native elm bark beetle, Hylurgopinus rufipes
[Eichhoff]) in the eastern United States and Canada. It was brought to
North America through the importation of unbarked elm logs containing
live brood of the pest. It breeds in various species of elm, but the greatest
damage is done to American elm (Ulmus americana L.). It is widely
distributed in the United States. The biology of this species is much
the same as for the native elm bark beetle. Adults fly to healthy elm
trees in the spring, where they feed on twigs, infecting them with the
fungal pathogen (Ophiostoma ulmi [Buisman] Melin and Nannf.) that
causes Dutch elm disease. Trees that are stressed by this disease or other
factors are attacked by females for breeding. Eggs are laid in galleries
that females dig under the bark of the tree. Larvae develop in galleries,
feeding in the cambium. Mature larvae overwinter in their galleries,
pupating in spring and producing a new flight of adults. Emergence
holes have a characteristic “shot hole” appearance. Mass attack on
stressed trees is mediated, as in many bark beetles, via an aggregation
pheromone. This beetle has displaced the native elm bark beetle in many
parts of North America, except in the coldest areas.

Hosts Commonly Attacked Smaller European bark beetle attacks all species of Elm (Ulmus) and the
Japanese species Zelkova serrata (Thunb.) Makino.

Distribution This invasive bark beetle is found in southern Canada and most of the
United States except some parts of Florida and Maine.

Images of Smaller
European Bark Beetle

Figure 1. Adult of smaller European elm bark beetle, Scolytus multistriatus.

(Left: J.R. Baker and S.B. Bambara, North Carolina State University,
Bugwood.org; right: Maja Jurc, University of Ljubljana, Bugwood.org)
A Photographic Guide 183

Images of Smaller European

Bark Beetle (continued)

Figure 2. Elm branch with feeding in crotch from adults of smaller European elm
bark beetle. (Joseph O’Brien, USDA Forest Service, Bugwood.org)

Figure 3. Galleries and brood of smaller European elm bark beetle; central
galleries between “wings” are oviposition galleries and the two “wings” are groups
of larval galleries; larvae are visible at ends of many galleries in photo on left.
(Left: Joseph O’Brien, USDA Forest Service, Bugwood.org; right: Beat Forster,
Swiss Federal Institute for Forest, Snow and Landscape Research,
Bugwood.org)

Figure 4. Round
emergence holes of the
smaller European elm
bark beetle. (J.R. Baker
and S.B. Bambara,
North Carolina State
University, Bugwood.org)
184 Forest Pest Insects in North America:

Images of Smaller European

Bark Beetle (continued)

Figure 5. Two views of American elms dying of Dutch elm disease. (Left:
Linda Haugen, USDA Forest Service, Bugwood.org; right: Roland J. Stipes,
Virginia Polytechnic Institute and State University, Bugwood.org)

Figure 6. Two views of the streaking and discoloration of the vascular tissue
in elms infected with Dutch elm disease. (Left: Joseph LaForest, University
of Georgia, Bugwood.org; right: North Carolina Forest Service Archive,
Bugwood.org)
A Photographic Guide 185

Important Biological Control In Europe, several parasitoids are known to attack S. multistriatus.
Agents Related to this Three of these—Dendrosoter protuberans (Nees), Ecphylus silesicacus
Pest Species (Ratzburg), and Coeloides scolyticida Wesm.—have been introduced
into the United States. However, densities of this invasive bark beetle
have not been compared between Europe and North America, to see
if they were higher in North America than Europe at the time of these
introductions. Nor have studies determined if these introductions (some
of which did result in establishment) reduced the abundance of this bark
beetle. Until recently whether or not this biological control effort had any
effect on the target insect was a moot point, as sufficient elm bark beetles
remained to allow the Dutch elm disease epidemic to go unchecked,
destroying most native elms. The picture is further complicated by
the fact that the pathogen, believed to be of Asian origin, has recently
evolved to form a new, more aggressive form, Ophiostoma novo-ulmi
Brasier. This new pathogen has spread from the American midwestern
states into other regions of North America, replacing the form of the
pathogen introduced in North America in the 1940s. Nevertheless, the
dynamics of this system in the context of replanting disease-resistant
American elms in wildlands should now be revisited.

Web Links for Information http://www.ento.okstate.edu/ddd/insects/smeurelmbark.htm; fact sheet of

on Smaller European Oklahoma State University.
Bark Beetle http://crawford.tardigrade.net/bugs/BugofMonth38.html; Bug of the
Month fact sheet.
http://aces.nmsu.edu/ces/plantclinic/documents/o-03-barkbeetles.pdf;
New Mexico fact sheet on bark beetles in general.

Articles Schroder, D. 1974. Investigations on the prospects for biological

control of scolytids on elms as a means of reducing Dutch elm disease.
Zeitschrift für Angewandte Entomologie 76(2): 150-159. (In German).
186 Forest Pest Insects in North America:

60. Spruce Beetle, Dendroctonus rufipennis (Kirby) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Spruce beetle, Dendroctonus rufipennis (Kirby), is a native North

American bark beetle that usually develops in wind-thrown, over mature,
or weakened trees, resources that normally occur at low densities in
forests. The biology of the beetle is similar to that of many bark beetles,
with females laying eggs in tunnels (egg galleries), from which larvae
then begin to feed, forming larval galleries. In this species young
larvae often feed in common galleries, but then produce individual
galleries as they mature. Larvae pupate at the end of their galleries, and
overwinter as either larvae or new adults in the galleries. Spruce beetles
generally require two years to complete their life cycle. However, when
temperatures are warmer than normal a generation can be completed in
one year. When two years are required, development is not synchronized
and beetles emerge and attack trees each year. Spruce beetle outbreaks
are associated with warmer than usual weather, and are often triggered by
disturbance events such as avalanches, storms, and logging. Outbreaks
tend to occur in areas with an abundance of large spruce trees. Outbreaks
occur several times each century, thinning large trees from stands over
extensive areas. More severe outbreaks in which trees of most sizes
and vigor classes are killed occur less frequently. Important outbreaks
historically have occurred in Alaska and Utah.

Hosts Commonly Attacked Spruce beetle attacks various species of native North America spruce
(Picea).

Distribution This bark beetle is found throughout the spruce forests of North America.

Images of
Spruce Beetle

Figure 1. Adult of spruce beetle, Dendroctonus rufipennis. (David McComb,

USDA Forest Service, Bugwood.org)
A Photographic Guide 187

Images of Spruce
Beetle (continued)

Figure 2. Female spruce beetle digging an oviposition gallery, with white eggs
visible in upper left. (Edward H. Holsten, USDA Forest Service, Bugwood.org)

Figure 3. Spruce beetle larvae at ends of tunnels. (Edward H. Holsten, USDA

Forest Service, Bugwood.org)

Figure 4. Pupae of spruce beetle in galleries. (Edward H. Holsten, USDA Forest

Service, Bugwood.org)
188 Forest Pest Insects in North America:

Images of Spruce Figure 5. Oviposition

(vertical) and larval
Beetle (continued) (horizontal) galleries of
spruce beetle etched
in wood under bark of
spruce tree. (Darren
Blackford, USDA Forest
Service, Bugwood.org)

Figure 6. Pitch tubes (left) and frass (right) on spruce trunks are signs of spruce
beetle attack. (Left: Darren Blackford, USDA Forest Service, Bugwood.org; right:
A. Steven Munson, USDA Forest Service, Bugwood.org)
A Photographic Guide 189

Images of Spruce
Beetle (continued)

Figure 7. Adult spruce beetles in galleries. (A. Steven Munson, USDA Forest
Service, Bugwood.org)

Figure 8. Burning of stumps (left) or logging slash (right) after felling infested
trees is intended to limit spruce beetle outbreaks. (Left: A. Steven Munson, USDA
Forest Service, Bugwood.org; right: USDA Forest Service - Region 2 - Rocky
Mountain Region Archive, USDA Forest Service, Bugwood.org)

Figure 9. Views of spruce trees killed by spruce beetle, at various physical

scales (dead trees denoted in artificial color in photo on right). (Left: USDA
Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org; right:
William M. Ciesla, Forest Health Management International, Bugwood.org)
190 Forest Pest Insects in North America:

Important Biological Control A variety of natural enemies have been recorded in the literature
Agents Related to this attacking spruce beetle brood (see Van Driesche et al., 1996 in USDA
Pest Species FS publication FHTET-96-19, URL given below), including several
species of braconid parasitoids in the genus Coeloides. Also the role of
woodpecker predation on brood has been studied. But the fundamental
determinants of outbreaks of spruce beetle do not seem to be driven by
natural enemies, but rather to be responses to weather, the amount of
breeding material for the beetle at a site, and local and regional spruce
stand conditions.

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=2819; fact

on Spruce Beetle sheet of Natural Resources Canada.
http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf; USDA Forest Service
publication FHTET-96-19, Van Driesche et al., 1996; see pages 55-57
for information on natural enemies of this species.
http://na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm; USDA
Forest Service Forest Insect and Disease Leaflet on spruce beetle.

Articles Schmid, J.M. and R.H. Frye. 1976. Stand ratings for spruce beetles. Res.
Note RM-309. USDA Forest Service, Rocky Mountain Forest and Range
Experiment Station, Ft. Collins, Colorado: 4 p.
Schmid, J.M. and R.H. Frye. 1977. Spruce beetle in the Rockies. USDA
Forest Service, Rocky Mountain Forest and Range Experiment Station,
General Technical Report RM-49. Ft. Collins, Colorado: 38 p.
Linton, D. A. and L. Safranyik, L. 1988. The spruce beetle Dendroctonus
rufipennis (Kirby): an annotated bibliography 1885-1987. Information
Report - Pacific Forestry Centre, Canadian Forestry Service (BC-X-298):
39 p.
A Photographic Guide 191

61. Black Turpentine Beetle, Dendroctonus terebrans (Olivier) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Black turpentine beetle, Dendroctonus terebrans (Olivier), is a native,

North American bark beetle that attacks various pines. Adults of this
species are strongly attracted to odors of freshly cut pine stumps,
where they are able to breed. In whole trees, eggs are primarily laid in
the bottom 1-2 meters of the boles as well as roots of weakened trees.
Eggs are laid in large oviposition galleries created by females with
the help of males. Larvae feed collectively in large galleries. Pupation
takes place in pupal cells constructed in the corky bark or between
the bark and the wood. In the northern part of the range, adults are the
overwintering stage. In southern areas, all stages may be present all year.
Damage can occur in healthy trees as well as stressed trees, and damage
in turpentine orchards can be severe. The black turpentine beetle is
often found attacking trees in association with the southern pine beetle
(Dendroctonus frontalis Zimmermann) or various species of southern
pine engraver beetles (Ips spp.).

Hosts Commonly Attacked Hosts of black turpentine beetle include all species of southern pines, but
especially loblolly (Pinus taeda L.) and slash (P. elliottii Engelm.) pines.
In the northern part of its range, black turpentine beetle attacks various
conifers, including black (P. thunbergiana Franco) and pitch (P. rigida
Miller) pines and red spruce (Picea rubens Sargent).

Distribution This bark beetle is found in the eastern United States from Florida
west to Missouri and Texas and north to coastal areas of southern New
England.

Images of Black
Turpentine Beetle

Figure 1. Adults of black turpentine beetle, Dendroctonus terebrans. (Left:

David T. Almquist, University of Florida, Bugwood.org; right: Clemson University
- USDA Cooperative Extension Slide Series, Bugwood.org)
192 Forest Pest Insects in North America:

Images of Black Figure 2. Black

turpentine beetle
Turpentine Beetle (bottom) is the largest
(continued) of the five species
of southern pine
bark beetles. The
other species (from
top down) are: Ips
avulsus (Eichhoff), Ips
grandicollis (Eichhoff),
Ips calligraphus
(Germar), and
Dendroctonus frontalis
(Zimmermann).
(Gerald J. Lenhard,
Louiana State
University,
Bugwood.org)

Figure 3. Eggs of black turpentine beetle (white, above beetle on left).

(Terry S. Price, Georgia Forestry Commission, Bugwood.org)
A Photographic Guide 193

Images of Black
Turpentine Beetle
(continued)

Figure 4. Larvae of black turpentine beetle in common feeding gallery (left);

larvae enlarged (right). (Left: James R. Meeker, USDA Forest Service,
Bugwood.org; right: Gerald J. Lenhard, Louiana State University, Bugwood.org)

Figure 5. Pupae of black turpentine beetle, in pupal cells, lower left.

(Ronald F. Billings, Texas Forest Service, Bugwood.org)

Figure 6. Pitch tubes (left) are created by the tree where black turpentine beetles
bore in the tree; during mass attack, there may be many pitch tubes in the lower
bole area (right). (Left: USDA Forest Service Archive, USDA Forest Service,
Bugwood.org; right: Ronald F. Billings, Texas Forest Service, Bugwood.org)
194 Forest Pest Insects in North America:

Images of Black
Turpentine Beetle
(continued)

Figure 7. Dying loblolly pines infested with both Ips beetles and black turpentine
beetles. (Ronald F. Billings, Texas Forest Service, Bugwood.org)

Figure 8. The clerid Thanasimus dubius (red larva in lower center, top; and
close up, bottom) is a common predator of brood of black turpentine beetles.
(Top: North Carolina State University Archive, North Carolina State University,
Bugwood.org; bottom: Gerald J. Lenhard, Louiana State University,
Bugwood.org)
A Photographic Guide 195

Important Biological Control Little is known about the specific natural enemies of this species. The
Agents Related to this clerid Thanasimus dubius (Fabricius) and the trogositid Temnochila
Pest Species virescens (Fabricius), predatory beetles that prey on many southern pine
bark beetles, are found attacking brood of black turpentine beetles.

Web Links for Information http://entnemdept.ufl.edu/creatures/trees/beetles/black_turpentine_beetle.

on Black Turpentine Beetle htm; fact sheet of the University of Florida.

Articles Barras, S.J. and T. Perry. 1971. Leptographium terebrantis sp. nov.
associated with Dendroctonus terebrans in loblolly pine. Mycopathologia
43: 1-10.
Foltz, J.L., E.P. Merkel, and R.C. Wilkinson. 1984. Annotated
bibliography of Dendroctonus terebrans (Oliver), Ips avulsus (Eichhoff),
Ips grandicollis (Germar) in the southeastern USA. Monograph No. 12.
Agricultural Experiment Station, University of Florida: 47 p.
196 Forest Pest Insects in North America:

62. Eastern Larch Beetle, Dendroctonus simplex (LeConte) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Eastern larch beetle, Dendroctonus simplex (LeConte), is a native North
American bark beetle whose only important host is eastern larch, Larix
larcina (Du Roi ) K. Koch. The species overwinters as larvae or adults
in galleries. Eggs are laid in groups along an egg gallery made by the
female in the inner bark and outer sapwood. Larval tunnels are short
and occur in the inner bark. There are up to three generations per year.
Before the 1970s, this bark beetle was only known to attack dying or
recently felled trees. Since then, however, a series of large outbreaks in
healthy stands have occurred in both Canada and the United States. Some
of these bark beetle outbreaks followed outbreaks of the defoliating
larch sawfly Pristiphora erichsonii (Hartig), which had produced many
stressed or dying larch trees. While the exact causes of these population
fluctuations are unknown, some authors have attributed outbreaks to the
maturing of larch stands over large areas, which lowers host resistance.

Hosts Commonly Attacked Hosts of this species are eastern larch (L. larcina), and occasionally red
spruce (Picea rubens Sargent).

Distribution This bark beetle is found in the United States from New England south
to West Virginia, and west to Minnesota. It is also found in Alaska and
in Canada from the Atlantic to the Pacific coast.

Images of Eastern
Larch Beetle

Figure 1. Adults of eastern larch beetle, Dendroctonus simplex, in galleries.

(Thérèse Arcand - Natural Resources Canada, Canadian Forest Service)
A Photographic Guide 197

Images of Eastern
Larch Beetle (continued)

Figure 2. Eggs of eastern larch beetle. (Thérèse Arcand - Natural Resources

Canada, Canadian Forest Service)

Figure 3. Larva of eastern larch beetle. (Thérèse Arcand - Natural Resources

Canada, Canadian Forest Service)

Figure 4. Pupa of eastern larch beetle. (Thérèse Arcand - Natural Resources

Canada, Canadian Forest Service)
198 Forest Pest Insects in North America:

Images of Eastern
Larch Beetle (continued)

Figure 5. Larval galleries at surface of wood made by eastern larch beetle.

(Thérèse Arcand - Natural Resources Canada, Canadian Forest Service)

Important Biological Control Eastern larch beetle has a large complex of natural enemies that have
Agents Related to this been studied in a number of locations (see Van Driesche et al. 1996 for a
Pest Species review). Some of the natural enemies reported include the dolichopodid
fly, Medetera gaspensis Bickel, and the rhizophagid beetle, Rhizophagus
dimidiatus Mannerheim, (both predators), and the parasitoids Spathius
canadensis Ashmead (Braconidae), Rhopalicus tutela (Walker), and
Roptrocerus xylophagorum (Ratzeburg).

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/elb/elb.htm; a USDA Forest

on Eastern Larch Beetle Service fact sheet, Forest Insect and Disease Leaflet #175, providing
extensive detail on biology, hosts, distribution, and control of eastern
larch beetle.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=2820; a
Natural Resources Canada fact sheet.

Articles Duncan, B. 1987. An illustrated guide to the identification and

distribution of the species of Dendroctonus Erichson (Coleoptera:
Scolytidae) in British Columbia. Journal of the Entomological Society
of British Columbia 84: 101-112.
Langor, D.W. and A.G. Raske. 1988. Annotated bibliography of the
eastern larch beetle, Dendroctonus simplex LeConte (Coleoptera:
Scolytidae). Information Report - Newfoundland Forestry Centre,
Forestry Canada (N-X-266), 1988: 38 p.
Langor, D.W. and A.G. Raske. 1988. Mortality factors and life tables of
the eastern larch beetle, Dendroctonus simplex (Coleoptera: Scolytidae),
in Newfoundland. Environmental Entomology 17: 959-963.
A Photographic Guide 199

Articles (continued) Langor, D.W. and A.G.A. Raske. 1989. A history of the eastern larch
beetle, Dendroctonus simplex (Coleoptera: Scolytidae), in North
America. Great Lakes Entomologist 22: 139-154.
Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
control of arthropod pests of the northeastern and north central
forests in the United States: a review and recommendations. USDA
FS, FHTET-96-19, Morgantown, West Virginia: 58-59. (Available
at http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
200 Forest Pest Insects in North America:

63. Southern Pine Beetle, Dendroctonus frontalis (Zimmermann)

(Coleoptera: Curculionidae, Scolytinae)

Orientation to Pest Southern pine beetle, Dendroctonus frontalis (Zimmermann), is the most
destructive bark beetle attacking pines in the southern United States. It is
a native North American species that also occurs in Mexico and Central
America. Its principal hosts are yellow pines, especially shortleaf (Pinus
echinata Mill.), loblolly (P. taeda L.), Virginia (P. virginiana Mill.), and
pitch (P. rigida Mill.) pines. Beetles overwinter in all stages and timing
of adult emergence in spring varies with location, from mid-March to
June. Bark areas from which adults have emerged have a shot-hole-like
appearance. Attack by adults during warm days in winter is usually on
previously attacked trees and is focused initially on the mid to upper part
of the tree. Females construct a nuptial chamber, where they are later
joined by males. An S-shaped oviposition gallery is then constructed,
along which eggs are laid. Larval galleries are at right angles to the
oviposition gallery and score the wood. Larval tunnels are primarily in
the bark and pupation occurs at the end of the larval gallery. With each
new generation, infested areas expand, sometimes covering very large
areas. In western North Carolina there are three to five generations per
year and up to seven per year further south. The generations overlap,
so in active infestations, all life stages of the beetle are present at any
one time and adult beetles emerge and attack new trees daily when
temperatures allow. Trees under 15 years of age or 5 cm in diameter are
rarely attacked. Infestations typically begin in stands of trees stressed
by high tree density, logging damage, lightning, dryness, flooding, or
disease. Trees die either because they are girdled by larval galleries,
or because of the effects of a blue stain fungus (Ceratocystis minor
[Hedgc.] Hunt), which is vectored by the beetle. Damage from outbreaks
is reduced somewhat by managing stands to reduce tree stress and by
halting the expansion of initial spots using salvage (cut-and-remove) or
cut-and-leave control methods.

Hosts Commonly Attacked Principal species in which reproduction of southern pine beetle occurs
are the yellow pines, especially shortleaf (P. echinata), loblolly (P.
taeda), Virginia (P. virginiana), and pitch (P. rigida) pines. The beetle
also attacks species such as longleaf (P. palustris Miller) and slash
(P. elliottii Engelm.) pines, but reproduction is often less successful
due to heavy resin flow.

Distribution This bark beetle is found throughout the southeastern and southern parts
of the United States, as well as Mexico and parts of Central America.
A Photographic Guide 201

Figure 1. Distribution of southern

pine beetle, Dendroctonus frontalis.
(Eastern Forest Environmental
Threat Assessment Center • Threat
Summary)

Images of
Southern Pine Beetle

Figure 2. Adults of southern pine beetle; left, beetle in flight. (Both photos:
Erich G. Vallery, USDA Forest Service - SRS-4552, Bugwood.org)

Figure 3. The head of the southern pine beetle is diagnostic for the species. The
head is broad, with a distinct notch or frontal groove on male beetles, bordered
by distinct projections or tubercules on either side of the groove. The antennae
are seven-segmented, consisting of the basal pedicel, elongated scape, four-
segmented funicle, and an enlarged club. (Pest and Diseases Image Library,
Bugwood.org)
202 Forest Pest Insects in North America:

Images of Southern Pine

Beetle (continued)

Figure 4. Egg of southern pine beetle. (USDA Forest Service Archive, USDA
Forest Service, Bugwood.org)

Figure 5. Life cycle diagram of southern pine beetle (left), showing various
larval stages; view of southern pine beetle larva in wood (right). (Left: Ronald F.
Billings, Texas Forest Service, Bugwood.org; right: Erich G. Vallery, USDA Forest
Service - SRS-4552, Bugwood.org)

Figure 6. Pupae of southern pine beetle. (Ronald F. Billings, Texas Forest

Service, Bugwood.org)
A Photographic Guide 203

Images of Southern Pine

Beetle (continued)

Figure 7. Young adults of southern pine beetle in galleries, ready to emerge.

(Erich G. Vallery, USDA Forest Service - SRS-4552, Bugwood.org)

Figure 8. Pitch tubes on pines (left) are a sign of bark beetles such as southern
pine beetle; close-up of pitch tube (center), and female southern pine beetle
with entrance hole in pitch tube, right. (All photos: Erich G. Vallery, USDA Forest
Service - SRS-4552, Bugwood.org)

Figure 9. Galleries of southern pine beetle and associated blue stain (left), and
view of blue stain fungus (right) in cross section slice of trunk. (Left: Erich G.
Vallery, USDA Forest Service - SRS-4552, Bugwood.org; right: Ronald F. Billings,
Texas Forest Service, Bugwood.org)
204 Forest Pest Insects in North America:

Images of Southern Pine

Beetle (continued)

Figure 10. Initial point of outbreak (“spot”) of southern pine beetle (left) and
larger area of damage, to loblolly pine (right). (Left: Erich G. Vallery, USDA
Forest Service - SRS-4552, Bugwood.org; right: William M. Ciesla, Forest Health
Management International, Bugwood.org)

Figure 11. Tree felling is a control method (left) used to prevent expansion of spot
infestations (right, spot after infested trees have been removed). Infested trees
and uninfested trees in a buffer zone are either cut and removed or cut and left
in place, in either case for the purpose of stopping spot expansion. (Both photos:
Ronald F. Billings, Texas Forest Service, Bugwood.org)

Figure 12. Larva of checkered beetle, Thanasimus dubius (Fabricius) (Cleridae)

eating a southern pine beetle larva (top left) and close up of T. dubius larva (top
right); adult of T. dubius with prey (bottom). (Top left: John Moser, USDA Forest
Service, Bugwood.org; top right: Gerald J. Lenhard, Louiana State University,
Bugwood.org; bottom: Erich G. Vallery, USDA Forest Service - SRS-4552,
Bugwood.org)
A Photographic Guide 205

Important Biological Control The most important natural enemies of southern pine beetles are
Agents Related to this predacious clerid beetles. These predators are attracted to the aggregation
Pest Species pheromone and host volatiles used in traps to assess southern pine
beetle densities in particular stands. Comparison of pest and predator
numbers caught in traps is used each spring throughout the southern and
eastern United States to forecast southern pine beetle population trends
(outbreaks or population declines) (Figure 13).

Figure 13. A decision chart to forecast southern pine beetle outbreaks, using
information on density of the bark beetle and its clerid predator in traps baited
with aggregation pheromone (developed by Ron Billings, Texas Forest Service)
(see Billings and Upton, 2010).

Web Links for Information http://entnemdept.ufl.edu/creatures/trees/southern_pine_beetle.htm;

on Southern Pine Beetle University of Florida fact sheet.
Fidgen, J. (2000). Southern pine beetle information directory.
http://everest.ento.vt.edu/~salom/SPBinfodirect/spbinfodirect2.html
(5 July 2001).
http://www.barkbeetles.org/spb/spbbook/Index.html; a web rendering
of the U.S. Department of Agriculture, Expanded Southern Pine Beetle
Research and Applications Program, Forest Service, Science and
Education Administration, Technical Bulletin 1631: [1980].
ftp://ftp.fao.org/docrep/fao/011/i0640e/i0640e10e.pdf; Bulletin of the
United Nations, Food and Agriculture Organization.
http://www.barkbeetles.org/centralamerica/0605e.html; management
of southern pine beetle in Central America.
206 Forest Pest Insects in North America:

Articles on Chellman, C.W. and R.C. Wilkinson. 1975. Recent history of southern
Southern Pine Beetle pine beetle, Dendroctonus frontalis Zimm. (Col.; Scolytidae), in Florida.
Florida Entomologist 58: 22.
Dixon, W.N. and T.L. Payne. 1979. Aggregation of Thanasimus dubius
on trees under mass-attack by the southern pine beetle. Environmental
Entomology 8: 178-181.
Billings, R.F. and H.A. Pase III. 1979. A field guide for ground checking
southern pine beetle spots. U.S. Department of Agriculture, Combined
Forest Pest Research and Development Program, Agriculture Handbook
No. 558: 19 p.
Swain, K.M. and M.C. Remion. 1981. Direct control methods for the
southern pine beetle. U.S. Department of Agriculture, Combined Forest
Pest Research and Development Program, Agriculture Handbook
No. 575: 15 p.
Billings, R.F. and W.W. Upton. 1993. Effectiveness of synthetic
behavioral chemicals for manipulation and control of southern pine
beetle infestations in East Texas. USDA Forest Service, Southern Forest
Experiment Station. General Technical Report: 555-568.
Clarke, S.L. and J.T. Nowak. 2009. Southern pine beetle. Forest Insect
and Disease Leaflet No. 49, USDA Forest Service, Portland, Oregon,
United States. 8 p.
Billings, R.F. and W.W. Upton. 2010. A methodology for assessing
annual risk of southern pine beetle outbreaks across the Southern Region,
using pheromone traps, p. 73-85. In: Pye, J.M., H.M. Rauscher, Y. Sands,
D.C. Lee, J.S. Beatty (tech. eds.). 2010. Advances in threat assessment
and their application to forest and rangeland management. General
Technical Report PNW-GTR-802. U.S. Department of Agriculture,
Forest Service, Pacific Northwest and Southern Research Stations,
Portland, Oregon, United States. 708 p., 2 vol. (Image available at
http://www.threats.forestencyclopedia.net/p/p3291).
A Photographic Guide 207

64. Western Pine Beetle, Dendroctonus brevicomis LeConte (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Western pine beetle, Dendroctonus brevicomis LeConte, is a native

bark beetle of the western United States that attacks and kills ponderosa
(Pinus ponderosa Douglas ex P & C. Lawson) and Coulter (P. coulteri
D. Don) pines greater than 15 cm in diameter, regardless of age or
vigor, including apparently healthy trees. Outbreaks are most common
in dense, even-age stands, but also occur in areas of mixed conifer
species. Mortality is often subtle yet significant when large old trees
are killed individually by the beetle. Large outbreaks are common and
over one million trees (more than 1 billion board feet of timber) may be
killed each year during outbreaks. Outbreaks have been known to occur
following wildfires in ponderosa pine stands. Beetle attacks are first
visible due to pitch tubes that form at the sites where female beetles bore
into the tree to create oviposition galleries. However, pitch tubes of this
species are less conspicuous than for other bark beetles and frequently
are not seen at all. Attacking adult beetles carry spores of a blue-staining
fungus, Ceratocystis minor (Hedg.), which contributes to tree death by
blocking the water-conducting vessels in the wood. The egg galleries are
about the width of an adult beetle and are usually tightly packed with
boring dust. There are two to four generations per year, depending on
latitude and altitude. Eggs are laid singly along the oviposition gallery
and larvae then make short lateral galleries as they feed. As is true for
many bark beetles, aggregation pheromones produced by the beetle
coordinate a mass attack on selected trees. Thinning of trees to reduce
density can be used to reduce stand susceptibility to mortality from
western pine beetle.

Hosts Commonly Attacked This beetle attacks ponderosa (P. ponderosa Douglas ex P & C. Lawson)
and Coulter (P. coulteri D. Don) pines.

Distribution Western pine beetle is most damaging

in California, but its range extends
northward into Oregon, Washington,
Idaho, and southern British Columbia, as
well as eastward into Montana, Nevada,
Utah, Colorado, Arizona, New Mexico,
and western Texas. It also occurs in
northwestern Mexico.

Figure 1. Distribution (in western

North America) of the western pine
beetle, Dendroctonus brevicomis.
(USDA Forest Service • Forest
Insect & Disease Leaflet 1)
208 Forest Pest Insects in North America:

Images of
Western Pine Beetle

Figure 2. Adult of western pine beetle, Dendroctonus brevicomis (left), and close
up of head (right); note lack of rows of tubercules on either side of the frontal
groove, as are present in the southern pine beetle, Dendroctonus frontalis
(Zimmermann). (Left: Erich G. Vallery, USDA Forest Service - SRS-4552,
Bugwood.org; right: Pest and Diseases Image Library, Bugwood.org)

Figure 3. Pitch tubes

of western pine beetle.
(Kenneth E. Gibson,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 209

Images of Western Figure 4. Larvae of

western pine beetle.
Pine Beetle (continued) (Ladd Livingston, Idaho
Department of Lands,
Bugwood.org)

Figure 5. Galleries of western pine beetle in ponderosa pine. (Left: USDA

Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org; right:
William M. Ciesla, Forest Health Management International, Bugwood.org)

Figure 6. Shot holes

formed in ponderosa
pine by emergence of
western pine beetles.
(Donald Owen,
California Department
of Forestry and Fire
Protection,
Bugwood.org)
210 Forest Pest Insects in North America:

Images of Western Figure 7. Two views of

stands of ponderosa
Pine Beetle (continued) pines killed by western
pine beetle. (Top:
William M. Ciesla,
Forest Health
Management
International,
Bugwood.org;
bottom: James Everitt,
Bugwood.org)

Figure 8. The predacious clerid beetle Enoclerus lecontei, a predator of western

pine beetle. (Brytten Steed, USDA Forest Service, Bugwood.org)
A Photographic Guide 211

Important Biological Control Important predators of brood of western pine beetle are the clerid beetles
Agents Related to this Enoclerus lecontei (Wolcott) and Enoclerus sphegeus Fabricius, as well
Pest Species as the blue-green ostomid Temnochila chlorodia Mannerheim and the
fly Medetera aldrichii Wheeler. Common parasites are Roptrocerus
xylophagorum Ratzeburg, Dinotiscus burkei Crawford, and Coeloides sp.
nr. brunneri Vierick.

Web Links for Information http://www.forestpests.org/acrobat/fidl1.pdf; USDA Forest Service,

on Western Pine Beetle Forest Insect and Disease Leaflet #1.
http://forestry.nv.gov/forestry-resources/forest-health/western-pine-
beetle/; fact sheet of the Nevada Division of Forestry.

Articles Miller, J.M. and F.P. Keen. 1960. Biology and control of the western pine
beetle. U.S. Department of Agriculture Misc. Publ. 800, Washington,
DC: 381 p.
Stark, R.W. and D.L. Dahlsten (eds.). 1970. Studies on the population
dynamics of the western pine beetle, Dendroctonus brevicomis
LeConte (Coleoptera: Scolytidae). University of California, Division of
Agricultural Science, Berkeley, California, USA: 174 p.
Wallin, K.F., T.E. Kolb, K.R. Skov, and M. Wagner. 2008. Forest
management treatments, tree resistance, and bark beetle resource
utilization in ponderosa pine forests of northern Arizona. Forest Ecology
and Management 255(8/9): 3263-3269.
212 Forest Pest Insects in North America:

65. Jeffrey Pine Beetle, Dendroctonus jeffreyi Hopkins (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Jeffrey pine beetle, Dendroctonus jeffreyi Hopkins, is a native North
American bark beetle that attacks only Jeffrey pine (Pinus jeffreyi Grev.
and Balf.) and it is that tree’s most serious insect pest. This pine beetle
occurs in the Sierra Nevada wherever Jeffrey pine is present. In most
cases this beetle occurs at low densities and attacks only small numbers
of slow growing trees with reduced vigor. However, large outbreaks can
occur when forest conditions are favorable, especially during extended
droughts. Adults emerge from infested trees in April or May. Mass attack
is initiated by colonizing beetles, which release aggregation pheromones.
Females bore in and excavate a vertical gallery where they lay their
eggs. The oviposition gallery is packed with boring dust, and larvae
later form short individual lateral tunnels where they feed. Jeffrey pine
beetles overwinter as larvae or as pupae in chambers excavated at the
end of larval galleries. Depending on local climate there are one or two
generations per year. Beetles infect trees with a blue-stain fungus, which
kills the tree by clogging the tree’s vascular system. Signs of attack
include pitch tubes on the mid or lower trunk. In weak trees pitch tubes
may be absent.

Hosts Commonly Attacked This beetle attacks only Jeffrey pine (P. jeffreyi).

Distribution Jeffrey pine beetle is found within the range of Jeffrey pine, principally
in California, with the exceptions of the San Jacinto, Santa Rosa, and
Luguna mountain ranges of southern California, where this bark beetle
has not been observed.

Images of
Jeffrey Pine Beetle

Figure 1. Adults of Jeffrey pine beetle, Dendroctonus jeffreyi. (Both photos:

Erich G. Vallery, USDA Forest Service - SRS-4552, Bugwood.org)
A Photographic Guide 213

Images of Jeffrey
Pine Beetle (continued)

Figure 2. Eggs of Jeffrey pine beetle in oviposition gallery (above dark line of
gallery). (Dave Powell, USDA Forest Service, Bugwood.org)

Figure 3. Larva of Jeffrey pine beetle, removed from gallery. (Erich G. Vallery,
USDA Forest Service - SRS-4552, Bugwood.org)
214 Forest Pest Insects in North America:

Images of Jeffrey Figure 4. Pupa of Jeffrey

pine beetle. (tia smith,
Pine Beetle (continued) Bugwood.org)

Figure 5. Galleries of
adults (vertical) and
larvae (horizontal) of
Jeffrey pine beetle.
(Darren Blackford,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 215

Images of Jeffrey
Pine Beetle (continued)

Figure 6. Pines killed by Jeffrey pine beetle near Lake Tahoe, California.
(Darren Blackford, USDA Forest Service, Bugwood.org)

Important Biological Control Natural enemies of this bark beetle have received little attention,
Agents Related to this but likely are similar to those groups attacking western pine beetle
Pest Species (Dendroctonus brevicomis LeConte).

Web Links for Information http://forestry.nv.gov/forestry-resources/forest-health/jeffery-pine-beetle/;

on Jeffrey Pine Beetle fact sheet of the Nevada Division of Forestry.
http://www.fs.fed.us/r6/nr/wildlife/decaid/IandDSpecies/
Jeffrey%20pine%20beetle.html; USDA Forest Service fact sheet.
http://www.unce.unr.edu/publications/files/ho/other/fs9840.pdf; fact
sheet of the University of Nevada at Reno.

Articles Smith, R.H., B.E. Wickman, R.C. Hall, C.J. DeMars, and G.T. Ferrell.
1981. The California pine risk-rating system: its development, use and
relationship to other systems. In: Hedden, R.L., S.J. Barras, J.E. Coster.
(tech. coords.). Hazard-rating systems in forest insect pest management:
symposium proceedings. Athens, Georgia, July 31-August 1, 1980.
General Technical Report WO-27, USDA Forest Service, Washington,
DC: 53-69.
Smith, S.L., R.B. Borys, and P.J. Shea. 2008. Jeffrey pine beetle. USDA
Forest Service, Forest Pest Leaflet 11: 7 p.
216 Forest Pest Insects in North America:

66. Mountain Pine Beetle, Dendroctonus ponderosae Hopkins (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Mountain pine beetle, Dendroctonus ponderosae Hopkins, a native North
American species, is the most destructive bark beetle of the western
United States and Canada, where it occurs widely on a variety of pines,
including lodgepole (Pinus contorta Douglas), ponderosa (P. ponderosa
Douglas ex C. Lawson), western white (P. monticola Douglas ex D.
Don), whitebark (P. albicaulis Engelmann) and sugar (P. lambertiana
Douglas) pines. Historically, highly destructive outbreaks of this species
have occurred repeatedly, decimating mature forests of such species as
lodgepole pine over extensive areas. All trees above 10 cm in diameter
are susceptible to attack. Beetle attacks are usually concentrated along
the main trunk, from about one meter above the ground up to the middle
branches. As with many bark beetles, signs of infestations include
pitch tubes on the trunk, red boring dust in bark crevices, discoloration
of foliage as trees die, and blue discoloration of the wood caused by
several associated fungi: Grosmannia clavigera (Robinson-Jeffrey and
Davidson) Zipfel, de Beer, and Wingfield; and Ophiostoma montium
(Rumbold) von Arx. Females dig egg galleries under the bark and lay
eggs on alternate sides of the gallery. Larvae excavate short feeding
tunnels at right angles to the egg gallery. Mature larvae construct pupal
cells at the ends of the larval galleries, where larvae pupate and from
which new adults later emerge. Most commonly there is one generation
per year but there may be only one every two years in the coldest parts
of the species’ range. Larvae and adults are the overwintering stages.

Hosts Commonly Attacked The principal hosts of this beetle are lodgepole (P. contorta), sugar
(P. lambertiana), western white (P. monticola), ponderosa (P. ponderosa),
and whitebark (P. albicaulis) pines, but Pinus aristata Engelmann,
P. balfouriana Balfour, P. coulteri D. Don, P. edulis Engelmann,
P. flexilis E. James, P. strobiformis Engelm., P. longaeva D.K. Bailey,
and P. monophylla Torr. and Frém. have also been recorded as hosts.

Distribution Mountain pine beetle is found

throughout the pine forests of western
Canada, the western United States and
northern Mexico (Baja California).

Figure 1. Distribution of mountain pine

beetle, Dendroctonus ponderosae,
in North America. (USDA Forest
Service • Forest Insect
& Disease Leaflet 2)
A Photographic Guide 217

Images of
Mountain Pine Beetle

Figure 2. Adults of mountain pine beetle. (Left: USDA Forest Service - Region 2
- Rocky Mountain Region Archive, USDA Forest Service, Bugwood.org; right:
Ron Long, Simon Fraser University, Bugwood.org)

Figure 3. Pitch tubes of mountain pine beetle during initiation of attack. (Left:
Steven Katovich, USDA Forest Service, Bugwood.org; right: Leslie Chong,
Simon Fraser University, Bugwood.org)

Figure 4. Larvae of mountain pine beetle in their galleries. (Left: USDA Forest
Service - Ogden Archive, USDA Forest Service, Bugwood.org; right: Ladd
Livingston, Idaho Department of Lands, Bugwood.org)
218 Forest Pest Insects in North America:

Images of Mountain
Pine Beetle (continued)

Figure 5. Pupae and adults (left) of mountain pine beetle; callow adult (right).
(Left: USDA Forest Service - Ogden Archive, USDA Forest Service,
Bugwood.org; right: USDA Forest Service - Region 2 - Rocky Mountain
Region Archive, USDA Forest Service, Bugwood.org)

Figure 6. Galleries of mountain pine beetle (left) and galleries and blue staining
of wood (right). (Left: Leslie Chong, Simon Fraser University, Bugwood.org;
right: Carl Jorgensen, USDA Forest Service, Bugwood.org)

Figure 7. Lodgepole pines (left) killed by mountain pine beetle; view of infested
pines (right). (Left: USDA Forest Service - Ogden Archive, USDA Forest Service,
Bugwood.org; right: USDA Forest Service - Region 2 - Rocky Mountain Region
Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 219

Images of Mountain
Pine Beetle (continued)

Figure 8. Adult of Enoclerus sphegeus Fabricus, a predator of mountain pine

beetle. (Brytten Steed, USDA Forest Service, Bugwood.org)

Important Biological Control Natural enemies of mountain pine beetle include predaceous insects such
Agents Related to this as Enoclerus sphegeus Fabricius, Temnochila chlorodia Mannerheim,
Pest Species and Medetera aldrichii Wheeler; and the parasitoid Coeloides
dendroctoni Cushman.

Web Links for Information http://en.wikipedia.org/wiki/Mountain_pine_beetle; Wikipedia article,

on Mountain Pine Beetle including comments on implications of current massive outbreak in
Canada.
http://www.ext.colostate.edu/pubs/insect/05528.html; factsheet of
Colorado State University.
http://forestry.nv.gov/forestry-resources/forest-health/mountain-pine-
beetle/; Nevada Division of Forestry.
http://www.fs.fed.us/r6/nr/fid/fidls/fidl-2.pdf; USDA Forest Service
Forest Pest and Disease Leaflet #2. 2009 revision; provides updated,
more detailed distribution map.
http://www.usu.edu/beetle; USDA Forest Service, Rocky Mountain
Research Station, Western Bark Beetles website including literature.

Articles Safranyik, L., A.L. Carroll, and B. Wilson. 2006. The biology and
epidemiology of the mountain pine beetle in lodgepole pine, p. 3-66.
In: Safraynyik, L. and W.R. Wilson (eds.). The Mountain Pine Beetle:
A synthesis of the biology, management and impacts on lodgepole pine.
Natural Resources Canada, Canadian Forest Service, Pacific Centre,
Victoria, British Columbia: 304 p.
Robertson, C., T.A. Nelson, D.E. Jelinski, M.A. Wulder, and B. Boots.
2009. Spatial-temporal analysis of species range expansion: the case
of the mountain pine beetle, Dendroctonus ponderosae. Journal of
Biogeography 36(8): 1446-1453.
220 Forest Pest Insects in North America:

67. Douglas-fir Beetle, Dendroctonus pseudotsugae Hopkins (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, a native

North American bark beetle, is the most important pest of Douglas-fir
(Pseudotsuga menziesii [Mirbel] Franco) in western North America
and the only bark beetle attacking this important timber species. Like
many bark beetles, this species often persists at low levels, breeding
primarily in felled, injured or diseased trees and doing little damage to
any particular stand. However, at times populations develop that are able
to kill even healthy trees, over large areas. Such outbreaks may develop
after extensive wind damage, droughts, or large fires that greatly increase
the number of susceptible trees. The signs of this bark beetle include
reddish or yellowish boring dust caught in bark crevices. Live trees may
respond to attack with abundant resin flow. A male and female beetle
work together to dig an oviposition gallery, where the female lays her
eggs, which are deposited in groups (10-36), spaced out at intervals along
the oviposition gallery. Larval dig feeding galleries that diverge from the
oviposition gallery in groups that resemble fans. Larvae pupate in cells
constructed at the ends of the larval galleries. The Douglas-fir beetle
has one generation per year. The insect mostly overwinters as adults,
but with some large larvae present as well. Adult emergence from the
wood occurs from April to June, depending on location and altitude, and
newly emerged beetles attack new trees in July and August. In coastal
areas, host tree resistance is usually adequate to check population growth
of this species. But in inland areas, where trees are subject to greater
environmental stress, outbreaks are more common.

Hosts Commonly Attacked The only breeding host of this beetle is Douglas fir (P. menziesii).

Distribution Douglas-fir beetle is found throughout the range of its principal host in
the western United States, Mexico, and Canada.

Images of Figure 1. Pair of

adult Douglas-fir
Douglas-fir Beetle beetles, Dendroctonus
pseudotsugae, inside
gallery. (Constance Mehmel,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 221

Images of Douglas-fir
Beetle (continued)

Figure 2. Larvae of Douglas-fir beetle in their galleries. (Left: Scott Tunnock,

USDA Forest Service, Bugwood.org; right: Malcolm Furniss, Bugwood.org)

Figure 3. Red boring dust (left) and resin flow (right) on the bark of a Douglas-fir
produced by adult Douglas-fir beetles. (Left: Doug Page, USDI Bureau of Land
Management, Bugwood.org; right: Constance Mehmel, USDA Forest Service,
Bugwood.org)

Figure 4. Galleries of Douglas-fir beetles. (Left: USDA Forest Service - Ogden

Archive, USDA Forest Service, Bugwood.org; right: Kenneth E. Gibson, USDA
Forest Service, Bugwood.org)
222 Forest Pest Insects in North America:

Images of Douglas-fir
Beetle (continued)

Figure 5. Douglas-fir killed by Douglas-fir beetle. (A. Steven Munson, USDA

Forest Service, Bugwood.org)

Figure 6. Adult of
Enoclerus sphegeus
Fabricus. (Brytten
Steed, USDA Forest
Service, Bugwood.org)

Figure 7. The braconid

parasitoid, Coeloides
vancouverensis (Dalla
Torre), ovipositing
through shaved
Douglas-fir bark onto
Douglas-fir beetle
larvae. (Roger Ryan,
USFS PNW Station,
Bugwood.org)
A Photographic Guide 223

Important Biological Control Among the important natural enemies of the Douglas-fir beetle are
Agents Related to this Enoclerus sphegeus Fabricius, Thanasimus undatulus Say, Temnochila
Pest Species chlorodia Mannerheim, Coeloides brunneri Viereck, and Medetera
aldrichii Wheeler.

Web Links for Information http://www.forestry.ubc.ca/fetch21/FRST308/lab6/dendroctonus_

on Douglas-fir Beetle pseudotsugae/douglas.html; fact sheet of the University of British
Columbia.
http://www.forestpests.org/acrobat/fidl5.pdf; fact sheet of the USDA
Forest Service, Insect pest and disease leaflet #5.

Articles Furniss, M.M. 1979. An annotated bibliography of the Douglas-fir beetle

(Dendroctonus pseudotsugae Hopkins). USDA Forest Service General
Technical Report (INT-48): 40 p.
224 Forest Pest Insects in North America:

68. Engraver Beetle, Ips typographus (L.) (Coleoptera: Curculionidae,

Scolytinae)

Orientation to Pest The engraver beetle, Ips typographus (L.), is native to Europe and
has not yet invaded North America, but is a species of concern as a
potentially serious pest of spruce (Picea). It is also referred to as the
European spruce bark beetle. This species is the most damaging bark
beetle in Europe, and historically many widespread infestations of long
duration have occurred. As with many bark beetles, attack on living trees
is facilitated by chemically coordinated mass attack and the vectoring
of a tree-killing fungus, in this case Ceratocystis polonica Siem. The
biology and life cycle of the insect is typical of bark beetles. Females
dig oviposition galleries and deposit eggs along their length. Larvae then
dig feeding galleries at right angles to the oviposition gallery, creating
a fan-like appearance. Larvae pupate in chambers at the ends of larval
galleries. Outbreaks are apparently initiated by events or conditions
that increase quickly the number of stressed or dead trees available as
breeding sites. Storms, drought, and logging (which increases the number
of wind-thrown trees subsequently) have all been suggested as factors
able to stimulate outbreaks. It has been suggested that natural enemies
do not control low density populations, and that the regulatory factor is
mortality from competition among larvae when breeding trees are scarce.
However, other authors find that natural enemies cause up to 82 percent
mortality to brood, suggesting they may indeed be important. Damage
may be reduced by prompt removal of recently attacked trees because as
many as 80 percent of parent beetles start a “sister brood” within weeks.
Mass trapping with a synthetic pheromone blend is also employed.

Hosts Commonly Attacked This beetle attacks spruces (Picea), such as Norway spruce (Picea abies
[L.] H. Karst.).

Distribution Engraver beetles are found throughout much of central Europe,

Scandinavia, and east into Russia, northeastern China, Korea, and Japan.

Images of
Engraver Beetle

Figure 1. Adults of the engraver beetle, Ips typographus (left: light brown, young
adults; right: fully darkened, mature specimen). (Left: Bjørn Økland, Norwegian
Forest Research Institute, Bugwood.org; right: Maja Jurc, University of Ljubljana,
Bugwood.org)
A Photographic Guide 225

Images of Engraver Figure 2. Close-up of

one egg of the engraver
Beetle (continued) beetle. (Daniel Adam,
Office National des
Forêts, Bugwood.org)

Figure 3. Larvae of engraver beetle (in a medium density infestation).

(Left: SRPV, Limoges Archive, Les Services Régionaux de la Protection des
Végétaux, Bugwood.org; center: Louis-Michel Nageleisen, Département de la
Santé des Forêts, Bugwood.org; right: Daniel Adam, Office National des Forêts,
Bugwood.org)
226 Forest Pest Insects in North America:

Images of Engraver
Beetle (continued)

Figure 4. Pupae (left) of the engraver beetle in chambers at tips of larval

galleries (in a low density infestation); right, close up of one pupa. (Left: Daniela
Lupastean, University of Suceava, Bugwood.org; right: Maja Jurc, University of
Ljubljana, Bugwood.org)

Figure 5. Galleries of engraver beetles: left, adults cutting oviposition galleries;

right, finished larval galleries branching off oviposition gallery with pupation
chambers at tips. (Both photos: Milan Zubrik, Forest Research Institute -
Slovakia, Bugwood.org)

Figure 6. Spruce trees in Slovakia killed by the engraver beetle. (Milan Zubrik,
Forest Research Institute - Slovakia, Bugwood.org)
A Photographic Guide 227

Images of Engraver
Beetle (continued)

Figure 7. Traps used to detect engraver beetles (left) or for mass trapping (right,
in logged area). (Both photos: Milan Zubrik, Forest Research Institute - Slovakia,
Bugwood.org)

Figure 8. Left, larva of external parasitoid on larva of engraver beetle; right,

multiple cocoons of ichneumonid parasitoid formed in larval galleries of engraver
beetle. (Also visible in photo on right in upper and lower areas are the galleries
of the smaller, very common European species, Pityogenes chalcographus [L.]).
(Left: Gyorgy Csoka, Hungary Forest Research Institute, Bugwood.org; right:
Petr Kapitola, State Phytosanitary Administration, Bugwood.org)

Figure 9. Larva of the long-legged fly Medetera signaticornis Loew, a

specialized predator of bark beetle larvae, known only from Ips typographus,
usually in high abundance in Western Europe. (Jiri Hulcr, Michigan State
University, Bugwood.org)
228 Forest Pest Insects in North America:

Images of Engraver
Beetle (continued)

Figure 10. The red-bellied clerid beetle, Thanasimus formicarius (L.), is an

important predator of the engraver beetle in Europe (left, adult; right, larva).
(Left: Scott Bauer, USDA Agricultural Research Service, Bugwood.org; right:
Daniela Lupastean, University of Suceava, Bugwood.org)

Important Biological Control The natural enemies of the engraver beetle have been reported from
Agents Related to this many locations and a large number of species of parasitoids and
Pest Species predators have been observed. Among the commonly reported species
are the red-bellied clerid beetle (Thanasimus formicarius [L.]), the
long-legged fly Medetera signaticornis Loew, and various braconid and
ichneumonid parasitoids. See the review by Mills and Schlup (1989).

Web Links for Information http://www.barkbeetles.org/exotic/htypgrph.html; fact sheet describes

on Engraver Beetle characters for species recognition.
http://www.fs.fed.us/foresthealth/technology/invasives_ipstypographus_
riskmaps.shtml; USDA website that describes forests in the United States
at potential risk from this bark beetle.
http://www.issg.org/database/species/ecology.asp?si=1441&fr=1&sts
=&lang=EN; factsheet of the Global Invasive Species Database, with
information on biology and method of movement between countries.

Articles Mills, N.J. and M. Schlup. 1989. The natural enemies of Ips typographus
in central Europe: Impact and potential use in biological control. In:
Kulhavy, D.L. and M.C. Miller (eds.). Potential for Biological Control
of Dendroctonus and Ips Bark Beetles. Published by the Center for
Applied Studies, School of Forestry, Stephen F. Austin State University,
Nacogdoches, Texas, USA: 131-146.
Wermelinger, B. 2004. Ecology and management of the spruce bark
beetle Ips typographus – a review of recent research. Forest Ecology and
Management 202(1/3): 67-82.
Grégoire, J.-C. and H.F. Evans. 2004. Damage and control of Bawbilt
organisms - an overview, pp. 19-37. In: Lieutier, F., K.R. Day, A. Battisti,
J.-C. Grégoire, and H.F. Evans (eds.). Bark and Wood Boring Insects in
Living Trees in Europe, a Synthesis. Springer, Netherlands.
A Photographic Guide 229

69. Pine Engraver Beetle, Ips pini (Say) (Coleoptera: Curculionidae

[formerly Scolytidae])

Orientation to Pest The pine engraver beetle, Ips pini (Say), is a native bark beetle with a
very wide distribution in North America. In natural forests, this species
is of little importance and is mainly associated with smaller diameter
(12-20 cm) pine or spruce trees that are dying or have been recently
killed by storms or logging. Under these conditions the species is
beneficial and contributes to recycling of stressed or dying trees. Its
numbers may increase when logging, thinning, or storms increase the
amount of breeding material that is locally available. When this happens,
healthy trees in the area may also be attacked. This species is considered
the most important bark beetle attacking red pine (Pinus resinosa Sol.
ex Aiton) in the Great Lakes states of the USA. Male beetles bore into
trees and create a nuptial chamber. Each male produces pheromones
that attract multiple females of the same species. After mating, 3 to 6
females construct oviposition galleries in the inner bark that radiate out
from the nuptial chamber in a “Y” or “H” shaped pattern. Larval galleries
branch off the oviposition galleries, and pupation takes place in cells at
the end of the larval galleries. Pine engraver beetles can vector a blue
stain fungus, which aids the beetle in killing the host tree by clogging
the water-transporting elements in the tree. Symptoms and signs of pine
engraver attack are essentially the same as those of many bark beetles,
including yellow, red, and brown foliage of dying trees, together with
boring dust and pitch tubes on the bark. Commonly, there are 3 to 4
generations per year.

Hosts Commonly Attacked Ips pini breeds in all the species of pines and spruce in its range and
is of greatest concern in lodgepole (P. contorta Douglas), ponderosa
(P. ponderosa Douglas ex C. Lawson), Jeffrey (P. jeffreyi Balf.), and red
(P. resinosa) pines.

Distribution The pine engraver is found from northern Canada and Alaska to northern
Mexico. It occurs throughout most of the coniferous forests of North
America except the Pacific Coastal forests, the southern pine forests, and
Mexico. In the eastern United States, it is widespread in the north and
extends along the Appalachian Mountains south to Georgia.

Images of Figure 1. Adult of pine

engraver beetle, Ips
Pine Engraver Beetle pini. (Pennsylvania
Department of
Conservation and
Natural Resources
- Forestry Archive,
Bugwood.org)
230 Forest Pest Insects in North America:

Images of Pine Engraver Figure 2. Nuptial

chamber with five
Beetle (continued) oviposition galleries
branching off, with
females in galleries at 2
and 10 o’ clock. (Scott
Tunnock, USDA Forest
Service, Bugwood.org)

Figure 3. Larval galleries of the pine engraver projecting at right angles to the
oviposition galleries. (USDA Forest Service - Region 6 - Pacific Northwest
Archive, USDA Forest Service, Bugwood.org)

Figure 4. Frass ejected from bore holes by adult pine engravers. (Brytten Steed,
USDA Forest Service, Bugwood.org)
A Photographic Guide 231

Images of Pine Engraver Figure 5. Adult

emergence holes of
Beetle (continued) pine engraver beetles.
(Steven Katovich,
USDA Forest Service,
Bugwood.org)

Figure 6. Lundgren
funnel trap used to
detect pine engraver
beetles at a slash
pile. (Brytten Steed,
USDA Forest Service,
Bugwood.org)
232 Forest Pest Insects in North America:

Images of Pine Engraver Figure 7. Group of red

pines (Pinus resinosa
Beetle (continued) Soland.) in Minnesota
killed by pine engraver
beetle. (Steven
Katovich, USDA Forest
Service, Bugwood.org)

Figure 8. Larva (left) and adult (right) of Thanasimus dubius, a common

predator of the pine engraver beetle. (Left: Gerald J. Lenhard, Louiana State
Univ, Bugwood.org; right: USDA Forest Service Archive, USDA Forest Service,
Bugwood.org)

Figure 9. Larva (left) and adult (right) of Enoclerus sphegeus, a common predator
of the pine engraver beetle. (Left: USDA Forest Service - Region 2 - Rocky
Mountain Region Archive, USDA Forest Service, Bugwood.org; right: Brytten
Steed, USDA Forest Service, Bugwood.org)
A Photographic Guide 233

Important Biological Control The natural enemies of the pine engraver beetle are similar to those
Agents Related to this of many bark beetles and include several predatory clerids, such as
Pest Species Thanasimus dubius (Fabricius) and Enoclerus sphegeus (Fabricius), as
well as such parasitoids as the braconid Coeloides dendroctoni Cushman
and the pteromalids Pachyceras xylophagorum Ratz., Rhopalicus
pulchripennis (Crawford), and Tomicobia tibialis Ashmead.

Web Links for Information http://wiki.bugwood.org/Ips_pini; a Bugwiki fact sheet on Ips pini.
on Pine Engraver Beetle http://www.fs.fed.us/r6/nr/fid/fidls/f122.htm; a USDA Forest Service
Forest and Insect Disease Leaflet: Gibson, K.E., Kegley, S.J., and
Livingston R.L. 1997. Pine Engraver, Ips pini, in the Western United
States. Forest Insect and Disease Leaflet No. 122. USDA Forest Service.
http://www.ext.colostate.edu/pubs/insect/05558.html; perspective on
other species of Ips (11 species in Colorado, featured on this website).

Articles Erbilgin, N., E.V. Nordheim, B.H. Aukema, and K.F. Raffa. 2002.
Population dynamics of Ips pini and Ips grandicollis in red pine
plantations in Wisconsin: within- and between-year associations with
predators, competitors, and habitat quality. Environmental Entomology
31: 1043-1051.
Dahlsten, D.L., D.L. Six, N. Erbilgin, K.F. Raffa, A.B. Lawson, and
D.L. Rowney. 2003. Attraction of Ips pini (Coleoptera: Scolytidae) and
its predators to various enantiomeric ratios of ipsdienol and lanierone in
California: Implications for the augmentation and conservation of natural
enemies. Environmental Entomology 32: 1115-1122.
234 Forest Pest Insects in North America:

70. Southern Pine Engraver, Ips grandicollis (Eichhoff) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The southern pine engraver (also called the eastern five-spined Ips),
Ips grandicollis (Eichhoff), is a native bark beetle that occurs in the
eastern parts of Canada and the United States, in Central America, and
on some Caribbean islands. It is also invasive in Australia, where it is a
pest of plantations of Monterey pine (Pinus radiata D. Don). The elytra
(hardened forewings) end in a scoped out area with typically 5 teeth on
each side. It breeds most often in recently felled trees and logging slash,
but can also breed in limbs and trunks of live trees that are under attack
by other bark beetle species. Most breeding occurs in limbs and the upper
crown. Drought and logging both can potentially stimulate outbreaks of
this species, causing small infestations to expand in area. The biology
of this species is similar to that of Ips pini (Say). Male beetles create a
nuptial chamber beneath the bark and produce pheromones that attract
multiple females. After mating, the females construct 3 to 5 oviposition
galleries that branch off from the nuptial chamber. The parent galleries
are kept free of boring dust, unlike those of the southern pine beetle
(Dendroctonus frontalis Zimmermann). Larval galleries branch off the
oviposition galleries and are usually run horizontally across the tree
trunk. Pine engraver beetles can vector a blue stain fungus, which aids
the beetle in killing the host tree by clogging the water-transporting
elements in the tree. Symptoms and signs of pine engraver attack are
essentially the same as those of many bark beetles, including the yellow,
red or brown foliage characteristic of dying trees, together with boring
dust and pitch tubes on the bark. There are six or more generations per
year in the southern United States.

Hosts Commonly Attacked Ips grandicollis breeds in most of the pine species in the southern United
States. North of the southern pines region, this species is principally
found associated with pitch pine (Pinus rigida Mill.).

Distribution The southern pine engraver is found in eastern Canada and in the eastern
United States from Massachusetts west to Minnesota, Nebraska and
Texas, and throughout the southern states.
A Photographic Guide 235

Images of
Southern Pine Engraver

Figure 1. Adult of southern pine engraver, Ips grandicollis. Right, see the scoped
out area of the fore wings and the surrounding teeth. (Both photos: Pest and
Diseases Image Library, Bugwood.org)

Figure 2. Comparative size of southern pine engraver (Ips grandicollis) relative

to other southern pine bark beetles. (John L. Foltz, University of Florida,
Bugwood.org)

Figure 3. Egg of southern pine engraver. (Lacy L. Hyche, Auburn University,

Bugwood.org)
236 Forest Pest Insects in North America:

Images of Southern Pine

Engraver (continued)

Figure 4. Callow adult (left), larva (middle), and pupa (right) of southern pine
engraver. (Roger Anderson, Duke University, Bugwood.org)

Figure 5. Left, nuptial chambers (see arrows) of southern pine engraver, with
five and four oviposition galleries branching off; right, larval galleries branching
off oviposition galleries, with pupation cells at end of each gallery. (Left:
Jeffrey Eickwort, Florida Department of Agriculture and Consumer Services,
Bugwood.org; right: Texas Forest Service Archive, Texas Forest Service,
Bugwood.org)

Figure 6. Group of
Monterey pines (P.
radiata) in Australia,
killed by southern pine
engraver. (Dennis
Haugen, Bugwood.org)
A Photographic Guide 237

Images of Southern Pine Figure 7. The predatory

checkered beetle
Engraver (continued) Thanasimus dubius is a
common predator of the
southern pine engraver.
(USDA Forest Service
Archive, USDA Forest
Service, Bugwood.org)

Important Biological Control The natural enemies of the southern pine engraver include the predatory
Agents Related to this clerid Thanasimus dubius (Fabricius), the predatory trogositid beetle
Pest Species Temnochila virescens (Fabricius), as well as various parasitoids,
including Roptrocerus xylophagorum Ratzeburg. Thanasimus dubius and
Temnochila virescens have been introduced into Australia against this
bark beetle.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/ips/ipsfidl.htm; a USDA Forest

on Southern Pine Engraver Service leaflet on Ips pine bark beetles in the southern USA, comparing
several species, including I. grandicollis.
http://www.eppo.org/QUARANTINE/insects/Ips_grandicollis/IPSXGR_
ds.pdf; EPPO data sheet on Ips grandicollis.

Articles Foltz, J.L., E.P. Merkel, and R.C. Wilkinson. 1984. Annotated
bibliography of Dendroctonus terebrans (Oliver), Ips avulsus (Eichhoff),
Ips grandicollis (Germar) in the southeastern USA. Monograph #12,
Agricultural Experiment Station, University of Florida: 47 p.
Lawson, S.A. and F.D. Morgan. 1992. Rearing of two predators,
Thanasimus dubius and Temnochila virescens, for the biological control
of Ips grandicollis in Australia. Entomologia Experimentalis et Applicata
65: 225-233.
Abbott, I. 1993. Review of the ecology and control of the introduced
bark beetle Ips grandicollis in Western Australia, 1952-1990.
CALMScience 1: 35-46.
238 Forest Pest Insects in North America:

71. Six-spined Ips, Ips calligraphus (Germar) (Coleoptera: Curculionidae,

Scolytinae)

Orientation to Pest The six-spined Ips, Ips calligraphus (Germar), is a native bark beetle
found in both the eastern and western United States. Two subspecies
of I. calligraphus are recognized that occur in the United States. Ips
calligraphus ponderosae is found attacking ponderosa pine (Pinus
ponderosa Douglas ex C. Lawson) in the Black Hills of South Dakota,
the eastern Rocky Mountains, and the northern Sierra Madre Oriental
of Mexico. Ips calligraphus calligraphus is found in the eastern
United States and California (where it likely was introduced from the
eastern United States). The subspecies found in the eastern United
States may attack all species of southern pines, including pitch pine
(P. rigida Millar). The eastern and western subspecies of I. calligraphus
breed in trees that have been felled, are dying, under drought stress, or
attack from other bark beetles. Trunks, stumps, and large limbs may
be attacked. Trunks of apparently healthy pines may sometimes be
attacked, especially in concert with other Ips or Dendroctonus species.
In the southern United States, this is one of the first species to attack
drought-stressed trees. Infestations may consist of scattered trees or fairly
large groups of trees, particularly during periods of drought. In general,
however, infestations of Ips species do not expand continuously as is
typical of southern pine beetle (Dendroctonus frontalis Zimmermann)
infestations. In the western United States, this species is most often
found in the thick-barked portion of the boles of ponderosa pines. Male
beetles construct a nuptial chamber where they attract and mate with two
to six females. The females then dig individual oviposition galleries that
radiate out from the nuptial chamber and are oriented vertically along the
trunk. Larval galleries are broad, long, and winding, and typically run
horizontally across the trunk. In the southern United States there are up
to 6 or more generations per year. In California, there are typically four
generations per year.

Hosts Commonly Attacked Ips calligraphus in the eastern United States may attack all native pine
species, but is found most commonly in loblolly (P. taeda L.), slash
(P. elliotii Engelm.), shortleaf (P. echinata Mill.), longleaf (P. palustris
Mill.), and pitch (P. rigida) pine. In the western United States, the
principal host is ponderosa pine (P. ponderosa).

Distribution The six-spined Ips subspecies found in the eastern United States occurs
from Florida, north to Massachusetts, and west to Minnesota and eastern
Texas. The western subspecies is found in Montana and South Dakotas,
south along the eastern Rocky Mountains.
A Photographic Guide 239

Images of
Six-spined Ips

Figure 1. Adult of six-spined Ips, Ips calligraphus, the largest of the southern U.S.
Ips species. (Left: David T. Almquist, University of Florida, Bugwood.org; right:
John L. Foltz, University of Florida, Bugwood.org)

Figure. 2. The six teeth

found at the end of the
depressed area at the
end of the body are
diagnostic for six-spined
Ips populations (see
discussion of subspecies
in Orientation to Pest
section). (J.R. Baker
and S.B. Bambara,
North Carolina
State University,
Bugwood.org)

Figure 3. Eggs of six-spined Ips. (Lacy L. Hyche, Auburn University,

Bugwood.org)
240 Forest Pest Insects in North America:

Images of Six-spined Ips

(continued)

Figure 4. Pupae of six-spined Ips. (Lacy L. Hyche, Auburn University,

Bugwood.org)

Figure 5. Nuptial
chamber (at tip of pen)
of the six-spined Ips,
with four oviposition
galleries. (Ronald F.
Billings, Texas Forest
Service, Bugwood.org)
A Photographic Guide 241

Images of Six-spined Ips Figure 6. Larval

galleries branching
(continued) off the oviposition
galleries (vertical) of
six-spined Ips.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)

Figure 7. Small pitch tube made by six-spined Ips. (G. Keith Douce, University of
Georgia, Bugwood.org)
242 Forest Pest Insects in North America:

Important Biological Control The natural enemies of the six-spined Ips include Thanasimus dubius
Agents Related to this (Fabricius), Temnochila virescens (Fabricius), and Medetera spp.,
Pest Species species that prey on a variety of Ips and Dendroctonus bark beetles.
Woodpeckers (Picidae), including the endangered red-cockaded
woodpecker (Picoides borealis), also may destroy I. calligraphus broods.
The impact of woodpecker feeding on infested trees is greatest during
winter months, when few other sources of food are available.

Web Links for Information http://www.barkbeetles.info/index.php; technical information on

on Six-spined Ips recognition, distribution and biology of bark beetles, especially those
of the southern United States and Mexico.
http://www.na.fs.fed.us/spfo/pubs/fidls/ips/ipsfidl.htm; USDA Forest
Service Pest and Disease Leaflet No. 129 “Ips bark beetles in the South”.
http://entnemdept.ufl.edu/creatures/trees/beetles/ips_beetles.htm; fact
sheet of the University of Florida covering various species of Ips bark
beetles.

Articles Wilkinson, R.C. and J.L. Foltz. 1980. A selected bibliography

(1959-1979) of three southeastern species of Ips engraver beetles.
Bulletin of the Entomological Society of America 26: 375-380.
Conner, M.D. and R.C. Wilkinson. 1983. Ips bark beetles in the
South. Forest Insect and Disease Leaflet No.129. U.S. Department of
Agriculture, Forest Service. 12 p.
Miller, M.C. 1984. Mortality contribution of insect natural enemies
to successive generations of Ips calligraphus (Germar) (Coleoptera,
Scolytidae) in loblolly pine. Zeitschrift für Angewandte Entomologie
98(5): 495-500.
Lanier, G.N., S.A. Teale, and J.A. Pajares. 1991. Biosystematics of the
genus Ips (Coleoptera: Scolytidae) in North America: review of the Ips
calligraphus group. The Canadian Entomologist 123: 1103-1124.
A Photographic Guide 243

72. Small Southern Pine Engraver, Ips avulsus Eichhoff (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The small southern pine engraver, Ips avulsus Eichhoff, is a native North
American bark beetle found throughout the eastern United States. It is
the smallest of the various species in this genus that breed in eastern
pines. Attacks are concentrated on sections of trees with thin-bark, such
as young pines or the limbs and tops of older trees. Attacks may occur
in young trees and the tops of large living trees, which may be killed.
However, when large living trees are attacked, the infestation is usually
limited to the upper crown, and the mid- and lower boles of the same
tree are often colonized by other species of Ips (Ips calligraphus Germar,
Ips grandicollis Eichhoff, or the southern pine beetle [Dendroctonus
frontalis Zimmermann]). Any tree cutting or accidental injury that creates
a pitch flow can induce attack by this species. Adults are very small
(2-3 mm) and the tip of the abdomen (the depressed area) is shallow,
with deep knobs and four small teeth on each margin. Males create a
nuptial chamber and produce a pheromone that attracts one or more
females. After mating, each female constructs a linear oviposition gallery
in the inner bark that parallels the grain of the wood. Larval galleries are
short and transverse, and each ends in a pupal cell in the phloem when
the larva completes development. Pine engraver beetles can vector a blue
stain fungus, which aids the beetle in killing the host tree by clogging
the water-transporting elements in the tree. Symptoms and signs of pine
engraver attack are essentially the same as those of many bark beetles,
including the yellow, red, or brown foliage characteristic of dying trees,
together with boring dust and pitch tubes on the bark. In the southern
United States, there may up to 10 or more generations per year.

Hosts Commonly Attacked Ips avulsus breeds in various southern pines, including eastern white
(Pinus strobus L.), loblolly (P. taeda L.), longleaf (P. palustris Miller),
pitch (P. rigida Miller), pond (P. serotina Michx.), sand (P. clausa
[Chapm. ex Engelm.] Sarg.), shortleaf (P. echinata Miller), slash
(P. elliottii Engelm.), and Virginia (P. virginiana Miller) pines.

