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INSECTS OF EASTERN FORESTS

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Miscellaneous Publication No. 1426
December 1985

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This publication supersedes ‘‘Eastern Forest Insects,”
Miscellaneous Publication No. 1175

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Washington, D.C.

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For sale by the Superintendent of Documents, U.S. Government Printing Office 137

Washington, D.C. 20402

iii

Library of Congress Catalog Card Number: 82-600590

U.S. Department of Agriculture, Forest Service.
Insects of eastern forests. Misc. Publ. 1426. Washington, DC:
U.S. Department of Agriculture, Forest Service; 1985. 608 p.

Highlights the description and biology of numerous species of harmful and useful
insects found in forests and forest products east of the 100th meridian. This manual
provides the forest practitioner and others with the means for identifying these
insects and their related organisms.

KEY WORDS: damaging arthropods of America north of Mexico; entomological

manual; forest pests in the United States: sap feeders, wood borers, bark beetles,
and defoliators in North America; tree and wood-product pests.

ii

Contents

Gauci,
ACKnOWIEdEIMeMUS 62.2 aes Gere a Hes a eho Rei dithe eae ateasret eon ated Medd one 8 Vv
PIRSIIS™ 35 SARS elie ery ceitee ed OR Re Or ce oe 1X
ROneS iMG GUSH eats Sint sacat alanis Such, eds aut aie MG me Ro Seed tau l
assesvcause diy fOneSt MNSECtS ssi. Sv eicy passe 4s a east chet ts sake, oye Ree 2
OTe stMMSCCM SUV. Cie core dien 2 sh. Stee he apes Hae pies A emetyae ha De ake Z
ForestaincecteCOmmOh 2 i 2s tee ti ea Sales wis Sa Gog Bale oe oe ee es 5
IN MUA EC OMELOMSM en reritaeaes nS ne. Mab One eR eda We uae aia gah eum ou 5
EDP Ga COMO ls trast nts Heide Lote SS eek ars. aa wee Ta Se pele ee 8
SINC WIC RCOMULON Me ctkee war aie vets Sa sa wisiny Ly otha ee a ee 9
Physicalcand mechanical methods of control... . 2. 66s eet ee ee 2
RerUlAlOmmeconerO leer att xr ct aus en das.4 Oat aad net coe @ hme a Mg Ee 10
BIOlOICAlBCOMEOl Oe ee ek wy kee Dea ek Gs bn Sees eA wa 4 13
ChiermcalPeominoleeces Fre ah le gas Ge es a baa eee whe bade 14
Othcrmapproacnes-to. COntrol.<. Foye we te se he Pd a ca ee eee 16
intesratedspestemanapemient |. Fe ee a en cea eee ee en 18
InSectseandeaireeDISCASES. ans eb Lawigive ewe Bone settee ee mete aed ade 20
Insectscandrixelated Ore anisms. se eles Pe a ened sale eda ke eee na ene 25
BhylumeWollusca—=-Shipworms -. e200. 2 ce ee Mee cee een ee be 2
iy AME ACO O PO aust: kak aes Ss Go ey tate gl pulp ain e Ge ons Pale eee dee 26
Classa@nustacea—_WOOd Tice: eis kh Sey tn wees dew ee ea dg eeu ee px |
GlasssDiplopoda=—Millipedes:\to.. ne gs Ba lee Ra ene a es 27
Class*Ghilopoda=-Centipedes ees ii eis oe eee yb ota eee ee a ees 28
Class Arachnida—Scorpions, spiders, mites, ticks, and allies ........ 28
Chasssiniscc ta MMSCCIS iw pach tes wy Miklal cen oo See ee sb we as hawt 33
Keys to the Orders and Families of Forest Insects and Allies ........... 34
lmportantzandeSeleeted: Orders of Forest Insects: e.8 be eee ee ee we
Ordenslhysanmura=—Bristletaus. soo. he nS ee ee oe 44
OrdemiCollcmbola——Springtails | i012 is led ee Ne ek ee ee eds Ad
@idersEphemeroptera==Mayilies® tases Pia is le ce eb ene Ae}
Order Odonata—Dragonflies and damselflies ...................... 45
@rdewPlecopiera——Stoneflies 5. alo oe Se ea ee a de 45
OrdermPsocoptera—Booklice and psocids 0.02)... ee eed. 46
Order Miallophaga—Chewine iCe- 3 oy che Fe a ee be es ee 46
OndemAnoplura=——Sucking Hee ioe ey we ce cally ee ee ee ee wee 46
Ordewmhysanoptera——T hrips: 2... we a a ee 46
Order Neuroptera—Dobsontlies, lacewings, antlions, and allies ...... 47
Ondem Siphomaptera—Fleas Goat oak Geos occa eens gee ee tk be es 48
Order Orthoptera—Grasshoppers, crickets, mantids, and allies ....... 48
Ordcmlsoplera.— lermitess ae ee ia ee eke bs Ea eae es ool 54
@idemblemiptera— mie DUS ei ce eG Sen ees ca wae os 63
~ Order Homoptera—Aphids, spittlebugs, scale insects, and allies ...... 69
Order Lepidoptera—Butterflies, moths, skippers ................... 123
Order Coleapteras- Beetles soo ic ie ee eee ew ew i eae es 237

ii

Order Hymenoptera—Ants, bees, sawflies, wasps, and allies ........ S74.

Order"Diptera--PHCS. oak mca eae ne eas eg ee fee ee ee 440
Peteranre Cited). 2.2 sccane sea gree ee te ee 455
Common and Scientific Names of Host Plants ....................... 529
Index tou lnsects bY FlOst- Plans 5 ot agaansaes een eas Oe ee 537
ISCCE SOK <P ie reek Sid eo ales eoh kana ar ig ae Ri we ca Ge Be 579

Acknowledgments

Arnold T. Drooz, principal research entomologist, Southeastern Forest Experi-
ment Station, Asheville, N.C., edited and coordinated materials from a team of
specialists, who called upon others as needed. The original intent was to use as
much of W. L. Baker’s “Eastern Forest Insects” as possible, making only a
minimum of changes. This proved impractical, as a number of specialists suggested
that major up-dating was needed. These requests were allowed.

Only a few members of the team are qualified taxonomists. Therefore, the text
was submitted to systematists in the U.S. Department of Agriculture and Smithso-
nian Institution for verification of the scientific names. Warm and grateful thanks
and appreciation are due all of these people and their organizations for providing
their time, expertise, and valuable suggestions toward completing this volume.

The names, organizations, and contribution of each member of this team are
listed to cordially recognize their help. The following specialists revised and
rewrote the subject matter of the text that follows their names:

Canadian Forestry Service, St. John’s, Newfoundland

Arthur G. Raske—Staphylinidae
Connecticut Agricultural Experiment Station, New Haven
John F. Anderson—Lymantriidae
Duke University, School of Forestry, Durham, N.C.
Roger F. Anderson, retired—Forest Insects to Isoptera
Iowa State University, Department of Entomology, Ames
James W. Mertins—Lepidoptera to Noctuidae
North Carolina State University, Department of Entomology, Raleigh
Herbert H. Neunzig—Pyralidae
State University of New York, College of Environmental Science and Forestry,
Syracuse
Douglas C. Allen—Geometridae, Notodontidae
Gerald N. Lanier—most of Scolytidae
Roy A. Norton—Insects and Related Organisms
U.S. Department of Agriculture, Forest Service
David E. Donley, Morgantown, W. Va.—Diptera
Bernard H. Ebel, Olustee, Fla. —Pyralidae
Gerhard F. Fedde, formerly Athens, Ga.—Aphididae
Cynthia K. Franklin, formerly Research Triangle Park, N.C.—Mutillidae
through Apoidea
Paul B. Godwin, Hamden, Conn.—Rhyncophora
Michael I. Haverty, Berkeley, Calif. —Isoptera
Cyrus J. Hay, retired, Delaware, Ohio—Cerambycidae
William E. Miller, formerly St. Paul, Minn.—Tortricidae
Jack C. Nord, Research Triangle Park, N.C.,—ambrosia beetles and
Platypodidae
John W. Peacock, Delaware, Ohio—Insects and Tree Diseases
James D. Solomon, Stoneville, Miss.—Cossidae, Cerambycidae

Robert C. Thatcher, Pineville, La.—southern pine beetle through red tur-
pentine beetle
Lonnie H. Williams, Gulfport, Miss.—Coleoptera to Bruchidae
Louis F. Wilson, East Lansing, Mich.—Orthoptera
U.S. Department of Agriculture, Agricultural Research Service (ARS), System-
atic Entomology Laboratory, Beltsville, Md.
Douglass R. Miller—Coccoidea
University of Florida, Department of Entomology and Nematology, Gainesville
Robert C. Wilkinson—Symphyta |
University of Georgia, Department of Entomology, Athens
John All—Bruchidae to Buprestidae |
Rudolph T. Franklin—Buprestidae
Ching H. Tsao—Yponomeutidae |
Robert H. Turnbow— Buprestidae
University of Wisconsin, Department of Entomology, Madison
Harry C. Coppel—Apocrita
Virginia Polytechnic Institute and State University, Department of Entomology,
Blacksburg
Herman J. Heikkenen—Noctuidae to Lymantriidae
Others who kindly gave of their time and expertise in reviewing revised and new
materials are:
Brigham Young University, Zoology and Entomology Laboratory, Provo
Stephen L. Wood—Scolytidae
U.S. Department of Agriculture, Forest Service
Arnold T. Drooz, Olustee, Fla.—Forest Insects, Symphyta, Geometridae
E. George Kuhlman, Research Triangle Park, N.C.—TInsects and Tree
Diseases

Robert L. Talerico, formerly Broomall, Pa.—Choristoneura spp.
Harry 0. Yates HI, Athens, Ga.—Olethreutinae, Phycitinae
U.S. Department of Agriculture, ARS Systematic Entomology Laboratory,
Beltsville, Md.
R. W. Poole—Noctuidae
Louise M. Russell, retired, cooperating scientist— Aphididae
Manya B. Stoetzel—Aphididae
Virginia Polytechnic Institute and State University, Department of Entomology,
Blacksburg
Herman J. Heikkenen—Olethreutinae
University of Florida, Department of Entomology and Nematology, Gainesville
Harold N. Greenbaum—Symphyta
University of Wisconsin, Department of Entomology, Madison
Daniel M. Benjamin—Symphyta
Harry C. Coppel—Symphyta
There have been numerous changes in nomenclature since the earlier publications
were printed, and keeping up with these changes is a never-ending process. Edna
M. Hoover (retired) of the U.S. Department of Agriculture, ARS, Systematic
Entomology Laboratory (SEL) at Beltsville, Md., transmitted the manuscript to
various SEL and other specialists for verification of scientific names. She also
verified some names when no specialist was available and assembled and transmit-
ted the corrected texts. Her interest and devotion to this chore are greatly appreci-
ated. The following list acknowledges the invaluable help of the SEL systematists at

vi

Beltsville and Washington, D.C., and indicates their area of expertise:
D. M. Anderson—Scolytidae
E. W. Baker—Acari
S. W. T. Batra—Apidae
R. W. Carlson—Ichneumonidae, Evaniidae
D. C. Ferguson—Nymphalidae, Papilionidae, Hesperiidae, Sphingidae, Geo-
metridae, Lymantriidae, Lasiocampidae, Mimallonidae, Pyralidae, Phycitinae
R. H. Foote—Tephritidae
R. J. Gagné—Cecidomyliidae, Asilidae, Muscidae, Calliphoridae, Sar-
cophagidae, Mycetophagidae, Oestridae, Cuterebridae
R. D. Gordon—Staphylinindae, Passalidae, Lucanidae, Scarabaeidae,
Meliodae, Coccinellidae
E. E. Grissell—Chalcidoidea
A. B. Gurney, resident cooperating scientist—Orthoptera
J. L. Herring—Hemiptera except Tingidae
R. W. Hodges—Limacodidae, Megalopygidae, Epipaschiinae, Tortricidae, Cos-
sidae, Agonexonidae, Gelechiodea, Oechoporidae, Blastobasidae,
Choreutidae, Sesiidae, Yponomeutidae, Heliozelidae, Gracillariidae,
Lyonetiidae, Psychidae, Nepticulidae, Incurvariidae, Hepialidae,
Erioncraniidae
J. M. Kingsolver—Trogositidae and Agri/us, Bruchidae, Buprestidae, Derodon-
tidae, Colydiidae
J. P. Kramer—Homoptera
P. M. Marsh—Braconidae, Proctotrupoidea
A. S. Menke—Cynipoidea, Chrysidoidea, Scolioidea, Vespoidea, and
Sphecoidea
D. R. Miller—Coccoidea
C. W. Sabrosky, resident cooperating scientist—Tachinidae, Chloripidae
D. R. Smith—Symphyta and Formicidae
T. J. Spilman—Cupedidae, Silphidae, Elateridae, Cerambycidae, Cucujidae,
Melandryidae, Oedemeridae, Lymexylidae, Mordellidae, Lyctidae,
Bostrichidae, Ptinidae
G. C. Steyskal, resident cooperating scientist—Chamaemyidae, Lonchaeidae
M. P. Stoetzel—Aphididae, Adelgidae, Phylloxeridae, Aleyrodidae
F. C. Thompson—Tabanidae, Hippoboscidae, Bibionidae
E. L. Todd—Ctenuchidae, Arctiidae, Noctuidae, Notodontidae,
R. E. White—Chrysomelidae, Cleridae, Anobiidae, Anthribidae, Breniidae
D. R. Whitehead—Rhyncophora
W. W. Wirth—Xylophagidae, Stratiomyidae, Rhagionidae
For verification of scientific names, thanks are also due to S. Nakahara, U.S.
Department of Agriculture, Animal and Plant Health Inspection Service—Thy-
sanoptera; R. C. Froeschner and T. L. Erwin, Smithsonian Institution, Department
of Entomology—Tingidae and Carabidae, respectively.

Vil

Preface

The focus of “Insects of Eastern Forests” 1s primarily on insect pests of trees and
wood products east of 100° longitude, north of Mexico. In a similar manner, the
region west of this meridian is treated in ““Western Forest Insects,”’ by R. L. Furniss
and V. M. Carolin, published in 1977 (U.S. Department of Agriculture Mis-
cellaneous Publication No. 1339) (457). Although the information included in these
books is directed to professional foresters and forest entomologists, others in related
fields and students should find them valuable references.

Insects are not alone in causing damage to trees and wood products. Therefore, it
is important to include other small animals such as certain mites and mollusks that
also injure trees and their products. Common arthropod pests that are found in the
forest environment are mentioned briefly. Many kinds of insects and related
arthropods are beneficial because they suppress populations of damaging species,
so this book also includes information on these groups.

The original treatment of this subject was given in F. C. Craighead’s “Insect
Enemies of Eastern Forests” (U.S. Department of Agriculture Miscellaneous Pub-
lication 657), published in 1950 (26/). This valuable handbook was written in the
late 1930’s but publication was halted by the wartime emergency. When it finally
was made available, DDT was the most promising development for the control of
many forest pests, and a small amount of information on this ““wonder chemical”
was fitted into the text. By the ttme W. L. Baker’s “Eastern Forest Insects” (U.S.
Department of Agriculture Miscellaneous Publication 1175) was published in 1972
(45), DDT was no longer permitted for agricultural and forestry purposes. Empha-
sis shifted from long-lived insecticides to those whose effectiveness was brief and
less harmful to the environment, and even to the development of more sophisticated
systems for regulating pest populations. This is where research and application are
now concentrating.

The popularity of Baker’s ‘Eastern Forest Insects” soon made it unavailable. It 1s
superseded by this volume, which closely follows Baker’s format but provides
additional information that is the result of another decade of research in entomology
and forest pest management.

With the firm conviction that insect classification is the proper responsibility of
specialists, keys to species are omitted. However, the user is helped to the degree
possible by several keys to damage symptoms. The inevitable desire to apply
standardized names to all things led to the biennial publication of approved
common names of insects by the Entomological Society of America (/4, /272).
“Insects of Eastern Forests” follows the protocol of placing accepted common
names before the scientific name and unapproved common names after the scien-
tific names.

The rapidly changing situation for controlling insects mitigates against naming
specific insecticides for this purpose. Current information can be obtained from
agricultural extension specialists.

The biology, life history, and damage of eastern forest insects is emphasized in
the text and illustrations to provide the reader with immediate background informa-
tion required for intelligent decisionmaking. To further assist the reader in this

ix

endeavor, an extensive list of literature citations is included’ as are indexes to the
scientific and common names of the insects and their tree and shrub hosts. Where
possible, hosts follow the nomenclature of E. L. Little, Jr, “Checklist of United
States Trees (Native and Naturalized)” (U.S. Department of Agriculture, Agri-
culture Handbook 541, 1979). A list of corresponding common and scientific
names of plant hosts is also given.

' Italic numbers in parentheses refer to Literature Cited.

Forest Insects

Insects, the most abundant form of life on earth, also are numerically dominant
in the forest. For over a period of some 350 million years they have evolved to
occupy an incredibly wide variety of ecological niches. They serve many functions
in the economy of the forest and are as essential a part of the complex associations
as are the trees themselves (674). Competition between people and insects for the
forest and its products increased with the rise of human population. We demand
much more of the shrinking forest environment now than we did in previous
centuries. So, this competition is an ever-increasing problem that must be under-
stood before it can be resolved.

By no means are all forest insects economically destructive. A multitude of
species are beneficial. Along with other organisms they are important decomposers
of forest debris—dead trees, fallen leaves, and such (542, 543). Thus, they help
return nutrients to the soil, increasing its fertility (306). Others hasten the death of
decadent, diseased, or overmature trees, making way for vigorous younger growing
stock (S3/). Many species of insects are parasites or predators of the destructive
ones (62/). Some innocuous species are hosts for parasites of damaging insects,
enabling the parasite population to remain at times when the population of destruc-
tive insects has declined or is in an unsusceptible stage.

Harmful forest insects and related forms are those that are responsible for
economic loss. They include (1) species that damage or destroy the flowers and
seeds of trees, and are particularly important pests in seed orchards and seed
production areas; (2) species that stunt, deform, or kill nursery stock or young trees
by damaging or destroying the buds, shoots, or roots; (3) species that cause loss of
vitality, growth reduction, and often the death of trees by eating the foliage; (4)
species that feed under the bark or in the wood of living trees and girdle and kill
them or riddle them with tunnels; and (5) other destructive species that bore into
and damage or destroy green logs, storm-felled timber, green-sawed and seasoned
lumber, rustic construction, poles, posts, crossties, mine props, and all manner of
finished products from flooring to furniture.

Most of our forest insects are native to the continent and are usually distributed
throughout the ranges of their hosts. Some are destructive at normal population
levels, but the majority commonly occur in such low numbers as to be of little or no
consequence. A few of the latter, however, are capable of great and rapid increases
in numbers when favorable environmental conditions prevail.

Eastern forests are also inhabited by many species of introduced insects, a few of
which are widely distributed and extremely destructive. These include the gypsy
moth, European pine shoot moth, balsam woolly adelgid, European pine sawfly,
and the smaller European elm bark beetle. Many species of natural enemies of
several introduced pests have also been imported and established in eastern forests.

‘Conditions conducive to forest insect outbreaks are only partly understood. It
appears, though, that outbreaks are most likely to occur (1) in pure stands rather
than in stands of mixed composition, (2) in overmature rather than in immature
stands, and (3) in plantations rather than in natural stands. They may also develop

in stands weakened or decimated by hailstorms, flood, wind, drought, disease, fire,
defoliation, or during logging operations. Outbreaks are likely to occur any time
there is a breakdown in the effectiveness of natural control factors, or when changes
occur in the genetic composition of populations, or in the age, composition, and
density of stands.

Forest insect outbreaks vary greatly in frequency, size, and duration. Fortunately,
the majority are small and short-lived, and usually consist of only one or a few
“spots” in a stand or region. Unfortunately, some may expand until they encompass
a few thousand square kilometers and last for many years before subsiding. From
1977 to 1981, more than 75 different species were recorded in outbreak status in
eastern forests. Some 40 of these are considered to be of major importance (587,
750). Many general discussions on forest insects are available (2/7, 74, 76, 83, 314,
492, 605, 644, 674, 691, 767, 768, 1078, 1196).

Losses Caused by Forest Insects

Insects are among the most destructive agents affecting forest and shade trees.
They are not only responsible for killing valuable trees and for the loss of growth or
recoverable volumes from surviving damaged trees, but also they weaken or destroy
wood structures. They are responsible for serious losses of other kinds, both
tangible and intangible. Management plans are often upset and fire hazards in-
creased, watersheds and wildlife habitats are impaired or destroyed, water in
streams and lakes is polluted, the incidence and severity of floods are increased,
and the attractiveness of parks and other recreational areas 1s reduced or destroyed.
Heavy investments in tree improvement programs also are jeopardized or upset by
the destruction of seeds and cones or valuable seed trees in seed orchards. Stand
composition is often changed, leading to the displacement of valuable tree species
by others of lesser value. Innumerable shade and ornamental trees around homes or
along roadsides, and street trees of towns and cities, are killed or their attrac-
tiveness is greatly reduced or destroyed.

Forest Insect Surveys

It has long been recognized that successful control of forest insect outbreaks
depends not only on the control method used but also on knowledge of the insect
situation in immediate, surrounding areas. When an outbreak is discovered, it is
necessary to know whether it is increasing or decreasing. It is also important to
locate the center of greatest concentration and the direction of spread (262). When
an outbreak occurs, it is also important to Know the value of the resources at stake
and the status of natural control factors before decisions to apply control are made.
Of like importance is the possession of a general knowledge of insect conditions
throughout a timber type or region. This is helpful in developing plans for the
detection of outbreaks in their early stages, a time when they may be suppressed
most easily and economically and before heavy losses are incurred. Similarly, it is
helpful to have a continuing record of the presence and abundance of the less
spectacular types of insects to determine if, when, and where their number or
damage is reaching economic levels of concern, even though not an outbreak.
Where intensive management of the forest for timber or other objectives is being
practiced, it is also important to have up-to-date information on all destructive pests
to allow for their early control where needed. Such information is sought or
acquired through surveys.

2

Forest insect surveys of various types have been made in the United States since
the early days of this century (/37) but, before the passage of the Forest Pest
Control Act in 1947, they were usually conducted on a more or less unorganized
basis. Passage of the act made it possible for the first time to conduct surveys on an
organized, systematic basis on forest lands of all ownerships. In part, the act
authorized the Secretary of Agriculture, either directly or in cooperation with other
Departments of the Federal Government, with State and other public agencies, and
with private organizations and individuals, to conduct surveys to detect and ap-
praise insect infestations before they develop to outbreak proportions, to appraise
their potential destructiveness, and to determine the needs for their control. Section
5 of the 1978 Cooperative Forestry Assistance Act (PL 95-313) retains these same
provisions. Forest insect surveys are also conducted in Canada. There, the aim 1s
not only to detect and appraise infestations but also to obtain information on the
distribution, biology, and cycles of abundance of insect species for taxonomic and
other purposes (756).

Basically, forest insect surveys are of two kinds—detection and evaluation—and
they may be conducted separately or together. Detection surveys are primarily for
the purpose of discovering threatening infestations. Evaluation surveys are usually
far more intensive and complex, being concerned with such factors as the intensity
and the trends of pest populations, the identification and evaluation of natural
control factors, the size and boundaries of infestations, the value of the resources at
stake, and the possibilities of economic loss and an estimate of its magnitude if
suppressive measures are not taken.

Many methods and techniques are employed in conducting surveys, the choice
depending on many factors such as the insect species, its life history and habits, the
nature of the damage it causes, the forest type and terrain, the size and accessibility
of the area, a working knowledge of natural control factors affecting it, and the
availability of trained personnel and funds. Aircraft are widely used in detection
surveys and to a lesser extent in evaluation surveys. Conditions may be recorded by
trained observers riding with the pilot (6, 557, 559) or by aerial photography (4, 5,
108, 212, 296, 556, 558, 1256, 1257). Computers process the data, and radio-
navigational aids keep the craft on the proper course (346, 5/3). Panoramic aerial
photography (2//) and satellite imagery (324) are under study for delimiting
outbreaks. The former is effective and cost competitive with earlier techniques, but
the latter system is expensive and shows only heavy defoliation accurately. Unfortu-
nately, evidence of infestation by many insect species is difficult or impossible to
detect from the air. Where these insects are concerned, the only recourse is to use
ground survey methods (/09, 1339). To the extent that it is possible to use them,
aerial surveys are far less time consuming and costly than comparable ground
surveys.

Ground surveys are not only required to detect infestations of many species but
are essential for evaluation purposes. In most cases, they are complex and require
the services of highly trained personnel. Ground surveys for evaluation purposes
present many different sampling problems, such as when, where, and how to
sample in order to obtain needed information within desired confidence limits. The
choice of procedure is governed largely by the insect species and the intensity and
size of the infestation. Pheromone traps are very useful to detect low populations of
insects (/0/, 1/99). In addition to data on insect population density and size, data
also may be collected to identify and record the abundance of natural control
factors. In all cases information is sought and analyzed to determine two things:

3

what is likely to happen if no action is taken to suppress the infestation, and what
might be expected to happen if suppressive measures are applied.

Sequential sampling techniques provide for flexibility in sample size in contrast
to conventional techniques that usually specify a fixed number of sampling units to
be examined. Units chosen at random from a sample are examined in sequence until
the sample falls into one or more classes distinguished by specified limits (/253).
Sampling plans have been published for several important forest insect pests (234,
23), 031, 032, 630, 101577 156):

Forest Insect Control

The impact of naturally occurring adverse environmental factors on forest insects
is so great that the majority of species never occur in sufficient numbers to
constitute an economic hazard. Even those insects that become sufficiently abun-
dant at times to be considered “pests” are also subject to control by these factors,
though usually to a lesser degree and frequency (62/, 630). Depending on the
insect and the value of the forest resources affected, a given degree of natural
control may reduce damage to an acceptable level.

Natural Control

Natural contro! results when naturally occurring adverse environmental factors
prevent insect populations from reaching or exceeding harmful proportions. It may
be effected by a single factor, such as abnormally low winter temperatures, or by
several factors working in combination; it never ceases to function entirely, but its
impact may fluctuate greatly from place to place and time to time (2/, 492, 62/,
674).

Physical factors.—Temperature is one of the most important physical factors
affecting forest insects. It not only sets limits to their distribution but also often
profoundly affects their abundance. The optimum temperature for many eastern
species appears to lie between 24° and 27° C; outside this range, insect activity
gradually declines and eventually ceases. When temperatures climb to 50° C or
drop to around — 18° C or lower, death results. Rapid changes in temperatures, such
as occur in the fall and spring, also may cause mortality even though absolute lethal
temperatures are not reached.

Insects react in many ways to escape the rigors of adverse temperatures. During
the summmer, leaf-feeding larvae may move from the upper to lower surfaces of
leaves, or from the periphery of the crowns to the shaded interior to escape
excessive heat. Solar-sensitive eggs may be deposited in the shade (340). Adults
find relief from heat by moving into the interior of stands, to the shaded portions of
trunks, or to the interior of crowns. During cooler-than-normal days, they may seek
out individual, exposed trees, the sunny side of trunks, or trees along the edges of
stands. To escape the effects of winter cold, a species may hibernate in the egg,
larval, pupal, or adult stage in areas where the likelihood of lethal temperatures
occurring is greatly lessened. Because of physiological changes that occur in the
dormant state, many northern species are able to survive temperatures well below
—18° C. Southern species are much less likely to survive exposure to such
temperatures.

Outbreaks of leaf-eating insects may be suppressed by late-spring frosts that kill
the young, tender foliage which the young larvae depend on for food. Late springs
may result in greatly delayed emergence from hibernation, and cool summers may
result in a slowdown in the rate of larval development. Prolonged developmental
periods add up to an extended period of exposure to parasites, predators, and
disease pathogens, which may lead to unusually heavy losses. The number of
generations produced per year may be related to photoperiod and the length of the
season of biologically effective temperatures. In the South, a species may produce

5

five or more generations per year, whereas in the North, it may produce only one or
less. Outbreaks of such a species often appear to develop with explosive suddenness
in the South. In the North, it may require several years for an equivalent outbreak to
occur.