Distribution The small southern pine engraver occurs from Pennsylvania to Florida
and west to Texas.
244 Forest Pest Insects in North America:

Images of Small
Southern Pine Engraver

Figure 1. Adult (left) of the small southern pine engraver, Ips avulsus, which is
the smallest of the eastern Ips species, as seen in relation to the other southern
pine bark beetle species (right). (Left: J.R. Baker and S.B. Bambara, North
Carolina State University, Bugwood.org; right: John L. Foltz, University of Florida,
Bugwood.org)

Figure 2. The depressed

region at the posterior
of the small southern
pine engraver is shallow,
with strong knobs on its
surface and four teeth
along each side. (J.R.
Baker and S.B. Bambara,
North Carolina State
University, Bugwood.org)

Figure 3. The pitch tube of the small southern pine engraver (center of photo,
small). (David J. Moorhead, University of Georgia, Bugwood.org)
A Photographic Guide 245

Images of Small Southern

Pine Engraver (continued)

Figure 4. A pair of small southern pine engraver beetles in their nuptial chamber,
showing associated oviposition galleries. (Erich G. Vallery, USDA Forest Service
- SRS-4552, Bugwood.org)

Figure 5. Short
galleries made by
larvae radiating out
from linear oviposition
galleries of the small
southern pine engraver.
(Jeffrey Eickwort,
Florida Department
of Agriculture and
Consumer Services,
Bugwood.org)

Figure 6. Galleries, pupae, and larvae of the small southern pine engraver.
(Ronald F. Billings, Texas Forest Service, Bugwood.org)
246 Forest Pest Insects in North America:

Images of Small Southern

Pine Engraver (continued)

Figure 7. Pupal cells of small southern pine engraver give a paddle-like

appearance to the larval galleries. (Robert L. Anderson, USDA Forest Service,
Bugwood.org)

Figure 8. Attacks of the

small southern pine
engraver on larger trees
are concentrated in
the tops of trees (here,
a loblolly pine [Pinus
taeda]). (Ronald F.
Billings, Texas Forest
Service, Bugwood.org)

Important Biological Control The natural enemies of the small southern pine engraver have not been
Agents Related to this reported but likely are similar to those of other Ips species.
Pest Species
A Photographic Guide 247

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/ips/ipsfidl.htm; a fact sheet on

on Small Southern southern Ips in general; gives list of host trees and map of range by Ips
Pine Engraver species.
http://edis.ifas.ufl.edu/in701; a University of Florida fact sheet on various
southern Ips species.

Articles Foltz, J.L., E.P. Merkel, and R.C. Wilkinson. 1984. Annotated
bibliography of Dendroctonus terebrans (Oliver), Ips avulsus (Eichhoff),
Ips grandicollis (Germar) in the southeastern USA. Monograph #12,
Agricultural Experiment Station, University of Florida: 47 p.
Birch, M.C., P. Svihra, T.D. Paine, and J.C. Miller. 1980. Influence
of chemically mediated behavior on host tree colonization by four
cohabiting species of bark beetles. Journal of Chemical Ecology 6:
395-414.
248 Forest Pest Insects in North America:

73. Silver Fir Beetle, Pseudohylesinus sericeus (Mannerheim) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The silver fir beetle, Pseudohylesinus sericeus (Mannerheim), is a minor
pest that attacks windthrown, felled, injured, and severely suppressed
trees. The damage it causes is rarely important. However, in the 1940s
and 1950s, large outbreaks occurred on Pacific silver fir (Abies amabilis
Douglas ex J.Forbes) in the Pacific Northwest of the United States. In
Oregon, there is one generation per year, but in Washington, a generation
requires two years. The egg gallery is transverse. In large trees, attacks
occur in the upper bole and limbs. The beetle often attacks trees that
are infected with brown-stain or root-rotting fungi such as Armillaria
sp., Heterobasidion annosum (Fr.) Bref., and Phellinus weirii (Murrill)
Gilbert.

Hosts Commonly Attacked The principal hosts of the silver fir beetle are various fir (Abies), western
hemlock (Tsuga heterophylla [Raf.] Sarg.), and Douglas-fir (Pseudotsuga
menziesii [Mirb.] Franco).

Distribution The silver fir beetle ranges principally from British Columbia to
California and eastward to northern Idaho. It also may occur in southeast
Alaska and western Montana.

Image of Figure 1. Adult of

Pseudohylesinus sp.
Silver Fir Beetle (Province of British
Columbia, Minstry of
Forests and Range,
Forest Practices Branch)
A Photographic Guide 249

Important Biological Control The natural enemies of silver fir beetle are unknown.
Agents Related to this
Pest Species
Web Links for Information http://www.for.gov.bc.ca/hfp/publications/00198/silver_fir_beetle.htm;
on Silver Fir Beetle fact sheet of the forestry department of the province of British Columbia,
Canada.
http://www.fs.fed.us/r6/nr/fid/fidls/fidl-60.pdf; USDA Forest Service pest
leaflet on silver fir beetle.

Articles Gara, R.I. 1982. Insect pests of true firs in the Pacific Northwest. In:
Oliver, C.D. and R.M. Kenady (eds.). Proceedings of the biology and
management of true fir in the Pacific Northwest. Seattle-Tacoma,
Washington, February 24-26, 1981. Contribution, Institute of Forest
Resources, University of Washington (45): 157-160.
250 Forest Pest Insects in North America:

74. Fir Engraver, Scolytus ventralis LeConte (Coleoptera: Curculionidae,

Scolytinae)

Orientation to Pest The fir engraver, Scolytus ventralis LeConte, is a pest of true firs in
western North America. Adults are about 4.0 mm long, making this
one of the largest species in the genus Scolytus. The egg galleries dug
by females are transverse to the trunk and galleries are deep enough to
deeply score the wood, especially close to the nuptial chamber. Eggs
are laid along the oviposition galleries, from which larvae dig lateral
galleries up or down the trunk after hatching. Pupation occurs in the
inner bark at the end of the larval galleries. Fir engraver attacks trees
of varying ages and transmits the brown-stain fungus Trichosporium
symbioticum Wright, which is essential for successful brood
development. Stressed trees that are infected with various fungi or that
have been damaged by outbreaks of other insects are likely to be attacked
by the fir engraver. Effects of this bark beetle vary, with some trees dying
quickly, while others are just top-killed or recover, even from repeated
attacks. There is one generation per year, or in some areas, only one in
two years. In intensively managed areas, prompt removal or treatment
of infested trees may help protect remaining trees for a time, but in most
areas control is best achieved by trying to lower sources of stress and
maintain high tree vigor.

Hosts Commonly Attacked The principal hosts of the fir engraver are various species of fir (Abies),
especially white (Abies concolor [Gordon] Lindley ex Hildebrand), grand
(A. grandis [Douglas ex D. Don] Lindley), and red fir (A. magnifica
A. Murray). Occasionally Douglas-fir (Pseudotsuga menziesii [Mirbel]
Franco), hemlock (Tsuga), or spruce (Picea) are attacked.

Distribution The fir engraver ranges from British Columbia to California, Wyoming,
Colorado, Arizona, and New Mexico.

Images of Figure 1. Adult of the

fir engraver, Scolytus
Fir Engraver ventralis. (Donald Owen,
California Department
of Forestry and Fire
Protection,
Bugwood.org)
A Photographic Guide 251

Images of Fir Engraver Figure 2. Pitch

streamers on trunk are
(continued) a sign of attack on fir by
the fir engraver. (Donald
Owen, California
Department of Forestry
and Fire Protection,
Bugwood.org)

Figure 3. Bubbled
bark is a sign of
previous fir engraver
attacks survived by a
tree. (Chris Schnepf,
University of Idaho,
Bugwood.org)
252 Forest Pest Insects in North America:

Images of Fir Engraver Figure 4. Galleries of the

fir engraver. Note the
(continued) horizontal oviposition
gallery, with multiple
larval galleries above
and below. (Dave
Holland, USDA Forest
Service, Bugwood.org)

Figure 5. White fir (Abies concolor) killed by the fir engraver. (William M. Ciesla,
Forest Health Management International, Bugwood.org)
A Photographic Guide 253

Images of Fir Engraver Figure 6. Top kill is

a sign of attack of fir
(continued) engraver on large,
otherwise healthy, trees.
(Kenneth E. Gibson,
USDA Forest Service,
Bugwood.org)

Figure 7. Damage at
the stand level from the
fir engraver. (A. Steven
Munson, USDA Forest
Service, Bugwood.org)

Important Biological Control The natural enemies of fir engraver are relatively unstudied.
Agents Related to this
Pest Species
Web Links for Information http://forestry.nv.gov/forestry-resources/forest-health/fir-engraver-beetle/;
on Fir Engraver Nevada Division of Forestry fact sheet.
www.oregon.gov/ODF/privateforests/docs/fh/FirEngraver.pdf?ga=t;
Oregon Department of Forestry website.
http://www.fs.fed.us/r6/nr/fid/fidls/fidl-13.pdf; USDA Forest Service
Forest Insect and Pest Leaflet on fir engraver.

Articles Berryman, A.A. 1973. Population dynamics of the fir engraver, Scolytus
ventralis (Coleoptera: Scolytidae). I. Analysis of population behavior
and survival from 1964 to 1971. The Canadian Entomologist 105:
1465-1488.
254 Forest Pest Insects in North America:

75. Columbian Timber Beetle, Corthylus columbianus Hopkins (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest Columbian timber beetle, Corthylus columbianus Hopkins, is a native

North American ambrosia beetle, one of a group of beetles that vector
wood-destroying fungi, which the beetle larva then eats. Unlike most
ambrosia beetles, Columbian timber beetles attack live trees rather than
dead, dying, or newly felled trees. Trees, however, do not die, since trees
are able to fill galleries with callus tissue. However, losses occur from
lowered quality of timbers cut from infested trees, due to the presence of
callus-filled galleries and fungal-stained wood.

Hosts Commonly Attacked Columbian timber beetles attack a wide range of hardwoods, including
various oaks (Quercus) and maples (Acer), sycamore (Platanus
occidentalis L.), poplar (Populus), elm (Ulmus), beech (Fagus), and
others.

Distribution The Columbian timber beetle ranges in the United States from
Massachusetts to Georgia, west to Michigan and Missouri.

Images of Columbian
Timber Beetle

Figure 1. Callow adult (left) and pupae of the Columbian timber beetle, Corthylus
columbianus. (Jack C. Nord, USDA Forest Service, Bugwood.org)
A Photographic Guide 255

Images of Columbian
Timber Beetle (continued)

Figure 2. Egg of Columbian timber beetle in chamber prepared by adult. Note

the ambrosia fungus (Ambrosiella eylebori Brader) coating the chamber, which
is inoculated into the gallery by the adult. (Jack C. Nord, USDA Forest Service,
Bugwood.org)

Figure 3. Galleries of Columbian timber beetle. (James Solomon, USDA Forest

Service, Bugwood.org)
256 Forest Pest Insects in North America:

Images of Columbian
Timber Beetle (continued)

Figure 4. Emergence holes and sap-soaked patches on trunk are signs of

Columbian timber beetles. (Left: Jack C. Nord, USDA Forest Service,
Bugwood.org; right: James Solomon, USDA Forest Service, Bugwood.org)

Figure 5. Milled maple timber shows the callus-filled galleries of Columbian

timber beetle, and blue staining of associated ambrosia fungus. (Left: Jack C.
Nord, USDA Forest Service, Bugwood.org; right: James Solomon, USDA Forest
Service, Bugwood.org)

Important Biological Control Natural enemies of Columbian timber beetle have not been reported.
Agents Related to this
Pest Species
Web Links for Information http://wiki.bugwood.org/Archive:South/Corthylus_columbianus;
on Columbian Timber Beetle Bugwiki fact sheet.

Articles Milne, D.H. and R.L. Giese. 1969. The Columbian timber beetle,
Corthylus columbianus (Coleoptera: Scolytidae). IX. Population biology
and gallery characteristics. Entomological News 80(9): 225-237.
A Photographic Guide 257

76. Ambrosia Beetles, Xyleborus spp. (Coleoptera: Curculionidae, Scolytinae)

Orientation to Pests Xyleborus ambrosia beetles are a genus of wood-boring beetles that carry
fungi that colonize their galleries and serve as food for their larvae. Other
genera of beetles that have similar symbiotic relationships with fungi
are also referred to as ambrosia beetles. The genus Xyleborus appears
to be a collection of poorly related beetles that is in need of revision. As
currently constructed, there are at least 17 species in the United States
and Canada (16 in the eastern United States), mostly native but with
several invasive species. Many of these species breed in both coniferous
and deciduous trees and shrubs of all sizes. Trees preferred for attack
are ones that are unhealthy, dying, or that have recently been wounded
or felled. Dead areas in living trees can also be attacked. Because this
group of beetles inoculate their larval galleries with wood-decay fungi,
which are eaten by their larvae, these beetles are able to make their
larval galleries in heartwood (a zone of low nutrition), in contrast to bark
beetles, which feed in the more nutritious sapwood. Notes on the biology
and distribution of four species of Xyleborus are given here. Many new
invasive species within this group have been detected recently in North
America. One invasive species, the redbay ambrosia beetle (Xyleborus
glabratus Eichhoff), is discussed separately elsewhere in this publication.

Species Comments on Biology and Gallery Shape

Xyleborus celsus Eichh. The largest member of the genus in the
United States, this species’ galleries
extend directly into the wood to a depth
of 12 to 18 mm, then branch one or
several times.
Xyleborus ferrugineus (F.) Galleries similar to X. celsus but smaller;
galleries may also be longer, more
winding, and branch less frequently.
Xyleborus affinis Eichh. Galleries consist of elongate tunnels,
off of which many transverse galleries
branch.
Xyleborus xylographus (Say) Attacks are commonly initiated at edges
of wounds, deep crevices, or burrows
made by other insects. Galleries run
obliquely across the grain to depths of
an inch or more.
258 Forest Pest Insects in North America:

Hosts Commonly Attacked Species Hosts

Xyleborus celsus Eichh. Breeds in dead, dying, and recently
felled trees and stumps of hickory.
Xyleborus ferrugineus (F.) Breeds in dead, dying, or felled trees
of a wide range of species including
oak, hickory, ash, cypress, walnut, pine,
beech, and sweetgum.
Xyleborus affinis Eichh. Breeds in dying trees and in green
logs and lumber of various hardwood
trees such as oak, hickory, sweetgum,
river birch, hackberry, mimosa,
persimmon, cypress, and black locust.
Very destructive of sweet gum in
southeastern United States along the
Gulf of Mexico.
Xyleborus xylographus (Say) Attacks the lower portions of dying
trees and the stumps, roots, and slash of
its hosts. Found especially in oak, but
also in hickory, chestnut, walnut, and
pine.

Distribution Species Distribution

Xyleborus celsus Eichh. Found in eastern United States, west to
Kansas.
Xyleborus ferrugineus (F.) Occurs from New York and Michigan,
south to Florida and Texas.
Xyleborus affinis Eichh. Occurs in eastern United States east
of a line from Michigan to Texas and
south of New York.
Xyleborus xylographus (Say) Occurs in eastern Canada and in the
United States, west to Kansas.
A Photographic Guide 259

Images of Xyleborus
Ambrosia Beetles

Figure 1. Adults of Xyleborus celsus (top), Xyleborus ferrugineus (middle), and

Xyleborus xylographus (bottom). (All photos: J.R. Baker and S.B. Bambara,
North Carolina State University, Bugwood.org)
260 Forest Pest Insects in North America:

Images of Xyleborus
Ambrosia Beetles
(continued)

Figure 2. Galleries of Xyleborus celsus in cross sections of hickory (Carya)

stems, showing intial deep pentration in the wood, with formation of branched
galleries in the heart wood. (James Solomon, USDA Forest Service,
Bugwood.org)

Important Biological Control Natural enemies of these beetles are poorly known.
Agents Related to this
Pest Species
Web Links for Information None
on Xyleborus
Ambrosia Beetles
Articles Atkinson, T.H., R.J. Rabaglia, and D.E. Bright. 1990. Newly detected
exotic species of Xyleborus (Scolytidae) in eastern North America, with a
revised key to species. The Canadian Entomologist 122: 93-104.
Haack, R.A. 2006. Exotic bark- and wood-boring Coleoptera in the
United States: recent establishments and interceptions. Canadian Journal
of Forest Research 36(2): 269-288.
A Photographic Guide 261

77. Redbay Ambrosia Beetle, Xyleborus glabratus Eichhoff (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The redbay ambrosia beetle, Xyleborus glabratus Eichhoff, was detected
in the United States for the first time in 2002 and has since caused
extensive mortality of redbay (Persea borbonia [L.] Spreng.) and
sassafras (Sassafras albidum [Nutt.] Nees) in coastal areas of South
Carolina, Georgia, and parts of Florida. This ambrosia beetle is the vector
of a lethal fungal pathogen (Raffaelea lauricola Harrington, Fraedrich
and Aghayeva) that affects various species in the Lauraceae. Infected
trees wilt and die within a few weeks or months of initial infection. This
disease has been designated as “laurel wilt,” and it is believed to have
arrived along with the redbay ambrosia beetle in wood packing material
(crates and pallets) used in trade. Xyleborus glabratus is about 2 mm
long, dark brown to almost black in color, nearly glabrous (shiny, without
hairs), and has an abruptly slanting depression at the tip of it abdomen.
However, positive confirmation of this species’ presence in a new area
requires confirmation by a taxonomic specialist. Very little is known
specifically about the biology of this beetle but it is likely similar to that
of other members of this genus (see other insects in this publication).
Adults and larvae inoculate new galleries with spores of a symbiotic
pathogenic fungus, which are carried in special pockets at the base of
the beetles’ mandibles. Males are flightless. Females can attack healthy
hosts, and, in Florida and Georgia, female beetles fly throughout the year
although at very low numbers in the winter. Signs of redbay ambrosia
beetle attack include small strings of compacted sawdust protruding from
the bark at the point of attack, but these strings soon disintegrate and
are not unique to this species. Small holes the diameter of a paperclip
wire in redbay are usually redbay ambrosia beetle entrance holes.
Foliage of infested trees wilts because the fungal infection is systemic
in the vascular elements of the tree, and foliage turns reddish brown. As
infected trees start to die, they may also be attacked by other species of
ambrosia beetle, particularly Xylosandrus crassiusculus (Motschulsky).
In Florida, up to 90 percent of redbay trees died in less than two years
following initial beetle invasion of a new site. Loss of redbay will
likely severely harm populations of the Palamedes swallowtail (Papilio
palamedes [Drury]), which feeds primarily on species of Persea. There
is also considerable concern about the potential impact of laurel wilt on
avocado (P. americana Miller), which is an economically important crop
in Florida and California.

Hosts Commonly Attacked In addition to redbay (P. borbonia) and sassafras (S. albidum), the wilt
pathogen spread by the redbay ambrosia beetle has been recovered in
the United States from avocado (P. americana), swampbay (P. palustris
[Raf.] Sarg.), pondspice (Litsea aestivalis [L.] Fernald), pondberry
(Lindera melissifolia [Walter] Blume) and camphor (Cinnamomum
camphora [L.] Sieb.). Pondspice and pondberry are of special concern
because they are rare or endangered species. Avocado is of concern
because of its economic importance.
262 Forest Pest Insects in North America:

Distribution The redbay beetle is native to South Asia (India, Myanmar) and East
Asia (Japan, Taiwan). The invaded range in the United States includes
coastal and adjacent areas of Georgia, South Carolina, and Florida, as
well as parts of North Carolina, one area in Mississippi, and two counties
in Alabama.

Figure 1. Map of the range of redbay ambrosia beetle, Xyleborus glabratus, in

the United States. (USDA Forest Service - Southern Region • Forest Health
Protection - Laurel Wilt)

Images of Figure 2. Adult of

the redbay ambrosia
Redbay Ambrosia Beetle beetle; bottom, insect in
natural position. (Top:
Michael C. Thomas,
Florida Department
of Agriculture and
Consumer Services,
Bugwood.org; bottom:
Shipher Wu)
A Photographic Guide 263

Images of Redbay Figure 3. Egg of redbay

ambrosia beetle, inside
Ambrosia Beetle gallery cut by female
(continued) beetle. (Karolynne
Griffiths, USDA APHIS
PPQ, Bugwood.org)

Figure 4. Larva of
redbay ambrosia beetle.
(Bettaman/Shipher at
[email protected])

Figure 5. Ambrosia
beetles produce
frass tubes at points
of entrance for adult
beetles during the
construction of their
galleries. The long
tubes are Xyleborus
crassiusculus, while
the very short tubes in
the debarked area are
redbay ambrosia beetle.
(Ronald F. Billings,
Texas Forest Service,
Bugwood.org)
264 Forest Pest Insects in North America:

Images of Redbay
Ambrosia Beetle
(continued)

Figure 6. Galleries of redbay beetle in redbay tree trunk, seen in cross section.
(James Johnson, Georgia Forestry Commission, Bugwood.org)

Figure 7. Dark staining

of the vascular tissue
is a sign of the fungus
associated with the
redbay ambrosia
beetle. (Both photos:
Albert [Bud] Mayfield,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 265

Images of Redbay
Ambrosia Beetle
(continued)

Figure 8. Dead redbay killed by redbay ambrosia beetle and its associated
fungus. The percentage of redbay mortality in this section of Florida forest
increased from 10 percent to 81 percent within a year. (Albert [Bud] Mayfield,
USDA Forest Service, Bugwood.org)

Important Biological Control Natural enemies of redbay ambrosia beetle are unknown at this time.
Agents Related to this
Pest Species
Web Links for Information http://www.freshfromflorida.com/pi/enpp/ento/x.glabratus.html; fact
on Redbay Ambrosia Beetle sheet of the Florida Department of Agriculture and Consumer Services,
Division of Plant Industry.
http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=148&langd
isplay=english; fact sheet of Exotic Forest Pest Information System for
North America.
http://www.invasivepests.org/products/gallery/xylgl1; fact sheet about
risks to trees of moving firewood.
http://www.redorbit.com/news/science/1685690/tiny_beetle_threatens_
floridas_avocado_industry/; redbay ambrosia beetle as a threat to
avocado industry in the United States.
http://www.invasive.org/browse/subinfo.cfm?sub=10998; fact sheet of
Bugwood Wiki.
http://www.public.iastate.edu/~tcharrin/LaurelWilt.html; factsheet on the
wilt pathogen and links to PDF versions of relevant articles.
266 Forest Pest Insects in North America:

Articles on Redbay Douce, G.K. and J. Johnson. 2005. Xyleborus glabratus in Georgia’s
Ambrosia Beetle coastal forests. Georgia Forestry Commission Pest Alert, October 31,
2005.
Hanula, J.L., A.E. Mayfield, III, S.W. Fraedrich, and R.J. Rabaglia. 2008.
Biology and host associations of redbay ambrosia beetle (Coleoptera:
Curculionidae: Scolytinae), exotic vector of laurel wilt killing redbay
trees in the southeastern United States. Journal of Economic Entomology
101: 1276-1286.
Fraedrich, S.W., T.C. Harrington, R.J. Rabaglia, M.D. Ulyshen, A.E.
Mayfield, III, J.L. Hanula, J.M. Eickwort, and D.R. Miller. 2008. A
fungal symbiont of the redbay ambrosia beetle causes a lethal wilt in
redbay and other Lauraceae in the southeastern United States. Plant
Disease 92: 215-224. (Available at http://www.public.iastate.edu/
~tcharrin/LaurelWilt.html).
Harrington, T.C., S.W. Fraedrich, and D.N. Aghayeva. 2008. Raffaelea
lauricola, a new ambrosia beetle symbiont and pathogen on the
Lauraceae. Mycotaxon 104: 399-404. (Available at http://www.public.
iastate.edu/~tcharrin/LaurelWilt.html).
Mayfield, A.E, T.C. Harrington, S. Fraedrich, J. Hanula, and others.
2009. Recovery plan for laurel wilt on redbay. In: Plant Diseases that
Threaten U.S. Agriculture. Prepared for the National Plant Disease
Recovery System, USDA and the American Phytopathological Society.
27 p. (Available at http://www.public.iastate.edu/~tcharrin/LaurelWilt.
html).
A Photographic Guide 267

78. Walnut Twig Beetle, Pityophthorus juglandis Blackman (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The walnut twig beetle, Pityophthorus juglandis Blackman, is the vector
of a newly recognized fungal species, Geosmithia morbida Kolarík,
Freeland, Utley, and Tisserat (Ascomycota: Hypocreales). This insect-
pathogen complex causes a fatal disease of eastern black walnut (Juglans
nigra L.) and certain other species of walnut (Juglans). The common
name for the disease is “thousand cankers” because death results from
the coalescing of a great many small, separate cankers that originate at
multiple points of twig beetle entry into twigs, branches, and main stems.
Walnut twig beetle, a type of bark beetle native to the southwestern
United States and parts of Mexico, is the only confirmed vector of the
pathogen. In its early stages this disease appears as circular-to-oblong
cankers that develop at the site of each walnut twig beetle gallery.
Over time, cankers expand in the phloem, coalesce, and girdle twigs
or branches. Geosmithia morbida does not move systemically within
the tree and the pathogen is moved between trees only by the insect
vector. Relatively large numbers of walnut twig beetles are needed to
cause enough cankers to kill trees. The origin of this newly recognized
pathogen is uncertain, but isolates from walnut species in the western
United States show high genetic diversity, suggesting the fungus may be
native to that region. Why the walnut twig beetle has recently increased
to levels able to unleash a wave of walnut mortality is unknown, but is
likely linked to its invasion of new parts of the country and attacks there
on new species of walnut, some of which are highly susceptible. On
black walnut in Colorado, the walnut twig beetle overwinters as adults in
cavities excavated in the bark. Beetles colonize new hosts by attacking
branches with rough bark, on the warmer side (exposed to sun). There
are two overlapping generations per year in Colorado. Mass attacks are
mediated by aggregation pheromones. Females dig oviposition galleries,
inoculate with the galleries with their Geosmithia fungus, and then lay
eggs along the gallery. Larvae emerge and dig tunnels, where they feed
on fungus-infected wood.

Hosts Commonly Attacked Trees that are highly susceptible to walnut twig beetle attack and fungal
infection in the western United States include the California walnut (J.
californica S. Wats.), Hinds walnut (J. hindsii [Jeps.] Jeps. ex R.E. Sm)
(both native to California), and black walnut (J. nigra) (native only to
the eastern United States). English walnut (J. regia L.) is moderately
susceptible, while Arizona walnut (J. major [Torr.] A. Heller)—believed
to be the original host for the beetle—appears to be quite tolerant. Two
other important members of the walnut family—butternut (J. cinerea
L.) and pecan (Carya illinoinensis [Wangenh.] K. Koch)—appear to be
resistant to the pathogen.
268 Forest Pest Insects in North America:

Distribution Walnut twig beetle and its associated pathogen are now reported from
eight states in the western USA and three in the east (Tennessee, Virginia,
and Pennsylvania). To what extent the western distribution reflects the
native range versus recent spread into new locations is not certain.

Figure 1. Distribution of the thousand cankers disease (plus, one eastern location
in Knoxville, Tennessee). (USDA APHIS)

Images of
Walnut Twig Beetle

Figure 2. Adult of the walnut twig beetle, Pityophthorus juglandis. (Steven Valley,
Oregon Department of Agriculture, Bugwood.org)
A Photographic Guide 269

Images of Walnut
Twig Beetle (continued)

Figure 3. Piles of boring dust at entrance of points of adult walnut twig beetle
attack. (Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 4. Adults of walnut twig beetles, in nuptial chamber. (Curtis Utley, CSUE,
Bugwood.org)

Figure 5. Egg gallery of walnut twig beetle. (Whitney Cranshaw, Colorado State
University, Bugwood.org)
270 Forest Pest Insects in North America:

Images of Walnut
Twig Beetle (continued)

Figure 6. Larvae of walnut twig beetle. (Whitney Cranshaw, Colorado State

University, Bugwood.org)

Figure 7. Galleries of walnut twig beetle in branch. (Whitney Cranshaw, Colorado

State University, Bugwood.org)

Figure 8. Larva and pupa of walnut twig beetle, with a view of its fungal
associate, Geosmithia morbida, which is sporulating profusely around the pupal
cells. (Curtis Utley, CSUE, Bugwood.org)
A Photographic Guide 271

Images of Walnut Figure 9. Weeping

wounds are signs of
Twig Beetle (continued) attack by walnut twig
beetle on English
walnut, Juglans regia.
Symptoms on black
walnut are different.
(Ned Tisserat, Colorado
State University,
Bugwood.org)

Figure 10. Emergence holes of walnut twig beetle. (Whitney Cranshaw, Colorado
State University, Bugwood.org)
272 Forest Pest Insects in North America:

Images of Walnut
Twig Beetle (continued)

Figure 11. Cankers that have developed around the galleries of walnut twig
beetle. (Ned Tisserat, Colorado State University, Bugwood.org)

Figure 12. Branch death and dieback are early symptoms of walnut twig beetle
attack (seen here on Arizona walnut). (Whitney Cranshaw, Colorado State
University, Bugwood.org)

Figure 13. Black walnut

trees dying or dead due
to thousand cankers
disease. (Curtis Utley,
CSUE, Bugwood.org)
A Photographic Guide 273

Important Biological Control Natural enemies of walnut twig beetle have not been reported.
Agents Related to this
Pest Species
Web Links for Information http://tn.gov/agriculture/publications/regulatory/tc_pathwayanalysis.
on Walnut Twig Beetle pdf; a Tennessee website where a USDA APHIS document is posted that
gives background information on vector and thousand cankers pathogen,
plus an analysis of likely means of further spread.
http://caforestpestcouncil.org/wp-content/uploads/2009/05/steven-
seybold-walnut.pdf; a Power Point presentation from a meeting that
covers various aspects of impact of walnut twig beetle in California.
http://caps.ceris.purdue.edu/webfm_send/854; provides a pictorial
key for identification of walnut twig beetle from its many con-generic
species.
http://www.invasivepests.org/products/gallery/pitju1.html.
http://www.michiganloggingcompany.com/Black-Walnut-Black-Death;
contains a video on aspects of the problem.
http://www.ext.colostate.edu/pubs/insect/0812_alert.pdf; fact sheet of the
Colorado State University.
http://www.fs.fed.us/foresthealth/fhm/sp/tcd/tcd.shtml; USDA Forest
Service page on thousand cankers disease and walnut twig beetle.

Articles Tisserat, N., W. Cranshaw, D. Leatherman, C. Utley, and K. Alexander.

2009. Black walnut mortality caused by the walnut twig beetle and
thousand cankers disease. Plant Health Progress. August 11, 2009. 10 p.
Grant, J.F., M.T. Windham, W.G. Haun, G.J. Wiggins, and P.L. Lambdin.
2011. Initial assessment of thousand cankers disease on black walnut,
Juglans nigra, in eastern Tennessee. Forests 2: 741-748.
274 Forest Pest Insects in North America:

79. Striped Ambrosia Beetle, Trypodendron lineatum (Olivier) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The striped ambrosia beetle, Trypodendron lineatum (Olivier), has long
been considered the most damaging ambrosia beetle in the western
United States (but see also the walnut twig borer, Pityophthorus
juglandis Blackman). It is particularly important in coastal areas of
British Columbia in Canada. Attacks are principally on dead and dying
timbers created by wind storms, fire, bark beetle attack, or long in-field
storage of cut logs. Females bore into the xylem and then larvae make
lateral galleries where they feed, pupate, and later emerge. There is
generally one generation per year.

Hosts Commonly Attacked Striped ambrosia beetles attack conifers in Abies, Picea, Pseudotsuga,
Tsuga, Larix, and Pinus. They rarely attack hardwoods.

Distribution The striped ambrosia beetle is Holarctic in distribution. In the western

United States, it is found from New Mexico and southern California to
the northern edge of the boreal forest in Canada. In the eastern United
States, it is found in New England and along the Appalachians to the
mountains of western North Carolina.

Images of
Striped Ambrosia Beetle

Figure 1. Adult of the striped ambrosia beetle, Trypodendron lineatum.

(Maja Jurc, University of Ljubljana, Bugwood.org)
A Photographic Guide 275

Images of Striped
Ambrosia Beetle
(continued)

Figure 2. Galleries of striped ambrosia beetle. Top left: galleries made by females
to access interior of log; bottom left: the up and down galleries off the adult’s
gallery made by larvae; right: diagram of the spatial relation of the two types of
galleries inside a log. (Top left: Wayne Brewer, Auburn University, Bugwood.org;
bottom left: Petr Kapitola, State Phytosanitary Administration, Bugwood.org;
right: Robert Dzwonkowski, Bugwood.org)

Figure 3. Emergence
holes (see sawdust) of
striped ambrosia beetles
in felled Norway spruce
(Picea abies [L.] Karst.).
(Both photos:
Beat Forster, Swiss
Federal Institute for
Forest, Snow and
Landscape Research,
Bugwood.org)
276 Forest Pest Insects in North America:

Images of Striped Figure 4. Emergence

holes of striped
Ambrosia Beetle ambrosia beetles. (Milan
(continued) Zubrik, Forest Research
Institute - Slovakia,
Bugwood.org)

Figure 5. Thanasimus formicarius, a clerid predator that likely feeds on

striped ambrosia beetle. (Scott Bauer, USDA Agricultural Research Service,
Bugwood.org)
A Photographic Guide 277

Important Biological Control Little is known of the natural enemies of this ambrosia beetle. The clerid
Agents Related to this Thanasimus formicarius (L.) responds to the pheromone of T. lineatum,
Pest Species which suggests that it may use the striped ambrosia beetle as food.

Web Links for Information http://www.forestry.ubc.ca/fetch21/FRST308/lab7/trypodendron_

on Striped Ambrosia Beetle lineatum/striped.html; fact sheet of the University of British Columbia,
which includes a short video of damage from this beetle to stored logs.
http://www.barkbeetles.org/biocontol/stripedambrosiabeetle.html; fact
sheet from Bugwood Wiki on this beetle.

Articles Lindgren, B.S., S.E.R. Hoover, A.M. MacIsaac, C.I. Keeling, and K.N.
Slessor. 2000. Lineatin enantiomer preference, flight periods, and effect
of pheromone concentration and trap length on three sympatric species
of Trypodendron (Coleoptera: Scolytidae). The Canadian Entomologist
132: 877-887.
Park, J. and M.L. Reid. 2007. Distribution of a bark beetle, Trypodendron
lineatum, in a harvested landscape. Forest Ecology and Management
242(2/3): 236-242.
278 Forest Pest Insects in North America:

80. Black Twig Borer, Xylosandrus compactus (Eichhoff) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The black twig borer, Xylosandrus compactus (Eichhoff), is an invasive
ambrosia beetle from Asia present in Hawaii and coastal regions of the
southeastern United States. Females bore into healthy twigs of over 200
hardwood species, where they lay loose clusters of eggs and inoculate
their galleries with a symbiotic fungus (Fusarium solani [Mart.] Sacc.)
on which their larvae later feed. Twigs may be colonized by one or
several females, depending on twig size. Mature larvae pupae in the
gallery and males, which are flightless, mate with females in the twig
before new females emerge. Males never leave the twig. Attacked trees
or other plants are usually not killed but may suffer considerable damage.
In Hawaii, this species is an important pest of coffee (Coffea canephora
Pierre [esp. var. robusta Ineac]).

Table 1. Characteristics of 4 Xylosandrus species present in the

southeastern United States (from University of Florida fact sheet on
black twig borer seen at http://entnemdept.ufl.edu/creatures/trees/black_
twig_borer.htm)

Characteristic X. compactus X. crassiusculus X. germanus X. zimmermanni

Female size 1.4-1.9 mm 2.1-2.9 mm 2.0-2.4 mm 1.3-1.5 mm

Surface of Shining and Dull and Shining Shining

declivity smooth granulate

Hair tuft Forms — — Tuft oriented

on base of transverse row longitudinally
pronotum

Geographic Throughout Throughout From Subtropical

location Florida and Florida and from Connecticut to South Florida
from North North Carolina Missouri, east and Mexico to
Carolina to to eastern Texas Texas, and Venezuela.
eastern Texas central Georgia.
Might appear in
North Florida

Common host In small twigs In wood of large In wood of large Only in

material on healthy, cut twigs, small twigs, small unhealthy,
and stressed branches and branches and cut, or broken
plants stems stems branches 1-3
cm diameter
A Photographic Guide 279

Hosts Commonly Attacked Hosts are quite varied but include maple (Acer spp.), hickory (Carya
spp.), magnolia (Magnolia spp.), dogwood (Cornus spp.), oaks (Quercus
spp.), and willows (Salix spp.), among others.

Distribution Black twig borer has been reported from Hawaii and in North America
from coastal parts of North Carolina to Texas (USA). Globally this
species has a wide distribution in tropical and subtropical areas.

Images of
Black Twig Borer

Figure 1. Adult of black twig beetle, Xylosandrus compactus; right, adult on

coffee. (Left: J.R. Baker and S.B. Bambara, North Carolina State University,
Bugwood.org; right: Scot Nelson, University of Hawaii at Manoa, Bugwood.org)

Figure 2. Entrance and exit holes of adult black twig beetles in a branch of a koa
tree (Acacia koa Gray) in Hawaii. (Scot Nelson, University of Hawaii at Manoa,
Bugwood.org)
280 Forest Pest Insects in North America:

Images of Black Figure 3. Galleries of

black twig beetle larvae
Twig Borer (continued) in magnolia. (Forrest L.
Oliveria, USDA Forest
Service, Bugwood.org)

Figure 4. Cross-sectional views of galleries of Xylosandrus germanus (Blandford)

in pecan. (Jerry A. Payne, USDA Agricultural Research Service, Bugwood.org)
A Photographic Guide 281

Images of Black Figure 5. “Toothpick”

frass ejected from bore
Twig Borer (continued)
hole of Xylosandrus
crassiusculus (Eichhoff).
(G. Keith Douce,
University of Georgia,
Bugwood.org)

Figure 6. Death of small cypress branch (left) caused by black twig borer; injury
to Acacia koa branch (right). (Left: Andrew J. Boone, South Carolina Forestry
Commission, Bugwood.org; right: Scot Nelson, University of Hawaii at Manoa,
Bugwood.org)

Important Biological Control Little is known of the natural enemies of black twig borer.
Agents Related to this
Pest Species
282 Forest Pest Insects in North America:

Web Links for Information http://www.extento.hawaii.edu/kbase/crop/type/xylosand.htm; fact sheet

on Black Twig Beetle of the University of Hawaii Extension Service.
http://entnemdept.ufl.edu/creatures/trees/black_twig_borer.htm; fact
sheet of University of Florida.

Articles Ngoan, N.D., R.C. Wilkinson, D.E. Short, C.S. Moses, and J.R.
Mangold. 1976. Biology of an introduced ambrosia beetle, Xylosandrus
compactus, in Florida. Annals of the Entomological Society of America
69: 872-876.
Masuya, H. 2007. Note on the dieback of Cornus florida caused by
Xylosandrus compactus. Bulletin of the Forestry and Forest Products
Research Institute, Ibaraki 402: 59-63.
A Photographic Guide 283

81. Larger Pine Shoot Beetle, Tomicus piniperda (L.) (Coleoptera:

Curculionidae, Scolytinae)

Orientation to Pest The larger pine shoot beetle, Tomicus piniperda (L.), is an invasive
species in North America that is found throughout much of Europe, as
well as parts of North Africa (Algeria, Canary Islands) and Asia (China,
Japan, Korea, and Turkey). Both larvae and adults are damaging to a
wide variety of pines. Larvae develop under bark of branches and trunks
of suppressed or recently fallen trees, as well as in recently cut stumps.
Adults cause more important damage by tunneling in live shoots and
leaders of healthy pine trees, reducing growth and deforming tree shape,
which can cause important losses in Christmas tree production. Larger
pine shoot beetles overwinter as adults in short galleries in the outer
bark of host trees near the ground. Adults emerge in late winter or early
spring and construct brood galleries in suitable host material. At times,
these parent adults (P1) will emerge, shoot feed, and then construct
another brood gallery. The brood adults (F1) emerge in summer and fly
to the shoots of pine trees where they feed until fall. A single brood adult
(F1 generation) will feed in multiple shoots during the summer months.
Brood adults do not become sexually mature until they overwinter.

The larger pine shoot beetle is also believed to vector or be associated

with several fungi that are pathogens in conifers, including Ophiostoma
minus (Hedgc.) Syd. et P. Syd., and various blue-staining fungi such as
species of Leptographium. Outbreaks of this pest are triggered when
there is a sudden increase in the amount of breeding material available
in early spring, such as following severe winter storms, winter logging
operations, or Christmas tree plantations with recently cut stumps or
piles of unsold trees. Silvicultural practices during fall and winter such as
reducing the amounts of pine logging slash left in the forest and cutting
stumps low to the ground will limit number of breeding sites available
for this bark beetle.

Hosts Commonly Attacked The preferred host in Europe and so far in North America is Scots pine
(Pinus sylvestris L.), but other native pines such as red pine (Pinus
resinosa Sol. ex Aiton) are commonly infested. However, all native
North American pines are potential hosts.
284 Forest Pest Insects in North America:

Distribution Native to Eurasia, this species is invasive in parts of the United States,
from Wisconsin to Maine and south to West Virginia.

Figure 1. Distribution of the larger pine shoot beetle as of 2005. (National

Agricultural Pest Information System)

Images of Larger
Pine Shoot Beetle

Figure 2. Adult of larger shoot beetle, Tomicus piniperda. (Pest and Diseases
Image Library, Bugwood.org)
A Photographic Guide 285

Images of Larger Pine Figure 3. Diagnostic

characters for
Shoot Beetle (continued) recognition of larger
pine shoot beetle: (a)
absence of setae in the
second row (counting
from the midline),
toward the rear of
the beetle (top) and
(b) a six-segmented
funicle preceeding the
anntennal club (bottom).
(Top: Pest and
Diseases Image Library,
Bugwood.org; bottom:
Eric Allen - Canadian
Forest Service, Pacific
Forestry Centre,
Victoria, British
Columbia)

Figure 4. Seasonal life history of the larger shoot beetle in the Great Lakes region
of the United States. (E. Richard Hoebeke, Cornell University, Bugwood.org)
286 Forest Pest Insects in North America:

Images of Larger Pine

Shoot Beetle (continued)

Figure 5. Adults of larger shoot beetle, showing how adults bore into shoots
causing then to die and fall off. Top, adult boring into shoot; bottom left, adult
inside feeding tunnel in shoot; bottom right, opening in shoot made by adult
beetle to enter feeding site. (Top: Steve Passoa, USDA APHIS PPQ,
Bugwood.org; bottom left and right: Gyorgy Csoka, Hungary Forest Research
Institute, Bugwood.org)

Figure 6. Pitch tubes of larger shoot beetle on trunk. (Petr Kapitola, State
Phytosanitary Administration, Bugwood.org)
A Photographic Guide 287

Images of Larger Pine

Shoot Beetle (continued)

Figure 7. Galleries of the larger pine shoot beetle; left, see the deeply etched
vertical oviposition galleries and right, larval galleries radiating out horizontally
from the vertical oviposition gallery. (Left: Hannes Lemme, Bugwood.org; right:
Milan Zubrik, Forest Research Institute - Slovakia, Bugwood.org)

Figure 8. Blue stain damage in logs affected by the larger shoot beetle. (USDA
Forest Service - Northeastern Area Archive, USDA Forest Service, Bugwood.org)
288 Forest Pest Insects in North America:

Images of Larger Pine

Shoot Beetle (continued)

Figure 9. Exit holes of the larger shoot beetle. (Milan Zubrik, Forest Research
Institute - Slovakia, Bugwood.org)

Figure 10. Tops of pines (left) showing loss of branches from feeding of adults of
larger shoot beetle; fallen branches (right) caused by adult feeding. (Left:
E. Richard Hoebeke, Cornell University, Bugwood.org; right: Hannes Lemme,
Bugwood.org)

Figure 11. An important predator in Europe of the engraver beetle, the red-
bellied clerid beetle, Thanasimus formicarius (L.) (left, adult; right, larva). (Left:
Scott Bauer, USDA Agricultural Research Service, Bugwood.org; right: Jiri Hulcr,
Michigan State University, Bugwood.org)
A Photographic Guide 289

Important Biological Control A variety of natural enemies have been recorded attacking the larger
Agents Related to this pine shoot beetle in Europe, especially the clerid predator Thanasimus
Pest Species formicarius (L.) and the pteromalid parasitoid Rhopalicus suspensus
Ratz., both of which at some times and places have been observed to kill
70-80 percent of the larger shoot beetles’ larvae or pupae.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/pest_al/shootbeetle/shootbeetle.htm;

on Larger Pine Shoot Beetle USDA Forest Service pest leaflet.
http://warehouse.pfc.forestry.ca/pfc/5134.pdf; fact sheet of the Canadian
Forest Service, British Columbia.
http://entnemdept.ufl.edu/creatures/trees/beetles/pine_shoot_beetle.htm;
fact sheet of the University of Florida.
http://en.wikipedia.org/wiki/Tomicus_piniperda; page from Wikipedia.
http://www.na.fs.fed.us/spfo/pubs/pest_al/shootbeetle/shootbeetle.htm;
pest alert of the USDA Forest Service, Northeastern Area.
http://www.issg.org/database/species/ecology.asp?si=1200; fact sheet
of the Global Invasive Species Database, with a discussion of routes of
introduction of this species to new areas.

Articles Morgan, R.E., P. de Groot, and S.M. Smith. 2004. Susceptibility of

pine plantations to attack by the pine shoot beetle (Tomicus piniperda)
in southern Ontario. Canadian Journal of Forest Research 34(12):
2528-2540.
Kennedy, A.A. and D.G. McCullough. 2002. Phenology of the larger
European pine shoot beetle Tomicus piniperda (L.) (Coleoptera:
Scolytidae) in relation to native bark beetles and natural enemies in pine
stands. Environmental Entomology 31: 261-272.
290 Forest Pest Insects in North America:

82. White Pine Cone Beetle, Conophthorus coniperda (Schwarz)

(Coleoptera: Curculionidae, Scolytinae)

Orientation to Pest The white pine cone beetle, Conophthorus coniperda (Schwarz), infests
cones of eastern white pine (Pinus strobus L.) and is an important pest
of seed production in natural stands and in seed orchards. Cones are
initially attacked by adult female beetles that start tunneling in the cone
stalk or base. This severs the conductive tissue that connects the twig
and cone, killing the cone. A small pitch tube generally marks the entry
point. Individual female beetles extend an egg gallery down the central
axis of a cone. Eggs are laid along this gallery. Larvae feed on seeds
and associated tissues until they reach maturity. Larvae then pupate, and
later emerge as adults, which darken over a few days. Development from
egg to adult occurs in one growing season, and there is one generation
per year. Female beetles can attack and oviposit in more than one cone.
Infested cones stop growing in length after they are attacked. So, cones
attacked in the early spring will be very small when they die, while
cones attacked late in the summer are much larger in size. Infested
cones shrivel, turn brown, and become hard. Because stalks of infested
cones are severed, they can easily be removed by hand from a branch.
Uninfested cones or cones attacked by other seed and cone insects, in
contrast, adhere tightly to branches and may require a hand-clipper to
remove. Infested cones eventually fall from trees, and beetles spend
the winter within infested cones on the forest floor. In early spring,
beetles chew emergence holes and fly from cones on the ground into tree
crowns where they locate new host material. This is the most important
pest of white pine seed orchards, sometimes destroying most or all of
a developing seed crop. Damage is concentrated on second year cones,
but first year conelets are also attacked. Beetles, when they are on the
ground, can be killed using a low-intensity ground fire, a practice that is
employed for control in seed orchards.

Hosts Commonly Attacked Eastern white pine (P. strobus) is the only host of this species, but there
are other species of Conophthorus associated with other pines.

Distribution The white pine cone beetle is found throughout the range of eastern
white pine, from eastern Canada (Quebec, Ontario, Nova Scotia), south
to North Carolina, and west to Minnesota.
A Photographic Guide 291

Images of White
Pine Cone Beetle

Figure 1. Adult white pine cone beetle, Conophthorus coniperda. (J.R. Baker and
S.B. Bambara, North Carolina State University, Bugwood.org)

Figure 2. A female beetle can be seen initiating an attack at the base of a white
pine cone where the cone stalk enters the cone. (Larry R. Barber, USDA Forest
Service, Bugwood.org)
292 Forest Pest Insects in North America:

Images of White Pine Figure 3. Egg-laying

gallery of a cone beetle
Cone Beetle (continued) (Conophthorus sp.) in
the center of a cone
that has been cut in
half. (Steven Katovich,
USDA Forest Service,
Bugwood.org)

Figure 4. Damage
inside a white pine cone
caused by feeding of
larvae of white pine
cone beetle. (Robert L.
Anderson, USDA Forest
Service, Bugwood.org)

Figure 5. Dead white

pine cones, infested
by larvae of white pine
cone beetle may hang
on trees for a period of
time before dropping
to ground. (Steven
Katovich, USDA Forest
Service, Bugwood.org)
A Photographic Guide 293

Images of White Pine

Cone Beetle (continued)

Figure 6. The length of dead infested cones can vary greatly. Small dead cones
are those attacked in early spring and larger dead cones are those attacked in
late summer. (Steven Katovich, USDA Forest Service, Bugwood.org)

Figure 7. Low intensity burns can be used in spring to control adults of white pine
cone beetle before they emerge from cones on the ground. (Larry R. Barber,
USDA Forest Service, Bugwood.org)

Important Biological Control Natural enemies of this insect have not been reported.
Agents Related to this
Pest Species
Web Links for Information None
on White Pine Cone Beetle
294 Forest Pest Insects in North America:

Articles on White Wade, D.D., G.L. Debarr, L.R. Barber, and E. Manchester. 1989.
Pine Cone Beetle Prescribed fire - a cost effective control for white pine cone beetle. In:
MacIver, D.C., H. Auld, and R. Whitewood (eds). Proceedings of the
10th Conference on Fire and Forest Meteorology. Forestry Canada,
Petawawa National Forestry Institute (PNFI), Chalk River, Ontario:
117-121.
De Groot, P. 1990. The taxonomy, life history and control of
Conophthorus (Coleoptera: Scolytidae) in eastern North America.
Proceedings - Cone and seed pest workshop, 4 October 1989.
Information Report N-X-274, Forestry Canada, Newfoundland and
Labrador Region, St. John’s, Newfoundland, Canada: 37-46.
Trudel, R., C. Guertin, and P. De Groot. 2004. Use of pityol to reduce
damage by the white pine cone beetle, Conophthorus coniperda
(Col., Scolytidae) in seed orchards. Journal of Applied Entomology
128: 403-406.
A Photographic Guide 295

83. Lodgepole Cone Beetle, Conophthorus ponderosae Hopkins

(Coleoptera: Curculionidae, Scolytinae)

Orientation to Pest Western cone beetles (Conophthorus spp.) were historically described
based on their principal host trees. The sugar pine cone beetle
(Conophthorus lambertianae Hopkins) was described from Pinus
lambertiana Douglas, but it was later declared a junior synonym of
Conophthorus ponderosae Hopkins, which was formerly known as the
ponderosa cone beetle, but is now referred to as the lodgepole cone
beetle (Wood 1977). Later work based on similarities among beetle
cuticular hydrocarbons (Page et al. 1990), however, suggested that the
sugar pine-feeding population of C. ponderosae may indeed be a separate
species (the former C. lambertianae), but it was not formerly reinstated
taxonomically. The possible validity of a sugar pine-feeding species is
consistent with recent DNA studies that shown that there likely have
been more speciation events in this genus than were formerly suspected.
But these appear to have been driven by geographic isolation, not affinity
to particular host species (Cognato et al., 2005), although sometimes
the two go together. The ponderosa cone beetle (C. ponderosae in the
wide sense), as discussed here, is composed of several populations as
noted above, whose relatedness is not certain. The population associated
with ponderosa (Pinus ponderosa Douglas ex C. Lawson), lodgepole
(P. contorta var. latifolia Douglas), and Jeffrey (P. jeffreyi Balf.) pines is
found from Washington to California, and in Arizona and New Mexico.
The population associated with sugar pine (P. lambertiana Douglas) is
found in California, Nevada and Oregon, where it feeds mostly on
P. lambertiana and sometimes on western white pine (P. monticola
Douglas ex D. Don).

For purposes of this presentation, we discuss C. ponderosae in the wide

sense, inclusive of C. lambertianae. Regardless of their specie-level
status, the biologies of these populations are very similar. In broad
terms, their biology is similar to that of the white pine cone beetle
(Conophthorus coniperda [Schwarz]) in the eastern United States. Adults
bore into the pedicles of immature, second year pine cones in the spring
and dig a tunnel along the length of the cone, where the female deposits
her eggs. Larvae feed on the scales, seeds, and tissues of the cone.
Pupation occurs within the cone and adults overwinter on the soil in
fallen cones. For the sugar-pine feeding population (“C. lambertianae”),
adults emerge from cones in the fall and move to twigs, boring into them
to feed, where they remain over winter.
296 Forest Pest Insects in North America:

Orientation to Pest The C. ponderosae population associated with sugar pine is the most
(continued) destructive of the western Conophthorus cone beetles. It sometimes
destroys up to 75 percent of sugar pine seeds in some years, making it an
important pest in sugar pine seed orchards. Similarly, the C. ponderosae
population associated with western white pine in British Columbia,
Washington, and Idaho has caused important losses in seed collection
stands and seeds orchards of that species. The other populations of
C. ponderosae have similar effects on other species of pines in seed
orchards, but the typical rate of seed destruction is lower.

Hosts Commonly Attacked The lodgepole cone beetle, in the wide sense, attacks ponderosa
(P. ponderosa), lodgepole (P. contorta var. latifolia), Jeffrey (P. jeffreyi),
sugar (P. lambertiana), and western white (P. monticola) pines.

Distribution The lodgepole cone beetle, in the wide sense, is found from British
Columbia to California, as well as in Idaho, Nevada, Arizona, and
New Mexico.

Images of
Lodgepole Cone Beetle

Figure 1. Adult of lodgepole cone beetle (Conophthorus ponderosae).

(D. Manastyrski, Bugwood.org)

Figure 2. Adult cone

beetles (Conophthorus
sp.) tunnel in pine cones
and their larvae feed on
the immature seeds and
other tissues. (Steven
Katovich, USDA Forest
Service, Bugwood.org)
A Photographic Guide 297

Images of Lodgepole
Cone Beetle (continued)

Figure 3. Pitch tube of Conophthorus ponderosae at base of cones in western

white pine (P. monticola) at site of adult entrance. (Ward Strong, BC Ministry of
Forests, Bugwood.org)

Figure 4. Internal
appearance of pine
cones infested by
larvae of lodgepole
cone beetles. (USDA
Forest Service Archive,
USDA Forest Service,
Bugwood.org)
298 Forest Pest Insects in North America:

Images of Lodgepole
Cone Beetle (continued)

Figure 5. Late instar Conophthorus ponderosae larva exposed in a dissected

western white pine (P. monticola) cone. (Canadian Forest Service Archive,
Canadian Forest Service, Bugwood.org)

Figure 6. Reduced
size of cones of
ponderosa pine (top)
and western white pine
(bottom) infested by
the lodgepole cone
beetle compared to
the larger, normal
cones. (Top: Sandy
Kegley, USDA Forest
Service, Bugwood.org;
bottom: Ward Strong,
BC Ministry of Forests,
Bugwood.org)
A Photographic Guide 299

Important Biological Control Little is known of the natural enemies of populations of lodgepole
Agents Related to this cone beetle. But larvae of the population found on sugar pine
Pest Species (“C. lambertianae”) are attacked in the cone by the bethylid wasp
Cephalonomia utahensis Brues. Adult beetles are attacked by both the
predatory clerid Enoclerus lecontei Wolcott and are parasitized by the
pteromalid wasp Tomicobia tibialis Ashmead.

Web Links for Information http://www.barkbeetles.org/biocontol/sugarpineconebeetle.html; fact

on Lodgepole Cone Beetle sheet from University of California with an emphasis on potential for
biological control of this pest.
http://www.fgcouncil.bc.ca/PM-Factsheet11-Conophthorus-ponderosae.
pdf; fact sheet from the provincial forest council of British Columbia,
Canada.

Articles Wood, S.L. 1977. New synonymy and new species of American bark
beetles (Coleoptera: Scolytidae), part V. Great Basin Naturalist 37:
383-394.
Page, M., L.J. Nelson, M.I. Haverty, and G.J. Blomquist. 1990.
Cuticular hydrocarbons of eight species of North American cone beetles,
Conophthorus Hopkins. Journal of Chemical Ecology 16: 1173-1198.
Furniss, M.M. 1997. Conophthorus ponderosae (Coleoptera: Scolytidae)
infesting lodgepole pine cones in Idaho. Environmental Entomology 26:
855-858.
Bennett, R. 2000. Management of cone beetles (Conophthorus
ponderosae, Scolytidae) in blister rust resistant western white pine seed
orchards in British Columbia. Seed and Seedlings Extension Topics
12: 16-18. (Available at http://www.for.gov.bc.ca/hti/publications/
newsletters/Vol1201Aug20.pdf).
Cognato, A.I., N.E. Gillette, R. Campos Bolaños, and F.A.H. Sperling.
2005. Mitochondrial phylogeny of pine cone beetles (Scolytinae,
Conophthorus) and their affiliation with geographic area and host.
Molecular Phylogenetics and Evolution 36: 494-508.
300 Forest Pest Insects in North America:

85. May and June Beetles, Phyllophaga spp. (Coleoptera: Scarabaeidae)

Orientation to Pest May and June beetles (Phyllophaga spp.) are a group of scarab beetles
whose larvae feed on the roots of trees and other plants, which can
significantly affect survival of seedlings or young trees in plantations.
This is a very large group of beetles native to North America. There
are more than 250 Phyllophaga species in North America. Most U.S.
species occur in the eastern half of the country. In the western part of the
country, this group is most numerous in Arizona. Many species occur
in Mexico. Here, we present a general view of the biology of the group
and mention a few species that are sometimes forestry pests. Adults of
several important species emerge in spring and early summer (May and
June, especially, hence their name). Adults are large, often brown, beetles
that are active in evening or early night and are big enough to be easily
noticed (1-3 cm long). Eggs are laid in groups in the soil, about 8-18 cm
deep, with each egg enclosed in a ball of dirt. Newly hatched larvae feed
on organic matter in the soil but soon switch to feeding on rootlets of tree
seedlings, grasses, or other plants. Winter is spent in the soil and larvae
move up and down to stay below the frost line. Mature larvae pupate in
the soil. One to three years are required to reach maturity in the southern
states, 2-3 years in mid-latitudes of the United States, and 3-4 years in
northern states and Canada. Because of the multi-year life cycle and large
number of species, relatively little is known of the population dynamics
of particular species.

Damage from Phyllophaga species can occur from either adults or

larvae. Adults feed at night on the foliage of many plants including
broad-leaved trees, shrubs, and some conifers. Damage from adults is
spotty, but when populations are high trees can be heavily defoliated in
woodlots or forest edges. Larvae, called white grubs (a name applied
to the whole scarab family), feed in the soil on roots of both grasses
and woody plants. Damage from Phyllophaga grubs occurs in forest
nurseries and plantations the southern and eastern United States. In the
western United States, damage from Phyllophaga grubs is less common
but is occasionally significant to coniferous seedlings in plantations
and nurseries on newly broken sandy land. In the eastern United States,
damage to forest nurseries and young plantations can be severe at times.
In Michigan and Wisconsin in red pine plantations on sandy soils, as few
as 0.75 grubs per cubic foot of soil were found to kill 15-45 percent of
red pine seedlings.
A Photographic Guide 301

Hosts Commonly Attacked

Common Species Comments on Biology (from Drooz 1985)
and Distribution of Some
Phyllophaga Species Phyllophaga crenulata P. crenulata occurs throughout the eastern United
(Froclich) States. Adults feed on a wide variety of hardwoods,
especially persimmon, hickory, basswood, willow,
birch, and buckeye. The larvae are often serious pests
in coniferous nurseries in the Lake States.

Phyllophaga drakei P. drakei occurs throughout most of the eastern United

(Kirby) States and southern Canada. Adults feed on the
leaves of beech, birch, dogwood, maple, basswood,
elm, and willow. The larvae are important pests in
forest nurseries and plantations in the Lake States and
Canada.

Phyllophaga forsteri P. forsteri occurs throughout the eastern United

(Burmeister) States. Adults feed on a wide variety of hardwoods
such as beech, birch, elm, magnolia, maple, tupelo,
walnut, and willow. There are also reports of their
feeding on pine. The larvae are often destructive in
nurseries in the southern United States.

Phyllophaga implicita P. implicita occurs mostly in the Mississippi and Ohio

(Horn) River Valleys. Adults feed on beech, dogwood, elm,
sycamore, tupelo, walnut, willow, basswood, maple,
and other plants. The larvae killed millions of seedlings
in nurseries in Iowa in the 1930s.

Phyllophaga luctuosa P. luctuosa occurs primarily along the Atlantic and Gulf
(Horn) coasts in sandy, oak-pine regions, but also farther
north and inland to Tennessee, Oklahoma, and Iowa.
Adults feed on persimmon, mulberry, tupelo, walnut,
willow, beech, birch, and loblolly and longleaf pines.
The larvae are often destructive in nurseries and,
probably, plantations.

Phyllophaga P. prununculina occurs in the South Atlantic and Gulf

prununculina Coast States. It is especially common in the Sand Hills
(Burmeister) of South Carolina. Adults feed on pines, especially
loblolly and longleaf, and sometimes oaks and
persimmon. The larvae have caused serious losses in
pine nurseries and plantations in South Carolina.

Phyllophaga prunina P. prunina occurs throughout the central United

(LeConte) States east of the Rocky Mountains. Adults feed on
hardwoods, including beech, elm, walnut, basswood,
and willow. Feeding on pine has also been observed.
Larvae are injurious in nurseries in the Lake states.

Phyllophaga rugosa P. rugosa occurs mostly in the northern United States

(Melsheimer) and southern Canada. Adults feed on a wide variety
of hardwoods. The larvae are often destructive in
coniferous nurseries in the Lake States.

Phyllophaga tristis (F.) P. tristis occurs throughout the eastern United States
and in southern Canada. Adults prefer the foliage of
oaks but also feed on maple, persimmon, hickory,
elm, and willow. Larvae have caused serious losses in
nurseries in the Lake States.
302 Forest Pest Insects in North America:

Images of May Figure 1. Adults of two

species of Phyllophaga
and June Beetles beetles, showing
general body form
shared by most species
in genus. (Top:
Steven Katovich,
USDA Forest Service,
Bugwood.org; bottom:
Whitney Cranshaw,
Colorado State
University,
Bugwood.org)

Figure 2. Species identification of larval scarabs (white grubs) is largely based

on the setal patterns on the tip of the abdomen (the raster) (here a white grub in
another genus, the oriental beetle Anomala orientalis [Waterhouse]). (H. Sawada,
Bugwood.org)
A Photographic Guide 303

Images of May and Figure 3. Diagram of life

cycle of Phyllophaga
June Beetles (continued) implicita. (North Dakota
State University
Extension Service)

Figure 4. The C-shape

form of a “white grub”
and visible legs are
typical of Phyllophaga
larvae. (Clemson
University - USDA
Cooperative Extension
Slide Series,
Bugwood.org)

Figure 5. Damage
from adult Phyllophaga
species consists of
defoliation, usually
on hardwoods, but
sometimes on conifers.
(Top: Darren Blackford,
USDA Forest Service,
Bugwood.org; bottom:
Whitney Cranshaw,
Colorado State
University,
Bugwood.org)
304 Forest Pest Insects in North America:

Images of May and

June Beetles (continued)

Figure 6. The most important damage from Phyllophaga species as forest pests
is death of seedlings or small trees whose roots are eaten by larvae in the soil.
Here, pine seedings whose roots have been eaten by Phyllophaga larvae.
(W.H. Bennett, USDA Forest Service, Bugwood.org)

Important Biological Control Parasitoids of May and June beetles include wasps in the genus Tiphia
Agents Related to this (Tiphiidae), especially Tiphia inornata (Say), as well as several species
Pest Species of tachinid parasitoids such as Microphthalma disjuncta Wiedman.
Grubs of Phyllophaga species are also attacked in the soil by various
bacteria, fungi and nematodes, some of which have been formulated as
biopesticides with varying levels of success in practical use.

Web Links for Information http://www.ag.ndsu.nodak.edu/aginfo/entomology/entupdates/whitegrub/

on May and June Beetles whitgrub.htm; fact sheet of the North Dakota State University of
prevention and management of white grub damage.
http://en.wikipedia.org/wiki/Phyllophaga_(genus); Wikipedia article
providing overview of white grubs in the genus Phyllophaga.
http://www.unl.edu/museum/research/entomology/Guide/Scarabaeoidea/
Scarabaeidae/Melolonthinae/Melolonthinae-Generic-pages/Melolonthini/
Phyllophaga/Phyllophaga.html; a guide to New World scarab genera,
including profiles of several Phyllophaga species.
http://www.freshfromflorida.com/pi/enpp/ento/scarab-pest-alert.html;
two pest Phyllophaga species newly invasive in Florida.
http://www.forestpests.org/nursery/whitegrubs.html; USDA Forest
Service fact sheet on white grubs as pests in forest nurseries.
A Photographic Guide 305

Articles Luginbill, P. and H.R. Painter. 1953. May beetles of the United States
and Canada. U.S. Department of Agriculture Technical Bulletin
No. 1060: 102 p.
Ives, W.G.H. and G.L. Warren. 1965. Sequential sampling for white
grubs. The Canadian Entomologist 97: 596-604.
Ritcher, P.O. 1966. White grubs and their allies. A study of North
American scarabaeoid larvae. Oregon State Monographs, Studies in
Entomology No. 4: 214 p.
Fowler, R.F. and L.F. Wilson. 1971. White grub populations,
Phyllophaga spp. in relation to damaged red pine seedlings in Michigan
and Wisconsin plantations (Coleoptera: Scarabaeidae). Michigan
Entomologist 4: 23-28.
Liesch, P.J. and R.C. Williamson. 2010. Evaluation of chemical controls
and entomopathogenic nematodes for control of Phyllophaga white
grubs in a Fraser fir production field. Journal of Economic Entomology
103: 1979-1987.
306 Forest Pest Insects in North America:

86. Black Vine Weevil, Otiorhynchus sulcatus (Fabricius) (Coleoptera:

Curculionidae)

Orientation to Pest The black vine weevil, Otiorhynchus sulcatus (Fabricius), is a weevil that
is likely native to some or all of Europe that has invaded North America,
Japan, New Zealand, Australia, and parts of southern South America. The
mechanism of these invasions has been movement of the insect in balled
nursery stock plants. This species feeds on over 100 species of plants.
Larvae are the main damaging stage, feeding in the soil on plant roots.
Young larvae eat the fine roots, but the following year, mature larvae
destroy the larger roots as well. Unlike the white grubs of scarabs such
as Japanese beetle (Popillia japonica Newman), weevil larvae such as
those of black vine weevil, are less C-shaped and lack legs. Adults feed
at night, notching the leaves of their host plants, but damage from adult
feeding is not severe. Leaf notching is, however, a useful indicator of
the presence of this nonflying, nocturnal insect whose larvae are out of
sight in the soil. The most common overwintering stage of this insect is
the young larva, but sometimes adults too overwinter. In general, in the
eastern United States, adults emerge in June. Black vine weevils are pests
in greenhouses and forestry nurseries where planting stock is produced,
to both potted and field-grown plants.

Hosts Commonly Attacked Plants fed on by black vine weevil include many woody and non-woody
species, including strawberry (Fragaria), yew (Taxis), spruce (Picea),
hemlock (Tsuga), rhododendron (Rhododendron), grape (Vitis), and
cyclamen (Cyclamen).

Distribution The black vine weevil is found in the northeastern and north central
parts of the United States and adjacent parts of eastern Canada, and from
Alaska south to California.

Images of
Black Vine Weevil

Figure 1. Adults of black vine weevil, Otiorhynchus sulcatus, are flightless (wing
covers do not open) and nocturnal. (Left: Kent Loeffler, Cornell University,
Bugwood.org; right: Cheryl Moorehead, individual, Bugwood.org)
A Photographic Guide 307

Images of Black
Vine Weevil (continued)

Figure 2. Larvae of black vine weevil are cream colored, with a brown head and
no legs. (Left: Peggy Greb, USDA Agricultural Research Service, Bugwood.org;
right: Mike Reding and Betsy Anderson, USDA Agricultural Research Service,
Bugwood.org)

Figure 3. Pupae of black vine weevil occur naked in the soil. (Left: Mike Reding
and Betsy Anderson, USDA Agricultural Research Service, Bugwood.org;
right: Jim Baker, North Carolina State University, Bugwood.org)

Figure 4. Adult black vine weevils notch edges of host leaves (left, sedum plants
and right, rhododendrons). (Left: Mike Reding and Betsy Anderson, USDA
Agricultural Research Service, Bugwood.org; right: Jim Baker, North Carolina
State University, Bugwood.org)
308 Forest Pest Insects in North America:

Images of Black Figure 5. Damage to

yew seedling roots from
Vine Weevil (continued) black vine weevil larvae.
(Jim Baker,
North Carolina
State University,
Bugwood.org)

Important Biological Control Specialized parasitoids or predators of this species have not been
Agents Related to this reported. Most attention has been on various species of nematodes
Pest Species (e.g., Heterorhabditis heliothidis [Kahn, Brooks, and Hirschman]) and
fungal entomopathogens (e.g., Metarhizium anisopliae [Metchnikoff]
Sorokin), which have been developed for use as biopesticides against this
species with varying degrees of success depending on such things as soil
temperature.

Web Links for Information http://woodypests.cas.psu.edu/factsheets/insectfactsheets/html/black_

on Black Vine Weevil vine_weevil.html; fact sheet of Pennsylvania State University.
http://www.umassgreeninfo.org/fact_sheets/root_shoot_feeders/black_
vine_weevil.html; fact sheet of the University of Massachusetts.
http://www.coopext.colostate.edu/TRA/PLANTS/blackvine.shtml; fact
sheet of the Colorado State University.
A Photographic Guide 309

Articles Haukeland, S. and T. Lola-Luz. 2010. Efficacy of the entomopathogenic

nematodes Steinernema kraussei and Heterorhabditis megidis against the
black vine weevil Otiorhynchus sulcatus in open field-grown strawberry
plants. Agricultural and Forest Entomology 12(4): 363-369.
Ansari, M.A., F.A. Shah, and T.M. Butt. 2008. Combined use of
entomopathogenic nematodes and Metarhizium anisopliae as a new
approach for black vine weevil, Otiorhynchus sulcatus, control.
Entomologia Experimentalis et Applicata 129: 340-347.
von Reibnitz, C. and G.F. Backhaus. 1992. Analysis of the incidence and
control of Otiorhynchus sulcatus in tree nurseries. Results from a survey
in Lower Saxony and Schleswig-Holstein. Gesunde Pflanzen 45(2):
54-60. (In German).
310 Forest Pest Insects in North America:

87. Strawberry Root Weevil, Otiorhynchus ovatus (L.) (Coleoptera:

Curculionidae)

Orientation to Pest The strawberry root weevil, Otiorhynchus ovatus (L.), is invasive
in North America. It is polyphagous, and larvae feed on plant roots,
while adults feed nocturnally on the foliage, buds and young shoots
of a wide range of host plants. Adults do not fly and all are females.
Unlike white grubs (larvae of scarabs), weevil larvae such as those
of strawberry root weevil are less C-shaped and lack legs. Black vine
weevil, Otiorhynchus sulcatus (Fabricius), is a similar appearing species,
but whose larvae are much larger than those of strawberry root weevil.
In Oregon, strawberry root weevil overwinters as partially grown larvae
or as adults. Adults emerge in late May and early June and begin laying
eggs about two weeks later. Larvae feed from September through April
of the following year and cause the most damage from late March to
mid-May. Strawberry root weevil is a forestry pest only in the context of
forest nurseries, where infestations may build up. Larvae feed on roots
of tree seedlings and can be controlled by soil applications of chemical
insecticides, nematodes, or entomopathogenic fungi.

Hosts Commonly Attacked Adult beetles feed on foliage, especially of arborvitae (Thuja). The larvae
feed on the roots of hemlock (Tsuga) and various other conifers, such as
spruce (Picea), pine (Pinus), yew (Taxus), and arborvitae, as well as a
variety of agricultural crops.

Distribution The strawberry root weevil is found in most parts of the United States
and southern Canada.

Images of
Strawberry Root Weevil

Figure 1. Adult of strawberry root weevil, Otiorhynchus ovatus. (Left: Pest and
Diseases Image Library, Bugwood.org; right: Whitney Cranshaw, Colorado State
University, Bugwood.org)
A Photographic Guide 311

Images of Strawberry Figure 2. Larva of

strawberry root weevil.
Root Weevil (continued) (Russell Karow, Oregon
State University)

Figure 3. Pupa of
strawberry root weevil.
(Russell Karow, Oregon
State University)

Figure 4.Girdling of bare root

spruce seedling by adults of
strawberry root weevil. (Natural
Resources Canada, Canadian
Forest Service)
312 Forest Pest Insects in North America:

Important Biological Control The only parasitoid known to attack this species in North America
Agents Related to this is the braconid Triaspis kurtogaster Martin, but that parasitoid’s
Pest Species importance in controlling populations of the weevil is unknown. Various
entomopathogenic fungi have been found attacking larvae of strawberry
root weevil, such as Isaria fumosorosea Wize (Ifr.) (formerly known as
Paecilomyces fumosoroseus [Wize] Brown and Smith) and Metarhizium
anisopliae (Metchnikoff) Sorokin. Various nematodes also attack larvae
of this species, Heterorhabditis bacteriophora Poinar being the species
most promising for control. Several species of both fungi and nematodes
have been assessed as biopesticides against larvae of strawberry root
weevil in various settings, including forest nurseries, cranberry bogs, and
potted nursery stock in and around greenhouses. In general, control has
been better in potted plants than open fields, and in warmer soils versus
cold soils. Cool temperatures (11-14 °C) reduce efficacy of some species
of nematodes.