Insect populations are also affected by atmospheric moisture and by moisture
conditions in host trees and in the soil. For example, various species of wood
borers, such as powderpost beetles, breed successfully only in very dry wood,
whereas others, such as the ambrosia beetles, require wood with a much higher
moisture content. Damage to black locust by the locust borer and to pines by
various species of bark beetles may be greatly intensified during or following
periods of drought. 3

A considerable degree of control also may be effected by other adverse weather
conditions. For example, heavy, beating rains may dislodge and destroy large
numbers of larvae that feed on the foliage of trees; moist weather may result in the
development of disease epizootics and the decimation of insect populations.

Host factors.—Many species of forest insects feed on only a few species of
trees. Some of the most serious bark beetle pests belong to this group. Many species
of destructive wood borers also confine their attacks to several closely related
species. In contrast, numerous other species feed on a great many tree species,
often in different families. Many leaf-eating species belong to this group. The
females of some insects deposit their eggs in all kinds of places; others deposit them
on host trees only. When the former condition prevails, it often leads to heavy losses
among newly hatched larvae, especially where acceptable food is not readily
available. Infestations of species that develop in weak or decadent trees may
disappear or be greatly reduced when the ratio of these trees to healthy vigorous
trees drops below a critical level in a stand.

Populations of multiple-host species, many of which attack trees in all stages of
health or vigor, may be held to low levels in mixed stands that contain a low ratio of
preferred to nonpreferred host species.

Biotic factors.—Parasites, predators, and pathogenic micro-organisms play
important roles in the natural control of insects. Their effects may not always be
evident, but they always bring some degree of pressure to bear on their hosts, and it
is often severe. Occasionally, when a virus epizootic develops, the results may be
catastrophic for the host insects. Operating singly or in combination, biotic factors
may limit the duration and magnitude of an outbreak, prolong the intervals between
outbreaks, or prevent outbreaks entirely. A large number of these control agents
have been collected and identified in the forests of eastern America, and new ones
are being added to the list each year.

Insect control by biotic factors has several advantages not offered by many of the
other approaches to control now available. Where these factors are well established,
they usually are self-perpetuating, barring natural catastrophies or interference by
humans: they adjust to changes in the size of host populations, and they operate
with practically no adverse side effects.

The majority of the parasites and predators of insects are other insects. Many
thousands of species belonging to well over 200 families in 15 orders are known to
be either parasitic or predacious; many of these attack forest insects. The majority
of parasitic forms belong to only a few families in the orders Hymenoptera and
Diptera. Important predators are found in several families of the orders Coleoptera,
Diptera, Neuroptera, Hemiptera, and Hymenoptera.

Parasitic and predacious insects differ widely in habits and behavior. A parasite
usually requires only one host in which to complete its development; a predator
usually requires several to many hosts for its development. A parasite usually does
not kill its host until it has completed its own development; a predator usually kills
its host as soon as it is encountered. Host selection by parasites is generally a
function of the egg-laying female adult only; whereas each individual predator must
search out hosts in all of the feeding stages.

Forest insects are also fed upon by many other forms of animal life, principally
spiders, mites, birds, and many mammals. Birds, especially woodpeckers, are
sometimes effective in suppressing outbreaks. Many other species of birds are also
thought to play important roles in control (1/90, 470, 564). Small mammals such as
shrews are often effective in controlling species that spend part of their lives on the
ground (/70).

The literature contains many references to the role of parasites and predators in
insect control. A good introduction to the subject is in several publications (26, 48,
IFO S205 218-2403, 284, 621, 1183, 1203, 1213).

Many species of forest insects are also subject to a considerable degree of control
by pathogenic micro-organisms. Some of the latter, especially the viruses in the
family Baculoviridae, frequently occur in the form of epizootics and decimate or
eliminate outbreaks over large areas. The baculo viruses are particularly useful
because they infect only invertebrates. Notable examples are (1) a polyhedrosis
virus disease that ultimately plays a leading role in the control of gypsy moth
outbreaks in the Northeast, and (2) a disease of the same type that caused the
collapse of an outbreak of the European spruce sawfly over a region of several
thousand square kilometers in eastern Canada and the Northeastern States during
the late 1930's.

There are more than 1,300 different kinds of micro-organisms, most of which are
pathogenic, that have been found associated with insects. This includes 22 types of
viral diseases in 826 hosts, and a total of 1,271 host-virus records (8/5), 100
species of bacteria (907), 750 species of fungi (907), 300 species of protozoa (907),
several rickettsiellas, and about 1,000 species of nematodes (99/). Generally
speaking, they gain entry into the insect by being ingested with food, during
hatching from contaminated eggs, through wounds or other damaged areas in the
integument, or through the tracheae. Most of them rely on wind, rain, streams,
healthy or contaminated insects, small mammals, or birds for dispersal throughout
an insect infestation or outbreak. Spread is occasionally so rapid that all of the
insects in an infestation may appear to be dying at the same time.

Many species of forest insects are particularly susceptible to viruses of the
nuclear-polyhedrosis type (873). Once inside the host these viruses enter the cell
nuclei, replicate, and some virions become encapsulated in many-sided, rodlike
crystals, known as polyhedra. Invaded cells are soon destroyed, the virions and
polyhedra are then released in the body cavity, and the host dies. Diseased larvae
usually become siuggish, cease feeding, and in some instances move upward in the
trees. After death they may be seen hanging by their forelegs, with the body
darkened, decomposed, and liquified. Eventually they completely disintegrate and

dry up on the trees.

Forest insects are also affected by several other kinds of viruses—cytoplasmic
polyhedrosis, granulosis, polymorphic inclusion, and noninclusion. Generally
speaking, these viruses appear to be less effective and less specific in control than

those of the nuclear-polyhedrosis type. Further information on viruses is available
(96,215,022, S16) 11455 J:195).

Several species of bacteria produce so-called milky diseases in the insect host,
the best known being the milky disease of Japanese beetle larvae (350). The larvae
of many closely related species of beetles are also affected by this disease. Scores of
species of Lepidoptera and apparently several species of Hymenoptera, Coleoptera,
Diptera, and Orthoptera are susceptible to diseases caused by various
crystalliferous bacteria, the best known of which are several varieties of Bacillus
thuringiensis Berliner (343, 344, 347, 549, 550, 709).

Some species of fungi occasionally cause significant reductions in insect popula-
tions. Aphids are particularly susceptible to attack by species of Entomophthora.
The species E. megasperma Cohn was credited with causing a high degree of
control of the forest tent caterpillar in Ontario. Another species, Beauveria bas-
siana (Balsamo) Vuillemin, reportedly killed more than 90 percent of the larvae of
the smaller European elm bark beetle in a number of infested elm trees in Connecti-
cut (3//). Pathogenic fungi, such as Paecilomyces sp., Beauvaria bassiana, Meta-
rhizium anisopliae, etc., killed nearly one-third of the southern pine beetles in the
field (S67). Laboratory testing has confirmed the activity of these fungi against the
southern pine beetle (954).

Many species of pathogenic Protozoa are also associated with forest insects (652,
1094, 1125). Generally speaking, these pathogens prolong the length of the larval
stage and reduce the fecundity of surviving adults.

Numerous species of nemotodes may be found in the gut, the hemocoel, or in
particular organs such as the Malpighian tubules of insects. Some of these kill the
host when they emerge, some cause death by initiating the action of lethal microbial
disease agents (/265). Some cause injury but not death (9/). Certain species
infesting bark beetles riddle the intestines and gonads of their hosts, block the
ducts, and cause the ovaries to shrivel. This may lead to a marked reduction in egg
production by infested female beetles and a sharp decline in beetle populations
(S26). The literature on the subject has been reviewed (/270).

Applied Control

When natural control factors fail to hold populations to economically tolerable
levels, it may be necessary to apply silvicultural, chemical, or other controls to
enhance, supplement, or serve as substitutes for natural control. Depending on the
insect and its biology and ecological relationships, a wide variety of materials and
methods are available for suppressing populations. These may be applied directly
against the insect to interfere with its growth or reproduction, or to kill it; or they
may be applied to the environment to render it less favorable for the insect through
alteration in food supply, in microclimatic factors, or in the abundance and effec-
tiveness of natural enemies.

The choice of materials or methods in artificial control depends on several things:
whether the insect is a native or introduced species, the factors known or thought to
be responsible for its abundance, the need for quick and effective reduction in its
numbers, and cost. If a native species is involved, usually the aim is not to eradicate
it, but rather to reduce its numbers to a tolerable level; if it is a recently introduced
pest, the aim may be to eradicate it or to slow down or prevent its spread.

Artificial controls designed to suppress existing outbreaks usually have short-
term effects only; those that are used for the prevention of outbreaks may be long

vt

lasting. They may also cause increases in nontarget pest populations (755, 9/2).
Control costs vary considerably, depending on the insect and its habits, the methods
and materials used, and the size and accessibility of infested stands. It may cost
only several dollars per hectare for infested stands treated with aerial application of
insecticides or it may cost several dollars per tree for individual treatments.
Regardless of the control method employed, the benefits from its use should usually
outweigh costs. Different methods of control involve different hazards; care must
be taken in the use of any of these control methods. Occasionally, control costs may
be retrieved through the salvage and sale of infested timber.

Silvicultural Control

Insect population density is influenced by the composition and condition of forest
stands. Because of this, it is sometimes possible to create unfavorable conditions
through the application of carefully designed cultural or management practices.
Efforts directed toward this end are commonly called silvicultural control.

Possibilities of silvicultural control occur during the establishment and through-
out the lives of stands by properly timed planting, by selecting the more resistant
tree species for planting, by putting them on sites best suited for them, or by
controlling their composition and density. Stand conditions may be created or
modified as needed by thinnings or cuttings. Pure stands may be broken up into
mixtures of age classes in small units, with no two contiguous units of the same age
class. Mixed stands may be broken up by cutting in small groups to maintain and
promote diversification in species composition and density. Overmature trees may
be removed from stands and the stands harvested as soon as they mature. High-risk
trees may be removed in sanitation-salvage cuttings (659, /05/). Further discus-
sions of silvicultural control are available (85, 457, 674, 1000, 1139, 1198).

Physical and Mechanical Methods of Control

Many forest, shade tree, and wood products insect pests are amenable to control
by physical or mechanical methods. The method chosen is dictated by such factors
as the habits and behavior of the insect pest involved, the location and value of the
trees or products attacked, and cost.

Trap trees and trap logs are sometimes effective in the control of bark beetles
(903). Bands of tanglefoot or similar materials may be placed around the trunks of
shade trees to prevent larvae or wingless female adults of several species, such as
the fall cankerworm, from ascending the trees to feed or oviposit. Young, newly
planted shade trees may be protected from borers of various kinds by enclosing the
trunks in wrapping paper. Bagworms on ornamental trees may be controlled by
handpicking. Subterranean termites can be kept out of structures by capping
foundations with a 10-cm layer of high-grade concrete or with properly applied
metal shields, by keeping untreated wood from contact with the ground, by using
solid foundation masonry, and by providing good ventilation between the ground
and timbers (/2). Similar treatments prevent damage by the old house borer (870).

Logging and milling infested timber and destroying the slabs that contain the
broods are often effective in preventing bark beetle outbreaks. Such methods may
be feasible only where the trees are of merchantable size, are accessible, and when
they can be harvested promptly. Bark beetle broods in or under the bark may also be
killed by storing infested logs in water, by peeling infested logs and burning the

bark, by sawing infested trees into short lengths and burning them, and by peeling
the bark of infested trees and exposing the brood to desiccation and to natural
enemies such as ants, birds, and small mammals. Felling infested trees in a north-
south direction in the open and periodically rotating them kills the beetles by solar
heat.

Ambrosia beetle damage to summer-cut logs can be reduced or prevented by
quickly removing the logs from the woods, by quickly manufacturing them or
storing them in ponds, by quickly drying the lumber cut from them, and by
removing the bark from rough-hewn pieces.

Ips bark beetle populations that sometimes develop in slash following cutting
operations can be reduced by limiting small-scale cuttings to the fall and winter
months, or by continuing large-scale cuttings throughout the summer months. In
situations where it is necessary to cut on a small scale or discontinuously during the
summer, control may be obtained by piling and burning the slash before the beetles
complete their development in it. Gathering and burning severed branches late in
the fall, winter, or early spring when the eggs and grubs are in the twigs is an
effective method of controlling twig-girdling species.

Wood-borer damage to field-piled pulpwood in the Lake States has been reduced
by piling the wood in the shade or by placing it in standard compact piles. Lyctus
powderpost beetle damage can be reduced by storing susceptible sapwood in water
for a lengthy period or by steaming it at high temperatures. Bark beetle damage to
bark-covered poles and slabs used in rustic construction can be reduced or pre-
vented by cutting the material during the fall and winter and then seasoning it off
the ground and under cover. Damage to young pines by the European pine shoot
moth in the North can be reduced by removing the lower limbs, so that the
overwintering stage of the shoot moth will occur on branches above the snow line
where they will be killed by lethal low temperatures during the winter (/337).

Regulatory Control

Regulatory control is aimed at the prevention of entry and establishment of
foreign plant and animal pests, or at the suppression, containment, or eradication of
such pests as may have become established in limited areas. This form of control
became possible with the passage of the Federal Insect Pest Act in 1905, which
enabled the Federal Government to regulate the importation and interstate move-
ment of articles that might spread insect pests. It was reinforced in 1912 by passage
of the Plant Quarantine Act, which authorized the Secretary of Agriculture to
enforce necessary regulations to protect the agricultural ecomomy of the United
States by preventing the introduction of insects and plant diseases. Later the
McNary-McSweeney Act of 1928 established a Federal policy on the use of
legislative means for combating forest insects and diseases. In 1947, passage of the
Forest Pest Control Act provided authority for the United States Government to act
alone or in cooperation with States, territories, or private timber owners to control
destructive forest insects or diseases.

Before the passage and enforcement of quarantine laws, many species of insects
associated with forest, shade, and ornamental trees, several of which are highly
destructive, gained entry into the Eastern United States. Since then the rate of entry
of additional species has been drastically reduced. The following is a list of species
known to have been introduced into this area:

10

Order and Species

Orthoptera

Gryllotalpa gryllotalpa (L.)
Hemiptera

Stephanitis rhododendri Horvath
lsoptera

Coptotermes formosanus Shiraki

Homoptera

Adelges abietis L.

A. piceae (Ratzeburg)

A. laricis Vallot

Asterolecanium variolosum (Ratzeburg)
Cryptococcus fagisuga Lindinger
Gossyparia spuria (Modeer)
Matsucoccus resinosae Bean & Godwin
Pealius azaleae (Bean & Moles)
Phenacoccus acericola King

Pineus strobi (Hartig)

Pseudococcus comstocki (Kuwana)

Quadraspidiotus perniciosus (Comstock)

Coleoptera
Anobium punctatum (De Geer)

Anomala orientalis Waterhouse
Brachyderes incanus (L.)
Callirhopalus bifasciatus (Roelofs)
Chlorophorus annularis (F.)

Chrysomela interrupta F.

C. crotchi Brown

Cryptorhynchus lapathi (L.)
Cyrtepistomus castaneus (Roelofs)
Heterobostrychus aequalis (Waterhouse)

Hylotrupes bajulus (L.)
Maladera castanea (Arrow)
Minthea rugicollis (Walker)

Nacerdes melanura (L.)

Otiorhynchus ovatus (L.)
O. sulcatus (F.)

Origin

Europe

Europe

Far East

Europe
Europe
Europe
Europe
Europe
Europe

N

Asia
Europe
Europe
Far East

China

Europe

Philippines
Europe
Japan
Japan, India

Europe
Europe
Europe
Japan
Far East

Europe
Orient
Antilles(?)

Europe

Europe
Europe

Principal food
plant

Nursery stock

Rhododendron

Wood
products

Spruce
True firs

Red pine

Azalea

Sugar maple

Pine

Deciduous
trees

Deciduous
trees

Wood
products
Nursery stock

Pine
Ornamentals
Stored
bamboo
Willow, poplar
Willow, poplar
Poplar
Oak
Oak &
mahogany
boards
Seasoned
coniferous
wood
Deciduous
trees
Oak flooring
Wooden
wharves,
piling
Conifers
Confers,
broadleaf
plants

Phyllobius oblongus (L.)

P. intrusus Kono

Plagiodera versicolora (Laicharteg)

Polydrusus impressifrons (Gyllenhall)

Popillia japonica Newman

Pyrrhalta luteola (Muller)

Rhizotrogus (= Amphimallon) majalis

Sciaphillus asperatus Bonsdort

Scolytus multistriatus (Marsham)
Xestobium rufovillosum De Geer

Zengophora scutellaris Suffrian

Lepidoptera

Di

12

Aethes rutilana (Hubner)
Anthophila pariana (Clerck)
Archips rosanus (L.)

Chrysoclista linneella Clerck
Cnidocampa flavescens (Walker)
Coleophora /aricella (Hubner)

C. ulmifoliella McDunnough

C. serratella (L.)

Dichomeris marginella (Denis &
Schiffermuller)

Epinotia nanana (Treitschke)

Etainia sericopeza (Zeller)

Euproctis chrysorrhoea (L.)

Gracillaria syringella (F.)

Homadaula anisocentra Meyrick

Leucoma Salicis (L.)
Lymantria dispar (L.)
Orgyia antiqua (L.)

Rhyacionia buoliana (Denis &
Schiffermuller)
Samia cynthia (Drury)

Sesia apiformis (Clerck)

Spilonota ocellana (Denis &
Schiffermuller)

Zeuzera pyrina (L.)

otera

Monarthropalpus buxi (Laboulbene)

Rhabdophaga Ssalicis (Schrank)

Europe

Japan
Europe,
Japan
Europe
Japan

Europe
Europe
Europe

Europe
Europe

Europe

Europe
Europe
Europe

Europe
Asia
Europe,

Japan
Europe
2

Europe
Europe
Europe
Europe
Europe
2

Europe
Europe
Europe

Europe
Asia

Europe

Europe

Europe
Europe

Deciduous
trees
Thuja, juniper

Willow

Poplar

Wide variety of
plants

Elm

Various plants

Maples, birch,
hazel,
hophorn-
bean

Elm

Wood
products

Poplar

Juniper
Apple, thorn
Deciduous
trees,
privet
Linden
Norway maple

Larch
Elm
Birch

Juniper
Spruce
Norway maple
Apple, pear
Lilac
Mimosa,
honeylocust
Poplar, willow
Oak, etc.
Deciduous
trees

Pines

Ailanthus,
cherry

Poplar

Elm, maple

Boxwood
Willow

Hymenoptera

Acantholyda erythrocephala (L.) Europe,
Japan Pine

Caliroa cerasi (L.) Europe Cherry,

shadbush
Diprion similis (Hartig) Europe Pine
Eriotremex formosanus (Matsumura) Asia, Taiwan Oak
Fenusa dohrnii (Tischbein) Europe Alder
F. pusilla (Lepeletier) Europe Birch
F. ulmi Sundevall Europe Elm
Gilpinia frutetorum (F.) Europe Pine
G. hercyniae (Hartig) Europe Spruce
Hemichroa crocea (Geoffroy) Europe Alder
Heterarthrus nemoratus (Fallen) Europe Birch
Neodiprion sertifer (Geoffroy) Europe Pine
Pristiphora erichsonii (Hartig) Eurasia Larch
P. geniculata (Hartig) Europe Mountain-ash
Profenusa thomsoni (Konow) Eurasia Birch
Sirex juvencus juvencus (L.) Europe Pine, fir,

spruce
Solenopsis invicta (Buren) South

America General

feeder

Trichiocampus viminalis (Fallen) Europe Poplar

Biological Control

Biological control, in the classical sense, implies the introduction of alien
parasites, predators, or disease organisms to suppress pest populations below an
economically damaging level. The forest environment is the most stable plant
community; therefore, it should be admirably suited to the practice of biological
control. In fact, a number of introduced forest insects have been subjected to this
type of control with great success, and some others with less success. The exposed
feeders—defoliators and sucking insects—are most readily suppressed; insects that
feed cryptically—such as bark beetles and borers—are least likely to succumb to
biological control.

Success in this field of endeavor is favored by proper packaging, rapid transpor-
tation, good communication between shipper and receiver, and also among
customs, quarantine, and postal officials. Contrarily, poor communication between
parties, inefficient customs, quarantines, and postal services, may ruin biological
control attempts. Political strife between nations may also interfere with the con-
duct of biological control operations.

Generally speaking, biological control efforts against forest insects have been
limited to (1) the importation and establishment of foreign parasites and predators
of introduced pests; (2) the transfer of parasites, predators, and disease pathogens
from one region to another within the country; (3) the augmentation of established
parasite and predator populations with field-collected or laboratory-reared indi-
viduals; and (4) the use of microbial sprays to control outbreaks.

The majority of importations of parasites and predators made before World War
‘Il emphasized enemies of the gypsy moth, browntail moth, satin moth (/73, 1/74,
6/3), European pine shoot moth, European pine sawfly, European spruce sawfly,
balsam woolly adelgid, and larch casebearer. Since then, importations have been
limited mostly to enemies of the gypsy moth, balsam woolly adelgid, larch

13

casebearer, and the smaller European elm bark beetle. By 1982 a total of 264
species of parasites and predators had been imported and liberated against 60
species of introduced and native pests. Of these, a total of 50 are known to be
established. Beginning in the early 1930’s, the Canadian government also imported
large numbers of many species of parasites and predators against a number of pests,
many of which also occur in the United States (227, 788). Some of these biological
control agents have spread into adjoining areas of the United States. Large numbers
of others have been shipped to this country and liberated in infested stands.

A polyhedrosis virus disease of the gypsy moth accidentally introduced into the
United States in the early 1900's has since played an important role in the control of
outbreaks of its host. Like others of its kind, this virus is most effective in dense
populations and is almost unnoticeable in light infestations. A microbial spray
formulation of this virus has been developed and registered under the name
Gypchek (732).

Applications of polyhedrosis virus sprays have been used to suppress populations
of the European pine sawfly (//2), the Virginia pine sawfly (790), the Swaine jack
pine sawfly (/095), and the redheaded pine sawfly (292, 1/45). Virus epizootics
have been initiated in populations of the forest tent caterpillar by disseminating the
virus during one generation (//44). Other important eastern forest insects that are
subject to polyhedrosis virus diseases are European spruce sawfly, eastern tent
caterpillar, fall webworm, linden looper, whitemarked tussock moth, winter moth,
jack pine sawfly, and the blackheaded pine sawfly.

Viruses are usually applied as sprays, but they may also be applied as dusts if
first incorporated with powder. Sprays may be applied by hand-operated sprayers,
mist blowers, or aircraft. Experimental vectoring with contaminated parasites or
males may simplify distribution of viruses in the future.

Considerable progress has been made in the control of a number of lepidopterous
defoliators with Bacillus thuringiensis. The HD-1 isolate of B. thuringiensis is now
used commercially against many forest insect defoliators, including gypsy moth,
spruce budworm, Douglas-fir tussock moth, tent caterpillars, webworms, and
cankerworms.

Chemical Control

The use of chemicals to suppress forest insect populations is usually a method of
last resort. It should be the practice to use them only when other forms of control,
either natural or artificial, fail or threaten to fail in the prevention or control of
destructive populations. Chemical insecticides sometimes may complicate the reso-
lution of an insect problem (642,755). Depending on the situation, chemicals may
be applied to a single tree or to forested areas covering thousands of hectares or
square kilometers. The aim, therefore, is usually limited to the suppression of
injurious populations to tolerable levels.

Many different types of equipment and techniques are available for applying
insecticides (993). Aerial applications are made to large areas by fixed-wing
aircraft (fig. 1) or helicopters (5/, 59, 90, 354, 627, 641). Individual trees, small
groups of trees, and seed orchard trees (fig. 2) may be treated by ground equipment
such as mist blowers (290, 994) or by knapsack sprayers. Large individual trees
may be treated by mist blowers and by hydraulic sprayers (S38). Logs are treated by
knapsack and power sprayers (/040). Fogging machines are sometimes used around

14

resorts and campgrounds. Systemic insecticides may be applied by trunk implanta-
tion or injection (837, 538), by banding or spraying the circumference of the trunk,
by spraying the foliage, by treating the soil around the base of trees (55, /364), or
by dipping seedlings or cuttings before planting (57, 320, 703, 537, 874, 940,
EST):

F-482299
Figure 1.—Airplane-spraying for spruce budworm
control.

“
x

F-517833

Figure 2.—Mist-blower application of insecticide to pine
trees in a seed orchard.

15

The effectiveness of chemicals in suppressing many forest insect populations has
been amply demonstrated. A large number of new insecticides and methods of
application have come into extensive use, and outbreaks covering thousands of
square kilometers of forest have been suppressed. As effective as insecticidal
control has been, however, it has not proved to be an unmixed blessing. Chemical
insecticides have been associated with known or suspected adverse side effects
caused by some of the more commonly used chemicals. This stems from the fact
that these chemicals are nonspecific and may be harmful to at least some other
exposed animal species (254). The problem is compounded in that some chemical
insecticides are very persistent (953, /358), that all of the material applied in a
given environment may not remain there, and that free-ranging animals cannot be
excluded from sprayed areas (560). The deleterious effects of these insecticides on
food chains are known by the failure of carnivorous birds to reproduce (585) and by
the appearance of fish unfit for human consumption. Public awareness of known
and possible hazards associated with the use of these chemicals has given rise to
concern over their continued widespread use.

Efforts are unceasing to discover and develop new and safer insecticides (/00S).
Insecticidal accumulations in the tissues of wildlife need to be determined as a basis
for indicating when undesirable damage to the biota might occur. Application
techniques and equipment are being refined to provide better control of the place-
ment of insecticides in the environment and to further lessen the dangers of
undesirable side effects. Studies are being made to improve sampling and biolog-
ical evaluation techniques as a basis for improving the timing of application and to
ensure that insecticides are applied in the proper amount, and only when and where
they are needed. Joint genetic and toxicological studies should lead toward efficient
application and minimal environmental contamination (//66, 1167).

The rapid changes occurring in the development and use of insecticides in forest
insect control make it inadvisable to include control recommendations in this
publication. Furthermore, no insecticides should be used to control an insect unless
recommendations are registered by the Environmental Protection Agency (/225).
When using any pesticide, remember to: (1) read the entire label before opening the
container, (2) meticulously follow application and disposal directions, and (3) be
certain to consult your County Agent or another specialist regarding any use not
explicitly mentioned on the label (1/225, 1242, 1273).

Other Approaches to Control

Many conventional methods of suppressing forest insect populations are effec-
tive, but they are all found wanting in important respects. Insecticides are often
entirely effective, but they usually cannot be applied to large areas without en-
dangering other forms of animal life. Furthermore, they usually do not hold pest
populations to subeconomic levels for long, nor do they reduce the vulnerability of
stands to future outbreaks. Most forms of biological control cannot be depended
upon to suppress an outbreak before damage has occurred. They may also be
incapable of preventing outbreaks, although they may reduce their frequency of
occurrence and their magnitude, intensity, and duration. Because of these deficien-
cies, aS well as those of other current methods of control, intensive research 1s
underway to develop new or improved methods and materials which may be used as
complements, supplements, or substitutes for them.

16

Sexual sterilization is receiving much current attention because of its potential
use as a new method of control. The effectiveness of releasing males sterilized by
gamma radiation was established in the mid-1950’s when they were used to
eradicate the screwworm, Cochliomyia hominivorax (Coquerel), from Curacao,
West Indies, and from the southeastern portions of its range in the United States.
The principles of sexual sterilization of insects are discussed in several papers (676,
677, 678, 679, 680). Several authors reported the research that led to the method
and its use against the screwworm (/79, /S0, 18/1, 734, 735). Research on the
sterilization of some important forest insects has not been promising (5/, 637).
However, in new work on the gypsy moth, use of the sterile-male technique with
wholly or partially sterile males, shows good promise for eliminating isolated
populations. Partially sterile males produce sterilized F, generations (/073). Further
research must be conducted before the full possibilities of using the sterile-male
release technique for forest insect control can be determined.