Web Links for Information http://mint.ippc.orst.edu/srwfact.pdf; fact sheet of Oregon State

on Strawberry Root Weevil University on biology and management.
http://whatcom.wsu.edu/ipm/manual/rasp/weevil.html; fact sheet of
Washington State University comparing several species in the genus
Otiorhynchus.
http://www.pfc.cfs.nrcan.gc.ca/diseases/nursery/pests/rootweev_e.html;
fact sheet of Natural Resources Canada discussing root weevils as pests
in forest nurseries.

Articles Nielsen, D.G. 1989. Minimizing Otiorhynchus root weevil impact in

conifer nurseries. In: Alfaro, R.I. and S.G. Glover (eds.). Proceedings
of Conference: Insects affecting reforestation: biology and damage.
Forestry Canada, Pacific and Yukon Region, Victoria, British Columbia:
71-79.
Vainio, A. and H.M.T. Hokkanen. 1993. The potential of
entomopathogenic fungi and nematodes against Otiorhynchus ovatus
L. and O. dubius Ström (Col., Curculionidae) in the field. Journal of
Applied Entomology 115(4): 379-387.
Berry, R.E., J. Liu, and E. Groth. 1997. Efficacy and persistence of
Heterorhabditis marelatus (Rhabditida: Heterorhabditidae) against
root weevils (Coleoptera: Curculionidae) in strawberry. Environmental
Entomology 26: 465-470.
A Photographic Guide 313

88. Japanese Beetle, Popillia japonica Newman (Coleoptera: Scarabaeidae)

Orientation to Pest The Japanese beetle, Popillia japonica Newman, is an invasive insect
in North America (United States and Canada) that is native to Japan.
It is also invasive in China, Russia and Portugal. Adults fly during
the summer and may occur in large numbers, defoliating raspberries,
strawberries, roses, grapes, and other plants. Adults burrow into the soil,
especially in grassy areas, and lay eggs several inches underground,
where larvae then develop, feeding on the roots of grass plants and
damaging the turf. There is usually one generation per year and partly
grown larvae in the soil are the overwintering stage.

Hosts Commonly Attacked Adults defoliate a wide range of plants including grapes, roses, and
various shade and fruit trees. Larvae eat the roots of grasses and are
major pests of turf, especially on golf courses. Larvae can be pests in tree
nurseries or on ornamental nursery stock.

Distribution In North America, the Japanese beetle is found in most states east of
the Mississippi River and the invaded area continues to expand. One
important pathway by which this pest is spread is the movement of larvae
in nursery stock shipped with soil. The beetle is not found in California
or the Pacific Northwest states.

Images of
Japanese Beetle

Figure 1. Adult Japanese beetle, Popillia japonica. (David Cappaert, Michigan

State University, Bugwood.org)
314 Forest Pest Insects in North America:

Images of Japanese
Beetle (continued)

Figure 2. Larva of the Japanese beetle (left); three species of “white grubs”
(right) to show relative size, from left to right: Japanese beetle, European chafer
(Rhizotrogus majalis [Razoumowsky]), and June beetle (Phyllophaga sp.).
(Both photos: David Cappaert, Michigan State University, Bugwood.org)

Figure 3. The V-shaped pattern of the “raster” (a group of hairs on underside of

the rear tip of larval body) is used to identify larvae of Japanese beetles.
(Mike Reding and Betsy Anderson, USDA Agricultural Research Service,
Bugwood.org; inset: Clemson University, Cooperative Extension Fact Sheet)

Figure 4. Adult Japanese beetles feeding on rose flower (left) and foliage of other
plants (right). (Left: Clemson University - USDA Cooperative Extension Slide
Series, Bugwood.org; right: Ronald S. Kelley, Vermont Department of Forests,
Parks and Recreation, Bugwood.org)
A Photographic Guide 315

Images of Japanese
Beetle (continued)

Figure 5. Turf infested by Japanese beetle larvae and then dug up by skunks or
other mammals to eat grubs. (M.G. Klein, USDA Agricultural Research Service,
Bugwood.org)

Important Biological Control A large scale effort to achieve biological control of this pest, based
Agents Related to this on searching for natural enemies in Japan and China, was made in the
Pest Species 1920s and 1930s. Several parasitoids were imported and established,
the most important of which were three tachinid flies (especially
Istocheta aldrichi [Mesnil]) and several tiphiid wasps (especially Tiphia
popilliavora Rohwer and Tiphia vernalis Rohwer). Several pathogens
of this beetle were also discovered and attempts were made to turn them
into biopesticides. These included the bacterium that causes milky spore
disease (Paenibacillus popilliae [Dutky]) and the nematode Steinernema
glaseri (Steiner). It was the study of this nematode that stimulated the
development of this family of nematodes as successful biopesticides
against soil insects. For a detailed history of the biological control efforts
against Japanese beetle, see Van Driesche et al. (1996).

Web Links for Information http://en.wikipedia.org/wiki/Japanese_beetle; Wikipedia article on

on Japanese Beetle biology and control.
http://edis.ifas.ufl.edu/in630; fact sheet of the University of Florida,
discussing biology and control.
http://www.aphis.usda.gov/publications/plant_health/content/printable_
version/jbidcard5-07.pdf; USDA fact sheet including a life cycle
diagram.
316 Forest Pest Insects in North America:

Articles on Van Driesche, R.G., S. Healy, and R.C. Reardon. Biological Control
Japanese Beetle of Arthropod Pests of the Northeastern and North Central Forests
in the United States: A review and recommendations. FHTET-96-19.
USDA Forest Service, Morgantown, West Virginia: 43-48. (Available
at http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
Power, K.T., R.S. An, and P.S. Grewal. 2009. Effectiveness of
Heterohabditis bacteriophora strain GPS11 applications targeted against
different instars of the Japanese beetle Popillia japonica. Biological
Control 48: 232-236.
Oliver, J.B., M.E. Reding, N.N. Youssef, M.G. Klein, B.L. Bishop, and
P.A. Lewis. 2009. Surface-applied insecticide treatments for quarantine
control of Japanese beetle, Popillia japonica Newman (Coleoptera:
Scarabaeidae), larvae in field-grown nursery plants. Pest Management
Science 65: 381-390.
A Photographic Guide 317

89. Twolined Chestnut Borer, Agrilus bilineatus (Weber) (Coleoptera:

Buprestidae)

Orientation to Pest Twolined chestnut borer, Agrilus bilineatus (Weber), is native to North
America and occurs throughout southeastern Canada and the eastern
and central United States. Adults are recognizable as dark colored
buprestids with two golden stripes running lengthwise along their back.
Adult beetles fly from April to August, depending on the location, and
there is one generation per year. In Michigan and surrounding areas,
adult emergence peaks in mid-to late June. Females lay eggs in small
clusters in bark cracks and crevices. Newly hatched larvae burrow into
the tree and form feeding galleries under the bark. Larvae are light
colored and up to 2.5 cm long when mature, and have two spines at the
tip of the abdomen, as is typical for all Agrilus larvae. Larvae construct
meandering galleries that are packed tightly with feces mixed with boring
dust. These feeding galleries interrupt the transport of food and water
in the phloem (inner bark) and xylem (outer sapwood) and eventually
girdle individual branches or the entire tree. Attacks occur in stressed or
dying chestnut (Castanea dentata [Marsh.] Borkh.) or oak (Quercus).
Live healthy trees are typically not infested. Attacks usually begin in the
crown of the tree, with some branches dying in the first year. Infestations
progress downward, and the trees usually die in the second or third
year. Incidence of attack by twolined chestnut borer increases following
stress such as drought or defoliation from insects such as gypsy moth
(Lymantria dispar [L.]).

Hosts Commonly Attacked American chestnut (C. dentata) and various species of oak, especially
white (Quercus alba L.), scarlet (Q. coccinea Muenchh.), northern pin
(Q. ellipsoidalis E. J. Hill), bur (Q. macrocarpa Michx.), chestnut
(Q. prinus L.), northern red (Q. rubra L.), post (Q. stellata Wangenh.),
black (Q. velutina Lamb.), and live (Q. virginiana Miller) oaks are hosts
of twolined chestnut borer.

Distribution Twolined chestnut borer is

widespread in the eastern
and central United States
and southeastern Canada.
Figure 1. Probable range of
the twolined chestnut borer,
Agrilus bilineatus, based on
the combined distribution of
its oak host species. (USDA
Forest Service • Forest Insect
and Disease Leaflet 168)
318 Forest Pest Insects in North America:

Images of Twolined
Chestnut Borer

Figure 2. Adult of twolined chestnut borer, showing the two golden stripes along
the back. (Robert A. Haack, USDA Forest Service, Bugwood.org)

Figure 3. Larvae of twolined chestnut borer in galleries (left) and close up of larva
(right), showing the two projections from the rear end of the body (found in all
Agrilus species). (Left: James Solomon, USDA Forest Service, Bugwood.org;
right: Robert A. Haack, USDA Forest Service, Bugwood.org)

Figure 4. The exit hole of the twolined chestnut

borer, like that of other species of Agrilus, is D
shaped rather than round or oval. (USDA Forest
Service • Forest Insect & Disease Leaflet 168)

Figure 5. View of the long winding larval galleries of twolined chestnut borer.
(Minnesota Department of Natural Resources Archive, Minnesota Department of
Natural Resources, Bugwood.org)
A Photographic Guide 319

Images of Twolined Figure 6. The earliest

signs of twolined
Chestnut Borer (continued) chestnut borer attack
are wilting of foliage in
later summer and then
death of upper limbs of
tree. (Steven Katovich,
USDA Forest Service,
Bugwood.org)

Figure 7. Phasgonophora sulcata is a

common larval parasite of the twolined
chestnut borer in Wisconsin. (USDA
Forest Service • Forest Insect & Disease
Leaflet 168)

Important Biological Control Larval parasites provide limited natural control of the twolined chestnut
Agents Related to this borer. One larval parasite, the chalcid wasp Phasgonophora sulcata
Pest Species Westwood, causes up to 10 percent larval mortality.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/chestnutborer/chestnutborer.htm;

on Twolined Chestnut Borer USDA Forest Service Forest Insect and Disease Leaflet No. 168.

Articles Dunbar, D.M. and G.R. Stephens. 1976. The bionomics of the twolined
chestnut borer. In: Anderson, J.F. and H.K. Kaya (eds.) Perspectives in
Forest Entomology. New York, New York. Academic Press: 73-83.
Cote, W.A., III and D.C. Allen. 1980. Biology of twolined chestnut
borer, Agrilus bilineatus, in Pennsylvania and New York. Annals of the
Entomological Society of America 73: 409-413.
320 Forest Pest Insects in North America:

Articles (continued) Haack, R.A., D.M. Benjamin, and B.A. Schuh. 1981. Observations on
the biology of Phasgonophora sulcata (Hymenoptera: Chalcididae),
a larval parasitoid of the twolined chestnut borer, Agrilus bilineatus
(Coleoptera: Buprestidae), in Wisconsin. The Great Lakes Entomologist
14: 113-116.
Haack, R.A. and D.M. Benjamin. 1982. The biology and ecology of the
twolined chestnut borer, Agrilus bilineatus (Coleoptera: Buprestidae),
on oaks, Quercus spp., in Wisconsin. The Canadian Entomologist 114:
385-396.
Dunn, J.P., T.W. Kimmerer, and G.L. Nordin. 1986. The role of host tree
condition in attack of white oaks by the twolined chestnut borer, Agrilus
bilineatus (Weber) (Coleoptera: Buprestidae). Oecologia 70: 596-600.
Muzika, R.M., A.M. Liebhold, and M.J. Twery. 2000. Dynamics of
twolined chestnut borer, Agrilus bilineatus, as influenced by defoliation
and selection thinning. Agricultural and Forest Entomology 2:
283-289. (Available at http://onlinelibrary.wiley.com/doi/10.1046/j.1461-
9563.2000.00077.x/pdf).
A Photographic Guide 321

90. Bronze Birch Borer, Agrilus anxius Gory (Coleoptera: Buprestidae)

Orientation to Pest Bronze birch borer, Agrilus anxius Gory, adults are slender, olive-
bronze beetles, with a coppery reflection. The species is native to North
America and is widespread in the eastern and central United States
and southeastern Canada. Before 1950, bronze birch borer and another
species, now called aspen borer (Agrilus liragus Barter and Brown) were
considered to be the same species; so records from before this period
need to be segregated based on the genus of tree attacked, with only
Betula-attacking records being bronze birch borer. The biology of this
species is similar to that of other Agrilus species, with adults emerging
in late spring or early summer, laying eggs in bark cracks or under bark
flaps, and larvae tunneling through the bark to the phloem and cambium
layers, where they construct and feed in long meandering galleries.
Full-grown larvae construct shallow cells in the xylem (outer sapwood)
in autumn where they overwinter. They pupate the following spring
and adults emerge shortly thereafter. There is usually one generation
per year, but it may require two years to reach maturity in some areas.
Bronze birch borer develops in weakened trees of various species of
birch (Betula), especially the white-barked species. Individual branches
or entire trees are killed by the girdling action of the larvae. Bronze
birch borer is considered the most serious pest of paper birch (Betula
papyrifera Marsh.) and extensive outbreaks occurred in New Brunswick
in 1939 and in the Great Lakes states in the 1990s. In both cases,
outbreaks were in large stands of older birch trees stressed by defoliating
insects or drought. Bronze birch borer may attack healthy trees during
outbreaks, especially isolated trees, and ones on the edges of clearings.
Ornamental birches, especially those of Asian and European origin, are
often highly susceptible to bronze birch borer attack.

Hosts Commonly Attacked Species that are severely attacked include European white birch
(B. pendula Roth) and whitebarked Himalayan birch (B. jacquemontii D.
Don). Commonly attacked species include paper birch (B. papyrifera),
gray birch (B. populifolia Marsh.), sweet birch (B. lenta L.), and yellow
birch (B. alleghaniensis Britt.).

Distribution Bronze birch borer occurs across Canada from Newfoundland to British
Columbia and south to New Jersey, Ohio, Colorado, Idaho, and Utah in
the United States.
322 Forest Pest Insects in North America:

Images of
Bronze Birch Borer

Figure 1. Adult bronze birch borer, Agrilus anxius. (John A. Weidhass, Virginia
Polytechnic Institute and State University, Bugwood.org)

Figure 2. Larva of bronze birch borer. (Whitney Cranshaw, Colorado State

University, Bugwood.org)

Figure 3. Larval galleries

of bronze birch borer
in a limb (top) and the
trunk (bottom). (Top:
William M. Ciesla, Forest
Health Management
International,
Bugwood.org; bottom:
Whitney Cranshaw,
Colorado State
University, Bugwood.org)
A Photographic Guide 323

Images of Bronze
Birch Borer (continued)

Figure 4. Mature larvae tunnel a short way into the wood (left) to overwinter
and pupate in spring; right, pupa of bronze birch borer in outer sapwood. (Left:
E. Bradford Walker, Vermont Department of Forests, Parks and Recreation,
Bugwood.org; right: Jim Baker, North Carolina State University, Bugwood.org)

Figure 5. D-shaped
exit hole made by adult
bronze birch borer.
(Whitney Cranshaw,
Colorado State
University, Bugwood.org)

Figure 6. Landscape trees with infestations of bronze birch borer. (Left: Steven
Katovich, USDA Forest Service, Bugwood.org; right: Randy Cyr, Greentree,
Bugwood.org)
324 Forest Pest Insects in North America:

Images of Bronze
Birch Borer (continued)

Figure 7. Damage from bronze birch borer in a forest birch stand in Minnesota.
(USDA Forest Service - Northeastern Area Archive, USDA Forest Service,
Bugwood.org)

Important Biological Control Natural enemies of bronze birch borer have been studied in New
Agents Related to this Brunswick and Pennsylvania. Egg parasitism by several species (the
Pest Species signiphorid Thysanus sp. and the encrytid Coccidencyrtus sp.) was
common (about 50 percent) and was considered important. Important
larval parasites included the braconid Atanycolus charus (Riley) and the
chalcidid Phasgonophora sulcata Westwood, which together killed about
18 percent of the larvae.

Web Links for Information http://ipm.illinois.edu/landturf/insects/bronze_birch_borer/index.html;

on Bronze Birch Borer IPM fact sheet from Illinois.
http://www.na.fs.fed.us/spfo/pubs/fidls/bbb/bbb.htm; USDA Forest
Service Forest Insect & Disease Leaflet 111.
http://wiki.bugwood.org/Agrilus_anxius; Bugwood Wiki fact sheet.
http://www.eppo.org/QUARANTINE/Alert_List/insects/agrilus_anxius.
htm; EPPO fact sheet on risk of bronze birch borer as a potential risk for
invasion to Europe.
http://www.eppo.org/QUARANTINE/Pest_Risk_Analysis/PRA_
documents.htm; EPPO fact sheet on risk of bronze birch borer to
European birches.
A Photographic Guide 325

Articles Anderson, R.F. 1944. The relation between host condition and attacks by
the bronze birch borer. Journal of Economic Entomology 37: 588-596.
Barter, G.W. 1957. Studies of the bronze birch borer, Agrilus anxius
Gory, in New Brunswick. The Canadian Entomologist 89: 12-36.
Ball, J. and G. Simmons. 1980. The relationship between bronze birch
borer and birch dieback. Journal of Arboriculture 6: 309-314.
Akers, R.C. and D.G. Nielsen. 1984. Predicting Agrilus anxius Gory
(Coleoptera: Buprestidae) adult emergence by heat unit accumulation.
Journal of Economic Entomology 77: 1459-1463.
Jones, E.A., D.D. Reed, G.D. Mroz, H.O. Liechty, and P.J. Cattelino.
1993. Climate stress as a precursor to forest decline: paper birch in
northern Michigan, 1985-1990. Canadian Journal of Forest Research
23: 229-233.
326 Forest Pest Insects in North America:

91. Emerald Ash Borer, Agrilus planipennis Fairmaire (Coleoptera:

Buprestidae)

Orientation to Pest Emerald ash borer, Agrilus planipennis Fairmaire, is an invasive borer
from northeast Asia threatening North American ash trees (Fraxinus). It
was first detected near Detroit, Michigan and likely was introduced in
the 1990s. It is now found in 14 other states and two Canadian provinces,
and the infested range is expanding rapidly. Emerald ash borer attacks
and kills healthy ash trees from ones several inches in diameter to
mature trees. Massive mortality to ash of several species has occurred
since the species’ invasion in both landscape plantings and natural ash-
dominated communities, especially in riparian areas. Larvae feed on
phloem and make serpentine galleries that girdle and kill trees when the
larval densities are high. Mature larvae tunnel into sapwood to pupate.
In northern areas (e.g., Michigan), a single generation may require two
years, but in mid-Atlantic states (e.g., Maryland), a generation can be
completed each year. In dense populations, woodpeckers consume many
larvae.

Hosts Commonly Attacked Most North American Fraxinus species are susceptible, but so far the
most affected species have been white (Fraxinus americana L.), green
(F. pennsylvanica Marshall), and black (F. nigra Marshall) ash.

Distribution The North America infested area is centered on Michigan. Extensive

infestations exist in Ontario, Illinois, Indiana, Ohio, Kentucky,
Pennsylvania, and Maryland, and smaller infested areas are found in
Quebec, Minnesota, Iowa, Wisconsin, Missouri, Tennessee, New York,
Virginia, and West Virginia (as of 2010).

Figure 1. Distribution of emerald ash borer, Agrilus planipennis, in 2010.

(Cornell University, Cooperative Extension • The New York Invasive Species
Clearinghouse)
A Photographic Guide 327

Images of
Emerald Ash Borer

Figure 2. Adults of emerald ash borer. (Top left: Howard Russell, Michigan State
University, Bugwood.org; top right and bottom: David Cappaert, Michigan State
University, Bugwood.org)

Figure 3. Emerald ash

borer eggs are white
when freshly laid (top),
but turn tan as they age
(bottom). (Both photos:
Houping Liu, Michigan
State University,
Bugwood.org)
328 Forest Pest Insects in North America:

Images of Emerald
Ash Borer (continued)

Figure 4. Feeding stage larvae of emerald ash borer: full grown 4th instar (left);
second, third and 4th instars (right). (Both photos: David Cappaert, Michigan
State University, Bugwood.org)

Figure 5. Larval galleries of emerald ash borer. (Top left: Troy Kimoto, Canadian
Food Inspection Agency, Bugwood.org; top right: Michigan Department of
Agriculture, Bugwood.org; bottom: Edward Czerwinski, Ontario Ministry of
Natural Resources, Bugwood.org)
A Photographic Guide 329

Images of Emerald
Ash Borer (continued)

Figure 6. Prepupae in chambers formed in sap wood by mature larvae. (Brian

Sullivan, USDA APHIS PPQ, Bugwood.org)

Figure 7. Prepupa of emerald ash borer in chamber in sap wood. (David

Cappaert, Michigan State University, Bugwood.org)

Figure 8. D shaped emergence hole of emerald ash borer (left) and hole made
by woodpecker (right) where an emerald ash borer larva was removed. (David
Cappaert, Michigan State University, Bugwood.org)
330 Forest Pest Insects in North America:

Images of Emerald
Ash Borer (continued)

Figure 9. Signs of infestation: left, suckering; right, bark splits. (Left: Joseph
O’Brien, USDA Forest Service, Bugwood.org; right: Michigan Department of
Agriculture, Bugwood.org)

Figure 10. Dead and dying ash in forest area due to emerald ash borer. (Troy
Kimoto, Canadian Food Inspection Agency, Bugwood.org)

Figure 11. Dying or dead landscape trees affected by emerald ash borer. (Left:
Steven Katovich, USDA Forest Service, Bugwood.org; right: David Cappaert,
Michigan State University, Bugwood.org)
A Photographic Guide 331

Images of Emerald Figure 12. Tree

removal was tried,
Ash Borer (continued) unsuccessfully, as a
means to eradicate
emerald ash borer
populations. (Top: David
Cappaert, Michigan
State University,
Bugwood.org; bottom:
Daniel Herms, The
Ohio State University,
Bugwood.org)

Figure 13. Girdling trees can be an effective survey method to detect emerald
ash borers. (Pennsylvania Department of Conservation and Natural Resources
- Forestry Archive, Bugwood.org)
332 Forest Pest Insects in North America:

Images of Emerald Figure 14. The Chinese

parasitoid of emerald
Ash Borer (continued) ash borer eggs, Oobius
agrili. (Debbie Miller,
USDA Forest Service,
Bugwood.org)

Figure 15. The Chinese

larval parasitoid
Tetrastichus planipennisi.
(Houping Liu, Michigan
State University,
Bugwood.org)

Figure 16. The Chinese

emerald ash borer larval
parasitoid, Spathius
agrili. (Jennifer Ayer,
Bugwood.org)
A Photographic Guide 333

Images of Emerald
Ash Borer (continued)

Figure 17. Larva, pupa (in cocoon), and adult of the native North American
parasitoid Atanycolus cappaerti Marsh and Strazanac. (Top and middle:
David Cappaert, Michigan State University, Bugwood.org; bottom: Jian Duan,
Bugwood.org)
334 Forest Pest Insects in North America:

Important Biological Control Natural enemies of emerald ash borer that have been collected in
Agents Related to this the native range (especially in China and Russia) include two larval
Pest Species parasitoids (the eulophid Tetrastichus planipennisi Yang and the braconid
Spathius agrili Yang), and an egg parasitoid, Oobius agrili Zhang and
Huang (Hymenoptera: Encyrtidae). Additional parasitoids have been
identified and are under consideration for importation, including Spathius
n. sp. and Antanycolus picipes Telenga from Russia. One group of native
North American parasitoids, braconids in the genus Atanycolus, has
adopted emerald ash borer and causes up to 20-40 percent mortality of
the larvae in some areas.

Web Links for Information http://www.invasiveforestinsectandweedbiocontrol.info/target_pests/

on Emerald Ash Borer insects_mites/Emeraldashborer.htm; University of Massachusetts fact
sheet on biological control of emerald ash borer.
http://www.nrs.fs.fed.us/disturbance/invasive_species/eab/control_
management/biological_control/; USDA Forest Service website on
biological control of emerald ash borer.
http://www.dec.ny.gov/animals/7253.html; fact sheet of the New York
Department of Environmental Conservation.

Articles Liu, H-P, L.S. Bauer, D.L. Miller, L-W. Song, Q-S. Luan, S-H. Sun,
and R.Z. Jin. 2007. Seasonal abundance and population dynamics of
Agrilus planipennis (Coleoptera: Buprestidae) and its natural enemies
Oobius agrili (Hymenoptera: Encyrtidae) and Tetrastichus planipennisi
(Hymenoptera: Eulophidae) in China. Biological Control 42: 61-71.
Yang, Z-Q, J.S. Strazanac, P.M. Marsh, C. van Achterberg, and W-Y.
Choi. 2005. First recorded parasitoid from China of Agrilus planipennis:
A new species of Spathius (Hymenoptera: Braconidae, Doryctinae).
Annals of the Entomological Society of America 98: 636-642.
Yang, Z-Q., J.S. Strazanac, Y-X. Yao, and X-Y. Wang. 2006. A new
species of emerald ash borer parasitoid from China belonging to the
genus Tetrastichus Haliday (Hymenoptera: Eulophidae) parasitizing
emerald ash borer from China. Proceedings of the Entomological Society
of Washington 108: 550-558.
Zhang,Y.Z., D.W. Huang, T.H. Zhao, H.P. Liu, and L.S. Bauer. 2005.
Two new species of egg parasitoids (Hymenoptera: Encrytidae) of wood-
boring beetle pests from China. Phytoparasitica 33: 253-260.
Duan, J.J., L.S Bauer, J.R. Gould, and J.P. Lelito. 2011. Biological
control of emerald ash borer in North America: Current progress and
potential for success. IOBC Newsletter, November 2011.
Duan, J.J., L.S Bauer, K.J. Abell, and R.G. Van Driesche. 2011.
Population responses of hymenopteran parasitoids to the emerald
ash borer (Coleoptera: Buprestidae) in north central United States.
BioControl 57: 199-209.
A Photographic Guide 335

92. Goldspotted Oak Borer, Agrilus auroguttatus Schaeffer (Coleoptera:

Buprestidae)

Orientation to Pest Goldspotted oak borer, Agrilus auroguttatus Schaeffer, is an oak-

attacking buprestid native to mountains in southern Arizona. This pest
invaded southern California, likely having been moved in firewood taken
by campers into public campgrounds in the region. It has killed more
than 80,000 oaks in California’s native oak-savannahs and is of concern
as an ecological pest. It is still spreading due to unrestricted movement
of firewood out of the affected parts of southern California, and the
ultimate extent of potential damage is not yet known. The biology of this
species is very similar to that of other Agrilus species, such as the better
known emerald ash borer (Agrilus planipennis Fairmaire). In southern
California, most goldspotted oak borers complete their development
in one year, but some may require longer. Adults have an extended
emergence period, but most do so in late June or early July. Adults have
a pre-oviposition period during which they feed on oak foliage. Eggs are
likely laid in crevices in the bark, and larvae after hatching bore through
the bark. Young larvae feed in the outer phloem, while older larvae feed
in the cambial layer. Pupation occurs in a chamber made by larvae in the
outer phloem. Affected trees experience death of limbs, die back, and
loss of vigor. Attacked trees often die within 2-3 years.

Hosts Commonly Attacked The species most affected are coast live oak (Quercus agrifolia Née)
and California black oak (Q. kelloggii Newb.). Canyon live oak (Q.
chrysolepis Liebm.) is affected to a lesser degree. See http://en.wikipedia.
org/wiki/Agrilus_coxalis - cite_note-npag-0.

Distribution Goldspotted oak borer occurs as a native species in southern Arizona and
as an invader in southern California. A related species, now known as
Agrilus coxalis Waterhouse, is native to southern Mexico and Guatemala.

Figure 1. Known
distribution of
goldspotted oak borer,
Agrilus auroguttatus,
in California (invaded
range-large circle) and
Arizona (native range-
small circles). (Tom
Coleman, USDA Forest
Service, Bugwood.org)
336 Forest Pest Insects in North America:

Images of
Goldspotted Oak Borer

Figure 2. Adult goldspotted oak borer. (Mike Lewis, Center for Invasive Species
Research, Bugwood.org)

Figure 3. Feeding larva of goldspotted oak borer, as seen in the larval gallery.
(Mark S. Hoddle, University of California - Riverside, Bugwood.org)

Figure 4. Mature, “J”-shaped larva of goldspotted oak borer (see folded

“hairpin” position of body). (Mark S. Hoddle, University of California - Riverside,
Bugwood.org)
A Photographic Guide 337

Images of Goldspotted
Oak Borer (continued)

Figure 5. Larval galleries of goldspotted oak borer under bark. (Tom Coleman,
USDA Forest Service, Bugwood.org)

Figure 6. Wet stains on bark (left) caused by feeding of larvae, and scaled areas
(right) where woodpeckers have removed larvae, are signs of goldspotted oak
borer infestations. (Left: Tom Coleman, USDA Forest Service, Bugwood.org;
right: Mark S. Hoddle, University of California - Riverside, Bugwood.org)
338 Forest Pest Insects in North America:

Images of Goldspotted Figure 7. Exit holes of

goldspotted oak borer
Oak Borer (continued) have the “D”-shape
typical of buprestids.
(Mark S. Hoddle,
University of California -
Riverside, Bugwood.org)

Figure 8. Dead oaks in California oak-savannah due to attack by goldspotted oak

borer. (Mike Lewis, Center for Invasive Species Research, Bugwood.org)

Figure 9. The eupelmid Calosota elongata, a larval ectoparasitoid of Agrilus

auroguttatus (left, female; right, male). (Mike Lewis, Center for Invasive Species
Research, Bugwood.org)
A Photographic Guide 339

Important Biological Control Natural enemies of this species are relatively unknown, but surveys
Agents Related to this are being conducted in the pest’s native range in southern Arizona
Pest Species to discover species suitable for importation to California. To date,
insects reared from logs infested with goldspotted oak borer have
mostly been generalist predators or parasitoids use not suitable for use
as classical biological control agents (for California). The euplemid
Calosota elongata Gibson is a larval parasitoid, collected from
Arizona, that appears to be more specialized and whose host range is
under investigation. The same species has been found in California in
association with goldspotted oak borer and may have been introduced
along with its host. DNA studies are underway to compare the California
population of C. elongata to ones in Arizona. No egg parasitoids are
known.

Web Links for Information http://cisr.ucr.edu/goldspotted_oak_borer.html; University of California,

on Goldspotted Oak Borer Riverside, Center for Invasive Species Research article on goldspotted
oak borer.
http://en.wikipedia.org/wiki/Agrilus_coxalis; Wikipedia article, under a
related but mistaken name of Agrilus coxalis.
http://ucanr.org/sites/SoCalOaks_NeedsAssessment2011/files/71253.pdf;
bibliographic list of articles about goldspotted oak borer.

Articles Coleman, T.W. and S.J. Seybold. 2008. New pest in California: The
goldspotted oak borer, Agrilus coxalis Waterhouse. USDA Forest
Service, Pacific Southwest Region, State and Private Forestry. Pest Alert
R5-RP-022: 4 p.
Hishinuma, S., T.W. Coleman, M.L. Flint, and S.J. Seybold. 2011.
Gold-spotted oak borer field identification guide. University of
California Agriculture and Natural Resources Statewide Integrated Pest
Management Program: 6 p. (Available at http://www.ipm.ucdavis.edu/
PDF/MISC/GSOB_field-identification-guide.pdf).
340 Forest Pest Insects in North America:

93. Soapberry Borer, Agrilus prionurus Chevrolat (Coleoptera: Buprestidae)

Orientation to Pest The soapberry borer, Agrilus prionurus Chevrolat, is a Mexican wood-
boring buprestid about which very little is known. It has moved out of
its native range in Mexico into Texas, where it is now killing western
soapberry trees (Sapindus saponaria var. drummondii [Hook. and
Arn.] L.D. Benson). How it got to Texas is not known, but movement
of firewood is a likely explanation. The biology of the insect is not
well known but is likely very similar to other invasive Agrilus (e.g.,
goldspotted oak borer [A. auroguttatus Schaeffer] and emerald ash borer
[A. planipennis Fairmaire]). The adults are about 1.3 cm long, black,
with four distinctive white dots on the wing covers. The larvae feed
immediately under the bark. While not certain, it is likely that the insect
has one generation per year, with adults emerging between May and
August. The insect probably overwinters as a mature larva or pupa in a
cell in the wood of the soapberry trees, with adult beetles later emerging
through a D-shaped exit hole and laying their eggs on the tree trunk.

Hosts Commonly Attacked The only reported host in Texas is western soapberry (S. saponaria var.
drummondii). Western soapberry trees larger than 5 cm in diameter at
breast height are susceptible and infestation usually results in tree death
within 1 to 3 years following initial attack. Hosts in Mexico have not
been recorded.

Distribution Native to Mexico, within the United States this species is currently found
only in Texas in 42 counties.

Figure 1. Distribution of soapberry borer, Agrilus prionurus, in the United States.

(USDA Forest Service • AFPE Online Mapping)
A Photographic Guide 341

Images of
Soapberry Borer

Figure 2. Adults of soapberry borer. (Ronald F. Billings, Texas Forest Service,

Bugwood.org)

Figure 3. Damage on
the trunk (top) of a
western soapberry tree
from soapberry borer is
highly visible, appearing
as scaled patches where
birds have picked off the
bark in search of larvae,
exposing galleries; larva
visible in photo below.
(Both photos: Ronald F.
Billings, Texas Forest
Service, Bugwood.org)
342 Forest Pest Insects in North America:

Images of Soapberry
Borer (continued)

Figure 4. Emergence hole of the soapberry borer, showing typical “D” shape
characteristic of buprestids. (Ronald F. Billings, Texas Forest Service,
Bugwood.org)

Figure 5. Western
soapberry tree showing
branches killed by the
soapberry borer and
epicormic sprouts along
lower trunk. (Ronald F.
Billings, Texas Forest
Service, Bugwood.org)
A Photographic Guide 343

Images of Soapberry
Borer (continued)

Figure 6. Damage to group of western soapberry trees by soapberry borer in

Houston, Texas. (Ronald F. Billings, Texas Forest Service, Bugwood.org)

Important Biological Control Natural enemies of western soapberry borer are unknown.
Agents Related to this
Pest Species
Web Links for Information http://tfsweb.tamu.edu/main/popup.aspx?id=5316; fact sheet of the Texas
on Soapberry Borer Forest Service.
http://elpasonaturally.blogspot.com/2009/06/wanted-location-of-
soapberry-borer.html; website of “El Paso Naturally,” which includes
photos of western soapberry borer.
http://www.texasinvasives.org; website providing description of insect
and questionnaire to report new infestations.

Articles Wellso, S.G. and J.A. Jackman. 2006. A new species of Anthaxia
(Haplanthaxia) Reitter (Coleoptera: Buprestidae) and new North
American buprestid distributional and host records. Pan-Pacific
Entomologist 82(2): 262-268. (First record in the USA).
Billings, R.F. and H.A. Pase, III. 2010. Soapberry borer continues to
spread in Texas. Texas Forestry, February issue: 7 p.
344 Forest Pest Insects in North America:

94. Hickory Spiral Borer, Agrilus torquatus LeConte (Coleoptera:

Buprestidae)

Orientation to Pest Hickory spiral borer, Agrilus torquatus LeConte, is a native beetle found
over a wide area in the eastern United States, feeding principally on
pecan (Carya illinoinensis [Wangenh.] K. Koch) and hickory (other
Carya spp.). There are a group of four closely species that previously
were considered subspecies of Agrilus arcuatus, but all of which have
now been raised to full species status, using their former subspecies
names. These include A. arcuatus torquatus, the focus of this page,
which attacks hickory and pecan; A. arcuatus fulgens from hazel
(Corylus); A. arcuatus corylicola—also from hazel (Corylus); and A.
arcuatus arcuatus from oak (Quercus) and American beech (Fagus
grandifolia Ehrh.). Agrilus torquatus requires two growing seasons
to complete its development, emerging in the spring or summer (May
to July) of the third year. After emerging, adults feed on host foliage,
making large irregular holes in the leaves. Eggs are deposited singly
on the bark surface of terminals or lateral twigs, usually near the base
of small shoots of the current season’s growth and are covered with a
transparent secretion that glues them in place. Larvae tunnel beneath
the bark, where they feed for two growing seasons. In late autumn, they
begin spiral burrows, encircling the stem until reaching the center and
severing the twigs by spring. Larvae pupate in cells constructed in the
pith of the stem. The most distinctive aspect of the damage of this species
is the spiral track made by the larvae, which is characteristic in form and
good for identification of the pest. This track is a winding concentric
gallery that moves from the inner bark to the heart of the branch or stem.
Branches and terminals are usually severed in late winter or spring and
the portion beyond the girdle typically dies before new foliage appears in
spring, making damaged terminals easy to spot. Most severed branches
break and fall to the ground around the time that buds open. Attacked
twigs are usually from ca 1-4 cm in diameter and 0.5 to 2.5 m long.
While most damage is to twigs and small branches, when seedlings
are attacked they may be killed. Extensive damage to large pecan trees
reduces nut production, and gives trees a ragged appearance. Repeated
attacks on young trees may make them stunted, misshapen, or crooked.

Hosts Commonly Attacked The hosts of this species are pecan (C. illinoinensis) and species of
hickory (Carya).

Distribution This species has been recorded from New York, Ohio, North Carolina,
and Mississippi.
A Photographic Guide 345

Images of
Hickory Spiral Borer

Figure 1. Adult of the closely related Agrilus arcuatus. (Tom Murray, Bugwood.org)

Figure 2. Larva of hickory spiral borer Agrilus torquatus. (James Solomon, USDA
Forest Service, Bugwood.org)

Figure 3. Distinctive spiraling pattern of larval galleries of hickory spiral borer.

(James Solomon, USDA Forest Service, Bugwood.org)
346 Forest Pest Insects in North America:

Important Biological Control Five insect parasitoids are known to attack this borer: Labena apicalis
Agents Related to this Cresson, Labena grallator (Say), Monogonogastra agrili (Ashmead),
Pest Species Tetrastichus rugglesi Rohwer, and Zatropis sp. near nigroaeneus
(Ashmead).

Web Links for Information None

on Hickory Spiral Borer

Articles Nelson, G.H. and H.A. Hespenheide. 1998. A re-evaluation of some

Agrilus Curtis species (Coleoptera: Buprestidae). Coleopterists Bulletin
52(1): 31-34.
A Photographic Guide 347

95. Flatheaded Appletree Borer, Chrysobothris femorata (Olivier)

(Coleoptera: Buprestidae)

Orientation to Pest Flatheaded appletree borer, Chrysobothris femorata (Olivier), is a

common, well known native borer affecting many species of hardwood
trees in North America. Adults emerge in summer and feed on foliage
of their host trees. Eggs are deposited under bark scales or in crevices
on the main trunk or larger branches. The larvae bore into the bark
and feed in the phloem and outer sapwood. In older trees, tunnels are
most often in the thick inner bark. Mature larvae build pupation cells in
the outer wood in late summer, where they pass the winter and pupate
the following spring. There is one generation per year. This borer is
especially damaging to newly planted trees and trees stressed by drought
or other factors. Young trees may be girdled and killed. Larger trees may
show injuries through loss of large patches of bark on trunks.

Hosts Commonly Attacked This species attacks a wide range of trees species, including sycamore
(Platanus occidentalis L.), silver maple (Acer saccharinum L.), boxelder
(A. negundo L.), black walnut (Juglans nigra L.), willow (Salix),
white oak (Quercus alba L.), black oak (Q. velutina Lamb.), yellow-
poplar (Liriodendron tulipifera L.), elm (Ulmus), American beech
(Fagus grandifolia Ehrh.), hickory (Carya), hackberry (Celtus), apple
(Malus domestica Borkh.), and pear (Pyrus).

Distribution This species is found throughout most of Canada and the United States.

Images of Flatheaded
Appletree Borer

Figure 1. Adult of the flatheaded appletree borer, Chrysobothris femorata.

(Joseph Berger, Bugwood.org)
348 Forest Pest Insects in North America:

Images of Flatheaded Figure 2. Mature larva

of flatheaded apple tree
Appletree Borer (continued) borer. (James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 3. Sap spot

over feeding site of
flatheaded appletree
borer larva in Nuttall oak
(Quercus texana Buckl.).
(James Solomon,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 349

Images of Flatheaded Figure 4. Damage to

red maple (Acer rubrum
Appletree Borer (continued) L.) from flatheaded
appletree borer. (John
Ruter, University of
Georgia, Bugwood.org)

Important Biological Control Natural enemies known to attack the flatheaded apple borer include
Agents Related to this the ichneumonids Labena grallator Say and Crytohelcostizus
Pest Species chrysobothridis Cushman, the chalcid Phasgonophora sulcata
Westwood, and the braconid Atanycolus rugosiventris Ashmead.
Predators include the clerids Chariessa pilosa (Foster) and Chariessa
pilosa onusta Say, the asilid Andrenosoma fulvicauda Say, and various
woodpeckers.

Web Links for Information http://insects.tamu.edu/extension/publications/epubs/eee_00027.cfm; fact

on Flatheaded Appletree sheet of the Texas A&M University covering biology and control.
Borer http://ipm.illinois.edu/greenhouse/insects/flat_headed_apple_tree_borer/
index.html; fact sheet of the University of Illinois.
http://www.utextension.utk.edu/publications/spfiles/SP503-I.pdf; guide
to the identification of apple flathead borer and other buprestids in
Kentucky.
http://wiki.bugwood.org/Archive:Hickory/Chrysobothris_femorata;
Bugwiki factsheet.

Articles Potter, D.A., G.M. Timmons, and F.C. Gordon. 1988. Flatheaded apple
treeborer (Coleoptera: Buprestidae) in nursery-grown red maples:
phenology of emergence, treatment timing, and response to stressed
trees. Journal of Environmental Horticulture 6(1): 18-22.
350 Forest Pest Insects in North America:

96. Red Oak Borer, Enaphalodes rufulus (Haldeman) (Coleoptera:

Cerambycidae)

Orientation to Pest Red oak borer, Enaphalodes rufulus (Haldeman), is a native borer
to North America, which attacks various species of oak (Quercus),
primarily those in the red oak subgenus. Red oak borer has a 2-year life
cycle that is unique in that adults only emerge during odd numbered
years. Mating takes place on the host tree and females lay eggs in
midsummer on roughened areas or near wounds, in bark crevices, or
under lichen or vines. Young larvae bore through the bark and spend
their first year in the phloem making small tunnels. The 2-year-old
larvae enlarge these phloem galleries and then bore into the xylem where
pupation takes place. The adult eventually emerges near the original
oviposition site. Fine frass is one of the first signs of attack. As the larvae
bore into the tree, wet spots can appear on the bark as sap oozes from the
bore hole in the bark. Larval galleries increase in size over time, reaching
1.5 cm in diameter. Tunnels are 15 to 25 cm long and penetrate directly
through the oak xylem. Larval tunnels cause lumber cut from infested
logs to be downgraded, reducing log value by up to 40 percent compared
to top quality wood.

Hosts Commonly Attacked This species attacks various oaks, including black (Quercus velutina
Lamb.), northern red (Q. rubra L.), and scarlet (Q. coccinea Muenchh.)
oaks.

Distribution Red oak borer is found throughout southern Canada and the eastern
United States.

Images of
Red Oak Borer

Figure 1. Adult of the red oak borer, Enaphalodes rufulus. (Jessica Lawrence,
Eurofins Agroscience Services, Bugwood.org)
A Photographic Guide 351

Images of Red Oak

Borer (continued)

Figure 2. Eggs of red oak borer. (University of Arkansas Forest Entomology Lab
Archive, University of Arkansas, Bugwood.org)

Figure 3. Young 1st-year red oak borer larva creating gallery in phloem.
(University of Arkansas Forest Entomology Lab Archive, University of Arkansas,
Bugwood.org)
352 Forest Pest Insects in North America:

Images of Red Oak Figure 4. Late stage red

oak borer larva in xylem
Borer (continued) gallery. (Fred Stephen,
University of Arkansas)

Figure 5. Gallery of red oak borer. (Fred Stephen, University of Arkansas)

Figure 6. Galleries of older red oak borer larvae in the heart wood, seen in cross
section. (University of Arkansas Forest Entomology Lab Archive, University of
Arkansas, Bugwood.org)
A Photographic Guide 353

Images of Red Oak

Borer (continued)

Figure 7. Emergence holes of red oak borer. (University of Arkansas Forest

Entomology Lab Archive, University of Arkansas, Bugwood.org)

Figure 8. Healed scars showing damage from red oak borer. (James Solomon,
USDA Forest Service, Bugwood.org)

Figure 9. Lumber cut from infested trees is downgraded because of the galleries
of red oak borer. (Joseph O’Brien, USDA Forest Service, Bugwood.org)
354 Forest Pest Insects in North America:

Images of Red Oak

Borer (continued)

Figure 10. View of trees killed by red oak borer. (Gerald J. Lenhard, Lousiana
State Univ, Bugwood.org)

Figure 11. View of

red oaks killed in the
Arkansas Ozarks during
outbreak, September
2000. (Fred Stephen,
University of Arkansas)

Important Biological Control Parasitoids of this species are mostly generalists on wood borers and are
Agents Related to this not well known. Woodpeckers are an important predator of red oak borer
Pest Species larvae.

Web Links for Information http://www.eppo.org/QUARANTINE/Alert_List/insects/enaphalodes_

on Red Oak Borer rufulus.htm; fact sheet of EPPO on red oak borer as a potential invasion
threat to Europe.

Articles Riggins, J.J., L.D. Galligan, and F.M. Stephen. 2009. Rise and fall of
red oak borer (Coleoptera: Cerambycidae) in the Ozark mountains of
Arkansas, USA. Florida Entomologist 92: 426-433.
Haavik, L.J. and F.M. Stephen. 2010. Historical dynamics of a native
cerambycid, Enaphalodes rufulus, in relation to climate in the Ozark and
Ouachita Mountains of Arkansas. Ecological Entomology 35: 673-683.
A Photographic Guide 355

97. Black Fir Sawyer Beetle, Monochamus urussovii (Fischer von

Waldheim) (Coleoptera: Cerambycidae)

Orientation to Pest Black fir sawyer beetle, Monochamus urussovii (Fischer von Waldheim),
is native to Eurasia and is not present in North America. Male black
fir sawyer beetles have antennae approximately twice as long as body,
whereas those of females are just a bit longer that the body. This beetle
attacks all species in the family Pinaceae. Firs (Abies) are especially
vulnerable to damage. It is the capability of M. urussovi to vector
phytopathogenic fungi that poses the greatest threat to North American
forests. Reproduction of this beetle occurs in weakened or dying trees
that have been affected by drought or fire, or have been wind thrown or
recently cut. Outbreaks can occur, however, in live, healthy trees. The
life cycle begins when beetles feed in the crown, removing strips of bark.
This feeding infects branches with the blue-stain fungus Leptographium
sibiricum Jacobs and Wingfield. Dieback of branches in the crown
weakens the tree and reduces resin flow. These changes make the
tree suitable for beetle oviposition and larval feeding. In the summer,
female beetles excavate oviposition niches, often in the branches, and
lay a single egg beneath the bark. Larvae dig sinuous galleries, up to
2.5 cm wide, under. These galleries are filled with frass. In the spring,
overwintered larvae dig into the sapwood and create a pupal chamber,
from which adults later emerge. As an infested host tree dies, its needles
turn yellow and then red.

Hosts Commonly Attacked This species is most damaging to fir (Abies), but larch (Larix), spruce
(Picea), pine (Pinus), and birch (Betula) species are also commonly
attacked.

Distribution This beetle occurs in Sweden, Norway, Finland, eastern Poland, Estonia,
Latvia, Lithuania, Belarus, Ukraine, Russia, Kazakhstan, Mongolia,
northeastern China, Korea, and northern Japan.

Images of Black Fir Figure 1. Adult of

black fir sawyer beetle,
Sawyer Beetle Monochamus urussovii.
(Vitaly Gumenuk,
Bugwood.org)
356 Forest Pest Insects in North America:

Images of Black Fir

Sawyer Beetle (continued)

Figure 2. The circular emergence holes of black fir sawyer beetles are large,
ca 1.2 cm in diameter. (Stanislaw Kinelski, Bugwood.org)

Figure 3. Brown foliage

is sign of attack by
black fir sawyer beetle.
(Stanislaw Kinelski,
Bugwood.org)
A Photographic Guide 357

Important Biological Control Some information is available in Russian language publications on the
Agents Related to this natural enemies of this species in Russia (see Isaev et al., 1988).
Pest Species

Web Links for Information http://www.inspection.gc.ca/english/plaveg/pestrava/monuru/tech/

on Black Fir Sawyer Beetle monurue.shtml; fact sheet of Canadian food inspection agency, proving
information on life history and damage.

Articles Vetrova, V.P., A.S. Isaev, N.V. Pashenova, and N.Yu Konstantinov. 1998.
Estimating the threat of a mass outbreak of Monochamus urussovii in
the dark coniferous stands of the Lower Angara region after damage by
Dendrolimus sibiricus. Lesovedenie 3: 58-67.
Jacobs, K., M.J. Wingfield, N.V. Pashenova, and V.P. Vetrova. 2000.
A new Leptographium species from Russia. Mycological Research 104:
1524-1529.
Isaev, A.S., T.M. Ovchinnikova, and V.G. Sukhovol’skii. 2001. Modeling
the population dynamics of the longhorn Monochamus urussovii in the
dark central-taiga forests of Siberia. Lesovedenie 4: 15-24.
Isaev, A.S., A.S. Rozhkov, and V.V. Kiselev. 1988. The fir longhorn
Monochamus urussovi. (Chernyi pikhtovyi usach Monochamus urussovi
[Fisch.]). Novosibirsk: Nauka, No. 270. (In Russian)
358 Forest Pest Insects in North America:

98. Japanese Pine Sawyer, Monochamus alternatus Hope (Coleoptera:

Cerambycidae)

Orientation to Pest Japanese pine sawyer, Monochamus alternatus Hope, is native to Asia,
principally China, Japan and Korea. In Asia, this species has proven
to be an important vector of an introduced pathogenic nematode, the
pine wood nematode, Bursaphelenchus xylophilus (Steiner and Buhrer)
Nickle, which kills Asian pines. Japanese pine sawyer is not yet present
in the United States. Japanese pine sawyers overwinter as large larvae
in galleries in infested trees. Pupation occurs at the end of winter, and
adults emerge in spring and feed on foliage of hosts (termed maturation
feeding). This feeding occurs on two-year old shoots of healthy trees.
This feeding is important because it provides locations for nematode
invasion of live healthy trees. Nematode larvae emerge from beetle
spiracles, fall onto damaged twigs, and penetrate the woody tissue
through the feeding wounds. Once mature, adult beetles lay their eggs
in trees that are stressed or have recently been killed by bark beetles,
lightening, disease, or other factors. Eggs are placed singly in slots in
bark cut by females. Each female may lay 100-200 eggs. Young larvae
feed in the cambium and mature larvae tunnel into the heartwood to
complete the cycle. There is one generation per year. Infestations damage
wood of recently dead and downed conifers, causing degradation and
loss of value for lumber.

The consequences of a potential introduction of Japanese pine sawyer

to the United States by itself may not have severe consequences for
two reasons. First, there are a number of other species of conifer-
infesting Monochamus beetles already present in North America (e.g.
M. caroliniensis, M. marmorator Kirby in Richardson, M. notatus
Drury, M. titillator Fabricius, M. scutellatus Say), all of which attack
recently killed or severely stressed trees. If introduced and established in
North America, Japanese pine sawyer would likely compete with these
species for the same set of hosts. Secondly, the pine wood nematode,
B. xylophilus, which is native to North America and is vectored by native
Monochamus, does not kill North American conifers. However, an
invasion of Japanese pine sawyer might also introduce an Asian species
of Bursaphelenchus, such as Bursaphelenchus mucronatus (Mamiya and
Enda) or Bursaphelenchus kolymensis Korentchenko, occur in Asia on
species of Abies, Larix and Pinus (the former species has been found in
Canada). These are not known to cause widespread mortality of their
native hosts, but may be more pathogenic to some North American
conifers, due to lack of evolutionary association and hence lack of
selection for resistance. Such an effect is predicted by the damage caused
to Asia conifers by an introduced American species of Bursaphelenchus.
Detection of invasion must be based on collection and identification of
adult beetles, as symptoms of infestation by M. alternatus are identical to
those caused by native North American Monochamus species.
A Photographic Guide 359

Hosts Commonly Attacked Japanese pine sawyer is known to attack seventeen species of pines
(Pinus), including, in Japan, Japanese red pine (Pinus densiflora Siebold
and Zucc.), Japanese black pine (P. thunbergii Parl.), luchu pine
(P. luchuensis Mayr), and, in China, Mason pine (P. massoniana
Lambert). It has also been reported to attack three species of spruce
(Picea spp.), and one species each of fir (Abies), true cedar (Cedrus),
and larch (Larix). Under laboratory conditions, fresh logs of the North
American slash pine (Pinus elliottii Engelmann) were preferred over
the native Japanese P. densiflora for oviposition, but larval survival
was lower.

Distribution Japanese pine sawyer is native to Asia and is found mainly in Japan,
China, and Korea.

Figure 1. World
distribution of
Japanese pine sawyer,
Monochamus alternatus.
(EPPO • EPPO Plant
Quarantine Data
Retrieval System [PQR])

Images of Japanese Figure 2. Adult of

Japanese pine sawyer
Pine Sawyer showing adult feeding
on needles. (Jijing Song
and Juan Shi, Beijing
Forestry University,
Bugwood.org)
360 Forest Pest Insects in North America:

Images of Japanese
Pine Sawyer (continued)

Figure 3. Various North

American species are
similar in appearance
and biology to Japanese
pine sawyer; for example,
Monochamus notatus
(Drury) (top), Monochamus
scuttelatus (Say) (middle),
and Monochamus
marmorator Kirby (bottom).
(Top: J. Tomminen,
University of Helsinki,
Bugwood.org; middle:
Laura Lazarus, North
Carolina Division of Forest
Resources, Bugwood.org;
bottom: Natasha Wright,
Florida Department of
Agriculture and Consumer
Services, Bugwood.org)
A Photographic Guide 361

Images of Japanese Figure 4. Adults of

Japanese pine sawyer
Pine Sawyer (continued) feed on foliage after
emergence to obtain
food to develop eggs.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)

Figure 5. Egg of
Japanese pine
sawyer (top), and bark
chewing at oviposition
site (bottom). (Top:
Jijing Song and Juan
Shi, Beijing Forestry
University, Bugwood.org;
bottom: USDA Forest
Service - North Central
Research Station
Archive, USDA Forest
Service, Bugwood.org)
362 Forest Pest Insects in North America:

Images of Japanese Figure 6. Larva of

Japanese pine sawyer.
Pine Sawyer (continued) (Jijing Song and Juan
Shi, Beijing Forestry
University, Bugwood.org)

Figure 7. Pupa of Japanese pine sawyer in the pupal chamber. The pupa is the
stage that attracts the pine wood nematode, which attacks the trachea system
of the pupa. (USDA Forest Service - Region 2 - Rocky Mountain Region Archive,
USDA Forest Service, Bugwood.org)

Figure 8. Tail of male

pine wood nematode,
Bursaphelenchus
xylophilus, showing
characteristic spicule.
(L.D. Dwinell, USDA
Forest Service,
Bugwood.org)
A Photographic Guide 363

Images of Japanese Figure 9. Exit hole of

Japanese pine sawyer
Pine Sawyer (continued) (top), and adult exiting
hole (bottom). (Top:
Jijing Song and Juan
Shi, Beijing Forestry
University, Bugwood.org;
bottom: USDA Forest
Service - Region 2 -
Rocky Mountain Region
Archive, USDA Forest
Service, Bugwood.org)

Figure 10. Mason pine (Pinus massoniana) in China killed by Japanese pine
sawyer. (Jijing Song and Juan Shi, Beijing Forestry University, Bugwood.org)
364 Forest Pest Insects in North America:

Important Biological Control In China, a recently recognized predator affecting Japanese pine
Agents Related to this sawyer is the Cryptalaus berus (Coleoptera: Elateridae), and in Japan,
Pest Species the predator Dastarcus helophoroides (Fairmaire) is believed to be an
important source of mortality.

Web Links for Information http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=77&langdispl

on Japanese Pine Sawyer ay=english; a web site of the USDA Forest Service, with extensive detail
on biology and risks.

Articles Kobayashi, F., A. Yamane, and T. Ikeda. 1984. The Japanese pine sawyer
beetle as the vector of pinewood nematode disease. Annual Review of
Entomology 29: 115-135.
Mota, M.M. and P. Vieira (eds.). 2008. Pine Wilt Disease: a Worldwide
Threat to Forest Ecosystems. Springer, New York.
A Photographic Guide 365

99. Asian Longhorned Beetle, Anoplophora glabripennis Motschulsky

(Coleoptera: Cerambycidae)

Orientation to Pest Asian longhorned beetle, Anoplophora glabripennis Motschulsky,

is native to Asia (principally China) and invasive in parts of western
Europe, and in several American cities, where eradication efforts are
underway. This borer attacks a variety of hardwood trees. In China,
Asian longhorned beetles are not damaging in forests, but because of
extensive planting of certain poplars (exotic varieties) that proved highly
susceptible to the species, the insect increased in abundance. This action
facilitated the beetle’s dissemination to other countries because infested
wood was used for packing material. The biology of the species is typical
of many longhorn beetles. Adults emerge over an extended period from
spring to fall, but especially in late June to early July. Adults remain
on or near their emergence tree and engage in maturation feeding on
leaves, petioles, and tender bark. Eggs are laid singly under the bark, in
egg sites chewed by females. Larvae feed in the cambium layer of the
tree and later into the heartwood. Larvae dig pupation chambers inside
the tree, which can be filled with frass. Adults emerge via large (1 cm
dia) round exit holes, which are a visible sign of infestation. In southern
Asia, a generation requires one year, but in northern areas, two years are
required. Generations may be overlapping. Unlike many cerambycids, A.
glabripennis attacks healthy trees as well as those under stress. Several
generations can develop within an individual tree, eventually killing it.

Hosts Commonly Attacked In China (the native range), the major hosts of A. glabripennis are
certain species of Populus, especially Populus nigra L., P. deltoides
W. Bartram ex Marshall, Populus x canadensis and the Chinese hybrid
P. dakhuanensis. Other important hosts are the willows Salix babylonica
L. and S. matsudana Koidzumi. Other species also recorded as hosts
in China include species of Acer, Alnus, Malus, Morus, Platanus,
Prunus, Pyrus, Robinia, Rosa, Sophora, and Ulmus. In urban areas in
North America invaded by this species, the hosts attacked by this beetle
have been species of maple (Acer negundo L., A. platanoides L.,
A. pseudoplatanus L., A. rubrum L., A. saccharinum L., and
A. saccharum Marshall) and Aesculus hippocastanum L. However, it has
also been found on a range of other hardwoods, including Liriodendron
tulipifera L., Morus alba L., Robinia pseudacacia L., and species of
Betula, Fraxinus, Populus, Salix, and Ulmus.

Distribution Asian longhorned beetle is native to Asia. It has been detected in parts of
Europe and in North America (Toronto, Canada, Illinois, the greater New
York City area, New Jersey, and Massachusetts).
366 Forest Pest Insects in North America:

Figure 1. Current
infestations (blue
dots, top map) of
Asian longhorned
beetle, Anoplophora
glabripennis, in the
United States; potential
areas at risk of
becoming infested (red,
on bottom map). (Top:
USDA Forest Service •
AFPE Online Mapping;
bottom: USDA APHIS)

Images of Asian Figure 2. Adult of Asian

longhorned beetle.
Longhorned Beetle (Michael Bohne,
Bugwood.org)
A Photographic Guide 367

Images of Asian
Longhorned Beetle
(continued)

Figure 3. The invasion

of North America by
Asian longhorned
beetle was likely caused
by the shipment of
massive quantities of
raw wooden packing
material from Asia (top).
To reduce risks of further
such invasions, wooden
packing material must
now be fumigated
(bottom) or heat treated
to kill larvae or other
stages in wood. (Both
photos: Larry R. Barber,
USDA Forest Service,
Bugwood.org)
368 Forest Pest Insects in North America:

Images of Asian Figure 4. Chewed bark

on twig, caused by
Longhorned Beetle maturation feeding of
(continued) adult Asian longhorned
beetles. (Dean
Morewood, Health
Canada, Bugwood.org)

Figure 5. Left, sign of oviposition (chewed pit) on the trunk; right, eggs of Asian
longhorned beetle. (Left: Kenneth R. Law, USDA APHIS PPQ, Bugwood.org;
right: Melody Keena, USDA Forest Service, Bugwood.org)

Figure 6. Mixed age

(top) and mature
(bottom) larvae of Asian
longhorned beetle.
(Top: Steven Katovich,
USDA Forest Service,
Bugwood.org; bottom:
Kenneth R. Law,
USDA APHIS PPQ,
Bugwood.org)
A Photographic Guide 369

Images of Asian
Longhorned Beetle
(continued)

Figure 7. Pupa of Asian longhorned borer: left, extracted; right, in a pupal

chamber, with frass. (Left: Michael Bohne, Bugwood.org; right: Kenneth R. Law,
USDA APHIS PPQ, Bugwood.org)

Figure 8. Larva feeding tunnels of Asian longhorned beetles in small trunk

(top), and tunnels as seen in cross section of cut branch (bottom). (Top: Larry
R. Barber, USDA Forest Service, Bugwood.org; bottom: E. Richard Hoebeke,
Cornell University, Bugwood.org)
370 Forest Pest Insects in North America:

Images of Asian
Longhorned Beetle
(continued)

Figure 9. Emergence holes of Asian longhorned beetles. (E. Richard Hoebeke,

Cornell University, Bugwood.org)

Figure 10. Dead

branches in tops of
trees are a sign of
Asian longhorned beetle
infestation. (Dennis
Haugen, USDA Forest
Service, Bugwood.org)
A Photographic Guide 371

Figure 11. Suvey for Asian longhorned beetle infestations are based on visual
detection of all signs of infestation; eradication efforts includes implementing
a regulated area (quarantine), conducting surveys of host trees, removing of
infested trees, and using treatment applications. (Top: Thomas B. Denholm,
New Jersey Department of Agriculture, Bugwood.org; bottom: USDA Agricultural
Research Service Archive, USDA Agricultural Research Service, Bugwood.org)

Important Biological Control Compared with other longhorned beetles, relatively few natural enemies
Agents Related to this of ALB have thus far been identified. Egg parasitoids are unknown.
Pest Species Larval parasitoids include Dastarcus longulus Sharp (Coleoptera:
Colydiidae), Scleroderma guani Xiao et Wu (Hymenoptera: Bethylidae),
Bullaea sp. (Diptera: Tachinidae), and Megarhyssa sp. (Hymenoptera:
Ichneumonidae). Pupal parasitoids include D. longulus, S. guani, and
Aprostocetus sp. (Hymenoptera: Eulophidae). Among these, D. longulus
and S. guani appeared to be the most promising for biological control
purposes (see websites mentioned on next page for further details).
372 Forest Pest Insects in North America:

Web Links for Information http://www.eppo.org/QUARANTINE/insects/Anoplophora_glabripennis/

on Asian Longhorned Beetle ANOLGL_ds.pdf; EPPO factsheet on Asian longhorned beetle as
European quarantine pest.
http://www.umassgreeninfo.org/fact_sheets/wood_attackers/asian_
longhorned_beetle.html.
http://www.massnrc.org/pests/pestFAQsheets/asianlonghorned.html;
Massachusetts fact sheet focusing on biology and recognition.
http://en.wikipedia.org/wiki/Asian_long-horned_beetle; Wikipedia
article.
http://www.uvm.edu/albeetle/research/biocontrol.html#NaturalEnemies;
discussion of natural enemies of Asian longhorn borer found in surveys
in China.
www.beetlebusters.info; USDA Asian longhorned beetle website.
www.aphis.usda.gov; eradication program information.

Articles Cavey, J.F., E.R. Hoebeke, S. Passoa, and S.W. Lingafelter. 1998.
A new exotic threat to North American hardwood forests: an Asian
longhorned beetle, Anoplophora glabripennis Motschulsky (Coleoptera:
Cerambycidae): I. Larval description and diagnosis. Proceedings of the
Entomological Society of Washington 100(2): 373-381.
Haack, R.A., K.R. Law, V.C. Mastro, H.S. Ossenbruggen, and B.J.
Raimo. 1997. New York’s battle with the Asian long-horned beetle.
Journal of Forestry 95(12): 11-15.
Li, W. and C. Wu. 1993. Integrated Management of Longhorn beetles
Damaging Poplar Trees. Beijing, China. Forest Press. 290 p.
Nowak, D.J., J.E. Pasek, R.O. Sequeira, D.E. Crane, and V.C. Mastro.
2001. Potential effect of Anoplophora glabripennis (Coleoptera:
Cerambycidae) on urban trees in the United States. Journal of Economic
Entomology 94(1): 116-122.
Hu, J.F., S. Angeli, S. Schuetz, Y.Q. Luo, and A.E. Hajek. 2009.
Ecology and management of exotic and endemic Asian longhorned beetle
Anoplophora glabripennis. Agricultural and Forest Entomology 11:
359-375.
A Photographic Guide 373

100. Poplar Borer, Saperda calcarata Say (Coleoptera: Cerambycidae)

Orientation to Pest Poplar borer, Saperda calcarata Say, is a native North American
insect found throughout Canada and the United States, attacking
poplars (Populus) and on occasionally willows (Salix). It can become
a significant pest of both eastern cottonwood (Populus deltoides Bartr.
Ex Marsh) in southern regions and quaking aspen (P. tremuloides
Michx.) further north. Overwintered adults emerge from June until
August. Females lay eggs in slits chewed into the bark, most often in
isolated, trees growing in full sun or in trees along stand edges where
they are more exposed to sunlight. Trees stressed by over-maturity,
grass competition, or poor site conditions are more likely to be attacked.
Larval development typically takes 3-4 years, but can vary from two to
five. Signs and symptoms of this pest include swollen areas on trunks
and larger branches, exit holes, and woodpecker excavations. Infested
trees may have varnish-like stains on the bark below the points of attack,
where reddish sap has oozed out. Fresh sap and coarse, fibrous frass are
good indicators of an active larval gallery. Infestations often do not kill
trees outright, but the trunk or branches may be weakened and break
in wind, ice, or snow storms. Small trees may be killed by girdling
from joining of several larval galleries. Surveys in northern states and
Canadian provinces have found attack rates on quaking aspen by poplar
borer to be quite high (>50 percent), especially in more open park-like
settings where trees are open-grown. Maintaining well-stocked stands
and avoiding poor-quality sites reduces the likelihood of infestation.
Removal of infested trees was not shown to be helpful in reducing
infestation levels since it resulted in lower stand density that encouraged
further attack. It is recommended that poplar stands be well-stocked and
clear cut at maturity.

Hosts Commonly Attacked Poplars (Populus) are the preferred hosts of this borer. Quaking aspen
(P. tremuloides) is favored in northern areas, cottonwood in southern
areas. Balsam poplar (P. balsamifera L.) is another favored host where
it occurs. Willow (Salix) has been reported as a host.

Distribution This species is found throughout Canada and the United States where
poplars are present.

Images of
Poplar Borer

Figure 1. Adults of poplar borer, Saperda calcarata. (Left: Whitney Cranshaw,

Colorado State University, Bugwood.org; right: James Solomon, USDA Forest
Service, Bugwood.org)
374 Forest Pest Insects in North America:

Images of
Poplar Borer (continued)

Figure 2. Larvae of
poplar borer in galleries
and pupa in pupal
chamber, surrounded
by characteristic coarse
fibrous frass (left); close
up of poplar borer larvae
(right). (Left: James
Solomon, USDA Forest
Service, Bugwood.org;
right: Whitney
Cranshaw, Colorado
State University,
Bugwood.org)

Figure 3. Sap leakage

and bark staining on
quaking aspen is a
sign of poplar borer
infestation. (Steven
Katovich, USDA Forest
Service, Bugwood.org)
A Photographic Guide 375

Images of Figure 4. Internal

appearance of poplar
Poplar Borer (continued) borer damage, with
bark cut open. (USDA
Forest Service - Ogden
Archive, USDA Forest
Service, Bugwood.org)

Figure 5. Frass being expelled from larval gallery of poplar borer. (Whitney
Cranshaw, Colorado State University, Bugwood.org)
376 Forest Pest Insects in North America:

Images of
Poplar Borer (continued)

Figure 6. Eastern cottonwood trees (Populus deltoides ssp. deltoides Bartram ex

Marsh.) broken due to tunneling of poplar borer. (James Solomon, USDA Forest
Service, Bugwood.org)

Important Biological Control Natural mortality of the poplar borer is frequently high and a variety
Agents Related to this of parasitoids have been reared from this species in various locations.
Pest Species Studies in Canada revealed that egg infertility, climate, excessive sap
flow, and natural enemies destroyed 81 percent of the population of
this borer. Woodpeckers can remove many late-stage larvae. However,
while descriptive work has been done in some locations, no fundamental
understanding exists of the importance of natural enemies as control
agents for this borer.

Web Links for Information http://www.colostate.edu/Dept/CoopExt/4dmg/Pests/popborer.htm; fact

on Poplar Borer sheet of the Colorado State University extension service.
http://wiki.bugwood.org/Archive:Borers/Saperda_calcarata; fact sheet of
Bugwood Wiki.

Articles Solomon, J.D. 1995. Guide to insect borers of North American broadleaf
trees and shrubs. Agricultural Handbook 706. Washington D.C.: U.S.
Department of Agriculture, Forest Service: 735 p. (Saperda calcarata,
pages 334-338).
Broberg, C.L. and J.H. Borden. 2005. Host preference by Saperda
calcarata Say (Coleoptera: Cerambycidae). Journal of the Entomological
Society of British Columbia 102: 27-34.
A Photographic Guide 377

102. Cottonwood Borer, Plectrodera scalator (Fabricius) (Coleoptera:

Cerambycidae)

Orientation to Pest Cottonwood borer, Plectrodera scalator (Fabricius), is a native insect

that attacks cottonwoods and poplars (Populus), willows (Salix), and
species of Platanus, Albizia, and Helianthus. Adults first emerge in late
spring and feed on the shoots of young trees. Mating and egg-laying
occur throughout the summer. Females dig away soil from the base of
the tree, where they cut a niche in the bark, and deposit one or several
eggs. Larvae feed in the bases and roots of live trees of all sizes. At first
larvae mine the inner bark, but by fall larvae bore into the wood. Two or
more growing seasons are required for larval development. In their last
year, the large larvae create tunnels at the base of the tree, where they
pupate the falling spring. New adults emerge by chewing through the
pupal chamber and digging their way out of the soil. Young trees may be
hollowed out, partially severed, or girdled near the root collar, causing
breakage. Greatest damage occurs to trees in plantations and nurseries,
and young stands on sandy soils.

Hosts Commonly Attacked This borer attacks cottonwoods and poplars (Populus), willows (Salix),
and species of Platanus, Albizia, and Helianthus.

Distribution The cottonwood borer is found New York to Montana and south to Texas.
Damage is greatest in the southern United States.

Images of Figure 1. Adult of

cottonwood borer,
Cottonwood Borer Plectrodera scalator.
(Top: Charles T. Bryson,
USDA Agricultural
Research Service,
Bugwood.org; bottom:
Donald Duerr, USDA
Forest Service,
Bugwood.org)
378 Forest Pest Insects in North America:

Images of Cottonwood Figure 2. Adult female

of cottonwood borer
Borer (continued) deposting an egg
below the soil line into
a young cottonwood
shoot. (James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 3. Larvae of cottonwood borer in root of eastern cottonwood (Populus

deltoides ssp. deltoides Bartram ex Marsh.) (left) and in stem of young tree
(right). (Both photos: James Solomon, USDA Forest Service, Bugwood.org)
A Photographic Guide 379

Images of Cottonwood Figure 4. Pupa of

cottonwood borer in
Borer (continued) cottonwood shoot.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 5. Two views of damage from cottonwood borer at base of cottonwood

rootstocks. (Both photos: James Solomon, USDA Forest Service, Bugwood.org)
380 Forest Pest Insects in North America:

Important Biological Control Little is known of the natural enemies of this species, except that
Agents Related to this cottonwood borer is sometimes parasitized by the sarcophagid fly
Pest Species Sarcophaga vericauda Hungerford, or parasitic wasps in the genera
Ibalia, Rhyssa, or Megarhyssa. It is also susceptible to applications of the
fungal biopesticide Beauveria bassiana.

Web Links for Information http://www.ento.okstate.edu/ddd/insects/cottonwoodborer.htm; fact sheet

on Cottonwood Borer of the Oklahoma State University.
http://www.forestpests.org/nursery/cwborers.html; fact sheet of the
USDA Forest Service.

Articles Solomon, J.D. 1980. Cottonwood borer (Plectrodera scalator) - a

guide to its biologv, damage, and control. Research Paper SO157. New
Orleans, LA, U.S. Department of Agriculture, Forest Service, Southern
Forest Experiment Station: 10 p.
Forschler, B.T. and G.L. Nordin. 1989. Impact of Beauveria bassiana on
the cottonwood borer Plectrodera scalator (Coleoptera: Cerambycidae),
in a commercial cottonwood nursery. Journal of Entomological Science
24(2): 186-190.
A Photographic Guide 381

103. Banded Hickory Borer, Knulliana cincta (Drury) (Coleoptera:

Cerambycidae)

Orientation to Pest Banded hickory borer, Knulliana cincta (Drury), is native to North
America and is found in the southern United States, west to Texas, and
in northern Mexico. Three subspecies of this beetle have been named:
Knulliana c. cincta (Drury), found in eastern North America west to
Texas; Knulliana cincta ochracea (Bates), found in the southeastern
United States, northeastern Mexico, and the Bahamas; and Knulliana
cincta sonorensis (Schaeffer), found from Texas to Arizona, and adjacent
regions in Mexico. This borer breeds in the dead branches and trunks of
many hardwoods. The adult is dark brown and 1.6-3.0 cm long. The body
is covered with gray hairs, and there are short sharp spines on the corners
of the thorax. Each wing cover has a yellow spot across it, giving the
appearance of a band across the back. Eggs are laid in summer beneath
the bark or directly on the wood of recently felled, dying, or dead trees.
The larvae feed under the bark during the remainder of the summer,
forming galleries in the wood and ejecting frass through openings in the
bark. During the fall and following summer larvae continue to feed in
the wood and pupate in the fall or spring between lumps of frass at the
end of the larval gallery. The life cycle most likely requires two years for
completion. Cordwood, logs, posts, and rustic work may be damaged by
this species. Adults are attracted to lights.

Hosts Commonly Attacked This borer attacks hickory (Carya), pecan (Carya illinoinensis
[Wangenh.] K. Koch), walnut (Juglans), oak (Quercus), eastern
hophornbeam (Ostrya virginiana [Mill.] K. Koch ), plum (Prunus), apple
(Malus), chestnut (Castanea), hackberry (Celtis), pear (Pyrus), willow
(Salix), mesquite (Prosopis), orange (Citrus), Sapindus, and Leucaena.
Hickory appears to be preferred.

Distribution This species is found in the United States from Virginia south to Florida
and west to Oklahoma and Texas, and in northern Mexico.

Images of Banded Figure 1. Adult banded

hickory borer, Knulliana
Hickory Borer
cincta. (Maury J. Heiman)
382 Forest Pest Insects in North America:

Images of Banded
Hickory Borer (continued)

Figure 2. Larva of banded hickory borer. (Lacy L. Hyche, Auburn University,

Bugwood.org)

Figure 3. Larval galleries of banded hickory borer. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Important Biological Control Little is known of the natural enemies of this species, except that it is
Agents Related to this sometimes parasitized by the ichneumonid wasp Labena grallator (Say).
Pest Species

Web Links for Information http://www.texasento.net/Knulliana.htm; a collector’s website about

on Banded Hickory Borer beetles from Texas.
http://www.insectsofwestvirginia.net/b/knulliana-cincta.html.

Articles None
A Photographic Guide 383

104. Locust Borer, Megacyllene robiniae (Forster) (Coleoptera:

Cerambycidae)

Orientation to Pest Locust borer, Megacyllene robiniae (Forster), is native to the United
States and bores in black locust (Robinia pseudoacacia L.) trunks.
Attacks occur most often in young trees or in older trees stressed by
drought or poor soil, or that have been overtopped by other trees. This
borer formerly was limited to its native range in Pennsylvania, south in
the Appalachian Mountains to Georgia and in the Ozark Mountains of
Arkansas and Missouri. However, black locust has been widely planted
outside this range and the borer is now found in those locations as well.
Black locusts have been widely planted for restoration of strip-mined
areas. Adults of locust borer emerge in late summer and are commonly
seen feeding on pollen on field goldenrod (Solidago canadensis L.).
Females lay their eggs in crevices of rough bark and around wounds of
living trees. Larvae bore into the bark and construct overwintering cells.
In the following spring, larval activity is resumed and sap may be seen
oozing from the larval entry holes. Larvae bore into the wood and feed
until they mature in mid-summer. During their feeding period, larvae
tunnel extensively throughout the heartwood. Maturing larvae construct
a tunnel to the exterior, through which adults later emerge. There is one
generation per year. Attack is most common on stressed trees, such as
those on reclaimed mining sites. Borer activity often results in wind
breakage of trees, followed by excessive branching. Attack rates increase
as the proportion of black locust in the stand increases, and larval to adult
survivorship increases as the attack rate increases.

Hosts Commonly Attacked Locust borer is a specialist that feeds only on black locust
(R. pseudoacacia).

Distribution Locust borer is found throughout the United States wherever black locust
grows naturally or has been introduced.

Images of Figure 1. Adult locust

borer, Megacyllene
Locust Borer robiniae. (Jon Yuschock,
Bugwood.org)
384 Forest Pest Insects in North America:

Images of Locust Figure 2. Locust borer

feeding on pollen of
Borer (continued) goldenrod (Solidago
sp.). (David Cappaert,
Michigan State University,
Bugwood.org)

Figure 3. Larva of locust borer in feeding gallery. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Figure 4. Pupa of locust borer in pupation chamber. (Lacy L. Hyche, Auburn

University, Bugwood.org)
A Photographic Guide 385

Images of Locust
Borer (continued)

Figure 5. Cracks, frass at base of tree, and emergence holes are signs of of
locust borer activity. (Left: James Solomon, USDA Forest Service, Bugwood.org;
right: Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 6. Locust borer larval galleries seen in cross and lateral sections.
(James Solomon, USDA Forest Service, Bugwood.org)

Important Biological Control Two species of wasps, Dolichomitus irritator (Fabricius) and
Agents Related to this Habrolepoidea depressa (Girault), have been recorded parasitizing locust
Pest Species borer larvae at low rates (<1 percent). Natural enemies of locust borer
have not been studied extensively.
386 Forest Pest Insects in North America:

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/locust/locust.htm; USDA Forest

on Locust Borer Service leaflet; extensive information on the biology, life cycle, and
damage of the insect.
http://www.treesforyou.org/Planting/TreeCare/Healthy/Locust-B.htm;
Washington state extension service leaflet.
http://www.dec.ny.gov/docs/lands_forests_pdf/locustb.pdf; extension
article of the New York Department of Environmental Conservation.
http://www.ces.ncsu.edu/depts/ent/notes/O&T/trees/note18/note18.html;
fact sheet of North Carolina State University Entomology Department;
information on chemical control provided.

Articles Harman, D.M., M.A. van Tyne, and W.A. Thompson. 1985. Comparison
of locust borer Megacyllene robiniae Forster (Coleoptera: Cerambycidae)
attacks on coal strip-mined lands and lands not mined. Annals of the
Entomological Society of America 78: 50-53.
Harman, D.M., P. Rudolf, and K.R. Dixon. 1985. Influence of stand
composition on locust borer (Coleoptera: Cerambycidae) attack rates.
Journal of Entomological Science 20: 207-211.
Echaves, V.D., D.M. Harman, and A.L. Harman. 1998. Site quality in
relation to damage by locust borer, Megacyllene robiniae Forster in black
locust. Journal of Entomological Science 33: 106-112.
A Photographic Guide 387

105. White Oak Borer, Goes tigrinus (De Geer) (Coleoptera: Cerambycidae)

Orientation to Pest White oak borer, Goes tigrinus (De Geer), is native to North America
and is found in much of the eastern United States in association with
white (Quercus alba L.) and overcup (Q. lyrata Walter) oaks. Damage
is greatest to younger trees and losses from this species are important
in oak stands grown for veneer or cooperage (barrels). Reproduction is
often concentrated in selected trees (brood trees) and removing brood
trees can help control the pest at the stand level. Adult beetles emerge in
May and June. Eggs are deposited in niches 6 mm in diameter that are
cut singly into the bark. As the larvae tunnel directly into the wood,
sap often oozes from the entrance holes in the tree and fine moist frass
is pushed out of the tunnels to the outside. Larval galleries are about
12 mm in diameter and 15 cm long. Pupation occurs within the gallery.
One generation takes about 3 to 5 years.

Hosts Commonly Attacked White oak borer breeds in white (Q. alba) and overcup (Q. lyrata) oaks.

Distribution White oak borer is found in the eastern United States from New York to
Florida and west to Michigan and Louisiana.

Images of Figure 1. Adult white

oak borer, Goes tigrinus.
White Oak Borer (James B. Hanson,
USDA Forest Service,
Bugwood.org)
388 Forest Pest Insects in North America:

Images of White Figure 2. Egg of white

oak borer, exposed
Oak Borer (continued) by removal of bark
flap. (James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 3. Larva of white

oak borer. (USDA Forest
Service Fact Sheet)

Figure 4. Signs of of white oak borer activity: (left) frass pushed out of borer hole
in tree trunk; (right) emergence holes in tree trunk. (Left: James Solomon, USDA
Forest Service, Bugwood.org; right: USDA Forest Service Fact Sheet)
A Photographic Guide 389

Important Biological Control Natural enemies of white oak borer have not been studied.
Agents Related to this
Pest Species
Web Links for Information http://www.fs.fed.us/r8/foresthealth/pubs/oakpests/p19.html; USDA
on White Oak Borer Forest Service factsheet covering biology and control.

Articles Donley, D.E. 1978. Distribution of the white oak borer Goes tigrinus
Degeer (Coleoptera: Cerambycidae) in a mixed oak stand. In: Pope, P.E.
(ed.). Central hardwood forest Conference II. Proceedings of a meeting
held at Purdue University at West Lafayette, Indiana, November 14-16,
1978. West Lafayette, Indiana, USA. Department of Forestry and Natural
Resources, Purdue University: 529-539.
Solomon, J.D. and D.E. Donley. 1983. Bionomics and control of the
white oak borer. Research Paper SO-198, Southern Forest Experiment
Station, USDA Forest Service.
390 Forest Pest Insects in North America:

106. Whitespotted Sawyer, Monochamus scutellatus (Say) (Coleoptera:

Cerambycidae)

Orientation to Pest Whitespotted sawyer, Monochamus scutellatus (Say), is a native North

American borer that attacks dying or recently dead conifers that have
dried out somewhat but have not been heavily attacked by other wood-
feeding insects. There are two recognized subspecies, M. scutellatus
scutellatus and M. scutellatus oregonensis. Females lay their eggs in slits
chewed in the bark near branch scars or rough areas of bark. When eggs
hatch, young larvae bore through the phloem into the cambium. Young
larvae mine beneath the bark. Later instars tunnel toward the heartwood.
Prior to pupation, the larva turns its tunnel toward the surface, where
it pupates behind a chip plug. There is one generation per year in the
northern part of its range and two in the southern portion. Logs left in
the forest untended over a summer season are especially suitable for
attack by this borer. This sawyer beetle is also able to vector the parasitic
pinewood nematode, Bursaphelenchus xylophilus (Steiner and Buhrer)
Nickle, which causes pine-wilt disease in Minnesota and Wisconsin.
Damage to logs can be reduced by cutting timber in fall or winter and
removing logs from woods before summer. Pulpwood logs can be
partially protected by piling them in the shade or covering them with
slash.

Hosts Commonly Attacked The whitespotted sawyer attacks a variety of conifers but prefers eastern
white pine (Pinus strobus L.). This beetle will also attack jack pine
(P. banksiana Lamb.), red pine (P. resinosa Ait.), balsam fir (Abies
balsamea [L.] Mill.), white spruce (Picea glauca (Moench) Voss), black
spruce (P. mariana (Mill.) B.S.P.), and red spruce (P. rubens Sarg.).
Tamarack (Larix laricina (DuRoi) K. Koch) occasionally serves as a
host.

Distribution Whitespotted sawyer is found in the United States, north of North

Carolina and westward through the Great Lakes states, across Canada
and north to Alaska.
A Photographic Guide 391

Images of
Whitespotted Sawyer

Figure 1. Adult whitespotted sawyer, Monochamus scutellatus; right, adult

feeding on twigs. (Left: Natasha Wright, Florida Department of Agriculture and
Consumer Services, Bugwood.org; right: Joseph Berger, Bugwood.org)

Figure 2. Adult emergence hole (with head of adult visible) of whitespotted sawyer.
(Laura Lazarus, North Carolina Division of Forest Resources, Bugwood.org)

Figure 3. Flagging of branches killed by adult feeding of whitespotted sawyer

beetles. (Ronald S. Kelley, Vermont Department of Forests, Parks and
Recreation, Bugwood.org)
392 Forest Pest Insects in North America:

Important Biological Control Natural enemies of whitespotted sawyer include tachinids in the genus
Agents Related to this Eutheresia and ichneumonids in the genera Rhyssa and Dolichomitus,
Pest Species but their actual influence on population levels of whitespotted sawyer is
unknown.

Web Links for Information http://wiki.bugwood.org/HPIPM:Pine_Sawyers; biology of pine sawyers;

on Whitespotted Sawyer Bugwood Wiki fact sheet.
http://www.forestpests.org/vermont/whitespottedsawyer.html; forestry
fact sheet from Vermont.
http://www.na.fs.fed.us/spfo/pubs/fidls/ws_sawyer/ws_sawyer.htm;
USDA Forest Service fact sheet on biology and control.

Articles Raske, A.G. 1973. Relationship between felling date and larval density of
Monochamus scutellatus. Bi-monthly Research Notes 29(4): 23-24.
Wingfield, M.J. and R.A. Blanchette. 1983. The pine-wood nematode,
Bursaphelenchus xylophilus, in Minnesota and Wisconsin: insect
associates and transmission studies. Canadian Journal of Forest
Research 13(6): 1068-1076.
A Photographic Guide 393

107. Bagworm, Thyridopteryx ephemeraeformis (Haworth) (Lepidoptera:

Psychidae)

Orientation to Pest Bagworm, Thyridopteryx ephemeraeformis (Haworth), is native to North

America, and larvae defoliate arborvitae (Thuja occidentalis L.), juniper
(Juniperus virginiana L.), eastern white pine (Pinus strobus L.), and
blue spruce (Picea pungens Engelm.), on which they are often abundant.
Male moths have wings but females are wingless and are unable to
move. Males and females emerge in the fall and males fly to females.
Females never emerge from their larval shelters (the “bag”), and mate
while still in their bags. Females then develop their eggs, which are laid
inside the bag and these are the overwintering stage. Eggs hatch in spring
and some larvae spin silk threads and are blown by the wind. Upon
alighting on plants, neonate larvae accept or reject plants, and if the
latter spin a new thread and repeat the dispersal process. Many larvae,
however, skip this dispersal phase and settle on their natal plants. Once
a larva has accepted a plant, it constructs a bag and drags it around as
it feeds. Mature larvae attach their bags to the plant and pupate inside
them. There is one generation per year. Bagworms can be locally dense
on ornamental plantings, causing defoliation and killing trees or shrubs.
Damage to forest stands is not usually important, because larvae prefer
sunny sites. Some damage does, however, occur in Atlantic white cedar
(Chamaecyparis thyoides [L.] Britton, Sterns and Poggenb.) and black
locust (Robinia pseudoacacia L.) stands, especially in the southern
United States.

Hosts Commonly Attacked Bagworm commonly attack arborvitae (T. occidentalis), juniper (J.
virginiana), eastern white pine (P. strobes), blue spruce (P. pungens),
Atlantic white cedar (C. thyoides), and black locust (R. pseudoacacia).
On other hosts, bagworm larvae are rare.

Distribution Bagworm is widely distributed in the eastern United States.

Images of Bagworm Figure 1. Adult

male bagworm
moth, Thyridopteryx
ephemeraeformis.
(Pennsylvania
Department of
Conservation and
Natural Resources
- Forestry Archive,
Bugwood.org)
394 Forest Pest Insects in North America:

Images of Bagworm Figure 2. Bagworm

eggs dissected out of
(continued) female inside case.
(Lacy L. Hyche, Auburn
University, Bugwood.org)

Figure 3. Young
bagworm in small
case (top); older
bagworm with larger,
but still mobile, case
(bottom). (Top: Rayanne
Lehman, Pennsylvania
Department of
Agriculture,
Bugwood.org; bottom:
Connecticut Agricultural
Experiment Station
Archive, Connecticut
Agricultural Experiment
Station, Bugwood.org)
A Photographic Guide 395

Images of Bagworm Figure 4. Bagworm

caterpillar partialy
(continued) extended outside of
protective bag (top),
and caterpillar pulled
out of bag (bottom).
(Both photos:
Gerald J. Lenhard,
Louiana State University,
Bugwood.org)

Figure 5. Bagworm prepupa (bottom) and pupa (top) in bags cut open for
viewing. (Gerald J. Lenhard, Louiana State University, Bugwood.org)
396 Forest Pest Insects in North America:

Images of Bagworm
(continued)

Figure 6. Male bagworm mating with flightless female inside her larval bag.
(Jerry A. Payne, USDA Agricultural Research Service, Bugwood.org)

Figure 7. Bag of mature bagworm, fixed in place (left); many bagworm cases
(right) on heavily infested plant. (Left: John H. Ghent, USDA Forest Service,
Bugwood.org; right: Pennsylvania Department of Conservation and Natural
Resources - Forestry Archive, Bugwood.org)
A Photographic Guide 397

Images of Bagworm Figure 8. High densities

of bagworm bags
(continued) on partly defoliated
conifer. (Pennsylvania
Department of
Conservation and
Natural Resources
- Forestry Archive,
Bugwood.org)

Important Biological Control Natural enemies of bagworms have been frequently studied because
Agents Related to this the bags are large and stationary, inviting their examination. The
Pest Species ichneumonid Itoplectis conquisitor (Say), a generalist pupal parasitoid,
is commonly found attacking bagworms and is often the most abundant
species reared from this host. Many other species of parasitoids have also
been reared from bagworms.

Web Links for Information http://ento.psu.edu/extension/factsheets/bagworm; fact sheet of

on Bagworm Pennsylvania State University.
http://www.uark.edu/ua/arthmuse/bagworm.html; University of Arkansas
fact sheet.
http://njaes.rutgers.edu/pubs/publication.asp?pid=FS1144; fact sheet of
New Jersey Rutgers NJAES Cooperative Extension.

Articles Horn, D.J. and R.F. Sheppard. 1979. Sex ratio, pupal parasitism, and
predation in two declining populations of the bagworm, Thyridopteryx
ephemeraeformis (Haworth) (Lepidoptera: Psychidae). Ecological
Entomology 4: 259-265.
Moore, R.G. and L.M. Hanks. 2004. Aerial dispersal and host plant
selection by neonate Thyridopteryx ephemeraeformis (Lepidoptera:
Psychidae). Ecological Entomology 29: 327-335.
398 Forest Pest Insects in North America:

108. Birch Casebearer, Coleophora serratella (L.) (Lepidoptera:

Coleophoridae)

Orientation to Pest Birch casebearer, Coleophora serratella (L.), is a European moth that is
invasive in North America and is associated with birches (Betula), alders
(Alnus), and elms (Ulmus). High densities sometimes defoliate birches,
and this insect is considered one of the most important pests of birch in
some parts of Canada such as Newfoundland. Because of misapplication
of names, literature records from 1938 to 1972 under this name very
likely do not refer to the true C. serratella as described here. The adult
appears in late spring. Eggs are laid on host leaves and the larva feeds
by inserting its head into small mines that it creates. The larva constructs
a protective case, which it drags with it as it moves around to feed.
Pupation occurs in early summer, and occurs in the larval case, which is
fixed to the upper surface of a leaf in a sunny situation.

Hosts Commonly Attacked Birch casebearer feeds on birches (Betula), alders (Alnus), and elms
(Ulmus).

Distribution Birch casebearer is found in the northeastern United States, eastern

Canada from Newfoundland through southern Ontario into Manitoba,
and in western Canada, in British Columbia.

Images of Figure 1. Adult birch

casebearer, Coleophora
Birch Casebearer serratella. (Ian Kimber)

Figure 2. Several
larvae (in cases) of
birch casebearer on
young white birch
leaf. (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)
A Photographic Guide 399

Images of Birch Figure 3. Larva of birch

casebearer, with the
Casebearer (continued) case removed. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)

Figure 4. White birch

defoliated by birch
casebearer. (Claude
Monnier - Natural
Resources Canada,
Canadian Forest
Service)

Important Biological Control Natural enemies of birch casebearer have been studied in Quebec and
Agents Related to this Newfoundland, and as many as 38 species of parasitoids attack this
Pest Species invasive species there. In the native range (central Europe) 25 species
are reported associated with this case bearer. Some of these species
(e.g., Campoplex borealis [Zett.] and Apanteles coleophorae [Wilk.],
Apanteles mesoxanthus Ruschka, and Apanteles cornvinus Reinh.)
have been released in North America, but there are no records of their
establishment.

Web Links for Information http://www.exoticpests.gc.ca/ins_details_eng.asp?pestType=ins&lang=en

on Birch Casebearer &geID=8229; Canadian factsheet on forest invasive alien species.

Articles Pschorn-Walcher, H. 1980. Population fluctuations and parasitisation

of the birch-alder casebearer (Coleophora serratella L.) in relation to
habitat diversity. Zeitschrift für Angewandte Entomologie 89(1): 63-81.
(In German).
Guevremont, H. and J. Juillet. 1975. Parasites of the birch casebearer,
Coleophora fuscedinella [= Coleophora serratella] Zeller (Lepidoptera:
Coleophoridae), in the Sherbrooke region, Quebec. Phytoprotection
56 (1): 1-17. (In French).
400 Forest Pest Insects in North America:

109. Larch Casebearer, Coleophora laricella (Hübner) (Lepidoptera:

Coleophoridae)

Orientation to Pest The larch casebearer, Coleophora laricella [Hübner]), is an invasive

species from Europe that caused extensive defoliation of North American
larch before it was brought under biological control. It has one generation
per year. Adults emerge in late spring and deposit their eggs singly on
needles in early summer. Newly hatched larva bore into needles and feed
as needle miners. After larvae consume a portion of their first needle,
a section of a mined-out needle is lined with silk, cut free, and carried
as a case. Larvae in cases then move to other needles and continue to
feed. Fed-on needles are only partly consumed. They remain attached
to branches but turn brown, making damage readily visible. Cases are
enlarged as larvae grow. Larvae overwinter in cases fixed to branches
and resume feeding in spring on new foliage. Upon reaching maturity,
larvae pupate in their cases from which moths later emerge.

Hosts Commonly Attacked In North America, this moth attacks both eastern (Larix laricina [Du Roi]
K. Koch) and western (Larix occidentalis Nuttl.) larch.

Distribution Larch casebearer occurs in Canada from the Maritmes to western Canada
and in the United States from New England to Minnesota and, separately,
in Washington, Oregon, Idaho and western Montana. This distribution
corresponds to that of eastern and western larch.

Images of
Larch Casebearer

Figure 1. Adults of larch casebearer (Coleophora laricella) mating. (Roger Ryan,

USFS PNW Station, Bugwood.org)

Figure 2. Young larva of larch casebearer in needle-mining phase. (Roger Ryan,

USFS PNW Station, Bugwood.org)
A Photographic Guide 401

Images of Larch
Casebearer (continued)

Figure 3. Older larva of larch casebearer with case. (Roger Ryan, USFS PNW
Station, Bugwood.org)

Figure 4. View of larch

casebearer larva with
case partially removed.
(Victor Ryabincov,
Bugwood.org)

Figure 5. Close view

of damage to needles
caused by larch
casebearer larval
feeding. (Ferenc
Lakatos, University
of West-Hungary,
Bugwood.org)
402 Forest Pest Insects in North America:

Images of Larch Figure 6. Branch

showing yellow foliage
Casebearer (continued) that has been partly
consumed by larch
casebearer. (Roger
Ryan, USFS PNW
Station, Bugwood.org)

Figure 7. Larch stand showing reddish-brown foliage due to feeding of larch

casebearer. (William M. Ciesla, Forest Health Management International,
Bugwood.org)

Figure 8. Cluster of overwintering larch casebearer in cases at tip of twig.

(Roger Ryan, USFS PNW Station, Bugwood.org)
A Photographic Guide 403

Images of Larch
Casebearer (continued)

Figure 9. The braconid Agathis pumila (Ratzburg), one of two key species giving
successful biological control of larch casebearer in North America. (Roger Ryan,
USFS PNW Station, Bugwood.org)

Figure 10. The eulophid Chrysocharis laricinellae Ratzburg, showing egg-laying

behavior. (Roger Ryan, USFS PNW Station, Bugwood.org)

Important Biological Control While a larger number of parasitoids were introduced, two species—the
Agents Related to this braconid Agathis pumila (Ratzburg) and the eulophid Chrysocharis
Pest Species laricinellae Ratzburg—were responsible for effective biological control
of larch casebearer in North America.
404 Forest Pest Insects in North America:

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/larch/larch.htm; USDA Forest

on Larch Casebearer Service Leaflet with information on biology, life history, damage, and
control.
http://www.fs.fed.us/r1-r4/spf/fhp/mgt_guide/larch_casebearer/larch_
casebearer.pdf; USDA Forest Service bulletin.
http://www.fs.fed.us/r1-r4/spf/fhp/field_guide/124lrchcb.htm.
http://www.dec.ny.gov/docs/lands_forests_pdf/cbearer.pdf.
http://www.oregon.gov/ODF/privateforests/docs/fh/larchcasebearer.
pdf?ga=t.

Articles Jagsch, A.1973. Population dynamics and parasite complex of the

larch miner, Coleophora laricella Hb., in the natural distribution area
of European larch, Larix decidua Mill. Zeitschrift für Angewandte
Entomologie 73: 1-42.
Niwa, G.G. 1983. Environmental factors influencing the effectiveness
of Agathis pumila Ratz. (Hymenoptera: Braconidae), an introduced
parasite of the larch casebearer, Coleophora laricella Hbn. (Lepidoptera:
Coleophoridae). Forestry Abstracts 44(11): 718.
Niwa, C.C., R.W. Stark, D.G. Burnell, and D.M.J. Knox. 1986.
Annotated bibliography of larch casebearer (Coleophora laricella)
parasitoids. Bulletin No. 41. Forest, Wildlife and Range Experiment
Station, University of Idaho: 135 p.
Ryan, R.B. 1990. Evaluation of biological control: introduced
parasites of larch casebearer (Lepidoptera: Coleophoridae) in Oregon.
Environmental Entomology 19: 1873-1881.
Ryan, R.B. 1997. Before and after evaluation of biological control of the
larch casebearer (Lepidoptera: Coleophoridae) in the Blue Mountains
of Oregon and Washington, 1972-1995. Environmental Entomology 26:
703-715.
A Photographic Guide 405

110. Pecan Cigar Casebearer, Coleophora laticornella (Clemens)

(Lepidoptera: Coleophoridae)

Orientation to Pest The pecan cigar casebearer, Coleophora laticornella (Clemens), is a

case-bearing moth native to North America. Its hosts include pecan
(Carya illinoinensis [Wangenh] K. Koch), walnut (Juglans), and several
species of hickory (Carya). It often causes serious damage to these trees.
This insect overwinters as a partially grown larva in a case about 6 mm
long that is attached to a limb or twig. In the spring, the larvae resume
feeding on the newly opened buds and young foliage and when mature,
larvae pupate within their cases. Moths emerge and lay eggs on leaves.
Young larvae feed as leafminers, but later construct cases in which they
remain as they move about, feeding on leaves and, the following year,
buds. After leaf drop, larvae move back onto twigs or limbs, where they
attach their cases for overwintering. There may be several generations
per year. The pecan cigar casebearer occurs from New England south to
Florida and as far west as Texas.

Hosts Commonly Attacked Hosts of pecan cigar casebearer include pecan (Carya illinoinensis
[Wangenh] K. Koch), walnut (Juglans), and several species of hickory
(Carya).

Distribution This moth is found in the eastern United States, from New England south
to Florida and west to Texas.

Images of Pecan Figure 1. Adult pecan

cigar casebearer,
Cigar Casebearer Coleophora laticornella.
(Mark Dreiling)
406 Forest Pest Insects in North America:

Images of Pecan
Cigar Casebearer
(continued)

Figure 2. Larva of pecan cigar casebearer in case. (H.C. Ellis, University of

Georgia, Bugwood.org)

Important Biological Control None

Agents Related to this
Pest Species
Web Links for Information None
on Pecan Cigar Casebearer

Articles None
A Photographic Guide 407

111. Oak Skeletonizer, Bucculatrix ainsliella Murtfeldt (Lepidoptera:

Bucculatricidae)

Orientation to Pest The oak skeletonizer, Bucculatrix ainsliella Murtfeldt, is a native

North American moth whose larvae skeletonize the leaves of some
oaks (Quercus). Eggs are laid on the upper side of leaves, adjacent to
a major vein. The first instar larvae are leafminers, forming a strongly
contorted serpentine mine about 1 cm long. Older larvae feed externally
on the lower leaf surfaces, causing “windows” in leaves or skeletonizing
them. There are two generations per year. Adults are active in April and
May (generation one) and again in July and August (generation two).
In the southern United States fly into September. The oak skeletonizer
overwinters as a pupa in a white cocoon about 3 mm in length.
Occasionally outbreaks occur over large areas.

Hosts Commonly Attacked Oak skeletonizer feeds on red (Quercus rubra L.) and black (Q. velutina
Lam.) oaks. Records on chestnut oak (Q. montana Willdenow) and
chestnut (Castanea) are unconfirmed.

Distribution The moth occurs in southern Canada and in the United States from Maine
to North Carolina and west to Mississippi. It has also been introduced
into British Columbia and, since 2006, into western Europe.

Images of
Oak Skeletonizer

Figure 1. Mine of first instar larva of oak skeletonizer, Bucculatrix ainsliella. (Erik
J. van Nieukerken, Netherlands Centre for Biodiversity Naturalis, Bugwood.org)
408 Forest Pest Insects in North America:

Images of
Oak Skeletonizer
(continued)

Figure 2. Oak skeletonizer larva feeding on the underside of an oak leaf.

(G. Keith Douce, University of Georgia, Bugwood.org)

Figure 3. Oak skeletonizer cocoons attached to leaf veins; see also larvae
and leaf damage (brown areas). (James Solomon, USDA Forest Service,
Bugwood.org)
A Photographic Guide 409

Images of Figure 4. Leaf damage

from larval feeding of
Oak Skeletonizer oak skeletonizer.
(continued) (James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 5. Damage to
an oak tree due to
larval feeding of oak
skeletonizer.
(Terry S. Price, Georgia
Forestry Commission,
Bugwood.org)
410 Forest Pest Insects in North America:

Images of
Oak Skeletonizer
(continued)

Figure 6. Damage to a stand of oak trees due to the oak skeletonizer.

(Richard Jernigan, Georgia Forestry Commission, Bugwood.org)

Important Biological Control Pupal parasitism (either by Ichneumonidae or Chalcidoidea) reached

Agents Related to this high levels (>40 percent parasitism) during the first generation of a
Pest Species population outbreak in British Colombia (see Gelok et al. 1998), but
larval parasitism was very low.

Web Links for Information http://www.oakwilt.com/skeletonizer/index.html; website of Texas

on Oak Skeletonizer private nursery company.
http://www.forestpests.org/vermont/oakskeletonizer.html; fact sheet from
Vermont Department of Forests, Parks and Recreation on lifecycle and
management.
http://www.fs.fed.us/r8/foresthealth/pubs/oakpests/p13.html; USDA
Forest Service fact sheet on biology and control.
http://www.microlepidoptera.nl/nieuws/art_2011.7.11.php;
Microlepidoptera of the Netherlands: first report on its introduction into
Europe.

Articles Gelok, E., R. McGregor, D. Henderson, and L. Poirier. 1998. Seasonal

occurrence and parasitism of Bucculatrix ainsliella (Lepidoptera:
Lyonetiidae) on Quercus rubra in Burnaby, British Columbia. Journal of
the Entomological Society of British Columbia 95: 111-116.
Gibbons, C.F. and J.W. Butcher. 1961. The oak skeletonizer, Bucculatrix
ainsliella, in a Michigan woodlot. Journal of Economic Entomology 54:
681-684.
A Photographic Guide 411

112. Palm Leaf Skeletonizer, Homaledra sabalella (Chambers) (Lepidoptera:

Coleophoridae)

Orientation to Pest The palm leaf skeletonizer, Homaledra sabalella (Chambers), is a native
moth that is an important pest of many species of palms in the southern
United States. It is an important pest in Florida, where it sometimes
causes serious damage to palm trees. An outbreak of what appeared to
be this species occurred on coconut palms (Cocos nucifera L.) in Florida
starting in about 1995 but curiously did not affect cabbage palmetto.
While identical in appearance to the cabbage palmetto-feeding moth, the
coconut palm-feeding population is perhaps an invasive cryptic species.
The matter is under investigation. The following notes on life history are
for the traditional species (the population found on cabbage palmetto).
Female moths lay batches of, on average, 36 eggs glued to the surface of
older palm leaves and covered with brown, papery material. The larvae
mine in groups on both the upper and lower surfaces of older leaves,
under webs of silk. Excrement from the larvae is deposited on top of
the silk. Pupation takes place in the larval mines. There may be up to
five generations per year, and there is no diapause. Cutting and burning
infested leaves is an effective method of control.

Hosts Commonly Attacked Palm leaf skeletonizer feeds on several species of palms, including
Canary Island date palm (Phoenix canariensis [Chabaud]) and cabbage
palmetto (Sabal palmetto [Walter] Schultes and Schultes), the latter
being an important native species in Florida. In Puerto Rico, the principal
palm attacked is the Puerto Rico hat palm, Sabal causiarum (O.F. Cook)
Becarri. This species also feeds on Latan palms (Latania spp.) and to a
lesser extent on Washingtonia robusta H.Wendl.

Distribution This insect is found across the southern United States from Florida to
California, this being the northern extension of its range, which lies
principally in Cuba, Puerto Rico, and Hispaniola.

Images of Palm Leaf Figure 1. Adult palm leaf

skeletonizer, Homaledra
Skeletonizer sabalella. (Bo Zaremba)
412 Forest Pest Insects in North America:

Images of Palm Leaf

Skeletonizer (continued)

Figure 2. Damage from feeding of larvae of the palm leaf skeletonizer.

(Tim Borschat, Bugwood.org)

Important Biological Control An undescribed tachinid fly in the genus Phytomyptera was observed
Agents Related to this in Florida associated with the decline of serious outbreaks of this insect
Pest Species in the 1990s. In older records from Florida, larvae of the carabid beetle
Plochionus amandus Newman and ants are listed as predators of this
species. Also, listed are six species of hymenopterous parasitoids.

Web Links for Information http://fshs.org/Proceedings/Password%20Protected/

on Palm Leaf Skeletonizer 2007%20vol.%20120/FSHS%20vol.%20120/356-359.pdf; an in-depth
article on this species.

Articles Creighton, J.T. 1937. Homaledra sabalella Chambers, the major pest of
palms in Florida. Journal of Economic Entomology 30: 590-595.
Howard, F.W. and E. Abreu. 2007. The palm leaf skeletonizer,
Homaledra sabalella (Lepidoptera: Coleophoridae): status and potential
pest management options. Proceedings of the Florida Horticultural
Society 120: 356-359.
A Photographic Guide 413

113. Mimosa Webworm, Homadaula anisocentra Meyrick (Lepidoptera:

Galacticidae)

Orientation to Pest Mimosa webworm, Homadaula anisocentra Meyrick, is an invasive

moth from China that was first recorded in North America in
Washington, D.C., in 1940. Larvae feed on flowers and foliage of the
introduced Asian silktree (mimosa) (Albizia julibrissin Durazz.) and the
native North American honeylocust (Gleditsia triacanthos L.). Larvae
web together areas on which they are feeding. This insect over-winters
as pupae encased in cocoons on bark of their host tree or in the leaf
litter below. Eggs of first generation moths are laid on leaves and hatch
in early to mid-June. There are two generations each year and second
generation moths emerge in August.

Hosts Commonly Attacked The caterpillars of this moth feed on the introduced Asian silktree (A.
julibrissin) and the native North American honeylocust (G. triacanthos).

Distribution This insect occurs from Massachusetts and Pennsylvania, south to

Florida and west to Mississippi, Kansas and Nebraska.

Images of Figure 1. Adult of

mimosa webworm,
Mimosa Webworm Homadaula anisocentra.
(Ashley Bradford,
Northern Virginia,
Wikipedia Commons)

Figure 2. Larva of mimosa webworm.

(Stan Gilliam)
414 Forest Pest Insects in North America:

Images of Mimosa Figure 3. A cocoon of

mimosa webworm, the
Webworm (continued) overwintering stage, in
a bark crack. (Michael
Masiuk, Bugwood.org)

Figure 4.Web produced by larvae of the mimosa webworm. (Brian Kunkel,

University of Delaware, Bugwood.org)
A Photographic Guide 415

Images of Mimosa Figure 5. Host tree

heavily damaged by
Webworm (continued) mimosa webworm.
(Michael Masiuk,
Bugwood.org)

Important Biological Control Some North American parasitoids attack this species, but parasitism rates
Agents Related to this are very low (2-4 percent). However, at least in Iowa one parasitoid, the
Pest Species eulophid Elasmus albizziae, attacked 39-47 percent of the pupae.

Web Links for Information http://woodypests.cas.psu.edu/factsheets/insectfactsheets/html/Mimosa_

on Mimosa Webworm Webworm.html; factsheet of Pennsylvania State University.

Articles Heppner, J.B. and G.W. Dekle. 1975. Mimosa webworm, Homadaula
anisocentra Meyrick (Lepidoptera: Plutellidae). Entomology Circular
No. 157, Division of Plant Industry, Florida Department of Agriculture
and Consumer Services.
Bastian, R.A. and E.R. Hart. 1989. First-generation parasitism of the
mimosa webworm (Lepidoptera: Plutellidae) by Elasmus albizziae
(Hymenoptera: Eulophidae) in an urban forest. Environmental
Entomology 19: 409-414.
Sadof, C.S. and R.T. Snyder. 2005. Seasonal abundance of Homadaula
anisocentra (Lepidoptera: Plutellidae) and two parasitoids, Elasmus
albizziae (Hymenoptera: Elasmidae) and Parania geniculata
(Hymenoptera: Ichneumonidae), in an urban forest. Environmental
Entomology 34: 70-74.
416 Forest Pest Insects in North America:

114. Lodgepole Needleminer, Coleotechnites milleri (Busck) (Lepidoptera:

Gelechiidae)

Orientation to Pest Lodgepole needleminer, Coleotechnites milleri (Busck), is a native North

American gelechiid moth whose larvae mine needles of lodgepole pine
(Pinus contorta Douglas ex Loud.). Larvae of all ages mine needles.
Sustained and repeated outbreaks of this species have occurred in
Yosemite National Park in California, causing widespread tree mortality.
The adults are active from mid-July to mid-August in odd numbered
years (each generation requires two years). Eggs are laid in late summer
and hatch the same year. First instars mine a single needle, near the
growing tip, and overwinter inside the mined needle. The following year,
larvae feed on several needles (as needle miners) and develop to the
fourth instar by the end of their second growing season. Fourth instars
overwinter in mined needles and in the third year, fifth instars complete
their feeding in spring, pupate in mined needles, and emerge as adults by
mid-summer.

Hosts Commonly Attacked The only known host is lodgepole pine (P. contorta).

Distribution This insect is found in western North American (United States and
Canada) within the range of lodgepole pine.

Images of Figure 1. Adult of

lodgepole needleminer,
Lodgepole Needleminer Coleotechnites milleri.
(Kipling Will)

Figure 2. Larva of
lodgepole needleminer.
(Scott Tunnock,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 417

Images of Lodgepole
Needleminer (continued)

Figure 3. Damage from lodgepole needleminer. (USDA Forest Service - Ogden

Archive, USDA Forest Service, Bugwood.org)

Figure 4. Stand of lodgepole pine damaged by lodgepole needleminer.

(Louisiana State University Archive, Louisiana State University, Bugwood.org)

Important Biological Control Many parasitoids and other natural enemies attacked this species
Agents Related to this during the declining phase of an outbreak in California (Struble, 1967).
Pest Species Collectively the larval and pupal parasitoids caused 28-46 percent
mortality.

Web Links for Information None

on Lodgepole Needleminer
418 Forest Pest Insects in North America:

Articles on Struble, G.R. 1967. Insect enemies in the natural control of the lodgepole
Lodgepole Needleminer needle miner. Journal of Economic Entomology 60: 225-228.
Struble, G.R. 1973. Biology, ecology, and control of the lodgepole needle
miner. Technical Bulletin, U.S. Department of Agriculture No. 1458:
38 p.
A Photographic Guide 419

115. Banded Ash Clearwing, Podosesia aureocincta Purrington

and Nielson, and Ash/Lilac Clearwing, Podosesia syringae (Harris)
(Lepidoptera: Sesiidae)

Orientation to Pest Banded ash clearwing, Podosesia aureocincta Purrington and Nielson,
and the ash/lilac borer Podosesia syringae (Harris), are similar native
North American clearwing moths with overlapping host ranges.
Podosesia syringae attacks lilac (Syringa), ash (Fraxinus), privet
(Ligustrum) and other trees and shrubs. Podosesia aureocincta breeds
solely in ash, principally green ash (Fraxinus pennsylvanica Marshall)
and white ash (F. americana L.). Before 1975, P. aureocincta was
thought to be a subspecies or just a late brood of the ash/lilac borer
emerging from September to December. Adults of P. aureocincta can be
recognized and separated from P. syringae by the presence of a narrow
orange-yellow partial band on the fourth abdominal segment. The larvae
of P. aureocincta can be distinguished from those of P. syringae because
P. aureocincta larvae have fewer crochets on their abdominal prolegs
(12 to 16 per row vs. 16 to 20 per row for P. syringae). The life histories
of these two species are similar. In both, females lay eggs in cracks on
the bark of ash (or for P. syringae, various hosts). Larvae tunnel through
the wood but continue to expel sawdust and frass, which accumulates
around the gallery opening from which the moth will eventually emerge.
The overwintering stage in both species is the partly grown larva, which
becomes active in spring. Pupation occurs at the end of the larval gallery.
Adults of banded ash clearwing emerge later in the year (September-
December) than those of P. syringae. There is only one generation of
P. syringae per year. Both species cause damage in forest trees and
landscape or nursery trees. The damage caused to ash by either species,
however, is negligible compared to that from the invasive emerald
ash borer, Agrilus planipennis Fairmaire. However, P. aureocincta is
regarded as an important timber pest in the southern United States, and
in the prairie states both species are damaging to trees in shelterbelts.

Hosts Commonly Attacked Podosesia syringae attacks lilac (Syringa), ash (Fraxinus), privet
(Ligustrum), and other trees and shrubs, while P. aureocincta breeds
principally in green (F. pennsylvanica) and white (F. americana) ash.

Distribution Podosesia syringae occurs more widely than P. aureocincta, being found
from Texas to Saskatchewan and east throughout the United States
and Canada. Banded ash clearwing, P. aureocincta, is known just from
the United States, from New York to Florida and west to Oklahoma.
Populations of P. aureocincta are also more scattered than are those
of P. syringae.
420 Forest Pest Insects in North America:

Images of Figure 1. Adult of the

ash/lilac clearwing,
Ash/Lilac Clearwing Podosesia syringae;
the adult of the closely
related species, the
banded ash clearwing,
Podosesia aureocincta
(not shown) is very
similar but has a
distinctive narrow yellow
band around the middle
of the abdomen.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 2. Larva of ash/lilac borer, Podosesia syringae, in green ash twig.

(James Solomon, USDA Forest Service, Bugwood.org)

Figure 3. Death of twigs killed by larvae of ash/lilac borer; note frass in photo on
right. (Both photos: James Solomon, USDA Forest Service, Bugwood.org)
A Photographic Guide 421

Images of Ash/Lilac
Clearwing (continued)

Figure 4. Pupal skins of ash/lilac borer left behind on ash trunk following moth
emergence. (Both photos: Whitney Cranshaw, Colorado State University,
Bugwood.org)

Figure 5. Larval galleries of the ash/lilac borer seen in cross section, in finished
ash lumber. (James Solomon, USDA Forest Service, Bugwood.org)

Important Biological Control Natural enemies of these clearwings have received little attention.
Agents Related to this Woodpeckers feed extensively on the immature stages. Several
Pest Species parasitoids have also been reared from them, but their importance has
not been assessed. The ichneumonid Phaeogenes ater Cresson and
the braconid Macrocentrus marginator (Nees) were reared from P.
aureocincta, while the ichneumonid Lissonota sp. was reared from both
P. aureocincta and P. syringae. Nematode applications can reduce borer
damage in nurseries.
422 Forest Pest Insects in North America:

Web Links for Information http://ipm.illinois.edu/landturf/insects/ash_lilac_borer/index.html; fact

on Ash/Lilac Borer sheet from the University of Illinois covering both species.
and Banded Ash Clearwing http://images.bugwood.org/mediawiki/pdf.cfm?title=Archive:Ash/
Podosesia_aureocincta; Bugwood fact sheet.
http://independent.academia.edu/VernonAntoineBrouJr/Papers/298747/
The_Flight_Periods_of_Podosesia_Syringae_Harris_Ana_Podosesia_
Aureocincta_Purrington_and_Nielsen_In_Louisiana; comparative data
on flights of P. aureocincta and P. syringae.

Articles McKnight, M.E. and S. Tunnock. 1973. The borer problem in green ash
in North Dakota shelterbelts. North Dakota Farm Research 30(5): 8-14.
Purrington, F.F. and D.G. Nielsen. 1979. Genitalic difference between
males of Podosesia aureocincta and P. syringae (Lepidoptera: Sesiidae).
Annals of the Entomological Society of America 72: 552-555.
Solomon, J.D. 1983. Lilac borer (Podosesia syringae) discovered
causing terminal mortality and resulting forks in young green ash trees.
Journal of the Georgia Entomological Society 18: 320-323.
Purrington, F.F. and D.G. Nielsen. 1987. New host records of parasitic
Hymenoptera in clearwing moths (Lepidoptera: Sesiidae). Great Lakes
Entomologist 20: 141-142.
Gill, S., J. Davidson, W. MacLachlan, and W. Potts. 1994. Controlling
banded ash clearwing moth borer using entomopathogenic nematodes.
Journal of Arboriculture 20: 146-149.
A Photographic Guide 423

116. Red Oak Clearwing Borer, Paranthrene simulans (Grote) (Lepidoptera:

Sesiidae)

Orientation to Pest Red oak clearwing borer, Paranthrene simulans (Grote), is a native North
American clearwing moth that affects American chestnut (Castanea
dentata [Marsh.] Borkh.) and some oaks. It is widespread in both eastern
Canada and the eastern United States. Adult moths emerge from April to
June in the southern parts of the species’ range and June and July in the
northern portion. Females lay eggs in bark crevices on the lower trunk.
Larvae burrow in the inner bark and create ovoid chambers. As larvae
grow, they extend their galleries. Larvae overwinter for two winters in
their galleries. In spring, after the second winter, larvae bore to the bark
surface, cap the gallery exits with a thin membrane, and then pupate. The
whole life cycle requires two years. Open-grown trees are attacked most
often. Bark swelling, pellet-like frass in bark cracks, and empty pupal
cases on the trunk are visible signs of attack by this borer. Tunneling
causes losses to nursery tree production and promotes decay in the lower
butt, below the cut made for harvest. Pheromone traps are available to
monitor moth flight to time pesticide applications.

Hosts Commonly Attacked This borer attacks American chestnut (C. dentata) and species in both the
red and white oak groups. Moth preference among hosts varies by region.
In the southern United States older Nuttall (Quercus texana Buckl.),
cherrybark (Q. pagoda Rafinesque), Shumard (Q. shumardii Buckland),
and eastern black (Q. velutina Lamb.) oaks are the principal hosts. In the
northeastern United States, injury is greatest in nursery stock and sapling
red or white oak group trees.

Distribution This borer occurs in the eastern parts of Canada and the United States,
north to Minnesota and south and west to Texas.

Images of Red Oak Figure 1. Adult red

oak clearwing borers,
Clearwing Borer Paranthrene simulans.
(Top: Daniel Herms, The
Ohio State University,
Bugwood.org; bottom:
James Solomon,
USDA Forest Service,
Bugwood.org)
424 Forest Pest Insects in North America:

Images of Red Oak Figure 2. Larva of red oak clearwing

borer in its gallery. (USDA Forest
Clearwing Borer Service Fact Sheet)
(continued)

Figure 3. Emergence holes of red oak clearwing borer on lower trunk of Nuttall
oak. (James Solomon, USDA Forest Service, Bugwood.org)

Important Biological Control No information is available on the natural enemies of this species.
Agents Related to this
Pest Species
Web Links for Information on http://www.fs.fed.us/r8/foresthealth/pubs/oakpests/p17.html; USDA
Red Oak Clearwing Borer Forest Service fact sheet.
http://images.bugwood.org/mediawiki/pdf.cfm?title=Archive:Oak/
Paranthrene_simulans; Bugwood Wiki fact sheet.

Articles McKern, J.A. and A.L. Szalanski. 2007. Molecular diagnostics of

economically important clearwing moths (Lepidoptera: Sesiidae).
Florida Entomologist 90: 475-479.
A Photographic Guide 425

117. Persimmon Borer, Sannina uroceriformis Walker (Lepidoptera:

Sesiidae)

Orientation to Pest Persimmon borer, Sannina uroceriformis Walker, is a native North

American clearwing moth found in the eastern United States. The
larvae feed in the wood of the lower trunk and tap roots of the American
persimmon tree, Diospyros virginiana L. Moths emerge in spring and
mate. Females either lay their eggs on tree bark or drop them at the base
of the host tree. After hatching, larvae move to suitable parts of the host
tree and burrow into the bark. Larvae tunnel in the cambium downward,
and when they are at or near the soil line, tunnel into the wood toward
the center of the tap or lateral roots. Larvae overwinter in their galleries
below the soil line and when ready to pupate extend their galleries
upward in the roots to the ground line or just above. Trees growing
in cutover areas and in hedgerows have higher infestation rates, as do
young trees in nurseries. The life cycle takes 2-3 years to complete.

Hosts Commonly Attacked This borer attacks only persimmons, particularly the native wild
persimmon of the United States (D. virginiana).

Distribution This borer occurs is found in the United States along the Atlantic coast
from New Jersey to Florida, and west to Kansas, Texas, Missouri, Ohio,
and Indiana.

Images of Figure 1. Adult of

persimmon borer,
Persimmon Borer Sannina uroceriformis.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 2. Larva of
persimmon borer.
(James Solomon,
USDA Forest Service,
Bugwood.org)
426 Forest Pest Insects in North America:

Images of Figure 3. Pupation tubes

of persimmon borer
Persimmon Borer pushed out of larval
(continued) galleries.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 4. Galleries of persimmon borer larvae in roots of host tree. (James

Solomon, USDA Forest Service, Bugwood.org)
A Photographic Guide 427

Important Biological Control No natural enemies of this species have been recorded.
Agents Related to this
Pest Species
Web Links for Information http://edis.ifas.ufl.edu/in669; University of Florida fact sheet on biology
on Persimmon Borer and control.
http://images.bugwood.org/mediawiki/pdf.cfm?title=Archive:Borers/
Sannina_uroceriformis; Bugwood factsheet.

Articles Solomon, J.D. 1995. Guide to Insect Borers in North American Broadleaf
Trees and Shrubs. Agriculture Handbook No. AH-706, see p. 56-57.
USDA Forest Service, Washington, DC: 735 p.
428 Forest Pest Insects in North America:

118. Poplar Clearwing Borer, Paranthrene dollii (Neumoegen) (Lepidoptera:

Sesiidae)

Orientation to Pest Poplar clearwing borer, Paranthrene dollii (Neumoegen), is a native

North American clearwing moth that occurs principally in the southern
United States, where it infests the base and root-collar area of young
cottonwoods (Populus) and black willow (Salix nigra Marshall). Eggs
are laid in bark crevices, and larvae bore in the wood and pith, making
open tunnels up to 15 cm long. Heavily infested trees are mechanically
weakened and subject to wind and ice breakage. Damage is sometimes
severe in cottonwood plantations and nurseries, and in the southern states
losses may reach 12 percent of nursery stock. There is one generation
per year.

Hosts Commonly Attacked This borer attacks cottonwoods (Populus) and black willow (S. nigra).

Distribution This borer occurs in the southern United States, from Florida to Texas.

Images of Figure 1. Adult of

poplar clearwing borer,
Poplar Clearwing Borer Paranthrene dollii.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 2. Larva of poplar

clearwing borer.
(James Solomon,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 429

Images of Poplar Figure 3. Frass being

expelled from larval
Clearwing Borer (continued) tunnels of poplar
clearwing borer.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 4. Enlarged
entrance holes in stem
of cottonwood seedling,
caused by larvae of
Paranthrene dollii.
(USDA Forest Service
Fact Sheet)
430 Forest Pest Insects in North America:

Important Biological Control One braconid, Apanteles paranthrenidis Muesebeck, is known from this
Agents Related to this borer’s larvae.
Pest Species

Web Links for Information http://www.forestpests.org/nursery/cwborers.html; USDA Forest Service

on Poplar Clearwing Borer bulletin on biology and control of several cottonwood borers, including
P. dollii.
http://wiki.bugwood.org/Archive:Borers/Paranthrene_dollii; Bugwood
Wiki fact sheet giving details of biology.

Articles Eroles-Harkins, L. 1983. Biology and life history of the cottonwood

clearwing borer, Paranthrene dollii (Neum.) (Lepidoptera: Sesiidae),
in the laboratory and a cottonwood nursery in Mississippi. MS thesis.
Mississippi State University. 106 p.
Nebeker, T.E. and L. Eroles-Harkins. 1988. Daily and seasonal flight
of male Paranthrene dollii (Lepidoptera: Sessiidae), monitored by
pheromone traps. Florida Entomologist 71(3): 376-380.
A Photographic Guide 431

119. Carpenterworm, Prionoxystus robiniae (Peck.) (Lepidoptera: Cossidae)

Orientation to Pest Carpenterworm, Prionoxystus robiniae (Peck.), is a native, wood-boring

moth that attacks various hardwoods. Adults emerge between May and
August and lay their eggs in large groups (200-1,000) on bark of hosts,
often near wounds or objects on the bark such as vines or moss. Young
larvae bore into the inner bark and feed there until half grown, at which
time they bore into the sapwood and then the heartwood. In southern
part of this species’ range, immature stages may complete development
in a single year, but in northern areas as many as 3 or 4 years may be
required. Tunnels to the outside are kept open and enlarged as larvae
grow. Tunnels are lined with silk by mature larvae and pupation occurs
in the upper end of the tunnel near the exit hole. Pupae, when ready
for adult emergence, wriggle to the exit where the adult emerges. The
pupal case remains at the entrance of this exit hole. Carpenterworms
seldom kill trees, but due to tunneling, infested trees are more likely to
break in wind storms, may become more vulnerable to other pests, and
the value of affected logs is reduced. No tree size preference has been
noted for attack, with trees as small of 5 cm in diameter being infested.
However, open grown trees, especially trees in urban landscapes and
rural shelterbelts, are at greatest risk of attack.

Hosts Commonly Attacked In the southern United States, oaks (Quercus), particularly members
of the red oak group, are heavily damaged. In the prairies and Rocky
Mountains of North America, poplar species (Populus) and green ash
(Fraxinus pennsylvanica Marshall) are major hosts. In California, live
oak (Quercus agrifolia Née) and introduced elms (Ulmus) are hosts.
Other hosts include black locust (Robinia pseudoacacia L.), maple
(Acer), and willow (Salix), among others.

Distribution This borer is widely distributed in the United States and southern
Canada.

Images of Figure 1. Adults of

the carpenterworm,
Carpenterworm Prionoxystus robiniae;
female (left) and male
(right). (James Solomon,
USDA Forest Service,
Bugwood.org)
432 Forest Pest Insects in North America:

Images of
Carpenterworm
(continued)

Figure 2. Female carpenterworm, center, in natural position. (Gerald J. Lenhard,

Louiana State Univ, Bugwood.org)

Figure 3. Larvae of
carpenterworm. (Top:
William H. Hoffard,
USDA Forest Service,
Bugwood.org; bottom:
James Solomon,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 433

Images of Figure 4. Larval gallery

of carpenterworm (top)
Carpenterworm and surface injury from
(continued) lateral gallery in trunk
(bottom). (Both photos:
James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 5. Staining
and oozing are
signs marking the
point of entrance for
carpenterworms. (Larry
R. Barber, USDA Forest
Service, Bugwood.org)
434 Forest Pest Insects in North America:

Images of
Carpenterworm
(continued)

Figure 6. Tunnels of carpenterworms in elm (left) and ash (right) logs at harvest.
(Left: USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org;
right: Theodor D. Leininger, USDA Forest Service, Bugwood.org)

Figure 7. Defects
in milled wood from
carpenterworm tunnels
and staining.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Important Biological Control Little is know about the natural control of carpenterworms. Woodpeckers
Agents Related to this prey on larvae. Parasitism appears to be minimal.
Pest Species
A Photographic Guide 435

Web Links for Information http://wiki.bugwood.org/Archive:Borers/Prionoxystus_robiniae;

on Carpenterworm Bugwood wiki fact sheet on biology and control.
http://entomology.unl.edu/ornamentals/pestprofiles/carpenterworm.
shtml; fact sheet of the University of Nebraska.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=8249&ind=C;
fact sheet of Natural Resources Canada.
http://www.entomology.ualberta.ca/searching_species_details.
php?s=2294; University of Alberta insect collection species page.

Articles McKnight, M.E. and S. Tunnock. 1973. The borer problem in green ash
in North Dakota shelterbelts. Shelterbelt Laboratory, Rocky Mountain
Forest & Range Experiment Station, Forest Service, USDA, Bottineau,
North Dakota, USA. North Dakota Farm Research 30(5): 8-14.
Solomon, J.D. and C.J. Hay. 1974. Annotated bibliography of the
carpenterworm, Prionoxystus robiniae. USDA Forest Service General
Technical Report SO-4, Southern Forest Experiment Station: 13 p.
Forschler, B.T. and G.L. Nordin. 1998. Suppression of carpenterworm,
Prionoxystus robiniae (Lepidoptera: Cossidae), with the entomophagous
nematodes, Steinernema feltiae and S. bibionis. Journal of the Kansas
Entomological Society 61: 396-400.
436 Forest Pest Insects in North America:

120. Pecan Carpenterworm, Cossula magnifica (Stecker) (Lepidoptera:

Cossidae)

Orientation to Pest Pecan carpenterworm, Cossula magnifica (Stecker), is a native wood

boring moth that occurs throughout the southeastern United States, where
it reproduces in pecan (Carya illinoinensis [Wangenh.] K. Koch), oak
(Quercus), and hickory (Carya). Adults emerge in spring and lay their
eggs on bark of small twigs. Young larvae bore in the pith of twigs.
Larger larvae move into the branches or the trunk, where they excavate
tunnels several inches in length. Frass is expelled through the larval
tunnel, which is kept open and enlarged as the larva grows. Pupation
occurs in the larval tunnel in the spring. There is one generation per year.

Hosts Commonly Attacked This borer attacks pecan (C. illinoinensis), oak (Quercus), and hickory
(Carya).

Distribution This borer occurs in the southeastern United States.

Images of
Pecan Carpenterworm

Figure 1. Adult of pecan carpenterworm, Cossula magnifica; right, in natural

pose. (Left: James Solomon, USDA Forest Service, Bugwood.org; right:
Theresa Thom, U.S. National Park Service)

Figure 2. Larva of pecan

carpenterworm.
(James Solomon,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 437

Images of
Pecan Carpenterworm
(continued)

Figure 3. Entrance of pecan carpenterworm tunnel in twig. (James Solomon,

USDA Forest Service, Bugwood.org)

Figure 4. Larval tunnels

of pecan carpenterworm
seen in longitudinal
section in pieces of
milled wood.
(James Solomon,
USDA Forest Service,
Bugwood.org)
438 Forest Pest Insects in North America:

Images of Figure 5. Frass pellets

and staining below the
Pecan Carpenterworm tunnel opening of the
(continued) pecan carpenterworm.
(James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 6. Larval tunnels

of pecan carpenterworm
seen in cross section in
a log. (James Solomon,
USDA Forest Service,
Bugwood.org)

Important Biological Control Two tachinid parasitoids—Phorocera comstocki Williston and Phorocera
Agents Related to this signata Aldrich and Webber—have been reared from this species, but not
Pest Species much is known of their effects on the insect’s population dynamics.

Web Links for Information http://wiki.bugwood.org/Archive:Borers/Cossula_magnifica; fact sheet

on Pecan Carpenterworm of Bugwood Wiki on biology and control.

Articles None
A Photographic Guide 439

121. Spruce Budworm, Choristoneura fumiferana (Clemens) (Lepidoptera:

Tortricidae)

Orientation to Pest Spruce budworm, Choristoneura fumiferana (Clemens), is a native North

American moth whose larvae feed on balasam fir (Abies balsamea
[L. Mill.), white (Picea glauca [Moench] Voss), red (P. rubens Sarg.),
and black spruce (P. mariana [Mill.] Britton, Sterns & Poggenburg), and
sometimes larch (Larix), pine (Pinus), and hemlock (Tsuga). Adults fly in
summer and deposits masses of eggs (2-60) on needles in the periphery
of the host tree’s crown. Eggs hatch and produce larvae that create a
hibernacula in which they overwinter as second instars. Hibernaculae
are found on branches and other sites on or near the host tree. In the
spring of the following year, larvae emerge at or before bud break,
disperse aerially, and begin to feed. Buds of staminate flowers are eaten
if available, and then old needles are consumed. Larvae then feed on the
new vegetative buds, and later, the new foliage. When about half grown,
larvae tie several branch tips together with silk to make a nest, where
mature larvae later pupate. Females deposit eggs before flying to new
sites. Defoliation events from this moth occur regularly over wide areas
in eastern Canada and economic losses can be great if uncontrolled. The
population dynamics of this species have been the focus of extensive
and intense investigation for many decades. For a general review of
economic losses and population dynamics see Van Driesche et al. (1996)
under articles below.

Hosts Commonly Attacked Balsam fir (A. balsamea [L.] Mill.) is the preferred host of this
species, but it also feeds on white (P. glauca [Moench] Voss), red
(P. rubens Sarg.), and black spruce (P. mariana [Mill.] Britton, Sterns
& Poggenburg), as well as to some degree on species of larch (Larix),
pine (Pinus), and hemlock (Tsuga).

Distribution This moth is found throughout the range of its main hosts (balsam fir
and white spruce) from Virginia to Labrador and west to Alaska. Related
budworms are found in the western part of North America that were
formerly thought to be the same species but now are considered distinct
species. Damage in the United States is concentrated along the northern
tier of states in the eastern half of the country, from Maine to Minnesota.
440 Forest Pest Insects in North America:

Images of Spruce Budworm

Figure 1. Adults of spruce budworm, Choristoneura fumiferana. (K.B. Jamieson,

Canadian Forest Service, Bugwood.org)

Figure 2. Eggs of spruce budworm on needle. (USDA Forest Service -

Northeastern Area Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 441

Images of Spruce Budworm Figure 3. Young larva

of spruce budworm
(continued) hanging from thread,
preparing for dispersal
by wind. (Jerald E.
Dewey, USDA Forest
Service, Bugwood.org)

Figure 4. Mature larva of spruce budworm. (Connecticut Agricultural Experiment

Station Archive, Connecticut Agricultural Experiment Station, Bugwood.org)
442 Forest Pest Insects in North America:

Images of Spruce Budworm Figure 5. Nests of

mature spruce budworm
(continued) larvae. (Top: A. Steven
Munson, USDA Forest
Service, Bugwood.org;
bottom: Joseph O’Brien,
USDA Forest Service,
Bugwood.org)

Figure 6. Pupa of spruce budworm (left) and cast pupal skin (right). (Both photos:
USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)

Figure 7. Needle mining

damage caused by
young larva of spruce
budworm. (USDA
Forest Service - Ogden
Archive, USDA Forest
Service, Bugwood.org)
A Photographic Guide 443

Images of Spruce Budworm Figure 8. Needle feeding

damage caused by
(continued) older larvae of spruce
budworm. (USDA
Forest Service - Ogden
Archive, USDA Forest
Service, Bugwood.org)

Figure 9. Damage (browning) to stand of host trees by spruce budworms.

(A. Steven Munson, USDA Forest Service, Bugwood.org)

Figure 10. Aerial pesticide applications to large areas damaged by spruce

budworm. (Jerald E. Dewey, USDA Forest Service, Bugwood.org)
444 Forest Pest Insects in North America:

Important Biological Control More than 90 species of parasitoids are known to attack spruce budworm
Agents Related to this in Canada. In Van Driesche et al. (1996) there are details about some
Pest Species of these species and references to their life histories. In addition, some
predator groups (birds) are believed to be important to the population
dynamics of spruce budworm. While some 15 species of pathogens
attack spruce budworm, none are important in the natural population
dynamics of the pest. However, one species, Bacillus thuringiensis
Berliner, is applied as a biopesticide for areawide control, having
fewer nontarget impacts on native species than conventional pesticides.
Hypotheses concerning the causes driving changes in spruce budworm
numbers are discussed in Van Driesche et al. (1996). It is widely believed
that one important influence is variation in spring weather that leads to
different degrees of synchrony between spring larval emergence and bud
break of host trees, with late bud break being unfavorable.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/sbw/budworm.htm; USDA Forest

on Spruce Budworm Service Pest and Disease leaflet covering biology and control.
http://en.wikipedia.org/wiki/Spruce_Budworm; Wikipedia article with
links to other Choristoneura species.
http://www.sampforestpest.ento.vt.edu/defoliating/spruce-budworm/
index.php; links to other documents related to sampling spruce budworm
in various contexts.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=12018; fact
sheet of Natural Resources Canada.
http://www.entomology.ualberta.ca/searching_species_details.
php?s=5272#; contains information on taxonomy of this species.

Articles Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations. FHTET
96-19, December 1996, USDA Forest Service, Morgantown, WV;
see http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf.
A Photographic Guide 445

122. Jack Pine Budworm, Choristoneura pinus pinus Freeman

(Lepidoptera: Tortricidae)

Orientation to Pest Jack pine budworm, Choristoneura pinus pinus Freeman, is closely
related to eastern spruce budworm, Choristoneura fumiferana
(Clemens), and is part of a complex of eight closely related species (see
Lumley et al. 2010, 2011 below). Jack pine budworm is a native North
American moth found in eastern Canada, westward to Alberta and in
the northeastern and north central United States. Its larvae feed various
pines, especially jack (Pinus banksiana Lamb.) and red (P. resinosa Sol.
ex Aiton) pine. Adults fly in mid-to-late summer and lay their eggs in
clusters (about 40) on needles. Eggs hatch and the young larvae, without
feeding, create hibernaculae under the bark scales on tree trunks or limbs,
or between needles. They then molt to second instars and overwinter.
In the spring of the following year, larvae emerge near the time when
staminate flower buds begin to swell. Staminate flowers are usually the
larvae’s first food, and thereafter most larvae move to new foliage, where
they feed and develop. Needles are clipped and webbed together by older
larvae, and needles are only partially consumed. Pupation occurs among
the needles or webbed shoots. Feeding of jack pine budworm usually
does not kill attacked trees, but may cause top killing and deform the
tree, reducing quality of logs for lumber purposes. During outbreaks,
younger trees may be killed outright. For further information see Van
Driesche et al. (1996).

Hosts Commonly Attacked Jack pine budworm larvae feed on various pines, but especially jack
(P. banksiana) and red (P. resinosa) pine.

Distribution This moth is found in Canada throughout the range of jack pine, from
the Maritimes west to Alberta. In the United States, it is found in the
northeastern and north central parts of the country, from New England to
the Great Lakes states.

Images of Figure 1. Adult of

jack pine budworm,
Jack Pine Budworm Choristoneura pinus
pinus. (USDA Forest
Service - Northeastern
Area Archive, USDA
Forest Service,
Bugwood.org)
446 Forest Pest Insects in North America:

Images of Jack Pine Figure 2. Hatched

egg mass (shiny area
Budworm (continued) on central needle) of
jack pine budworm.
(USDA Forest Service
- Northeastern Area
Archive, USDA Forest
Service, Bugwood.org)

Figure 3. Young larvae of jack pine budworm, feeding on male flower of host
plant. (D.M. Benjamin, University of Wisconsin, Bugwood.org)
A Photographic Guide 447

Images of Jack Pine

Budworm (continued)

Figure 4. Mature larva of jack pine budworm. (Steven Katovich, USDA Forest
Service, Bugwood.org)

Figure 5. Pupa of jack pine budworm, in area webbed by larvae.

(Steven Katovich, USDA Forest Service, Bugwood.org)

Figure 6. Webbing and feeding on shoots of host by jack pine budworm larvae.
(Minnesota Department of Natural Resources Archive, Minnesota Department of
Natural Resources, Bugwood.org)
448 Forest Pest Insects in North America:

Images of Jack Pine

Budworm (continued)

Figure 7. Damage (see browning) to stand of host trees by jack pine budworm
larvae. (James B. Hanson, USDA Forest Service, Bugwood.org)

Figure 8. Damage to
understory seedling pine
by jack pine budworm.
(Minnesota Department
of Natural Resources
Archive, Minnesota
Department of
Natural Resources,
Bugwood.org)
A Photographic Guide 449

Images of Jack Pine

Budworm (continued)

Figure 9. Aerial view of area defoliated by jack pine budworm. (A. Steven
Munson, USDA Forest Service, Bugwood.org)

Important Biological Control From 13 to 46 parasitoid species have been recovered in studies of
Agents Related to this jack pine budworms in various locations (see Van Driesche et al. 1996
Pest Species for more detail). The species of greatest importance appear to be the
larval parasitoids Apanteles fumiferanae Viereck (Braconidae) and
Glypta fumiferana (Viereck) (Ichneumonidae), and the pupal parasitoid
Itoplectis conquisitor (Say) (Ichneumonidae). The abundance of male
flowers in a stand is believed to be linked to outbreaks of this species
(see Van Driesche et al. 1996 for a discussion of the literature on this
point).

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/howtos/ht_jack/ht_jack.htm; USDA

on Jack Pine Budworm Forest Service fact sheet on biology and management.
http://en.wikipedia.org/wiki/Jack_Pine; Wikipedia article on basic
biology.
http://www.entomology.ualberta.ca/searching_species_print.
php?s=5271&sn=; provides details on the taxonomy of this species.

Articles Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations. FHTET
96-19, December 1996, USDA Forest Service, Morgantown, WV;
see http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf.
450 Forest Pest Insects in North America:

Articles on Jack Pine Lumley, L.M. and F.A.H. Sperling. 2010. Integrating morphology and
Budworm (continued) mitochondrial DNA for species delimitation within the spruce budworm
(Choristoneura fumiferana) cryptic species complex (Lepidoptera:
Tortricidae). Systematic Entomology 35: 416-428.
Nealis, V.G. 1991. Parasitism in sustained and collapsing populations of
the jack pine budworm, Choristoneura pinus pinus Free. (Lepidoptera:
Tortricidae), in Ontario, 1985-1987. The Canadian Entomologist 123:
1065-1075.
Lumley, L.A. and F.A.H. Sperling. 2011. Utility of microsatellites and
mitochondrial DNA for species delimitation in the spruce budworm
(Choristoneura fumiferana) species complex (Lepidoptera: Tortricidae).
Molecular Phylogenetics and Evolution 58: 232-243.
A Photographic Guide 451

123. Large Aspen Tortrix, Choristoneura conflictana (Walker) (Lepidoptera:

Tortricidae)

Orientation to Pest Large aspen tortrix, Choristoneura conflictana (Walker), is a native

North American tortricid moth found throughout the range of quaking
aspen (Populus tremuloides Michx.) in Canada and the United States.
In Canada, adults lay their eggs in June or July in flat clusters, usually
on the leaves. Eggs hatch and the first instars feed together on leaves,
webbing foliage together. Fed first instars then move to the trunk where
they create hibernaculae and overwinter as second instars. In the spring
of the following year, larvae emerge and feed by mining the swelling
buds. Later, larvae feed in leaf rolls. Pupation occurs in the leaf rolls,
and adults emerge in a few weeks to lay eggs of the new generation.
Defoliating outbreaks have occurred in the aspen stands in Canada,
Alaska, New England, New York, and Michigan. Reasons for outbreaks
have not been studied, but the existence of overwintering young larvae
that feed on buds the following year suggests that variation in the degree
of synchrony (driven by annual weather events) between bud break and
larvae resumption of feeding is likely to be a factor.