Chemically induced sterility also offers promise as a control measure. The
appropriate chemicals are called chemosterilants because of their capacity to de-
prive insects of the ability to reproduce (/39, /0/8). Most of the chemosterilants
that offer promise for practical control are mutagenic and can be used only in ways
that will avoid all contact between them and nontarget animals (907). Chemo-
sterilants may be added to the insects’ food or applied to surfaces to which the
insects are attracted. Those mixed with food appear to produce maximum effective-
ness and minimum hazard. So far, none is available for use in forest insect control.

Insect repellents can be useful alternatives to the use of chemical insecticides.
Considerable research, development, and use of repellents is associated with insects
and related animals of importance to public health. However, relatively little
emphasis has been placed on insects that damage trees or their products.

Tests of organotins indicate that they deter feeding by the pales weevil (//97).
Certain products of the neem tree also show deterrent capability (/82, 708). Pine
oil, a by-product of sulfate pulping, shows promise as a repellent of several
Dendroctonus spp. (931).

Insect attractants are being given increased research attention to improve or
reduce the cost of surveys and to open up new opportunities in integrated pest
management of forest insects (99, 1/86). Some of these materials are powerful
enough to lure insects over considerable distances. Attractants of many kinds are
being studied, but those consisting of natural chemicals produced by host plants or
the insects themselves are receiving the most attention.

Many important forest insects in the orders Lepidoptera, Coleoptera, and
Hymenoptera produce sex attractants. They may be emitted directly from the
insect, or they may emanate from the frass or excrement left from insects feeding
under the bark (/0S2). The material emitted by certain species 1s enormously
attractive. One female of the introduced pine sawfly attracted more than 11,000
males (239).

Sex attractants have long been used to some extent in surveys of a few forest
insects, particularly the gypsy moth (225). For many years the tips of abdomens of
virgin female moths were placed in specially designed traps to capture male moths.
Research to improve on this technique led to the isolation and synthesis of the
attractant material (634), and the subsequent synthesis of a closely related, com-
paratively inexpensive, highly attractive material called disparlure (/06). Synthetic
attractants are now used widely for detecting and evaluating gypsy moth as well as
other forest insects, such as Douglas-fir tussock moth, Nantucket pine tip moth,

17

European pine shoot moth, introduced pine sawfly, western pine shoot borer
(Eucosma sonomana Kearfott), and spruce budworms.

Beginning with the isolation, identification, and synthesis of the pheromone of
the western pine beetle (Dendroctonus brevicomis LeConte) (/082), aggregating or
sex pheromones of a number of bark beetles important to forestry have been isolated
and identified. These include compounds from four species of /ps and five species
of Dendroctonus. In most instances, a complex of two or more substances has been
shown to exist. Individual chemicals in a particular species complex may be
produced by either males or females and have highly specific behavioral functions.
Among these functions are sex attraction, aggregation, antiaggregation, attack
termination, and species isolation. In the case of both the western pine beetle and
the elm bark beetle, the synthesized form of attractant/aggregant has been evaluated
in large-scale field studies. A broader coverage of progress and developments is
available (1/0, 633).

Several other approaches to control are also being investigated. These include the
selection and breeding of hybrids and varieties of trees resistant to attack by a
number of important insects (472, 849, 1/114), as well as the discovery or breeding
of more vigorous strains of natural enemies, the introduction of detrimental genetic
traits into pest insects, and the identification and isolation of chemicals responsible
for host resistance or attraction (///6, 1/17, 1118). Toa great extent, knowledge of
the enzymatic composition of pest and parasite insects is basic to understanding
success with alternatives to chemical control. The value of such studies will be
recognized increasingly (200). Studies are also being conducted on the use of insect
growth regulators that interrupt larval development by preventing the normal
molting process. Laboratory and field tests show that these materials have great
promise (/019, 1020, 1031, 1286).

Integrated Pest Management

The publication of “Silent Spring” (/99) in 1962 focused the attention not only
of Americans but also of people around the world on the serious question of the
continued use of broad-spectrum pesticides that persist in the environment. Within
a few years the use of many pesticides of this type was either banned or severely
restricted worldwide. As a result of all this, an urgent need for alternative materials
and methods was made sharply evident. Although many associated problems are
yet to be solved, modern instrumentation, analytical chemistry, and essentially
artificial diets for insects, have strongly supported the advances that are in progress.
The twin goals of sophisticated systems to suppress populations of threatening pests
are now based upon the application of a range of techniques designed to maximize
pest management within economic tolerance and minimize or eliminate environ-
mental trauma. The term “Integrated Pest Management” is designed to identify
these systems. Therefore, for the purpose of definition, integrated pest management
(IPM), is a system that utilizes all suitable techniques and methods of pest preven-
tion and suppression, in the context of good ecological practice, to maintain pest
population levels below those causing economic injury (3/3, 1195).

Most of the important pest species in eastern forests lend themselves to some
form and degree of IPM. The long life of the forest crop, and the fact that many
species of commercially valuable trees can withstand some degree of infestation
without serious damage, provide the opportunity to use different methods of
management in the majority of cases.

18

Since early in this century, considerable emphasis has been placed on importing
and augmenting natural enemies of many species of introduced pests (2/8, 6/3). In
many cases, the dispersal of these imported enemies has been accelerated by the
transfer of colonies to distant infested areas (299, 34/, 704). The increase in
numbers after successful establishment of several species has also been helped by
the use of large numbers of field-collected or laboratory-reared individuals (647,
928).

The presence and abundance of natural enemies is taken into consideration in
deciding whether to suppress an outbreak and in planning how to accomplish it. For
example, the sudden appearance of a virus disease among the damaging insects in
heavily infested stands usually portends the imminent collapse of the outbreak.
Therefore, when evidence of this disease is encountered during surveys, it is often
decided to withhold other methods of control, particularly the application of
insecticide. Observation of abnormally high percentages of natural control provided
by insect parasites and predators leads to a similar decision. In other situations
where insecticidal control is deemed necessary, it is often possible to limit applica-
tion to designated portions of infested areas because of the abundance of natural
enemies in other portions (673).

Knowledge of the population dynamics of a pest species provides opportunities
for various other means of preventing or reducing losses, thereby eliminating the
need for direct, suppressive measures to control outbreaks. The genetic basis for
insect behavior is being explored to suggest and explain responses to pest manage-
ment practices (428). Depending on the situation, outbreaks of certain leaf-feeding
insects may be prevented or minimized by modifying the composition and density
of susceptible stands (3/3, 75/). It is often possible to prevent or suppress out-
breaks of some bark beetles by thinning the stand, removing infested slash, or
salvaging infested trees before the emergence of beetle broods from the bark
(1195). The utilization of mature trees growing on poor sites before the trees begin
to deteriorate and become attractive to borers, such as the hemlock or bronze birch
borers, is a recommended control practice. The avoidance of unfavorable sites in
planting programs is recommended for preventing damage by certain species (5/5,
548). Adherence to good construction methods when building structures reduces
the need for chemical control of termites later (JO50). The destruction of elm
material suitable for breeding by the smaller European elm bark beetle reduces the
need for the use of insecticides in Dutch elm disease control (/287).

Some of the new approaches to biological control, many of which are yet to be
perfected, may be expected to provide additional opportunities for IPM in the
coming years. These include the combined use of insect growth regulators or
microbial insecticides with pheromone traps to rid the sprayed zone of remaining
male insects. Application of these new-generation insecticides can be followed by
the liberation of massive numbers of parasites, some species of which are known
not to overwinter in the region. Comprehensive treatments of insect-pest manage-
ment have been published (3/3, 664, 907, 1008, 1195).

19

Insects and Tree Diseases

Insects are involved directly or indirectly in the transmission of many serious
diseases of forest and shade trees in the Eastern United States. Some of these
diseases, such as Dutch elm disease and elm phloem necrosis, are among the most
devastating tree diseases known. The majority of tree diseases are caused by various
species of fungi. However, phanerogams, bacteria viruses, mycoplasmas, and
rickettsiae also cause plant diseases. Some disease organisms, such as those
causing Dutch elm disease and chestnut blight, attack living, healthy trees and kill
them quickly. Other diseases, such as stains and heartrots, occur in living trees
weakened or injured by environmental conditions.

Fungal diseases can be transmitted by insects through the direct transfer of spores
from infested to healthy trees. The bodies of insects emerging from diseased hosts
become contaminated with the spores and when the insect attacks healthy trees,
spores are introduced into the wounds. Viral, mycoplasmal, and rickettsial organ-
isms are most often directly transmitted by species of sucking insects, such as
leafhoppers. Wounds made by insects can indirectly serve as courts of entry for
wind- and rain-borne spores.

Dutch elm disease is caused by the introduced fungus, Ceratocystis ulmi
(Buisman) C. Moreau. The disease was first recorded in North America in Ohio in
1930, and by 1975 had been reported in all of the contiguous 48 States except
Arizona, Florida, Louisiana, Nevada, New Mexico, and Utah (/069). Both native
and exotic species of elm are hosts for the disease, with the common American elm
as its most susceptible host. Principal insect vectors are the smaller European elm
bark beetle and the native elm bark beetle (224, 274). Other insects carry spores of
the fungus, but the life cycle of the elm bark beetle is ideal for disease spread. Adult
beetles emerging from diseased trees carry spores to healthy parts of the same tree
and to adjacent healthy trees where the beetles feed and lay eggs. The spores
germinate in the feeding wounds and initiate new infections, which weakens the
tree and aids the larval feeding (/09/). Much of the effort to control Dutch elm
disease has involved insect control. Early efforts were made by using insecticides.
Sanitation efforts have been more long lasting by reducing the opportunities for
beetles to move the fungus. However, the common occurrence of elms has made
this a costly method (970, 1069, 1097).

The first symptoms of Dutch elm disease are wilting and yellowing or drying of
foliage. Defoliation and death of the affected branches soon occurs. Diseased trees
commonly die gradually, branch by branch, over a period of several months or even
years. A brown discoloration in the water-conducting vessels develops in affected
trees. In early spring, this occurs as brown streaks in the wood just under the bark of
diseased branches (fig. 3). Later in the growing season, it appears as brown spots or
as a partial or complete brown circle in one or more outer rings of the wood.

Elm phloem necrosis is a disease of American, slippery, cedar, winged, and
September elms. It occurs in parts of Ohio, Indiana, Hlinois, Missouri, Lowa,
Nebraska, Kansas, Oklahoma, Arkansas, Mississippi, Alabama, Georgia, Ten-
nessee, Kentucky, West Virginia, Pennsylvania, New York, and New Jersey. Evi-
dence indicates the disease is not caused by a virus, as earlier suspected, but rather

20

by a mycoplasmal organism (/3/5). Mycoplasmas are transmitted through root
grafts, or by the whitebanded elm leafhopper (43) that is widely distributed in the
Eastern United States. The leafhopper ingests the mycoplasma while feeding on the
sap of a diseased tree. Later, the disease organism is transmitted when the insect
feeds on a healthy tree.

Courtesy G. N. Lanier, SUNY Coll. Environ.
Sci. & For., photo by George Snyder

Figure 3.—Elm twig with streaking symptomatic of Dutch
elm disease.

Foliar symptoms of elm phloem necrosis vary but are somewhat similar to those
produced by drought, girdling, and certain other tree diseases. Symptoms may first
appear on a single branch or a portion of the top, but most often the entire tree is
symptomatic. In large trees, there 1s usually an apparent thinning of foliage in the
top or at the outer tips of branches. This is followed by a general thinning of foliage
throughout the crown. Generally, the leaves become chlorotic but some leaves
become dry and brown. Defoliation and tree death occasionally happen within 3 to
4 weeks but it usually requires from | to 1’ years. Before the tree dies, a typical
yellow to butterscotch discoloration of the phloem develops (fig. 4). In large trees
this discoloration is usually found only in large roots and the lower part of the trunk.
In most small trees, and occasionally in large ones, it may be found in the upper
part of the trunk and in branches. The butterscotch-colored phloem of American
elm with elm necrosis has a faint odor of wintergreen, especially if the plant part is
enclosed briefly in a bottle.

During ephiphytotics such as occurred in the Central States in the late 1930’s and
1940’s, elm phloem necrosis killed numerous valuable shade tree elms in many
cities and towns (fig. 5). During the 1950’s the disease was at a low ebb, but more
recently the disease is again on the increase in New York, Ohio, and Pennsylvania.

Beech bark disease is a destructive disease of American and European beeches
and all their varieties. It is caused by related fungi, Nectria coccinea var. faginata
Lohman, Watson, & Ayres and N. galligena Bres. (250). These pathogens gain
entry into the tree through tiny ruptures in the bark caused by the feeding of the
beech scale. The scale and probably the fungus are of foreign origin. The disease,

21

which was first recorded at Halifax, Nova Scotia, in 1920, now occurs throughout
the Maritime Provinces of Canada, west to Quebec, and south through the beech-
growing areas of New England to West Virginia (847, /080). Surveys indicate that
the disease is moving south and west at a steady rate.

F-520108 F-519951
Figure 4.—Typical discoloration Figure 5.—Shade tree elms killed by elm
of the phloem of an elm tree phloem necrosis mycoplasm.
infected with elm phloem

necrosis.

The fungus may infect large areas on scale-infested trees, completely girdling
and killing them. Trees only partially girdled may remain alive in a weakened state
for many years unless broken by the wind. On some trees, the fungus 1s confined to
strip or spot cankers on the bole. Parts of the crowns of these trees become chlorotic
and die.

In many stands, beech bark disease has killed more than half the beech, and the
commerical value of many of the survivors has been seriously reduced. Disease
control has not been attempted; the presence of disease-free trees in the midst of
heavy infestations suggests resistance in the host population (/080).

Oak wilt disease, caused by the fungus Ceratocystis fagacearum (Bretz) Hunt,
occurs Over a wide area from Nebraska and Kansas to the Carolinas and Pennsylva-
nia. The fungus attacks all native species of oak regardless of size, age, or vigor.
Local spread from infected trees to adjacent healthy trees occurs through natural
root grafts; insects, however, apparently are responsible for spread over longer
distances. Oak bark beetles, Pseudopityophthorus spp., are recognized as being the
most important vectors of the oak wilt fungus (/68, 506, 1021, 1022). In areas of
the country where mycelial mats of the fungus develop on diseased trees, species of
sap-feeding and mycophagus nitidulid beetles probably are vectors of the fungus
(323, 811).

Oak wilt is potentially a very serious disease (fig. 6). Given the proper conditions
for an epiphytotic, it could cause heavy losses in the vast oak forests of the Eastern

22

United States, but it has not, probably because the vectors are not efficient
transmitters of the disease. It has, however, caused considerable damage in many
oak woodlots and forest stands in Wisconsin, Minnesota, West Virginia, and lowa
(438). Control measures include sanitation by removal of infected trees and trench-
ing between infected and healthy trees to eliminate root grafts.

Persimmon wilt is a fast-killing disease of common persimmon that occurs in
the Southeastern States west to Texas and Oklahoma. It is caused by the fungus,
Cephalosporium diospyri Crandall, which produces masses of spores beneath the
bark of infected trees. In smooth-barked trees the spores occur in such large masses
that the overlying bark is raised in the form of blisters. When these blisters break,
the spores are released and then blown away by the wind. In rough-barked trees the
spores are produced in the cambial region and are released when the bark begins to
disintegrate or is removed or broken off.

Wind is undoubtedly the major agent of spore dissemination, but some spread is
also probably effected by insects such as the bostrichid, Xylobiops basilaris (Say).
The adults of this species, some of which attack healthy trees, become contami-
nated when they emerge through spore masses on dying trees. Feeding injuries on
the terminals and twigs by adults of this twig girdler also serve as entry courts for
windborne spores (263).

Fungi that cause blue stains are introduced into pines by bark beetles, and the
fungi kill trees (65, 9/8). Blue stain occurs in felled timber, in beetle-infested trees,
and in trees weakened by fire, drought, or other adverse factors.

F-519913

Figure 6.—Oak tree killed by the oak wilt
fungus, Ceratocystis fagacearum.

23

Ceratocystis minor (Hedgc.) Hunt, one of the most important of the blue-stain
fungi in the Eastern United States, is introduced into pine trees by the southern pine
beetle. Blue-stain fungi interfere with normal water movement in the tree, and the
tree shows wilt symptoms as it dies. The presence of blue stain in wood reduces its
market value.

Ceratocystis ips (Rumbold) C. Moreau is introduced into southern pines by the
engraver beetles, /ps calligraphus and I. grandicollis, and it produces a stain that
spreads inward from the beetle galleries toward the heartwood (fig. 7). This fungus
has also been found in living and felled red pines attacked by /. pini and /.
grandicollis in the Lake States (7/9).

Several authors discuss the interrelationships of bark beetles and blue-stain Sie
(146, 917, 1044). A number of species of ambrosia beetles carry staining fungi on
their bodies and, because they attack hardwood logs and lumber in large numbers,
are considered to be important disseminators of stain-inducing agents (/23/).

The cyclic occurrence of pitch canker disease of slash and loblolly pines

suggests that it is dependent on insects for spread (35/). Pitch canker disease is
eaiced by Fusarium moniliforme Sheld. var. subglutinans Wollenw. & Reink. Tip
moth damage and pitch canker damage are sometimes associated. The deodar
weevil feeds on young shoots in the upper crown of slash and loblolly pines,
creating wounds, which are colonized by the pitch canker organism. Since spores of
the fungus are wind- and water-borne, any insect-induced wound is probably used
as a court of entry by the fungus.

Courtesy Duke Univ. Sch. For.

Figure 7.—Cross section of bole of shortleaf pine killed
by Ips calligraphus, showing development of blue-
stain fungus, Ceratocystis ips, in sapwood.

24

Insects and Related Organisms

No discussion of organisms that damage or destroy trees or forest products is
complete without reference to some species of mollusks and arthropods other than
insects. A number of others in the hierarchy of arthropods, such as crustacea,
millipedes, centipedes, scorpions, spiders, mites, ticks, etc., are commonly en-
countered in woodland settings, so they are included in this book. Some species of
mites, in particular, may damage, even destroy trees, while others are beneficial as
predators of pest insects.

Phylum Mollusca—Shipworms

Wood submerged in salt water is attacked by several species of bivalve mollusks,
commonly known as shipworms. These relatives of clams, mussels, and oysters can
cause severe damage to pilings, especially along the Gulf Coast and along the
Atlantic Coast south of the Chesapeake Bay. Boats are also damaged occasionally.
These organisms are discussed briefly because the damage they cause is similar to
and often confused with that caused by wood-boring insects.

The genera 7eredo and Bankia contain the so-called shipworms. Their bodies are
long and soft. They are armed with small, chisellike shells at the anterior end and
there are two siphon tubes at the posterior end. They are free-swimmers in the early
larval stage, and during this period the body is protected by a bivalve shell. Free-
swimming larvae seek out wood and attach themselves to it near the mud line. Then
they bore into the wood, leaving very small openings to the outside. Once inside
they develop rapidly, enlarging and lengthening their tunnels as they grow. During
this period when the shell is no longer needed for protection, it is used as a boring
tool. As they develop, shipworms secrete a calcareous material with which they line
their tunnels. The openings made by the young larvae are never enlarged. As a
result, the only external signs of infestation of a piece of heavily damaged wood are
tiny holes in its surface.

Because a single piling may contain several thousand shipworms, it may be
literally honeycombed (fig. 8). In such situations, tunnels may be no more than 6
mm in diameter and only a few centimeters long. Under the most favorable
conditions, however, the tunnels may reach a diameter of 2.5 cm and a length of 1.2
m.

The genus Martesia also contains a number of destructive species. They differ
from the shipworms in being clamlike. The young are also free-swimmers and seek
out and attack submerged wood, making small, inconspicuous entrance holes.
Once inside the wood, they grow until they are approximately 25 mm in diameter
and 60 mm long. As they grow, they enlarge their cavities to accommodate their
bodies. Members of this genus are found along the shores of the Gulf of Mexico.

Damage to pilings by shipworms can be prevented by impregnating the wood
with a suitable preservative. Coal tar creosote (American Wood-Preservers’ Asso-
ciation specification) is recommended for treating waterfront timbers of Douglas-fir
and southern pine. Copper naphthenate 1s usually used for the protection of wood in
boats. Coatings of paint and metal are also effective as long as they remain intact.

25

F-519930

Figure 8.—Cross section of a piece of timber riddled
with tunnels made by a molluscan shipworm of the
genus Bankia.

Phylum Arthropoda

Insects belong to the phylum Arthropoda, one of the major groups of the animal
kingdom. In the hierarchy of animal phyla it stands near the top, far removed from
the simplest, one-celled organisms in the phylum Protozoa. The phylum also
contains many other well-known forms such as crawfish, shrimp, millipedes,
centipedes, spiders, and mites. Members of the phylum are distinguished by a body
that is composed of a series of more or less similar rings or segments joined
together, some of which bear jointed legs. In certain forms, the segmentations of
the body may be obscure and not evident from cursory examination. In most
instances this is due to a secondary modification of form, a result of adaptation to
special modes of life (230).

Insects constitute the most abundant and important group of arthropods, but
many other members of the phylum are also common and often important. Many of
the latter are harmful, either to trees and other vegetation or to wildlife and humans.
Many are so small they are seldom seen, although enormously abundant; other,
larger ones, are easily seen. Some of the latter are also many-legged and wormlike
and bear little or no resemblance to insects; others are often mistakenly identified as
insects. The arthropods discussed here belong to the classes Insecta (insects),
Crustacea (crawfish, shrimp, crabs), Diplopoda (millipedes), Chilopoda (cen-
tipedes), and Arachnida (spiders, mites, scorpions).

26

Class Crustacea— Wood Lice

Crustaceans are primarily marine organisms. They occur throughout the world,
and several species attack the wood of most species of trees when the wood is
placed in saltwater. The surface of heavily infested wood may be so completely
honeycombed that it resembles a sponge. When this damaged wood 1s removed by
wave action, deeper layers of uninfested wood are exposed. They, in turn, are
attacked and the process of destruction is repeated. Continuous infestation may
result in the loss of the outer 2 cm of wood per year. Infested pilings often have an
hourglass shape as a result of the erosion of infested portions between low- and
high-tide marks. Heaviest attacks occur between mean tide level and low tide, but
serious damage sometimes occurs at water depths of 12 to 21 m or more (2/). In
contrast, Oniscus asellus L., the common terrestrial sowbug, and related species
are usually beneficial. In certain habitats they rapidly ingest decaying leaves and
wood, releasing nutrients needed for plant growth.

Limnoria lignorum (Rathke), the woed louse, is one of the most destructive
species. It has seven pairs of legs, sharp claws for holding onto the wood, and a pair
of toothed mandibles for boring into it. Full-grown specimens are 3 to 6 mm long
and resemble sowbugs. This species occurs in clear saltwater only and spreads
slowly, usually in infested driftwood.

Members of the genus Sphaeroma are beetlelike in appearance and are up to 12
mm long. They occur in both salt and fresh water and may be found in such places
as crevices, empty barnacle shells, and in burrows made by molluscan borers. They
also damage pilings, usually between tide marks but sometimes all the way down to
the mud line. Damage is usually less severe than that caused by Limnoria, even
though the burrows are somewhat wider and penetrate to depths of 8 to 10 cm.

The genus Chelura contains the largest of the wood-boring crustaceans. None of
these initiates attacks on wood, but they do invade and enlarge burrows made by
shipworms. C. terebrans Philippi is a well-known species along the Atlantic Coast.

Methods for protecting timbers from wood-boring crustaceans are the same as
those used against molluscan borers.

Class Diplopoda—Millipedes

Millipedes are slow-moving, elongate, wormlike organisms, usually with 30 or
more pairs of legs. Generally they are found under bark, stones, old boards, or in
damp rubbish. Their food consists of decaying vegetable matter. The adult has two
body regions: (1) the head, which bears a pair of short antennae, usually seven-
segmented; and (2) the rest of the body, which consists of a large number of similar
cylindrical segments. The first four or five body segments are not fused, and each
bears a single pair of legs. The remaining segments are fused into ringlike joints,
each of which bears two pairs of legs. Narceus americanus (Beauvois), one of the
larger species, is dark brown, narrowly ringed with red, and about 10 cm long.

Millipedes sometimes become abundant. At such times they may invade camps,
old buildings, and residences in large numbers. Many species emit a fluid with a
cyanidelike odor through openings along the sides of the body. Removal of moisture
and of accumulations of vegetable matter from infested areas should be helpful in
control of millipedes. Discussion of numerous species occurring in the Eastern
United States may be found elsewhere (/3/2).

27

Class Chilopoda—Centipedes

Centipedes are wormlike animals with elongate, flattened, segmented bodies.
They have 15 or more pairs of strong legs, one pair per segment. The antennae have
14 or more segments, and the appendages of the first body segment behind the head
are Clawlike and function as poison jaws. The last two pairs of legs at the posterior
of the body are directed backward. Centipedes are usually found under bark, in
rotting logs, or under stones or boards. They are swift runners and feed on various
small animals such as snails, insects, and spiders. All species possess poison jaws
with which to paralyze their prey. They will also bite humans.

The largest centipedes belong to the family Scolopendridae, some of which may
be more than 30 cm long. The largest species found in the United States, however,
is only 12.5 cm long. Members of this family are also the most venomous of all
centipedes. The bite of the largest ones is not only quite painful but also occasion-
ally fatal (/40). The larger and more dangerous species are southern in distribution;
those occurring in the North are usually too small to be harmful to humans. The
centipedes of New York have been reviewed (37).

The removal of debris used as hiding places is helpful in centipede control.

Class Arachnida—Scorpions, Spiders, Mites,
Ticks, and Allies

The class Arachnida is a large group of air-breathing arthropods whose bodies
usually comprise two regions: (1) the cephalothorax, and (2) the abdomen. They
generally have six pairs of appendages (the chelicera, pedipalps, and four pairs of
legs) but are without antennae.

Order Scorpiones—Scorpions

Scorpions are fairly common in the Eastern United States, especially in the
South. However, because they are active only at night and usually remain hidden
during the day, they are seldom seen. Indoors, they usually hide in such places as
closets, attics, folded blankets, shoes, and papers. Scorpions vary considerably in
size, ranging from about | to 18 cm long. The abdomen is divided into a large
anterior portion of seven segments and a long, narrow taillike posterior portion of
five segments. The latter ends in a vesicle that bears a poisonous sting. When a
scorpion runs, it holds its large clawlike pedipalps forward, and the posterior end,
bearing the sting, is usually curved upward.

Scorpions are capable of inflicting painful but rarely fatal stings. Very young and
very old people appear to be the most vulnerable. Scorpions’ food consists of a
wide variety of animal life including other scorpions, spiders, flies, beetles,
cockroaches, grasshoppers, crickets, termites, centipedes, and earthworms. A
common species in many parts of the Eastern United States is Centruroides vittatus
(Say), the so-called striped scorpion (fig. 9). This 1s a fairly small species, being
only about 58 to 60 mm long. Its sting results in a sharp pain that usually lasts about
15 to 20 minutes. Stings inflicted from late March to early May, however, may
remain painful for several hours (36).

Scorpions may be trapped during dry seasons by spreading wet burlap bags on
the ground in infested areas.

Order Pseudoscorpiones—Pseudoscorpions

Pseudoscorpions bear a striking resemblance to their larger relatives, the true
scorpions, but differ in having wider, shorter abdomens and no terminal poison
glands. They are seldom more than 5 mm long.

28

F-532018
Figure 9.—Centruroides vittatus, the striped scorpion.

Pseudoscorpions may be found in a wide variety of places—in soil cover, under
bark, under the wings of beetles, on the bodies of birds and in their nests, in
buildings, in chicken houses, and in beehives. Their food consists of mites, ants,
and various other insects. One species, Chelifer cancroides (L.), is frequently found

in association with humans. Adults are about 2.5 to 3 mm long (582).
Order Opiliones—Daddylonglegs

Daddylonglegs are somewhat similar in appearance to spiders but differ in having
small, compact, nearly globular bodies and extremely long legs. They also are
rather slow in movement, usually appearing to totter about, and they occur com-
monly in most parts of the United States. Their food consists chiefly of plant juices
or dead insects. Some apparently feed on living insects. When crushed, their bodies

give off a disagreeable odor. None is harmful to humans.
Order Araneae—Spiders

Spiders are found almost everywhere, both indoors and outdoors, and are so
familiar as to need no description (229). Objects of fear and revulsion to many
people, their presence in large numbers in and around places of habitation, or other
areas frequented by people, is often considered intolerable. Fortunately, the major-
ity of species pose no hazard to humans, even though all have venom glands.
Actually, they generally are more beneficial than harmful since their food consists
mainly of insects and other small organisms. A few species, however, are poisonous
to humans and should be avoided (35).

The black widow spider, Latrodectus mactans (F.), is probably the most
poisonous spider in the United States. The effect of its bite is described as being
extremely painful throughout all the muscles of the body, and it occasionally results
in death. The black widow occurs from southern Canada southward through the
United States, Mexico, Central America, and deep into South America. In the
Eastern States it is most common in the South, but also occurs as far north as
Maine.

Black widows are usually found in such places as garages, sheds, outdoor toilets,
under rocks and old boards, and in hollow logs or animal burrows. The female is
shining jet black and bears an hourglasslike red mark on the underside of the

29

abdomen. The body of a full-grown specimen is about 12 mm long exclusive of the
long legs. Males are much smaller and are seldom seen.

The brown recluse spider, Loxosceles reclusa Gertsch and Mulaik, is also quite
poisonous to humans, its bite producing a condition known as “North American
Loxoscelism.” Adults are about 9 mm long, have long legs, and vary from light
fawn to dark brown. A distinguishing mark is a dark fiddle-shaped band on the
anterior portion of the carapace, which narrows to a thin center line and extends
almost to the abdomen. It is only within recent years that this species has been
recognized as poisonous to humans. It has been recorded from the Southern and
Central States. It may be found in almost any location where protection, food, and
dryness are adequate, but it probably occurs most commonly inside buildings. Its
web is medium-size and irregular, with a maze of threads extending in all directions
without definite pattern (573).

It is usually impractical or impossible to eliminate spiders completely. Popula-

tions can be reduced, however, by the removal of favorite breeding places.
Order Acari—Mites and Ticks

The order Acari contains a large number of important pests of plants, humans,
and other animals. The majority are extremely small and seldom seen; others, such
as the ticks, are large enough to be seen with the naked eye. Members of this order
differ from other arachnids in having the mouth parts more or less distinctly set off
from the rest of the body on a false head, and the body is never so divided that a
distinct cephalothorax and abdomen are clearly recognizable (39).

Mites.—Numerous species of mites feed on and weaken or kill a wide variety of
valuable plants, including trees (640). Many others attack humans and various other
forms of animal life, often causing extreme irritation, and sometimes illness and
death. A considerable number are either parasitic or predacious on various species
of destructive insects and harmful species of their own kind (750, /076). Many
others feed on dead materials of all kinds on the forest floor.

The family Tetranychidae (465, 7S/) contains a large number of species com-
monly known as spider mites, many of which are important pests of trees and
shrubs. These mites vary from yellowish, greenish, orangish, and reddish to red,
and all are less than | mm long. Infested leaf surfaces are usually covered with a
fine netting of silk and spotted with tiny spherical eggs or broken egg shells.
Heavily infested foliage may be discolored, disfigured, or killed.

The spruce spider mite, Oligonychus ununguis (Jacobi), feeds on a number of
conifers, especially spruce, thuja, and some of the pines, and is widely distributed
in the United States. Infested trees may become brownish gray and appear un-
healthy, or they may be completely defoliated. It is considered the most destructive
conifer-feeding mite known (/43). Outbreaks over large forested areas have been
recorded. Periods of drought appear to be most favorable for population buildups.
Outbreaks have also occurred following widespread spraying with DDT (642).
Young nursery stock and recently planted trees are especially subject to serious
injury.

Oligonychus milleri (McGregor) feeds on pines throughout the South and North
to Pennsylvania and the Lake States. Outbreaks covering millions of hectares have
been recorded from North Carolina to Florida, in northern Louisiana and southern
Arkansas, and in Mississippi and Texas. Young, open-grown, even-aged stands
appear to be susceptible to severe attack.

Other important species in the genus Oligonychus include O. bicolor (Banks),
long recognized as a pest of oaks, especially shade oaks, and sometimes injurious

30

to beech; the southern red mite, O. ilicis (McGregor), damages azalea and
camellia; O. aceris (Shimer) is sometimes a serious pest of maples; O. letchworthi
Reeves occasionally seriously injures hophornbeam; O. newcomeri (McGregor)
often seriously infests serviceberry; O. cunliffei Pritchard & Baker feeds on longleaf
pine in Florida; O. boudreauxi Pritchard & Baker feeds on baldcypress in the South;
and O. propetes Pritchard & Baker feeds on oaks from Washington, D.C., to North
Carolina.

The carmine spider mite, 7etranychus cinnabarinus (Boisduval), 1s a common
and widely distributed species that feeds on a great many species of plants, trees,
and ornamentals. Several generations are produced during the summer months,
often giving rise to tremendous populations. Heavily infested plants may be entirely
defoliated, especially during hot, dry weather. 7. homorus Pritchard & Baker feeds
on hickory and ash in North Carolina; 7. magnoliae Boudreaux occurs on magnolia
and yellow-poplar in Louisiana; the fourspotted spider mite, 7. canadensis
(McGregor), feeds on elm, basswood, horsechestnut, Osage-orange, and poplar
throughout the Eastern United States and southern Canada; and the Schoene spider
mite, 7. schoenei McGregor, infests elm and black locust throughout the Eastern
United States.

The genus Eotetranychus also contains a number of common and frequently
important species. F. populi (Koch) and E. weldoni (Ewing) are found on poplars
and willows; E. hicoriae (McGregor) occurs on pecan, hickory, horsechestnut, and
various Oaks; F. matthyssei Reeves attacks elm in New York. Heavy infestations
may cause severe browning and cupping of the undersides of leaves. E. querci
Reeves has caused severe browning of pin oaks in New York.

Platytetranychus multidigituli (Ewing) feeds on the leaves of honeylocust, caus-
ing them to turn yellow, and P. thujae (McGregor) attacks northern white-cedar,
juniper, and baldcypress. Eurytetranychus buxi (Garman) is a serious pest of
European boxwood. The leaves of infested plants become bronzed, then wither and
sometimes fall prematurely.

The family Eriophyidae contains a number of tree-infesting species (66/). Many
produce open pouchlike or blisterlike galls on the twigs and leaves of their hosts.
Some cause a rusting of infested leaves, and certain others feed on buds. A few of
the more common and important tree-infesting species are discussed here.

The maple bladdergall mite, Vasates quadripedes Shimer, is a common spe-
cies. It ranges in length from about 0.05 to 0.2 mm and feeds on the undersurface of
silver maple leaves, causing the formation of pouchlike or bladderlike galls up to 3
mm in diameter (fig. 10). At first, these galls are light colored or yellowish green.
Later, they are reddish to almost black and look for all the world like miniature
green peppers standing on stalks above the leaf surface. Heavily infested leaves are
often distorted. The related species, V. aceris-crummena (Riley), produces slender,
fusiform, or spindle-shaped galls about 5 mm long on the upper surfaces of silver
and sugar maple leaves.

A few of the other eriophyids infesting trees and some of their hosts are as
follows: Eriophyes fraxiniflora Felt feeds in the staminate flowers of ash. Infested
clusters become deformed and remain on the tree as green masses until fall. The
pearleaf blister mite, E. pyri (Pagenstecher), an introduced pest of pear and apple,
also has been recorded on mountain-ash and serviceberry. It is sometimes abundant
enough on mountain-ash to cause noticeable injury. E. parapopuli (Keifer) stunts
the growth of poplar by producing woody galls around the buds. FE. caulis Cook
causes large, deforming, hairy growths on the petioles of black walnut leaves.

Bil

Acalitus phloeococotes (Nalepa) detorms fruit spurs and produces woody galls on
plum. Nalepella tsugifoliae Keifer has been reported to damage hemlock in nurser-
ies in New York. The pine bud mite, 7risetacus pini (Nalepa), causes the yellowing
and dropping of pine needles. 7. cupressi (Keifer) attacks southern redcedars,
causing distortion and deformation of young trees.

F-506694

Figure 10.—Galls of the maple bladdergall mite,
Vasates quadripedes, on leaf of mapie.

Mites of the family Sarcoptidae are skin parasites of warm-blooded animals. The
itch mite, Sarcoptes scabiei (De Geer), is a well-known species. It attacks humans
and causes severe itching as it burrows into the skin.

The family Trombiculidae contains the notorious redbugs or chiggers. The
common chigger in the Eastern United States is the first instar of the species,
Eutrombicula alfreddugesi (Oudemans). In its later stages, it feeds on insects,
snakes, birds, lizards, and rodents. When it attaches itself to the skin of humans and
insects, it inserts its mouth parts to feed, causing intense itching and sores. Heavy
attacks may cause fever, and secondary infections may occur. Chiggers are often
abundant in the forests of the Central and Southern States and along the Atlantic
seaboard north to New Jersey.

Many other species of mites occur in association with bark beetles (89/), and
some are known to be parasitic. So far, very little information is available on their
effectiveness in natural control of the beetles. In some instances, it may be
substantial.

Ticks.—Ticks are all parasitic, chiefly on mammals, birds, and reptiles. All
American species have a number of features in common, such as large size, a
piercing hypostome with recurved teeth, and chelicera with lateral teeth on the
movable digits. Some ticks are mostly covered by a hard dorsal plate called the
scutum; in others, this hard plate is absent.

The American dog tick, Dermacentor variabilis (Say), occurs throughout the
Eastern United States, but is most abundant in coastal areas and in the Mississipp1
River Valley. The larvae and nymphs feed largely on rodents; the adults feed on

32

dogs, horses, hogs, cattle, and many species of wild animals. Humans also may be
bitten, but they are not a preferred host (777). Unfed adults are brown, variously
marked with white, and about 5 mm long. Engorged females are bluish gray and
sometimes reach a length of 12 mm. Females lay eggs in various places but never
on the host; the young seek out their host after hatching. Heaviest infestations are
usually found on vegetation along game trails, paths, and roadways. Adults are
most abundant during spring and early summer in the North. After August they are
usually very scarce. In the South, they occur throughout the year. This species
transmits Rocky Mountain spotted fever and tularemia, both serious diseases of
humans. It also transmits anaplasmosis, a disease of cattle.

The blacklegged tick, /xodes scapularis Say, occurs along the Atlantic Coast.
The larvae and nymphs feed on rodents and reptiles, and the adults feed on deer,
cattle, sheep, dogs, and other large animals. People are also bitten. This species has
long mouth parts and inflicts a very painful bite. Infestations are usually found on
vegetation along game trails, paths, and roadways where people and other animals
come by. Adult ticks are most prevalent during fall and early winter. This species
transmits anaplasmosis to cattle and piroplasmosis to dogs (/2/9).

Class Insecta—Insects

An insect is an air-breathing arthropod with a distinct head, thorax, and ab-
domen. It has one pair of antennae, three pairs of legs, and usually one or two pairs
of wings in the adult state. The majority of species are terrestrial and are found in an
almost endless variety of microhabitats on the land. Many other species spend part
of their lives in water.

The majority of insects hatch into wormlike larvae that grow by periodically
shedding the outer skin, finally transforming into an inactive pupal stage from
which the adult emerges. This type of development is known as complete meta-
morphosis. The immature stages of some others are very similar in appearance to
the adults and are known as nymphs. Members of the latter group are said to have
simple or incomplete metamorphosis.

Approximately 900,000 species of insects have been described. These species
represent 80 to 90 percent of all the known kinds of animals. The actual number of
insect species, however, 1s believed to be much larger and to run into the millions
(1048). Countless numbers feed on plants of all kinds, attacking all parts from the
roots in the ground to the flowers and seeds in the tops. Thousands of others feed on
other insects or other animals, including people.

Fortunately for humans and their interests, the majority of insects are either
innocuous or beneficial. The remainder, unfortunately, include some of our most
important enemies—not only do they feed on humans and other animals and often
transmit deadly or debilitating disease organisms, but they also devour crops and
decimate forests. Because of their abundance, their fantastic reproductive powers,
and their remarkable capacities for adapting to changing conditions, insects present
a continuing challenge to us in our efforts to limit their numbers to tolerable levels.

Several general references to insects are available for consultation (/40, /55,
LOD" 23083605025, 725% 828, 1043).

33

Keys to the Orders and Families of Forest
Insects and Allies, Based on Types of Injury

The following keys are designed for the use of those unfamiliar with the orders
and families of insects. They are first separated into primary divisions according to
the portions of trees attacked, the size of the trees, and the timber products infested.
These main divisions are in turn subdivided into other groups or subdivisions.

I.

III.

Primary Divisions of Key

Insects injurious to seeds, seedlings, small reproduction, and young planta-
tions.

A. Seeds, cones, and fruits.

B. Seedlings and small reproduction.

Insects, etc., injurious to large living trees and to small trees more than | or 2
meters tall.

A. Yellowing foliage.

B. Defoliators, leafminers, etc.

C. Twig and tip damage, etc.

D. Borers in wood and bark.

E. Galls, swellings, etc.

F. Sucking insects.

Insects injurious to forest products.

Defects in green timber.

Insects in round logs.

Insects in lumber.

Insects in wood in ground.

Defects in wood in salt and brackish water.

SS

Division I
Insects Injurious to Seeds, Seedlings, Small Reproduction,
and Young Plantations

This group includes the insects that attack the seeds, cones. and fruits of forest
trees. and young seedlings in the nursery, plantation, or forest. Insects attacking
trees more than 4 or 5 years old (large reproduction, forest and shade trees) are

discussed under Dvision II.

l.

34

A. Insects Attacking Seeds, Cones, and Fruits

Larvae without well-developed head capsule: maggotlike:
With a sclerotized structure like a “breastbone” near anterior end: in
seeds of fir, baldcypress. birch, and fruit of chokecherry..... Diptera,
Cecidomyiidae
Without “breastbone”: mouth parts well developed: in fruits of cherry.
apple. plum. hawthorn: in berries of dogwood. holly, and others: and in
WalnUL IDUSKS: 24 23.6 5c Seen pee et eee eee Diptera. Tephritidae

i)

Te.

10.

11.

. In the shucks of pecans, hickories, and walnuts and in acorns... .. Lepi-
doptera, Blastobasidae, Tortricidae

IMEC ONE Si en RBar Neti. a ciatrec seri t veal eke se is Lepidoptera, Pyralidae
In fruits of wild cranberries and blueberries ... Lepidoptera, Pyralidae
In acorns, walnuts, chestnuts, hickory nuts, and filberts ............
Iinele OuMMIMNOUS -SCCUSs texte aero ee ee Coleoptera, Bruchidae
IMG COMES arene net eee ouvert Sy tem Mua, Lane late) oe Suns Aloe ont cema dg Bid 6G
First abdominal spiracle vestigial; body spindle-shaped .. Hymenoptera,
Cynipidae

First abdominal spiracle normal; body curved like a closed finger ....
Coleoptera, Curculionidae

. Cones abnormal and dropping prematurely .... Coleoptera, Scolytidae
Cones maturing; damage to seeds alone

WeaIVAC SIS CI ESS rire sais sued Saw Sete eck Hymenoptera, Chalcididae

anvacawathrtrue less eid dint) le hu he wats Coleoptera, Anobiidae

B. Insects, etc., Attacking Seedlings and Small Reproduction

Plants cut off near ground line or stems lacerated and shriveled ......
Plants wilting or fading, easily plucked from ground because of severed
Stemmol Lools*DElOWSTOUNG: so..4 2) Sarecs vole eh Pa he Gwe He a oe os
Callelikesswellangs,On-the StOmMs Ao iiie ee wate Papas ea oo elie wie iw a
Bark gnawed in patches along the stem of conifers ................
Foliage off-color, yellowish or rusty, and often covered with very fine
cobweblike threads or matting .............. Acari, Tetranychidae
Roots showing small cottony or globular objects at time of transplanting
Scat, onctta 2a). cothe ea ae R gr n RC e O Homoptera, Aphididae
Leaves or cotyledons cut off and carried away; Southern States
i Sey een en eee aa a Hymenoptera, Formicidae
Other types of damage. See Division II.
Small, tender plants:
Small, legless, maggotlike larvae causing injury ... Diptera, Muscidae
Large, smooth, dark-colored caterpillars with legs and prolegs present
Re eel e Uiaacctle se Saag hak, ot hates Lepidoptera, Noctuidae
Larger plants, hardwoods, with woody part tunneled:

ftunnelsslongitudimal <2. eS ieue vile sd ae 3 Coleoptera, Cerambycidae
Tunnels across the grain, wood stained ..... Coleoptera, Scolytidae
Tunnels irregular, chiefly mderground ................. Isoptera
. Curved, grublike larvae in the soil ........ Coleoptera, Scarabaeidae
Elongate, cylindrical, hard-shelled larvae ..... Coleoptera, Elateridae
Small, molelike tunnels usually near the surface of the soil ... Orthoptera,
Gryllotalpidae

Bark swollen and gnarled at or just below ground line
WMimimendamage ser oo. oes a es oe 3 Lepidoptera, Pyralidae
Suckinevdamaces. isl. Ls eka se. Homoptera, Fulgoridae
INearbyganmtlilly res hy ie i See. Hymenoptera, Formicidae
Northern, Southern, and Lake States ...... Coleoptera, Curculionidae
Southemuandslake; States). 7252.6. oh Orthoptera, Acrididae

10
1]

35

Division II
Insects, etc., Injurious to Large Living Trees and to
Small Trees More Than | or 2 Meters Tall
This group includes insects and related organisms that commonly attack living
forest and shade trees but excludes those that attack fruit and seeds or that attack
small plants not more than 4 or 5 years old. Insects responsible for damage to wood
products, many of which primarily inhabit dead or dying trees or which attack logs,
lumber, and other wood products, are discussed under Division III.
Injury consisting of discolored, yellowish, rusty, or mottled foliage covered

with fine cobweblike threads or matting ....................2.0005- » A
Injury consisting of defoliation, leaf rolling, leaf tying, leaf and petiole
mining, or bast or epidermis miners on green-barked stems ........... B
Injury occurring on new growth, twigs, branches, or small trees, consisting of
mining, pruning, withering, or flagging ....................0.0005- é:
Injury caused by larvae or beetles boring in the bark, under the bark, or in the
WOO wis oer sit tsk ccseily Sew S, ¢ Stale cue tac, whe Ome ece ane aetna D
Injury consisting of a gall or swelling on stem, branch, or leaf .......... E
Injury caused by sucking insects feeding on leaves, twigs, or bark surfaces,
usuallythe SOfter tissues Of (he plant 2... 4<26 2. 2G eas eee h oe ee ee F

A. Yellowing Foliage

Injury consisting of discolored, yellowish, rusty, or mottled foliage covered
with fine cobweblike threads or matting .......... Acari, Tetranychidae

B. Defoliation and Other Injury

L. Injury-caused by beetles and/or larvae xn. 8 Gs co ba Fe She Me oe 2
Fy Y Case AE VAC Lo coyote «9 Se ae xed eRe ee eee 4
Injury caused by other types of insects or insect not present ......... S

2. Adults and larvae associated on the leaves... Coleoptera, Chrysomelidae
PUES Only PRCSONE, ea aches 2a ia ee are a 3

3. Beetles usually found feeding at night ..... Coleoptera, Scarabaeidae
Small, bright-colored jumping beetles .... Coleoptera, Chrysomelidae
Dull black, purplish, or gray, soft-bodied beetles. . Coleoptera, Meloidae
Siiall snout Wecles: caves we, ».z\d<.2 Aer had Coleoptera, Curculionidae

4 SProleos, Uusually-two Or FIVE padlls 4.2) s,s hs a oe ee oe Lepidoptera
Prolegs, usually six or more pairs—or none ......... Hymenoptera,

Tenthredinidae
Prolegs inconspicuous; leaf- or bast-mining forms”

>; Ciretlar holes cut the leaves 2.4.3... Hymenoptera, Megachilidae

Trees defoliated, ant mounds nearby; Texas, Louisiana .. Hymenoptera,
Formicidae

Leaves rolled into a small, compact bundle Coleoptera, Curculionidae

> There seems to be no simple and practical method of separating the leafminers of the four orders that
have species with this habit. Those found in conifers are probably either Lepidoptera or Hymenoptera:
those on hardwoods may be Lepidoptera or Hymenoptera, or of the families Curculionidae.
Chrysomelidae. or Buprestidae of the Coleoptera: or they may be Diptera of the families Agromyzidae or
Cecidomyiidae.

36

Grasshoppers associated with injury ......... Orthoptera, Acrididae
Walkingsticks associated with injury ....... Orthoptera, Phasmatidae

C. Twig Pruning and Other Injury

. Injured portion hollowed or mined; injury caused by larvae or bark

beetles sawiichyane wsuallivapresente trek ewe ee essen eb ees
Injury caused by external feeding or ovipositing, which removes a portion
of the bark or causes a definite mechanical injury or a resin-infiltrated
SC Algae er) ere men nae Tt eta tino doa Or Oh Mar SMe hte are cache Oa aid, tele
Cottony masses on tops of twigs concealing the insects; conifers .....
Homoptera, Coccidae, Phylloxeridae
Injury on two or more whorls of the terminal of conifers; inactive,
curved larvae under bark or in pupal cells in wood........ Coleoptera,
Curculionidae
Twigs or branches of hardwoods or conifers containing bark beetles or
powderpost beetles or a cylindrical shotlike hole, usually darkly
stained, directly entering injured portion ....Coleoptera, Scolytidae,
Bostrichidae
(NUT ELE UTS UT Se ae ea aR Ren ag
ihwicsnotmined=below fading portion 2... 06.52. basse. ee Se boa eee
Twigs mined far below fading portion, tunnel often extending to the
ROU Oe meg Aree ae ate a Ee ws fe oe Coleoptera, Cerambycidae

. Larvae with prolegs; often pitch masses at point of injury . . Lepidoptera,

Olethreutinae
Larvae without conspicuous prolegs; usually a spine on last segment
5 olGe MEDS Bik. A a Gaeta OO a ie ee RE AT Ie Hymenoptera, Siricidae

. Larvae with well-developed prolegs; usually colored; usually in more tender

parts of twigs ...... Lepidoptera, Olethreutinae, Sesiidae, Cossidae

Larvae with underdeveloped prolegs; in woody portions of twigs .....

> Elongate; iat larvae; mines filled’s.-. ts. .0... Coleoptera, Buprestidae

Cylindrical larvae; mines open ........... Coleoptera, Cerambycidae

. Obvious scar and pitching of wood at base of injury or along twigs;
conifers

Scale ‘bodies present on twig. ..-: 0.02. Homoptera, Margarodidae

Scalesbodiessabsemt.> 8 xe Coleoptera, Cerambycidae

Numerous phloem scars on twigs; spittle masses may be present .. .
Homoptera, Cercopidae
(Hail injury is similar except that the scars are always on top side of

branch)
Twigs slit with a lacerated wound at base of injury or at point of
LON TSU Ge ase ee ee ee Homoptera, Cicadidae, Membracidae

Orthoptera, Gryllidae

D. Borers in Wood and Bark

14. Borers in the phloem and outer corky bark of living trees rarely scarring

(LOVE -SVOCONGYG Se Un VE eR Be 8 RRS Se pa SR one ae ea

10
11

15.

16.

20.

22

38

Borers in callous tissue around wounds

On various hardwoods ............... Coleoptera, Curculionidae
On maples; 15+; Lepidoptera, Sesiidae Coleoptera, Cerambycidae
Olr Copiers. os 2a hee estes poet Lepidoptera, Sesiidae, Pyralidae

Borers in the dead wood beneath fire scars, turpentined faces, blazes,
cavities, and: Similar wWOundS.23.3 = «.< akat.ancene cca scevads eb e es
Borers under the bark or in the wood (other than beneath scars or catfaces)
Ol IMVIMG WEES. <.ot ooe ua eae Maas uah Rare eee ee ee See ee
Bark beetles associated with their larvae under the bark ............

Root Doters:of mining at baSe “Ol Wee 24 bo sc5n toca nc stat ae Seees
White, unpremented Jarvae ..4...4<5¢58. Coleoptera, Cerambycidae
Highly pigmented larvae in galls ...... Hymenoptera, Tenthredinidae
Serpentine mines just under the epidermis of chestnut and oak .......
Lepidoptera, Nepticulidae

Beetle larvae; wood stained around holes’... Coleoptera, Scolytidae,
Platypodidae

White, fleshy, cylindrical larvae in hardwoods ....................
Coleoptera, Cerambycidae, Brentidae
White, fleshy, flat-headed larvae in turpentined faces in fire scars on

COMMCES 6 Serr ae en ae = ae oe eevee eee Coleoptera, Buprestidae
Larvae with heavy, chitinous armature on last segment; chestnut, oak,
TDS Se aonatenccapy cuensaaanere a: ecient Coleoptera, Lymexylonidae

. Larval mines extended under the bark and also deep into the wood in later

SIO ES. liu: at arn aces oes Gea nes oe wee re ee ein eee rere cee
Larval mines entirely under the bark or only in wood of current annual
3) i ee ee en eee ee ee ee nae err a oe

. Pitch exuding from larval mines; larvae with prolegs present; conifers

en ae a ee ee eee Lepidoptera, Sesiidae, Pyralidae
No pitch, ‘but often water and tass exuding 2..2%,255 524 2s45-4¢e 503
Head of larvae globular protuberant ..°.6:2....0:.20nssduns ewes
Head of larvae somewhat flattened and embedded in prothorax . . Coleop-
tera, Cerambycidae
Prolegs absent; last segment often heavily armed ........ Coleoptera,
Tenebrionidae

Prolegs absent; larvae curved, grublike; in willow, poplar, and palmetto
Cine wile Bee uae at ee ee Coleoptera, Curculionidae
Prolegs present; last segment not heavily armed ...Lepidoptera,
Hepialidae, Sesiidae

. Larvae depressed, flat-headed or pestle-shaped. . Coleoptera, Buprestidae

Larvae curved, orublike ..242..2.2s924.: Coleoptera, Curculionidae
Larvae slender; thoracic segments not noticeably enlarged; cause pitch
tlecks-mt wood: birch. Cle: 2.45.4 sae oa Diptera, Agromyzidae

Larvae with prolegs:; poplar, willow, alder, ash, persimmon..........
Lepidoptera, Hepialidae, Sesiidae
Larvae without prolegs

In hardwoods ...............---000+ Coleoptera, Cerambycidae
In conifers; associated with pitch mass ... Coleoptera, Curculionidae,
Scolytidae

18

le

iy
20

23%

24.

25:

26.

Dil

28.

29:

30.

Sl

SY

E. Galls?

Galls of more or less open, exposed, simple structure, or, when enclosed,
the insects maintain permanent openings, or the galls are dehiscent to
permit the escape of the numerous insects inhabiting them ........