Hosts Commonly Attacked The large aspen tortrix is mainly associated with quaking aspen
(P. tremuloides) but also feeds on balsam poplar (Populus sect.
tacamahaca), bigtooth aspen (P. grandidentata Michaux), paper birch
(Betula papyrifera Marsh.), willow (Salix), and alder (Alnus).

Distribution This moth is found throughout the range of quaking aspen in Canada and
the United States.

Images of Large
Aspen Tortrix

Figure 1. Adult of large aspen tortrix, Choristoneura conflictana. (Steven

Katovich, USDA Forest Service, Bugwood.org)
452 Forest Pest Insects in North America:

Images of Large Aspen Figure 2. Egg mass

of large aspen tortrix.
Tortrix (continued) (Steven Katovich,
USDA Forest Service,
Bugwood.org)

Figure 3. Young larvae

of large aspen tortrix,
feeding on webbed
leaves. (Minnesota
Department of Natural
Resources Archive,
Minnesota Department
of Natural Resources,
Bugwood.org)
A Photographic Guide 453

Images of Large Aspen

Tortrix (continued)

Figure 4. Mature larva of large aspen tortrix. (William M. Ciesla, Forest Health
Management International, Bugwood.org)

Figure 5. Rolled leaf,

feeding site of older
larvae of large aspen
tortrix. (Steven Katovich,
USDA Forest Service,
Bugwood.org)
454 Forest Pest Insects in North America:

Images of Large Aspen Figure 6. Pupa of large

aspen tortrix.
Tortrix (continued) (William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)

Figure 7. Quaking aspen

defoliated by large
aspen tortrix.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)
A Photographic Guide 455

Images of Large Aspen

Tortrix (continued)

Figure 8. Aerial view of area defoliated by large aspen tortrix. (USDA Forest
Service - Region 2 - Rocky Mountain Region Archive, USDA Forest Service,
Bugwood.org)

Important Biological Control A large number of species of parasitoids (>20) have been reared from
Agents Related to this larvae and pupae in several locations where studies have been conducted
Pest Species (Alberta, Canada; Alaska). Collectively, these cause significant levels
of mortality, up to 50 percent in some cases. Among the more important
species are Macrocentrus iridescens French, Glypta inversa Cresson,
Glypta fumiferanae (Viereck), and Agathis annulipes (Cresson).

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=12016; fact

on Large Aspen Tortrix sheet of Natural Resources Canada.
http://www.fs.fed.us/r10/spf/fhp/leaflets/Larasptor.htm; USDA Forest
Service fact sheet.
http://www.fs.fed.us/r10/spf/fhp/leaflets/Fidl139.htm; USDA Forest
Service leaflet.
http://www.entomology.ualberta.ca/searching_species_details.
php?s=5251; fact sheet of the University of Alberta.

Articles Torgersen, T.R. and R.C. Beckwith. 1974. Parasitoids associated with the
large aspen tortrix, Choristoneura conflictana (Lepidoptera: Tortricidae),
in interior Alaska. The Canadian Entomologist 106: 1247-1265.
Jones, B.C., J. Roland, and M.L. Evenden. 2009. Development of a
combined sex pheromone-based monitoring system for Malacosoma
disstria (Lepidoptera: Lasoicampidae) and Choristoneura conflictana
(Lepidoptera: Tortricidae). Environmental Entomology 38: 459-471.
456 Forest Pest Insects in North America:

124. Sugar Pine Tortrix, Choristoneura lambertiana Busck (Lepidoptera:

Tortricidae)

Orientation to Pest Sugar pine tortrix, Choristoneura lambertiana Busck, is a tortricid

moth native to western North American that feeds on various conifers,
especially pines. This species includes many different races that vary
in appearance or aspects of biology, some of which have been given
subspecies names. This moth’s life history is similar to that of other
better known members of the genus. Adults fly in summer (July and
August) and lay eggs on the foliage of the host plant. Young larvae
overwinter in hibernaculae on the tree trunk and reemerge the following
spring when they feed by mining needle sheaths and staminate cones.
New foliage buds are also consumed. Several larvae (1-5) may feed
together on the same shoot. Larvae web the needles into feeding shelters
and pupae are formed among the webbed needles. Damaged trees are not
likely to die, but tops may be killed.

Hosts Commonly Attacked The sugar pine tortrix feeds on various pines, including sugar (Pinus
lambertiana Douglas), lodgepole (P. contorta Douglas), limber (P. flexilis
E. James), and ponderosa (P. ponderosa Douglas ex. C. Lawson) pines,
as well as some other conifers, including Douglas-fir (Pseudotsuga
menziesii [Mirb.] Franco) and juniper (Juniperus).

Distribution This moth is found in several western U.S. states (including California,
Colorado, Idaho, Montana, Oregon, and Montana). Ranges by subspecies
have been recorded as follows: Choristoneura lambertiana lambertiana
(confirmed: Siskiyou County and Ashland, Oregon; uncertain: Alberta
and British Columbia in Canada and Idaho, Montana, eastern Oregon,
and Wyoming in the United States); C. lambertiana ponderosana
(Arizona, Colorado, Nevada, New Mexico, western New England, South
Dakota); C. lambertiana subretiniana (California and Oregon).

Images of
Sugar Pine Tortrix

Figure 1. Adults (two forms with different color patterns) of the sugar pine
tortrix, Choristoneura lambertiana. (Left: Mark McGregor, USDA Forest Service,
Bugwood.org; right: USDA Forest Service - Ogden Archive, USDA Forest
Service, Bugwood.org)
A Photographic Guide 457

Images of Sugar Pine

Tortrix (continued)

Figure 2. Egg mass of the sugar pine tortrix. (USDA Forest Service -
Northeastern Area Archive, USDA Forest Service, Bugwood.org)

Figure 3. Larvae of
the sugar pine tortrix.
(Top: Scott Tunnock,
USDA Forest Service,
Bugwood.org; bottom:
Bernard J. Raimo,
USDA Forest Service,
Bugwood.org)
458 Forest Pest Insects in North America:

Images of Sugar Pine

Tortrix (continued)

Figure 4. Needles that have been clipped and webbed together by sugar pine
tortrix larvae. (Left: Scott Tunnock, USDA Forest Service, Bugwood.org; right:
USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)

Figure 5. Pupa of the

sugar pine tortrix (top)
and skin of emerged
pupa (bottom). (Top:
USDA Forest Service
- Ogden Archive,
USDA Forest Service,
Bugwood.org; bottom:
Bernard J. Raimo,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 459

Images of Sugar Pine

Tortrix (continued)

Figure 6. Damage to tips of small pine from larvae of the sugar pine tortrix.
(Dave Powell, USDA Forest Service, Bugwood.org)

Important Biological Control As with other Chorisoneura species, the sugar pine tortrix is affected by
Agents Related to this a large complex of natural enemies, especially parasitoids, of which some
Pest Species 22 species are known from rearing. However, the effect, if any, of these
species on fluctuations in density of this pest is unknown.

Web Links for Information http://www.ireference.ca/search/Choristoneura%20lambertiana/;

on Sugar Pine Tortrix discusses host plants of some of the subspecies of the sugar pine tortrix.
http://en.wikipedia.org/wiki/Choristoneura_lambertiana; Wikipedia
articles includes information on ranges of subspecies.
http://www.ext.colostate.edu/pubs/insect/05567.html; factsheet of
Colorado State University giving general information of subspecies
associated with ponderosa pine.

Articles Stevens, R.E., T.K. Borg, and T.O. Thatcher. 1977. Notes on a
pine-feeding budworm, Choristoneura lambertiana ponderosana
(Lepidoptera: Tortricidae), in the Colorado Rockies. The Canadian
Entomologist 109: 1269-1274.
Powell, J.A. 1995. Biosystematic Studies of Conifer-feeding
Choristoneura (Lepidoptera: Tortricidae) in the Western United States.
University of California Publications, Entomology, Berkeley, California.
(Contains range information.)
460 Forest Pest Insects in North America:

125. Western Spruce Budworm, Choristoneura occidentalis Freeman

(Lepidoptera: Tortricidae)

Orientation to Pest Western spruce budworm, Choristoneura occidentalis Freeman,

is a tortricid moth native to western North American that feeds on
various conifers, including species of fir (Abies), larch (Larix), spruce
(Picea), and Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco).
The life history of this moth is similar to that of the spruce budworm
(Choristoneura fumiferana [Clemens]). Adults fly and lay eggs in
summer (July and August) on the host plant. Young larvae do not feed
but rather immediately spin a hibernacula under bark scales, where they
overwinter. The next spring, larvae become active and feed by mining
old needles until the buds swell. Larvae then bore into the buds and feed
on the expanding needles. Later in the spring, they loosely web together
growing tips and feed upon the new needles, where they later pupate.
There is one generation per year. This is considered one of the most
damaging pests in western North American forests, and large, sustained
outbreaks sometimes occur. Damaged trees may be killed entirely or just
the tops may die, destroying the commercial value of the trees.

Hosts Commonly Attacked This species feeds on various confers including Abies concolor (Gordon)
Lindley ex Hildebrand, A. grandis (Douglas ex D. Don) Lindley,
A. lasiocarpa (Hooker) Nuttall, Larix occidentalis Nutt., Picea
engelmannii Parry ex Engelm., P. glauca (Moench) Voss, P. pungens
Engelm., Douglas-fir, and juniper (Juniperus).

Distribution This moth is found in several western U.S. states, including Colorado,
Idaho, Montana, Wyoming, Colorado, Arizona, Utah, New Mexico,
Oregon, and Washington, as well of British Columbia and southwest
Alberta, Canada.

Figure 1. Distribution
of the western spruce
budworm, Choristoneura
occidentalis, in western
North America. (USDA
Forest Service •
Forest Insect &
Disease Leaflet 53)
A Photographic Guide 461

Images of Western
Spruce Budworm

Figure 2. Adult of the western spruce budworm. (USDA Forest Service - Region 4
- Intermountain Archive, USDA Forest Service, Bugwood.org)

Figure 3. Silvery egg mass of the western spruce budworm on needle. (USDA
Forest Service - Region 2 - Rocky Mountain Region Archive, USDA Forest
Service, Bugwood.org)

Figure 4. Very young larva of western spruce budworm feeding in bud (left) and
on new foliage (right). (Both photos: USDA Forest Service - Region 2 - Rocky
Mountain Region Archive, USDA Forest Service, Bugwood.org)
462 Forest Pest Insects in North America:

Images of Western
Spruce Budworm
(continued)

Figure 5. Webbed foliage, the feeding site of young larvae of the western spruce
budworm. (Ladd Livingston, Idaho Department of Lands, Bugwood.org)

Figure 6. Mature larva of western spruce budworm. (Scott Tunnock, USDA Forest
Service, Bugwood.org)

Figure 7. Pupae of the western spruce budworm. (Left: USDA Forest Service -
Region 2 - Rocky Mountain Region Archive, USDA Forest Service, Bugwood.org;
right: David McComb, USDA Forest Service, Bugwood.org)
A Photographic Guide 463

Images of Western Figure 8. Damage

caused by western
Spruce Budworm spruce budworm to a
(continued) grand fir.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)

Figure 9. Stand of grand fir in Oregon defoliated by western spruce budworm.

(William M. Ciesla, Forest Health Management International, Bugwood.org)
464 Forest Pest Insects in North America:

Images of Western
Spruce Budworm
(continued)

Figure 10. Stand with dead trees 10 years after a western spruce budworm
outbreak in Oregon. (Dave Powell, USDA Forest Service, Bugwood.org)

Figure 11. Eggs of western spruce budworm parasitized by the egg parasitoid
Trichogramma minutum Riley. (USDA Forest Service - Region 2 - Rocky
Mountain Region Archive, USDA Forest Service, Bugwood.org)

Important Biological Control As with other Choristoneura species, the western spruce budworm is
Agents Related to this affected by a large complex of natural enemies, especially parasitoids,
Pest Species which, along with adverse weather, partially limit the pest’s density.
However, population outbreaks occur when climatic and forest stand
conditions are favorable. There are more than 40 species of insect
parasitoids of the western spruce budworm, of which four or five species
are most common. See Bellows et al. (1998) for species names.
A Photographic Guide 465

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/westbw/fidl-wbw.htm; USDA

on Western Spruce Budworm Forest Service fact sheet covering biology, control and other topics.
http://www.ext.colostate.edu/pubs/insect/05543.html; fact sheet of
Colorado State University.
http://ext.nrs.wsu.edu/forestryext/foresthealth/notes/westernbudworm.
htm; fact sheet of Washington State University.
http://www.oregon.gov/ODF/privateforests/docs/fh/
WesternSpruceBudworm.pdf?ga=t; fact sheet of the Oregon Department
of Forestry.

Articles Schmidt, W.C. and D.C. Fellin. 1973. Western spruce budworm damage
affects form and height growth of western larch. Canadian Journal of
Forest Research 3(1): 17-26.
Jennings, D.T., F.B. Knight, S.C. Hacker, and M.E. McKnight. 1979.
Spruce budworms bibliography. Misc. Rep. 213. University of Maine,
School of Forest Resources, Life Science and Agricultural Experiment
Station. Orono, Maine: 687 p.
Bellows, T.S., C. Meisenbacher, and R.C. Reardon. 1998. Biological
Control of Arthropod Forest Pests of the Western United States: A
review and recommendations. FHTET-96-21. USDA Forest Service,
Morgantown, West Virginia, USA.
466 Forest Pest Insects in North America:

126. Spruce Bud Moth, Zeiraphera canadensis Mutuura and Freeman

(Lepidoptera: Tortricidae)

Orientation to Pest Spruce bud moth, Zeiraphera canadensis Mutuura and Freeman, is a
native tortricid moth associated predominantly with spruce (Picea) in
eastern North America. It overwinters as eggs, which hatch in the spring.
There are four larval instars. First and second instar larvae mine needles
of recently burst buds. Third and fourth instars feed on the stems of
shoots. Pupation usually takes place in the soil. There is one generation
per year. Larval feeding on shoots of the upper crown and on the leader
can stunt tree growth, causing economically important damage in spruce
plantations. Injury is most common on open grown spruces (before
crown closure) and this species is not a pest in natural forest stands.

Hosts Commonly Attacked This moth feeds mainly on white spruce (Picea glauca [Moench]), but
sometimes attacks black spruce (P. mariana [Mill.] Britton, Sterns and
Poggenburg) or balsam fir (Abies balsamea [L.] Mill.).

Distribution This moth occurs throughout the range of spruce in the northern United
States and eastern Canada.

Images of
Spruce Bud Moth

Figure 1. Adult of spruce bud moth, Zeiraphera canadensis Mutuura and

Freeman. (Bo Zaremba - [email protected])
A Photographic Guide 467

Images of Spruce Figure 2. Parasitized

egg of spruce bud
Bud Moth (continued) moth (black) at base of
shoot. (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Figure 3. Mature larva

of spruce bud moth.
(Dion Manastyrski -
Pacific Forestry Centre,
Natural Resources
Canada, Canadian
Forest Service)

Figure 4. Buds damaged by spruce bud moth. (Left: Connecticut Agricultural

Experiment Station Archive, Connecticut Agricultural Experiment Station,
Bugwood.org; right: Dion Manastyrski - Pacific Forestry Centre, Natural
Resources Canada, Canadian Forest Service)

Figure 5. Buds damaged

by spruce bud moth; see
cap of bud held by silk
threads. (Jean-François
Mouton - Natural
Resources Canada,
Canadian Forest
Service)
468 Forest Pest Insects in North America:

Important Biological Control In Newfoundland, Canada, the ichneumonid Tycherus osculator

Agents Related to this parasitizes up to 50 percent of the larvae of spruce bud moth.
Pest Species

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=11208; fact

on Spruce Bud Moth sheet of Natural Resources Canada, including information on how to
scout for this pest.
http://www.fs.fed.us/r10/spf/fhp/leaflets/Sprlarbudmot.htm; fact sheet
of USDA Forest Service, Alaska on Zeiraphera spp. and methods for
reducing their damage on spruce and larch.

Articles Turgeon, J.J. 1985. Life cycle and behaviour of the spruce bud moth,
Zeiraphera canadensis (Lepidoptera: Olethreutidae) in New Brunswick.
The Canadian Entomologist 117: 1239-1247.
Turgeon, J.J. and J. Régnière. 1987. Development of sampling
schemes for the spruce bud moth, Zeiraphera canadensis (Lepidoptera:
Tortricidae). The Canadian Entomologist 119: 239-249.
Turgeon, J.J., N. Nelson, and E.G. Kettela. 1987. Reproductive biology
of the spruce bud moth, Zeiraphera canadensis (Lepidoptera: Tortricidae:
Olethreutinae), in New Brunswick. The Canadian Entomologist
119: 361-364.
Carroll, A.L. and D.T. Quiring. 1993. Influence of feeding by Zeiraphera
canadensis (Lepidoptera: Tortricidae) on growth of white spruce: larval
density - damage and damage - shoot production relationships. Journal
of Applied Ecology 30: 629-639.
West, R.J., M. Kenis, G.W. Butt, and S.M. Bennett. 1999. Parasitoid
complex of Zeiraphera canadensis (Lepidoptera: Tortricidae) and
evaluation of Tycherus osculator (Hymenoptera: Ichneumonidae) as a
biological control agent. The Canadian Entomologist 131: 465-474.
West, R.J., M. Kenis, R.S. Bourchier, S.M. Smith, and G.W. Butt.
2001. Zeiraphera canadensis Mutuura and Freeman, spruce bud
moth (Lepidoptera: Tortricidae). In: Mason, P.G. and J.T. Huber
(eds.). Biological Control Programmes in Canada, 1981-2000.
CABI Publishing, Wallingford, UK: 279-283.
A Photographic Guide 469

127. Cottonwood Twig Borer, Gypsonoma haimbachiana (Kearfott)

(Lepidoptera: Tortricidae)

Orientation to Pest Cottonwood twig borer, Gypsonoma haimbachiana (Kearfott), is a

native tortricid found throughout the eastern United States and parts
of southern Canada. It bores in the stems of young eastern cottonwood
(Populus deltoides Bartr.) and is considered an important pest of young
cottonwoods that are grown in plantations. Winter is spent as young
larvae that are found on trees in silk-lined shallow pits excavated in
healed over borer entrance holes or other depressions on trees. Some
older larvae overwinter in hollowed-out buds. In spring, larvae bore
into tender new shoots. After pupation and adult emergence, eggs are
laid singly or in small groups on the foliage. New larvae of summer
generations cover themselves with silk mixed with trash, then bore into
the midrib. After a few days, the larvae abandon their midrib galleries
and tunnel in shoots to complete their development. In the southern
United States, there may be four to five generations per year. Damaged
trees are stunted, crooked, and have many limbs, reducing tree value.

Hosts Commonly Attacked This borer attacks various poplars, especially eastern cottonwood.

Distribution Cottonwood twig borer is found in the eastern United States, west to
Texas and in Ontario, Canada. It is common in the southern United
States.

Images of Figure 1. Adults of

the cottonwood twig
Cottonwood Twig Borer borer, Gypsonoma
haimbachiana (Kearfott).
(Top: Jim Vargo; bottom:
Steve Scott, Illinois
Eastern Community
College, Bugwood.org)
470 Forest Pest Insects in North America:

Images of Cottonwood Figure 2. Egg of

cottonwood twig
Twig Borer (continued) borer. (R.C. Morris,
USDA Forest Service,
Bugwood.org)

Figure 3. Larvae of cottonwood twig borer. (Both photos: Whitney Cranshaw,

Colorado State University, Bugwood.org)
A Photographic Guide 471

Images of Cottonwood
Twig Borer (continued)

Figure 4. Cottonwood twig broken at larval tunnel of cottonwood twig borer.

(Whitney Cranshaw, Colorado State University, Bugwood.org)

Figure 5. Frass tubes at cottonwood twig borer entrance holes are a sign of this
pest’s presence. (James Solomon, USDA Forest Service, Bugwood.org)

Important Biological Control Some natural enemies of have been noted attacking cottonwood twig
Agents Related to this borer (e.g., those in Steward and Payne (1972) and Steward (1973), cited
Pest Species below. Also, see comments in Van Driesche et al. (1996). However, these
have not been studied sufficiently to get a general understanding of their
importance or distribution.
472 Forest Pest Insects in North America:

Web Links for Information http://wiki.bugwood.org/Archive:South/Gypsonoma_haimbachiana;

on Cottonwood Twig Borer Bugwood Wiki fact sheet on biology.
http://www.fs.fed.us/r8/foresthealth/idotis/insects/cwtwigbr.html; USDA
Forest Service, Southern Region fact sheet.

Articles Morris, R.C. 1967. Biology of Gypsonoma haimbachiana (Lepidoptera:

Olethreutidae), a twig borer in eastern cottonwood. Annals of the
Entomological Society of America 60: 423-427.
Stewart, J.W. and T.L. Payne. 1972. Parasites and predators of the
cottonwood twig borer in Texas. Environmental Entomology 1: 669-670.
Stewart, J.W. and T.L. Payne. 1975. Seasonal abundance and impact of
the cottonwood twig borer on cottonwood trees. Journal of Economic
Entomology 68: 599-602.
Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations. FHTET
96-19, December 1996, USDA Forest Service, Morgantown, WV.
(See http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
McMillin, J.D., M.J. Anderson, E.E. Butin, and E.R. Hart. 1998.
Phenology and infestation patterns of the cottonwood twig borer
(Lepidoptera: Tortricidae) in Iowa. Great Lakes Entomologist
31: 181-190.
A Photographic Guide 473

129. Oak Leafroller, Archips semiferanus (Walker) (Lepidoptera: Tortricidae)

Orientation to Pest Oak leafroller, Archips semiferanus (Walker), is native to North America,
where it is widely distributed in the eastern half of the continent. Females
lay eggs in masses of 40 to 50 eggs on branches or rough bark in July.
Eggs overwinter and larvae emerge in spring and feed on buds and young
leaves. Mid-to-late instar larvae roll leaves together with silk to form a
nest in which they feed. Mature larvae pupate in the leaf rolls or crevices
in June and adults emerge a few weeks later. Periodic outbreaks of this
species have occurred in oak forests in North America since the 1960s.
In Pennsylvania and Michigan, hundreds of thousands of acres have been
defoliated in some years, with substantial tree mortality and economic
loss.

Hosts Commonly Attacked Larvae of oak leafroller feed on a range of oaks (Quercus), especially
northern red (Quercus rubra L.), scarlet (Q. coccinea Muenchh.),
chestnut (Q. prinus L.), and white (Q. alba L.) oaks. In addition, this
moth sometimes feeds on species of witch hazel (Hamamelis) and apple
(Malus).

Distribution Archips semiferanus is found in southeastern Canada and the eastern

United States, west to Colorado and Texas.

Images of
Oak Leafroller

Figure 1. Adult of oak leafroller, Archips semiferanus. (Todd Gilligan, CSU,

Bugwood.org)
474 Forest Pest Insects in North America:

Images of Oak
Leafroller (continued)

Figure 2. Larva of oak leafroller. (A. Steven Munson, USDA Forest Service,
Bugwood.org)

Important Biological Control Natural enemies of this species have been studied in Pennsylvania where
Agents Related to this pupae were attacked by two ichneumonids, Itoplectis conquistor (Say)
Pest Species and Phaeogenes gilvilabris Allen, and larvae were attacked by various
tachinids. Larval parasitism was more common than pupal parasitism.

Web Links for Information http://en.wikipedia.org/wiki/Archips_semiferanus; Wikipedia fact sheet

on Oak Leafroller on biology and taxonomy of species.

Articles Freeman, T.N. 1958. The Archipinae of North America. The Canadian
Entomologist, Supplement No. 7: 89 p.
Chapman, P.J. and S.E. Lienk. 1971. Tortricid fauna of apple in New
York; including an account of apples’ occurrence in the State, especially
as a naturalized plant. New York State Agricultural Experiment Station,
Special Publication, Geneva, New York: 142 p.
Wilson, L.F. 1972. Life history and outbreaks of an oak leafroller,
Archips semiferanus (Lepidoptera: Tortricidae), in Michigan. Great
Lakes Entomologist 5: 71-77.
Mumma, R.O. and A.S. Zettle. 1977. Larval and pupal parasites of the
oak leafroller, Archips semiferanus. Environmental Entomology 6:
601-605.
A Photographic Guide 475

130. Longleaf Pine Seedworm, Cydia ingens (Heinrich) (Lepidoptera:

Tortricidae)

Orientation to Pest Longleaf pine seedworm, Cydia ingens (Heinrich), is a native tortricid
that is part of the C. toreuta complex whose members cannot be
separated morphologically and whose species status is uncertain
(Gilligan et al. 2008). Species are currently recognized based on host
and range. C. ingens (in the narrow sense) has been interpreted as being
restricted to Florida. This species, or closely related forms, occur in the
southeastern United States where their larvae feed in immature cones of
long leaf pine (Pinus palustris Mill.), destroying the seeds.

Hosts Commonly Attacked There are many seed cone worms associated with various pines. In the
southern United States this species, or a closely related one, feeds on
long leaf pine (P. palustris).

Distribution This species complex is found in the southeastern United States, with
C. ingens (in the narrow sense) perhaps being restricted to Florida.

Image of Longleaf
Pine Seedworm

Figure 1. Adult of longleaf pine seedworm, Cydia ingens. (Todd Gilligan, CSU,
Bugwood.org)

Important Biological Control No information is available on natural enemies of this species.

Agents Related to this
Pest Species
Web Links for Information http://www.tortricidae.com/identificationPhoto.asp?NAV=10; website of
on Longleaf Pine Seedworm tortricd.net, resources on the Tortricidae.
476 Forest Pest Insects in North America:

Articles on Abrahamson, L.P. and K.J. Kraft. 1965. A population study of the cone
Longleaf Pine Seedworm moth Laspeyresia toreuta Grote in Pinus banksiana stands. Ecology
46: 561-563.
Harbo, J.R. and K.J. Kraft. 1969. A study of Phanerotoma toreutae,
a parasite of the pine cone moth Laspeyresia toreuta. Annals of the
Entomological Society of America 62: 214-220.
Gilligan, T.M., D.J. Wright, and L.D. Gibson. 2008. Olethreutine
Moths of the Midwestern United States, An Identification Guide.
Ohio Biological Survey 16(2): 334 p.
A Photographic Guide 477

131. European Pine Shoot Moth, Rhyacionia buoliana (Denis and

Schiffermiiller) (Lepidoptera: Tortricidae)

Orientation to Pest European pine shoot moth, Rhyacionia buoliana (Denis and
Schiffermiiller), is an invasive tortricid native to Europe, the eastern
Mediterranean region, and Japan. It is invasive in the United States,
Canada, Chile, and Uruguay. It is a serious pest of pines, especially
in pine plantations because it kills the leaders of pines, destroying
their growing form. Larvae feed on pines, especially red pine (Pinus
resinosa Sol. ex Aiton) or introduced European pines, and, in Southern
Hemisphere plantations, on Monterrey pines (P. radiata D. Don). Buds
of branches are also attacked. Moths emerge in early summer and lay
their eggs on new needles. Young larvae construct webs that are coated
with resin below the current year’s needle sheaths and stems. Feeding
initially occurs within the sheath at the base of needles. By midsummer,
larvae switch and feed on buds until August. Larvae overwinter in webs
at the bases of buds and in spring move to undamaged buds or new
shoots, where they complete development. Pupation takes place during
the second summer in the dead buds and shoots, and when adults emerge,
empty pupal cases remain sticking out of the pupal chambers in the
hollowed-out buds. One generation occurs each year. Death of leaders
leads to deformed, bushy trees. In Chile, as much as a third of trees in
Monterrey pine plantations may be infested.

Hosts Commonly Attacked In North America, this moth attacks various native pines, especially
red pine (P. resinosa), as well as several introduced European species,
particularly Mugo (P. mugo Turra) and Scots (P. sylvestris L.) pines. The
five-needle pines are relatively resistant. In the Southern Hemisphere, an
important host is the North American species P. radiata, when grown in
plantations.

Distribution In North America, the moth occurs in southern Canada from

Newfoundland to the Great Lakes and in the United States south to
Maryland and Illinois. A separate infestation exists in British Columbia,
Oregon, and Washington.

Images of European
Pine Shoot Moth

Figure 1. Adults of European pine shoot moth, Rhyacionia buoliana. (Left: USDA
Forest Service - Northeastern Area Archive, USDA Forest Service, Bugwood.org;
right: USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)
478 Forest Pest Insects in North America:

Images of European
Pine Shoot Moth
(continued)

Figure 2. Eggs (right) and first instar larva (left) of European pine shoot moth.
(USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)

Figure 3. Larva of European pine shoot moth feeding in shoot. (Fabio Stergulc,
Università di Udine, Bugwood.org)
A Photographic Guide 479

Images of European Figure 4. Live pupa

of European pine
Pine Shoot Moth shoot moth in place in
(continued) damaged shoot (top);
pupa dissected from
shoot (middle), and
cast pupal skin (bottom)
from damaged leader.
(Top: David McComb,
USDA Forest Service,
Bugwood.org; middle
and bottom: USDA
Forest Service - Ogden
Archive, USDA Forest
Service, Bugwood.org)
480 Forest Pest Insects in North America:

Images of European Figure 5. Deformation of

shape of pine tree; top,
Pine Shoot Moth in young tree; bottom,
(continued) eventual tree shape
(here, “Y”) due to leader
death from feeding of
European pine shoot
moth. (Top: Jan Liska,
Forestry and Game
Management Research
Institute, Bugwood.org;
bottom: Fabio Stergulc,
Università di Udine,
Bugwood.org)
A Photographic Guide 481

Important Biological Control Many studies have described the parasitoids that can be reared from
Agents Related to this larvae and pupae of European pine shoot moth in various countries
Pest Species (United States, Canada, Chile, parts of Europe) and on various tree
species. More than a hundred species of parasitoids have been recorded.
Studies conducted in coastal areas of Germany in the 1950s and 1960s,
for example, found larval and pupal parasitism of this species to be
in the 28-65 percent range in pine plantations. The most common
parasitoids observed were the ichneumonid Cremastus confluens Grav.
and the braconid Orgilus obscurator Nees. Similar studies have been
done in various locations, with some variation in the parasitoid guild.
Several European parasitoid species were released in North America
for suppression of this pest, but only a few have become established.
Of these, only O. obscurator appears to be widespread and of some
importance. Releases of European parasitoids in North America have,
however, had limited effect on densities of this pest in most areas.
Possible explanations for this lack of control may include low vegetation
diversity in North American pine plantations and a possible need
to import additional species of parasitoids from Europe. Parasitoid
importations against this species have also been undertaken in Chile.

Web Links for Information on http://www.forestry.ubc.ca/fetch21/FRST308/lab4/rhyacionia_buoliana/

European Pine Shoot Moth europe.html; fact sheet of the University of British Columbia.
http://wiki.bugwood.org/Archive:Northeast/Rhyacionia_buoliana;
fact sheet of BugwoodWiki, covering biology.
http://www.pfc.forestry.ca/diseases/nursery/pests/europea1_e.html;
Natural Resources Canada fact sheet.
http://www.cabi.org/isc/?compid=5&dsid=23641&loadmodule=
datasheet&page=481&site=144; fact sheet of CABI from the UK,
showing worldwide distribution.

Articles Schultz, A., D. Häussler, K.H. Apel, and E.K. Groll. 1997. RHYA-
SIM - a model illustrating the population dynamics of the pine
shoot moth (Rhyacionia buoliana). Beiträge für Forstwirtschaft und
Landschaftsökologie 31(2): 88-91. (In German).
Daterman, G., A. Eglitis, D. Czokajlo, C. Sack, and P. Kirsch. 2001.
Attract and kill technology for management of European pine shoot
moth (Rhyacionia buoliana) and Western pine shoot borer (Eucosma
sonomana). In: Knízek, M., B. Forster, and W. Grodzki (eds.).
Methodology of forest insect and disease survey in central Europe.
Proceedings of the Fourth Workshop of the IUFRO Working Party.
Prague, Czech Republic, 17-20 September 2001. Journal of Forest
Science 47 (Special Issue 2): 66-69.
Huerta, A., F. Robredo, J. Diez, and J.A. Pajares. 2006. The parasitoid
complex selection of the European pine shoot moth (Rhyacionia
buoliana Den. et Schiff.) (Lepidoptera: Tortricidae) for the biological
control in Chile. Boletín de Sanidad Vegetal, Plagas 32(4): 95-607.
(In Spanish).
482 Forest Pest Insects in North America:

132. Nantucket Pine Tip Moth, Rhyacionia frustrana (Comstock)

(Lepidoptera: Tortricidae)

Orientation to Pest Nantucket pine tip moth, Rhyacionia frustrana (Comstock), is a native
shoot-boring tortricid associated with pines in the eastern United
States. There is a similar insect, found in Nebraska attacking pines in
plantations, that was formerly thought to be a subspecies of R. frustrana
but later was recognized as its own species, R. bushnelli Miller. Larvae of
Nantucket pine tip moth bore into and kill leaders of pines, both reducing
overall growth in wood volume and deforming the tree. The species is
a pest of various southern pines in plantations. Some species such as
shortleaf (Pinus echinata Miller) and loblolly (P. taeda L.) pines are
severely affected. The proportion of tips infested by this pest decreases
after trees reach 3-4 meters in height.

Hosts Commonly Attacked In North America, Nantucket pine tip moth attacks several southern pines
in plantations, especially shortleaf (P. echinata) and loblolly (P. taeda)
pines. In New England, pitch pine (P. rigida Miller) is a favorite host in
pine barrens.

Distribution Nantucket pine tip moth occurs in the United States from eastern Texas
north to Missouri, east to Florida and north to Massachusetts. There is
also an isolated, invasive population in southern California.

Images of Nantucket
Pine Tip Moth

Figure 1. Adult of Nantucket pine tip moth, Rhyacionia frustrana. (USDA Forest
Service Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 483

Images of Nantucket Figure 2. Egg of

Nantucket pine tip
Pine Tip Moth (continued) moth. (Darrell Ross,
Oregon State University,
Bugwood.org)

Figure 3. Larva of
Nantucket pine tip
moth. (Darrell Ross,
Oregon State University,
Bugwood.org)
484 Forest Pest Insects in North America:

Images of Nantucket Figure 4. Pupa of

Nantucket pine tip
Pine Tip Moth (continued) moth. (Darrell Ross,
Oregon State University,
Bugwood.org)

Figure 5. Dead tips

caused by feeding
of Nantucket pine tip
moth. (Top: Eric R. Day,
Virginia Polytechnic
Institute and State
University,
Bugwood.org; bottom:
Darrell Ross, Oregon
State University,
Bugwood.org)
A Photographic Guide 485

Images of Nantucket Figure 6. Stunting

seedling tree due to tip
Pine Tip Moth (continued) death from Nantucket
pine tip moth feeding.
(Darrell Ross, Oregon
State University,
Bugwood.org)

Important Biological Control Many studies have described the parasitoids that can be reared from
Agents Related to this larvae and pupae of Nantucket pine tip moth. Of the various larval
Pest Species parasitoids noted, the species that are consistently of greatest importance
in nearly all studies have been, in order of importance, the ichneumonid
Campoplex frustranae Cushman, the tachinid Lixophaga mediocris
Aldrich, and the eurytomid Eurytoma pini Bugbee (see Van Driesche
et al. 1996 for details on natural enemies of this moth). Introduction of
C. frustranae to California greatly reduced the density of the invasive
population of this species found in that area.

Web Links for Information http://edis.ifas.ufl.edu/in581; a fact sheet of the University of Florida on
on Nantucket Pine Tip Moth biology and control.
http://na.fs.fed.us/spfo/pubs/fidls/nantucket/nantucket.htm; USDA Forest
Service leaflet on biology and control.
486 Forest Pest Insects in North America:

Articles on Nantucket Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Pine Tip Moth Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations. FHTET
96-19, December 1996, USDA Forest Service, Morgantown, WV
(see http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
McCravy, K.W. and C.W. Berisford. 2001. Effects of vegetation control
on parasitoids of the Nantucket pine tip moth, Rhyacionia frustrana
(Lepidoptera: Tortricidae). Florida Entomologist 84: 282-287.
DeBarr, G.L., J.W. Brewer, R.S. Cameron, and C.W. Berisford. 2002.
Nantucket pine tip moth, Rhyacionia frustrana, lures and traps:
what is the optimum combination? The Nantucket pine tip moth: old
problems, new research. Proceedings of an Informal Conference, The
Entomological Society of America, Annual Meeting, Atlanta, Georgia,
USA, 12-16 December 1999. In: Berisford, C.W. and D.M. Grosman
(eds.). General Technical Report SRS-51, Southern Research Station,
USDA Forest Service, Ashville, North Carolina: 56-68.
A Photographic Guide 487

133. Pitch Pine Tip Moth, Rhyacionia rigidana (Fernald) (Lepidoptera:

Tortricidae)

Orientation to Pest Pitch pine tip moth, Rhyacionia rigidana (Fernald), is a member of
a group of very similar moths, and it resembles Nantucket pine tip
moth, R. frustrana (Fernald), in appearance and biology. But unlike R.
frustrana, this moth can attack and kill large pitch pines (Pinus rigida
Mill.). Besides pitch pines, this native North American tortricid attacks
several other native or introduced pines. The feeding of pitch pine tip
moth larvae kills branches or leaders, and may kill the entire tree. There
are three generations per year in the southern United States, but only one
in the north.

Hosts Commonly Attacked In North America, this moth attacks several native or exotic pines,
including pitch (P. rigida), Corsican (P. nigra J. F. Arnold), Virginia (P.
virginiana Mill.), red (P. resinosa Sol. ex Aiton), Scotch (P. sylvestris L.),
loblolly (P. taeda L.), and slash (P. elliottii Engelm.) pines.

Distribution This moth is found from Georgia to Texas, north to Missouri, New York,
and Maine.

Image of
Pitch Pine Tip Moth

Figure 1. Adult of pitch pine tip moth, Rhyacionia rigidana. (Jim Vargo)
488 Forest Pest Insects in North America:

Important Biological Control Little to no information is available on the natural enemies of this
Agents Related to this species, apart from some parasitoids reported from this species in
Pest Species Missouri (Kearby and Taylor, 1975).

Web Links for Information None

on Pitch Pine Tip Moth
Articles Berisford, C.W. 1974. Species isolation mechanisms in Rhyacionia
frustrana and R. rigidana. Annals of the Entomological Society of
America 67: 292-294.
Kearby, W.H. and B. Taylor. 1975. Larval and pupal parasites reared
from tip moths of the genus Rhyacionia in Missouri (Lepidoptera:
Olethreutidae). Journal of the Kansas Entomological Society 48:
206-211.
Gargiullo, P.M. and C.W. Berisford 1983. Life tables for the Nantucket
pine tip moth, Rhyacionia frustrana (Comstock), and the pitch pine
tip moth, Rhyacionia rigidana (Fernald) (Lepidoptera: Tortricidae).
Environmental Entomology 12: 1391-1402.
A Photographic Guide 489

134. Southwestern Pine Tip Moth, Rhyacionia neomexicana (Dyar)

(Lepidoptera: Tortricidae)

Orientation to Pest The southwestern pine tip moth, Rhyacionia neomexicana (Dyar), is a
native North American tip moth that distorts and kills terminals of young
ponderosa pine (Pinus ponderosa Douglas ex Lawson) in Arizona, New
Mexico, Colorado, the Dakotas, and Nebraska. Moths fly in spring laying
eggs as new needles emerge. Eggs hatch in 2-3 weeks, and small larvae
feed inside needles. Later, larvae feed inside needle sheaths or buds and
eventually the large larvae hollow out growing shoots. Larval feeding
under the bark of new shoots produces girdling wounds that cause shoots
to turn brown and become crooked. Mature larvae leave the tips during
summer and spin cocoons, usually in the bark crevices on the base of
the tree below the litter, where they pupate and overwinter. There is
one generation per year. Heavy infestation for consecutive years may
retard growth, leaving trees short and bushy. Trees are affected in both
plantations and natural forests. Injury is most severe where trees are
planted on poor sites.

Hosts Commonly Attacked In North America, this moth attacks ponderosa (P. ponderosa), Austrian
(P. nigra J. F. Arnold), Mugo (P. mugo Turra), Scots (P. sylvestris L.),
and foxtail (P. balfouriana Balf.) pines.

Distribution This moth occurs in the United States in Arizona, New Mexico, Utah,
Colorado, the Dakotas, Montana, and Nebraska.

Image of Southwestern
Pine Tip Moth

Figure 1. Adult of southwestern pine tip moth, Rhyacionia neomexicana.

(Jim Vargo)
490 Forest Pest Insects in North America:

Important Biological Control Little to no information is available on the natural enemies of this
Agents Related to this species.
Pest Species

Web Links for Information on http://wiki.bugwood.org/HPIPM:Rhyacionia_neomexicana; Bugwood

Southwestern Pine Tip Moth Wiki fact sheet on biology and control.
http://www.ext.colostate.edu/pubs/insect/05529.html; Colorado State
University fact sheet, which compares this species to related tip moths.
http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/fsbdev2_026053.pdf;
USDA Forest Service pest leaflet that includes a distribution map for
the species.

Articles Jennings, D.T. 1975. Life history and habits of the southwestern pine
tip moth, Rhyacionia neomexicana (Dyar) (Lepidoptera: Olethreutidae).
Annals of the Entomological Society of America 68: 597-606.
Stevens, R.E. and D.T. Jennings. 1977. Western pine-shoot borer: a threat
to intensive management of ponderosa pine in the Rocky Mountain area
and Southwest. USDA Forest Service, General Technical Report, Rocky
Mountain Forest and Range Experiment Station, RM-45.
Wagner, M.R. and Z. Chen. 2004. Long-term benefits to the growth
of ponderosa pines from controlling southwestern pine tip moth
(Lepidoptera: Tortricidae) and weeds. Journal of Economic Entomology
97: 1972-1977.
A Photographic Guide 491

135. Eastern Pine Shoot Borer, Eucosma gloriola Heinrich (Lepidoptera:

Tortricidae)

Orientation to Pest Eastern shoot borer, Eucosma gloriola Heinrich, is a tortricid moth,
native to North America, whose larvae attack shoots of various native
or introduced pines. Larvae tunnel in new shoots, killing them. The
shape of the tree’s main trunk becomes deformed when the terminal
leader is killed. This moth is more common in plantations and artificially
reforested areas than in native forests. In plantations, 7-41 percent of
shoots and up to 10 percent of terminal leaders may be infested.

Hosts Commonly Attacked In North America, this moth attacks various native or introduced pines,
including eastern white (Pinus strobus L.), jack (P. banksiana Lamb.),
red (P. resinosa Sol. ex Aiton), Austrian (P. nigra J. F. Arnold), mugo
(P. mugo Turra), and Scots (P. sylvestris L.) pines.

Distribution In North America, this moth is found from the northeastern United States
west to the Great Lakes states, as well as in southern Canada.

Images of Eastern Figure 1. Adult of

eastern pine shoot
Pine Shoot Borer borer, Eucosma gloriola.
(Tom Murray)

Figure 2. Larva of
eastern pine shoot
borer. (Louis Wilson,
USDA Forest Service)

Figure 3. Exit hole in

leader of mature larvae
of eastern pine shoot
borer. (Louis Wilson,
USDA Forest Service)
492 Forest Pest Insects in North America:

Images of Eastern Pine

Shoot Borer (continued)

Figure 4. Dead leader (seen as short stub at base of top whorl of branches)
caused by eastern pine shoot borer. (A. Steven Munson, USDA Forest Service,
Bugwood.org)

Figure 5. Deformity
of main trunk of pine
caused by eastern pine
shoot borer. (Minnesota
Department of Natural
Resources Archive,
Minnesota Department
of Natural Resources,
Bugwood.org)
A Photographic Guide 493

Important Biological Control Five species of natural enemies of this borer have been recorded in
Agents Related to this surveys in Pennsylvania, New York, Ontario, and Manitoba: Glypta
Pest Species sp., Rhorus sp. (both ichneumonids), Bracon rhyacioniae (Muesebeck)
(Braconidae), Elachertus cidariae Ashmead (Eulophidae), and
Habrocytus sp. (Pteromalidae). Of these, Glypta sp. (perhaps Glypta
eucosmae Walley and Baron) may be the most important. Why this
insect, which causes little damage in forests, becomes damaging in
plantations is not understood.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/eps_borer/eps_borer.htm; USDA

on Eastern Pine Shoot Borer Forest Service, Forest Insect and Disease Leaflet 134, covering biology
and control.
http://ento.psu.edu/extension/christmas-trees/information/pest-fact-
sheets/eastern-pine-shoot-borer-eucosma-gloriola-heinric; fact sheet of
Pennsylvania State University of biology and control.

Articles Drooz, A.T. 1960. White pine shoot borer (Eucosma gloriola Heinrich).
Journal of Economic Entomology 53: 248-251.
Newman, J.H. 1968. First records of the white pine shoot borer,
Eucosma gloriola (Lepidoptera: Olethreutidae), in Michigan. Michigan
Entomologist 1: 267-270.
Wong, H.R. and A.E. Campbell. 1967. The larval feeding habits of the
eastern pine-shoot borer, Eucosma gloriola Heinrich (Lepidoptera:
Tortricidae), in jack pine regeneration in Michigan. Manitoba
Entomologist 1: 42-46.
DeBoo, R.F., W.L. Sippell, and H.R. Wong. 1971. The eastern pine-shoot
borer, Eucosma gloriola (Lepidoptera: Tortricidae), in North America.
The Canadian Entomologist 103: 1473-1486.
494 Forest Pest Insects in North America:

136. Lodgepole Cone Moth, Eucosma rescissoriana Heinrich (Lepidoptera:

Tortricidae)

Orientation to Pest The lodgepole cone moth, Eucosma rescissoriana Heinrich, is found
in northwestern North America south to central California. It is highly
destructive to cones and seeds of western white pine (Pinus monticola
Douglas). In British Columbia, adults emerge in May, and oviposition
occurs in early June; in Idaho, oviposition is in June. Eggs are laid under
scale tips of second-year cones. Larvae burrow in cones of host plants,
consuming the seeds. Pupae overwinter, presumably, in the duff. There is
one generation per year. While a pest in seed orchards in North America,
it is considered to be a potential biological control agent of P. monticola
in New Zealand, where this pine is invasive outside of plantations.

Hosts Commonly Attacked In North America, this moth attacks western white pine (P. monticola).

Distribution This moth occurs in North America from British Columbia and western
Alberta south as far as central California and New Mexico.

Images of
Lodgepole Cone Moth

Figure 1. Adult of lodgepole cone moth, Eucosma rescissoriana. (Ward Strong,

BC Ministry of Forests, Bugwood.org)

Figure 2. Eggs of
lodgepole cone moth
on cone scale. (Ward
Strong, BC Ministry of
Forests, Bugwood.org)
A Photographic Guide 495

Images of Lodgepole
Cone Moth (continued)

Figure 3. Larvae of lodgepole cone moth hatching from eggs on cone scale.
(Ward Strong, BC Ministry of Forests, Bugwood.org)

Figure 4. Larvae of lodgepole cone moth inside cone. (Ward Strong, BC Ministry
of Forests, Bugwood.org)

Figure 5. Appearance of
early stage of damage
to cone of western white
pine by lodgepole cone
moth. (Ward Strong,
BC Ministry of Forests,
Bugwood.org)
496 Forest Pest Insects in North America:

Images of Lodgepole Figure 6. Internal

feeding and damage
Cone Moth (continued) to cone of western
white pine caused by
lodgepole cone moth.
(Ward Strong, BC
Ministry of Forests,
Bugwood.org)

Important Biological Control Twelve species of parasitic insects were found to be associated with E.
Agents Related to this rescissoriana in western white pine cones in northern Idaho, of which the
Pest Species three most abundant species were Pimplopterus n. sp. (Ichneumonidae),
Chelonus petrovae McComb (Braconidae), and Apanteles starki Mason
(Braconidae). Total parasitism ranged from 9.4 to 40.9 percent in a seed-
production area over three years.

Web Links for Information None

on Lodgepole Cone Moth

Articles Ollieu, M.M. and J.A. Schenk. 1966. The biology of Eucosma
rescissoriana Heinrich in western white pine in Idaho (Lepidoptera:
Olethreutidae). The Canadian Entomologist 98: 268-274.
Goyer, R.A. and J.A. Schenk. 1969. Parasitism of the cone moth
Eucosma rescissoriana (Lepidoptera: Olethreutidae) in northern Idaho.
The Canadian Entomologist 101: 1063-1069.
Brockerfoff, E.G. and M. Kay. 1998. Prospects and risks of biological
control of wilding Pinus contorta in New Zealand. Proceedings of the
51st New Zealand Plant Protection Conference: 216-223.
A Photographic Guide 497

137. Western Pine Shoot Borer, Eucosma sonomana Kearfott (Lepidoptera:

Tortricidae)

Orientation to Pest The western pine shoot borer, Eucosma sonomana Kearfott, is a
significant pest of ponderosa (Pinus ponderosa Douglas ex C. Lawson)
and lodgepole (P. contorta Douglas ex Louden) pines in many parts of
the western North America, and is especially damaging to young, open-
grown trees. Larvae bore in terminal shoots, feeding exclusively in
the pith. As a result, they stunt the growth but rarely kill the terminals.
Larvae leave shoots in late spring, drop to the ground, and pupate.
Repeated attacks lead to shorter trees and may cause up to a 20 percent
loss in volume growth over a rotation.

Hosts Commonly Attacked In North America, this moth attacks ponderosa (P. ponderosa) and
lodgepole (P. contorta) pines.

Distribution In North America, this moth occurs is recorded from Arizona, New
Mexico, Wyoming, Colorado, Utah, South Dakota, Montana, Idaho,
California, Oregon, Washington, and British Columbia (see map in
Sartwell et al., 1980).

Images of Western Figure 1. Adult of

western pine shoot
Pine Shoot Borer borer, Eucosma
sonomana.
(Scott Tunnock,
USDA Forest Service,
Bugwood.org)

Figure 2. Larva of
western pine shoot borer
in leader. (Darrell Ross,
Oregon State University,
Bugwood.org)
498 Forest Pest Insects in North America:

Images of Western
Pine Shoot Borer
(continued)

Figure 3. Tunnels of western pine shoot borer in lodgepole pine. (Left: USDA
Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org; right:
Darrell Ross, Oregon State University, Bugwood.org)

Figure 4. Terminal
of pine damaged by
western pine shoot
moth. (Darrell Ross,
Oregon State University,
Bugwood.org)
A Photographic Guide 499

Important Biological Control No information was available on natural enemies of this species.
Agents Related to this
Pest Species
Web Links for Information None
on Western Pine Shoot Borer
Articles Stevens, R.E. and D.T. Jennings. 1977. Western pine shoot borer: A
threat to intensive management of ponderosa pine in the Rocky Mountain
area and southwest. USDA Forest Service, Rocky Mountain Forest and
Range Experiment Station, Fort Collins, Colorado, General Technical
Report RM-45: 8 p.
Sartwell, C., G.E. Daterman, T.W. Koerber, R.E. Stevens, and L.L.
Sower. 1980. Distribution and hosts of Eucosoma sonomana in the
western United States as determined by trapping with synthetic sex
attractants. Annals of the Entomological Society of America 73: 254-256.
500 Forest Pest Insects in North America:

138. Douglas-fir Cone Moth, Barbara colfaxiana (Kearfott) (Lepidoptera:

Tortricidae)

Orientation to Pest The Douglas-fir cone moth, Barbara colfaxiana (Kearfott), is found
in western North America from California to British Columbia, east to
Montana, and Colorado. Adults emerge in the spring and lay their eggs
on young cones. Young larvae feed largely on cone scales, but older
larvae consume developing seeds. Pupae overwinter in papery, resin-
coated cocoons in center of cones. Pupae may remain in diapause for as
long as 3 years. Damage occurs on Douglas-fir (Pseudotsuga menziesii
[Mirb.] Franco) and true firs (Abies) and is greatest at inland (non-
coastal) sites, where summers are hotter and drier. One larva can destroy
60 percent of the seeds in a cone, while three larvae will likely destroy
all of the seeds. There is one generation per year. Pheromone traps using
the pheromone of Contarinia oregonensis Foote, can be used to monitor
the pest.

Hosts Commonly Attacked In North America, this moth attacks Douglas-fir (P. menziesii) and
various true firs (Abies).

Distribution In North America, this moth occurs is recorded from California to British
Columbia, east to Montana, and Colorado.

Images of Figure 1. Adults of

Douglas-fir cone moth,
Douglas-fir Cone Moth Barbara colfaxiana.
(D. Manastyrski,
Bugwood.org)

Figure 2. Larva of
Douglas-fir cone moth.
(Julie Brooks,
Bugwood.org)
A Photographic Guide 501

Images of Douglas-fir Figure 3. Pupa of

Douglas-fir cone moth.
Cone Moth (continued) (D. Manastyrski,
Bugwood.org)

Figure 4. Douglas-fir
cone moth, Barbara
colfaxiana, caught
in trap baited with
pheromone of Contarinia
oregonensis.
(Ward Strong,
BC Ministry of Forests,
Bugwood.org)

Important Biological Control The ichneumonid Glypta evetriae Cushman is reported as a parasitoid of
Agents Related to this this species.
Pest Species
Web Links for Information http://www.forestry.ubc.ca/fetch21/FRST308/lab2/barbara_colfaxiana/
on Douglas-fir Cone Moth conemoth.html; fact sheet of the University of British Columbia.
http://www.fgcouncil.bc.ca/PM-Factsheet08-Barbara-colfaxiana.pdf;
fact sheet on control of Douglas-fir cone moth.

Articles Clark, E.C., J.A. Schenk, and D.L. Williamson. 1963. The cone-infesting
moth Barbara colfaxiana as a pest of Douglas-fir in northern Idaho.
Annals of the Entomological Society of America 56: 246-250.
Hedlin, A.F. 1974. Cone and seed insects of British Columbia. Canadian
Forestry Service, Pacific Forestry Research Centre, Victoria, BC.
BC-X-90: 63 p.
Nebeker, T.E. 1977. A partial life table for the Douglas-fir cone moth,
Barbara colfaxiana (Lepidoptera: Olethreutidae). The Canadian
Entomologist 109: 943-951.
502 Forest Pest Insects in North America:

Articles on Douglas-fir Sweeney, J.D. and G.E. Miller. 1989. Distribution of Barbara colfaxiana
Cone Moth (continued) (Kearfott) (Lepidoptera: Tortricidae) eggs within and among Douglas-
fir crowns and methods for estimating egg densities. The Canadian
Entomologist 121: 569-578.
Koerber, T.W. and G.P. Markin. 1984. Metasystox-R® injections increase
seed yield of Douglas-fir in California, Oregon, and Washington.
In: Yates, H.O., III (ed.). Proceedings of the Cone and Seed Insects
Working Party Conference. Asheville, North Carolina, Southeastern
Forest Experiment Station: 137-146.
A Photographic Guide 503

139. Pine Webworm, Pococera robustella (Zeller) (Lepidoptera: Pyralidae)

Orientation to Pest Pine webworm, Pococera robustella (Zeller), is a native North American
moth. Females lay groups of one to 10 overlapping eggs on host needles.
Young larvae are pine needle miners, but older larvae feed externally,
building webs around groups of needles. Larvae leave the nest to
clip needles, which they bring back into the nest to consume. Mature
larvae move to the ground and spin cocoons in which they pupate.
Several species of pines are attacked. In southern slash pine (Pinus
elliottii Engleman) plantations, damage occurs mainly in the first year
after planting. In northern parts of the United States, damage occurs
Pine webworm
in jack (P. banksiana Lamb.) and red pine (P. resinosa Sol. ex Aiton)
plantations, but is mostly of concern to Christmas tree growers due to
the unsightliness of the webs. There are one (in the north) to three (in the
south) generations per year.

Hosts Commonly Attacked In North America, this moth attacks many pine species but is most
common on slash (P. elliottii), red (P. resinosa) and jack (P. banksiana)
pines.

Distribution This moth occurs in southern Canada and throughout most of the eastern
United States.

Images of Figure 1. Adult

pine webworm,
Pine Webworm Pococera robustella.
(Natasha Wright,
Florida Department
of Agriculture and
Consumer Services,
Bugwood.org

Figure 2. Larva of pine

webworm. (Connecticut
Agricultural Experiment
Station Archive,
Connecticut Agricultural
Experiment Station,
Bugwood.org)
504 Forest Pest Insects in North America:

Images of Pine Figure 3. Feeding

web of larvae of pine
Webworm (continued) webworm.
(Robert L. Anderson,
USDA Forest Service,
Bugwood.org)

Figure 4. Damage to
a young pine seedling
caused by pine
webworm.
(Robert L. Anderson,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 505

Important Biological Control The ichneumonid Syzeuctus elegans (Cresson) is a common parasitoid
Agents Related to this of this species. Other natural enemies include other parasitic wasps,
Pest Species parasitic flies, predatory insects and birds.

Web Links for Information http://entnemdept.ufl.edu/creatures/trees/moths/pine_webworm.htm;

on Pine Webworm fact sheet of the University of Florida, covering biology and control.
http://wiki.bugwood.org/Pococera_robustella; fact sheet of Bugwood
wiki covering biology and control.
http://www.freshfromflorida.com/pi/enpp/ento/entcirc/ent419.pdf;
fact sheet of Florida Department of Agriculture and Consumer Services,
Division of Plant Industry.

Articles Hertel, G.D. and D.M. Benjamin. 1979. Biology of the pine webworm
in Florida slash pine plantations. Annals of the Entomological Society of
America 72: 816-819.
Wallesz, D.P. and D.M. Benjamin. 1960. The biology of the pine
webworm, Tetralopha robustella, in Wisconsin. Journal of Economic
Entomology 53: 587-589.
506 Forest Pest Insects in North America:

140. Zimmerman Pine Moth, Dioryctria zimmermani (Grote) (Lepidoptera:

Pyralidae)

Orientation to Pest Zimmerman pine moth, Dioryctria zimmermani (Grote), is one member
of a group of closely related native North American species that have
been poorly distinguished and still await further clarification in some
cases. Names in the older literature may be unreliable. Adults of D.
zimmermani emerge in mid-summer and lay their eggs in various places
on the tree trunk or buds. Upon hatching, larvae immediately settle into
bark crevices and overwinter in hibernacula. Larvae resume feeding the
following spring by tunneling into newly formed terminals or lateral
shoots, which become hooked and discolored. Larvae feed on the
cambial layer and outer xylem of all common pine species. Feeding sites
are distinguished by large masses of accumulated frass and pitch. When
mature, larvae leave the terminals and tunnel into the whorl, girdling tips
or branches. Pupation takes place in the pitch masses associated with
larval feeding.

Hosts Commonly Attacked In North America, this moth attacks most common pines, but damage
is principally important in Christmas tree plantations of Scotch (Pinus
syvestris L.), red (P. resinosa Sol. ex Aiton), and Austrian (P. nigra J. F.
Arnold) pines in the north central United States and southeastern Canada.

Distribution In North America, this moth occurs in southeastern Canada and the
northeastern United States, as far west as Minnesota.

Images of
Zimmerman Pine Moth

Figure 1. Adult of Zimmerman pine moth, Dioryctria zimmermani. (Whitney

Cranshaw, Colorado State University, Bugwood.org)
A Photographic Guide 507

Images of Zimmerman Figure 2. Larva of

Zimmerman pine
Pine Moth (continued) moth. (Phil Pellitteri,
University of Wisconsin,
Entomology Dept,
Bugwood.org)

Figure 3. Feeding of Zimmerman pine moth larvae results in pitch accumulations

on the bark, usually near the branch nodes. (Left: Rayanne Lehman,
Pennsylvania Department of Agriculture, Bugwood.org; right: Whitney Cranshaw,
Colorado State University, Bugwood.org)

Figure 4. Pupal skin of Zimmerman pine moth inside pitch mass (opened).
(Phil Pellitteri, University of Wisconsin, Entomology Dept, Bugwood.org)
508 Forest Pest Insects in North America:

Important Biological Control Various parasitoids are listed in the literature as attacking this species.
Agents Related to this But many of these studies predate the clarification of species boundaries
Pest Species in this group and so it is unclear exactly which species these records refer
to. Regardless, total parasitism in most studies did not exceed 5 percent.

Web Links for Information http://greenindustry.uwex.edu/problemdetails.cfm?problemid=99;

on Zimmerman Pine Moth fact sheet of the University of Wisconsin on recognition and control.
http://ento.psu.edu/extension/christmas-trees/information/pest-fact-
sheets/zimmerman-pine-moth; fact sheet of Pennsylvania State
University on biology and control.
http://extension.psu.edu/ipm/program/christmas-tree/pest-fact-sheets/
stem-and-root-injury/zimmerman.pdf; fact sheet of Pennsylvania State
University on biology and control.

Articles Yonker, J.W. and D.L. Schuder. 1980. The biology of the Zimmerman
pine moth (Dioryctria zimmermani) in Indiana landscapes with reference
to its control. Proceedings of the Indiana Academy of Science 89:
207-208.
Mutuura, A. 1982. American species of Dioryctria (Lepidoptera:
Pyralidae). VI. A new species of Dioryctria from eastern Canada
and north-eastern United States. The Canadian Entomologist 114:
1069-1076.
A Photographic Guide 509

141. Fall Cankerworm, Alsophila pometaria (Harris) (Lepidoptera:

Geometridae)

Orientation to Pest Fall cankerworm, Alsophila pometaria (Harris), is a native moth capable
of defoliating many hardwood species. The species overwinters as
eggs, which hatch in early spring to coincide with bud break. Young
“inchworm” larvae skeletonize new leaves, while older larvae consume
all but the larger veins or midrib. By midsummer, mature larvae crawl
down or drop on silk threads to the soil where they pupate in the soil.
Adults emerge in early winter (November or early December) after initial
frosts to mate. The wingless females climb tree trunks and deposit their
eggs in rows in a single layer on smaller branches or trunk. There is one
generation per year. In some areas fall cankerworm periodically goes into
outbreak, causing widespread damage. Damage results from complete or
partial defoliation of forest or urban trees.

Hosts Commonly Attacked In North America, larvae of this defoliating moth attack a wide range
of hardwoods and shrubs, including elm (Ulmus), maple (Acer), ash
(Fraxinus), oak (Quercus), apple (Malus domestica Borkh.), hickory
(Carya tomentosa [Poir.] Nutt.), dogwood (Cornus), and many others.

Distribution This moth has a wide distribution in southeastern Canada and the eastern
United States, west to Alberta, Montana, and Missouri, and south to
North Carolina.

Images of Figure 1. Adults of fall

cankerworm, Alsophila
Fall Cankerworm pometaria; top, wingless
female; bottom, male.
(Top: James B. Hanson,
USDA Forest Service,
Bugwood.org; bottom:
USDA Forest Service
- Region 8 - Southern
Archive, USDA Forest
Service, Bugwood.org)
510 Forest Pest Insects in North America:

Images of Fall
Cankerworm (continued)

Figure 2. Egg mass of fall cankerworm. Note straight rows of eggs in single layer.
(Pennsylvania Department of Conservation and Natural Resources - Forestry
Archive, Bugwood.org)

Figure 3. Larvae of fall

cankerworm; top, young
larva; bottom, mature
larvae. (Top: A. Steven
Munson, USDA Forest
Service, Bugwood.org;
bottom: E. Bradford
Walker, Vermont
Department of Forests,
Parks and Recreation,
Bugwood.org)
A Photographic Guide 511

Images of Fall
Cankerworm (continued)

Figure 4. Feeding of fall cankerworm. (USDA Forest Service - Ogden Archive,

USDA Forest Service, Bugwood.org)

Figure 5. Defoliation
Fall cankerworm of hardwoods by fall
cankerworms. (James B.
Hanson, USDA Forest
Service, Bugwood.org)

Figure 6. Sticky bands

on shade trees to
trap fall cankerworm
females. (G. Keith
Douce, University of
Georgia, Bugwood.org)
512 Forest Pest Insects in North America:

Important Biological Control Relatively few parasitoids have been recorded attacking this species.
Agents Related to this Among these are the braconid Meteorus autographae Muesbeck and
Pest Species the scelionid egg parasitoid Telenomus alsophilae Viereck. The latter is
believed to be the most important parasitoid affecting this species, killing
15-20 percent of the eggs. A nuclear polyhedrosis virus is also associated
with this species and some strains of Bacillus thuringiensis Berliner are
effective against it.

Web Links for Information http://ento.psu.edu/extension/factsheets/fall-cankerworm; fact sheet of

on Fall Cankerworm Pennsylvania State University on biology and control.
http://www.maine.gov/doc/mfs/fallcank.htm; fact sheet of the Maine
Department of Conservation on biology and control.
http://www.umassgreeninfo.org/fact_sheets/defoliators/cankerworm.
html; Extension fact sheet from the University of Massachusetts.
http://ohioline.osu.edu/hyg-fact/2000/2558.html; Extension fact sheet of
the Ohio State University comparing spring and fall cankerworms.

Articles Fedde, G.F. 1980. Spring parasitism of fall cankerworm eggs in northern
Georgia by Telenomus alsophilae (Hymenoptera: Scelionidae). Journal
of the Georgia Entomological Society 15: 199-206.
Miller, F., K. Malmquist, and G. Ware. 2001. Evaluation of Asian,
European, and North American elm (Ulmus spp.) biotypes to feeding by
spring and fall cankerworms. Journal of Environmental Horticulture 19:
216-221.
LaFrance, K.R. and A.R. Westwood. 2006. An assessment of tree
banding techniques to capture cankerworm defoliators of elm and ash
trees in Winnipeg, Manitoba, Canada. Arboriculture and Urban Forestry
32: 10-17.
A Photographic Guide 513

142. Spring Cankerworm, Paleacrita vernata (Peck) (Lepidoptera:

Geometridae)

Orientation to Pest Spring cankerworm, Paleacrita vernata (Peck), is a native North

American moth that defoliates many hardwood species and in some
locations goes into periodic outbreaks. Damage is caused by young
“inchworm” larvae skeletonizing new leaves and older larvae consuming
all but the larger veins or midrib. This species may be confused with
the very similar fall cankerworm, Alsophila pometaria (Harris). Adults
emerge from pupal cells in the soil in early spring when the ground is
thawed. Females crawl up tree trunks to lay eggs, which are deposited
in loose clusters of 100 or more in bark crevices or under bark scales.
Eggs hatch in early spring and larvae feed on leaves through the end of
spring. Mature larvae drop to the soil on silk threads in summer. Larvae
remain dormant within their soil-covered pupal cells throughout the rest
of the summer and fall, pupating in late fall or early spring. There is one
generation per year.

Hosts Commonly Attacked In North America, larvae of this moth feed on many hardwood species,
including elm (Ulmus), green ash (Fraxinus pennsylvanica Marsh.),
maple (Acer), bur oak (Quercus macrocarpa Michx.), linden/basswood
(Tilia), silver birch (Betula pendula Roth), and apple (Malus domestica
Borkh.).

Distribution This geometrid is found in southeastern Canada, as well as throughout

the northeastern and north central United States, and west to parts of
Texas, Colorado, and California.

Images of Spring Figure 1. Adults of

spring cankerworm,
Cankerworm Paleacrita vernata; top,
wingless female; bottom,
male. (Top: Mark
Ascerno, University of
Minnesota, University of
Minnesota • Extension;
bottom: Nolie Schneider,
Bugwood.org)
514 Forest Pest Insects in North America:

Images of Spring Figure 2. Larvae of

spring cankerworm.
Cankerworm (continued) (Top: James B. Hanson,
USDA Forest Service,
Bugwood.org; bottom:
USDA Forest Service
- Northeastern Area
Archive, USDA Forest
Service, Bugwood.org)

Figure 3. Tree trunk with band trap to catch female spring cankerworms as they
crawl up trees. (William A. Carothers, USDA Forest Service, Bugwood.org)
A Photographic Guide 515

Important Biological Control Several species of parasitoids are known from spring cankerworm,
Agents Related to this including three braconids, Rogas geometrae Ashmead, Apanteles
Pest Species paleacritae Riley, and Meteorus hyphantriae Riley. It is also attacked
by the ichneumonids Phobcampe geometrae (Ashmead) and Hyposoter
fuscitarsis (Vierieck). Bacillus thuringiensis Berliner is bacterium that
has been formulated as a biopesticide that can be used to control larvae
of this species.

Web Links for Information http://ohioline.osu.edu/hyg-fact/2000/2558.html; Extension fact sheet of

on Spring Cankerworm the Ohio State University comparing spring and fall cankerworms.
http://www.extension.umn.edu/distribution/horticulture/dg0876.html;
fact sheet of University of Minnesota extension service comparing spring
and fall cankerworms.
http://insects.tamu.edu/fieldguide/cimg293.html; fact sheet of the Texas
A&M University on biology and control.
http://entomology.unl.edu/ornamentals/pestprofiles/scankerworm.shtml;
fact sheet of the University of Nebraska on biology and control.

Articles Frye, R.D., D.K. McBride, D.R. Carey, T.L. Elichuk, and R.L. Dregseth.
1977. Cankerworm control in shelterbelts. North Dakota Farm Research
34(6): 3-7.
LaFrance, K.R. and A.R. Westwood. 2006. An assessment of tree
banding techniques to capture cankerworm defoliators of elm and ash
trees in Winnipeg, Manitoba, Canada. Arboriculture and Urban Forestry
32: 10-17.
516 Forest Pest Insects in North America:

143. Eastern Hemlock Looper, Lambdina fiscellaria fiscellaria Guenée

(Lepidoptera: Geometridae)

Orientation to Pest Eastern hemlock looper, Lambdina fiscellaria fiscellaria Guenée, is a

native North American geometrid whose larvae defoliate eastern hemlock
(Tsuga canadensis [L.] Carrière), balsam fir (Abies balsamea [L.] Mill.)
and white spruce (Picea glauca [Moench] Voss). It is found in Canada
from Newfoundland to Alberta and in the United States it occurs widely
within the range of eastern hemlock. Adults fly in late summer or fall
and lay eggs singly or in small groups on bark, or associated moss and
lichens. Winter is spent in the egg stage and larvae begin to feed in the
late spring on newly opened clusters of needles. Older larvae also feed
on old needles. Some attacked needles are chewed but not severed and
remain attached, giving the tree a brown appearance. Pupation usually
takes place on the bark. Repeated periodic outbreaks have occurred over
wide areas in eastern Canada and the northeast and north central United
States, especially Newfoundland, Wisconsin, and Michigan.

Hosts Commonly Attacked This moth feeds mainly on eastern hemlock (T. canadensis) (especially
in the more southerly parts of it range) but in the north also feeds on
balsam fir (A. balsamea) and white spruce (P. glauca). During outbreaks
it sometimes feeds on other conifers and even some hardwoods.

Distribution This geometrid is found in Canada from Newfoundland to Alberta, and in

the United States it occurs widely within the range of eastern hemlock.

Images of Eastern Figure 1. Adult of

eastern hemlock looper,
Hemlock Looper Lambdina fiscellaria
fiscellaria. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)

Figure 2. Eggs of
eastern hemlock looper.
(Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)
A Photographic Guide 517

Images of Eastern
Hemlock Looper (continued)

Figure 3. Larva of eastern hemlock looper. (Connecticut Agricultural Experiment

Station Archive, Connecticut Agricultural Experiment Station, Bugwood.org)

Figure 4. Defoliation from eastern hemlock looper in Michigan. (USDA Forest

Service - Northeastern Area Archive, USDA Forest Service, Bugwood.org)

Important Biological Control Because outbreaks occur of this species defoliator, its natural enemies
Agents Related to this have been studied on various occasions. Before 1950, the most common
Pest Species parasitoid attacking it was the braconid larval parasitoid Apanteles
sp. nr. flavovariatus Muesbeck. But subsequently a parasitic tachinid,
Winthemia occidentis Reinhard, associated with another hemlock looper
in western North America, was introduced and is now the dominant
parasitoid of eastern hemlock looper. Pupal parasitism, while reaching up
to 30 percent or more, is due to a collection of polyphagous parasitoids.
518 Forest Pest Insects in North America:

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=8846; fact

on Eastern Hemlock Looper sheet of Natural Resources Canada on phenology and extent of damage
in Canada.
http://www.glfc.forestry.ca/VLF/ForumPresentations/
heberthemlocklooper_08e.pdf; detailed history of outbreaks in Canada.