Galls usually completely enclosing the inhabitant; one or, rarely, several
insects to a cavity; occasionally a permanent opening is maintained by
thre wice din Pel anvae quer cml taht eats Bou hard ak hms Lea aces ee

Mites present having two pairs of legs; galls of various shapes but always
provided with an opening to the exterior and lined on the inside with
NAIEWSOI IZZY CLOWNS oreo. e sevice tele tie a e\ et Acari, Eriophyidae

AMIS POMC TINAS Cee tay erasers eth BRS Aaot dere EO oY teciy mad aban Guat sed! SSB ge. Mate

INSECISNOtsILCE. TOR UIMPING a4h- 2 deems s dese, noe es beens tea Rade dws

Insects with hindlegs developed for jumping ... Homoptera, Psyllidae
Leaf galls on hardwoods, chiefly elm, poplar, hickory, ash, sumac, and

NVC MEM AAGM Ae clgitr tal, wd 68 ui terse Se made Homoptera, Aphididae
Conelike galls on tips of spruce twigs ..... Homoptera, Phylloxeridae
Pitlike galls on twigs of hard pines ....... Homoptera, Margarodidae
Pitlike galls on twigs of white oak ..... Homoptera, Asterolecaniidae

Galls inhabited by larvae with well-developed head capsules ........
Larvae without well-developed head capsules, maggotlike; white to yel-
lowish or reddish in color:
Larvae with a distinct structure like a “‘breastbone” near anterior end
Re Etta rane ta FF AS es sseiatens & 2 Diptera, Cecidomyiidae
Larvae without ““breastbone’’; mouth parts well developed .... Diptera,
Agromyzidae
lcanvac leelessaor -with-only minute Tees sa. te ieee Goo hae 2 ams
Bessswelledevcloped also; prolegs. present 7...2. 6) 5200s Sasol wa
Woody galls containing plain evidences of mining activity of the larvae;
larvae with well-developed head capsules and mandibles . Coleoptera,
Buprestidae, Cerambycidae, Curculionidae
Larval mines not obvious; white larvae, curved or grublike in form,
legless, and with distinct head capsule, each contained in a specialized

CONE ee reenact Be Hymenoptera, Cynipidae, Chalcididae
Onew illo gee ie Arce rs pr” ets ed Hymenoptera, Tenthredinidae
Onvlocust=poplar maples 23. ur. S ae sae. Lepidoptera, Tortricidae

F. Sucking Insects

InjuRVZOT ANSects presemt ON -TCAVES 2.6 ee gabe we cy Se
Injug¢spmimanilyacontined tO LW1SS Rae vo. at Pete ec ja ee ee eas
Injury primarily confined to branches and main stem ..............
Leaves off-color, yellowish or spotted from feeding punctures of active,

jumping insects .... Homoptera, Cicadellidae Hemiptera, Tingidae
Leaves bearing galls or abnormal spots:
(OTe NTS! XO Si AVE Sat on een an ara a ee Homoptera, Psyllidae

24

oa

25

26

28

pay)
30

32
33
34

* It seems impossible to devise a key that will separate all the varied types of galls into family or order

groups. However, many groups are fairly true to type and, if considered with the larvae or other stages of the
insect inhabiting them, you can make a fairly workable distinction.

39

On elm, poplar, willow, witch-hazel, hickory, oak, chestnut, etc.
ae Gre eee eee Homoptera, Aphididae, Phylloxeridae

OP COMMEIS( 2 hu 15- hee ote es Sees Homoptera, Phylloxeridae

Exposed insects on the leaves:

Scalelike, gall-like, or soft grublike insects covered with wax in the
form of powder or tufts ................ Homoptera, Coccidae

Fringed, scalelike immature forms associated on the leaves with white
four-winged “thes” 2.2... se 5ns005> Homoptera, Aleyrodidae

Soft-bodied insects with long conspicuous antennae . Homoptera, Ap-

hididae
33. Insects surrounded by a conspicuous frothy mass of spittle:
Ends of branches and trees slowly dying in severe infestations, numer-
ous small resin-soaked spots in inner bark and twig wood: pines
ee eee re a eee ea ae Homoptera, Cercopidae
Injury consisting of ragged slits in the twigs, often breaking at incision:
the tops of the branches hanging with withered leaves; hardwoods
pevindee cOrre te eee hee ree Homoptera. Cicadidae, Membracidae
Injury consisting of gall-like or gouty swelling on limbs and twigs of firs
re ar ee a ee eS ere eee Homoptera, Phylloxeridae
Tips of hard pines flagged (needles yellowing); scales embedded in pits in
DAU Ge oe seer ein ee ee ee Homoptera. Coccidae
Branches and twigs infested with scalelike, gall-like, or soft-bodied
insects covered with waxy powder or tufts; twigs often dying ......
Homoptera, Coccidae

Tips of branches swollen forming pineapplelike galls ..............
Homoptera, Phylloxeridae
Tips of new growth withering, infested with numerous soft-bodied insects

with prominent antennae ................ Homoptera, Aphididae

34. Fir trees unhealthy and dying; trunks infested with masses of soft-bodied
insects appearing as a whitish wool .... Homoptera, Phylloxeridae

Beech trees unhealthy and slowly dying, with dead areas of bark on stems
covered with whitish masses of soft-bodied insects .... Homoptera,
Margarodidae

Trees infested with scalelike, gall-like, or soft grublike insects, and
covered with wax in the form of powder or tufts ...... Homoptera,
Coccidae

Division III

Insects Injurious to Forest Products
This group includes insects causing the type of injury seen in the handling of
forest products, i.e., logs and lumber, poles, posts, piling, and manufactured
materials such as handles, gun stocks, stored wood, and wood in buildings. Certain |
types of damage found in green logs or freshly sawed lumber are the result of insects |
boring in the phloem or wood of the living tree. These are also treated here for |
convenience. They are usually distinguishable by the more or less stained condition |
of the surrounding wood, pitch infiltration, or the presence of scar (callous) tissue. |
Defects occurring in the wood of green logs or lumber, revealed as the logs are |

sawed, usually as darkly stained, pitch-infiltrated wood, or scar (callous)
CISSUC 2. a sek eo weet ee oad PRS bee Seat ae ee ee A

Injury occuring to material having the bark present (lumber excepted), such as
round logs after the trees are felled and left either in the woods or at the
mul On OSSeUmIZed fOr mMIStIC WORK CLC. f2Binc we ote bn. ea.

Injury to freshly sawed lumber, seasoned lumber, stored and manufactured
matenalsmOmWOOdmmMyDUNGINGS Wo. ava eee eee ee ee ae

Injury to materials in contact with the ground, such as crossties, posts, poles,
foundation materials, pilmg above’ water, etc: ....2 0... ek eee.

Defectsam wood in-salt and brackish water... 0. 5.22.00... eee cae

A. Defects in Green Timber

IP U TIME ARCO OU Sez ces chore heca ee mo ayo Gag abo ccs ane atabee Giapatle me klel aher tye @lanie4, a 4

ICOMBICTS MR iis atten rte a es Ta er ay ols w ole Wclutacdad & & Hae cw wha a

2. Holes small, “pinholes,’’ 6 mm or less in cross section; circular, open,

imemanever filled: with boring dust =~... eles... eee ee ee ed

Holes larger, ““grub holes,” up to 19 mm in diameter, usually oval in cross

section, usually open, not filled with boring dust ................

Pith flecks in wood; birch, maple, etc. ....... Diptera, Agromyzidae

3. Pinholes, about 3 mm in size, of uniform diameter throughout, wood

stained in streaks, in oaks and yellow-poplar Coleoptera, Scolytidae

Holes tapering, several sizes grouped together and originating in a

wound:

Holes up to 6 mm in diameter, in American chestnut and chinkapin

MPM er ests NE wis Case ena Coleoptera, Lymexylonidae

Holes up to 3 mm in diameter, in oak and other woods ... Coleoptera,

Brentidae

4. Variable-size holes grouped and radiating from wounds or cavities ....

Coleoptera, Cerambycidae, Tenebrionidae, Brentidae

Large grub holes, up to 25 mm in diameter, usually appearing singly and
not associated with wounds: |

In hickory ... Coleoptera, Cerambycidae Lepidoptera, Cossidae

In poplar and cottonwood... Coleoptera, Cerambycidae Lepidop-

tera, Cossidae, Sesiidae
In maple .... Coleoptera, Cerambycidae, Tenebrionidae Lepidop-

tera, Cossidae, Sesiidae
MRS lala tee ede eh fete wiartaara has te SHI doit Lepidoptera, Sesiidae
IMPS ESIMAMION: . ie 5: ce Ne lo. Lepidoptera, Sesiidae
In locust ..... Lepidoptera, Cossidae Coleoptera, Cerambycidae

5. Pitch pockets in the wood... Lepidoptera, Sesiidae, Pyralidae Coleop-
tera, Scolytidae
Holes filled with boring dust, associated with turpentine faces or fire

scars:
nthe SOUL APINES. "2 diab eck ete be Coleoptera, Buprestidae
In the North, pines or other conifers .... Coleoptera, Cerambycidae,

Buprestidae
B. Insects in Round Logs
6. Sawdust exuding from small, round ‘pinholes’ (2.5 mm or less in

diameter) on the surface of the bark; wood usually stained around the
NOG: se, sk Sate nian Coleoptera, Scolytidae

MM NO

41

Sawdust exuding from larger holes; larvae present under the bark or in
WOOG Fxg done shatter he et ae eee oe Coleoptera, Cerambycidae

Sawdust not exuded; the only evidence of work is the presence of larvae or
galleries under bark or in the wood:

Larvae elongate, cylindrical ........... Coleoptera, Cerambycidae
Earvae flat-headed:< 24:1. 65i-.+eda48 ess Coleoptera, Buprestidae
Larvae curved, legless; only one larva to a burrow ... Coleoptera,

Curculionidae

Larvae curved, legless: several larvae in a burrow, each usually sepa-

rated by a pith or clay partition across the gallery ... Hymenoptera,

Sphecidae, Vespidae, Apidae
Larvae and bark beetles associate ......... Coleoptera, Scolytidae

C. Insects in Lumber

7. Fine sawdust exuding from small “pinholes” (less than 2.5 mm in
diameter) in green lumber; holes usually darkly stained .. Coleoptera,
Scolytidae
Sawdust, if exuding, coming from larger holes in drier lumber, cut a
DEOMI OT ORS Satlerd pases Be: «ake ed ee ae 7 ewat ene eer eines tae arate waar
8. Damage to lumber with bark present:
Larvae ClO ANS x8 uke mi. Se GAs ee Coleoptera, Cerambycidae
Ary ae CUIVCU 2671.2 on nearness Sala Coleoptera, Bostrichidae
Damage not associated with presence of bark on material ...........
9. Fine sawdust exuding from circular or oval holes:

Siiall.. CuEvee Iainae =. dete dye eee ne Coleoptera, Lyctidae
Ploneste larvae «3.2. eect eee eee Coleoptera, Cerambycidae
Large, black ants associated with damage; sawdust accumulating in large

pilés from.damp wood ...n~ ere ceesse Hymenoptera, Formicidae

Damage concealed and sawdust usually not falling from holes .......
10. Larval tunnels packed with sawdust:
Larvae elongate scylundricalies..oc 259% « Coleoptera, Cerambycidae
Larvae elongate, flat-headed ............ Coleoptera, Buprestidae
Tunnnels open:
Irregular cavities following the grain of the wood loosely filled with

fie ammpressed Peels) .2 91.4 As de oh ogee oes oe 3 ee Isoptera
Round holes 12 mm or less in diameter, often with cross partitions or
COS. b.1.0:5-3 au anete of Hymenoptera, Apidae, Vespidae, Sphecidae

D. Insects in Wood in Ground

11. Large, elongate larvae associated with damage consisting of grub holes
extending through the wood............ Coleoptera, Cerambycidae,
Oedemeridae

Large irregular cavities eaten in the springwood, usually extending with

the grain of the wood; sides of cavities plastered with claylike excre-

ACI Weides eee sae ome ene he Aas eae cee oa s [soptera
Large. irregular cavities eaten into wood, usually cutting across grain,
surfaces smooth, no excrement, large piles of sawdust accumulating
outside; large. black ants associated with injury in moist or damp

WOOR. exis gne teehee Begs sales wen Penn Se Hymenoptera, Formicidae

42

10

E. Defects in Wood in Salt and Brackish Water

12. Pilings with spongelike damage resulting in hourglass shape due to wood
erosion between low and high tide ........ Crustacea, Limnoriidae
13. Pilings or other wood with holes showing calcareous shells in holes
eee ter nt ncroh Vat ial Mita Me iow | Gastropoda, Teredinidae

43

Important and Selected Orders of Forest
Insects

The major section of this book. which follows, discusses the most important
insects affecting forest and shade trees, shrubs, and wood products. The discussion
includes brief descriptions of the insects and their habits, primary hosts, economic
impact on forests, and possible control measures. Also included are a number of
other insects that are abundant in the forest but usually are not thought of as forest
insects. Given the ever-increasing use of forested areas for recreational purposes
and the growing public concern about our wildlife resources, it seems appropriate
to include such insects and describe their importance as pests of humans or other
forms of animal life.

Order Thysanura—Bristletails

Bristletails occur abundantly in the forest in rotting wood and debris, under
stones, and among fallen leaves, but none is injurious. They are distinguished by
the stylelike appendages on some of the abdominal segments, by the two or three
taillike appendages at their posterior ends, and usually by their elongate bodies.

Order Collembola—Springtails

Springtails are very small, primitive, wingless insects. The body is covered by a
soft exoskeleton and there is a single pair of antennae, each normally consisting of
four segments. The mouth is located ventrally, and the mandibles and maxillae are
either toothed for chewing or styliform for sucking. There are three pairs of legs,
each typically terminating in one or two claws. The first abdominal segment bears a
ventral tube or collophore; there is a pair of small appendages fused basally on the
venter of the third segment; and a furcula is appended to the ventral surface of the
fourth. The latter operates as a spring and 1s capable of propelling the insect into the
air as much as 10 cm (835).

Springtails are widely distributed and are among the most abundant of insects.
They are found in all kinds of places—many unexpected, such as on the surface of
snow and on the surface of water. In the forest they are found in moist soil, among
dead leaves. in dead and decaying logs, under loose bark, and in bark crevices of
living trees. The majority feed on algae, fungi, and lichens and other living or dead
plant matter. Pollen from conifers is favored by some species in the spring. Some
may be attracted in large numbers to decaying fruit and animal matter. Others may
be attracted to sap flowing from trees in the spring.

Springtails tend to be beneficial in the forest because they help to reduce litter
and form humus. Keys to Collembola occurring in eastern North America are
available (88, 429, 535, 862, 1359).

Order Ephemeroptera—Mayflies

Mayflies are frail, delicate insects with medium-size soft bodies that end in three
long threadlike tails, or caudal setae. The wings are membranous and many-veined
and are held upright while at rest. The front pair is large and triangular: the hind

44

pair, when present, is small and rounded. The antennae are bristlelike and in-
conspicuous; the mouth parts, vestigial. The nymphs are elongate and cylindrical or
flattened, and have leaflike gills along the sides of the body. In most species the
gills have three long tails.

Mayfly nymphs live under stones or among debris on the bottom of streams
where they feed on decaying vegetable matter, algae, and diatoms. When they
become full grown they leave the water and transform to adults on nearby vegeta-
tion. The adults occasionally appear in enormous numbers, but they seldom live
longer than a day to two. Occasionally, their dead bodies literally pile up along the
shore, on bridges, or in the streets of nearby towns (9//).

Beyond the occasional nuisance created by the presence of piles of dead insects in
areas frequented by humans, and the importance of the nymphs as fish food, which
is substantial, mayflies are of no economic importance.

Order Odonata—Dragonflies and Damselflies

Dragonflies and damselflies are relatively large and often beautifully colored
insects. The adults have two pairs of elongate, membranous, many-veined wings of
about equal size. The head and the compound eyes are large, the antennae are very
small and bristlelike, and the abdomen is long and slender. Dragonflies may be up
to 75 mm long and they hold their wings in a horizontal position while at rest.
Damselflies are usually somewhat smaller. Their wings are folded along the
abodomen or are tilted up while at rest. Adults of both groups feed on various
insects they capture while in flight. They are common and often abundant around
slow streams and ponds. Dragonflies are particularly noticeable because of their
large size and their rapid flight back and forth over the water.

Dragonfly and damselfly nymphs are all aquatic and feed on various small
aquatic organisms. Prey is captured through the use of a modified labrum contain-
ing two movable clawlike lobes at the tip. This device is held folded under the head
when not in use and is about one-third as long as the body when fully extended. The
nymphs breathe by means of gills. In the dragonflies, gills are in the rectum; in
damselflies, gills are three leaflike structures at the end of the abdomen. Mature
nymphs crawl out of the water and transform to the adult stage, usually on rocks or
vegetation.

Order Plecoptera—Stoneflies

Stoneflies are small to medium-size drab insects with soft flattened bodies. The
wings are membranous and have numerous cross veins. The front pair is rather
narrow and elongate; the hind pair is shorter but wider and is usually folded in
pleats while at rest. The antennae are long, slender, tapering, and many-segmented.
The cerci, when present, are usually long and many-jointed. The nymphs are flat-
bodied and somewhat elongate. They have long antennae, long cerci, and branched
gills on the thorax and about the base of the legs.

Stonefly adults occur near streams or along rocky shores of lakes. The nymphs
are aquatic and are usually found under stones in rapids of streams. When they
reach maturity, they leave the water and climb up on nearby objects to transform to
the adult stage (909).

Stoneflies may be a nuisance at times, especially when they appear in swarms in
recreational areas. The nymphs are an important source of fish food.

45

Order Psocoptera—Booklice and Psocids

Booklice and psocids are small, soft-bodied, winged or wingless insects, usually
less than 6 mm long. The more typical psocids have well-developed wings and bear
a striking resemblance to aphids of the order Homoptera. The wings are held
rootlike and almost vertically over the body while at rest. Booklice are either
wingless or possess only vestigial wings, and are about | mm long.

Psocids are found under stones, on or under the bark and on the foliage of trees or
shrubs. They are not injurious to trees but they may be a nuisance, especially when
they occur in large numbers around residences or in recreational areas. They feed
on fungi, lichens, and probably other vegetable matter.

Booklice occur most commonly in damp, dark rooms not generally used. They
are occasionally found in old books, where they feed on the paste of the bindings.
Sometimes they are abundant enough to cause serious damage.

Order Mallophaga—Chewing Lice

Chewing lice are all external parasites of birds and animals. The adults are small,
usually flattened, and wingless. They feed on the feathers, hairs, or skin of their
hosts. None is known to attack people. The order is divided into four families.
Members of the family Trichodestidae attack various species of domestic animals,
Gyropdidae feed on rodents, and the other two—Menoponidae and Philop-
teridae—feed on birds and poultry.

Order Anoplura—Sucking Lice

Sucking lice are small, wingless insects that live on the skin of various mammals
and suck their blood. Their bodies are flattened, the mouth parts consist of piercing
stylets, and there is a rostrum with many tiny hooks at the front of the head. The
tarsus consists of a single large claw. This claw is opposed by a toothed projection
on the tibia. The body louse, Pediculus humanus humanus L., and the crab louse,
Pthirus pubis (L.), attack humans, and a number of other species attack various
kinds of livestock and other animals. Published treatments of the order (390, 4/0)
are available, as are discussions on control of species attacking people (2/9, 370).

Order Thysanoptera—Thrips

Thrips are small slender-bodied insects, usually from 0.05 to 5.0 mm long.
Adults are either wingless or have four long, narrow, fringed wings with few or no
veins. The mouth parts are of the rasping, sucking type; the antennae are usually
short and 6- to 10-segmented; the tarsi are 1- to 2-segmented, with one or two
claws, and are bladderlike at the tip.

Thrips are frequently extremely abundant on flowers. Others occur on foliage,
fruit, bark, fungi, and in debris. A number of species cause considerable damage to
cultivated plants, but only a few have been reported injurious to trees (38).

Liothrips umbripennis (Hood) is sometimes abundant enough on chestnut oak to
cause the curling of leaves. The slash pine flower thrips, Gnophothrips fuscus
(Morgan), was first reported as damaging to pine seedlings in nurseries in New
York and Rhode Island and to jack pine on rocky slopes in Ontario. It is known as a
serious seed orchard pest of slash pine in Florida and southern Georgia (356, 547).
It infests the female flower buds, flowers, and very young conelets, puncturing and

46

abrading scales and bracts. Heavy infestations result in obvious resinosus and death
of flowers or conelets. Less intensive attacks kill scattered cone scales and result in
gnarled, distorted cones in which seed loss is both direct from lost cone scales and
indirect from poor seed release from the deformed cones. The flower thrips,
Frankliniella tritici (Fitch), feeds on the flowers and flower buds of hawthorn,
which sometimes prevents the buds from opening.

Order Neuroptera—Dobsonflies, Lacewings,
Antlions, and Allies

This order contains a wide variety of terrestrial and aquatic insects. The adults
have two pairs of large, membranous, leaflike wings that they hold rooflike over the
abdomen while at rest. The antennae are generally long and many-segmented; the
tarsi are five-segmented and there are no cerci. The larvae are practically all

campodeiform and are usually armed with very large, curved mandibles.
Family Corydalidae
Dobsonflies

The best known species of this family is the dobsonfly, Corydalus cornutus (L.).
The adult, especially the male, occasionally reaches a length of 100 mm, has two
long curving pincers or mandibles, and has a wingspread of 100 to 125 mm.
Females are similar in appearance except that the mandibles are smaller. The full-
grown larva, commonly known as hellgrammite, is also large and formidable in
appearance. Hellgrammites are found under stones in stream beds, especially
where the water runs swiftest. After about 22 years, they leave the water and
construct cells in which to pupate under stones, logs, or other objects on or near the
bank of the stream, usually during early summer. The hellgrammites are highly
prized as fish bait; otherwise, members of the family are of little or no economic
importance.

Family Chrysopidae
Green Lacewings

In both the adult and larval stages, members of the family Chrysopidae are all
predacious on soft-bodied insects. They occur commonly in late summer and fall on
the foliage of plants infested with these insects. Aphids and mealybugs appear to be
preferred as hosts, but leafhoppers, thrips, mites, and certain species of scale
insects are also attacked. The adults are green or yellowish green and have delicate,
lacelike wings. Egg are usually laid at the ends of 3- to 4-mm-long gelatinous stalks
firmly attached to the surfaces of leaves. The larvae are elongate, yellow or gray
mottled with brown, and taper toward each end. Some species have the odd habit of
covering their bodies with packets of trash woven together loosely with strands of
silk. The winter is spent usually as full-grown larvae in silken cocoons in bark
crevices or in such protected places as piles of leaves on the ground. There are one
to several generations per year, depending on climate (////). The goldeneye
lacewing, Chrysopa oculata Say, is an important predator of spruce gall adelgids in

the Lake States.
Family Hemerobiidae
Brown Lacewings

Brown lacewing adults have brown or dark-colored bodies often marked with
yellow; occasionally the abdomen is pale yellow. Otherwise, they resemble adult
green lacewings very closely. All species are predacious on other insects, prin-
cipally aphids, but also adelgids, mealybugs, whiteflies, mites, and occasionally
diaspine scales. The larvae are similar in general appearance to the larvae of green

47

lacewings, but they do not carry packets of trash on the dorsum. Eggs are laid on
the surface of leaves. Depending on the species, winter is spent in the larval, pupal.
or adult stage. The number of generations per year varies from one to many,
depending on species and climate.

Family Myrmeleontidae

Antlions

Antlion larvae, or doodlebugs, as they are also commonly called, live in tiny,
conical pits or craters in the ground in dry, dusty, or sandy areas. The pits are
usually about 37 to 50 mm wide and from 25 to 50 mm deep. The sides slant
sharply from the rim to a point in the bottom. The adults have long slender bodies
and two pairs of long, narrow, delicate, many-veined wings; larvae are broad,
somewhat flat, taper toward each end, and have long, curved mandibles armed with
strong spines and setae. The larva lies hidden under sand at the bottom of its pit and
feeds on ants or other insects that fall into the pit.

Order Siphonaptera—F leas

Fleas are small, wingless, hard-bodied, jumping insects. The body is strongly
flattened laterally and is armed with numerous backward-projecting spines or
bristles. The mouth parts are formed for sucking and the legs are long; there may be
no eyes. Adults feed on the blood of birds, wild and domestic animals, and people.
The larvae feed on organic matter, their own cast skins, and the feces of the adults.
Many species are economic pests. About 75 species of animals and birds in the
Eastern United States are attacked by more than 50 different species of fleas (439).

Some of the more important eastern species are: the human flea, Pulex irritans
(L.); the cat flea, Crenocephalides felis (Bouché): the dog flea, C. canis (Curtis);
and the oriental rat flea, Xenopsylla cheopis (Rothschild). The latter species 1s the
principal vector of bubonic plague and may also transmit endemic typhus to
humans.

Order Orthoptera—Grasshoppers, Crickets,
Mantids, and Allies

The order Orthoptera contains some of our most familiar insects, and many of
them are important household or agricultural crop pests. A few species are also
injurious to trees.

Members of the order may be either winged or wingless. The winged forms have
two pairs of wings, with the front pair generally long and narrow, many-veined, and
leathery or parchmentlike. The hindwings are membranous, much broader, and are
usually folded in fanlike pleats beneath the front wings while the insect is at rest.
The body is elongate, and cerci are usually well developed. The females of many
species have long ovipositors, often as long as the body.

Many publications have been issued on the Orthoptera (7, /29, 453, 455, 542,

543, 544, 545, 889, 1016, 1226, 1317).
Family Mantidae
Mantids

Mantids are predacious and feed on a wide variety of other insects. They are
large, elongate insects. The eyes are very large; the head is wider than it is long,
and movable; and the prothorax is very long—sometimes nearly as long as the
remainder of the body. The front coxae also are very long and the front femora and
tibiae are armed with strong spines that fold together to form a pincer. To capture

48

their prey, they usually lie in wait for it, holding their front legs in an upright
position. Once the prey comes within reach, the armed tibiae and femora shoot out
with lightninglike speed to grasp it. Because of their habit of holding their front legs
in an upright position, these insects are commonly called “praying mantis.”
Mantids lay their eggs in the fall in papier-maché-like egg cases or ootheca, each of
which contains 200 or more eggs; hatching occurs the following spring. There is
one generation per year.

The Carolina mantid, Stagmomantis carolina (L.), is the most common species
in the Southern States. It occurs from the Atlantic Coast to New Mexico and north
to Nebraska, Pennsylvania, and Illinois. Adults are 75 to 100 mm long. The male is
grayish brown with smoky-brown outer wings and often with a greenish-yellow
body and legs. Females either are colored like the males or are greenish yellow with
bright-green forewings. S. floridensis (Davis), a Somewhat more slender and longer
species, occurs in Florida.

The Chinese mantid, Tenodera aridifolia sinensis (Saussure), an introduced
species, occurs in the Eastern States west to Ohio and south to South Carolina. The
adults are elongate, robust, and about 100 mm long (fig. 11). Females are green or
greenish yellow; males are the same color or wholly brown, or brown with green
margins on the forewings. The narrowwinged mantid, 7. augustipennis (Sau-
ssure), also an introduced species, is similar to but more slender and smaller than
the Chinese mantid. It is widely distributed in the Eastern States.

Courtesy Conn. Agric. Exp. Stn.

Figure 11.—Adult and egg mass of the Chinese mantid,
Tenodera aridifolia sinensis.

Two other eastern species are the European mantid, Mantis religiosa L., an
introduced species (medium size and greenish yellow) and Litaneutria minor

(Scudder). The latter is about 25 mm long, and occurs in the Great Plains (509).
Family Phasmatidae
Walkingsticks

Eastern species of walkingsticks are long, slender, and subcylindrical. The head
is free and nearly horizontal, the antennae are long, the eyes small, the abdomen is
elongate, the legs very long and slender, and the wings are absent except for one
species in Florida with rudimentary wings. Walkingsticks are slow-moving Insects,
and all are plant feeders. The eggs are hard-shelled and are often dropped or laid on
the ground.

49

The waikingstick, Diapheromera femorata (Say), the only species of economic
importance, occurs in southern Canada and throughout most of the Eastern United
States west to the Great Plains and Texas. Its preferzed hosts appear to be black oak,
northern red oak, basswood, elm, black locust, and cherry, but it also feeds on
white oak, quaking aspen, paper birch, ash, dogwood, and hickory.

Adults average 75 mm long and, while motionless, closely resemble the twigs of
their hosts (fig. 12). The body color is variable. Some individuals are all brown or
green; others are mottled or multicolored with dark or light shades of grays, greens,
reds, or brown. Newly hatched nymphs are pale green, about 8 mm long, and look
like miniature adults. The egg is very hard, oval, seedlike, shiny black or brown,
and has a broad white or olive band on one edge. It is about 2 mm long.

F-504382
Figure 12.—Adult walkingstick, Diapheromera
femorata.

Winter is spent in the egg stage and hatching occurs in May or early June. The
nymphs feed at first on shrubs such as sweetfern, blueberry, strawberry, and
serviceberry. Later, they feed on leaves of the same trees as the adults. Adults
emerge in July or August and feed and lay eggs until the onset of cold weather. In
heavily infested stands, the sound of falling eggs striking the ground is much like
that produced by raindrops in a light shower. In the South most of the eggs hatch the
following year; in the North most hatch the second spring following their deposi-
tion.

Severe outbreaks occur occasionally in the Lake States. They also occur less
frequently south of a line drawn from Nebraska to Delaware. Trees may be
defoliated twice in the same season during severe outbreaks. Branch mortality
sometimes occurs in stands heavily defoliated three or four times; continued
defoliation for several years may lead to considerable tree mortality (493). Impor-
tant natural control factors include the hymenopterous parasite, Mesitiopterus
kahlii (Ashmead); various predators such as crows, robins, and other birds; and dry
weather during the period of egg hatch (/322).

50

Several other species of walkingsticks also occur in the Eastern United States.
The twostriped walkingstick, Anisomorpha buprestoides (Stoll), is found in the
Deep South, typically in oak stands growing on excessively drained, sandy soil in
Florida. Eggs are laid in groups of 8 to 10, each in a small pit dug in the soil. A.
ferruginea (Palisot de Beauvois) feeds on various trees and shrubs from south-
eastern Nebraska and Arkansas through the high country to Georgia and the
Carolinas. Diapheromera velii Walsh and D. blatchleyi (Caudell) feed on grasses
and tall shrubs. D. velii occurs in the Great Plains; D. blatchleyi, from the Great
Plains to the Atlantic Coast. Megaphasma denticrus (Stal), the giant walkingstick,
sometimes attains a length of 150 mm. Its habits are similar to those of D. femorata,

but it 1s apparently never abundant enough to be injurious.
Family Acrididae
Grasshoppers

This family contains most of the well-known grasshoppers, some of which are
frequently destructive to agricultural crops. Ordinarily, they are not very injurious
to trees, but they may seriously damage them during outbreaks. Young trees in
nurseries, shelterbelts, and plantations are particularly vulnerable, especially in the
Great Plains, in the upper Mississippi Valley, and in the Lake States. The adults are
distinguished by their short filiform antennae of 25 segments, short and in-
conspicuous Ovipositors, auditory organs on the sides of the first abdominal seg-
ment, greatly enlarged hindlegs, narrow forewings, and broad, membranous, fan-
like hindwings.

The majority of grasshoppers breed and live in the same general area throughout
the year. Certain others may build up in such vast numbers that they are forced to
leave their breeding grounds. At such times, they may travel considerable dis-
tances. All species have much the same life history. Nearly all lay their eggs in pods
at depths of 25 to 75 mm in the soil in late summer or fall, usually in grain stubble,
meadows, and along ditchbanks, fence rows, and roadsides. Hatching occurs in the
South as early as February; in the Northern States, it usually occurs in May or June.
Maturity is reached in 40 to 70 days, after which some adults live and feed until the
onset of cold weather. The majority of species spend the winter in the egg stage, but
a few overwinter as nymphs or adults, especially in the South.

The following are a few of the species that may be injurious in nurseries,
plantations, and shelterbelts during epidemics. The migratory grasshopper,
Melanoplus sanguinipes (F.)—the adult 1s about 20 mm long and reddish brown
with a distinct patch of black on the neck or collar. The differential grasshopper,
M. differentialis (Thomas )—the adult is 37 mm long. It is yellow with contrasting
black markings; has clear, glossy hindwings; and usually bears yellow and black
chevron-shaped bars on the sides of the thighs of the hindlegs. The twostriped
grasshopper, M. bivittatus (Say)—the adult is about 30 mm long. It is slightly
more robust than the differential grasshopper and is greenish yellow beneath, with
two yellow stripes running the full length of the dorsum. The redlegged grasshop-
per, M. femurrubrum (De Geer)—the adult is about 18 mm long. It is reddish
brown above, yellowish beneath, has colorless hindwings, and has red-tinged
hindlegs. The clearwinged grasshopper, Camnula pellucida (Scudder)—the adult
is about 12 mm long and yellow to brown. The forewings are blotched with large
brown spots, and the hindwings are clear.

A few species of grasshoppers are more closely associated with trees than others.
One, Dendrotettix quercus Packard, the postoak locust, is widely distributed from
east-central Texas to the Lake States, Long Island, and southeastern Canada. Oaks

51

are its preferred hosts, but it has also been observed on pines in the Lake States.
Several outbreaks have been recorded, some of which covered several square
kilometers. During intervals between outbreaks, it is very scarce. Adults are present
from June to September and lay their eggs in the soil in late summer. The nymphs
climb the trees to feed. Other tree-infesting species are: D. australis (Morse)—
apparently feeds exclusively on Virginia pine in the Southeastern States:
Melanoplus punctulatus (Scudder)—has been observed defoliating young eastern
white pines in plantations in Connecticut: M. bruneri (Scudder)—inhabits co-
niferous forests; and the eastern lubber grasshopper, Romalea microptera (Palisot

de Beauvois)—feeds on shrubs in Fiorida and Alabama.
Family Tettigoniidae
Longhorn Grasshoppers and Katydids

Longhorn grasshoppers and katydids are mostly large with hairlike antennae,
four-segmented tarsi, laterally flattened bladelike ovipositors, and auditory organs
sometimes at the base of the front tibiae. The males “sing” by rubbing a sharp edge
at the base of one front wing along a filelike ridge on the ventral side of the other
front wing. The songs of different species differ in the character of the pulses, the
pulse rate, and in the way the pulses are grouped.

Tree-inhabiting katydids are usually long-winged and green, matching the color
of the foliage on which they feed. Eggs are laid end to end in overlapping rows on
leaves or twigs, or are inserted into the edges of leaves. The winter is usually spent
in the egg stage and hatching occurs in the spring. Some of the more common
species, none of which is very injurious, are discussed below.

The forktailed bush Katydid, Scudderia furcata Brunner von Wattenwyl, so-
called because of the forked appendages at the top of the abdomen of the male, is
widely distributed. It occurs on but is not restricted to trees. The related species, S.
curvicauda (De Geer), lives commonly on oak.

The broadwinged katydid, Microcentrum rhombifolium (Saussure), is widely
distributed in the East. Adults are 50 to 60 mm long and leaf green. The angular-
winged Katydid, M. retinerve (Burmeister), a smaller species, is more southerly in
its distribution. Both species have long, narrow wings, and the vertex is narrowed
anteriorly.

Pterophylla camellifolia (F.) is the katydid so commonly heard on summer
evenings. The adult is large, green, and robust. The front wings are dark green,
leaflike, very broad, concave within, and wholly enclose the abdomen. Infestations
occur as small colonies in the dense foliage of trees. Eggs are laid in crevices of
loose bark or within the soft stems of woody plants.

Other long-winged species include: Hubbellia marginifera (Walker), a large
species with green front wings, sometimes spotted with brown, which occurs on
pines in the Southeastern States, and Conocephalus brevipennis (Scudder), the
shortwinged meadow katydid, which lays its eggs in willow in Canada.

Some members of the family have functional wings and live only on the ground.
The most familiar species is the Mormon cricket, Anabrus simplex Haldeman.
Although primarily western in distribution, it occasionally occurs in destructive
numbers as far eastward as the Dakotas and Kansas. It feeds mostly on various trees
and shrubs.

Family Gryllidae
Crickets

Crickets are medium-size insects, usually with long filiform antennae, three-

segmented tarsi, spear-shaped ovipositors, and hindlegs fitted for jumping. The

52

wings of certain species are fully developed, and lie flat on the back and bend down
abruptly at the sides of the body. In other species, wings are either abbreviated or
absent. The winged forms possess auditory tympana on one or both sides of the fore
tibia. The males also possess sound-producing organs near the base of the dorsal
surface of the front wings.

There are several different kinds of crickets, the most familiar perhaps being the
common field and house crickets of the genus Gryllus (7). Tree crickets, however,
are the only members of the family that are injurious to trees or shrubs.

Tree crickets are small, delicate, and pale colored. The tarsi are three-segmented,
with the second segment small and compressed. The wings of the male are broad
and lie flat over the abdomen, while those of the female are narrow and wrapped
closely about the body. Tree crickets feed on other insects such as aphids, treehop-
pers, and scales; and also on the leaves, flowers, and bark of trees.

The snowy tree cricket, Oecanthus fultoni TY. J. Walker, is a common species
throughout the United States. Adults are pale green and about 14 mm long. The
wings are transparent with a slight greenish tinge, and each of the first two antennal
segments bears a black spot. Adults are found on various trees and shrubs,
preferring those growing in the open. Eggs are laid singly in a row of punctures in
the bark of twigs or small branches. O. exclamationis Davis is similar in ap-
pearance, habits, and range to O. fultoni except for a black club-shaped mark on the
base of the first segment of the antenna. O. pini Beutenmiuller occurs on pines and
lays its eggs in regular rows on the bark. O. latipennis Riley occurs commonly on
shrubs and low trees, especially on scrub oaks in dry open areas in the Eastern
States.

Anurogryllus arboreus T. J. Walker, the arboreal short-tailed cricket, occurs
mainly along the Atlantic Coast from New Jersey to Florida and westward to
southeastern Texas. It has been observed damaging newly germinated slash pine
seedlings in Louisiana, Texas, and Arkansas. It cuts off the seedlings, pulls them
into its tunnels, and eats the tender foliage (/238, 1259).

Family Gryllotalpidae
Mole Crickets

Gryllotalpa gryllotalpa (L.), the European mole cricket, an introduced species,
occurs in a number of places along the East Coast. Adults are brownish yellow
tinged with fuscous above and are pale, brownish yellow underneath. They are
covered with velvety hairs and are up to 37 mm long. The forelegs terminate in four
dactyls that are used for digging. This species feeds at night by tunneling in the
upper 25 to 50 mm of soil. It cuts off the roots of seedlings, eats pits in underground
roots and stems, cuts off stems above the ground, and eats seeds. It is occasionally
injurious in nurseries.

The changa, Scapteriscus vicinus Scudder, a tropical species, occurs along the
Coastal Plain of the Southeastern States. The adult is brown above, light brown
below, and about 37 mm long. Its forelegs terminate in two dactyls. S$. abbreviatus
Scudder occurs in Georgia and Florida. Adults are brownish fuscous and blotched
with yellow. The southern mole cricket, S. acletus Rehn & Hebard, occurs from
Georgia to Texas. It 1s pinkish buff in color and somewhat more slender than S.
abbreviatus. All members of this genus feed on earthworms, roots, and insects.
They are occasionally injurious in nurseries.

Family Blattidae
Cockroaches

Several species of cockroaches may be found under the bark and in cavities in
dead trees. They feed chiefly on animal and vegetable refuse and, as far as known,

2)

are of no economic importance to forestry. The adults are distinguished by their
depressed, oval bodies, their nearly horizontal heads, their slender, depressed legs
of almost equal size, and the absence of long ovipositors in the females.

Order \Isoptera—Termites

Termites constitute the order Isoptera. They are one of the oldest and most
primitive groups of insects. Termites are social insects and live in colonies. In all
species there is a definite caste system, with each of the adult castes performing an
essential function in the life of the colony. Termites utilize cellulose, which they
normally obtain from wood or other plant material, as their main energy source.
Cellulose is normally indigestible to most animals; however termites have single-
celled protozoa and bacteria within their digestive tracts that break cellulose into
digestible units (58S). Many species of fungi are associated with termites in a
beneficial manner. This association varies from little dependence on fungal activity
in wood to a totally symbiotic relationship with fungi (/053). The influence of fungi
on the activity of termites is poorly understood but some species, such as
Gloeophyllum trabeum (Pers. ex Fr.) Murr. (Lenzites trabea Pers. ex Fr.), are known
to produce an attractant and a feeding stimulant (S29, //2/).

Only 38 species of termites are believed to be native to the continental United
States. Of these, 14 are known to occur in the eastern half of the country, exclusive
of western Texas (/267). In addition to these native species, two introduced species,
Cryptotermes brevis (Walker) and Coptotermes formosanus Shiraki, occur in the
East. C. brevis may have been introduced into southern Florida. C. formosanus is
believed to have been introduced into the Gulf Coast during World War II.

Termites are small, cylindrical, soft-bodied insects sometimes without eyes. The
antennae are moniliform and, in the winged adults, the number of antennal seg-
ments varies from 12 to 25 or more. The median ocellus is absent. In certain
species it 1s replaced by a more or less distinct opening of a gland known as the
fontanelle. The wings are long and narrow and, when folded on the back, extend
well beyond the end of the abdomen. In all North American species, the forewings
and hindwings are similar in form and in the general features of their venation. The
abdomen is broadly joined to the thorax.

Termites have very few effective natural enemies that attack individuals in the
nest. For example, there are no known internal insect parasites, and predacious
enemies are largely limited to ants, lizards, and birds. During and after the
swarming period the winged adults of many species are eaten by all sorts of
animals, particularly other arthropods (scorpions, solpugids, spiders, centipedes,
dragonflies, cockroaches. mantids, crickets, beetles, flies, and wasps) and verte-
brates (fish, frogs, toads, lizards, snakes, birds, and mammals, including humans)
(741). These predators may have some adverse effect on the establishment of new
colonies but probably have no effect at all on the parent colony. As yet, there are no
studies on the rate of predation and its possible significance in controlling the
populations of a single species of termite (94/). Fungi, nematodes, and bacteria
usually kill a small percentage of the individuals in a colony. Occasionally molds
cause much heavier losses.

In many parts of the Eastern United States, particularly the South, termites are
almost universally present in stumps, logs, and other woody materials in contact
with the ground. Termites are largely beneficial in forested areas because of their
value in the decomposition of dead wood and its reincorporation into the soil.

54

However, when termites infest wooden structures and other useful wood products,
they are often extremely destructive. The extent of the monetary losses they cause 1s
not known. Estimates vary, however, from $100 to more than $750 million per year
in the United States for damage, repairs, and preventive and remedial control (645,
TN SSD)

For further information on termites, the reader is referred to the following

studies: 365, 366, 536, 566, 617, 645, 686, 695, 696, 720, 1266, 1316.
Family Rhinotermitidae

Subterranean Termites

The subterranean termites of the Eastern United States all belong to the family
Rhinotermitidae. Because these termites all require a constant supply of moisture,
their colonies are found either entirely or partly in the ground. They may, however,
feed in wood located some distance from the ground, but they always maintain
connection with the ground unless a continuous supply of water 1s otherwise
available. In order to attack wood located away from the ground where a supply of

water 1s not available, they construct covered passageways, commonly called
shelter tubes (fig. 13A).

WER

os
»
4
GB
{
a

F-531242
Figure 13.—Subterranean termite: A, shelter tube on

foundation wall; B, damage to a pine stake.

Regardless of the extent of damage they cause to the interior of wood (fig. 13B),
these termites always leave a covering shell intact. Because of this shell, there
usually is no external evidence of infestation, even though the interior of the wood
may be destroyed. The first indication of infestation may be the swarming of

55

winged adults, the presence of shelter tubes over foundation walls, or the sudden
collapse of the surface of infested wood.

The principal food of subterranean termites is cellulose, which they obtain from
wood and other plant tissues. As a result, these termites are not only destructive to
the woodwork of buildings, utility poles, fence posts, and other wood products, but
also to paper, fiberboard, and various types of fabrics derived trom cotton and other
plants. Shrubs, nursery stock, ornamental plants, and many kinds of shade trees are
damaged occasionally. Trees killed by other insects, fire, or disease, particularly in
the South, are attacked and the timber rendered unmerchantable unless it is utilized
shortly after being killed.

In attacking wood, subterranean termites usually feed on the soft springwood
only. As a result, their galleries run with the grain. These galleries are charac-
teristically stained on the inside with grayish specks of excrement and earth but are
free of pellets like those found in the galleries of nonsubterranean termites.

Subterranean termites occur throughout most of the United States and in southern
Canada. They are common in most of the eastern half of the United States and along
the Pacific Coast. It is considered likely that the native species have occupied their
present ranges for millions of years; however, there seems to be an increase in their
destructiveness northward. This increase has probably resulted from the general
adoption of central heating plants in structures, from changes in building practices,
the wider use of termite-susceptible wood in construction, and from the tremendous
expansion of suburban homes into forested areas.

The eastern subterranean termite, Reticulitermes flavipes (Kollar), is the
predominant species in the Eastern United States. It occurs from the East Coast to
the Great Plains and from the Gulf of Mexico north to Ontario. In general, R.

flavipes occupies regions of high humidity. Dead wood in forested areas 1s com-

monly infested with this species. It will attack wood from most of the species of
trees in this part of the country. Infestations also occur in the woodwork of
buildings, in cellulose materials in storage, and in poles and posts.

Every mature colony of the eastern subterranean termite contains some or all of
seven castes during the year: larvae or immatures, nymphs, soldiers, workers (a
“true” worker caste is not believed to exist in Reticulitermes), alates or winged
reproductives, and primary and supplementary (or replacement) reproductives.
Larvae hatch from the eggs and are precursors of all the other castes (fig. 14A).
Workers (fig. 14B) are soft-bodied, grayish white, and slightly less than 6 mm
long. They are the individuals responsible for the maintenance of the colony and the
gathering of food. Soldiers (fig. 14C) are similar to the workers except for their
much larger, long heads, their larger and more formidable mandibles, and their
slightly greater length. Soldiers cannot feed themselves but contribute to colony
labor by defending it. Nymphs (fig. 14D) develop from larvae and eventually molt
into reproductives (fig. 14E). These reproductives are soft-bodied and brown to
black, have two pairs of long, whitish opaque wings of equal size, and are 10 to 12
mm long. The winged reproductives fly from their colony to form new colonies.

Colonizing flights occur during the day, most frequently after the first warm days
of spring, often following a warm rain in April or May. They may also occur during
the remainder of the warm season and sometimes even in the fall, especially in the
South. In heated buildings, flights occasionally occur during the winter. The
individuals in these flights are attracted by strong light. When they emerge within
buildings, they gather about windows and doors. Here, they soon shed their wings,
which may be found in large numbers even after the termites have disappeared.

56

Soon after shedding their wings, males and females pair off and search for a place
to form a new nest. The great majority of them are usually unsuccessful, but some
survive and manage to hollow out small cells in the ground near wood. They enter
the cell, seal the opening, and eventually mate.

A,B,E,F: F-531243
D: F-531902
C,G: Courtesy South. For. Exp. Stn.,
: Gulfport, Miss.
Figure 14.—Eastern subterranean termite,
Reticulitermes flavipes: A, \arvae or immatures; B,
worker; C, soldier; D, nymph; E, winged primary
reproductive; F, physiogastric female primary
reproductive; G, female supplementary reproductive.

a7

A few days after mating, the female lays from 6 to 12 eggs. Hatching occurs in
about 50 to 60 days, and the young larvae grow to maturity in less than | year. Later,
both soldiers and reproductive nymphs appear in the colony and reach maturity
within | and 2 years, respectively. Mating continues at irregular intervals, and the
colony continues to increase in size. The original pair of reproductives lives
together for life, sometimes for many years. The female increases greatly in size
(fig. 14F) and produces literally thousands of eggs in her lifetime. In well-
established colonies, there may be hundreds of thousands of individuals. In very
large colonies, supplementary reproductives (fig. 14G) will develop in a colony to
produce eggs if the primary reproductives die. These large colonies may be so
widely spread out that it is difficult or impossible to estunate their size or to locate
their main parts.

Reticulitermes tibialis Banks is the most widely distributed species of Re-
ticulitermes in North America. It is largely confined to the arid areas of the western
half of the United States but also occurs eastward in the North Central States as far
as the Chicago area. To the east, R. tibialis overlaps a portion of the ranges of R.
flavipes, R. hageni Banks, and R. virginicus (Banks). Along the southern shores of
Lake Michigan, R. tibialis coexists with R. arenincola Goellner among sand dunes,
where it infests small pieces of wood partly buried in the sand. Farther to the
southwest, in Kansas and Texas, it occurs in heavily sodded prairies and in hard-
packed and often alkali soil.

While R. tibialis is fully capable of damaging buildings, it is not as economically
important as R. flavipes because of the thinly populated regions in which it occurs.
Flights of this species occur over a large part of the year during the fall, winter, or
spring, depending on the locality. Winged adults, in general, are almost entirely
shiny black and about 10 mm long. The tibiae are also blackened and the pronotum
is broad.

Reticulitermes virginicus occurs in the southeastern and central portions of the
United States. Its range coincides closely with that of R. hageni in the east,
although it does not appear to extend as far to the west. It is often mistaken for R.
flavipes but is smaller. Throughout its range, R. virginicus overlaps the region
occupied by R. flavipes, but it does not extend into the northern regions occupied
by this latter species. In general, the line of northward extension of R. virginicus
coincides with the regions where the average annual minimum temperature does not
fall below — 23° C. Westward, this species extends through southern Illinois and
Missouri, south to the coast of the Gulf of Mexico, as far west as Houston, Tex.,
and eastward to the southern tip of Florida (/267).

Reticulitermes hageni is a small yellow-brown species and is the most distinctive
of the Reticulitermes. It occupies about the same territory as R. virginicus. These
two species are reported to occur more commonly in drier forested areas than does
R. flavipes.

Reticulitermes arenincola was described from collections from the sand dunes
along the southern shore of Lake Michigan in Indiana and Michigan. Little has been
detailed regarding the field biology of this species. It inhabits sandy areas, and its
flights occur toward the end of May.

Flights of R. virginicus tend to occur later in the spring than those of R. flavipes,
although some overlap occurs between these two species. Flights of R. hageni tend
to occur later than those of R. virginicus, although, again, there is some overlap
between species. Flights of R. virginicus in the southern portion of its range occur

58

earlier than flights in the northern portion, even within buildings. This is not the
case with in-building flights of R. flavipes (1267).

All species of Reticulitermes that occur in the Eastern United States are eco-
nomically important with the possible exception of R. arenincola. The taxonomy of
the genus is not well established. Although there can be little doubt that there are
physiological, behavioral, and ecological differences among species, there is no
evidence to suggest that control methods for one should be different from control
methods for another. One should be certain of the genus but, except as a matter of
interest, the particular species cannot now be considered critical from the stand-
point of control.

The Formosan subterranean termite, Coptotermes formosanus Shiraki, is an
introduced species. Active colonies were first recorded in North America in a
shipyard warehouse in Houston, Tex., in 1965. Since then additional colonies have
been discovered in Houston and Galveston, Tex.; Charleston, S.C.; and New
Orleans and Lake Charles, La. The species was described from individuals col-
lected in Formosa in 1909. Since then it has been reported from Ceylon, China,
Guam, Hawaii, Japan, and South Africa. Incipient colonies generally start in or
near the soil where moisture and damp wood are available. Nests are normally built
in soil near the base of tree stumps, utility poles, or other underground food
sources, but they may be found almost anywhere if conditions are favorable.
Colonies have been observed on boats, ships, barges, dredges, water tanks, piers,
floating drydocks, in living and dead trees, and in buildings.

Workers are grayish white overall. The head is pale yellow, with a white mark in
the center and a dark-brownish spot on each side of the clypeus. The pronotum is
nearly twice as broad as it is long, and the legs are slender and hairy. Soldiers (fig.
15) have oval heads and slender bodies. The pronotum is short, elliptical, and
notched at the middle of the frontal margin. A small, short tubelike process extends
from the frontal gland and exudes a milky, acidulous secretion. The abdomen is
slender, entirely pubescent, and bears a pair of three-segmented caudal appendages
(cerci). Alates have large, hyaline wings, 10.6 mm long, about three times as long
as the abdomen, and about three times as long as they are broad. The head is
hexagonal and brown; the frons 1s irregularly concave at the center where a globular
projection occurs; the pronotum is semicircular and as broad as the head; the legs
are short, heavy, and yellowish brown; and the abdomen is short, elliptical, and
yellowish brown. The bodies are marked with minute spots, and the wings are 12 to
14 mm long.

Nests of Formosan subterranean termites are constructed of a friable material
called carton. It consists of a mixture of masticated wood, saliva, and excrement. A
nest may be several cubic feet in volume. Tunnels radiate from nests constructed in
the soil. They can be found at depths of 3 m in the earth and extend horizontally up
to 60 m (665). Their walls are lined with essentially the same materials used in
constructing the nest and are nearly impervious to water. Primary queens can lay up
to 1,000 eggs per day, and a single colony may contain many hundreds of thousands
of individuals. Winged reproductives swarm during May and June. They are poor
fliers, and the majority drop to the ground within 90 m of the nests unless carried
farther by the wind. First evidence of a colony’s presence may be the appearance of
these winged adults at swarming time.

Large colonies can cause severe damage in a short time (fig. 16A). In Hawaii,
walls of new buildings have been hollowed out in 3 months’ time. Living trees are
also hollowed out (fig. 16B) and weakened. Known susceptible trees in the South

59

are the Chinese elm, several species of oak, hackberry, and velvet ash. Dead trees
are highly susceptible. Extensive damage all the way to the top of 20-m-tall
baldcypress snags has been observed in Louisiana (77).

F-531901

Figure 15.—Soldier of the Formosan
subterranean termite,
Coptotermes formosanus.

F-519935, 519934

Figure 16.—Formosan subterranean termite colony and
damage: A, carton nest and damaged wood above a
window inside a building; B, colony of workers in a
cypress log.

Families Kalotermitidae and Rhinotermitidae
Nonsubterranean Termites

Many species of nonsubterranean termites occur in the Eastern States. They are
found throughout Florida, also in a narrow strip along the Atlantic Coast as far
north as southeastern Virginia, and westward along the Gulf Coast to Mexico.
Infestations are found in structural timber and other woodwork in buildings,
furniture, utility poles, wooden derricks, piles of lumber, wood pulp or fiber

60

insulation boards, in other products containing cellulose, and in trees and other
plants. Because of their ability to live in wood that is frequently moved, nonsubter-
ranean termites are often found in regions far removed from their normal range,
including Canada. None of these termites is able to establish permanent infestations
if transported out of its normal range (366, 1/24).

Nonsubterranean termites usually are less injurious than subterranean species in
the Eastern United States. However, many years ago, large buildings in the South
were seriously damaged (//22). Nonsubterranean termites fly directly to and enter
untreated and unpainted wood at swarming time. They cut across the grain of the
wood and excavate broad pockets or chambers connected by tunnels of small
diameter. They feed on both the soft springwood and the harder summerwood.
Their cavities contain compressed fecal pellets which often have six darkened
grooves along their length (fig. 17). Some of these pellets are pushed to the exterior
through small holes and are found in piles on the floors of infested buildings.

F-519928
Figure 17.—Fecal pellets of nonsubterranean termites.

Nonsubterranean termites have a caste system similar to that of Reticulitermes.
However, few of the species, except Prorhinotermes, contain individuals that could
be considered workers. Most of the termites present are larvae or nymphs, and most
of the work of the colony is performed by the nymphs.