Articles MacLean, D.A. and E. Paul Ebe. 1999. The impact of hemlock looper
(Lambdina fiscellaria fiscellaria [Guen.]) on balsam fir and spruce in
New Brunswick, Canada. Forest Ecology and Management 120: 77-87
Hartling, L.K., N. Carter, and J. Proude. 1999. Spring parasitism of
overwintered eggs of Lambdina fiscellaria fiscellaria (Lepidoptera:
Geometridae) by Telenomus near alsophilae (Hymenoptera: Scelionidae).
The Canadian Entomologist 131: 421-422.
VanFrankenhuyzen, K., R.J. West, and M. Kenis. 2001. Lambdina
fiscellaria fiscellaria (Guenée), hemlock looper (Lepidoptera:
Geometridae). In: Mason, P.G. and J.T. Huber (eds.). Biological Control
Programmes in Canada, 1981-2000. CABI Publishing, Wallingford, UK:
141-144.
Iqbal, J., D.A. MacLean, and J.A. Kershaw, Jr. 2011. Impacts of hemlock
looper defoliation on growth and survival of balsam fir, black spruce and
white birch in Newfoundland, Canada. Forest Ecology and Management
261: 1106-1114.
A Photographic Guide 519

144. Western Hemlock Looper, Lambdina fiscellaria lugubrosa (Hulst)

(Lepidoptera: Geometridae)

Orientation to Pest Western hemlock looper, Lambdina fiscellaria lugubrosa (Hulst), is

a native North American geometrid whose larvae defoliate western
hemlock (Tsuga heterophylla [Raf.] Sarg.) in coastal areas of Oregon,
Washington, and British Columbia. Adults fly in late summer or fall
and lay eggs on bark, or associated moss and lichens or even downed
logs. Winter is spent in the egg stage and larvae begin to feed in the
late spring. Young larvae may feed on understory vegetation of various
kinds of trees or shrubs. Initial feeding on hemlock is concentrated in
the buds. In summer, older larvae attack old needles, some of which
are notched but not severed. These remain attached, giving the tree
a brown appearance. Mature larvae spin silk threads and drop to the
ground to pupate in the late summer. New moths emerge in fall to lay
overwintering eggs. Repeated large outbreaks occurred over wide areas
in western North America in the past, damaging large volumes of old
growth hemlock. However, since old growth hemlock stands in many
areas have been logged, outbreaks have become smaller.

Hosts Commonly Attacked This moth feeds mainly on western hemlock (T. heterophylla), but during
outbreaks other conifer species and hardwood shrubs are also attacked.

Distribution This geometrid is found in western North America in coastal forests of

Oregon, Washington, and British Columbia.

Images of Western
Hemlock Looper

Figure 1. Adult of western hemlock looper, Lambdina fiscellaria lugubrosa.

(Jerald E. Dewey, USDA Forest Service, Bugwood.org)
520 Forest Pest Insects in North America:

Images of Western Figure 2. Larva of

western hemlock looper.
Hemlock Looper (Jerald E. Dewey,
(continued) USDA Forest Service,
Bugwood.org)

Figure 3. Defoliation of a
western hemlock stand
by western hemlock
looper. (Bruce Hostetler,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 521

Images of Western
Hemlock Looper
(continued)

Figure 4. Defoliation of a western hemlock stand by western hemlock looper.

(USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)

Important Biological Control Pupal parasitoids reported from this species in Alaska included eight
Agents Related to this ichnuemonids: Pimpla pedalis Cresson, P. aquilonia Cresson, P. hesperus
Pest Species (Townes), Apechthis ontario (Cresson), Itoplectis quadricingulatus
(Prov.), Masttus laplantei Mason, Cratichneumon sp. (probably C.
ashmeadi [Schulz]), and Aoplus velox occidentalis (Harrington).

Web Links for Information http://www.fs.fed.us/r1-r4/spf/fhp/field_guide/109wsthml.htm;

on Western Hemlock Looper USDA Forest Service fact sheet on recognition and biology of pest.
http://www.fs.fed.us/r1-r4/spf/fhp/mgt_guide/hemlock_looper/page2.
html; USDA Forest Service fact sheet on recognition and biology of pest.
http://www.fs.fed.us/r1-r4/spf/fhp/mgt_guide/hemlock_looper/index.
html; a USDA Forest Service manual on control of western hemlock
looper.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=1000002;
a fact sheet of Natural Resources Canada on biology and phenology of
pest.
http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5191788.
pdf; a USDA Forest Service manual on control and management of
western hemlock looper.

Articles Torgersen, T.R. 1971. Parasites of the western hemlock looper, Lambdina
fiscellaria lugubrosa (Hulst), in southeast Alaska (Lepidoptera:
Geometridae). Pan-Pacific Entomologist 47: 215-219.
Harris, J.W.E., A.F. Dawson, and R.G. Brown. 1982. The western
hemlock looper in British Columbia 1911-1980. Report BC-X-234,
Pacific Forest Research Centre, Canada: 18 p.
Mills, N.J. and M. Räther. 1990. Hemlock loopers in Canada; biology,
pest status and potential for biological control. Biocontrol News and
Information 11(3): 209-222.
522 Forest Pest Insects in North America:

145. Greenstriped Forest Looper, Melanolophia imitata (Walker)

(Lepidoptera: Geometridae)

Orientation to Pest Greenstriped forest looper, Melanolophia imitata (Walker), is a native

geometrid found in moist conifer forests of western North America.
Adults fly in spring and lay eggs singly (up to 80 per female) on tree
branches and trunks. Larvae feed on foliage of all ages, but the previous
year’s foliage is preferred and damage is concentrated in tree crowns.
In late summer larvae drop to the ground and pupate in the litter, where
they overwinter. There is one generation per year. Larvae are solitary
feeders and generally are not economically damaging. However, periodic
outbreaks have occurred in Canada (especially British Columbia) that
were destructive to western hemlock (Tsuga heterophylla [Raf.] Sarg.).
Outbreaks in the past have ended abruptly due to natural causes.

Hosts Commonly Attacked This moth feeds mainly on western hemlock (T. heterophylla), Douglas
fir (Pseudotsuga menziesii [Mirbel] Franco), western redcedar (Thuja
plicata Donn ex D. Don), true firs (Abies), and spruce (Picea).

Distribution This geometrid is found in western North America from Alaska to

southern California and, in Canada, eastward to Alberta, in humid areas.

Images of Greenstriped
Forest Looper

Figure 1. Adult of greenstriped forest looper, Melanolophia imitata. (Marius

Aurelian, Bugwood.org)
A Photographic Guide 523

Images of Greenstriped Figure 2. Larva of

greenstriped forest
Forest Looper (continued) looper. (Natural
Resources Canada,
Canadian Forest
Service)

Important Biological Control Past studies have shown that fungal pathogens such as species of
Agents Related to this Cordyceps and Entomophthora cause significant mortality to pupae in
Pest Species the soil.

Web Links for Information on http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=1000022; fact

Greenstriped Forest Looper sheet of Natural Resources Canada on phenology and biology of insect.
http://www.for.gov.bc.ca/hfp/publications/00198/green_striped_forest_
looper.htm; short fact sheet of the Ministry of Forests and Range,
Province of British Columbia.
http://cfs.nrcan.gc.ca/pubwarehouse/pdfs/3317.pdf; forest pest leaflet
on greenstriped forest looper by Pacific Forestry Centre of Natural
Resources Canada.

Articles Evans, D. 1962. Descriptions and life history of Melanolophia imitata

(Walker) (Lepidoptera: Geometridae). The Canadian Entomologist 94:
594-605.
Humphreys, N. 1986. Green-striped forest looper in British Columbia.
Canadian Forest Service, Pacific Forestry Centre, FIDS Report No. 86-9:
43 p.
524 Forest Pest Insects in North America:

146. Saddleback Looper, Ectropis crepuscularia (Denis and Schiffermüller)

(Lepidoptera: Geometridae)

Orientation to Pest Saddleback looper, Ectropis crepuscularia (Denis and Schiffermüller),

is a native North American looper present across southern Canada, the
adjacent tier of U.S. states, and the eastern United States. This species
is a generalist, feeding more on conifers in western North American and
all of Canada, but on various hardwood species in the eastern United
States. Adults fly in spring, when they lay their eggs. Larvae are solitary
feeders able to mimic twigs. Young larvae often begin feeding first on
groundcover species and understory shrubs. Larvae later move up trees
where they complete development. Mature larvae drop to the soil to
pupate in the litter, where they overwinter. There at least two generations
per year over much of the eastern United States. Larvae of this species
are generally economically damaging. Outbreaks in British Columbia
and Alaska have occurred.

Hosts Commonly Attacked In Canada and the western United States, saddleback loopers feed mostly
commonly on conifers, including hemlock (Tsuga), true firs (Abies),
spruce (Picea), and larch (Larix). In the eastern United States, broadleaf
trees are the main hosts, including apple (Malus), ash (Fraxinus), birch
(Betula), dogwood (Cornus), maple (Acer), oak (Quercus), poplar
(Populus), walnut (Juglans), and willow (Salix).

Distribution This geometrid is found in a band across North America including

southern Canada and the adjacent tier of U.S. states. In the eastern United
States, it is also found south to Florida.

Images of
Saddleback Looper

Figure 1. Adult of saddleback looper, Ectropis crepuscularia. (John Davis,

Bugwood.org)
A Photographic Guide 525

Images of Saddleback
Looper (continued)

Figure 2. Larva of saddleback looper. (Connecticut Agricultural Experiment

Station Archive, Connecticut Agricultural Experiment Station, Bugwood.org)

Important Biological Control No information on the natural enemies of this species was found.
Agents Related to this
Pest Species
Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=8578; fact
on Saddleback Looper sheet of Natural Resources Canada on biology and phenology of species.

Articles Ferris, R.L. 1993. Saddleback looper. Forest Pest Leaflet No. 23. Pacific
Forestry Centre, Canadian Forest Service: 3 p.
526 Forest Pest Insects in North America:

147. Spearmarked Black Moth, Rheumaptera hastata (L.) (Lepidoptera:

Geometridae)

Orientation to Pest Spearmarked black moth, Rheumaptera hastata (L.), is a native North
American looper found from North Dakota to British Columbia and
Alaska. It feeds mainly on birch (Betula). Adults fly in spring. In Alaska,
eggs are laid singly or in clusters on tops of leaves, from mid-June to
early July. Young larvae feed gregariously between two leaves webbed
together to make a sandwich-type shelter. Larvae feed on both new and
old-growth foliage and usually feed just on the upper side, skeletonizing
leaves. Larvae mature in July and August, drop to the ground on silken
threads and pupate in the leaf litter, where they overwinter. There is
one generation per year. This moth is a serious defoliator of paper birch
(Betula papyrifera Marsh.) in interior Alaska. Epidemic populations
have occurred at 15- to 17-year intervals, persisted for 2 years, and then
collapsed from natural causes.

Hosts Commonly Attacked This species feeds mainly on mainly on birch (Betula spp.). In Alaska,
paper birch (Betula papyrifera Marsh.) is the preferred host, but larvae
also feed on species of alder (Alnus), willow (Salix), and rose (Rosa). In
Canada, the insect also feeds on sweetgale (Myrica).

Distribution This geometrid is found throughout Canada

and Alaska, and the northern United States,
with southerly range extensions along
mountain chains (Cascades, Rockies,
Appalachians).

Figure 1. North American distribution of

spearmarked black moth, Rheumaptera hastata.
(USDA Forest Service • Forest Insect & Disease
Leaflet 156)

Images of Spearmarked Figure 2. Adult of

spearmarked black
Black Moth moth. (S. Overby
- Swedish Museum of
Natural History)
A Photographic Guide 527

Images of Spearmarked Figure 3. Life stages

(adult, eggs, larva,
Black Moth (continued) pupae) of spearmarked
black moth. (USDA
Forest Service • Forest
Insect & Disease Leaflet
156)

Figure 4. Larva of
spearmarked black
moth. (Graham Finch)

Figure 5. Tent made by

larva of spearmarked
black moth, where it
feeds. (Graham Finch)

Figure 5. Birch
defoliated by
spearmarked black
moth. (Peter A. Rush,
USDA Forest Service,
Bugwood.org)
528 Forest Pest Insects in North America:

Important Biological Control Parasitoids that have been reared from the overwintering pupae of this
Agents Related to this moth include Aoplus ruficeps vagans Provancher (Ichneumonidae),
Pest Species Coccygomimus aquilonius (Ichneumonidae), and Cratichneumon
sp. (Ichneumonidae). The parasitoid Meteorus niveitarsis Cresson
(Braconidae) has been reared from larvae.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/spear/spear.htm; USDA Forest

on Spearmarked Black Moth Service fact sheet on biology and pest status in Alaska.

Articles McGuffin, W.C. 1973. The Rheumaptera of North America (Lepidoptera:

Geometridae). The Canadian Entomologist 105: 383-389.
Werner, R.A. and B.H. Baker. 1977. Spear-marked black moth. Forest
Insect & Disease Leaflet 156. Portland, OR. U.S. Dept. of Agriculture,
Forest Service, Pacific Northwest Forest and Range Experiment Station.
Available at http://www.ncrs.fs.fed.us/pubs/viewpub.asp?key=965.
Werner, R.A.1977. Biology and behavior of the spear-marked
black moth, Rheumaptera hastata, in interior Alaska. Annals of the
Entomological Society of America 70: 328-336.
A Photographic Guide 529

148. Elm Spanworm, Ennomos subsignarius (Hübner) (Lepidoptera:

Geometridae)

Orientation to Pest Elm spanworm, Ennomos subsignarius (Hübner), is a native geometrid

moth whose polyphagous larvae feed on many species of hardwood
trees, especially hickory (Carya), oak (Quercus), and ash (Fraxinus). It
is most important as a forest pest in the Appalachian Mountains, from
Pennsylvania south, but an urban forest outbreak has also occurred in St.
John’s, Newfoundland, and Labrador. This univoltine species overwinters
as eggs, which are laid in clusters on the bole and on the underside of
branches. Eggs hatch in early spring and young larvae feed on the lower
surfaces of leaves, producing “shot-hole” damage. Older larvae consume
the whole leaf apart from the major veins. After developing through five
instars, larvae pupate in loose silk cocoons formed on the partly eaten
foliage and in bark crevices. Outbreaks of this species occur at irregular
intervals that defoliate trees over wide areas.

Hosts Commonly Attacked This species feeds on elm (Ulmus), hickory (Carya), oak (Quercus),
maple (Acer), beech (Fagus), and ash (Fraxinus).

Distribution This geometrid is found in hardwood forests from Newfoundland and

Labrador to Saskatchewan in Canada and throughout the eastern United
States, west to Michigan, Colorado, and Texas.

Images of
Elm Spanworm

Figure 1. Adults of elm spanworm, Ennomos subsignarius, mating (left) and

laying eggs (right). (Both photos: Heidi Fry, Bugwood.org)
530 Forest Pest Insects in North America:

Images of Elm Figure 2. Egg mass of

elm spanworm. (Heidi
Spanworm (continued) Fry, Bugwood.org)

Figure 3. Early (left) and late (right) instar larvae of elm spanworm. (Both photos:
Heidi Fry, Bugwood.org)

Figure 4. Masses of
elm spanworm larvae
feeding on host plant
during an outbreak.
(Bugwood.org)

Figure 5. Pupae of elm

spanworm. (Arnold T.
Drooz, USDA Forest
Service, Bugwood.org)
A Photographic Guide 531

Images of Elm Figure 6. Shot hole

or tattered damage
Spanworm (continued) from feeding of elm
spanworms. (Heidi Fry,
Bugwood.org)

Figure 7. Trees
defoliated by elm
spanworm. (Heidi Fry,
Bugwood.org)

Figure 8. Defoliation
of a forest stand by
elm spanworm.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)
532 Forest Pest Insects in North America:

Images of Elm
Spanworm (continued)

Figure 9. Adults of the egg parasitoid Telenomus droozi emerging from eggs of
elm spanworm. (Arnold T. Drooz, USDA Forest Service, Bugwood.org)

Important Biological Control Some of the more important biological control agents affecting this
Agents Related to this species are the egg parasitoid Telenomus droozi Muesebeck and the
Pest Species predatory carabid Calosoma scrutator (F.), which eats larvae.

Web Links for Information http://ento.psu.edu/extension/factsheets/elm-spanworm; fact sheet of

on Elm Spanworm Pennsylvania State University on biology and control.
http://www.fs.fed.us/r8/foresthealth/idotis/insects/elmspan.html; short
fact sheet of the USDA Forest Service Forest Health Protection Southern
Region on biology and control.

Articles Anderson, J.F. and K.K. Harry. 1976. Parasitoids and diseases of the elm
spanworm. Journal of the New York Entomological Society 84: 169-177.
Drooz, A.T. 1980. A review of the biology of the elm spanworm
(Lepidoptera: Geometridae). Great Lakes Entomologist 13: 49-53.
Fry, H.R.C., D.T. Quiring, K.L. Ryall, and P.L. Dixon. 2008.
Relationships between elm spanworm, Ennomos subsignaria, juvenile
density and defoliation on mature sycamore maple in an urban
environment. Forest Ecology and Management 255: 2726-2732.
Fry, H.R.C., K.L. Ryall, P.L. Dixon, and D.T. Quiring. 2008. Suppression
of Ennomos subsignaria (Lepidoptera: Geometridae) on Acer
pseudoplatanus (Aceraceae) in an urban forest with bole-implanted
acephate. Journal of Economic Entomology 101: 822-828.
A Photographic Guide 533

149. Winter Moth, Operophtera brumata (L.) (Lepidoptera: Geometridae)

Orientation to Pest Winter moth, Operophtera brumata (L.), is a European geometrid that
is invasive in North America, in Nova Scotia, part of British Colombia,
and, most recently, southern New England. It is a defoliator that is
a generalist feeder, attacking a variety of hardwoods. Winter moths
overwinter as eggs, which hatch early in spring, and young larvae feed
on opening buds and developing leaves. Older larvae feed inside loose
leaf rolls and, when mature, drop to the ground, where they pupate late
May to early June. Adults emerge in late fall or early winter. Females,
which are wingless, climb trees to lay eggs. Persistent defoliation leads
to branch death, top kill, and tree death.

Hosts Commonly Attacked This species is a generalist feeder that attacks are variety of hardwoods,
but especially apple (Malus), red oak (Quercus rubra L.), American elm
(Ulmus americana L.), red maple (Acer rubrum L.), American basswood
(Tilia americana L.), and hop hornbeam (Ostrya virginiana [Mill.] K.
Koch).

Distribution This geometrid is native in Europe, but in North America invasive

populations exist in Nova Scotia, the Pacific Northwest (Oregon
to British Columbia) and the northeastern United States. Outbreak
populations currently exist in eastern Massachusetts and Rhode Island.
A survey with pheromone traps by Elkinton et al. (2010) recovered
winter moths mainly in coastal areas from Long Island to Maine.

Images of Figure 1. Adults of winter

moth, Operophtera
Winter Moth brumata; male (top)
and wingless female
(bottom). (Top: Louis-
Michel Nageleisen,
Département de la
Santé des Forêts,
Bugwood.org; bottom:
Gyorgy Csoka, Hungary
Forest Research
Institute, Bugwood.org)
534 Forest Pest Insects in North America:

Images of Winter Figure 2. Eggs of

winter moth. (Gyorgy
Moth (continued) Csoka, Hungary Forest
Research Institute,
Bugwood.org)

Figure 3. Larvae of
winter moth. (Both
photos: Milan Zubrik,
Forest Research
Institute - Slovakia,
Bugwood.org)

Figure 4. Pupa of winter

moth. (Hannes Lemme,
Bugwood.org)
A Photographic Guide 535

Images of Winter
Moth (continued)

Figure 5. Feeding of larvae of winter moth. (Hannes Lemme, Bugwood.org)

Important Biological Control This species has well known natural enemies, having been studied in part
Agents Related to this of its native range (the UK) and been the target of successful classical
Pest Species biological control programs in both eastern and western Canada. Similar
work is now underway in Massachusetts. The natural enemies of most
importance are the tachinid parasitoid, Cyzenis albicans Fallén, and the
ichneumonid Argypon flaveolatum (Gravenhorst). The tachinid is the
more important.

Web Links for Information http://www.umassgreeninfo.org/fact_sheets/defoliators/wm_overview_

on Winter Moth 09.pdf; University of Massachusetts fact sheet.
http://en.wikipedia.org/wiki/Winter_Moth; Wikipedia article.
http://www.youtube.com/watch?v=kVxzOl8zrUE; Youtube video
on Cyzinis albicans releases in Massachusetts by Joe Elkinton of the
University of Massachusetts.

Articles Ring, R.A. 1988. Pest management of the European winter moth,
Operophtera brumata (L.), in British Columbia. Northwest
Environmental Journal 41: 329-330.
Embree, D.G. 1991.The winter moth Operophtera brumata in eastern
Canada, 1962-1988. Forest Ecology and Management 39: 47-54.
Elkinton, J.S., G.H. Boettner, M. Sremac, R. Gwiazowski, R. Hunkins,
J. Callahan, S.B. Schuefele, C. Donahue, A.H. Porter, A. Khrimian, B.M.
Whited, and N.K. Campbell. 2010. Survey for winter moth (Lepidoptera:
Geometridae) in northeastern North America with pheromone-baited
traps and hybridization with the native Bruce spanworm. Annals of the
Entomological Society of America 103: 135-145.
536 Forest Pest Insects in North America:

150. Bruce Spanworm, Operophtera bruceata (Hulst) (Lepidoptera:

Geometridae)

Orientation to Pest Bruce spanworm, Operophtera bruceata (Hulst), is a native geometrid

moth in North America that closely resembles winter moth (Operophtera
brumata [L.]). It is found from coast to coast in Canada and from New
England to the Lake States in the United States. Bruce spanworm is
a defoliator and a generalist feeder, attacking a variety of hardwoods.
Bruce spanworm overwinters as eggs, which hatch early in spring.
Larvae either feed openly or in the shelter of leaves that are loosely
rolled and webbed together. When mature, larvae drop to the ground,
where they pupate. Adults emerge in fall. Females, which are wingless,
climb trees to lay eggs in bark crevices or under loose bark. Defoliating
outbreaks have been reported.

Hosts Commonly Attacked This species is a generalist feeder that attacks are variety of hardwoods,
but especially sugar maple (Acer saccharum Marshall), trembling aspen
(Populus tremuloides Michx.), willow (Salix), and American beech
(Fagus grandifolia Ehrh.), among others.

Distribution This geometrid is found from coast to coast in Canada and from New
England to the Lake states in the United States.

Images of
Bruce Spanworm

Figure 1. Adults of bruce spanworm, Operophtera bruceata; male (left) and

wingless female (right). (Left: Ronald S. Kelley, Vermont Department of Forests,
Parks and Recreation, Bugwood.org; right: Pennsylvania Department of
Conservation and Natural Resources - Forestry Archive, Bugwood.org)
A Photographic Guide 537

Images of Bruce Figure 2. Egg of bruce

spanworm. (Ronald
Spanworm (continued) S. Kelley, Vermont
Department of Forests,
Parks and Recreation,
Bugwood.org)

Figure 3. Larvae of
bruce spanworm.
(E. Bradford Walker,
Vermont Department
of Forests, Parks
and Recreation,
Bugwood.org)

Important Biological Control The natural enemies of this species are less well known than are those of
Agents Related to this the closely related winter moth.
Pest Species
Web Links for Information http://www.maine.gov/doc/mfs/brucspan.htm; fact sheet of the state
on Bruce Spanworm of Maine.

Articles Elkinton, J.S., G.H. Boettner, M. Sremac, R. Gwiazdowski,

R.R. Hunkins, J. Callahan, S.B. Scheufele, C.P. Donahue, A.H. Porter,
A. Khrimian, B.M. Whited, and N.K. Campbell. 2010. Survey for
winter moth (Lepidoptera: Geometridae) in northeastern North America
with pheromone-baited traps and hybridization with the native Bruce
spanworm (Lepidoptera: Geometridae). Annals of the Entomological
Society of America 103: 135-145.
538 Forest Pest Insects in North America:

151. Forest Tent Caterpillar, Malacosoma disstria Hübner (Lepidoptera:

Lasiocampidae)

Orientation to Pest Forest tent caterpillar, Malacosoma disstria Hübner, occurs throughout
Canada and the United States and is a generalist defoliator that feeds on
a variety of hardwood trees. In the north and west of the United States
(and southern Canada), trembling aspen (Populus tremuloides Michx.)
is preferred. In the southern United States, various gums (Nyssa spp.
and Liquidambar styraciflua L.) and oaks (Quercus) are the common
hosts. However, during outbreaks, larvae will feed on a wide variety
of hardwood species. Winter is passed as eggs, which hatch in spring.
Larvae begin feeding on buds and expanding leaves. Older larvae feed
on mature leaves. Young larvae feed in groups, but older larvae are
dispersed. Larvae lay silk trails, which they follow, but they do not form
a silken tent. Rather, they make silk mats on which larvae rest. Mature
larvae wander and pupate inside a yellowish-white cocoon fixed to
the host tree or other objects. Adults emerge in midsummer (north) or
spring (south) and lay their eggs in clusters that encircle twigs of the
host tree. Eggs are cemented together and covered with dark colored
material called spumaline. There is one generation per year. A number
of very large outbreaks, covering thousands of square miles and lasting
for several years, have occurred in North America. Tree mortality is
generally low, but loss of growth can be considerable (>70 percent) due
to effects of defoliation. In the southern United States, in bottomlands,
gum trees may die from defoliation. In the northeastern United States,
defoliation of sugar maple stands lowers maple syrup yields and quality.

Hosts Commonly Attacked This species is a generalist defoliator that feeds on a variety of hardwood
trees. In the north and west, trembling aspen (P. tremuloides) is preferred.
In the southern United States, gums (Nyssa spp. and L. styraciflua) and
oaks (Quercus) are common hosts.

Distribution Forest tent caterpillar occurs throughout Canada and the United States.

Images of Forest Figure 1. Adult of

forest tent caterpillar,
Tent Caterpillar Malacosoma disstria.
(Whitney Cranshaw,
Colorado State
University,
Bugwood.org)
A Photographic Guide 539

Images of Forest Figure 2. Eggs of forest

tent caterpillar (top)
Tent Caterpillar and newly hatched
(continued) caterpillars on egg mass
(bottom). (Top: Steven
Katovich, USDA Forest
Service, Bugwood.org;
bottom: E. Bradford
Walker, Vermont
Department of Forests,
Parks and Recreation,
Bugwood.org)

Figure 3. Larvae of forest tent caterpillar feeding. (Gerald J. Lenhard, Louiana

State University, Bugwood.org)
540 Forest Pest Insects in North America:

Images of Forest Figure 4. Larvae of

forest tent caterpillar
Tent Caterpillar resting together on
(continued) trunk. (Andrew J. Boone,
South Carolina
Forestry Commission,
Bugwood.org

Figure 5. Forest tent

caterpillar cocoon (with
pupa inside). (James B.
Hanson, USDA Forest
Service, Bugwood.org)
A Photographic Guide 541

Images of Forest
Tent Caterpillar
(continued)

Figure 6. Interior view of stand defoliated by forest tent caterpillar. (Herbert A.

‘Joe’ Pase III, Texas Forest Service, Bugwood.org)

Figure 7. Strips of water tupelo (Nyssa aquatica L.) defoliated by forest tent
caterpillar (green areas are strips of less favored host species). (William M. Ciesla,
Forest Health Management International, Bugwood.org)

Important Biological Control The natural enemies of this species are well known and include
Agents Related to this 14 species of Hymenoptera egg parasites and, attacking larvae or pupae,
Pest Species 52 Diptera, and 61 Hymenoptera species. In addition there are some
18 Hemiptera, nine Coleoptera, and one Dermaptera that are predators of
various life stages of forest tent caterpillar. In northern areas, the tachinid
Sarcophaga aldrichi Park is important in terminating outbreaks. In the
northeastern United States, the tachinids Zenilla protuberans A and W
and Compsilura concinnata Meigan are also important.
542 Forest Pest Insects in North America:

Web Links for Information http://facultyweb.cortland.edu/fitzgerald/ForestTentCaterpillar.html;

on Forest Tent Caterpillar website of T.D. Fitzgerald. “Social Caterpillars: Forest Tent Caterpillar
(Malacosoma disstria).”
http://na.fs.fed.us/spfo/pubs/fidls/ftc/tentcat.htm; Forest and Disease Pest
Leaflet No. 9 of the USDA Forest Service.
http://entnemdept.ufl.edu/creatures/trees/forest_tent_caterpillar.htm;
fact sheet of the University of Florida on biology and control.
http://www.youtube.com/watch?v=x81ba8-e_O8--Youtube; video of
forest tent caterpillars moving about.

Articles Witter, J.A. and H.M. Kuhlman. 1972. A review of the parasites and
predators of tent caterpillars (Malacosoma spp.) in North America.
Minnesota Agricultural Experiment Station. Technical Bulletin 289: 48 p.
Parry, D., J.R. Spence, and W.J.A. Volney. 1997. Responses of natural
enemies to experimentally increased populations of forest tent caterpillar,
Malacosoma disstria. Ecological Entomology 22: 97-108.
Wood, D.M., D. Parry, R.D. Yanai, and N.E. Pitel. 2010. Forest
fragmentation and duration of forest tent caterpillar (Malacosoma
disstria Hübner) outbreaks in northern hardwood forests. Forest Ecology
and Management 260: 1193-1197.
Wood, D., R. Yanai, D. Allen, and S. Wilmot. 2009. Sugar maple decline
after defoliation by forest tent caterpillar. Journal of Forestry 107: 29-37.
A Photographic Guide 543

152. Eastern Tent Caterpillar, Malacosoma americanum (F.) (Lepidoptera:

Lasiocampidae)

Orientation to Pest Eastern tent caterpillar, Malacosoma americanum (F.), occurs throughout
southern Canada and the eastern United States and is a defoliator that
feeds on a variety of hardwood trees, especially cherry (Prunus) and
apple (Malus). Winter is passed as eggs, which hatch in spring. Larvae
begin feeding on buds and expanding leaves. Older larvae feed on mature
leaves. Larvae lay silk trails that they follow to foliage, where they feed
in groups. When not feeding, larvae shelter in silk tents constructed
low in trees in the forks of branches. Mature larvae wander and pupate
inside a yellowish-white cocoon fixed to the host tree or other objects.
Adults emerge in midsummer (north) or spring (south) and lay their
eggs in clusters that encircle twigs of the host trees. Eggs are cemented
together and covered with dark colored material called spumaline. There
is one generation per year. This species is primarily a nuisance species,
causing little economic loss, with the exception of black cherry (Prunus
serotina Ehrh.) stands managed to produce veneer wood. The tents of this
species are commonly seen along roadsides on young cherry or apple.
Caterpillars that have fed on cherry, if then eaten by horses, can cause
mares to abort, making this pest of special concern in areas dedicated to
rearing race horses.

Hosts Commonly Attacked This species is a generalist defoliator on a variety of hardwood trees,
especially cherry (Prunus) and apple (Malus).

Distribution This species occurs throughout southern Canada and the eastern United
States.

Images of Eastern Figure 1. Adult of

eastern tent caterpillar,
Tent Caterpillar Malacosoma
americanum.
(Lacy L. Hyche,
Auburn University,
Bugwood.org)
544 Forest Pest Insects in North America:

Images of Eastern Tent

Caterpillar (continued)

Figure 2. Egg masses of eastern tent caterpillar. (A. Steven Munson, USDA
Forest Service, Bugwood.org)

Figure 3. Young larvae of forest tent caterpillar on a small web. (A. Steven
Munson, USDA Forest Service, Bugwood.org)
A Photographic Guide 545

Images of Eastern Tent

Caterpillar (continued)

Figure 4. Large larvae of eastern tent caterpillar. (Joseph O’Brien, USDA Forest
Service, Bugwood.org)

Figure 5. Close up of eastern tent caterpillar larva. (Clemson University - USDA

Cooperative Extension Slide Series, Bugwood.org)
546 Forest Pest Insects in North America:

Images of Eastern Tent Figure 6. Comparison

of caterpillars of eastern
Caterpillar (continued) tent caterpillar (right)
and forest tent caterpillar
(left). (Ronald S. Kelley,
Vermont Department
of Forests, Parks
and Recreation,
Bugwood.org)

Figure 7. Close up of
eastern tent caterpillar
tent with mature larvae.
(William H. Hoffard,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 547

Images of Eastern Tent

Caterpillar (continued)

Figure 8. Eastern tent caterpillar cocoon (with pupa inside). (Lacy L. Hyche,
Auburn University, Bugwood.org)

Figure 9. Webs of eastern tent caterpillar on defoliated trees. (Left: Robert L.

Anderson, USDA Forest Service, Bugwood.org; right: G. Keith Douce, University
of Georgia, Bugwood.org)
548 Forest Pest Insects in North America:

Important Biological Control A variety of natural enemies of this species have been recognized. See
Agents Related to this Van Driesche et al. (2002) for details. However, their importance in
Pest Species determining the density of this species has not been investigated.

Web Links for Information http://www.ca.uky.edu/entomology/entfacts/ef423.asp; fact sheet of the

on Eastern Tent Caterpillar University of Kentucky on biology and control.
http://ohioline.osu.edu/hyg-fact/2000/2022.html; fact sheet of Ohio State
University.
http://www.ag.auburn.edu/enpl/bulletins/easterntentcaterpillar/
easterntentcaterpillar.htm; fact sheet of Auburn University, Alabama.
http://www.youtube.com/watch?v=1qO6eCygcTE; a time lapse video of
web construction and feeding on Youtube.
http://www.dec.ny.gov/animals/7111.html; New York Department of
Conservation site comparing gypsy moths and eastern tent caterpillar.

Articles Progar, R.A., M.J. Rinella, D. Fekedulegn, and L. Butler. 2010. Nuclear
polyhedrosis virus as a biological control agent for Malacosoma
americanum (Lepidoptera: Lasiocampidae). Journal of Applied
Entomology 134: 641-646.
Haynes, K.F., J. McLaughlin, S. Stamper, C. Rucker, F.X. Webster,
D. Czokajlo, and P. Kirsch. 2007. Pheromone trap for the eastern tent
caterpillar moth. Environmental Entomology 36: 1199-1205.
Crump, D., R.M. Silverstein, H.J. Williams, and T.D. Fitzgerald. 1987.
Identification of trail pheromone of larva of eastern tent caterpillar
Malacosoma americanum (Lepidoptera: Lasiocampidae). Journal of
Chemical Ecology 13: 397-402.
VanDriesche, R.G., S. Healy, and R.C. Reardon. 2002. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States. A review and recommendations. FHTET-
96-19. USDA Forest Service, Morgantown, West Virginia. (Available at
http://www.fs.fed.us/foresthealth/technology/pdfs/FHTET_96_19.pdf).
A Photographic Guide 549

153. Siberian Moth, Dendrolimus sibiricus TschetvericKov (Lepidoptera:

Lasiocampidae)

Orientation to Pest Siberian moth, Dendrolimus sibiricus TschetveriKov, is a defoliator

of great importance in northern Asia, causing widespread, extensive
mortality to various conifers during outbreaks. It does not occur in North
American but is a species of great concern. Typically, this moth has a
two-year life cycle, spread over parts of three calendar years. Adults
fly and lay eggs from late June to mid-July. Eggs are laid in chains or
clusters on needles or branches. In their first year, larvae feed until they
reach the second or third instar and then overwinter in the forest litter. In
year two, partly grown larvae continue to feed, at a slower rate, until they
are in the fifth to seventh instar, at which stage they overwinter. In their
third calendar year, larva feed until mature in spring and pupate inside
cocoons fixed to trees. Adult moths appear in late June and July and
may fly several kilometers before laying eggs. Outbreaks occur at about
10 year intervals and last 2-3 years. An outbreak can defoliate thousands
of hectares and affect entire forests. Since the insect has overlapping
generations and life cycles extending more than one growing season, it
can defoliate stands in the spring, summer and fall, but most defoliation
occurs in spring when older larvae feed.

Hosts Commonly Attacked Larvae of D. sibiricus feed on the foliage of more than 20 species of
conifers in northern Asia, especially on species of larch (Larix) and fir
(Abies), but also on less favored hosts such as five-needle and two-needle
pines (Pinus) and spruce (Picea). Injury is greatest to species of fir and
certain pines (e.g., P. koraiensis Sieb. et Zucc. and P. sibirica Du Tour).
Although larch species are the most favored hosts, they can re-foliate and
are seldom killed. It has been shown that this polyphagous species can
develop on a wide range of European and North American conifers.

Distribution The Siberian moth does not occur in North America, but is considered
a very high risk invader because of its potential for invasion and the
magnitude of likely damage. Boreal and subalpine forests in North
America would be at risk. Its current distribution includes Siberia
(Russia), northeastern China, northern Mongolia, and part of North
Korea.
550 Forest Pest Insects in North America:

Images of
Siberian Moth

Figure 1. Male Siberian moth, Dendrolimus sibericus. (Natalia Kirichenko,

Bugwood.org)

Figure 2. Egg mass of Siberian moth. (John H. Ghent, USDA Forest Service,
Bugwood.org)

Figure 3. Larva of
Siberian moth. (John
H. Ghent, USDA Forest
Service, Bugwood.org)
A Photographic Guide 551

Images of Siberian
Moth (continued)

Figure 4. Pupal cocoons of Siberian moth. (John H. Ghent, USDA Forest Service,
Bugwood.org)

Figure 5. Stand of Siberian larch (Larix sibirica Ledeb.) defoliated by Siberian

moth. (John H. Ghent, USDA Forest Service, Bugwood.org)

Important Biological Control More than 40 species of parasitoids are known to attack D. sibiricus, of
Agents Related to this which the most important are the egg parasites Telenomus tetratomus
Pest Species (Thomson) (Scelionidae) and Ooencyrtus pinicola Matsumura
(Encyrtidae), the larval parasite Rogas dendrolimi Matsumura
(Braconidae), and the larval-pupal parasites Blepharipa schineri
(Mesnil.), Blepharipa pratensis Meigen, and Lespesia frenchii
(Williston) (all Tachinidae).
552 Forest Pest Insects in North America:

Web Links for Information http://www.aphis.usda.gov/plant_health/plant_pest_info/pest_detection/

on Siberian Moth downloads/pra/dsuperanspra.pdf; a mini risk assessment of Siberian silk
moth (from a North American perspective).
http://www.eppo.org/QUARANTINE/insects/Dendrolimus_sibiricus/
DS_Dendrolimus_spp.pdf; an EPPO data sheet for European quarantine
officials.
http://www.issg.org/database/species/ecology.asp?si=1428&fr=1&sts=&l
ang=EN; write up on species in Global Invasive Species Database.
ftp://ftp.fao.org/docrep/fao/011/i0640e/i0640e10h.pdf; fact sheet by
United Nations FAO group.
http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=45&langdispl
ay=english; Exotic forest pest data sheet of the USDA Forest Service.

Articles Kirichenko, N.I., J. Flament, Y.N. Baranchikov, and J.-C. Grégoire. 2008.
Native and exotic coniferous species in Europe – possible host plants for
the potentially invasive Siberian moth, Dendrolimus sibiricus Tschtv.
(Lepidoptera, Lasiocampidae). OEPP/EPPO Bulletin 38: 259-263.
Kirichenko, N.I., Y.N. Baranchikov, and S. Vidal. 2009. Performance of
the potentially invasive Siberian moth Dendrolimus superans sibiricus
on coniferous species in Europe. Agricultural and Forest Entomology
11: 247-254.
A Photographic Guide 553

154. Pinkstriped Oakworm, Anisota virginiensis (Drury) (Lepidoptera:

Saturniidae)

Orientation to Pest Pinkstriped oakworm, Anisota virginiensis (Drury), is a native moth

that feeds on foliage of oaks (Quercus) and other hardwoods. It occurs
throughout the eastern United States and southern Canada. Young larvae
feed in groups, skeltonizing leaves. Older larvae are less gregarious and
consume all but the main veins of leaves. Larvae can be found April
through September depending on the climate, with one generation per
year in its northern range and up to three generations per year in its
southern-most range. This insect is common but not abundant and is
usually considered a minor pest to landscape trees where damage to
larger trees is often limited to defoliation of single branches. However,
small trees can be completely defoliated.

Hosts Commonly Attacked Pinkstriped oakworm feeds primarily on oaks (Quercus), but it is also
reported from chestnut (Castanea), hazel (Corylus), maple (Acer), and
birch (Betula).

Distribution This species occurs in southern Canada and the eastern United States, as
far west as Manitoba and Texas.

Images of
Pinkstriped Oakworm

Figure 1. Adults of pinkstriped oakworm, Anisota virginiensis: male and female

mating. (Pennsylvania Department of Conservation and Natural Resources
- Forestry Archive, Bugwood.org)
554 Forest Pest Insects in North America:

Images of Pinkstriped
Oakworm (continued)

Figure 2. Eggs of pinkstriped oakworm. (James Solomon, USDA Forest Service,

Bugwood.org)

Figure 3. Young larvae of pinkstriped oakworm. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Figure 4. Close up of pinkstriped oakworm larva. (James Solomon, USDA Forest

Service, Bugwood.org)
A Photographic Guide 555

Images of Pinkstriped
Oakworm (continued)

Figure 5. Pupa of pinkstriped oakworm. (Lacy L. Hyche, Auburn University,

Bugwood.org)

Figure 6. A group of pinkstriped oakworm caterpillars defoliating an oak branch.

(James Solomon, USDA Forest Service, Bugwood.org)

Important Biological Control In Manitoba, five species of larval or pupal parasitoids have been
Agents Related to this recorded from this species: Houghia sternalis (Coquillett), Lespesia
Pest Species anisotae Webber, and Winthemia datanae (Tns.) (all Tachinidae); and
Habronyx magniceps (Cresson) and Hyposoter fugitivus (Say) (both
Ichneumonidae).
556 Forest Pest Insects in North America:

Web Links for Information http://www.ag.auburn.edu/enpl/bulletins/pinkoakworm/pinkoakworm.

on Pinkstriped Oakworm htm; fact sheet of Auburn University on recognition and biology.
http://okeechobee.ifas.ufl.edu/News%20columns/
Pinkstriped%20Oakworm.htm; University of Florida fact sheet on
biology and control.
http://www.fs.fed.us/r8/foresthealth/idotis/insects/oakworm.html; USDA
Forest Service fact sheet on two oakworm species.

Articles Wagner, D.L., V. Giles, R.C. Reardon, and M.L. McManus. 1998.
Caterpillars of Eastern Forests. USFS Technology Transfer Bulletin,
FHTET-96-34: 113 p.
Henne, D.C. 2004. Parasitoid survey of Anisota virginiensis
(Lepidoptera: Saturniidae) at Belair, Manitoba from 1989-1999.
Proceedings of the Entomological Society of Manitoba 60: 5-10.
A Photographic Guide 557

155. Orangestriped Oakworm, Anisota senatoria (J.E. Smith) (Lepidoptera:

Saturniidae)

Orientation to Pest Orangestriped oakworm, Anisota senatoria (J.E. Smith), is a native

silkmoth that feeds mainly on the foliage of oaks (Quercus). It has as a
wide distribution in eastern United States and southern Canada. Adults
appear in the June or July in the northern part of its range and lay groups
of eggs on the undersides of leaves of host trees. Young larvae feed in
groups, skeletonizing leaves. Older larvae consume all but the main veins
and are less gregarious. Mature larvae drop to the ground and wander
before pupating in the soil, where they overwinter. Outbreaks occur
infrequently. There is one generation per year in most parts of the range.
Defoliation can be significant in some years, but little harm results unless
trees are under additional stress from other factors.

Hosts Commonly Attacked This species feeds primarily on oaks (Quercus).

Distribution This species occurs in much of the eastern United States and southern
Canada, west to Minnesota and Texas, although it is not common in the
southeastern United States.

Images of Figure 1. Adults of

orangestriped oakworm,
Orangestriped Oakworm Anisota senatoria.
(John Wheatley, John B.
Wheatley, Bugwood.org)
558 Forest Pest Insects in North America:

Images of Orangestriped
Oakworm (continued)

Figure 2. Eggs of orangestriped oakworm. (John Wheatley, John B. Wheatley)

Figure 3. Young larvae of orangestriped oakworm. (Lance S. Risley, William

Paterson University, Bugwood.org)

Figure 4. Close up of mature orangestriped oakworm caterpillar. (Lacy L. Hyche,

Auburn University, Bugwood.org)
A Photographic Guide 559

Images of Orangestriped
Oakworm (continued)

Figure 5. Defoliation by larvae of orangestriped oakworm. (Erich G. Vallery,

USDA Forest Service - SRS-4552, Bugwood.org

Important Biological Control Parasitoids that have been reared in Virginia from orangestriped
Agents Related to this oakworm larvae or pupae include Hyposoter fugitivus (Say)
Pest Species (Ichneumonidae) (in larvae, 3.2 to 9.3 percent) and Lespesia anisotae
(Webber) (Tachinidae) and Belvosia bifasciata (Fabricius) (Tachinidae)
(in larvae at 10.7 and 0.7 percent, respectively).

Web Links for Information http://www.ct.gov/caes/lib/caes/documents/publications/fact_sheets/

on Orangestriped Oakworm entomology/orangestriped_oakworm.pdf; website of the Connecticut
Agricultural Experiment Station.
http://www.ag.auburn.edu/enpl/bulletins/orangeoakworm/
orangeoakworm.htm; fact sheet of Auburn University, Auburn, Alabama.
http://www.oakgov.com/msu/assets/docs/publications/e2654_oakworm.
pdf; MSU Extension Bulletin-2654, fact sheet of Michigan State
University.

Articles Coffelt, M.A. and P.B. Schultz. 1993. Population biology of

orangestriped oakworm (Lepidoptera: Saturniidae) in southeastern
Virginia. Journal of Entomological Science 28: 218-229.
Coffelt, M.A. and P.B. Schultz. 1993. Larval parasitism of orangestriped
oakworm (Lepidoptera: Saturniidae) in the urban shade tree environment.
Biological Control 3: 127-134.
Coffelt, M.A. and P.B. Schultz. 1989. Development of an aesthetic
injury level to decrease pesticide use against orange-striped oakworm
(Lepidoptera: Saturniidae) in an urban pest management project. Journal
of Economic Entomology 83: 2044-2049.
560 Forest Pest Insects in North America:

156. Pandora Moth, Coloradia pandora Blake (Lepidoptera: Saturniidae)

Orientation to Pest Pandora moth, Coloradia pandora Blake, is an important defoliator

of pines in the western United States. There are three recognized
subspecies. Two years are needed to complete the life cycle. Adults
appear in spring or early summer and eggs hatch in August. Larvae
crawl up trees and young larvae feed in groups on new foliage. Immature
larvae spend the winter hibernating in clusters at the base of the needles
and resume feeding the following spring. In June of the second year,
mature larvae crawl down trees and pupate in earthen cells in the ground.
Adults may emerge the following year, or remain as pupae for up to five
years. Outbreaks occur in regions with soils loose enough for larvae to
bury themselves for pupation, these being chiefly pumice or decomposed
granite soils. Large and economically important outbreaks have occurred
in the past and these seem to reoccur at 20-30 year intervals, lasting 6 to
8 years each. Because each generation requires two years for completion
and populations are synchronized, feeding is concentrated in alternate
years, which allows many trees to recover and survive. In non-outbreak
years, this insect is uncommon.

Hosts Commonly Attacked This species feeds primarily on ponderosa (Pinus ponderosa Douglas
ex C. Lawson), lodgepole (P. contorta Douglas) and Jeffrey (P. jeffreyi
Balf.) pines, but other pines are also sometimes attacked.

Distribution This species occurs in the United States west of the Rocky Mountains,
except for Idaho and Washington.

Images of Figure 1. Adults

of Pandora moth,
Pandora Moth Coloradia pandora
(top, female; bottom,
male). (Top: USDA
Forest Service Archive,
USDA Forest Service,
Bugwood.org; bottom:
Terry Spivey, USDA
Forest Service,
Bugwood.org)
A Photographic Guide 561

Images of Pandora Moth Figure 2. Eggs of

Pandora moth.
(continued) (Darrell Ross, Oregon
State University,
Bugwood.org)

Figure 3. Young larva

of Pandora moth (top
photo), and older larvae
(middle and bottom
photos; note: upper
larvae in bottom photo
is infected with a virus).
(Top: Darrell Ross,
Oregon State University,
Bugwood.org; middle:
USDA Forest Service
- Region 2 - Rocky
Mountain Region
Archive, USDA Forest
Service, Bugwood.org;
bottom: Donald Owen,
California Department
of Forestry and
Fire Protection,
Bugwood.org)
562 Forest Pest Insects in North America:

Images of Pandora Moth Figure 4. Pupae of

Pandora moth.
(continued) (Darrell Ross, Oregon
State University,
Bugwood.org)

Figure 5. Ponderosa
pines defoliated by
Pandora moth. (Bruce
Hostetler, USDA Forest
Service, Bugwood.org)

Important Biological Control During epidemics, populations of Pandora moth are affected by
Agents Related to this many natural mortality factors, including a wilting virus (likely a
Pest Species polynucleohedrosis virus) of the mature larvae, small mammal predators
of the pupae in the soil, and various parasitoids.

Web Links for Information http://www.forestpests.org/acrobat/pandora.pdf; full text of article:

on Pandora Moth Carolin, V.M., Jr., and J.A.E. Knopf. 1968. The Pandora Moth. U.S.
Department of Agriculture, Forest Service, Forest Pest Leaflet 114.
http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/fsbdev2_043666.
pdf; USDA Forest Service leaflet on Pandora moth.

Articles Schmid, J.M. and D.D. Bennett. 1986. The North Kaibab Pandora moth
outbreak 1978-1984. USDA Forest Service, Rocky Mountain Forest
and Range Experiment Station, Fort Collins, Colorado, USA, General
Technical Report RM-153: 18 p.
Speer, J.H., T.W. Swetnam, B.E. Wickman, and A. Youngblood. 2001.
Changes in Pandora moth outbreak dynamics during the past 622 years.
Ecology 82: 679-697.
Speer, J.H. and R.L. Holmes. 2004. Effects of Pandora moth outbreaks
on ponderosa pine wood volume. Tree-Ring Research 60: 69-76.
A Photographic Guide 563

157. European Gypsy Moth, Lymantria dispar (L.) (Lepidoptera:

Lymantriidae)

Orientation to Pest European gypsy moth, Lymantria dispar (L.), is an invasive species in
the United States that has been a major forest pest for over a century. Its
damage consists of defoliation of deciduous forests and trees in spring
and early summer. Efforts to control this species with introduced natural
enemies began around 1900 and currently it is under good control in
New England, but outbreaks still occur to the west and south, where the
pest continues to invade. There is one generation a year and the species
overwinters as eggs on trees or various objects. Spread occurs locally
by wind-blown young caterpillars and over long distances by human
movement of egg masses. Female moths cannot fly. Both adults and
caterpillars are easily recognized.

Hosts Commonly Attacked The larvae of this species feed on most species of deciduous trees,
especially oaks (Quercus).

Distribution Figure 1. World

distribution of European
gypsy moth, Lymantria
dispar. (USDA Forest
Service - Northeastern
Research Station;
“Gypsy Moth in North
America”)

Images of European Figure 2. Female

European gypsy moth
Gypsy Moth (on egg mass). Note
white wings with brown
zigzag lines. (Steven
Katovich, USDA Forest
Service, Bugwood.org)
564 Forest Pest Insects in North America:

Images of European Figure 3. Male European

gypsy moth. Note brown
Gypsy Moth (continued) wings and feather-like
antennae. (James
A. Copony, Virginia
Department of Forestry,
Bugwood.org)

Figure 4. Newly hatched caterpillars of European gypsy moth on egg mass,

preparing to disperse by crawling or blowing in the wind on silk threads.
(Milan Zubrik, Forest Research Institute - Slovakia, Bugwood.org)

Figure 5. Mature European gypsy moth caterpillar. Note series of blue or red
spots on back, plus clusters of hairs on sides. (USDA Forest Service Archive,
USDA Forest Service, Bugwood.org)
A Photographic Guide 565

Images of European Figure 6. Pupae of

European gypsy moth
Gypsy Moth (continued) in silken cradle (far
left). Threads cut
away to show detail.
Female pupa above,
male below. (Milan
Zubrik, Forest Research
Institute - Slovakia,
Bugwood.org)

Figure 7. Defoliation from European gypsy moth. (Landesforstpräsidium Sachsen

Archive, Bugwood.org)

Figure 8. Aerial view of defoliation from European gypsy moth. In the absence of
effective biological control, widespread defoliation of forest stands can occur in
late spring. Defoliation areas appear brown on the upper slope. (Pennsylvania
Department of Conservation and Natural Resources - Forestry Archive,
Bugwood.org)
566 Forest Pest Insects in North America:

Images of European Figure 9. The

nucleopolyhedrosis virus
Gypsy Moth (continued) of European gypsy moth
can cause epidemics
in dense European
gypsy moth populations
but comes too late to
prevent damage. Virus-
killed caterpillars hang
by their legs on tree,
and fluids from rotting
cadavers drip onto
foliage, spreading virus.
(William M. Ciesla,
Forest Health
Management
International,
Bugwood.org)

Figure 10. The fungus Entomophthora maimaiga has emerged as a key

biological control agent of European gypsy moth in the northeastern United
States (left, diseased caterpillar; right, fungal protoplasts). (Both photos:
Andy Liebhold - USDA Forest Service - Northeastern Research Station;
“Gypsy Moth in North America”)
A Photographic Guide 567

Images of European Figure 11. Compsilura

concinnata, an important
Gypsy Moth (continued) tachinid parasitoid of the
European gypsy moth.
(Joyce Gross, UCB,
Bugwood.org)

Figure 12. Cotesia

melanoscela, a
parasitoid of European
gypsy moth larvae.
(James Lindsey,
Ecology of Commanster)

Important Biological Control Of some 80 species of parasitoids imported and released into the United
Agents Related to this States against this pest, 11 species became established and six are
Pest Species believed to have contributed to control of the European gypsy moth:
(1) Ooencyrtus kuvanae (Howard) (Hymenoptera: Encrytidae);
(2) Cotesia melanoscela (Ratzeburg) (Hymenoptera: Braconidae);
(3) Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae);
(4) Compsilura concinnata (Meigen) (Diptera: Tachinidae);
(5) Parasetigena silvestris (Robineau-Desvoidy) (Diptera: Tachinidae);
and (6) Blepharipa pratensis (Meigen) (Diptera: Tachinidae). Effective
control was achieved in the northeastern United States following the
accidental introduction of the fungal pathogen Entomophthora maimaiga
Humber, Shimazu, and Soper.

Web Links for Information http://www.fs.fed.us/ne/morgantown/4557/gmoth/; a USDA Forest

on European Gypsy Moth Service site that provide information and links to other sites on a variety
of topics, including distribution, effects on forests, management, and
natural enemies.
http://www.na.fs.fed.us/spfo/pubs/fidls/gypsymoth/gypsy.htm; a
USDA Forest Service leaflet (Forest insect and disease leaflet No. 162)
covering life cycle, hosts, effects of defoliation, population dynamics,
management, use of pesticides, and silvicultural treatments.
568 Forest Pest Insects in North America:

Articles Hajek, A.E., J.S. Elkinton, and J.J. Witcosky. 1996. Introduction and
spread of the fungal pathogen Entomophaga maimaiga (Zygomycetes:
Entomophthorales) along the leading edge of gypsy moth (Lepidoptera:
Lymantriidae) spread. Environmental Entomology 25: 1235-1247.
Liebhold, A., J. Elkinton, D. Williams, and R.M. Muzika. 2000. What
causes outbreaks of the gypsy moth in North America? Population
Ecology 42: 257-266.
Elkinton, J.S., W.M. Healy, A.M. Liebhold, and J.P. Buonaccorsi. 2002.
Gypsy moths and forest dynamics. In: McShea, W.J. and W.M. Healy.
Oak Forest Ecosystems: Ecology and Management for Wildlife. John
Hopkins University Press. Baltimore, MD, USA: 100-112.
A Photographic Guide 569

158. Asian Pink Moth, Lymantria mathura Moore (Lepidoptera:

Lymantriidae)

Orientation to Pest Asian pink moth (formerly called “rosy gypsy moth”), Lymantria
mathura Moore, is an Asian defoliator not yet present in the United
States, but one that poses a high risk of invasion and potential damage.
It defoliates a variety of hardwood species and outbreaks may affect
large areas. During outbreaks, the pest population density may reach
1000 caterpillars per tree. The species overwinters as eggs containing
fully developed larvae, ready to hatch. Larvae emerge early in spring
and disperse and attack buds, then leaves. Most feeding occurs at night.
Mature larvae pupate in flimsy cocoons on the host tree. Females fly at
night (1:00-3:00 a.m.) and lay eggs in masses of 150-600 on bark of host
tree or other objects. There is one generation per year.

Hosts Commonly Attacked Asian pink moth attacks many species of Betula, Castanea, Juglans,
Malus, Quercus, Salix, Tilia, Ulmus, and other deciduous trees. Its
preferred hosts in the Russian Far East include Juglans mandshurica
Maxim., Quercus mongolica Fisch. ex Turcz., and Q. dentata Thunb.

Distribution Asian pink moth is native to Asia and is found in the Russian Far
East, Nepal, Japan, Korea, northern India, and parts of China (Hebei,
Heilongjiang, Jilin provinces, and western China).

Images of Figure 1. Adults of Asian

pink moth, Lymantria
Asian Pink Moth mathura (top, male;
bottom, female).
(Top: DAFF Archive,
Bugwood.org; bottom:
David Mohn, Critters
Page [Creatures
Great and Small],
Bugwood.org)
570 Forest Pest Insects in North America:

Images of Asian Figure 2. Comparison of

the female of Asian pink
Pink Moth (continued) moth (bottom) to that of
gypsy moth (Lymantria
dispar [L.]), top. (USDA
APHIS PPQ Archive,
USDA APHIS PPQ,
Bugwood.org)

Figure 3. Egg mass of Asian pink moth. (David Mohn, Critters Page [Creatures
Great and Small], Bugwood.org)
A Photographic Guide 571

Images of Asian Figure 4. Young larvae

of Asian pink moth.
Pink Moth (continued) (David Mohn,
Critters Page [Creatures
Great and Small],
Bugwood.org)

Figure 5. Mature larvae

of Asian pink moth.
(USDA APHIS PPQ
Archive, USDA APHIS
PPQ, Bugwood.org)

Figure 6. Pupa of Asian

pink moth. (David Mohn,
Critters Page [Creatures
Great and Small],
Bugwood.org)
572 Forest Pest Insects in North America:

Important Biological Control In Korea, natural enemies associated with Asian pink moth include
Agents Related to this Cotesia melanoscela (Ratzberg), the dominant larval parasitoid, and
Pest Species Brachymeria lasus (Walker), the most important pupal parasitoid.

Web Links for Information http://www.eppo.org/QUARANTINE/insects/Lymantria_mathura/DS_

on Asian Pink Moth Lymantria_mathura.pdf; detailed fact sheet of the European quarantine
organization EPPO.
http://www.aphis.usda.gov/plant_health/plant_pest_info/pest_detection/
downloads/pra/lmathurapra.pdf; APHIS (USDA) risk assessment for
Asian pink moth to the United States.

Articles Zlotina, M.A., V.C. Mastro, D.E. Leonard, and J.S. Elkinton. 1998.
Survival and development of Lymantria mathura on North American,
Asian, and European tree species. Journal of Economic Entomology
91: 1162-1166.
Anon. 2005. Lymantria mathura. EPPO Bulletin 35(3): 464-467.
A Photographic Guide 573

159. Nun Moth, Lymantria monacha Moore (Lepidoptera: Lymantriidae)

Orientation to Pest Nun moth, Lymantria monacha Moore, is a Eurasian species that has not
yet invaded North America but is a species of high concern. Outbreaks
of the nun moth are often observed in Scots pine (Pinus sylvestris L.)
and Norway spruce stands (Picea abies [L.] Karst.), but many conifers
and broadleaf trees are also hosts. The biology closely resembles that
of the gypsy moth (Lymantria dispar [L.]), except that females of nun
moth can fly, which they do at night. The species overwinters as eggs
containing fully developed larvae, which emerge when buds on host trees
are opening. Young larvae feed on buds in groups but after the first instar,
larvae consume mature needles and feed alone. Pupation takes place on
the host tree and by mid-summer adults are in flight. Adults lay eggs in
cracks of bark on the host tree or in crevices of other objects in masses of
70 to 300 eggs. Egg masses may be moved on firewood or other material.
There is one generation per year.

Hosts Commonly Attacked Outbreaks of the nun moth are often observed in Scots pine (P. sylvestris)
and Norway spruce (P. abies) stands. However, most conifers and many
broad-leaved tree species may also be used as food plants.

Distribution Nun moth is widely distributed and is found in China, Japan, Russia, and
Europe.

Images of Figure 1. Adults of

nun moth, Lymantria
Nun Moth monacha; top, male and
bottom, female. (Note:
this species has a wide
range of color variation,
which is not captured
in these photos.) (Both
photos: DAFF Archive,
Bugwood.org)
574 Forest Pest Insects in North America:

Images of Nun Moth Figure 2. Adult female

nun moth in resting
(continued) position. (Hannes
Lemme, Bugwood.org)

Figure 3. Egg mass of

nun moth. (Stanislaw
Kinelski, Bugwood.org)
A Photographic Guide 575

Images of Nun Moth Figure 4. Young larvae

of nun moth. (Note: like
(continued) adults, larvae vary in
color.) (Both photos:
Stanislaw Kinelski,
Bugwood.org)

Figure 5. Mature larva

of nun moth. (Stanislaw
Kinelski, Bugwood.org)
576 Forest Pest Insects in North America:

Images of Nun Moth Figure 6. Pupae of nun

moth. (Hannes Lemme,
(continued) Bugwood.org)

Figure 7. Small conifer

killed by nun moth.
(Landesforstpräsidium
Sachsen Archive,
Bugwood.org)

Figure 8. Norway spruce

stand defoliated by
nun moth. (Jan Liska,
Forestry and Game
Management Research
Institute, Bugwood.org)
A Photographic Guide 577

Important Biological Control Many natural enemies have been recorded attacking nun moth, usually
Agents Related to this during outbreaks or in the decline phase as populations collapse. These
Pest Species include many species of parasitoids (mostly tachinid and sarcophagid
flies and braconid wasps). See Van Driesche et al. (1996) for details. A
baculovirus is also known from this species. However, there is limited
experimental evidence about the role such agents play in regulation of
the species or curtailment of outbreaks.

Web Links for Information http://wiki.bugwood.org/Archive:Atlas/Lymantria_monacha#Biology;

on Nun Moth short fact sheet of Bugwood Wiki.
ftp://ftp.fao.org/docrep/fao/011/i0640e/i0640e10q.pdf; fact sheet of the
United Nations FAO group.
http://www.inspection.gc.ca/english/plaveg/pestrava/lymmon/tech/
lymmone.shtml; fact sheet of the Canadian Food Inspection Agency.
http://www.nrs.fs.fed.us/disturbance/invasive_species/nun_moth/;
contains information on the species’ biology, photos, a reference list,
and a pest alert.

Articles Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations.
FHTET 96-19, USDA Forest Service, Morgantown. (Available
at http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
Keena, M.A., K.S. Shields, and M. Torsello. 1998. Nun moth: potential
new pest. Pest Alert NA-PR-95-98. USDA Forest Service, Northeastern
Area, State and Private Forestry: 2 p.
Keena, M.A. 2003. Survival and development of Lymantria monacha
(Lepidoptera: Lymantriidae) on North American and introduced Eurasian
tree species. Journal of Economic Entomology 96: 43-52.
Prestemon, J.P., J.A. Turner, J. Buongiorno, S.S. Zhu, and R.H. Li.
2008. Some timber product market and trade implications of an invasive
defoliator: the case of Asian Lymantria in the United States. Journal of
Forestry 106: 409-415.
578 Forest Pest Insects in North America:

160. Browntail Moth, Euproctis chrysorrhoea L. (Lepidoptera: Lymantriidae)

Orientation to Pest Browntail moth, Euproctis chrysorrhoea L., is an invasive species in

North America whose effects on trees is similar to that of gypsy moth
(Lymantria dispar [L.]), being a defoliator of a wide range of deciduous
broadleaf trees. Introduced into North America in the later 19th century,
by 1914 it had infested a wide band of coastal and inland New England.
In addition to widespread defoliation of many types of deciduous
hardwood trees, including many fruit trees, the species also was a threat
to public health because of the severe skin rashes from the caterpillar’s
hairs. These, if inhaled, could cause illness leading to death in some
cases. By 1922, its range began to quickly collapse and currently the
moth is found only in a few small coastal enclaves. While the cause of
this decline was not understood at the time, it was subsequently proved
to be caused by the action of Compsilura concinnata (Meigen), a
polyphagous tachinid introduced against it and the gypsy moth.

Hosts Commonly Attacked Browntail moth feeds on most species of deciduous broadleaf trees,
especially species of apple (Malus), pear (Pyrus), plum (Prunus), oak
(Quercus), willow (Salix), elm (Ulmus), and maple (Acer).

Distribution At the height of its invasion, browntail moth was found in much of
New England, but it is currently found only at the very tip of Cape Cod,
Massachusetts, and on some islands in Casco Bay, near Portland, Maine.

Figure 1. Historical
and current distribution
of browntail moth,
Euproctis chrysorrhoea,
in North America.
A Photographic Guide 579

Images of
Browntail Moth

Figure 2. Female browntail moth. (Jan Samanek, State Phytosanitary

Administration, Bugwood.org)

Figure 3. Egg mass of browntail moth. (Jan Samanek, State Phytosanitary

Administration, Bugwood.org)

Figure 4. Young
browntail moth
caterpillars on web.
(Jan Samanek,
State Phytosanitary
Administration,
Bugwood.org)
580 Forest Pest Insects in North America:

Images of Browntail
Moth (continued)

Figure 5. Mature browntail moth caterpillars. (Jan Samanek, State Phytosanitary

Administration, Bugwood.org)

Figure 6. Side view of browntail moth caterpillar showing prominent hairs that
cause severe skin rash. (Jan Samanek, State Phytosanitary Administration,
Bugwood.org)

Figure 7. Skin rash from

contact with browntail
moth caterpillars.
(Jan Samanek,
State Phytosanitary
Administration,
Bugwood.org)
A Photographic Guide 581

Images of Browntail
Moth (continued)

Figure 8. Pupae of browntail moth in web. (Jan Samanek, State Phytosanitary

Administration, Bugwood.org)

Figure 9. Defoliated trees and old browntail moth webbing. (Milan Zubrik, Forest
Research Institute - Slovakia, Bugwood.org)

Figure 10. Compsilura

concinnata, an important
tachinid parasitoid of the
European gypsy moth.
(Joyce Gross, UCB,
Bugwood.org)
582 Forest Pest Insects in North America:

Important Biological Control Elkinton et al. (2006) demonstrated that C. concinnata (Diptera:
Agents Related to this Tachinidae) was likely the agent responsible for the extinction of
Pest Species browntail moth from much of its former range in North America in the
early part of the 20th century. Two other tachinids, Townsendiellomyia
nidicola and Carcelia laxifrons, both browntail moth specialists were
also introduced and successfully established at the same time as C.
concinnata. More recently, a nucleopolyhedrosis virus from England,
specific to the moth has been released and established in Maine, but
impacts on remaining populations were not yet clear as of 2010. Many
browntail moth populations in Maine have been heavily affected by a
naturally occurring fungal pathogen, Entomophaga aulicae (Reichardt in
Bail) Humber.

Web Links for Information http://www.maine.gov/doc/mfs/btm.htm; leaflet from Maine Department

on Browntail Moth of Conservation that includes recommendations for chemical controls
and other measures.

Articles Elkinton, J.S., D. Parry, and G.H. Boettner. 2006. Implicating an

introduced generalist parasitoid in the invasive browntail moth’s
enigmatic demise. Ecology 87: 2664-2672.
Elkinton, J.S., E. Preisser, G. Boettner, and D. Parry. 2008. Factors
influencing larval survival of the invasive browntail moth (Lepidoptera:
Lymantriidae) in relict North American populations. Environmental
Entomology 37: 1429-1437. (Available at http://cels.uri.edu/preisserlab/
media/publications/preisser/Env%20Ent%202008%20(37,6)%201429-
1437.pdf).
A Photographic Guide 583

161. Pine Tussock Moth, Dasychira pinicola (Dyar) (Lepidoptera:

Lymantriidae)

Orientation to Pest Pine tussock moth, Dasychira pinicola (Dyar), is a native North
American tussock moth. It is known for sudden eruptions in population
that quickly subside and may not occur again for many years. It is found
in southeastern Canada and from the northeastern United States west
to Minnesota. The larvae feed on various conifers, especially on jack
pine (Pinus banksiana Lamb.) Outbreaks, when they do occur, can be
extensive and damaging since the caterpillar stage consumes both new
and old foliage, leaving host conifers completely defoliated. In the last
century there have been several large outbreaks on jack pine in the Lakes
States. In the Lake States, moths fly in mid-summer and eggs are laid
in irregular clusters on or near the female pupal case, on the needles
or the trunk. Eggs hatch later in the summer and young larvae feed on
needles until they reach the second or third instar, when they go into
hibernation for the winter. Larvae resume feeding in the spring, at first
eating staminate flowers and young needles. Later, old-needles are also
eaten. Fully grown larvae spin cocoons on needles or twigs, where they
also pupate. Moths emerge shortly thereafter. There is one generation per
year.

Hosts Commonly Attacked The caterpillar stage of pine tussock moth feeds on various conifers,
including jack (P. banksiana), red (P. resinosa Sol. ex Aiton), and
eastern white (P. strobus L.) pines, spruce (Picea) and balsam fir (Abies
balsamea [L.] Mill.).

Distribution Pine tussock moth is found in southeastern Canada and from the
northeastern United States west to Minnesota.

Images of Figure 1. Adult of pine

tussock moth, Dasychira
Pine Tussock Moth pinicola. (Gordon
Harrison)
584 Forest Pest Insects in North America:

Images of Pine
Tussock Moth (continued)

Figure 2. Eggs of the pine tussock moth. (USDA Forest Service - Northeastern
Area Archive, USDA Forest Service, Bugwood.org)

Figure 3. Larva of
pine tussock moth.
(USDA Forest Service
- Northeastern Area
Archive, USDA Forest
Service, Bugwood.org)

Figure 4. Pupa of pine tussock moth. (USDA Forest Service - Northeastern Area
Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 585

Images of Pine
Tussock Moth (continued)

Figure 5. Jack pines damaged by pine tussock moth. (Minnesota Department

of Natural Resources Archive, Minnesota Department of Natural Resources,
Bugwood.org)

Important Biological Control Natural enemies of this species have been studied during outbreaks that
Agents Related to this occurred in Wisconsin in the 1960s. Thirty-one parasitic species were
Pest Species reared during that study. Eight insect predators and four pathogens were
also reported. Parasitism levels were significant in the egg, larval and
pupal stages (see Sreenivasum et al. 1972 for details).

Web Links for Information None

on Pine Tussock Moth

Articles Sreenivasam, D.D., D.M. Benjamin, and D.D. Walgenbach. 1972.

The bionomics of the pine tussock moth. Research Bulletin No. 282.
School of Natural Resources, College of Agriculture and Life Sciences,
University of Wisconsin, Madison: 36 p.
586 Forest Pest Insects in North America:

162. Whitemarked Tussock Moth, Orgyia leucostigma (J. E. Smith)

(Lepidoptera: Lymantriidae)

Orientation to Pest Whitemarked tussock moth, Orgyia leucostigma (J. E. Smith), is a

native North American tussock moth whose larvae feed on a variety of
deciduous and coniferous trees. The species is found in most of eastern
Canada and the eastern United States. This tussock moth passes the
winter as eggs, which hatch in spring and thereafter young larvae often
spin down on silk threads and balloon to new host plants. Young larvae
skeletonize leaves of host trees, while older larvae consume whole leaves
(eating all but the major veins). Cocoons are formed in bark crevices
or between branches, and adults emerge in a few weeks. Females are
wingless and lay eggs in masses. There are one or two generations per
year depending on the region. Damage to forest trees is minor and this
species is better known as a pest of shade or fruit trees in urban areas and
in intensively managed plantations of Christmas trees.

Hosts Commonly Attacked Whitemarked tussock moth exploits a broad range of plant species and
has been reported to cause significant damage to many tree species,
including apple (Malus), basswood (Tilia), elm (Ulmus), poplar
(Populus), Norway maple (Acer platanoides L.), silver maple (Acer
saccharinum L.), sycamore (Platanus), paper birch (Betula papyrifera
Marsh.), yellow birch (Betula alleghaniensis Britt.), larch (Larix), and
balsam fir (Abies balsamea [L.] Mill.).