Winged adults usually swarm during the early evening or morning hours in late
winter or spring. Unless carried by the wind, they fly for only short distances in
search of places to start a new colony. Once they have succeeded, they shed their
wings and bore directly into the wood. Then they plug the opening, and seal
themselves in. Colonies grow very slowly, are seldom confined to a single chamber,
and rarely contain more than a few thousand individuals. Piles of pellets on the
floor may be the first evidence of infestations. Other telltale evidence consists of
pitted and roughened surfaces of infested floors, doorframes, and other wood.

Cryptotermes brevis (Walker) (Family Kalotermitidae) occurs commonly in
southern Florida and is fairly widespread (although not abundant) on the coast of
the Gulf of Mexico as far west as Brownsville, Tex. Isolated infestations have also
been found elsewhere, including Tennessee, Maryland, Ohio, and New York. In
most cases these infestations have been directly linked to transport of furniture from
the Southern United States, particularly Florida and Hawaii, and from the Orient.
Quarantine officials commonly intercept this species in cargo entering the United
States from tropical and semitropical areas of the world. C. brevis has not been

61

recorded in any natural habitat, but only in fabricated structures. It can attack
buildings but is particularly noted for severe damage to floors, woodwork, and
especially furniture.

Cryptotermes cavifrons Banks (Family Kalotermitidae) occurs in southern Flor-
ida in dead trees, logs, stumps, and branches. Unlike C. brevis, which seems to be
confined to structures, C. cavifrons has rarely been associated with buildings or
furniture. The usual habitat of this species is in dry, sound, hardwoods in the
hammock areas in southern Florida (/267).

Incisitermes snyderi (Light) (Family Kalotermitidae) 1s the common dry-wood
termite of the Southeastern United States. It occurs from South Carolina to Florida
and west to Brownsville, Tex., mainly along the coast, but as far north as Alex-
andria, La. Infestations occur in the woodwork of buildings, in untreated utility
poles, and in dead trees, logs, and branches. It does not appear to have the wide
ecological range of 7. minor (Hagen) (Family Kalotermitidae), which occurs in
western North America. Collection records suggest that this species is not as
subject to transport and survival in new localities as are 7. minor and C. brevis. It
does not seem to be as great an economic problem, generally, as is 7. minor on the
West Coast.

Incisitermes schwarzi (Banks) (Family Kalotermitidae) is the common dry-wood
termite in southern Florida, occurring also in the northern portion of the State. It is
common in coastal and everglade hammocks, and has a fairly high moisture
requirement. Infestations are found in the woodwork of buildings and in dead trees,
logs, and stumps. Two rather distinctive forms of soldiers (“‘longheaded” and
“shortheaded”’) occur in colonies of this species (/267).

Incisitermes milleri (Emerson) (Family Kalotermitidae) is a very small termite
that occurs on the southern Florida Keys. It is encountered in very dry wood
(1267).

Kalotermes approximatus Snyder (Family Kalotermitidae) occurs primarily in
northern and central Florida. Other scattered collection records of this species
suggest it may have a much wider range than generally believed and perhaps is not
frequently collected because it does not frequent houses as much as some of the
other dry-wood termites. It has been reported to occur in dead wood of oak,
sweetgum, and magnolia in Florida. Apparently it occurs in dead wood and in dead
areas of living trees. There are few flight records for this species. It is reportedly a
diurnal flier, as its deep pigmentation suggests, which is not the normal case for
dry-wood termites (/267).

Neotermes castaneus (Burmeister) (Family Kalotermitidae) is a very large spe-
cies that occurs in the southern portion of Florida and on the eastern Keys. It has not
been found in woodwork in buildings. Infestations have been limited to dead wood
of trees, logs, stumps, and branches, and to living citrus trees. N. jouteli (Banks)
(Family Kalotermitidae) occurs in the coastal areas of southern Florida, including
the Keys. It is occasionally found in moist foundation timbers of buildings, but in
nature it lives in dead trees and in logs and branches lying on the ground. Both of
these species have a fairly high moisture requirement. Their workings are often
situated in fairly damp wood or in dead wood adjacent to the living wood in living
trees.

Calcaritermes nearcticus (Snyder) (Family Kalotermitidae) is also known only
from Florida. It is similar to Cryptotermes; however, it occurs in damper wood and
lines some of its galleries with brownish material.

62

a

Prorhinotermes simplex (Hagen) (Family Rhinotermitidae) is found only in
Florida in the United States. It has a very limited distribution on some of the
southeastern Keys and on the mainland occurs only in Dade County. [t occurs in
mangrove swamps on the coast and in pine forest areas in the lower Everglades. It is
fairly common in the Miami area but is not found north of Fort Lauderdale. It has
not been collected more than 10 km from the coast. This coastal distribution is
typical of this genus throughout the world (/267).

Order Hemiptera—True Bugs

The order Hemiptera consists of a large and widely distributed group of insects.
The majority of species are terrestrial in habit, but several are aquatic. Many of the
terrestrial forms are phytophagous and feed on a wide variety of trees and smaller
plants. Trees, fortunately, are seriously damaged by only a few species. The order
also contains a large number of predatory species, many of which feed on other
insects and their eggs. A number of others feed on blood of humans and other
animals and are decidedly obnoxious or harmful, especially those that transmit
disease-causing organisms.

The majority of the Hemiptera have the basal portion of the forewing thickened
and leathery, and only the apical portion is membranous. It is from this “‘half-
wing” appearance that the order gets its name. The hindwings are entirely mem-
branous and usually slightly shorter than the forewings. Both pairs of wings lie flat
over the abdomen with the membranous distal portion of the front ones overlap-
ping. The mouth parts consist of a bundle of stylets inside a segmented sheath. This
slender beak arises from the front part of the head and usually extends backward
along the underside of the body, sometimes to the base of the hind pair of legs.
Feeding is accomplished by inserting the stylets into the tissues of the plant or
animal host and sucking up the juices or blood. The beak sheath folds back beneath
the insect as the stylets pierce deeper into the tissues. The antennae are fairly long
and consist of four or five segments. Many species have scent glands which give off
offensive odors when the insect is disturbed. Comprehensive studies have been

made of the Hemiptera of eastern North America (/30, 157).
Family Pentatomidae
Stink Bugs

Pentatomids are commonly called “stink bugs” because of the disagreeable odor
they emit when disturbed. The adult is usually a broad, short, slightly convex
insect. The antennae are five-segmented, and the head and thorax sometimes form
a triangle. Many species are brightly colored or conspicuously marked. Green ones
may be difficult to detect on leaves. Dark-gray ones are also difficult to see when
they rest on the bark of limbs and trunks of trees. Most species are plant feeders, but
some feed on other insects, and some feed on both.

The shieldbacked pine seed bug, Jetyra bipunctata (Herrich-Schaffer), occurs
on loblolly, slash, shortleaf, longleaf, Virginia, eastern white, red, jack, and sand
pines and is an important pest in southern pine seed orchards (286, 360). The adults
and nymphs are oval and have a humpbacked appearance. The adults are about 15
mm in length. Their color varies from gray-brown in the early stages to a dark
reddish-brown with black markings as adults. Their shape and color provide
concealment while they feed on nearly mature cones. When motionless, they are
not readily noticeable. The eggs are small, green spheres 1.5 mm in diameter.
About a dozen eggs are laid in two alternate rows along a single needle or in a group

63

on a cone scale. Nymphs of all stages are gregarious. There is only one generation
each year. Nymphs and adults have piercing-sucking mouth parts that they insert
into cones to penetrate the seeds. Nymphs in the third stage and older, as well as
adults, destroy seeds in developing cones. Most of the seed damage occurs in late
summer and fall and can be detected on radiographs (287). In seed orchards, losses
are reflected by poor seed viability and low yields of sound seeds per cone.

Other plant-feeding species and their hosts include: Brochymena quadripustulata
(F.), the fourhumped stink bug—feeds through the bark of limbs and trunks of
elm, oak, and willow. Brownish, hairlike lines running across the grain in the
cambial region are evidence of its attack. Adults frequently overwinter in houses. B.
carolinensis (F.)—feeds through the bark of slash pine and longleaf pine. Elas-
muche lateralis (Say)—feeds on the leaves and catkins of yellow birch. Pitedia
uhleri (Stal) —feeds on the larger branches and trunks of eastern white pine in the
Lake States.

Predacious species include the spined soldier bug, Podisus maculiventris (Say),
which has been recorded feeding on more than 30 species of destructive insects,
many of which are forest pests. The adult is dull yellow above, with numerous dark-
brown punctures, and is about 12 mm long. Also, Stiretus anchorago (F.) feeds on
the larvae of many species, including the gypsy moth and tent caterpillars. Adults
are shiny, dark metallic, and 8 to 11 mm long.

Family Tingidae
Lace Bugs

Lace bugs, so-called because of their broad, lacelike forewings, are usually
whitish and 5 to 6 mm long. They are flat, oval or rectangular, and the head ts often
hidden beneath a large hood on the front of a greatly modified pronotum, which
projects beyond the sides of the body. The abdomen is completely beneath the
forewings, which are frequently transparent. The nymphs are black and often
covered with long spines.

Many species of lace bugs live and feed on the undersurfaces of leaves. A given
species usually occurs either on a single host or on closely related ones. The upper
surfaces of infested leaves become whitened or brownish, or dead in appearance.
The undersurfaces are speckled with eggs, excrement, and cast skins of the
developing nymphs. The leaves of heavily infested trees may turn entirely brown
and fall off. The winter is spent as adults under bark scales or other cover on the
host tree, or as eggs cemented to the undersurfaces of leaves or embedded in leat
tissues. Most species have two generations a year.

The sycamore lace bug. Corythucha ciliata (Say), occurs throughout the East-
ern United States (7235) and in southern Canada. Its preferred host is sycamore,
but it also feeds occasionally on ash, hickory, and mulberry. The adult (fig. 18) is
white and about 3 mm long. Overwintering adults emerge early in the spring and
deposit their eggs along the ventral surface of the midrib of a leaf. Hatching occurs
in 2 to 3 weeks, and the nymphs feed for 5 or 6 weeks. There are two generations
per year in the North, probably more in the South. Light feeding causes a stippling
on foliage. Heavily infested leaves of sycamore turn white and drop prematurely.
During dry weather this may result in severe injury.

The oak lace bug, C. arcuata (Say), feeds on white, bur, and chestnut oaks from
Alabama and the Carolinas to southern Canada. Its life history and control have
been described (233). The winter is spent in either the egg or adult stage. Infested
leaves appear grayish white. Heavily infested trees may be defoliated, especially
during dry weather. Bur oak in shelterbelt plantings is especially susceptible.

64

Ce — —=E— eee

Courtesy Conn. Agric. Exp. Stn.

Figure 18.—Adults of the sycamore lace bug,
Corythucha ciliata.

The elm lace bug, C. u/mi Osborn & Drake, feeds on American elm in many
Eastern States and southern Canada, and on Siberian elm on the northern Great
Plains. It 1s capable of defoliating its host.

Heavy infestations of C. mollicula Osborn & Drake may seriously injure willow,
its only known host, throughout the Eastern States. Heavy infestations of the
hackberry lace bug, C. ce/tidis Osborn & Drake, often occur on hackberry in the
Midwest. The species has also been reported from Florida. C. pallipes Parshley, the
birch lace bug, is often abundant on young yellow birch. Other hosts include paper
birch, beech, eastern hophornbeam, willow, mountain-ash, and maple. C. per-
gandei Heidemann feeds principally on alder and occasionally on hazel, elm, and
birch. C. pruni Osborn & Drake feeds on pin cherry; C. juglandis (Fitch), the
walnut lace bug, on black walnut, butternut, and basswood; C. elegans Drake on
willow, balsam poplar, quaking aspen, and bigtooth aspen; C. aesculi Osborn &
Drake on buckeye; C. associata Osborn & Drake on pin cherry; C. bellula Gibson
on hawthorn; and the hawthorn lace bug, C. cydoniae (Fitch), on hawthorn and
pyracantha.

The basswood lace bug, Gargaphia tiliae (Walsh), often occurs in large num-
bers on the undersides of the leaves of basswood. Adults overwinter either under

leaves on the ground or in bark crevices. There are two generations per year.
Family Reduviidae
Assassin Bugs

Most species of assassin bugs are predacious on other insects, but some are
blood-sucking and frequently bite people. Many species are capable of inflicting
painful bites and will do so if carelessly handled. Most species are found on various
parts of plants, but a few are found on the ground or under objects on or near the
ground. Adults are varicolored, usually black, brownish, green, or orange. They
range greatly in size, some being more than 25 mm long. The head is long, narrow,
and cylindrical with the part behind the eyes necklike. The head bears a stout, rigid,
three-jointed beak that usually curves downward in the form of a semiloop, with the
tip resting in a groove in front of and between the front legs. The margins of the
abdomen are often exposed beyond the edges of the wings.

65

The wheel bug, Ari/us cristatus (L.), 1s an important predator of various forest
insects. As a young nymph it feeds on aphids, and later it attacks lepidopterous
larvae such as the fall webworm and other insects such as the locust borer. The adult
is a large, striking insect, with coglike teeth projecting from a median, longitudinal
ridge on the thorax. The female is much larger than the male and may reach a length
of 30 mm.

Other species attacking forest insects include: Sinea spinipes (Herrich-Schaffer),
which feeds on the fall webworm in the South; Acholla multispinosa (De Geer), an
enemy of the pine webworm; Zelus exsanguis Stal, an enemy of the gypsy moth;
and Melanolestes picipes (Herrich-Schaffer), an enemy of May beetles and their

larvae. The biologies of various species of this family have been reported (/0/3).
Family Nabidae
Damsel Bugs

Damsel bugs appear to be entirely predatory on soft-bodied plant-feeding insects.
Adults are usually pale brown to straw colored and about 8 mm long. The forelegs
are quite slender and fitted for grasping, having the tibiae armed with minute spines
and the femora enlarged. Nabis sordidus Reuter is a common species in eastern

forests. It often occurs in large numbers on rank undergrowth.
Family Anthocoridae
Flower Bugs

The majority, if not all, of the members of this family of small bugs are
predacious on other insects. The adults may be found on flowers, under loose bark,
in leaf litter, or in decaying fungi. Anthocoris musculus (Say) feeds on soft-bodied,
leaf-feeding insects, principally lace bugs, on deciduous trees in the Northern
States. Elatophilus inimica (Drake & Harris) feeds on the red pine scale in
Connecticut; it has also been found on pines infested with Matsucoccus gallicolus
Morrison in Massachusetts. The majority of flower bugs are black with white »
markings and are only about 3 to 5 mm long. Orius insidiosus (Say) 1s a common
predator of insect eggs.

Family Miridae
Plant Bugs

This is the largest family in the order Hemiptera, with about 1,600 species
occurring in the United States and Canada alone (675). The majority of species
appear to be phytophagous. Many others are predacious and feed on a wide variety
of young or soft-bodied insects. The adults are 2 to 9 mm long. The antennae and
beak are each four-segmented, with the second segment of the beak longer than the
head. The tarsi are usually three-segmented. The hemelytra, when fully developed,
are separated into a clavus, corium, cuneus, and membrane. In some species, the
hemelytra are abbreviated and the membrane is either absent or reduced to a narrow
band.

The oaks, ashes, hickories, and birches serve as hosts for many species. Trop-
idosteptes amoenus Reuter has caused noticeable injury to ash seedlings in nurser-
ies in the Lake States: the tarnished plant bug. Lygus lineolaris (Palisot de Beau-
vois), is often injurious to ornamentals and to forest nursery trees. It causes split-
stem lesions on the stems of young Populus. The stems sometimes break at the
lesion (J054). Young, succulent growth of elm is frequently damaged by the
feeding of Neolygus invitus (Say).

Seven plant bugs restricted to honeylocust were investigated in Pennsylvania, and
the life history of the honeylocust plant bug, Diaphnocoris chlorionis (Say), is
given along with biological observations on Lopidea incurva Knight, Lygocoris
tinctus (Knight), Pilophorus walshii Uhler, Plagiognathus delicatus (Uhler), and

66

Taedia gleditsiae (Knight) (/28/). D. chlorionis has damaged and defoliated
honeylocust in Michigan, New York, North Carolina, Ohio, Pennsylvania, Ver-
mont, and Wisconsin. It 1s widely distributed from Quebec and Ontario to South
Carolina, Mississippi, Texas, and in the North, west to Illinois, Indiana, and Iowa,
and also California.

Plagiognathus albatus (Van Duzee), the sycamore plant bug, is the cause of
unique injury to Platanus spp. over much of eastern North America. Small holes
appear at feeding sites, giving the appearance of shotholes caused by defoliators,
hail, or bacterial leaf spot (/280).

Family Coreidae
Coreid Bugs

This is a large family of relatively large bugs, some members of which have the
legs flattened and leaflike. They are similar to the lygaeids but differ in having
numerous veins in the membrane of the hemelytra.

The leaffooted pine seed bug, Lepfoglossus corculus (Say), occurs throughout
the Eastern United States and is a major pest in southern pine seed orchards (286,
360). Hosts include loblolly, shortleaf, slash, longleaf, Virginia, eastern white,
pitch, Table Mountain, and spruce pines. Other pine species, native or introduced,
grown in this region are also likely hosts. The large, conspicuous adults take flight
with a loud buzzing sound when disturbed. Both adults and nymphs are reddish-
brown to gray and have long legs. Adults are 15 to 18 mm in length, with a white
zigzag line across the wings and flattened, leaflike hind tibiae. The cylindrical eggs
are 2 mm long and about | mm in diameter. They are cream colored when first laid,
but turn dark reddish-brown as the embryo develops. Usually about 20 eggs are laid
end to end in a line along a single needle. There are several generations each year in
the South. Nymphs and adults have piercing-sucking mouth parts that they insert
into the conelets or cones to penetrate the developing ovules and seeds. Attacked
cones show no external damage symptoms. In early stages, nymphs feed upon the
needles and conelets. Second-stage nymphs destroy ovules in conelets, and exten-
sive ovule destruction causes conelet abortion (289). Third- , fourth- , and fifth-
stage nymphs and adults feed primarily upon seeds within cones. Some seeds
damaged in late summer and fall can be detected on radiographs (287). However, in
seed orchards, losses are also reflected by poor survival of conelet crops, high
numbers of empty seeds, poor seed viability, and low yields of sound seeds per
cone.

A related species, L. occidentalis Heidemann, a serious pest of Douglas-fir seed
in California (685), has been observed feeding on the needles and green cones of

Austrian pine in Missouri.
Family Rhopalidae
Rhopalid Bugs

The boxelder bug, Leptocoris trivittatus (Say), often becomes a pest wherever
boxelder is grown as a shade tree in the United States and Canada. In heavily
infested areas, it will feed on ash and maple. Adults (fig. 19) are somewhat
flattened, brownish black on top and about 12 mm long. There are also three red,
longitudinal stripes on the thorax, the margins of the basal half of the wings are red,
and the abdomen is bright red. Nymphs are wingless but possess wing pads and are
dark toward the head. They have bright-red abdomens.

The winter is spent in the adult stage in dry, sheltered places, such as the attics of
houses. During warm winter days, adults become active and come out of hiding,
only to retreat again when it turns cold. During the spring, they emerge and fly to

67

F-508517
Figure 19.—Adult cf the boxelder bug,
Leptocoris trivittatus, on seed of
boxelder.

their hosts where they deposit eggs on the leaves. Eggs also are occasionally laid on
leaves or under the bark of other tree species, on stones, grass, litter, fences, and in
doorway crevices. Eggs hatch in 11 to 14 days and the nymphs feed on leaves,
fruits, or soft seeds by inserting their beaks into the tissues. Feeding continues
throughout the summer, or until the nymphs become adults. In some parts of the
country the adults may emerge by midsummer and give rise to a second generation
that matures in the fall.

The importance of the boxelder bug as a pest derives from its habit of invading
houses in large numbers in search of shelter. Householders and supervisors of
outdoor recreational areas and parks are often concerned. One way to reduce the
problem is to remove the boxelder trees, particularly the female seed-bearing trees.
The removal of leaf litter also discourages the insects from congregating (/350).
Family Lygaeidae
Lygaeid Bugs

This is a rather large family, most members of which feed on mature seed. The
adults are small insects, about 2 to 18 mm long. Many are conspicuously marked
with spots or bands of white, black, or red. Various plant-feeding species are found
on herbaceous vegetation in the forest. One species, Kleidocerys resedae geminatus

(Say), feeds on the catkins of yellow and gray birches, rhododendron, etc.
Family Thaumastocoridae
Thaumastocorid Bugs

Xylastodoris luteolus Barber, the royalpalm bug, is the only member of this
family known to occur in the United States. It feeds on the Florida royalpalm,
Roystonea elata (Bartr.) F. Harper, in southern Florida, as well as in its native land,
Cuba. Adults are pale yellow, flattened, and about 2 to 2.5 mm long. Eggs are
deposited on the undersurfaces of leaflet midribs, usually on older trees. Feeding
occurs on unfolded leaflets of newly emerging fronds and to some extent on the
spike or projecting part of the terminal bud. Fronds exposed to heavy feeding may
turn brown (54).

68

Order Homoptera—Aphids, Spittlebugs, Scale
Insects, and Allies

The order Homoptera consists of divergent groups, some of which are among the
most common and abundant of all insects. Except for the cicadas, they are mostly
small and inconspicuous. A few are brilliantly colored, and many are grotesque in
shape. They are closely related to members of the order Hemiptera, but are
distinguished by their uniformly textured wings and by the point of origin of the
beak at the back of the underside of the head. The mouth parts consist of four
piercing stylets (the mandibles and maxillae). Many species are wingless, at least in
the female. When wings are present, there are usually four with the front pair
longer and the hind pair often wider. The wings do not overlap much at the top, and
their bases are never abruptly thicker than their tops. They usually slope roof-
shaped over the abdomen while the insect is at rest. Members of many families are
able to conceal themselves beneath various protective coverings such as froth, waxy
tufts, hard waxy shells, and the sloughing epidermis of bark. The majority are also
able to produce honeydew, a sweet sticky excretion.

Homopterous insects differ greatly in their biologies; some species produce
several generations per year while others may require several years to complete one
life cycle. The life history of some species is also very complex, involving both
bisexual and parthenogenetic generations, winged and wingless individuals and
generations, and sometimes the regular alternation of food plants. However, all
species are phytophagous, and they feed on an almost endless variety of plants of all
sizes and ages. They feed by inserting their beaks into plant tissues and extracting
the sap. A large number of species are injurious both to cultivated crops and to

forest, shade, and ornamental trees.
Family Flatidae
Flatid Planthoppers

A number of species of flatid planthoppers feed on trees and shrubs, but they are
seldom of economic importance (949, 1229). Members of different genera differ
greatly in body and form. Those occurring in this country are usually less than 12
mm long, and many have the head greatly modified, with the part in front of the
eyes greatly enlarged and snoutlike. A useful characteristic for recognizing them is
the position and form of the antennae. The two basal segments are stout, whereas
the remainder consists of a nearly pear-shaped basal segment and a segmented,
bristlelike terminal part. They also have a few large spines on the hind tibiae. Two
fairly common species are Anormenis septentrionalis (Spinola) and Metcalfa pru-
inosa (Say). The nymphs, which feed on the undersurfaces and midribs of leaves,
resemble small masses of cotton and are about 6 mm in diameter. They jump when

disturbed.
Family Cicadellidae
Leafhoppers

This is one of the largest families of insects in the world; all appear to be plant-
feeders. The adults range in length from about 3 to 15 mm and come in an almost
endless variety of colors. Certain species resemble flatid planthoppers, but they
differ in having one or more rows of small spines extending the length of the hind
tibiae. Because of the nature of their feeding, which consists of piercing plant
tissues with their mouth parts and sucking the juices, the damage they cause is
usually not recognized or is attributed to other factors. About the only visible
effects of leafhopper feeding are the white stippling of foliage or the browning,

69

withering, and curling of leaves. Because of the difficulty in observing or evaluat-
ing the damage to trees, very little is known of its magnitude or importance. The
probability is that it is grossly underestimated. These insects, in fact, may contrib-
ute significantly to the general unproductiveness of many stands of valuable
hardwoods in the Eastern United States. Many species are also vectors of destruc-
tive plant diseases, especially diseases of viral, mycoplasmal, or rickettsial origin.

Some species of leafhoppers spend the winter in the egg stage; others, as adults.
Nymphs usually hatch from overwintering eggs in May or June, and then feed on
new, tender leaves. Overwintering adults emerge during the first warm days in
spring and lay eggs as soon as the leaves of their hosts are fully developed. Nymphs
hatch in about 10 days. As a rule there are one or two generations per year;
sometimes, more.

Forest and shade trees serve as hosts for many species of Erythroneura, Em-
poasca, and Typhlocyba (295). Several species of /diocerus feed on willow and
poplar. Various species of Macropsis feed on poplar, willow, and honeylocust.
Certain species of Scaphoideus, Gyponana, and Ponana occur on American elm;
oaks are infested by species of Alebra, Eutittix, and Penthimia. This is only a
partial listing of genera and species known to attack trees; the total number is far
greater.

The most important leafhopper as far as forest and shade trees are concerned Is
the whitebanded elm leafhopper, Scaphoideus luteolus Van Duzee, the vector of
elm phloem necrosis, a mycoplasmal disease of American elm (43). Adults are
difficult to separate from those of closely related species, but not so the nymphs
(fig. 20). After the second instar, practically all of the nymphs are dark brown with
a transverse white band across the dorsum. This band lies just behind the thorax and
covers the first two and part of the third abdominal segments. The species is widely
distributed in the Eastern United States, from New York west to Kansas, Nebraska,
and Iowa, and south to Georgia and Alabama.

F-531900

Figure 20.—Adult and nymph of the
whitebanded elm leafhopper,
Scaphoideus /uteolus.

70

The whitebanded elm leafhopper lays its eggs in the cork parenchyma of elm
bark. When the nymphs hatch, they wander in search of leaves on which to feed.
The first of these to be found usually are on tiny branchlets growing from the trunk.
A dozen or more young nymphs may be found clustered on the undersurface of one
of these leaves, where they feed on the midribs or larger veins. Excessive feeding
often causes the apical portions of the leaves to turn brown and die. Older nymphs
are more widely distributed throughout the crown of the tree (44).

Elm phloem necrosis mycoplasm is transmitted by the adults that feed first on the
leaves of diseased trees and then on the leaves of healthy trees. Studies have shown
that a period of several days must elapse after the insect feeds on a diseased tree
before it can transmit the mycoplasm. Not all of the details of the life cycle of the
species have been determined, but it is known that the winter is spent in the egg
stage and that there is only one generation per year.

Homalodisca coagulata (Say), Oncometopia orbona (F.), Cuerna costalis (F.),
and Graphocephala versuta (Say) transmit the rickettsia causing phony peach, a
destructive disease of peach trees in the South.

Family Membracidae
Treehoppers

Treehoppers are characterized by the prolongation of the pronotum backward and
above the abdomen. In some species it not only extends backward but also sidewise
and upward, and in some, it extends to the top of the abdomen and completely
covers the wings. The hindlegs are long and adapted for jumping, and the female’s
ovipositor 1s long and sawlike. The majority of species apparently live on trees,
most often in open stands but also in woods. Only a few species are of economic
importance. The species of treehoppers found in Connecticut and Ohio have been
discussed (456, 950).

The buffalo treehopper, Stictocephala bisonia Kopp & Yonke, 1s occasionally
injurious to young ash and elm trees. Injury results from two opposing slits cut in
the bark by the female during oviposition. Eggs are placed in the slit, and the
portion of the stem beyond it often dies. Adults are light green to yellowish and
about 9 mm long. The pronotum is sharply elevated, is widest at the tip, bears two
sharply pointed horns that extend at right angles to the body, and ends beyond the
tip of the abdomen in a narrowed acute process.