Distribution Whitemarked tussock moth is found throughout eastern Canada and the
eastern United States.

Images of Whitemarked
Tussock Moth

Figure 1. Adults of whitemarked tussock moth, Orgyia leucostigma: left, winged

male; right, wingless female. (Left: James Solomon, USDA Forest Service,
Bugwood.org; right: North Carolina Forest Service Archive, Bugwood.org)
A Photographic Guide 587

Images of Whitemarked
Tussock Moth (continued)

Figure 2. Flightless female of whitemarked tussock moth producing her egg

mass. (John L. Foltz, University of Florida, Bugwood.org)

Figure 3. Larvae of
whitemarked tussock
moth: top, dorsal view;
bottom, lateral view.
(Top: John H. Ghent,
USDA Forest Service,
Bugwood.org; bottom:
Connecticut Agricultural
Experiment Station
Archive, Connecticut
Agricultural Experiment
Station, Bugwood.org)
588 Forest Pest Insects in North America:

Images of Whitemarked Figure 4. Top, pupae of

whitemarked tussock
Tussock Moth (continued) moth (removed from
cocoons); bottom,
cocoons of whitemarked
tussock moth. (Top:
Robert L. Anderson,
USDA Forest Service,
Bugwood.org; bottom:
James Solomon,
USDA Forest Service,
Bugwood.org)

Figure 5. Feeding damage of whitemarked tussock moth larvae. (James

Solomon, USDA Forest Service, Bugwood.org)
A Photographic Guide 589

Important Biological Control Two strains of nucleopolyhedroviruses, one specific to whitemarked

Agents Related to this tussock moth and the other to Douglas fir tussock moth, Orgyia
Pest Species pseudotsugata (McDunnough), have been found to cause widespread
mortality in populations of whitemarked tussock moth and are being
developed as possible biological control agents in Christmas tree
plantations (balsam fir) in eastern Canada.

Web Links for Information on http://www.ag.auburn.edu/enpl/bulletins/whitemarkedtussock/

Whitemarked Tussock Moth whitemarkedtussock.htm; fact sheet of Auburn University.
http://www.ag.auburn.edu/enpl/bulletins/whitemarkedtussock/
whitemarkedtussock.htm; fact sheet of the USDA Forest Service,
Forest Health Protection, Southern Region.
http://insects.tamu.edu/fieldguide/cimg313.html; fact sheet of Texas
A&M University.

Articles Thurston, G.S. 2001. Orgyia leucostigma (J. E. Smith), whitemarked

tussock moth (Lepidoptera: Lymantriidae). In: Mason, P.G. and J.T.
Huber. Biological Control Programmes in Canada, 1981-2000. CABI
Publishing, Wallingford, UK: 201-203.
Medina, R.F. and P. Barbosa. 2002. Predation of small and large Orgyia
leucostigma (J. E. Smith) (Lepidoptera: Lymantriidae) larvae by
vertebrate and invertebrate predators. Environmental Entomology 31:
1097-1102.
590 Forest Pest Insects in North America:

163. Douglas-fir Tussock Moth, Orgyia pseudotsugata (McDunnough)

(Lepidoptera: Lymantriidae)

Orientation to Pest The Douglas-fir tussock moth, Orgyia pseudotsugata (McDunnough),

is a native defoliator of major importance in the interior Douglas-
fir and true fir forests of western North America, where is defoliates
Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and various true
firs (Abies). Adults emerge in late summer or early fall. The flightless
females remain on their cocoons, where they mate and lay their egg
mass. Eggs overwinter and hatch the following spring when new foliage
has appeared. Young larvae crawl to new needles and begin feeding,
but small larvae may disperse by dropping on silken threads and being
blown by the wind, especially when new foliage is not available. Young
larvae feed on the unfolding new needles, while older larvae feed on
both new and old foliage. Foliage of heavily infested forest stands turns
distinctly brownish. By late summer, mature larvae form cocoons either
on the foliage when densities are low or on tree trunks and objects on
the ground when densities are high. New adults emerge in a few weeks
to complete the life cycle. The cocoons and egg masses are among the
most conspicuous evidence of tussock moth abundance. There is one
generation a year. Periodic large scale outbreaks have occurred. These
outbreaks develop explosively but subside abruptly after about 3 years.
Between outbreaks this insect is seldom seen.

Hosts Commonly Attacked Douglas-fir tussock moth feeds on Douglas-fir (P. menziesii), and
various native firs, especially grand (Abies grandis [Douglas ex D. Don]
Lindley), white (A. concolor [Gordon] Lindley ex Hildebrand), and
subalpine fir (A. lasiocarpa [Hooker] Nuttal).

Distribution Douglas-fir tussock moth is found from southern British Columbia

through the eastern half of the Pacific coast states. It is also found in the
Rocky Mountain States, south to Arizona and New Mexico.

Images of Douglas-fir
Tussock Moth

Figure 1. Adults of Douglas-fir tussock moth, Orgyia pseudotsugata: left, male;

right, wingless female on her cocoon. (Both photos: Ladd Livingston, Idaho
Department of Lands, Bugwood.org)
A Photographic Guide 591

Images of Douglas-fir Figure 2. Egg masses

(top) of Douglas-fir
Tussock Moth (continued) tussock moth and egg
mass (bottom) with
newly hatched larvae.
(Top: Kenneth E.
Gibson, USDA Forest
Service, Bugwood.org;
bottom: USDA Forest
Service - Region 4
- Intermountain Archive,
USDA Forest Service,
Bugwood.org)

Figure 3. Mature larvae

of Douglas-fir tussock
moth. (Ladd Livingston,
Idaho Department of
Lands, Bugwood.org)

Figure 4. Cocoons of Douglas-fir tussock moth (left), and close up (right). (Left:
William M. Ciesla, Forest Health Management International, Bugwood.org; right:
USDA Forest Service - Ogden Archive, USDA Forest Service, Bugwood.org)
592 Forest Pest Insects in North America:

Images of Douglas-fir Figure 5. Feeding

damage of Douglas-
Tussock Moth (continued) fir tussock moth on
Douglas-fir trees.
(Dave Powell,
USDA Forest Service,
Bugwood.org)

Figure 6. Defoliation and tree mortality caused by Douglas-fir tussock moth.

(David McComb, USDA Forest Service, Bugwood.org)
A Photographic Guide 593

Important Biological Control The Douglas-fir tussock moth has many natural enemies including
Agents Related to this a nucleopolyhedrosis virus that is capable of dramatically reducing
Pest Species populations, but which usually appears after trees have been seriously
defoliated. Many parasitoids are known to attack this species, including
a Telonomus sp. egg parasitoid, and the larval parasitoids Phobocampe
pallipes (Provancher) (Ichneumonidae) and Carcelia yalensis Sellers
(Tachinidae). Parasitoids sometimes hasten the decline of an outbreak.

Web Links for Information http://insects.ippc.orst.edu/pdf/reb130.pdf.

on Douglas-fir Tussock Moth http://www.forestry.ubc.ca/fetch21/DFTM/dftmtot.html; fact sheet of the
University of British Columbia.
http://ext.nrs.wsu.edu/forestryext/foresthealth/notes/douglasmoth.htm;
fact sheet of Washington State University.
http://www.fs.fed.us/r6/nr/fid/fidls/fidl-86.pdf; USDA Forest Service
Forest Insect and Disease Leaflet.

Articles Mills, N.J. and F. Schoenberg. 1985. Possibilities for the biological
control of the Douglas-fir tussock moth, Orgyia pseudotsugata
(Lymantriidae), in Canada, using natural enemies from Europe.
Biocontrol News and Information 6(1): 7-18.
Dahlsten, D.L., D.L. Rowney, and W.A. Copper. 1992. Comparison of
artificial pupation shelters and other monitoring methods for endemic
populations of Douglas-fir tussock moth, Orgyia pseudotsugata
(McDunnough) (Lepidoptera: Lymantriidae). The Canadian
Entomologist 124: 359-369.
Otvos, I.S., R.F. Shepherd, and J.C. Cunningham. 2001. Orgyia
pseudotsugata (McDunnough), Douglas-fir tussock moth (Lepidoptera:
Lymantriidae). In: Mason, P.G. and J.T. Huber (eds.). Biological Control
Programmes in Canada, 1981-2000. CABI Publishing, Wallingford, UK:
204-212.
Otvos, I.S. and R.F. Shepherd. 1991. Integration of early virus treatment
with a pheromone detection system to control Douglas-fir tussock moth,
Orgyia pseudotsugata (Lepidoptera: Lymantriidae), populations at
pre-outbreak levels. Forest Ecology and Management 39: 143-151.
594 Forest Pest Insects in North America:

164. Poplar Tentmaker, Clostera inclusa (Hübner) (Lepidoptera:

Notodontidae)

Orientation to Pest Poplar tentmaker, Clostera inclusa (Hübner), is a native notodontid

moth that feeds on poplar (Populus) and willow (Salix) and is found
from southern Canada to Georgia, and west to Colorado. Adults fly from
March to July in the southern United States and from July to August
in the northern part of its U.S. range. Eggs are laid in clusters on the
undersides of leaves. Larvae are gregarious and live in tents, which they
construct by webbing together the edges of several leaves. The larvae
feed through the growing season and move to the ground in late fall,
where they pupate in loose cocoons and spend the winter. There are one
or two generations per year, depending on the location. Small groups of
open-grown trees may be defoliated.

Hosts Commonly Attacked This caterpillar feeds on poplar (Populus) and willow (Salix).

Distribution Poplar tentmaker is found from southern Canada to Georgia, west to

Colorado.

Images of Figure 1. Adult of poplar

tentmaker, Clostera
Poplar Tentmaker inclusa. (Marvin Smith,
Bugwood.org)
A Photographic Guide 595

Images of Poplar
Tentmaker (continued)

Figure 2. Egg mass of poplar tentmaker. (James Solomon, USDA Forest Service,
Bugwood.org)

Figure 3. Mature larvae of poplar tentmaker. (Robert L. Anderson, USDA Forest

Service, Bugwood.org)

Figure 4. Cocoon of poplar tentmaker. (William A. Carothers, USDA Forest

Service, Bugwood.org)
596 Forest Pest Insects in North America:

Images of Poplar
Tentmaker (continued)

Figure 5. Poplars defoliated by poplar tentmaker. (James Solomon, USDA Forest

Service, Bugwood.org)

Figure 6. Stand of eastern cottonwood (Populus deltoides ssp. deltoides Bartram

ex Marsh.) defoliated by poplar tentmaker. (James Solomon, USDA Forest
Service, Bugwood.org)

Important Biological Control Ladybird beetles may consume the eggs of this species.
Agents Related to this
Pest Species
Web Links for Information http://www.ag.auburn.edu/enpl//bulletins/poplartentmaker/
on Poplar Tentmaker poplartentmaker.htm; fact sheet of Auburn University on biology and
control.
http://www.entomology.umn.edu/cues/Web/190PoplarTentmaker.pdf;
fact sheet from the University of Minnesota.

Articles Solomon, J.D. and F.L. Oliveria. 1993. Evaluation of poplar tentmaker
defoliation on growth and survival of older cottonwood plantations.
Research Paper - Southern Forest Experiment Station, USDA Forest
Service (SO-271): 6 p.
A Photographic Guide 597

165. Saddled Prominent, Heterocampa guttivitta (Walker) (Lepidoptera:

Notodontidae)

Orientation to Pest Saddled prominent, Heterocampa guttivitta (Walker), is a native North

American notodontid that occurs in southeastern Canada and throughout
the eastern United States, feeding on beech (Fagus grandifolia Ehrh.),
paper birch (Betula papyrifera Marshall), sugar maple (Acer saccharum
Marshall) and many other hardwoods. Adults fly in spring and lay up to
500 eggs singly on the host’s leaves. Young larvae skeletonize leaves,
while older ones consume all but the larger veins. During outbreaks,
larvae may move from tree to tree. In mid-summer, larvae move into
the leaf litter, where they pupate and then pass the winter. There is one
generation per year in the northern part of the insect’s range. Periodic
outbreaks have occurred, especially in the northeastern United States,
and these may defoliate, and sometimes kill or top-kill affected trees.

Hosts Commonly Attacked This caterpillar feeds most commonly on American beech
(F. grandifolia), paper birch (B. papyrifera), and sugar maple
(A. saccharum).

Distribution The saddled prominent occurs in southeastern Canada and throughout the
eastern United States.

Images of Figure 1. Adult of

saddled prominent,
Saddled Prominent Heterocampa guttivitta.
(Mark Dreiling,
Bugwood.org)
598 Forest Pest Insects in North America:

Images of Saddled Figure 2. Eggs of

saddled prominent.
Prominent (continued) (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Figure 3. Young larvae

of saddle prominent
have structures near
head that resemble
horns. (Lacy L. Hyche,
Auburn University,
Bugwood.org)

Figure 4. Mature larva

of saddled prominent.
(Ronald S. Kelley,
Vermont Department
of Forests, Parks
and Recreation,
Bugwood.org)

Figure 5. Pupal cell

(left) and pupa (right,
removed from cell) of
saddled prominent.
(Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)
A Photographic Guide 599

Images of Saddled Figure 6. Appearance

of saddle prominent
Prominent (continued) feeding on maple.
(USDA Forest Service,
Forest Insect & Disease
Leaflet 167)

Important Biological Control The parasitoids recorded from various life stages of the saddled
Agents Related to this prominent include the scelionid egg parasitoid Telenomus coelodasidis
Pest Species Ashmead and the larval parasitoids Phobocampe pallida (Cushman)
(Ichneumonidae) and Eulophus anomocerus (J.C. Crawford)
(Eulophidae), both of which were uncommon (<1 percent parasitism).
Another ichneumonid, Cratichneumon sublatus (Cresson), is the main
pupal parasitoid, attacking 1-17 percent under outbreak conditions and
32-57 percent in the year following the collapse of its host’s population.

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/saddled/fidl-sp.htm; USDA

on Saddled Prominent Forest Service Pest and Disease Leaflet No. 167 on biology and control.
http://wiki.bugwood.org/Heterocampa_guttivitta; Bugwood Wiki fact
sheet on biology and distribution.

Articles Fisher, G.T. 1970. Parasites and predators of the species of a saddled
prominent complex at Groton, Vermont. Journal of Economic
Entomology 63: 1613-1614.
Allen, D.C. 1972. Insect parasites of the saddled prominent,
Heterocampa guttivitta (Lepidoptera: Notodontidae) in the northeastern
United States. The Canadian Entomologist 104: 1609-1622.
Allen, D.C. 1973. Fecundity of the saddled prominent, Heterocampa
guttivitta. Annals of the Entomological Society of America 66:
1181-1183.
Grimble, D.G. and R.G. Newell. 1973. Damage to sugar maple in
New York State from saddled prominent defoliation. Applied Forestry
Research Institute Note No. 7. Syracuse, NY, Syracuse University: 4 p.
Spear-O’Mara, J. and D.C. Allen. 2007. Monitoring populations of
saddled prominent (Lepidoptera: Notodontidae) with pheromone-baited
traps. Journal of Economic Entomology 100: 335-342.
600 Forest Pest Insects in North America:

166. Walnut Caterpillar, Datana integerrima Grote and Robinson

(Lepidoptera: Notodontidae)

Orientation to Pest Walnut caterpillar, Datana integerrima Grote and Robinson, is a native
North American notodontid whose larvae feed on walnut (Juglans),
butternut (J. cinerea L.), pecan (Carya illinoinensis [Wangenh.] K.
Koch), and hickory (C. tomentosa [Poir.] Nutt.). It is found in southern
Ontario (Canada) and throughout the eastern United States. Adults fly
in spring and summer and lay their eggs in masses on the undersides of
the leaves of the host plants. The larvae feed gregariously until almost
mature and are often seen in masses on the trunk and larger limbs, where
they congregate to molt. Mature larvae drop to the ground and wander
before pupating in the soil. There are one or two generations per year,
depending on the location.

Hosts Commonly Attacked This caterpillar feeds on walnut (Juglans), butternut (J. cinerea), pecan
(C. illinoinensis), and hickory (C. tomentosa).

Distribution The walnut caterpillar is found in southern Ontario (Canada) and

throughout the eastern United States.

Images of
Walnut Caterpillar

Figure 1. Adult of walnut caterpillar, Datana integerrima. (Jerry Armstrong,

Bugwood.org)
A Photographic Guide 601

Images of Walnut
Caterpillar (continued)

Figure 2. Eggs of walnut caterpillar. (Lacy L. Hyche, Auburn University,

Bugwood.org)

Figure 3. Larvae (middle instars) of walnut caterpillar. (Lacy L. Hyche, Auburn

University, Bugwood.org)

Figure 4. Larvae of walnut caterpillar feed in groups. (Jerry A. Payne, USDA

Agricultural Research Service, Bugwood.org)
602 Forest Pest Insects in North America:

Images of Walnut Figure 5. Walnut

caterpillars aggregating
Caterpillar (continued) on branch for molting.
(Jerry A. Payne, USDA
Agricultural Research
Service, Bugwood.org)

Figure 6. Leaf skeletonizing by young larvae of walnut caterpillar. (Lacy L. Hyche,

Auburn University, Bugwood.org)
A Photographic Guide 603

Images of Walnut Figure 7. Complete

defoliation of small tree
Caterpillar (continued) by walnut caterpillar.
(Lacy L. Hyche,
Auburn University,
Bugwood.org)

Important Biological Control Known parasitoids of the walnut caterpillar include Telenomus
Agents Related to this ichthyurae Ashmead and Tetrastichus sp. in Texas.
Pest Species

Web Links for Information http://www.na.fs.fed.us/spfo/pubs/fidls/walnutcat/walnutfidl.htm; USDA

on Walnut Caterpillar Forest Service, Pest and Disease Leaflet No. 41
http://www.ento.okstate.edu/ddd/insects/walnutdatana.htm; fact sheet of
Oklahoma State University on biology and life cycle.
http://www.ag.auburn.edu/enpl//bulletins/walnutcaterpillar/
walnutcaterpillar.htm; fact sheet of Auburn University (Alabama, USA)
on biology and control.
http://repository.tamu.edu/bitstream/handle/1969.1/87115/pdf_1558.
pdf?sequence=1; fact sheet of the Texas A&M University on biology
and control.
604 Forest Pest Insects in North America:

Articles on Tedders, W.L. and H.C. Ellis. 1977. Aerial application of Bacillus
Walnut Caterpillar thuringiensis var. kurstaki (HD-1) to shade and ornamental pecan trees
against Hyphantria cunea and Datana integerrima. Journal of the
Georgia Entomological Society 12: 248-250.

Farris, M.E. 1978. Bionomics of the walnut caterpillar, Datana

integerrima G. & R. Urbana, Il: University of Illinois, Dissertation.
106 p.

Farris, M.E. and J.E. Appleby. 1979. The walnut caterpillar, Datana
integerrima G. & R. In: Anon. Walnut insects and diseases, workshop
proceedings. General Technical Report NC-52. U.S. Department of
Agriculture, Forest Service, North Central Forest Experiment Station,
St. Paul, MN: 22-28.
A Photographic Guide 605

167. Variable Oak Leaf Caterpillar, Lochmaeus manteo (Doubleday)

(Lepidoptera: Notodontidae)

Orientation to Pest Variable oak leaf caterpillar, Lochmaeus manteo (Doubleday), is a native
North American moth whose larvae feed on a wide variety of deciduous
trees, especially oaks. The species is found across much of eastern North
America. It overwinters as a prepupa in the soil and pupation occurs in
early spring, with adult emergence following in May or June. Eggs are
laid singly on the leaves of the host plant. The color pattern of larvae
is variable. Young larvae skeletonize leaves, but older larvae eat entire
leaves apart from the larger veins. There are two generations per year in
the southern United States and one in the north. Outbreaks sometimes
occur that defoliate large areas; however, tree mortality is usually very
low.

Hosts Commonly Attacked All species of oaks are attacked, but white oak (Quercus alba L.) is
preferred. Other hosts include American beech (Fagus grandifolia
Ehrh.), American basswood (Tilia americana L.), paper birch (Betula
papyrifera Marshall), American elm (Ulmus americana L.), walnut
(Juglans), boxelder (Acer negundo L.), persimmon (Diospyros sp.), and
apple (Malus).

Distribution The variable oak leaf caterpillar is found in a wide area in eastern North
America, east of a line from western Ontario to eastern Texas.

Images of Variable
Oak Leaf Caterpillar

Figure 1. Adult of variable oak leaf caterpillar, Lochmaeus manteo. (USDA Forest
Service - Northeastern Area Archive, USDA Forest Service, Bugwood.org)
606 Forest Pest Insects in North America:

Images of Variable
Oak Leaf Caterpillar
(continued)

Figure 2. Eggs of variable oak leaf caterpillar. (Ronald F. Billings, Texas Forest
Service, Bugwood.org)

Figure 3. Young larvae of variable oak leaf caterpillar. (Ronald F. Billings, Texas
Forest Service, Bugwood.org)

Figure 4. Mature larvae of the variable oak leaf caterpillar (color patterns vary).
(Left: Gerald J. Lenhard, Lousiana State University, Bugwood.org; right:
Scott Tunnock, USDA Forest Service, Bugwood.org)
A Photographic Guide 607

Images of Variable
Oak Leaf Caterpillar
(continued)

Figure 5. Pupae of variable oak leaf caterpillar. (USDA Forest Service -

Northeastern Area Archive, USDA Forest Service, Bugwood.org)

Figure 6. Feeding damage of variable oak leaf caterpillar. (Ronald F. Billings,

Texas Forest Service, Bugwood.org)

Important Biological Control Among the nature enemies of this insect are egg parasitoids in the
Agents Related to this genera Trichogramma and Telenomus, which may kill 90 percent of
Pest Species the eggs. Nearly all egg masses have some parasitized eggs; only the
eggs concealed within a cluster escape. This high level of parasitism
and the failure of many prepupae to pupate in the spring, appear to be
the major factors affecting population density. At least seven species
of larval parasites attack variable oakleaf caterpillar larvae. The most
important species are Diradops bethunei Cresson (Ichneumonidae),
Protomicroplitus schizurae (Braconidae), and Lespesia schizurae
Sabrosky (Tachinidae), which collectively may kill up to 90 percent
of the larvae.
608 Forest Pest Insects in North America:

Web Links for Information on http://www.fs.fed.us/r8/foresthealth/pubs/oakpests/p5.html; USDA Forest

Variable Oak Leaf Caterpillar Service Southern Region fact sheet on biology and control.
http://www.maine.gov/doc/mfs/VariableOakLeafCaterpillar_new.htm;
fact sheet of the Maine Forest Service on biology and control.
http://na.fs.fed.us/spfo/pubs/fidls/variableOLC/voc.htm; USDA Forest
Service Forest Pest and Disease Leaflet No. 67.

Articles Surgeoner, G.A. and W.E. Wallner. 1978. Foliage consumption by

the variable oak leaf caterpillar, Heterocampa manteo (Lepidoptera:
Notodontidae), its use in defoliation predictions. The Canadian
Entomologist 110: 241-244.
A Photographic Guide 609

168. Fall Webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae)

Orientation to Pest Fall webworm, Hyphantria cunea (Drury), is a native North American
tiger moth, the larvae of which feed on the leaves of more than 100
species of forest and shade trees. It is found in southern Canada and
throughout the United States. Adults usually fly in late spring or early
summer and lay masses of eggs on the undersides of leaves. In the
southern United States, adults may be present from early spring through
late summer. Larvae web foliage together and feed in groups inside
a web, which is expanded as the larvae grow. Young larvae are upper
surface skeletonizers, but older larvae consume the entire leaf except
for the larger veins. Nearly mature larvae may feed alone outside the
web. Larvae move to the ground when ready to pupate, where they spin
a thin cocoon in the duff. Regional color variation exists in both adults
and larvae. There are one to four generations per year, with multiple
generations occurring at more southern latitudes. In its native range this
insect has little impact on tree health as defoliation occurs late in the
year, when most nutrients are in the roots. It is invasive in Eurasia, and
there the level of damage appears to be greater.

Hosts Commonly Attacked Fall webworm feeds on a more than 100 species of deciduous forest
or shade trees. Commonly defoliated plants include hickory (Carya
tomentosa [Poir.] Nutt.), pecan (C. illinoinensis [Wangenh.] K. Koch),
walnut (Juglans), elm (Ulmus), alder (Alnus), willow (Salix), mulberry
(Morus), oak (Quercus), American sweetgum (Liquidambar styraciflua
L.), poplar (Populus) and numerous species of Rosaceae, e.g., cherry
(Prunus), apple (Malus), and hawthorn (Crataegus).

Distribution Fall webworm is found in southern Canada and throughout the United
States.

Images of
Fall Webworm

Figure 1. Adults of fall webworm, Hyphantria cunea, may be pure white or

bear black spots. (Left: Jerry A. Payne, USDA Agricultural Research Service,
Bugwood.org; right: Gerald J. Lenhard, Louiana State University, Bugwood.org)
610 Forest Pest Insects in North America:

Images of Fall Figure 2. Top, eggs

of fall webworm near
Webworm (continued) adult female; bottom,
eggs with newly
hatched larvae.
(Top: Lacy L. Hyche,
Auburn University,
Bugwood.org; bottom:
Ministry of Agriculture
and Regional
Development Archive,
Ministry of
Agriculture and
Regional Development,
Bugwood.org)

Figure 3. Left, young larvae of fall webworm; right, group of young larvae feeding.
(Left: Milan Zubrik, Forest Research Institute - Slovakia, Bugwood.org; right:
Lacy L. Hyche, Auburn University, Bugwood.org)
A Photographic Guide 611

Images of Fall Figure 4. Top, mature

larvae of fall webworm;
Webworm (continued) bottom, larva of orange
race. (Top: James B.
Hanson, USDA Forest
Service, Bugwood.org;
bottom: Lacy L. Hyche,
Auburn University,
Bugwood.org)

Figure 5. Pupa of fall

webworm. (Ministry
of Agriculture and
Regional Development
Archive, Ministry of
Agriculture and Regional
Development,
Bugwood.org)

Figure 6. Close up of
feeding of fall webworm.
(Milan Zubrik, Forest
Research Institute -
Slovakia, Bugwood.org)
612 Forest Pest Insects in North America:

Images of Fall Figure 7. Feeding and

webs of fall webworm.
Webworm (continued) (Top: Clemson
University - USDA
Cooperative Extension
Slide Series,
Bugwood.org; bottom:
Steven Katovich,
USDA Forest Service,
Bugwood.org)
A Photographic Guide 613

Important Biological Control Many different species of parasitoids are associated with the larvae and
Agents Related to this pupae of this species in different locations.
Pest Species

Web Links for Information http://ohioline.osu.edu/hyg-fact/2000/2026.html; fact sheet of Ohio State

on Fall Webworm University on biology and control.
http://ento.psu.edu/extension/factsheets/fall-webworm; fact sheet of
Pennsylvania State University.
http://www.ag.auburn.edu/enpl/bulletins/fallwebworm/fallwebworm.htm;
fact sheet of Auburn University (Alabama, USA).
http://www.umassgreeninfo.org/fact_sheets/defoliators/fall_webworm.
html; fact sheet of the University of Massachusetts.
http://www.oregon.gov/ODF/privateforests/docs/fh/FallWebworm.
pdf?ga=t; fact sheet of the Oregon Department of Forestry.
http://edis.ifas.ufl.edu/in878; extension publication of the University of
Florida.

Articles Morris, R.F. 1963. Synonymy and color variation in the fall webworm,
Hyphantria cunea Drury (Lepidoptera: Arctiidae). The Canadian
Entomologist 95: 1217-1223.
Oliver, A.D. 1964. Studies on the biological control of the fall webworm,
Hyphantria cunea, in Louisiana. Journal of Economic Entomology 57:
314-318.
Warren, L.O. and M. Tadic. 1970. The fall webworm, Hyphantria cunea
(Drury). Arkansas Agricultural Experiment Station Bulletin No. 759:
106 p.
Yang, Z.Q., J.R. Wei, and X.Y. Wang. 2006. Mass rearing and
augmentative releases of the native parasitoid Chouioia cunea for
biological control of the introduced fall webworm Hyphantria cunea in
China. BioControl 51: 401-418.
Wang, Y., F.Q. Goo, Z. Xu, X. Chen, K. Li, Y.F. Xiao, S.H. Sun, and
J.Y. Qi. 2010. Parasitoids and mortality of the overwintering pupal
stage of Hyphantria cunea (Drury). Journal of Shenyang Agricultural
University 41(6): 686-689.
614 Forest Pest Insects in North America:

169. Arborvitae Leafminer, Argyresthia thuiella (Packard) (Lepidoptera:

Argresthiidae)

Orientation to Pest The arborvitae leafminer, Argyresthia thuiella (Packard), is a native

North American moth that forms mines in foliage of eastern arborvitae
(northern white cedar) trees (Thuja occidentalis L.) in forests and
landscapes. This moth is found in eastern Canada and in the United
States, south to Maryland and west to Missouri. It is also invasive in
Europe. Adults fly in spring to early summer and adults lay eggs in the
axils of branchlets or along the edges of leaves. Newly emerged larvae
bore into the leaves and feed the rest of the year there as leafminers.
Winter is passed as larvae in mines and pupation occurs in the mine.
Adults emerge the following spring. Outbreaks in Maine have severely
damaged arborvitae. Damage can also occur in nurseries or on landscape
arborvitae.

Hosts Commonly Attacked This caterpillar feeds on eastern arborvitae (northern white cedar) trees
(T. occidentalis).

Distribution The moth is found in eastern Canada and in the United States, south to
Maryland and west to Missouri.

Images of Figure 1. Adult of

arborvitae leafminer,
Arborvitae Leafminer Argyresthia thuiella
(moth wingspan is
8 mm); bottom, moth
in natural pose.
(Top: Petr Kapitola,
State Phytosanitary
Administration,
Bugwood.org; bottom:
Tom Murray)
A Photographic Guide 615

Images of Arborvitae
Leafminer (continued)

Figure 2. Mature larvae of arborvitae leafminer, removed from mine. (John A.

Weidhass, Virginia Polytechnic Institute and State University, Bugwood.org)

Figure 3. Top, mined

(brown) arborvitae
foliage; bottom, view
of larvae in mine (cut
away). (Top: Connecticut
Agricultural Experiment
Station Archive,
Connecticut Agricultural
Experiment Station,
Bugwood.org; bottom:
Robert Childs, University
of Massachusetts,
Bugwood.org)
616 Forest Pest Insects in North America:

Images of Arborvitae
Leafminer (continued)

Figure 4. Arborvitae leafminer pupa. (John A. Weidhass, Virginia Polytechnic

Institute and State University, Bugwood.org)

Figure 5. View of
damage on landscape
arborvitae; top, close
up; bottom, general
view of damaged plant.
(Both photos: Robert
Childs, University
of Massachusetts,
Bugwood.org)

Important Biological Control In Canada, natural control of arborvitae leafminers is provided by the
Agents Related to this widespread parasitoid Pentacnemus bucculatricis Howard and also by
Pest Species Apanteles bedelliae Viereck. Some 26 species of parasitoids are known
to attack this leafminer.
A Photographic Guide 617

Web Links for Information http://www.umassgreeninfo.org/fact_sheets/leaf_miners/arborvitae_

on Arborvitae Leafminer leafminer.html; fact sheet of the University of Massachusetts on biology
and control.
http://www.entomology.umn.edu/cues/Web/064ArborvitaeLeafminer.pdf;
fact sheet of the University of Minnesota on biology and control.
http://ccesuffolk.org/assets/Horticulture-Leaflets/Arborvitae-Leafminer.
pdf; fact sheet of Cornell Extension.

Articles Kurir, A. 1983. On the dispersal and biology of the North American
arborvitae leafminer, Argyresthia thuiella Packard (Lep., Argyresthiidae)
in Austria. Anzeiger für Schädlingskunde Pflanzenschutz Umweltschutz
56(7): 125-128. (In German).
Anon. 1983. Cedar leafminers. Argyresthia spp. Insect Identification
Sheet, Agriculture Canada (74): 2 p.
618 Forest Pest Insects in North America:

170. Pine Needle Sheathminer, Zelleria haimbachi Busck (Lepidoptera:

Yponomeutidae)

Orientation to Pest Pine needle sheathminer, Zelleria haimbachi Busck, is a native North
American moth that feeds on needles of jack pine (Pinus banksiana
Lamb.) in Ontario (Canada) and the Great Lakes region of the United
States, and on ponderosa (P. ponderosa P. & C. Lawson) and lodgepole
(P. contorta Douglas) in western North America north of Mexico. Larvae
create loose silken tunnels next to the bark of first-year shoots, from
where they bore through the sheaths and eat the needle bases. Needles
die and turn tan, and are loosely held in the sheath. Dead needles easily
slip out of the sheaths with a gentle pull. Larvae escape disturbance by
wriggling backwards and falling off the shoot, to hang suspended by a
strand of silk.

Hosts Commonly Attacked This caterpillar feeds on jack (P. banksiana), lodgepole (P. contorta), and
ponderosa pine (P. ponderosa).

Distribution The moth is found in Ontario (Canada), the Great Lakes region of the
United States, California, and other parts of the western United States.

Images of Figure 1. Adult of pine

needle sheathminer,
Pine Needle Sheathminer Zelleria haimbachi.
(Chris Grinter)

Figure 2. Oviposition
scar in needles where
eggs have been
deposited. (Donald
Owen, California
Department of Forestry
and Fire Protection,
Bugwood.org)
A Photographic Guide 619

Images of Pine Needle

Sheathminer (continued)

Figure 3. Young larvae of pine needle sheathminer, in its sheath feeding stage.
(Donald Owen, California Department of Forestry and Fire Protection,
Bugwood.org)

Figure 4. Mature larvae of pine needle sheathminer. (Left: Connecticut

Agricultural Experiment Station Archive, Connecticut Agricultural Experiment
Station, Bugwood.org; right: Donald Owen, California Department of Forestry
and Fire Protection, Bugwood.org)

Figure 5. Pupae of pine needle sheathminer (left), also showing close up of

webbing and frass associated with feeding; feeding damage seen from a
distance (right) on ponderosa pine. (Left: Scott Tunnock, USDA Forest Service,
Bugwood.org; right: Ward Strong, BC Ministry of Forests, Bugwood.org)
620 Forest Pest Insects in North America:

Important Biological Control No information on natural enemies of this species was found.
Agents Related to this
Pest Species
Web Links for Information None
on Pine Needle Sheathminer

Articles Stevens, H.E. 1959. Biology and control of the pine needle-sheath miner,
Zelleria haimbachi Busck (Lepidoptera; Hyponomeutidae). Pacific
Southwest Forest and Range Experiment Station: 20 p.
A Photographic Guide 621

171. Pine Butterfly, Neophasia menapia (Felder and Felder) (Lepidoptera:

Pieridae)

Orientation to Pest The pine butterfly, Neophasia menapia (Felder and Felder), is a native
North American butterfly that is an important pest of ponderosa pine
(Pinus ponderosa P. & C. Lawson) in the western United States, where
large outbreaks have occurred. Adults fly in late summer and fall, and
eggs are laid in rows in groups of 5-20 on current year needles. Eggs
overwinter and hatch the following June, or about the time that new
needles begin to appear on ponderosa pine. Young larvae eat the old
needles first, but new needles are eaten later by older larvae. Larvae
attach themselves to needles, twigs, bark, or other objects before
pupating. In 2 to 3 weeks, adults emerge and complete the cycle. There
is one generation per year. Older trees are more susceptible to injury than
younger thriftier trees. Outbreaks usually last 3 or 4 years; most cause
little mortality, although intense and widespread outbreaks may result in
reduction in tree growth. A few outbreaks have killed extensive stands of
older trees when aided by bark beetles.

Hosts Commonly Attacked In most areas, this caterpillar feeds mainly on ponderosa pine (P.
ponderosa), but during outbreaks other conifers may also be attacked if
intermixed with ponderosa pine, such as lodgepole (P. contorta Douglas)
and western white (P. monticola Douglas ex D. Don) pines, Douglas-fir
(Pseudotsuga menziesii [Mirb.] Franco), grand fir (Abies grandis [Dougl.
ex D. Don] Lindl.), and western larch (Larix occidentalis Nutt.). In
the coastal area of northwestern Washington and on Vancouver Island,
British Columbia, Canada, outbreaks have occurred in Douglas-fir
stands.

Distribution The butterfly is found throughout the western United States; but
outbreaks have occurred primarily in Oregon, Washington, Idaho,
Montana, and British Columbia.

Images of
Pine Butterfly

Figure 1. Adult female (left) and male (right) of pine butterfly, Neophasia
menapia. (Left: Ladd Livingston, Idaho Department of Lands, Bugwood.org;
right: Terry Spivey, USDA Forest Service, Bugwood.org)
622 Forest Pest Insects in North America:

Images of Pine Butterfly

(continued)

Figure 2. Eggs of pine butterfly. (USDA Forest Service - Ogden Archive, USDA
Forest Service, Bugwood.org)

Figure 3. Young larvae of pine butterfly. (Left: Scott Tunnock, USDA Forest
Service, Bugwood.org; right: USDA Forest Service - Ogden Archive, USDA
Forest Service, Bugwood.org)
A Photographic Guide 623

Images of Pine Butterfly Figure 4. Mature larva

of pine butterfly. (Ladd
(continued) Livingston, Idaho
Department of Lands,
Bugwood.org)

Figure 5. Pupa of pine

butterfly. (Jerald E.
Dewey, USDA Forest
Service, Bugwood.org)
624 Forest Pest Insects in North America:

Images of Pine Butterfly

(continued)

Figure 6. Feeding damage of pine butterfly on ponderosa pines. (USDA Forest

Service - Ogden Archive, USDA Forest Service, Bugwood.org)

Important Biological Control Several insects, including hemipterans (Podisus spp. and Apoecilus spp.),
Agents Related to this snakeflies (Raphidia spp.), sarcophagid flies (Agria spp.), and wasps
Pest Species (Ichneumonidae) have been reported preying on different stages of pine
butterfly. Decline of the 1922-23 outbreak of this butterfly was attributed
to mortality from some of these natural enemies, especially a native
ichneumon parasite, Theronia atalantae (Poda).

Web Links for Information http://butterfly.ucdavis.edu/butterfly/Neophasia/menapia; website on this

on Pine Butterfly insect from a natural history point of view.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=1000029&
ind=P; fact sheet of Natural Resource Canada on biology and phenology.
http://extension.oregonstate.edu/mwm/w10fhmbutterfly; extension note
of Oregon State University.

Articles Evenden, J.C. 1926. The pine butterfly, Neophasia menapia Felder.
Journal of Agricultural Research 33(4): 339-344.
Evenden, J.C. 1940. Effects of defoliation by the pine butterfly upon
ponderosa pine. Journal of Forestry 38(12): 949-955.
Cole, W.E. 1966. Effect of pine butterfly defoliation on ponderosa pine in
southern Idaho. Research Note INT-46. Ogden, UT: U.S. Department of
Agriculture, Forest Service, Intermountain Forest and Range Experiment
Station: 7 p.
Young, R.M. 1986. Mass emergences of the pine white, Neophasia
menapia menapia (Felder & Felder), in Colorado (Pieridae). Journal of
the Lepidopterists’ Society 40 (4): 314.
A Photographic Guide 625

172. Balsam Gall Midge, Paradiplosis tumifex Gagné (Diptera:

Cecidomyiidae)

Orientation to Pest Balsam gall midge, Paradiplosis tumifex Gagné, is a native North
American cecidomyiid fly that galls needles of balsam (Abies balsamea
[L.] Miller) and Fraser fir (A. fraseri [Pursh] Poir.). In older literature,
these galls were mistakenly attributed to another species, Dasineura
balsamicola (Lintner), which is an inquiline that invades the galls of
P. tumifex, killing it and feeding on the gall tissue. Paradiplosis tumifex
has a wide distribution, being found throughout the ranges of its two
main hosts. The gall midge overwinters as a larva in the soil under an
infested host tree. Pupation takes place in spring, and adult emergence
occurs shortly thereafter. Eggs are laid in developing needles, and larval
feeding causes quick development of gall tissue in immature needles.
In late fall, larvae leave the galls and drop to the ground for the winter.
There is one generation per year. Galling is of concern only in Christmas
tree plantations, not forests.

Hosts Commonly Attacked This gall midge attacks needles of balsam fir (A. balsamea) and Fraser fir
(A. fraseri).

Distribution This species is found throughout the ranges of balsam and Fraser
firs, being much of southeastern Canada and adjacent areas of the
northeastern United States (range of balsam fir), plus the sky islands of
the southern Appalachian Mountains (range of Fraser fir).

Images of
Balsam Gall Midge

Figure 1. Adult balsam gall midge, Paradiplosis tumifex. (Ronald S. Kelley,

Vermont Department of Forests, Parks and Recreation, Bugwood.org)
626 Forest Pest Insects in North America:

Images of Balsam
Gall Midge (continued)

Figure 2. Balsam gall midge galls on balsam fir. (Ronald S. Kelley, Vermont
Department of Forests, Parks and Recreation, Bugwood.org)

Figure 3. Larvae of
the balsam gall midge
exposed by opening the
gall. (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Important Biological Control Parasitoids of this gall maker include the encyrtids Pseudoencyrtus
Agents Related to this borealis MacGown, Tetrastichus cecidivorus MacGowan, Tetrastichus
Pest Species marcovitchi (Crawford), Tetrastichus whitmani (Firault), and the
platygasterids Platygaster abicollis MacGown and Osgood and
Playtgaster mainensis MacGown and Osgood.

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=3708; fact

on Balsam Gall Midge sheet of Natural Resources Canada on biology and phenology.
http://www.maine.gov/doc/mfs/balsgalmid.htm; fact sheet of the state
of Maine.
http://extension.unh.edu/resources/files/Resource000986_Rep2348.pdf;
highly detailed power point presentation on biology and control of this
gall maker.
A Photographic Guide 627

Articles Osgood, E.A. and R.J. Gagné. 1978. Biology and taxonomy of two
gall midges (Diptera : Cecidomyiidae) found in galls on balsam fir
needles with description of a new species of Paradiplosis. Annals of the
Entomological Society of America 71: 85-91.
West, R.J. and J.D. Shorthouse. 1982. Morphology of the balsam
fir needle gall induced by the midge Paradiplosis tumifex (Diptera:
Cecidomyiidae). Canadian Journal of Botany 60: 131-140.
Räther, M. and N.J. Mills. 1989. Possibilities for the biological control
of the Christmas tree pests, the balsam gall midge, Paradiplosis tumifex
Gagné (Diptera: Cecidomyiidae) and the balsam twig aphid, Mindarus
abietinus Koch (Homoptera: Mindaridae), using exotic enemies from
Europe. Biocontrol News and Information 10(2): 119-129.
Osgood, E.A., R.L. Bradbury, and F.A. Drummond. 1992. The balsam
gall midge - an economic pest of balsam fir Christmas trees. Maine
Agricultural Experiment Station Technical Bulletin No. 151: 30 p.
(Available at http://library.umaine.edu/maineaes/technicalbulletin/tb151.
pdf).
628 Forest Pest Insects in North America:

173. Poplar Leafcurl Midge, Prodiplosis morrisi Gagné (Diptera:

Cecidomyiidae)

Orientation to Pest The poplar leafcurl midge, Prodiplosis morrisi Gagné, is a native North
American cecidomyiid fly that lays its eggs on developing cottonwood
terminals. The larvae feed on surfaces of expanding leaves, causing
them to curl into elongated, largely cylindrical shapes. Leaves may stay
green with only curled edges turning brown, or leaf may be killed. With
sufficient feeding, terminal shoots may become stunted or killed. Trees
of all sizes may be attacked. There may be up to five or more generations
per year. Each generation damages 1 to 2 whorls of leaves, and because
there is a short period between generations, normal leaves alternate
with damaged leaves. The many larvae that develop in each curled leaf
drop to the soil beneath the tree to pupate. Larvae of the last generation
overwinter in the soil.

Hosts Commonly Attacked This midge attacks various species of Populus, including eastern
cottonwood (P. deltoides W. Bartram ex Humphry Marshall), quaking
aspen (P. tremuloides Michx.), and Populus hybrids.

Distribution This species is of greatest importance in the southern United States, in

cottonwood plantations.

Images of Figure 1. Injury to

cottonwood seedling
Poplar Leafcurl Midge from poplar leafcurl
midge, Prodiplosis
morrisi. (James
Solomon, USDA Forest
Service, Bugwood.org)
A Photographic Guide 629

Important Biological Control No information is available on natural enemies of this species.

Agents Related to this
Pest Species
Web Links for Information http://wiki.bugwood.org/Archive:Poplar/Prodiplosis_morrisi; Bugwood
on Poplar Leafcurl Midge Wiki fact sheet on biology.

Articles Gagné, R.J. 1966. A new species of Prodiplosis (Diptera: Cecidomyiidae)

found in leaf curls on Populus deltoides. Annals of the Entomological
Society of America 59: 1154-1157.
Gagné, R.J. 1989. The Plant-feeding Gall Midges of North America.
Cornell University Press. Ithaca, New York: 340 p.
Ostry, M.E., L.F. Wilson, H.S. McNabb, Jr., and L.M. Moore. 1988.
A Guide to Insect, Disease, and Animal Pests of Poplars. Agricultural
Handbook No. 677. U.S. Department of Agriculture, Washington, DC:
118 p.
Skuhravý, V., M. Skuhravá and J.W. Brewer. 1997. Gall midges (Dipt.,
Cecidomyiidae) associated with Populus tremula, P. tremuloides and
their hybrid (Salicaceae). Journal of Applied Entomology 121: 315-320.
630 Forest Pest Insects in North America:

174. Douglas-fir Needle Midge, Contarinia pseudotsugae Condrashoff

(Diptera: Cecidomyiidae)

Orientation to Pest Douglas-fir needle midge, Contarinia pseudotsugae Condrashoff, is one

of three species in this genus of native North American cecidomyiid flies
that lay their eggs in developing needles of Douglas-fir (Pseudotsuga
menziesii [Mirb.] Franco). This is the most abundant of the three. It is
a significant pest of Douglas-fir when it is grown as Christmas trees,
particularly in the dry-belt of open-grown interior forests in the Pacific
northwestern United States in California, Oregon, Washington, Idaho,
Montana, and in British Columbia, Canada. Adults emerge from the soil
in spring and lay eggs on needles. The larvae bore into needles to feed,
causing their growth to be deformed. In the fall, mature larvae drop back
into the soil to pupate and overwinter. Damaged needles bear small galls
and heavily galled Christmas trees have lower market value. There is one
generation of this fly per year.

Hosts Commonly Attacked The only known host of this insect is Douglas-fir (P. menziesii).

Distribution This species is of concern in the Pacific northwestern United States

(California, Oregon, Washington, Idaho, and Montana) and British
Columbia, Canada, but is also found in Pennsylvania.

Images of Douglas-fir
Needle Midge

Figure 1. Adult of Douglas-fir needle midge, Contarinia pseudotsugae. (USDA

Forest Service Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 631

Images of Douglas-fir Figure 2. Larva of

Douglas-fir needle
Needle Midge (continued) midge inside its gall,
which has been cut
open. (Ward Strong,
BC Ministry of Forests,
Bugwood.org)

Figure 3. Douglas-
fir needles (top)
showing injury due
to Douglas-fir needle
midge; and, bottom,
damage in an earlier
stage. (Top: Elizabeth
Willhite, USDA Forest
Service, Bugwood.
org; bottom: Rayanne
Lehman, Pennsylvania
Department of
Agriculture,
Bugwood.org)

Figure 4. Damage to Douglas-fir from Douglas-fir needle midge. (Left: USDA

Forest Service Archive, USDA Forest Service, Bugwood.org; right: Ward Strong,
BC Ministry of Forests, Bugwood.org)
632 Forest Pest Insects in North America:

Important Biological Control No information is available on natural enemies of this species.

Agents Related to this
Pest Species
Web Links for Information http://extension.psu.edu/ipm/program/christmas-tree/pest-fact-
on Douglas-fir Needle Midge sheets/needle-discoloration-and-injury/Douglas-fir.pdf; fact sheet of
Pennsylvania State University on biology and control.
http://oregonstate.edu/dept/nurserystartup/onnpdf/onn060108.pdf; fact
sheet of Oregon State University, helpful in determining a spray schedule
for this pest.
http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5187407.
pdf; fact sheet of the USDA FS Forest Health Protection unit on
recognition and biology.

Articles Condrashoff, S.F. 1961. Three new species of Contarinia (Diptera:

Cecidomyiidae) in Douglas-fir needles. The Canadian Entomologist
93(2): 123-130.
Condrashoff, S.F. 1962. Bionomics of three closely related species of
Contarinia Cond. (Diptera: Cecidomyiidae) from Douglas-fir needles.
The Canadian Entomologist 94: 376-394.
A Photographic Guide 633

175. Pine False Webworm, Acantholyda erythrocephala (L.) (Hymenoptera:

Pamphiliidae)

Orientation to Pest Pine false webworm, Acantholyda erythrocephala (L.), is an introduced

pamphiliid sawfly of European origin that is now present in several
parts of eastern North America and Canada. It feeds on several pines.
Winter is passed as prepupae, which pupate in the spring. Adults emerge
soon after and lay eggs in short rows of 3 to 10 eggs in small slits cut
into last year’s needles. Young larvae spin loose webs in which they
feed gregariously on old needles, which they cut off and then pull into
the web to eat. Older larvae are solitary and feed from silken tubes
bound to twigs. Frass and bits of needles stick to these silk tubes. When
mature, larvae drop to the ground, build earthen cells, and overwinter as
prepupae. Heavy infestations can defoliate hosts and cause tree mortality.

Hosts Commonly Attacked This sawfly feeds especially on eastern white (Pinus strobus L.) and red
(P. resinosa Sol. ex Aiton), but also Scotch (P. sylvestris L.), Austrian
(P. nigra J. F. Arnold), and Japanese red (P. densiflora Siebold & Zucc.)
pines.

Distribution This sawfly is found in New England, New York, New Jersey,
Pennsylvania, Wisconsin, Minnesota, Newfoundland, Alberta, Quebec,
and Ontario.

Images of Pine Figure 1. Top, adult

male (right) and female
False Webworm (left) of pine false
webworm, Acantholyda
erythrocephala, in
copula; and, bottom,
adult female ovipositing
on red pine foliage.
(Both photos: Barry
Lyons, Canadian Forest
Service, Bugwood.org)
634 Forest Pest Insects in North America:

Images of Pine
False Webworm
(continued)

Figure 2. Eggs of pine false webworm on red pine foliage. (Barry Lyons,
Canadian Forest Service, Bugwood.org)

Figure 3. Top, neonate larva of pine false webworm emerging from the egg;
and bottom, ultimate instar larva climbing on silk strands outside the web. (Both
photos: Barry Lyons, Canadian Forest Service, Bugwood.org)
A Photographic Guide 635

Images of Pine Figure 4. Damage

(shoots with clipped off
False Webworm needles) by pine false
(continued) webworm on young
plantation-grown red
pine. (Barry Lyons,
Canadian Forest
Service, Bugwood.org)

Figure 5. Defoliation in pine plantations in Simcoe Co., Ontario, due to pine false
webworm. (Barry Lyons, Canadian Forest Service, Bugwood.org)
636 Forest Pest Insects in North America:

Important Biological Control Since this is an invasive species in North America, the parasitoids from
Agents Related to this its native range in Europe have been investigated for introduction. The
Pest Species most important such species is the tachinid Myxexoristops hertingi
Mesnil, which has been released in Ontario, Canada.

Web Links for Information http://bugs.osu.edu/bugdoc/Shetlar/factsheet/christmasstree/pine_false_

on Pine False Webworm webworm.htm; fact sheet of Ohio State University on biology and
control of pine false webworm for Christmas tree growers.
http://www.dec.ny.gov/docs/lands_forests_pdf/pfw.pdf; article
concerning importance of pine false webworm in New York.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=1000130;
fact sheet of Natural Resources Canada.

Articles Lyons, D.B. 1995. Pine false webworm, Acantholyda erythrocephala. In:
Armstrong, J.A. and W.G.H. Ives (eds.). Forest Insect Pests in Canada.
Natural Resources Canada, Canadian Forest Service, Ottawa: 245-251.
Kessel, C. 1998. The impact of the pine false webworm on the
Ontario Christmas tree industry. In: Lyons, D.B., G.C. Jones, and T.A.
Scarr (eds.). Proceedings of a workshop on the pine false webworm,
Acantholyda erythrocephala (Hymenoptera: Pamphiliidae). University
of Toronto, Toronto, Ontario, Canada, 16 February 1998.
Asaro, C. and D.C. Allen. 2001. History of a pine false webworm
(Hymenoptera: Pamphiliidae) outbreak in northern New York. Canadian
Journal of Forest Research 31: 181-185.
Kenis, M. and K. Kloosterman. 2001. European parasitoids of the pine
false webworm (Acantholyda erythrocephala [L.]) and their potential for
biological control in North America. In: Liebhold, A.M., M.L. McManus,
I.S. Otvos, S.L.C. Fosbroke (eds.). Proceedings of a conference entitled
“Integrated management and dynamics of forest defoliating insects” held
1999 August 15-19, Victoria, British Columbia, and published as General
Technical Report NE-277. U.S. Department of Agriculture, Forest
Service, Northeastern Research Station, Newtown Square, Pennsylvania,
USA: 65-73. (Available at http://iufro-archive.boku.ac.at/wu70307/
victoria/kenis.pdf).
Lyons, D.B., M. Kenis, and R.S. Bourchier. 2002. Acantholyda
erythrocephala (L.), pine false webworm (Hymenoptera: Pamphiliidae ).
In: Mason, P.G. and J.T. Huber (eds.). Biological Control Programmes in
Canada, 1981-2000. CABI, Wallingford, UK: 22-28.
A Photographic Guide 637

176. Redheaded Pine Sawfly, Neodiprion lecontei (Fitch) (Hymenoptera:

Diprionidae)

Orientation to Pest Redheaded pine sawfly, Neodiprion lecontei (Fitch), is a native North
American sawfly that defoliates various pines. Conifers in other genera
may also be fed on when they grow intermixed with preferred species.
This sawfly is found in southeastern Canada and throughout the eastern
United States. It overwinters as a prepupa in the soil. Pupation occurs
in spring and adults emerge soon after. Eggs are inserted in needles of
either the current or previous year’s growth. A single female may lay up
to 150 eggs. The larvae feed gregariously on either old or new needles,
or even bark of tender twigs. Young larvae partially consume needles
and cause them to dry out and curl, giving a straw-like appearance.
Older larvae may consume entire needles. In some cases, trees may be
completely defoliated. To prepare for winter, mature larvae drop from the
foliage to the ground, where they spin a cocoon and molt to the pre-pupal
stage. There are three generations per year in the southern United States
but only one in the northern part of the United States and in Canada.
This is one of the most damaging sawflies in North America. Trees at
greatest risk are stressed trees growing on shallow soils, very wet or
dry sites, or ones where trees are experiencing severe competition from
other vegetation. Reducing vegetative competition, avoiding planting
on poor sites, and using tight spacing to encourage early canopy closure
are recommended to reduce damage from this sawfly to pines grown in
plantations.

Hosts Commonly Attacked The main hosts of redheaded pine sawfly are jack (Pinus banksiana
Lambert), red (P. resinosa Sol. ex Aiton), shortleaf (P. echinata Mill.),
loblolly (P. taeda L.), slash (P. elliottii Engelm.), longleaf (P. palustris
Mill.), and pitch (P. rigida Mill.) pines.

Distribution This sawfly is found in southeastern Canada and throughout the eastern
United States.

Images of Redheaded Figure 1. Adult

redheaded pine sawfly,
Pine Sawfly Neodiprion leconte.
(Lacy L. Hyche,
Auburn University,
Bugwood.org)
638 Forest Pest Insects in North America:

Images of Redheaded Figure 2. Eggs of

redheaded pine sawfly
Pine Sawfly (continued) being laid (top) and,
bottom, close up of
eggs in pine needle.
(Top: James McGraw,
North Carolina State
University, Bugwood.org;
bottom: Lacy L. Hyche,
Auburn University,
Bugwood.org)

Figure 3. Larvae of redheaded pine sawfly; left, close up of two, and right,
a large aggregation of feeding larvae. (Left: Gerald J. Lenhard, Louisiana
State University, Bugwood.org; right: Clemson University - USDA Cooperative
Extension Slide Series, Bugwood.org)

Figure 4. Cocoons of
redheaded pine sawfly.
(Lacy L. Hyche, Auburn
University, Bugwood.org)
A Photographic Guide 639

Images of Redheaded Figure 5. Characteristic

“straw” damage caused
Pine Sawfly (continued) by redheaded pine
sawfly larvae. (G. Keith
Douce, University of
Georgia, Bugwood.org)

Figure 6. Damage (branches with clipped off needles) to longleaf pines by

redheaded pine sawfly. (Andrew J. Boone, South Carolina Forestry Commission,
Bugwood.org)
640 Forest Pest Insects in North America:

Important Biological Control Important natural enemies of this sawfly include a nuclear polyhedrosis
Agents Related to this virus that has been developed in Canada as a biopesticide, the egg
Pest Species parasitoid Closterocerus cinctipennis Ashmead, and tachinid larval
parasitoids in the genus Spathimeigenia.

Web Links for Information http://www.forestpests.org/vermont/redheadsawfly.html; fact sheet of the

on Redheaded Pine Sawfly Vermont Department of Forests, Parks, and Recreation on biology and
control.
http://edis.ifas.ufl.edu/in882; fact sheet of the University of Florida on
biology and control.
http://pubs.ext.vt.edu/3006/3006-1453/3006-1453_pdf.pdf; fact sheet of
the Virginia Tech extension on biology and control.
http://ohioline.osu.edu/hyg-fact/2000/2554.html; fact sheet of Ohio State
University extension on biology and control.

Articles Benjamin, D.M. 1955. The biology and ecology of the redheaded pine
sawfly. USDA Forest Service Technical Bulletin No. 1118: 57 p.
Drooz, A.T., R.C. Wilkinson, and V.H. Fedde. 1977. Larval and cocoon
parasites of three Neodiprion sawflies in Florida. Environmental
Entomology 6: 60-62.
Cunningham, J.C., P. DeGroot, and J.R. McPhee. 1984. Lecontvirus:
a viral insecticide for control of redheaded pine sawfly, Neodiprion
lecontei. Technical Note No. 2, Forest Pest Management Institute,
Canada: 5 p.
Hyche, L.L. 1992.The redheaded pine sawfly (Neodiprion lecontei):
a guide to recognition and habits. Bulletin No. 617 of the Alabama
Agricultural Experiment Station. Auburn University, Auburn, Alabama:
11 p.
Wilson, L.F., R.C. Wilkinson, Jr., and R.C. Averill. 1992. Redheaded pine
sawfly - its ecology and management. USDA Forest Service Handbook
No. 694: 54 p.
A Photographic Guide 641

177. Swaine Jack Pine Sawfly, Neodiprion swainei (Middleton)

(Hymenoptera: Diprionidae)

Orientation to Pest Swaine jack pine sawfly, Neodiprion swainei (Middleton), is a native
North American sawfly. It is considered to be the most important sawfly
pest of pines in eastern Canada. Its preferred host is jack pine (Pinus
banksiana Lambert), but other species of pines will also be defoliated
if they grow near an outbreak of this sawfly on jack pine. This sawfly
is found in the Great Lakes states of the United States and throughout
most of the range of jack pine in Canada (from Nova Scotia west to
Alberta). Outbreaks, though, have only been recorded in Ontario and
Quebec. Swaine jack pine sawfly overwinters as a prepupa in cocoons
in the litter or topsoil. Pupation occurs in spring and adults emerge
soon after. In June or July, females lay one to three eggs per needle in
the current year’s growth. The larvae feed gregariously on old needles,
usually on trees in exposed locations. Mature larvae drop from foliage to
the ground, where they spin a cocoon in the duff and molt to the pre-
pupa, the overwintering stage. There is one generation per year. Many
outbreaks of this sawfly occurred in Ontario and Quebec up until the
1960s, usually at about 8 year intervals, on poor sites (for jack pine) on
outwash plains. The last significant defoliation (in Canada) occurred
over 440 ha in Quebec in 1994. In stands of mature trees, most trees may
either be killed or, if they survive, will be stag-headed (killed on top). In
the Great Lakes states (in the United States), outbreaks have occurred
mainly in plantations or windbreaks. A nuclear polyhedrosis virus that
is specific to this sawfly was discovered in the 1960s and developed
as a potential biopesticide. It provided effective control but was never
developed as a commercial pesticide.

Hosts Commonly Attacked The main host of the Swaine jack pine sawfly is jack pine (P. banksiana
Lambert), but other pines that may be defoliated if near an outbreak
of this sawfly include red (P. resinosa Sol. ex Aiton), eastern white
(P. strobus L.), and Scotch (P. sylvestris L.) pines.

Distribution This sawfly is found in Canada from Nova Scotia west to Alberta and
in the Great Lakes states of the United States.

Images of Swaine Figure 1. Adult Swaine

jack pine sawfly,
Jack Pine Sawfly Neodiprion swainei.
(Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)
642 Forest Pest Insects in North America:

Images of Swaine Figure 2. Eggs of

Swaine jack pine
Jack Pine Sawfly sawfly. (Thérèse Arcand
(continued) - Natural Resources
Canada, Canadian
Forest Service)

Figure 3. Larvae of
Swaine jack pine sawfly.
(Both photos: Catherine
Linnen, University of
Kentucky, Bugwood.org)

Figure 4. Damage to
jack pine by Swaine jack
pine sawfly. (Claude
Monnier - Natural
Resources Canada,
Canadian Forest
Service)
A Photographic Guide 643

Images of Swaine Figure 5. Jack pines

defoliated by Swaine
Jack Pine Sawfly jack pine sawfly.
(continued) (Robert Gagnon
- Natural Resources
Canada, Canadian
Forest Service)

Important Biological Control Important natural enemies of this sawfly include a nuclear polyhedrosis
Agents Related to this virus and various larval parasitoids. Red wood ants (Formicidae) were
Pest Species also introduced from Manitoba and Italy into Quebec as pupal predators.

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=6585;

on Swaine Jack Pine Sawfly fact sheet of Natural Resources Canada on biology and control.

Articles Smirnoff, W.A. 1961. Predators of larvae of Neodiprion swainei Midd.

(Hymenoptera: Tenthredinidae). The Canadian Entomologist 93:
272-275.
Smirnoff, W.A., J.J. Fettes, and W. Haliburton. 1962. A virus disease of
Swaine’s jack pine sawfly, Neodiprion swainei Midd., sprayed from an
aircraft. The Canadian Entomologist 94: 477-486.
McLeod, J.M. 1970. The epidemiology of the Swaine jack-pine sawfly,
Neodiprion swainei Midd. The Forestry Chronicle April issue: 126-130.
(Available at http://pubs.cif-ifc.org/doi/pdf/10.5558/tfc46126-2; provides
details of timing and stand location where outbreaks have occurred in
Quebec).
Price, P.W. and H.A. Tripp. 1972. Activity patterns of parasitoids
on the Swaine jack pine sawfly, Neodiprion swainei (Hymenoptera:
Diprionidae), and parasitoid impact on the host. The Canadian
Entomologist 104: 1003-1016.
Price, P.W., R.H. MacArthur, and E.O. Wilson. 1973. Parasitoid
strategies and community organization. Environmental Entomology
2: 623-626.
Finnegan, R.J. and W.A. Smirnoff. 1984. Neodiprion swainei
(Middleton), Swaine jack pine sawfly (Hymenoptera: Diprionidae). In:
Kelleher, J.S. and M.A. Hulme (eds.). Biological Control Programmes
against Insects and Weeds in Canada 1969-1980. Commonwealth
Agricultural Bureaux, Slough, United Kingdom: 341-348.
644 Forest Pest Insects in North America:

178. European Pine Sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera:

Diprionidae)

Orientation to Pest European pine sawfly, Neodiprion sertifer (Geoffroy), is a European

species that is invasive in North America. It feeds on many species of
pines. This sawfly occurs across Canada and widely in the eastern United
States. The winter is passed as eggs, which hatch in early spring. Larvae
feed gregariously on old needles. New foliage is never eaten. Larvae
move from tree to tree as necessary for food. Mature larvae drop from
foliage to the ground, where they spin a cocoon in the duff and pupate.
Adults emerge in late summer and lay 6 to 8 eggs per needle in new
foliage. There is one generation per year. Trees attacked by this sawfly
are never killed because new foliage is not eaten. Damage consists
of reduced tree growth in forests or loss of quality in Christmas tree
plantations.

Hosts Commonly Attacked European pine sawfly feeds on many species of pines, but especially
on Scots (Pinus sylvestris L.), red (P. resinosa Sol. ex Aiton), jack
(P. banksiana Lambert), Japanese red (P. densiflora Siebold & Zucc.),
and Table mountain pines (P. pungens Lamb.). Other species of pines
will also be defoliated if they grow near an outbreak of this sawfly.

Distribution This sawfly is found in Newfoundland, Nova Scotia, New Brunswick,

Ontario, Quebec, and British Columbia in Canada, and in the eastern
United States from New England west to North Dakota, and south to
Missouri.

Images of European Figure 1. Adults

of European pine
Pine Sawfly sawfly, Neodiprion
sertifer; top, female;
bottom, male (note
enlarged antennae).
(Both photos: Louis-
Michel Nageleisen,
Département de la
Santé des Forêts,
Bugwood.org)
A Photographic Guide 645

Images of European
Pine Sawfly (continued)

Figure 2. Egg scars of European pine sawfly in needles of Scots pine. (Andrea
Battisti, Universita di Padova, Bugwood.org)

Figure 3. Larvae of
European pine sawfly.
(Top: Gyorgy Csoka,
Hungary Forest
Research Institute,
Bugwood.org; bottom:
Steven Katovich,
USDA Forest Service,
Bugwood.org)

Figure 4. Pupal cocoons

of European pine sawfly.
(Jim Occi, BugPics,
Bugwood.org)
646 Forest Pest Insects in North America:

Images of European
Pine Sawfly (continued)

Figure 5. Damage to red pine by European pine sawfly larvae; note that only
old needles are eaten. (Both photos: Steven Katovich, USDA Forest Service,
Bugwood.org)

Figure 6. Larvae of European pine sawfly killed by a virus (note the characteristic
position, hanging head down). (John D. Kegg, New Jersey Department of
Agriculture, Bugwood.org)
A Photographic Guide 647

Important Biological Control Important natural enemies of this sawfly include an imported European
Agents Related to this nuclear polyhedrosis virus that is used effectively as a biopesticide, and
Pest Species several European larval parasitoids, which have not provided control of
this pest.

Web Links for Information http://ohioline.osu.edu/hyg-fact/2000/2555.html; fact sheet of Ohio State

on European Pine Sawfly University on biology and control.
http://ento.psu.edu/extension/factsheets/european-pine-sawfly; fact sheet
of Pennsylvania State University on biology and control.
http://www.forestpests.org/vermont/europeansawfly.html; fact sheet of
Vermont Department of Forests, Parks, and Recreation.
http://www.exoticpests.gc.ca/ins_details_eng.asp?pestType=ins&lang=lt
&geID=6583; website of Forest Invasive Alien Species (Government of
Canada).

Articles Finlayson, L.R. and T. Finlayson. 1958. Parasitism of the European

Pine Sawfly, Neodiprion sertifer (Geoff.) (Hymenoptera: Diprionidae),
in Southwestern Ontario. The Canadian Entomologist 90: 223-225.
(Available at http://pubs.esc-sec.ca/doi/abs/10.4039/Ent90223-4).
Entwistle, P.F., H.F. Evans, K.A. Harrap, and J.S. Robertson. 1978. Field
trials on the control of the pine sawfly, Neodiprion sertifer, using purified
nuclear polyhedrosis virus. Technical Report, Unit of Invertebrate
Virology, Natural Environment Research Council (UK): 62 p.
Gur’yanova, T.M. 1979. Significance of the diapause of the parasite
Exenterus abruptorius (Ichneumonidae) in the dynamics of numbers of
the red pine sawfly (Neodiprion sertifer). Zoologicheskii Zhurnal 58 (9):
1339-1349.
648 Forest Pest Insects in North America:

179. Virginia Pine Sawfly, Neodiprion pratti pratti (Dyar) (Hymenoptera:

Diprionidae)

Orientation to Pest Virginia pine sawfly, Neodiprion pratti pratti (Dyar), is a native North
American sawfly that feeds on Virginia (Pinus virginiana Mill.) and
shortleaf (P. echinata Mill.) pines. It is found from New Jersey south to
North Carolina and west to Illinois. The winter is passed as eggs, which
hatch in early spring. Larvae feed gregariously on old needles. Mature
larvae drop from foliage to the ground, where they spin cocoons in the
duff and pupate. Adults emerge in late summer and lay eggs individually
in needles. There is one generation per year. Trees attacked by this sawfly
are not killed, but tree growth in forests is reduced. A large outbreak
of this species (5.6 million acres) occurred in the 1950s in Maryland,
Virginia, and North Carolina. Note, this species should not be confused
with Neodiprion pratti (Dyar), a species associated with sand pine
(P. clausa [Chapm. ex Engelm.] Sarg.).

Hosts Commonly Attacked Virginia pine sawfly feeds on Virginia (P. virginiana Mill.) and shortleaf
(P. echinata Mill.) pines.

Distribution Virginia pine sawfly is found from New Jersey south to North Carolina
and west to Illinois.

Images of Virginia
Pine Sawfly

Figure 1. Adult male of Virginia pine sawfly, Neodiprion pratti pratti. (USDA Forest
Service - Region 8 - Southern Archive, USDA Forest Service, Bugwood.org)
A Photographic Guide 649

Images of Virginia
Pine Sawfly (continued)

Figure 2. Egg scars of Virginia pine sawfly on pine needles. (Caleb L. Morris,
Virginia Department of Forestry, Bugwood.org)

Figure 3. Larvae of Virginia pine

sawfly. (USDA Forest Service
- Forest Health Protection,
Southern Region)

Figure 4. Feeding damage of Virginia pine sawfly. (USDA Forest Service -

Ashville Archive, USDA Forest Service, Bugwood.org)
650 Forest Pest Insects in North America:

Images of Virginia
Pine Sawfly (continued)

Figure 5. Pine stand defoliated by Virginia pine sawfly. (Caleb L. Morris, Virginia
Department of Forestry, Bugwood.org)

Important Biological Control No information was found on the natural enemies of this species.
Agents Related to this
Pest Species
Web Links for Information http://www.fs.fed.us/r8/foresthealth/idotis/insects/vapinesf.html; USDA
on Virginia Pine Sawfly Forest Service fact sheet on biology and control.
http://pubs.ext.vt.edu/2911/2911-1424/2911-1424.html; Virginia
Extension Service fact sheet on biology and control.
http://entnemdept.ufl.edu/creatures/trees/sawfly/pine_sawflies.htm;
fact sheet of the University of Florida; compares some details among
a number of sawfly species that feed on pine, including Virginia pine
sawfly.
http://www.forestpests.org/acrobat/not%20linked/gfc-01.pdf; fact sheet
of the Georgia Forestry Commission; provides drawings that compare
body markings of larvae of various sawflies that feed on pines.

Articles None
A Photographic Guide 651

180. Hemlock Sawfly, Neodiprion tsugae Middleton (Hymenoptera:

Diprionidae)

Orientation to Pest The hemlock sawfly, Neodiprion tsugae Middleton, is an important

native defoliator of western hemlock (Tsuga heterophylla [Raf.] Sarg.)
in western North America. This sawfly overwinters as eggs, which are
laid in the fall in needles of the current year’s growth, usually one egg
per needle. Eggs hatch in spring and the larvae feed into early summer.
Young larvae usually feed gregariously, but change to feeding singly
as they mature. The larvae eat the old needles, reducing growth but
rarely killing trees. Mature larvae drop to the duff where they spin
cocoons. Usually there is one generation a year, but some larvae take
more than one year to complete their life cycle. Extensive outbreaks
on western hemlock have been recorded in natural forests in Oregon,
British Columbia, and Alaska, often in association with outbreaks of
blackheaded budworm (Acleris gloverana Walsingham). In coastal
Alaska, defoliation can be concentrated on large trees on southern and
western slopes near waterways. In British Columbia when hemlock
sawfly is associated with blackheaded budworm, thinned stands have
been reported to suffer heavier defoliation than unthinned stands.

Hosts Commonly Attacked The main host is western hemlock (Tsuga heterophylla [Raf.] Sarg.), but
in addition, mountain hemlock (T. mertensiana [Bong.] Carr). and Pacific
silver fir (Abies amabilis Douglas ex J. Forbes) are sometimes attacked
as well.

Distribution This sawfly is found in coastal forests

of western North America in Oregon,
Washington, British Columbia, and
Alaska. It also occurs in interior
forests of British Columbia, Idaho,
and Montana.

Figure 1. North
American distribution
of hemlock sawfly,
Neodiprion tsugae.
(USDA Forest Service
• Forest Insect and
Disease Leaflet 31)
652 Forest Pest Insects in North America:

Images of
Hemlock Sawfly

Figure 2. Adult of hemlock sawfly. (Edward H. Holsten, USDA Forest Service,

Bugwood.org)

Figure 3. Larvae of
hemlock sawfly. (Top:
USDA Forest Service
- Region 10 - Alaska
Archive, USDA Forest
Service, Bugwood.
org; bottom: Dion
Manastyrski - Natural
Resources Canada,
Canadian Forest
Service)

Figure 4. Cocoon of
hemlock sawfly, from
which the adult has
emerged. (USDA Forest
Service Insect and
Disease Leaflet 31)
A Photographic Guide 653

Images of Hemlock Figure 5. Feeding

damage (thin foliage)
Sawfly (continued) of hemlock sawfly on
western hemlock. (David
L. Overhulser, Oregon
Department of Forestry,
Bugwood.org)

Figure 6. Stand of hemlock showing extensive damage during outbreak of

hemlock sawfly. (USDA Forest Service - Region 10 - Alaska Archive, USDA
Forest Service, Bugwood.org)
654 Forest Pest Insects in North America:

Important Biological Control Parasitoids attacking hemlock sawfly include the ichneumonids
Agents Related to this Delomerista japonica diprionis Cushman, Oresbius tsugae tsugae
Pest Species (Cushman), and Itoplectis quadricingulatus (Provancher). A fungus
(Entomopthora sphaerosperma Fres.) was the most effective control
observed in studies in Alaska, particularly in wet years.

Web Links for Information http://www.fs.fed.us/r10/spf/fhp/leaflets/Fidl31.htm; USDA Forest

on Hemlock Sawfly Service Insect and Disease Leaflet 31 on biology and control of
hemlock sawfly.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=1000010;
fact sheet on this insect by Natural Resources Canada.

Articles Furniss, R. L. and P.B. Dowden. 1941. Western hemlock sawfly,

Neodiprion tsugae Middleton, and its parasites in Oregon. Journal of
Economic Entomology 34: 46-52.
Hard, J.S. 1976. Natural control of hemlock sawfly, Neodiprion tsugae
Middleton (Hymenoptera: Diprionidae), populations in southeast Alaska.
The Canadian Entomologist 108: 485-498.
Hard, J.S. and D.C. Schmiege. 1968. The hemlock sawfly in southeast
Alaska. USDA Forest Service Research Paper No. PNW-65. Pacific
Northwest Forestry and Range Exp. Station, Portland, Oregon: 11 p.
Torgersen, T.R. 1968. Parasites of the hemlock sawfly, Neodiprion
tsugae, in coastal Alaska. Annals of the Entomological Society of
America 61(5): 1155-1158.
A Photographic Guide 655

181. Introduced Pine Sawfly, Diprion similis (Hartig) (Hymenoptera:

Diprionidae)

Orientation to Pest Introduced pine sawfly, Diprion similis (Hartig), is an invasive species in
North America of Eurasia origin that defoliates eastern white pine (Pinus
strobus L.) and some other pines. It occurs widely in eastern Canada and
the United States. Winter is spent in the prepupal stage in a cocoon on
the tree or in the duff. Insects pupate in early spring and adults emerge
by May to mid-June. Females lay eggs in rows of up to 10 in slits cut
in needles of hosts. Young larvae feed in groups, but older larvae feed
alone. Larvae of the first generation feed on old needles only, but larvae
of the second generation eat both old and new needles. Normally there
are two generations per year, but in favorable areas, there may be a
partial third generation. Infested trees may be defoliated and trees
may be killed.

Hosts Commonly Attacked This species feeds on pines, especially eastern white pine (P. strobus).
Scotch (P. sylvestris L.), red (P. resinosa Sol. ex Aiton), jack
(P. banksiana Lamb.), and mugo (P. mugo Turra) pines are also attacked.

Distribution In the United States, this sawfly is found from Maine west to Minnesota
and south to North Carolina, Tennessee, and Virginia. In Canada, it is
present in Newfoundland, Nova Scotia, New Brunswick, Prince Edward
Island, Quebec, Ontario, and Manitoba.

Images of Introduced
Pine Sawfly

Figure 1. Adult of introduced pine sawfly, Diprion simili. (John H. Ghent, USDA
Forest Service, Bugwood.org)
656 Forest Pest Insects in North America:

Images of Introduced
Pine Sawfly (continued)

Figure 2. Eggs of introduced pine sawfly inserted into a pine needle. (John H.
Ghent, USDA Forest Service, Bugwood.org)

Figure 3. Larvae of introduced pine sawfly. (Steven Katovich, USDA Forest

Service, Bugwood.org)

Figure 4. Cocoon
(emerged) of introduced
pine sawfly. (Gyorgy
Csoka, Hungary Forest
Research Institute,
Bugwood.org)
A Photographic Guide 657

Images of Introduced
Pine Sawfly (continued)

Figure 5. Defoliation of eastern white pines due to introduced pine sawfly larvae.
(John H. Ghent, USDA Forest Service, Bugwood.org)

Figure 6. Emergence of the ichneumonid parasitoid Exenterus amictorius

(Panzer) from a cocoon of introduced pine sawfly. (John H. Ghent, USDA Forest
Service, Bugwood.org)
658 Forest Pest Insects in North America:

Important Biological Control In Europe, many species of parasitoids have been reared from D.
Agents Related to this similis, including most commonly Aptesis subguttatus (Grav.), A.
Pest Species basizonia (Grav.), Exenterus amictorius (Panzer), E. adspersus Hartig.,
Palexorista inconspicua (Meigen), Hypsantyx impressus (Grav.),
and Monodontomerus dentipes (Dalman). Palexorista inconspicua is
believed by some to be the most important tachinid parasitoid of D.
similis in Europe. In North America, D. similis is attacked by several
European parasitoids that were introduced against other closely related
invasive sawflies. In Connecticut, nearly 50 percent of D. similis larvae
were parasitized by a complex of eight parasitoid species, especially
M. dentipes, which was also one of the most important parasitoids
in Pennsylvania and New Jersey. In Minnesota, the most common
parasitoids were E. amictorius, M. dentipes, and Delomerista japonica
Cushman. Diprion similis did not invade the southern United States
until the 1970s, and, in that region, few parasitoid species were found
attacking the pest. Monodontomerus dentipes was introduced to the
region, where it became abundant and was credited with controlling the
pest in North Carolina.

Web Links for Information http://www.exoticpests.gc.ca/ins_details_eng.asp?pestType=ins&lang=lt

on Introduced Pine Sawfly &geID=6552; fact sheet of Environment Canada on biology of species.
http://www.forestpests.org/vermont/introducedsawfly.html; fact sheet
of the Vermont Department of Forests, Parks and Recreation on biology
and control.
http://na.fs.fed.us/spfo/pubs/fidls/intro_sawfly/intro_sawfly.htm;
USDA Forest Service Forest Insect and Disease Leaflet.

Articles Monroe, H.A.U. 1935. Observations on the habits of an introduced pine

sawfly Diprion simile Htg. The Canadian Entomologist 67: 137-140.
Tsao, C.H. and A.C. Hodson. 1956. The effect of different host species
on the oviposition and survival of the introduced pine sawfly. Journal of
Economic Entomology 49: 400-401.
Coppel, H.C., J.W. Mertins, and J.W.E. Harris. 1974. The introduced
pine sawfly, Diprion similis (Hartig) (Hymenoptera: Diprionidae):
A review with emphasis on studies in Wisconsin. University of
Wisconsin-Madison Research Bulletin R 2393: 74 p.
Drooz, A.T.; J.H. Ghent, and C.M. Huber. 1985. Insect parasites
associated with the introduced pine sawfly, Diprion similis (Hartig)
(Hymenoptera: Diprionidae), in North Carolina. Environmental
Entomology 14: 401-403.
A Photographic Guide 659

182. European Spruce Sawfly, Gilpinia hercyniae (Hartig) (Hymenoptera:

Diprionidae)

Orientation to Pest The European spruce sawfly, Gilpinia hercyniae (Hartig), is an

introduced sawfly from Europe that feeds on spruce (Picea). Currently,
this sawfly is found in Canada from the Maritimes west to Manitoba,
and on the island of Newfoundland. In the United States, it is present
from the northeast to the Great Lake states. White spruce (Picea glauca
[Moench] Voss) and other species in the genus are attacked. Adults
emerge in spring and lay eggs in slits cut in needles of host trees. Larvae
of all ages feed singly; old needles are preferred, but both old and new
needles are consumed. Mature larvae spin cocoons in which they pupate.
For the overwintering generation, cocoons are formed in the duff below
host trees, while non-overwintering cocoons are attached to foliage or
twigs. Depending on the location, there may be one to three generations
per year. About a decade or so after the detection of this species in
North America, outbreaks became widespread and continuous in eastern
Canada. In the 1940s, biological control of this species was achieved by
introducing a series of European parasitoids and, accidentally, a nuclear
polyhedrosis virus. Outbreaks of this species no longer occur in North
America due to the combined action of these natural enemies.

Hosts Commonly Attacked White spruce (P. glauca [Moench] Voss) is the preferred host, but
red (P. rubens Sarg.), black (P. mariana [Mill.] Britton, Sterns &
Poggenburg), and Norway (P. abies [L.] H. Karst.) spruce are attacked.

Distribution This sawfly is found from the Maritimes west to Manitoba, and on
the island of Newfoundland. In the United States, it is present in the
northeastern States, west to the Great Lakes region.

Images of European
Spruce Sawfly

Figure 1. Adult of the European spruce sawfly, Gilpinia hercyniae. (Gerhard

Elsner, Biologische Bundesanstalt für Land und Forstwirtschaft, Bugwood.org)
660 Forest Pest Insects in North America:

Images of European Figure 2. Eggs (seen

as swollen areas, see
Spruce Sawfly (continued) arrow) inside spruce
needles.
(Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Figure 3. Larvae of the European spruce sawfly. (Gerhard Elsner, Biologische

Bundesanstalt für Land und Forstwirtschaft, Bugwood.org)

Figure 4. Cocoons of a non-diapausing generation of the European spruce

sawfly. (Gerhard Elsner, Biologische Bundesanstalt für Land und Forstwirtschaft,
Bugwood.org)
A Photographic Guide 661

Important Biological Control Importations of parasitoids from Europe against the European spruce
Agents Related to this sawfly were begun after detection of a large outbreak in the Gaspé
Pest Species region of Quebec, and natural enemy introductions were carried out on
a massive scale. Millions of sawfly cocoons and eggs were collected in
Europe and shipped to Canada where their parasitoids were reared and
released. Eventually a rearing program was established in Canada that
produced over 800 million parasitoids between 1932 and 1958. Overall
some 30 species of parasitoids were identified from Europe and Japan
and introduced into North America. At least seven species established, of
which the first was Dahlbominus fuscipennis Zett. in Quebec, followed
by several species of Exenterus. Biological control was eventually
achieved not by parasitoids but by the accidental introduction of a
nucleopolyhydrosis virus (Borrelinavirus hercyniae) sometime before
1938. By 1943, G. hercyniae populations were in marked decline with
B. hercyniae identified as the most important cause. This virus, together
with the introduced parasitoids, caused a widespread reduction of
G. hercyniae of 90-95 percent by 1945. Once outbreaks had ceased
G. hercyinae populations were regulated by the combination of
B. hercyniae and two introduced parasitoids Drino bohemica (Mesn.)
and Exenterus vellicatus Cush. Minor outbreaks continued to occur for
some years, on the edge of the species’ range and in forests that had
been sprayed with DDT to control populations of the spruce budworm
(Choristoneura fumiferana Clem.); areas where the pest had temporarily
outpaced the expansion of the ranges of the introduced natural enemies
or where natural enemies had been killed by pesticide use.

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=6555;

on European Spruce Sawfly fact sheet of Natural Resources Canada.

Articles Clark, R.C., L.J. Clarke, and K.E. Pardy. 1973. Biological control of the
European spruce sawfly in Newfoundland. Bi-monthly Research Notes
29(1): 2-3.
Adams, P.H.W. and P.F. Entwistle. 1981. An annotated bibliography of
Gilpinia hercyniae (Hartig), European spruce sawfly. Occasional Papers
No. 11. Commonwealth Forestry Institute: 58 p.
McCugan, B.M. and H.C. Coppell. 1962. A review of biological control
attempts against insects and weeds in Canada. Part II. Biological
control of forest insects, 1910-1958. Technical Communication No. 2,
Commonwealth Institute of Biological Control, Trinidad. Commonwealth
Agriculture Bureaux, Farnahm Royal, Bucks, England: 35-177.
662 Forest Pest Insects in North America:

183. Larch Sawfly, Pristiphora erichsonii (Hartig) (Hymenoptera:

Tenthridinidae)

Orientation to Pest Larch sawfly, Pristiphora erichsonii (Hartig), was historically a serious
pest of larch (Larix) in North America. It is present throughout Canada,
Alaska, and most of the northern tier of US states. Distinct strains of
the species have been recognized, with different geographical origins.
A debate has existed over whether strains of this species found in North
American before 1920 were native to North America or were invasive
from Europe at an earlier date. Further complicating matters is the
accidental introduction of additional strains, definitely from Europe,
that occurred during a biological control program (1910-1930) against
the species that released cocoons (both parasitized and unparasitized)
collected in Europe in the field in Canada. This sawfly overwinters as
prepupae in cocoons in the duff. Adults emerge in spring and lay eggs in
new shoots, causing curling. All larch sawfly adults are females. Newly
hatched larvae feed in groups, mainly on the needle clusters of older
twigs. Dense populations can defoliate whole trees. Mature larvae drop
to the ground to spin cocoons. There is one generation per year in most
locations, but some individuals may remain in their cocoons for two
or three winters before emerging. Damaging outbreaks have occurred
periodically. During outbreaks, trees are not commonly killed, but wood
growth is reduced. Weakened surviving trees may also be at increased
risk of bark beetle attack.

Hosts Commonly Attacked Larch sawfly feeds on eastern larch (Larix laricina [Du Roi] K. Koch),
western larch (L. occidentalis Nutthal), and alpine larch (L. lyallii Parl.).

Distribution Larch sawfly occurs in all Canadian provinces, Alaska, all the northern
tier of U.S. states, as well as Maryland, North Carolina, and West
Virginia.

Images of Figure 1. Adult of larch

sawfly, Pristiphora
Larch Sawfly erichsonii. (Natural
Resources Canada,
Canadian Forest
Service)
A Photographic Guide 663

Images of
Larch Sawfly (continued)

Figure 2. Larvae of larch sawfly. (Steven Katovich, USDA Forest Service,

Bugwood.org)

Figure 3. Cocoons of larch sawfly, showing adult emergence holes. (Arnold T.

Drooz, USDA Forest Service, Bugwood.org)

Figure 4. Larch needles showing feeding by larch sawfly larvae. (Dayle D.

Bennett, Bugwood.org)
664 Forest Pest Insects in North America:

Images of
Larch Sawfly (continued)

Figure 5. A larch stand in Minnesota showing partial (50 percent) defoliation

caused by larch sawfly. (Arnold T. Drooz, USDA Forest Service, Bugwood.org)

Figure 6. Landscape level defoliation of larch in Minnesota in 1953 caused by

larch sawfly. (Arnold T. Drooz, USDA Forest Service, Bugwood.org)

Important Biological Control The history of biological control efforts against larch sawfly in North
Agents Related to this America is complex. See Van Driesche et al. (1996) for a detailed history.
Pest Species Natural enemies in North America that are important to the population
dynamics of larch sawfly include a tachinid (Bessa harveyi [Townsend]),
a guild of native small mammals, and two introduced ichneumonid
parasitoids, Mesoleius tenthredinis Morely and Olesicampe benefactor
Hinz. An important factor mediating outcomes in early stages of the
biological control program was variation in the ability of the several
sawfly strains to encapsulate the eggs of the parasitoid M. tenthredinis.
This was subsequently resolved by introduction of a second strain that
was able to resist encapsulation.
A Photographic Guide 665

Web Links for Information http://www.forestry.ubc.ca/fetch21/FRST308/lab5/pristiphora_erichsonii/

on Larch Sawfly sawfly.html; fact sheet of the University of British Columbia on biology.

Articles Turnock, W.J. 1960. Ecological life-history of the larch sawfly,

Pristiphora erichsonii (Htg.) (Hymenoptera: Tenthredinidae), in
Manitoba and Saskatchewan. The Canadian Entomologist 92: 500-516.
Hu, F. 1964. Population dynamics of the larch sawfly. The Canadian
Entomologist 96: 160-161.
Turnock, W.J. 1972. Geographical and historical variability in
population patterns and life systems of the larch sawfly (Hymenoptera:
Tenthredinidae). The Canadian Entomologist 104: 1883-1900.
Wong, H.R. 1974. The identification and origin of the strains of the larch
sawfly, Pristiphora erichsonii (Hymenoptera: Tenthredinidae), in North
America. The Canadian Entomologist 106: 1121-1131.
Maltais, P.P., J.J. Juillet, and D.D. de Oliveira. 1980. Ecology and
population dynamics of the larch sawfly, Pristiphora erichsonii (Htg.)
in Kouchibouguac National Park, New Brunswick. Annals of the
Entomological Society of Quebec 25: 141-162.
Krause, S.L. and K.F. Raffa. 1996. Defoliation tolerance affects the
spatial and temporal distributions of larch sawfly and natural enemy
populations. Ecological Entomology 21: 101-111.
Van Driesche, R.G., S. Healy, and R.C. Reardon. 1996. Biological
Control of Arthropod Pests of the Northeastern and North Central
Forests in the United States: A review and recommendations. FHTET
96-19, December 1996, USDA Forest Service, Morgantown, WV.
(See http://www.forestpestbiocontrol.info/fact_sheets/documents/
arthropodpestsnortheastern_northcentral.pdf).
666 Forest Pest Insects in North America:

184. Mountain Ash Sawfly, Pristiphora geniculata (Hartig) (Hymenoptera:

Tenthridinidae)

Orientation to Pest Mountain ash sawfly, Pristiphora geniculata (Hartig), is a European

tenthredinid sawfly that is invasive in North America. In North America,
it feeds on species of Sorbus. Mountain ash sawfly occurs in Canada
east of central Ontario and in the northeastern United States. The winter
is passed as a prepupa in a cocoon in the soil or leaf litter. Pupation
occurs in the spring and adults emerge soon after. Eggs are deposited
in slits cut in the edges of the leaves of the host trees. Newly hatched
larvae feed on the leaf edges. When disturbed, larvae raise their bodies
in an “s” position. Young larvae feed gregariously, but older larvae feed
individually. Larvae eat entire leaves except for the midribs. Mature
larvae of the overwintering generation drop to the soil and spin their
cocoons. Mountain ash sawfly is primarily a pest of ornamentals and
in the wild has largely been suppressed by the introduced parasitoid
Olesicampe geniculatae Quednau and Lim. In all but the northern part
of its range in North America, there are two generations per year.

Hosts Commonly Attacked This species feeds on American mountain ash (Sorbus americana
Marshall) and European mountain ash (S. aucuparia L.).

Distribution Mountain ash sawfly occurs in Canada east of central Ontario and in
the northeastern United States, including New York, New Jersey, and
Michigan.

Images of Figure 1. Adult of

mountain ash sawfly,
Mountain Ash Sawfly Pristiphora geniculata.
(James Lindsey,
Bugwood.org)

Figure 2. Eggs of
mountain ash sawfly,
inserted into edges of
leaves of the host plant.
(L.J. Lipovsky, Maine
Forest Service)
A Photographic Guide 667

Images of Mountain Ash Figure 3. Larvae of

mountain ash sawfly;
Sawfly (continued) young larvae feeding in
a group (top) and older
larvae feeding alone
(bottom). (Top: Steven
Katovich, USDA Forest
Service, Bugwood.org;
bottom: E. Bradford
Walker, Vermont
Department of Forests,
Parks and Recreation,
Bugwood.org)

Figure 4. Foliage of mountain ash stripped of all but mid ribs by mountain ash
sawfly larvae. (Joseph O’Brien, USDA Forest Service, Bugwood.org)

Important Biological Control This introduced sawfly has been controlled by releases of the parasitoid
Agents Related to this Olesicampe geniculatae.
Pest Species
668 Forest Pest Insects in North America:

Web Links for Information http://www.forestpests.org/vermont/mountainashsawfly.html; fact sheet

on Mountain Ash Sawfly of the Bugwood network.
http://maine.gov/doc/mfs/mtnash.htm; fact sheet of the Maine
Department of Conservation on biology and control of insect.
http://www.fs.fed.us/r6/nr/fid/invasives/mtn-ash-sawfly.shtml; fact sheet
of USDA Forest Service Forest Health Protection Pacific Northwest
Region—biology and control.

Articles Forbes, R.S. and L. Daviault.1964. The biology of the mountain ash
sawfly, Pristiphora geniculata (Htg.) (Hymenoptera: Tenthredinidae), in
eastern Canada. The Canadian Entomologist 98: 1117-1133.
Quednau, F.W. 1990. Introduction, permanent establishment, and
dispersal in eastern Canada of Olesicampe geniculatae Quednau and Lim
(Hymenoptera: Ichneumonidae), an important biological control agent of
the mountain ash sawfly, Pristiphora geniculata (Hartig) (Hymenopttera:
Tenthredinidae). The Canadian Entomologist 122: 921-934.
West, R.J., P.L. Dixon, F.W. Quednau, K.P. Lim, and K. Hiscock.
1994. Establishment of Olesicampe geniculatae Quednau and Lim
(Hymenoptera: Ichneumonidae) to control the mountain ash sawfly,
Pristiphora geniculata (Hartig) (Hymenopttera: Tenthredinidae) in
Newfoundland. The Canadian Entomologist 126: 7-11.
West, R.J., P.L. Dixon, F.W. Quednau, and K.P. Lim. 2001.
Pristiphora geniculata (Hartig), mountain ash sawfly (Hymenoptera:
Tenthredinidae). In: Mason, P.G. and J.T. Huber (eds.). Biological
Control Programmes in Canada, 1981-2000. CABI Publishing,
Wallingford, UK: 228-230.
A Photographic Guide 669

185. Yellowheaded Spruce Sawfly, Pikonema alaskensis (Rohwer)

(Hymenoptera: Tenthredinidae)

Orientation to Pest The yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer), is a

native sawfly found throughout northeastern and northwestern North
America that feeds on all species of native and introduced spruce (Picea).
It overwinters as prepupae in the duff and adults emerge in spring. Eggs
are laid in the current season’s needles just as buds begin to flush in
the spring. Larvae feed on the new needles primarily, but may exploit
old needles if new needles become scarce. In summer, fully developed
larvae drop to the ground and spin cocoons (in which they molt to the
prepupal stage) in the duff or topsoil. Damage occurs mainly in the upper
crown of young, open-grown spruce, especially on trees in ornamental
and shelterbelt plantings; mature trees are rarely attacked. Several years
of heavy feeding may kill the leader and upper crown branches of trees.
There is one generation per year.

Hosts Commonly Attacked Hosts of yellowheaded spruce sawfly include all native spruces in North
America: Engelmann (Picea engelmannii Parry ex Engelm.), white
(P. glauca [Moench] Voss), black (P. mariana [Mill.] Britton, Sterns &
Poggenburg), Sitka (P. sitchensis [Bong.] Carr.), and Colorado blue
(P. pungens Engelm.), and the introduced Norway spruce (P. abies [L.]
Karst.).

Distribution This sawfly occurs in throughout the range of spruce in North America,
and can be found from Alaska to Maine.

Images of Yellowheaded Figure 1. Adult of

yellowheaded spruce
Spruce Sawfly sawfly, Pikonema
alaskensis.
(D.K.B. Cheung)
670 Forest Pest Insects in North America:

Images of Yellowheaded Figure 2. Ovipostion

cuts in needles made
Spruce Sawfly (continued) by yellowheaded spruce
sawfly. (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Figure 3. Larvae of
yellowheaded spruce
sawfly; top, young
larvae and bottom,
mature larvae. (Top:
Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service; bottom:
E. Bradford Walker,
Vermont Department of
Forests, Parks
and Recreation,
Bugwood.org)

Figure 4. Prepupa of
yellowheaded spruce
sawfly in cocoon
(opened).
(D.K.B. Cheung)
A Photographic Guide 671

Images of Yellowheaded Figure 5. Damage

(top) to white spruce
Spruce Sawfly (continued) from yellowheaded
spruce sawfly; close
up (bottom) of larvae
feeding on spruce.
(Top: Steven Katovich,
USDA Forest Service,
Bugwood.org; bottom:
E. Bradford Walker,
Vermont Department
of Forests, Parks
and Recreation,
Bugwood.org)

Figure 6. Stand of
spruce showing injury
from yellowheaded
spruce sawfly. (USDA
Forest Service -
Northeastern Area
Archive, USDA Forest
Service, Bugwood.org)
672 Forest Pest Insects in North America:

Important Biological Control The natural enemies of this species have been studied in Minnesota,
Agents Related to this Maine and Nova Scotia. A monitoring trap has been developed for one
Pest Species parasitoid, the ichneumonid Syndipnus rubiginosus Walley.

Web Links for Information on http://www.na.fs.fed.us/spfo/pubs/gtr/sprucesawfly/biology.htm; a very

Yellowheaded Spruce Sawfly detailed fact sheet of the USDA Forest Service on biology of this species.
http://www.fs.fed.us/r10/spf/fhp/leaflets/yellowhd_spruce_sawfly.pdf;
fact sheet of the USDA Forest Service on biology and control.
http://gardenline.usask.ca/pests/yellow.html; fact sheet of the University
of Saskatchewan on biology and control.
http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=7875;
fact sheet of the Natural Resources Canada on biology and phenology.
http://www1.agric.gov.ab.ca/$department/deptdocs.nsf/all/opp3005; fact
sheet of the Alberta Department of Agriculture, on biology and control.
http://www.entomology.umn.edu/cues/Web/225YellowheadedSpruce
Sawfly.pdf; fact sheet of the University of Minnesota on biology and
control.
http://www.maine.gov/doc/mfs/yhss.htm; fact sheet of the Maine
Department of Conservation on biology and control.
http://www.forestpests.org/vermont/yellowheadsawfly.html; fact sheet
of the Vermont Department of Forests, Parks, and Recreation.
http://www.fs.fed.us/r6/nr/fid/fidls/fidl-69.pdf; USDA Forest Service
Forest Pest Leaflet No. 69.

Articles Thompson, L.C. and H.M. Kulman. 1980. Parasites of the yellowhead
spruce sawfly, Pikonema alaskensis (Hymenoptera: Tenthredinidae), in
Maine and Nova Scotia. The Canadian Entomologist 112: 25-29.
Katovich, S.A., D.G. McCullough, and R.A. Haack. 1995. Yellowheaded
spruce sawfly: its ecology and management. General Technical Report
No. NC-179. USDA Forest Service. Michigan.
Thurston, G.S. 2001. Pikonema alaskensis (Rohwer), yellowheaded
spruce sawfly (Hymenoptera: Tenthredinidae). In: Mason, P.G. and J.T.
Huber (eds.). Biological Control Programmes in Canada, 1981-2000.
CABI Publishing, Wallingford, UK: 219-221.
Johns, R.C., D.P. Ostaff, and D.T. Quiring. 2006. Sampling procedures
for evaluating yellowheaded spruce sawfly density and defoliation in
juvenile black spruce stands. Journal of the Acadian Entomological
Society 2: 1-12.
Johns, R.C., D.P. Ostaff, and D.T. Quiring. 2006. Relationships between
yellowheaded spruce sawfly density and defoliation on juvenile black
spruce. Forest Ecology and Management 228: 51-60.
A Photographic Guide 673

186. Birch Leafminer, Fenusa pumila (Leach) (Hymenoptera: Tenthridinidae)

Orientation to Pest The birch leafminer, Fenusa pumila (Leach) (formerly known as Fenusa
pusilla [Lepeletier]), is a European sawfly that was mines the leaves
of several species of birch (Betula) in North America, where it invaded
just prior to the 1920s. It is found principally in southeastern Canada
and the northeastern United States, in the range of its main hosts, gray
(Betula populifolia Marshal) and paper birch (B. papyrifera Marshal).
This sawfly overwinters as prepupae in earthen cells in the soil. Pupation
occurs in spring and adults emerge near bud break. Eggs are laid singly
in unfolding leaves, and larvae create leafmines where they feed. Mines
are blotch shaped and enlarge and eventually merge as larvae grow. In
New England, there is only one full generation per year, but a small
fraction of the first and second generations develop to adults without
diapause and emerge to produce a partial second and third generations.
Before the 1980s, birch leafminer densities were high enough so that
most leaves of white and gray birches in landscapes or road edges
were mined; mined leaves being completely browned. This created an
aesthetic problem on landscape trees and likely reduced the vigor of
forest trees. In the period of 1970-2000, birch leafminer was subjected to
biological control through introduction of its specialized parasitoids from
Europe and densities in the northeastern United States have now declined
to low pest levels (<5 percent leaves mined).

Hosts Commonly Attacked The principal birch species used as hosts are gray (B. populifolia) and
paper (B. papyrifera) birch. Yellow (B. alleghaniensis Britton) and
black (=sweet) (B. lenta L.) birch are not attacked. Several introduced
European birches are also attacked in landscape plantings.

Distribution This sawfly occurs from Newfoundland (Canada) south to Maryland and
west to Minnesota and Iowa, with isolated populations also in Oregon,
Washington, and Alaska, especially on introduced birches planted as
ornamentals.

Images of
Birch Leafminer

Figure 1. Adults of birch leafminer, Fenusa pumila; left, adult on young birch leaf;
right, close up. (Left: Steven Katovich, USDA Forest Service, Bugwood.org;
right: Cheryl Moorehead, individual, Bugwood.org)
674 Forest Pest Insects in North America:

Images of Birch Figure 2. Eggs of birch

leafminer (points),
Leafminer (continued) surrounded by yellow
halos of tissue. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)

Figure 3. Small mines of birch leafminer, showing young larvae feeding in

separate mines (top); bottom, close up on a single mine, with larva visible. (Top:
Steven Katovich, USDA Forest Service, Bugwood.org; bottom: Joseph O’Brien,
USDA Forest Service, Bugwood.org)
A Photographic Guide 675

Images of Birch
Leafminer (continued)

Figure 4. Middle-age (top) and older (bottom) larvae of birch leafminer, showing
fusion of mines as larvae increase in size. (Top: Brian Kunkel, University of
Delaware, Bugwood.org; bottom: E. Bradford Walker, Vermont Department of
Forests, Parks and Recreation, Bugwood.org)

Figure 5. Close up of damage on branch heavily mined by birch leafminer.

(Whitney Cranshaw, Colorado State University, Bugwood.org)
676 Forest Pest Insects in North America:

Images of Birch Figure 6. High density

of mining by birch
Leafminer (continued) leafminer. (Steven
Katovich, USDA Forest
Service, Bugwood.org)

Important Biological Control Two European ichneumonids (Lathrolestes nigricollis [Thompson] and
Agents Related to this Grypocentrus albipes Ruthe) and one eulophid (Chrysocharis nitetis
Pest Species [Wilkinson]) were introduced into North America for biological control
of the birch leafminer. Lathrolestes nigricollis is the only species to
have become widely established. At the level of individual release sites,
L. nigricollis has lowered pest density from 50 to <5 percent of first
generation leaves being mined. At the landscape level, this parasitoid
has suppressed this leafminer to insignificant levels throughout the
northeastern United States, with the exception of southern New Jersey.

Web Links for Information http://njaes.rutgers.edu/pubs/publication.asp?pid=FS1164; fact sheet of

on Birch Leafminer Rutgers University (New Jersey) on biology and control.
http://www.extension.umn.edu/distribution/horticulture/DG6134.html;
fact sheet of University of Minnesota Extension on biology and control.
http://www.invasiveforestinsectandweedbiocontrol.info/target_pests/
insects_mites/Birchleafminer.htm; fact sheet of the University of
Massachusetts on biological control of birch leafminer.
A Photographic Guide 677

Articles Cheng, H.H. and E.J. LeRoux. 1965. Preliminary life tables and notes
on mortality factors of the birch leaf miner, Fenusa pusilla (Lepeletier)
(Hymenoptera: Tenthredinidae), on blue birch, Betula caerulea grandis
Blanchard, in Quebec. Annals of the Entomological Society of Quebec
11: 81-104.
Eichhorn, O. And H. Pschorn-Walcher. 1973. The parasites of the birch
leaf-mining sawfly (Fenusa pusilla (Lep.), Hym.: Tenthredinidae) in
Central Europe. Technical Bulletin of the Commonwealth Institute of
Biological Control No. 16: 79-104.
Fuester, R.W., P.B. Taylor, W.H. Day, R.M. Hendrickson, Jr., and E.M.
Blumenthal. 1984. Introduction of exotic parasites for biological control
of the birch leafminer (Hymenoptera: Tenthredinidae) in the Middle
Atlantic states. Journal of Economic Entomology 77: 1565-1570.
Van Driesche, R.G., R. Childs, R.A. Casagrande, and L. Tewksbury.
1997. Establishment, distribution, and impact in southern New England
of Lathrolestes nigricollis (Thompson) (Hymenoptera: Ichneumonidae),
an introduced parasitoid of the birch leafminer, Fenusa pusilla
(Lepeletier) (Hymenoptera: Tenthredinidae). The Canadian Entomologist
129: 601-611.
Langor, D.W., S.C. Digweed, and J.R. Spence. 2001. Fenusa pusilla
(Lepeletier), birch leafminer, and Profenusa thomsoni (Konow),
ambermarked birch leafminer (Hymenoptera: Tenthredinidae). In:
Mason, P.G. and J.T. Huber (eds.). Biological Control Programmes in
Canada, 1981-2000. CABI Publishing, Wallingford, UK: 123-127.
Casagrande, R., R.G. Van Driesche, M. Mayer, R. Fuester, D. Gilrein,
L. Tewksbury, and H. Faubert. 2009. Biological control of Fenusa pusilla
(Hymenoptera: Tenthredinidae) in the northeastern United States: a
thirty-four year perspective on efficacy. Florida Entomologist 92:
243-247. (Available at http://www.fcla.edu/FlaEnt/fe92p243.pdf).
678 Forest Pest Insects in North America:

187. Ambermarked Birch Leafminer, Profenusa thomsoni (Konow)

(Hymenoptera: Tenthridinidae)

Orientation to Pest The ambermarked birch leafminer, Profenusa thomsoni (Konow), is

a European sawfly that feeds inside the leaves of several species of
birch (Betula). It invaded North America at the beginning of the 20th
century but was first reported in Canada in 1959. It is found both in
the northeastern United States (New England through the Great Lakes
region) and in eastern Canada from the Maritimes to Manitoba, where
it is not considered a pest because of its low densities. It also occurs in
western Canada and Alaska, where it reached high densities in Alberta
(1990s) and Alaska (2000-2008), especially in urban areas, before
being brought under biological control with an introduced parasitoid
and two native species. This sawfly overwinters as prepupae in earthen
cells in the soil. Pupation occurs in the spring, and adults emerge in
early summer. Eggs are inserted singly into tissue of mature leaves, and
larvae create leafmines where they feed. The blotch-shaped mines of this
species increase in size and several mines may merge as larvae grow.
In Alaska there is one generation per year; however, there are two in
Massachusetts.

Hosts Commonly Attacked The principal birch species used as hosts by ambermarked birch
leafminer are gray (Betula populifolia Marshal) and paper (B. papyrifera
Marshal) birch, but yellow birch (B. alleghaniensis Britt.) is also reported
as a host. Several introduced European birches are also attacked in
landscape plantings.

Distribution This sawfly occurs from the northeastern United States (New England to
the Great Lake States), eastern Canada from the Maritimes to Manitoba,
in western Canada (Alberta, Northwest Territories), and Alaska. Small
urban populations are found in additional locations (e.g., Bozeman,
Montana) and many of these may be unrecognized, being taken for
mines of Fenusa pumila Leach.

Images of Ambermarked Figure 1. Adult of

ambermarked birch
Birch Leafminer leafminer, Profenusa
thomsoni. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)
A Photographic Guide 679

Images of Ambermarked Figure 2. Eggs of

ambermarked birch
Birch Leafminer (continued) leafminer. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)

Figure 3. Mine of
nearly mature larva of
ambermarked birch
leafminer. (Thérèse
Arcand - Natural
Resources Canada,
Canadian Forest
Service)

Figure 4. Older instar

of ambermarked birch
leafminer, with mine
opened to reveal
larva. (Thérèse Arcand
- Natural Resources
Canada, Canadian
Forest Service)

Figure 5. Close up of
branch with mines of
ambermarked birch
leafminer. (René Pâquet
- Natural Resources
Canada, Canadian
Forest Service)
680 Forest Pest Insects in North America:

Images of Ambermarked Figure 6. Birch trees

(brown) in Alaska heavily
Birch Leafminer (continued) mined by ambermarked
birch leafminer. (Chris
MacQuarrie, University
of Alberta)

Figure 7. Important
parasitoids of
ambermarked birch
leafminer in Alaska:
Aptesis segnis
(top), Lathrolestes
soperi (middle), and
Lathrolestes thomsoni
(bottom). (All three
photos: Anna Soper,
Bugwood.org)
A Photographic Guide 681

Important Biological Control Three ichneumonids (Lathrolestes soperi Reshchikov, L. thompsoni

Agents Related to this Reshchikov, and Aptesis segnis [Provancher]) are important biological
Pest Species control agents of this leafminer. The two Lathrolestes species attack the
host larva in the mine. Lathrolestes soperi is known from Anchorage,
Alaska and is of unknown origin. This species is particularly common
in forested habitats. Lathrolestes thompsoni is a species that was
observed to provide control of this leafminer in Edmonton, Alberta, and
was moved to Anchorage, Alaska, where it was established to provide
biological control. Aptesis segnis is a facultative hyperparasitoid that
attacks the host (or parasitoid) in the host’s pupal cells in the soil.

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=7926;

on Ambermarked fact sheet of the Natural Resouces Canada, on biology and phenology.
Birch Leafminer
Articles Digweed, S.C., R.L. McQueen, J.R. Spence, and D.W. Langor. 2003.
Biological control of the ambermarked birch leafminer, Profenusa
thomsoni (Hymenoptera: Tenthredinidae), in Alberta. Information Report
NOR-X-389. Natural Resources Canada, Canadian Forest Service,
Northern Forestry Centre, Edmonton, Alberta: 8 p.
Digweed, S.C. and D.W. Langor. 2004. Distributions of leafmining
sawflies (Hymenoptera: Tenthredinidae) on birch and alder in
northwestern Canada. The Canadian Entomologist 136: 727-731.
Digweed, S.C., C.J.K. MacQuarrie, D.W. Langor, D.J.M. Williams,
J.R. Spence, K. Nystrom, and L. Morneau. 2009. Current status of
invasive alien birch-leafmining sawflies (Hymenoptera: Tenthredinidae)
in Canada, with keys to species. The Canadian Entomologist 141:
201-235.
Reshchikov, A.V., A. Soper and R.G. Van Driesche. 2010. Revision of the
Nearctic species of Lathrolestes (Hymenoptera: Ichneumonidae), with
special reference to species attacking Betula leaf mining Tenthredinidae
(Hymenoptera). Zootaxa 2614: 1-17.
682 Forest Pest Insects in North America:

188. Sirex Woodwasp, Sirex noctilio (Fabricius) (Hymenoptera: Siricidae)

Orientation to Pest Sirex woodwasp, Sirex noctilio (Fabricius), is a European woodwasp

that is widely invasive in the Southern Hemisphere. More recently it
has also become established in the United States and Canada, with the
initial discovery in New York in 2004. This species is a major threat to
pines. In contrast to native North American woodwasps that attack only
dead and dying trees, Sirex woodwasp can attack and kill living pines.
At low population densities, the wasp selects suppressed, stressed, or
injured trees for egg laying. Foliage of infested trees initially wilts and
turns red in 3-6 months following attack. Infested trees often have pearly
resin beads exuding from the drill holes where eggs were laid. Larval
galleries are tightly packed with fine white sawdust and adults chew
round exit holes that vary from 3 to 9 mm in diameter. Sirex noctilio is
expected to complete one generation per year throughout most of the
United States. Adult emergence occurs from July through September,
with peak emergence during August. When eggs are laid, the female
wasp also injects into the tree both a toxic mucus and a symbiotic fungus
(Amylostereum areolatum [Fr.] Boidin). The toxic mucus shuts down the
water transport, allowing the fungus to grow rapidly and kill the tree.
The fungus also provides nutrition for the developing woodwasp larvae.
Females can produce 25 to 450 eggs in their lifetime, depending upon
their size. The larval stage generally takes 10-11 months.

Hosts Commonly Attacked In addition to its preferred pine hosts (Pinus), S. noctilio can occasionally
use spruce (Picea), fir (Abies), larch (Larix), and Douglas fir
(Pseudotsuga menziesii [Mirb.] Franco).

Distribution Sirex noctilio is native to Eurasia and North Africa, but it has invaded the
Southern Hemisphere (New Zealand, Australia, Argentina, Chile, Brazil,
Uruguay, and South Africa). Most recently, it has also been introduced
to the United States (Michigan, New York, Ohio, Pennsylvania, and
Vermont) and Canada.

Figure 1. Worldwide
distribution of Sirex
noctilio, before its
invasion in North
America.
(Dennis Haugen,
Bugwood.org)
A Photographic Guide 683

Images of Figure 2. Mating pair

of Sirex woodwasps
Sirex Woodwasp (top: female left, male
right) and mounted
adult male (bottom).
(Top: Vicky Klasmer,
Instituto Nacional
de Tecnologia
Agropecuaria,
Bugwood.org;
bottom: Pest and
Diseases Image Library,
Bugwood.org)

Figure 3. Larvae of
Sirex noctilio in galleries
inside wood. (Top:
Vicky Klasmer, Instituto
Nacional de Tecnologia
Agropecuaria,
Bugwood.org; bottom:
Dennis Haugen,
Bugwood.org)
684 Forest Pest Insects in North America:

Images of Sirex
Woodwasp (continued)

Figure 4. Wood sawn from a tree damaged by Sirex noctilio. (Dennis Haugen,
Bugwood.org)

Figure 5. Cambial
staining caused by
the symbiotic fungus
(Amylostereum
areolatum) that is
transmitted by Sirex
noctilio. White plugs are
larval galleries packed
with excrement (frass).
(Vicky Klasmer,
Instituto Nacional
de Tecnologia
Agropecuaria,
Bugwood.org)
A Photographic Guide 685

Images of Sirex Figure 6. Emergence

holes of Sirex noctilio.
Woodwasp (continued) The round exit holes
vary from 3 to 9 mm
in diameter. (Dennis
Haugen, Bugwood.org)

Figure 7. Resin beads

from Sirex noctilio
oviposition drills in red
pine. (David Williams,
USDA APHIS,
Bugwood.org)
686 Forest Pest Insects in North America:

Images of Sirex Figure 8. Tree mortality

(80-90 percent) from a
Woodwasp (continued) Sirex noctilio outbreak in
a 15-year-old plantation
in South Australia in
1989. (Dennis Haugen,
Bugwood.org)

Figure 9. Life cycles

of the two forms of the
nematode Beddingia
siricidicola. (Dennis
Haugen, Bugwood.org)
A Photographic Guide 687

Images of Sirex
Woodwasp (continued)

Figure 10. Applying nematodes to a Sirex noctilio-infested tree for biological

control. Inoculation holes are made with a special punch hammer. (Dennis
Haugen, Bugwood.org)

Figure 11. Sirex noctilio parasitoids, Rhyssa persuasoria (left) and Rhyssa
lineolata (right). (Kevin Dodds, USDA Forest Service, Bugwood.org)
688 Forest Pest Insects in North America:

Important Biological Control Several parasitoids have been introduced from Europe and North
Agents Related to this America into New Zealand and Australia for biological control of Sirex
Pest Species woodwasps in pine plantations. The most effective species were Rhyssa
persuasoria (L.), Megarhyssa nortoni (Cresson), and Ibalia leucospoides
Hochenwald. However, the highest level of control was ultimately
provided in Australia by a parasitic nematode (Beddingia siricidicola
[Bedding]). This nematode invades the eggs of female woodwasps and is
transported by them to new egg laying sites. The growing nematode can
develop into either of two very different forms: one is a fungus-eating
form that lives on the symbiotic fungus transmitted by the woodwasp
and the other is a parasite that develops inside woodwasp larvae and
adults. Infected woodwasp larvae are not killed, but rather, the nematode
reproduces inside the larval stages and infects the eggs of the adult
female before she emerges from a tree. The nematode is effective in
controlling Sirex woodwasp because it sterilizes the female but does not
affect her flying ability, thus allowing her to disperse nematodes widely.

Web Links for Information http://na.fs.fed.us/spfo/pubs/pest_al/sirex_woodwasp/sirex_woodwasp.

on Sirex Woodwasp htm; a pest alert fact sheet of the USDA Forest Service on identification
and biology.
http://www.invasivespeciesinfo.gov/animals/sirexwasp.shtml; a list of
websites about Sirex noctilio.
http://www.massnrc.org/pests/pestFAQsheets/european%20woodwasp.
htm; a fact sheet of the University of Massachusetts, including links to
key sites for further information.
http://www.aphis.usda.gov/plant_health/plant_pest_info/sirex/
downloads/sirexfungus.pdf; a fact sheet on Amylostereum areolatum.

Articles Bedding, R.A. and E.T. Iede. 2005. Application of Beddingia siricidicola
for Sirex woodwasp control. In: Nematodes as Biocontrol Agents. CAB
International, Cambridge: 385-399.
Long, S.J., D.W. Williams, and A.E. Hajek. 2009. Sirex species
(Hymenoptera: Siricidae) and their parasitoids in Pinus sylvestris in
eastern North America. The Canadian Entomologist 14: 153-157.
Dodds, K.J., P. de Groot, and D.A. Orwig. 2010. The impact of Sirex
noctilio in Pinus resinosa and Pinus sylvestris stands in New York and
Ontario. The Canadian Journal of Forest Research 40: 212-223.
Slippers, B., P. DeGroot and M.J. Wingfield (eds.). The Sirex Woodwasp
and its Fungal Symbiont; Research and Management of a Worldwide
Invasive Pest. Springer Pub. In press.
Schiff, N.M., H. Goulet, D.R. Smith, C. Boudreault, A.D. Wilson, and
B.E. Scheffler. 2011. Siricidae (Hymenoptera: Siricoidea) of the Western
Hemisphere. Canadian Journal of Arthropod Identification. In press.
A Photographic Guide 689

189. Asian Chestnut Gall Wasp, Drycosmus kuriphilus Yasumatsu

(Hymenoptera: Cynipidae)

Orientation to Pest The Asian chestnut gall wasp, Drycosmus kuriphilus Yasumatsu, is a
gall making cynipid that is specific to chestnut (Castanea spp.). The gall
wasp is native to China but invasive in many countries around the world,
including parts of the United States. Galling disrupts shoot and twig
growth, impedes normal flowering, and weakens infested trees. Adult
females (there are no males) emerge from galls in early to mid summer
and lay several eggs in each chestnut bud they attack. Eggs hatch in mid
to late summer, and young larvae feed a little and then overwinter in
the infested buds. When buds begin to grow in spring, galls are formed.
Galls have one to several chambers, each with a developing gall wasp
larva. Larvae eat the tissue within the galls and pupate there. New gall
wasps emerge soon after. After adults emerge, galls turn brown and
woody and may remain on the tree for several years and are a clear sign
of infestation. In China, pest outbreaks occur about every 10 years and
last 2-3 years. There is one generation per year.

Hosts Commonly Attacked This species is specific to species of chestnuts (Castanea).

Distribution Asian chestnut gall wasp is native to China, but is also present in Korea,
Japan, Nepal, Europe (Italy, France, Slovenia, Hungary, Switzerland, and
Croatia) and the United States as an invasive pest.

Figure 1. Distribution of
the chestnut gall wasp,
Drycosmus kuriphilus,
in the United States.
(Ignazio Graziosi)

Images of Chestnut Figure 2. Adult of Asian

chestnut gall wasp.
Gall Wasp (Lynne Rieske)
690 Forest Pest Insects in North America:

Images of Chestnut Figure 3. Young gall

of Asian chestnut gall
Gall Wasp (continued) wasp in spring (top)
and old gall after wasp
emergence (bottom),
which may remain on
trees for several years.
(Both photos: Lynne
Rieske)

Figure 4. Interior view

of gall in cross section,
showing multiple
chambers and one gall
wasp pupa. (Melanie
Sprinkle, Bugwood.org)

Figure 5.View of
old galls on leafless
chestnut tree in winter.
(Lynne Rieske)
A Photographic Guide 691

Important Biological Control Classical biological control programs focusing on the parasitoid Torymus
Agents Related to this sinensis Kamijo (Hymenoptera: Torymidae) have been implemented in
Pest Species several countries, including Japan and the United States. Several native
species also utilize the gall wasp as a host.

Web Links for Information http://www.eppo.org/QUARANTINE/insects/Dryocosmus_kuriphilus/

on Chestnut Gall Wasp DS_Dryocosmus_kuriphilus.pdf; extensive document of EPPO
(European quarantine organization) on many aspects of the species’
biology.
http://www.agroscope.admin.ch/data/publikationen/1304403002_Aebi_
A_Paper_Dryocosmus_Kuriphilus_01-2011.pdf; an extensive document
on this species and its biological control in Switzerland.

Articles Moriya, S., K. Inoue, A. Otake, M. Shiga, and M. Mabuchi. 1989.

Decline of the chestnut gall wasp population, Dryocosmus kuriphilus
Yasumatsu (Hymenoptera, Cynipidae) after the establishment of Torymus
sinensis Kamijo (Hymenoptera, Torymidae). Applied Entomology and
Zoology 24: 231-233.
Aebi, A., K. Schonrogge, G. Melika, A. Quacchia, A. Alma, and G.N.
Stone. 2007. Native and introduced parasitoids attacking the invasive
chestnut gall wasp Dryocosmus kuriphilus. Bulletin of the European and
Mediterranean Plant Protection Organization (EPPO Bulletin) 37:
166-171.
Cooper, W.R. and L.K. Rieske. 2007. Community associates of an exotic
gallmaker, Dryocosmus kuriphilus (Hymenoptera: Cynipidae), in eastern
North America. Annals of the Entomological Society of America 100:
236-244.
Cooper, W.R. and L.K. Rieske. 2007. Review of the historic and current
status of the Asian chestnut gall wasp in North America. Journal of the
American Chestnut Foundation 11: 26-33.
Rieske, L.K. 2007. Success of an exotic gallmaker, Dryocosmus
kuriphilus, on chestnut in the USA: a historical account. Bulletin of the
European and Mediterranean Plant Protection Organization (EPPO
Bulletin) 37: 172-174.
692 Forest Pest Insects in North America:

190. Spruce Spider Mite, Oligonychus ununguis (Jacobi) (Acari:

Tetranychidae)

Orientation to Pest Spruce spider mite, Oligonychus ununguis (Jacobi), is a cosmopolitan

spider mite of uncertain origin associated with spruce (Picea) and other
conifers. All life stages (egg, immature stages, adult) occur together on
the host foliage, which is webbed over as infestation grows. Winter is
passed in the egg stage. Multiple generations occur per year. Outbreaks
are often induced by the use of pesticides but may occur naturally in
dry years. Outbreaks are more common on young trees, on which mites
prefer older foliage in the interior of the tree.

Hosts Commonly Attacked This mite feeds on Colorado (Picea pungens Engelm.) and white (P.
glauca [Moench] Voss) spruce, Douglas-fir (Pseudotsuga menzeseii
Mirb.), thuja (Thuja), jack (Pinus banksiana Lamb.) and some other
pines, eastern hemlock (Tsuga canadensis [L.] Carrière), and larch
(Larix).

Distribution Spruce spider mite is found in many countries around the world, and
its origin in unknown. Its occurrence on introduced plants in countries
lacking native conifers suggests that at least part of this distribution is
from invasion, likely due to movement of infested woody nursery stock.

Images of Spruce Figure 1. Adult of spruce

spider mite, Oligonychus
Spider Mite ununguis. (Ward Strong,
BC Ministry of Forests,
Bugwood.org)

Figure 2. Eggs of spruce spider mite. (Both photos: Ward Strong, BC Ministry of
Forests, Bugwood.org)
A Photographic Guide 693

Images of Spruce Figure 3. Colony, with

webbing, of spruce
Spider Mite (continued) spider mite. (USDA
Forest Service - Region
4 - Intermountain
Archive, USDA Forest
Service, Bugwood.org)

Figure 4. Close up of chlorotic spotting caused by spruce spider mite feeding.

(John A. Weidhass, Virginia Polytechnic Institute and State University,
Bugwood.org)
694 Forest Pest Insects in North America:

Images of Spruce Figure 5. Damage on

white spruce (top) from
Spider Mite (continued) spruce spider mite
and close up (bottom)
of webbing from mite.
(Both photos: Ward
Strong, BC Ministry of
Forests, Bugwood.org)

Important Biological Control As with most spider mites, control agents are various species of
Agents Related to this predatory mites (Phytoseiidae), predatory gall midges (Cecidomyiidae,
Pest Species genus Feltiella), and ladybird beetles (Coccinellidae), which vary by
region. Destruction of such natural enemies through widespread use
of pesticides for control of other forest pests can induce outbreaks of
O. ununguis. This occurred in Montana in the 1950s, for example,
when DDT was applied over a large area for control of western spruce
budworm (Choristoneura occidentalis Freeman). Such problems are
most likely in plantations, seed orchards, or Christmas tree farms that are
regularly treated for various insect pests.
A Photographic Guide 695

Web Links for Information http://imfc.cfl.scf.rncan.gc.ca/insecte-insect-eng.asp?geID=136; fact

on Spruce Spider Mite sheet of Natural Resources Canada.
http://wiki.bugwood.org/Oligonychus_ununguis; Bugwood Wiki
factsheet on biology and control.
http://ento.psu.edu/extension/christmas-trees/information/pest-fact-
sheets/Spspidermiteent069.pdf/view; Pennsylvania Department of
Agriculture circular on control of spruce spider mite.

Articles Löyttyniemi, K. and K. Heliövaara. 1991. Effect of forest fertilization

on the spruce spider mite Oligonychus ununguis (Jacobi) (Acarina,
Tetranychidae). Acarologia 32: 139-143.
Shrewsbury, P.M. and M.R. Hardin. 2003. Evaluation of predatory
mite (Acari: Phytoseiidae) releases to suppress spruce spider mites,
Oligonychus ununguis (Acari: Tetranychidae), on juniper. Journal of
Economic Entomology 96: 1675-1684.
696 Forest Pest Insects in North America:
A Photographic Guide 697

Acknowledgments
We thank the following for their review of the species listed:

1. Pear thrips, Taeniothrips inconsequens: Ken Raffa, [email protected]

2. Introduced basswood thrips, Thrips calcaratus: Ken Raffa, [email protected]
3. Redbanded thrips, Selenothrips rubrocinctus: Howard Frank, [email protected]
4. Slash pine flower thrips, Gnophothrips fuscus: Howard Frank, [email protected]
5. Northern walkingstick, Diapheromera femorata: Howard Frank, [email protected]
6. Oak lace bug, Corythucha arcuata: Paula Shrewsbury, [email protected]
7. Sycamore lace bug, Corythucha ciliata: Paula Shrewsbury, [email protected]
8. Shield-backed pine seed bug, Tetyra bipunctata: Jean Turgeon, [email protected]
9. Leaffooted pine seed bug, Leptoglossus corculus: Dan Miller, [email protected]
10. Royal palm bug, Xylastodoris luteolus: Howard Frank, [email protected]
11. Saratoga spittlebug, Aphrophora saratogensis: Ken Raffa, [email protected]
12. Pine spittlebug, Aphrophora parallela: Ken Raffa, [email protected]
13. Thorn bug, Umbonia crassicornis: Howard Frank, [email protected]
14. Periodical cicada, Magicicada septendecim: Paula Shrewsbury, [email protected]
15. Eastern spruce gall adelgid, Adelges abietis: Scott Salom, [email protected]
16. Balsam woolly adelgid, Adelges piceae: Mike Montgomery, [email protected]
and Scott Salom, [email protected]
17. Hemlock woolly adelgid, Adelges tsugae: Scott Salom, [email protected]
18. Cooley spruce gall adelgid, Adelges cooleyi: Scott Salom [email protected]
19. Pine bark adelgid, Pineus strobi: Scott Salom, [email protected]
20. Pine leaf adelgid, Pineus pinifoliae: Scott Salom, [email protected]
21. White pine aphid, Cinara strobi: Mike Montgomery, [email protected]
22. Woolly elm aphid, Eriosoma americanum: Mike Montgomery, [email protected]
23. Woolly beech aphid, Phyllaphis fagi: Mike Montgomery, [email protected]
24. Tuliptree aphid, Illinoia liriodendri: Mike Montgomery, [email protected]
25. Norway maple aphid, Periphyllus lyropictus: Mike Montgomery, [email protected]
26. Linden aphid, Eucallipterus tiliae: Mike Montgomery, [email protected]
27. Spruce aphid, Elatobium abietinum: Ann Lynch, [email protected]
28. Beech scale, Cryptococcus fagisuga: Ben Normark, [email protected]
29. Gillette’s eriococcin, Eriokermes gillettei (= Eriococcus gillettei): Ben Normark,
[email protected]
30. Pine bast scale, Matsucoccus matsumurae: Ben Normark, [email protected]
31. Tuliptree scale, Toumeyella liriodendri: Ben Normark, [email protected]
32. Pine tortoise scale, Toumeyella parvicornis: Steve Clarke, [email protected]
33. Tessellated scale, Eucalymnatus tessellatus: Ben Normark, [email protected]
698 Forest Pest Insects in North America:

Acknowledgments (continued)
We thank the following for their review of the species listed:

34. Elongate hemlock scale, Fiorinia externa: Kris Abell, [email protected]

35. Oystershell scale, Lepidosaphes ulmi: Ben Normark, [email protected]
36. San José scale, Diaspidiotus perniciosus: Ben Normark, [email protected]
37. Willow scale, Diaspidiotus gigas: Rob Progar, [email protected]
and Ben Normark, [email protected]
38. Camphor scale, Pseudaonidia duplex: Ben Normark, [email protected]
39. Coconut scale, Aspidiotus destructor: Ben Normark, [email protected]
40. Florida red scale, Chrysomphalus aonidum: Ben Normark, [email protected]
41. Magnolia white scale, Pseudaulacaspis cockerelli: Ben Normark, [email protected]
42. White peach scale, Pseudaulacaspis pentagona: Ben Normark, [email protected]
43. Citrus whitefly, Dialeurodes citri: Howard Frank, [email protected]
44. The mealybug Oracella acuta: Steve Clarke, [email protected]
45. Cottonwood leaf beetle, Chrysomela scripta: David Coyle, [email protected]
46. Imported willow leaf beetle, Plagiodera versicolora: Paula Shrewsbury, [email protected]
47. Elm leaf beetle, Xanthogaleruca luteola: Steve Dreistadt, [email protected]
48. Larger elm leaf beetle, Monocesta coryli: Steve Dreistadt, [email protected]
49. Giant palm weevil, Rhynchophorus cruentatus: Mark Hoddle, [email protected]
50. Red palm weevil, Rhynchophorus ferrugineus: Mark Hoddle, [email protected]
51. Pales weevil, Hylobius pales: Ken Raffa, [email protected]
52. Pine root collar weevil, Hylobius radicis: Ken Raffa, [email protected]
53. Pitch-eating weevil, Pachylobius picivorus: Ken Raffa, [email protected]
54. White pine weevil, Pissodes strobi: Ken Raffa, [email protected]
55. Eastern pine weevil, Pissodes nemorensis: Ken Raffa, [email protected]
56. Whitefringed beetles, Naupactus spp.: Ben Normark, [email protected]
57. Yellow-poplar weevil, Odontopus calceatus: David Coyle, [email protected]
58. Native elm bark beetle, Hylurgopinus ru ipes: Steve Katovitch, [email protected]
and Bob Rabaglia, [email protected]
59. Smaller European elm bark beetle, Scolytus multistriatus: Roger Fuester,
[email protected]
60. Spruce beetle, Dendroctonus ru ipennis: Ann Lynch, [email protected]
61. Black turpentine beetle, Dendroctonus terebrans: Ron Billings, [email protected]
62. Eastern larch beetle, Dendroctonus simplex: Robert Haack, [email protected]
63. Southern pine beetle, Dendroctonus frontalis: Ron Billings, [email protected]
64. Western pine beetle, Dendroctonus brevicomis: Andy Eglitis, [email protected]
65. Jeffrey pine beetle, Dendroctonus jeffreyi: Patrick Shea, [email protected]
A Photographic Guide 699

Acknowledgments (continued)
We thank the following for their review of the species listed:

66. Mountain pine beetle, Dendroctonus ponderosae: Barbara Bentz, [email protected] and
Patrick Shea, [email protected]
67. Douglas-fir beetle, Dendroctonus pseudotsugae: connie mehmel, [email protected]
68. Engraver beetle, Ips typographus: Fredrik Schlyter, [email protected]
69. Pine engraver beetle, Ips pini: Ron Billings, [email protected]
70. Southern pine engraver, Ips grandicollis: Ron Billings, [email protected]
71. Six-spined Ips, Ips calligraphus: Ron Billings, [email protected]
72. Small southern pine engraver, Ips avulsus: Ron Billings, [email protected]
73. Silver fir beetle, Pseudohylesinus sericeus: Ron Billings, [email protected]
and Bill Ciesla, [email protected]
74. Fir engraver, Scolytus ventralis: Ann Lynch, [email protected]
75. Columbian timber beetle, Corthylus columbianus: Howard Frank, [email protected]
76. Ambrosia beetles, Xyleborus spp.: Jim Hanula, [email protected]
77. Redbay ambrosia beetle, Xyleborus glabratus: Jim Hanula, [email protected]
78. Walnut twig beetle, Pityophthorus juglandis: Bud Mayfield, [email protected]
79. Striped ambrosia beetle, Trypodendron lineatum: Jim Hanula, [email protected]
80. Black twig borer, Xylosandrus compactus: Jim Hanula, [email protected]
81. Larger pine shoot beetle, Tomicus piniperda: Robert Haack, [email protected]
82. White pine cone beetle, Conophthorus coniperda: Steve Katovich, [email protected]
83. Lodgepole cone beetle, Conophthorus ponderosae: Ward Strong, [email protected]
85. May and June beetles, Phyllophaga spp.: Steve Katovich, [email protected]
86. Black vine weevil, Otiorhynchus sulcatus: Lance Osborne, [email protected]
87. Strawberry root weevil, Otiorhynchus ovatus: Lance Osborne, [email protected]
88. Japanese beetle, Popillia japonica: Lance Osborne, [email protected]
89. Two-lined chestnut borer, Agrilus bilineatus: Bob Haack, [email protected]
90. Bronze birch borer, Agrilus anxius: Bob Haack, [email protected]
91. Emerald ash borer, Agrilus planipennis: Jian Duan, [email protected]
92. Goldspotted oak borer, Agrilus auroguttatus: Tom Coleman, [email protected]
93. Soapberry borer, Agrilus prionurus: Ron Billings, [email protected]
94. Hickory spiral borer, Agrilus torquatus: Matt Ginzel, [email protected]
95. Flatheaded appletree borer, Chrysobothris femorata: Tom Coleman, [email protected]
96. Red oak borer, Enaphalodes rufulus: Fred Stephen, [email protected]
97. Black fir sawyer beetle, Monochamus urussovi: Mike Montgomery, [email protected]
98. Japanese pine sawyer, Monochamus alternatus: Mike Montgomery, [email protected]
700 Forest Pest Insects in North America:

Acknowledgments (continued)
We thank the following for their review of the species listed:

99. Asian longhorned beetle, Anoplophora glabripennis: Rhonda Santos, [email protected]

100. Poplar borer, Saperda calcarata: Steve Katovich, [email protected]
102. Cottonwood borer, Plectrodera scalator: David Coyle, [email protected]
103. Banded hickory borer, Knulliana cincta: Jon Sweeney, [email protected]
104. Locust borer, Megacyllene robiniae: Jon Sweeney, [email protected]
105. White oak borer, Goes tigrinus: Matt Ginzel, [email protected]
and Jon Sweeney, [email protected]
106. Whitespotted sawyer, Monochamus scutellatus: Jean Turgeon, [email protected]
107. Bagworm, Thyridopteryx ephemeraeformis: Marc Rhainds, [email protected]
108. Birch casebearer, Coleophora serratella: (no outside reviewer)
109. Larch casebearer, Coleophora laricella: Roger Ryan, [email protected]
110. Pecan cigar casebearer, Coleophora laticornella: (no outside reviewer)
111. Oak skeletonizer, Bucculatrix ainsliella: Erik van Nieukerken, [email protected]
112. Palm leaf skeletonizer, Homaledra sabalella: Howard Frank, [email protected]
113. Mimosa webworm, Homadaula anisocentra: Robert Childs, [email protected]
114. Lodgepole needleminer, Coleotechnites milleri: (no outside reviewer)
115. Banded ash clearwing, Podosesia aureocincta: Matt Ginzel, [email protected]
116. Red oak clearwing borer, Paranthrene simulans: Matt Ginzel, [email protected]
and Tom Coleman, [email protected]
117. Persimmon borer, Sannina uroceriformis: Matt Ginzel, [email protected]
118. Poplar clearwing borer, Paranthrene dollii: David Coyle, [email protected]
119. Carpenterworm, Prionoxystus robiniae: Marvin Harris, [email protected]
and Brytten Steed, [email protected]
120. Pecan carpenterworm, Cossula magnifica: Marvin Harris, [email protected]
121. Spruce budworm, Choristoneura fumiferana: Lisa Lumley, [email protected]
122. Jack pine budworm, Choristoneura pinus pinus: Lisa Lumley, [email protected]
123. Large aspen tortrix, Choristoneura conflictana: Lisa Lumley, [email protected]
124. Sugar pine tortrix, Choristoneura lambertiana: Lisa Lumley, [email protected]
125. Western spruce budworm, Choristoneura occidentalis: Lisa Lumley, [email protected]
126. Spruce bud moth, Zeiraphera canadensis: Jean Bev Turgeon, [email protected]
127. Cottonwood twig borer, Gypsonoma haimbachiana: David Coyle, [email protected]
129. Oak leafroller, Archips semiferanus: Todd Gilligan, [email protected]
130. Longleaf pine seedworm, Cydia ingens (= Lapeyresia ingens): Todd Gilligan, [email protected]
131. European pine tip moth, Rhyacionia buoliana: Darrell Ross, [email protected]
A Photographic Guide 701

Acknowledgments (continued)
We thank the following for their review of the species listed:

132. Nantucket pine shoot moth, Rhyacionia frustrana: Jim Lashomb, [email protected]
and Darrell Ross, [email protected]
133. Pitch pine tip moth, Rhyaciona rigidana: Darrell Ross, [email protected]
134. Southwestern pine tip moth, Rhyaciona neomexicana: Darrell Ross, [email protected]
135. Eastern pine shoot borer, Eucosma gloriola: Jean Turgeon, [email protected]
136. Lodgepole cone moth, Eucosma rescissoriana: Ward Strong, [email protected]
137. Western pine shoot borer, Eucosma sonomana: Darrell Ross, [email protected]
138. Douglas-fir cone moth, Barbara colfaxiana: Ward Strong, [email protected]
139. Pine webworm, Pococera robustella: Bud Mayfield, [email protected]
140. Zimmerman pine moth, Dioryctria zimmermani: Amanda Roe, [email protected]
141. Fall cankerworm, Alsophila pometaria: Brytten Steed, [email protected]
142. Spring cankerworm, Paleacrita vernata: Brytten Steed, [email protected]
143. Eastern hemlock looper, Lambdina fiscellaria fiscellaria: Darrell Ross, [email protected]
144. Western hemlock looper, Lambdina fiscellaria lugubrosa: Darrell Ross, [email protected]
145. Greenstriped forest looper, Melanolophia imitata: (no outside reviewer)
146. Saddleback looper, Ectropis crepuscularia: (no outside reviewer)
147. Spearmarked black moth, Rheumaptera hastata: (no outside reviewer)
148. Elm spanworm, Ennomos subsignarius: Peggy Dixon, [email protected],
Heidi Fry, [email protected] and Krista Ryall, [email protected]
149. Winter moth, Operophtera brumata: Joe Elkinton, [email protected]
150. Bruce spanworm, Operophtera bruceata: Joe Elkinton, [email protected]
151. Forest tent caterpillar, Malacosoma disstria (Lasiocampidae): Joe Elkinton, [email protected]
152. Eastern tent caterpillar, Malacosoma americanum (Lasiocampidae): Joe Elkinton, [email protected]
153. Siberian moth, Dendrolimus sibiricus: Mike Montgomery, [email protected]
154. Pinkstriped oakworm, Anisota virginiensis (Saturniidae): Toby Petrice, [email protected]
155. Orangestriped oakworm, Anisota senatoria (Saturniidae): Rose Hiskes, [email protected]
and Toby Petrice, [email protected]
156. Pandora moth, Coloradia pandora: Darrell Ross, [email protected]
157. European gypsy moth, Lymantria dispar (Lymantriidae): Roger Fuester, [email protected]
158. Asian pink moth, Lymantria mathura: Roger Fuester, [email protected]
159. Nun moth, Lymantria monacha (Lymantriidae): Melody Keena, [email protected]
160. Browntail moth, Euproctis chrysorrhoea (Lymantriidae): Joe Elkinton, [email protected]
161. Pine tussock moth, Dasychira pinicola (Lymantriidae): Steve Katovich, [email protected]
162. White-marked tussock moth, Orgyia leucostigma (Lymantriidae): Robert Johns, [email protected]
702 Forest Pest Insects in North America:

Acknowledgments (continued)
We thank the following for their review of the species listed:

163. Douglas-fir tussock moth, Orgyia pseudotsugata: Ann Lynch, [email protected]

164. Poplar tentmaker, Clostera inclusa: David Coyle, [email protected]
165. Saddled prominent, Heterocampa guttivitta (Notodontidae): Dave Wagner, [email protected]
166. Walnut caterpillar, Datana integerrima: Dave Wagner, [email protected]
167. Variable oak leaf caterpillar, Lochmaeus manteo (Notodontidae): Joe Pase, [email protected]
168. Fall webworm, Hyphantria cunea (Arctiidae): Bud Mayfield, [email protected]
169. Arborvitae leafminer, Argyresthia thuiella: Paul Opler, [email protected]
170. Pine needle sheathminer, Zelleria haimbachi: Ward Strong, [email protected]
171. Pine butterfly, Neophasia menapia: Don Scott, [email protected]
172. Balsam gall midge, Paradiplosis tumifex (Cecidomyiidae): Rick West, [email protected]
173. Poplar leafcurl midge, Prodiplosis morrisi (Cecidomyiidae): Brytten Steed, [email protected]
174. Douglas-fir needle midge, Contarinia pseudotsugae (Cecidomyiidae): Ward Strong,
[email protected]
175. Pine false webworm, Acantholyda erythrocephala (Pamphiliidae): Barry Lyons,
[email protected]
176. Redheaded pine sawfly, Neodiprion lecontei (Diprionidae): Bud Mayfield, [email protected]
177. Swaine jack pine sawfly, Neodiprion swainei (Diprionidae): Chris MacQuarrie,
[email protected]
178. European pine sawfly, Neodripion sertifer (Diprionidae): Barry Lyons, [email protected]
179. Virginia pine sawfly, Neodipirion pratti pratti (Diprionidae): Chris MacQuarrie,
[email protected]
180. Hemlock sawfly, Neodiprion tsugae: Chris MacQuarrie, [email protected]
181. Introduced pine sawfly, Diprion similis: Barry Lyons, [email protected]
182. European spruce sawfly, Gilpinia hercyniae: Chris MacQuarrie,
[email protected]
183. Larch sawfly, Pristiphora erichsonii (Tenthredinidae): Ken Raffa, [email protected]
184. Mountain ash sawfly, Pristiphora geniculata: Rick West, [email protected]
185. Yellowheaded spruce sawfly, Pikonema alaskensis: Rob Johns, [email protected].
186. Birch leafminer, Fenusa pumila: Richard Casagrande, [email protected]
187. Ambermarked birch leafminer, Profenusa thomsoni: Anna Soper, [email protected]
188. Sirex woodwasp, Sirex noctilio: Dave Williams, [email protected]
189. Asian chestnut gall wasp, Dryocosmus kuriphilus: L.K. Rieske, [email protected]
190. Spruce spider mite, Oligonychus ununguis: Ward Strong, [email protected]