The threecornered alfalfa hopper, Spissistilus festinus (Say), occasionally
damages black locust seedlings in nurseries in the South. Injury results from
nymphs feeding on the stems, usually 25 to 50 mm above the base. Gall-like
swellings or calluses that develop just above the feeding punctures kill many of the
seedlings. Other seedlings are lost when they break at these points. Stictocephala
militaris (Gibson & Wells) nymphs feed on the veins and petioles of sweetgum
leaves in north Georgia. Gall-like enlargements may develop at points of injury on
the petioles, or the leaves may die. Other common membracids and their tree hosts
are as follows: Micrutalis calva (Say)—abundant on honeylocust; Telamona re-
clivata Fitch—common to abundant on basswood; 7. decorata Ball—fairly com-
mon on quaking aspen; Vanduzeea arquata (Say) and Thelia bimaculata (F.)—
abundant on black locust; Cyrtolobus discoidalis (Emmons)—common on red oak;
Carynota stupida (Walker)—sometimes abundant on yellow birch; Platycotis
vittata (F.)—abundant on oak; and the twomarked treehopper, Enchenopa
binotata (Say)—common on butternut but also attacks walnut.

71

Family Cercopidae
Spittlebugs

Spittlebugs are represented in our fauna by 25 species (//48). As a group, the
adults are stout-bodied, rarely over 12 mm long, oval or oval-elongate, and dull
colored. The nymphal or immature stages are spent in frothy masses of spittle on
their host plants. The genus Aphrophora is undergoing revision (5/8).

The pine spittlebug, Aphrophora parallela (Say), occurs in southern Canada
and throughout most of the Eastern States from New England to the Lake States,
Arkansas, Florida, and Alabama. Its favorite host appears to be Scotch pine, but it
is also known to attack pitch, eastern white, Virginia, jack, slash, loblolly, and
Japanese pines; Norway, white, and red spruces; and balsam fir, larch, and eastern
hemlock. Trees of all ages and sizes are attacked. During outbreaks, infested Scotch
pines may be severely stunted or killed (//38). Adults are tan to dark reddish-brown
with two narrow, oblique, light bands, usually bordered by darker bands, on each
wing cover, and are about 8 to 11 mm long (fig. 21).

Eggs are deposited in dead woody tissue or just under the bark of twigs during
July and August. In the northern parts of its range, the species spends the winter in
the egg stage. The nymphs usually hatch in May, and then feed on the twigs, where
they soon cover themselves with spittle. As they grow, they usually change lo-
cations and form new masses of spittle at each stop. Upon the approach of
maturity they often move the the main trunk where several may occupy a single
spittle mass. When full grown, they migrate to the needles and transform to
adults. Adults are present during July and August and feed on the same hosts
without producing spittle masses. There is one generation per year.

The fungus, Entomophora aphrophorae Rostrup, and high temperatures during
the nymphal period occasionally cause heavy mortality of the pine spittlebug. The
pine spittlebug is often associated with Diplodia pini, a fungus that apparently
invades the tree when weakened and through the insect’s feeding punctures. Most

| Courtesy Conn. Agric. Exp. Stn.

Figure 21.—A, adult; B, nymphal spittle masses of the
pine spittlebug, Aphrophora parallela.

V2

flagging injury attributed to the spittlebug, in the Lake States at least, is due to the
fungus rather than the insect. Controlling the insect halts the disease spread. The
problem is worst in weakened large sapling and pole-size Scotch pine stands.
Thinning closed stands to make them more vigorous is also recommended as a
control practice.

The Saratoga spittlebug, Aphrophora saratogensis (Fitch), occurs in south-
eastern Canada and the East Central United States from Maine to Minnesota.
Nymphs feed on a wide variety of herbs and shrubs. Sweetfern is preferred and
necessary for most outbreaks. Outbreaks can occur, however, when 80 percent of
the forest floor is covered by other broadleaf plants. The adults feed on red, jack,
eastern white, Virginia, pitch, and loblolly pines, and occasionally on white
spruce, balsam fir, and tamarack. In the Lake States, red pine is hit hardest and jack
pine ranks second. Heavy infestations in the Lake States have destroyed entire red
pine plantations.

The adult is about 9 to 10 mm long, light brown to tan, with a light, irregular
arrow-shaped stripe on the head and pronotum and with oblique, wavy markings on
the wing covers. The abdomen of the young nymph is scarlet, edged with black.
Fifth instars are dark brown.

Adults are active from late June to late September. They feed by inserting their
mouth parts into the cortex of new and old shoots of the host, mostly on 2-year-old
internodes. They extract sap and inject toxic saliva which causes the formation of
necrotic resin-filled pockets in the pholem and xylem tissues. Damaged pines are
characterized by the presence of reddish-brown “‘flags” of dead foliage, by numer-
ous puncture wounds on the twigs covered with small drops of resin, and by light-
tan flecks in the wood and inner bark at feeding points. Extensive feeding kills
branches, stunts and deforms growing shoots, and may kill entire trees. Mortality
usually begins 2 or 3 years after the first flags are seen (389).

On red pine, eggs are laid under the outer scales of buds on the upper part of the
tree. On jack pine, they are laid in the sheaths of current year’s needles or under the
bark of twigs. The winter is spent in the egg stage and the nymphs hatch in the
spring just about the time red and jack pine buds begin to elongate. Young nymphs
drop to the ground and begin feeding at the base of alternate host stems, either
singly or in small groups. As they feed, they form masses of spittle (fig. 22). To
find these masses it is frequently necessary to brush the litter away from the base of

/\

F-489365

Figure 22.—Spittle mass of the Saratoga spittlebug,
Aphrophora saratogensis, at base of sweetfern.

73

the stem. The nymphs reach maturity and transform to adults in about 40 to 70
days, depending on the weather. The adults then fly to the pine hosts and begin
feeding on needle-bearing twigs. In the Lake States, adult transformation is about
80 percent complete by early to mid-July.

Nymphal populations are sometimes reduced by late spring frosts in the Lake
States. Hot, dry weather also kills many young nymphs, especially in open planta-
tions. Insect parasites and predators provide a certain amount of control but are
unable to prevent outbreaks. Recommended preventive control measures include
selection of sites for red or jack pine plantations that are comparatively free of hosts
favored by the nymphs, dense planting to secure early crown closure, and planting
on good sites. Methods of direct control of adults and nymphs are available (7334,
135).

Prosapia bicincta (Say) occurs from Massachusetts to Florida and west to Kansas
and Texas. Adults are dark brown and about 9 mm long. There usually are two
distinct bands across the wings and a narrower orange band on the thorax between
the humeral angles. Adults have been recorded feeding on holly, redbud, cherry,
and a wide variety of other woody and herbaceous plants. Infested leaves of holly
become distorted, stunted, and discolored and may have necrotic areas at feeding
sites.

Clastoptera undulata Uhler nymphs and adults have been observed feeding on
the young twigs of horsetail casuarina in Florida. In heavily infested areas, the
adults may be attracted to lights in such large numbers that they become a nuisance.
The alder spittlebug, C. obtusa (Say), a common species, feeds on various shrubs
and trees, including hickory, birch, and alder. The pecan spittlebug, C. achatina
Germar, is occasionally a serious pest of pecan in the Midwest. It feeds on the
terminals, sometimes killing fruit-producing shoots. The dogwood spittlebug, C.
proteus Fitch, has been recorded on pine. C. salicis (De Geer) frequently occurs in
large numbers on willow. The meadow spittlebug, Philaenus spumarius (L.),
occasionally feeds on Scotch pine in the Northeast.

Family Cicadidae
Cicadas

Cicadas are the largest members of the order Homoptera in the United States.
The adults are stocky, heavy-bodied insects with large compound eyes and mem-
branous wings, and some reach a length of 50 mm. There are two common types,
(1) the dog-day cicadas, often called harvest flies, and (2) the periodical cicadas,
also known as 17-year locusts. The dog-day group contains large blackish species,
usually with greenish markings. The life cycle lasts from 2 to 5 years but, because
of overlapping broods, some adults appear every year. Periodical cicadas differ
from the dog-day group in being smaller and in having reddish eyes, reddish legs,
and reddish wing veins. The life cycle is 13 years in the South and 17 years in the
North.

Cicadas deposit their eggs in the twigs of trees and shrubs and often damage
twigs so severely that their terminal portions die. When the eggs hatch, the young
nymphs drop to the ground, enter the soil, and feed on roots. Here the nymph
remains until ready to molt for the last time, years later. Before molting, it emerges
from the ground and climbs upon some object, usually the trunk of a tree, fastens its
claws in the bark, and molts. The adults of some species live 5 to 6 weeks.

The periodical cicada, Magicicada septendecim (L.) (fig. 23A), is widely
distributed in the Eastern United States, and it lays its eggs in more than 70 species
of trees and other plants. The most susceptible of the trees appear to be the oaks,

74

hickory, honeylocust, dogwood, apple, and peach; however, many others such as
sweetgum, elm, ash, yellow-poplar, walnut, sycamore, and redbud may also be
heavily attacked. Adults are about 40 mm long. The female is completely black on
top, but the male has 4 or 5 orange-brown abdominal segments on top.

“SS

F-532845, F-532846
Figure 23.—The periodical cicada, Magicicada
septendecim: A, adult; B, oviposition scars in white
oak twig.

The female uses a sawlike ovipositor to puncture the bark and make a pocket in
the wood in which she deposits from 24 to 28 eggs in two rows. She may then
proceed along the twig and repeat the process until she has deposited about 5 to 20
batches of eggs. Sometimes the punctures are placed so close together that the
wounds appear as a single slit up to 75 mm long (fig. 23B). When the eggs hatch,
the nymphs fall and enter the ground where they feed on suitable roots. Usually
they are found at depths of about 0.5 m beneath the surface. When they become full
grown, they emerge to begin a new cycle, usually leaving the ground during the
night. Emergence may begin as early as the last week of April in the South and as
late as the last week in May in the North.

A few days after the adults appear, the males begin drumming or singing. During
outbreaks the sound is loud and incessant, literally deafening in wooded areas. The
chorus begins at dawn and the volume increases as the temperature rises. It ceases
at evening.

Cicada adults cause no visible feeding damage. In contrast, the egg-laying habits
of the female may cause serious damage, especially to young, transplanted trees in
nurseries and orchards. Some damage also results to older trees. Damaged twigs
wilt, and some break at damaged points. Methods of control are discussed (/220).

Magicicada cassini (Fisher) also occurs in the Eastern United States. In Kansas,
it is found mainly in streamside habitats in lowland forests. Tibicen canicularis
(Harris) has been reported causing serious damage in white spruce plantations on
poor sites in Quebec.

75

Family Psyllidae
Jumping Plantlice

Jumping plantlice or psyllids are very small, about 2 to 5 mm long, and look very
much like minature cicadas. There is also some resemblance to winged aphids but
psyllids differ in having stouter legs, with the hind pair adapted to jumping. The
nymphs of certain species which secrete large quantities of wax resemble woolly
aphids. The adults are very active and jump or fly when disturbed.

Psyllids fall into two species groups: (1) the leaf gall makers, and (2) the woody
gall makers. Adults of the leaf-feeding forms emerge from galls in the fall: the
wood-feeding forms emerge in spring. Leaf gall makers are further subdivided into
two groups, blister gall makers and nipple gall makers.

Pachypsylla celtidisvesicula Riley, the blistergall psyllid, forms a small, mono-
thalamous (one psyllid per gall) blister gall. It is believed to occur throughout the
range of hackberry in the United States and is the most abundant of the hackberry
psyllids. Fifty or more may infest a single leaf. Adults often become a household
nuisance when thousands accumulate on screens in the autumn before hibernating.
Two undescribed species of blistergall psyllids occur on sugarberry and netleaf
hackberry.

Pachypsylla celtidisastericus Riley, the hackberry stargall, produces a gall that
closely resembles the blistergall formed by the blistergall psyllid, except that it has
starlike growth on the lower side of the leaf. It can occur on all hackberry tree
species except Celtis occidentalis L. |

The hackberry nipplegall maker, P. celtidismamma (Riley), is perhaps the best
known hackberry psyllid. It is found only on C. occidentalis. There are other
nipplegall makers on the other four species of hackberry trees, but they are not this
species. It is monothalamous, although at times it may appear to be polythalamous
because as many as seven galls of another psyllid, P. celtidisvesicula, may be
incorporated into a single nipplegall of P. ce/tidismamma (890). Like the blistergall
psyllid, adults cluster on screens during the fall. The nipplegall psyllid may
disfigure leaves when infestations are heavy (///2).

Pachypsylla celtidisgemma Riley, the budgall psyllid, occurs throughout the
range of hackberry trees in the United States, and may become a pest when
populations are high. Adults appear during the latter part of June in the New York
City area and lay their eggs on the young leaves. Young nymphs enter the buds and
initiate gall formation. The polythalamous galls contain several nymphs, each in its
own chamber. The winter is passed as a fifth nymphal instar in the gall (/243).

Pachypsylla venusta (Osten Sacken), the petiolegall psyllid, is the largest of the
hackberry psyllids, and it forms the largest galls. The galls are polythalamous and
usually contain 6 cells, although there may be as many as 30. These galls may
persist on the trees for several years and seriously damage their appearance.
Distribution of the species is mostly limited to below the 40th parallel.

Pachypsylla celtidisinteneris Mally forms small, inconspicuous, monothalamous
galls under the bark of hackberry twigs.

Several other species in the genus Psy/la feed on various trees, shrubs, and
ornamentals, and produce large quantities of honeydew that drips and covers their
hosts. A black mold growing in this material often makes infested plants look less
healthy than they are.

The boxwood psylud, Psyl/a buxi (L.), attacks American and English varieties
of boxwood and probably occurs wherever they grow. The adult is greenish and has
transparent wings; nymphs are gray-green and covered with a white cottony or

76

waxy material. The first winter is spent in the nymphal stage. Feeding is resumed in
the spring and adults begin to emerge in early May. Infested leaves curl and form
cups in which nymphs are concealed. Feeding also results in reduced growth of
young twigs. Eggs are laid under bud scales during July and August. Control
recommendations are available (/269). Other species of Psylla and their hosts
include: P. annulata Fitch—paper birch and maple; P. carpinicola Crawtord—
birch; P. floccosa Patch and P. galeaformis Patch—alder; and P. trimaculata
Crawtord—cherry.

Trioza tripunctata (Fitch) has been observed feeding on the needles of pines in
ornamental plantings in Connecticut. Damage consists of yellowish or reddish spots
that frequently coalesce, causing the needles to die and drop prematurely. The
persimmon psylla, 7. diospyri (Ashmead), and 7. magnoliae (Ashmead) feed on
persimmon and magnolia, respectively, in Florida.

More than 150 species of hymenopterous and dipterous parasites attack psyllids
in the United States (639).

Family Aphididae
Aphids, Plantlice

The aphids or plantlice constitute a very large group of small, soft-bodied insects
that feed by sucking the sap from the leaves, stems, and roots of plants and trees.
They are pear-shaped or globular, have fairly long antennae, and usually a pair of
tubelike cornicles rise from the top of the fifth or sixth abdominal segment. Winged
males are common. The wings are very delicate and membranous, with only a few
simple veins, and they are usually held vertically above the body while at rest. Egg-
laying females of many species are also winged.

Ordinarily, most of the aphids feed while exposed on their hosts. Others feed in
Sheltered locations such as inside leaves, which they cause to curl or to become
distorted, or inside galls. Trees of all kinds, sizes, and ages are attacked, but they
normally are not seriously injured. Serious damage sometimes results, however,
~ especially to shade and ornamental trees and young trees in plantations. Honeydew
dropping from heavily infested shade and ornamental trees is often a nuisance
because it forms a sticky coating on everything below, especially sidewalks, parked
automobiles, and park benches. Additional damage may result from the growth of a
sooty mold in the honeydew. It not only detracts from the esthetic value of shade
trees and ornamentals, but also interferes with the food-manufacturing process of
their leaves. However, honeydew is also useful as food for many species of
beneficial insects. Numerous publications have been issued on the identity, dis-
tribution, and hosts of aphids (603, 955, 962, 963, 964, 965, 978). Nomenclature
cited in this section 1s based upon that set forth in “Survey of the World’s Aphids”
(B53):

The giant bark aphid, Longistigma caryae (Harris), one of the largest of all
North American aphids, occurs throughout much of the Eastern United States and
feeds on a variety of deciduous trees such as elm, pecan, sycamore, oak, maple,
basswood, birch, beech, walnut, and willow. Adults are about 6 mm long, and are
covered with a bluish-white bloom. Males are winged; egg-laying females are
wingless. Eggs are laid on twigs, a single female laying many. The aphids usually
occur in clusters on the undersides of twigs and small limbs. Depending on locality,
there may be several generations per year. Heavily infested twigs may be seriously
injured or killed.

The genus Cinara contains a number of conifer-infesting species that feed on the
bark; only a few of these are mentioned here.

1)

The white pine aphid, C. strobi (Fitch), occurs from New England to the Lake
States and Carolinas and feeds on eastern white pine. Winged forms are almost 4
mm long. The body 1s shiny dark brown, with a white stripe down the middle of the
dorsum and white powdery spots on the sides, and it bears long stiff hairs.

During the fall, winged aphids lay their eggs end to end along the needle. As
many as 27 may be found on a single needle, although 5 or 6 are generally more
common. Hatching occurs in the spring, and wingless females produce living
young which live in colonies up to 75 or 100 mm long clustered around a branch or
the leader. Several generations later, winged females are produced and they migrate
and also produce living young. Toward the fall, winged males and females mate,
and a new crop of overwintering eggs 1s laid. Young trees or individual branches of
large trees may be killed by heavy infestations or their growth may be seriously
reduced.

Cinara sabinae (Gillette & Palmer), a small yellowish species about 3 mm long
and covered with a white powdery secretion, feeds on redcedar in the Eastern
United States. Colonies are usually found on twigs and small branches. Heavily
infested trees often become unsightly as a result of black mold developing in
honeydew on the foliage. The related species, C. canadensis Hottes & Bradley and
C. juniperivora (Wilson), also feed on redcedar.

Cinara pinea (Mordwilko) is a large reddish-brown species with numerous dark
specks and a pair of large spots behind the cornicles. It has been recorded feeding
on young Scotch, red, and Virginia pines. Additional pine-infesting species of
Cinara include C. atlantica (Wilson) on loblolly, shortleaf, Virginia, slash, long-
leaf, pond, sand, and spruce pines in the South, and Scotch, pitch and Table
Mountain pines in the North; C. taedae Tissot on loblolly, Virginia, slash, sand,
spruce, pond, Table Mountain, and pitch pines; C. watsoni Tissot on loblolly,
shortleaf, slash, spruce, pond, red, Scotch, Virginia, sand, jack, pitch, and Table
Mountain pines; C. pinivora (Wilson) on loblolly, slash, Virginia, shortleaf, sand,
spruce, pond, pitch, red, jack, and Table Mountain pines; and C. pergandei
(Wilson) on loblolly, Virginia, shortleaf, sand, spruce, pitch, and jack pines. C.
pergandei is perhaps the most common bark aphid on Virginia pine. Pine-infesting
aphids that attack the needles include Eulachnus rileyi (Williams), the powdery
pine needle aphid, on Scotch, red, pitch, Virginia, shortleaf, loblolly, slash, and
eastern white pines; FE. agilis (Kaltenbach) on red, Scotch, Austrian, and eastern
white pines; Essigella pini (Wilson), the speckled pine needle aphid, on loblolly,
slash, and pond pines in Florida (also recorded from Maryland and Pennsylvania);
Schizolachnus piniradiatae (Davidson), the woolly pine needle aphid, on jack and
red pines in the Eastern States.

Several other species of aphids have also been recorded on other conifers. For
example, Cinara confinis (Koch) has been recorded on fir; C. pilicornis (Hartig) on
Norway spruce and various firs; C. tujafilina (Del Guercio) on northern white-
cedar, baldcypress, and other species (heavy infestations have been recorded in
nurseries in Florida and Delaware); Prociphilus bumelia (Schrank) on balsam fir, as
well as roots of eastern white pine; and the balsam twig aphid, Mindarus abietinus
(Koch), on a wide variety of hosts including balsam fir, Fraser fir, Siberian fir,
subalpine fir, white spruce, and juniper. On fir, damaged needles curl and the bark
of heavily infested twigs becomes roughened (9/9).

The boxelder aphid, Periphyllus negundinis (Thomas), feeds on the leaves and
twigs of boxelder wherever it grows in the United States and Canada. The aphid’s
body is usually yellowish green with brownish marks on the thorax and abdomen.

78

Walks, benches, or cars parked beneath infested trees are often badly soiled. The
related species, the Norway maple aphid, P. /yropictus (Kessler), often completly
defoliates Norway and sugar maple trees. Honeydew dripping from infested leaves
is a nuisance. P. americanus (Baker); Drepanaphis carolinensis Smith; D.
nigricans Smith; D. sabrinae Miller; the painted maple aphid, D. acerifoliae
(Thomas); and Drepanosiphum platanoidis (Schrank), the sycamore maple aphid,
are also found on maples.

Numerous species of Myzocallis such as M. bellus (Walsh), the greater striped
red oak aphid; M. punctatus (Monell); M. discolor (Monell); and M. melanocera
Boudreaux & Tissot feed on various oaks. Other common members of related
genera and their hosts include: Prerocallis alnifoliae (Fitch)—alder; Eucallipterus
tiliae (L.), the basswood aphid—native and introduced lindens; the crapemyrtle
aphid, 7inocallis kahawaluokalani (Kirkaldy)—crapemyrtle; the black pecan ap-
hid, Melanocallis fumipennellus (Fitch)—hickory and pecan; and the elm leaf
aphid, Jinocallis ulmifolii (Monell)—elm.

Many other species of free-living aphids also occur commonly on various
deciduous trees. These include the black-margined aphid, Monellia costalis
(Fitch), M. caryella (Fitch), M. microsetosa Richards, and M. nigropunctata
Granovsky on hickories; Chaitophorus stevensis Sanborn, C. populicola Thomas,
and Tuberolachnus salignus (Gmelin) on poplars; Pterocomma populifoliae (Fitch)
and P. smithiae (Monell), the black willow aphid, on poplars and willows; P.
salicis (L.), on willow; the tuliptree aphid, ///inoia liriodendri (Monell), on
yellow-poplar and magnolia; the cowpea aphid, Aphis craccivora Koch, on black
locust seedlings; Phyllaphis fagi (L.), on nearly all species of beech; Calaphis
betulaecolens (Fitch), the common birch aphid, on birch (often in large numbers);
C. betulella Walsh, on birch and beech; Euceraphis punctipennis (Zetterstedt), the
European birch aphid, on various birches, especially yellow birch; FE. lineata
Baker, on gray birch; and E. mucida (Fitch), on river birch.

Three species of woolly aphids are commonly found on elms in the Eastern
United States. The woolly apple aphid, Eriosoma lanigerum (Hausmann), feeds on
new terminal leaves, causing them to curl or appear in the form of rosettes. It
attacks apple, pear, hawthorn, and mountain-ash. Damage to elms is not especially
severe. The presence of large numbers of distorted leaves on shade trees is
unsightly. The woolly elm aphid, EF. americanum (Riley), has elm as a primary
host and serviceberry as an alternate host. It feeds at the edges of young elm leaves
in the spring, causing them to swell, curl, and roll inward from the edge. Two
generations are produced on elm. Members of the second generation then fly to
serviceberry and attack its roots. The summer is spent here. During the fall, a
winged generation develops and flies back to the elm, where it lays a cluster of
overwintering eggs. E. rileyi Thomas, the woolly elm bark aphid, attacks Ameri-
can and slippery elms. It occurs in dense woolly clusters on the limbs and trunks,
causing knotty growths to form at the sites of injury. Heavily infested trees may be
seriously injured. Hawthorn is infested by the closely related species, E. crataegi
(Oestlund). It occurs in dense colonies in twigs and branches and is occasionally
injurious to ornamentals.

‘Several other woolly aphids are also found on various species of hardwoods. The
beech blight aphid, Fagiphagus imbricator (Fitch), feeds primarily on the bark of
twigs and small branches, but also can be found on the trunk and undersides of
leaves of beech trees from New England to Georgia and Illinois. Its body is covered
with a white, cottony substance that strings out in fairly long threads and forms a

79

tuft at the rear end. Heavy infestations occasionally develop on forest-grown trees.
The woolly alder aphid, Prociphilus tessellatus (Fitch), occurs throughout the
Eastern United States and is often abundant on alder and silver maple. P. fraxinifolii
(Riley) feeds on ash and P. corrugatans (Sirrine) on serviceberry. P. longianus
Smith is associated with root galls caused by cancer-root on black oak.

Certain species of aphids produce galls by their feeding. The elm cockscombgall
aphid, Colopha ulmicola (Fitch), is probably the most important tree-infesting
species. It occurs throughout most of the United States and Canada wherever its
hosts, American, rock, and slippery elms, grow. In early summer, it feeds on
leaves, causing the formation of galls up to 25 mm long and 6 mm in height. The
gall is irregular in shape and resembles a rooster’s comb. The winter 1s spent in the
egg stage on elm, and by midsummer the aphids have left for a secondary host
thought to be a grass. There are six generations per year. Damage is not severe, but
when large numbers of galls occur on the leaves of young trees and ornamentals, the
trees may become unattractive. Another species, 7etraneura ulmi (L.), produces
pedunculated, bladderlike galls up to 25 mm long on the upper surfaces of leaves in
New England.

The poplar vagabond aphid, Mordwilkoja vagabunda (Walsh), feeds at the tips
of twigs of cottonwoods and occurs from New England to Utah, causing the
formation of convoluted galls (fig. 24) up to 13 cm in diameter. These galls may
occur singly or in clusters of three to five each. The winter 1s usually spent in the
egg stage in old galls or occasionally in nearby bark crevices (623). Other gall-
producing aphids include Hormaphis hamamelidis (Fitch), which forms conical
galls on the upper surfaces of witch-hazel leaves: Hamaelistes spinosus Shimer, the
witch-hazel gall aphid, which causes galls to form on the stem buds of witch-
hazel: Kaltenbachiella ulmifusa (Walsh & Riley), which forms spindle-shaped,
saclike galls on the upper surfaces of leaves of slippery elm; and the poplar petiole
gall aphid, Pemphigus populitransversus Riley, which occurs on various species of
poplar where it overwinters and produces oval galls on the leaf petiole.

F-506747

Figure 24.—Galls of the poplar vagabond aphid,
Mordwilkoja vagabunda, on poplar.

80

Family Adelgidae
Adelgids

Members of this family differ morphologically from true aphids 1n having shorter
antennal segments and lacking cornicles. Unlike the Aphididae, both sexually
perfect and imperfect female adelgids lay eggs. All species occur only on conifers.
Feeding sites of important eastern adelgids include needles, twigs, limbs, trunks, or
the inside of galls.

The eastern spruce gall adelgid, Ade/lges abietis (L.), an introduced species
from Europe, occurs in southeastern Canada and in the Northeastern and Lake
States. Its preferred host appears to be Norway spruce, but it is also found
occasionally on white, red, and blue spruces. [t overwinters as small nymphs under
coverings of waxy threads at the bases of buds on the undersides of twigs (/290).
The nymphs molt in the spring and become stem-mothers that lay eggs on the
needles about the time the buds are ready to break. Hatching occurs in about 2
weeks and the young nymphs crawl to the bases of new needles. Here they feed,
causing the formation of pineapple-shaped galls (958) in which they live and
continue their development (fig. 25). During late August to October, the galls open
and the nearly mature nymphs craw] out onto the needles. The nymphs transform to
winged adults in a couple of days but, because they are weak fliers, many remain on
the tree. The females insert their mouth parts through the bark, deposit 100 or more
eggs each, and then die. These eggs hatch within 16 days and the young nymphs
immediately crawl to overwintering sites. There is one generation per year. This
insect is a serious pest in nurseries and Christmas tree plantations and on park and

Courtesy D. C. Allen,
SUNY, Coll. Environ. Sci. & For.

Figure 25.—Galls of the eastern spruce gall adelgia,
Adelges abietis, on spruce.

81

other ornamental trees. It damages trees by reducing their vitality, detracting from
their esthetic value, and weakening shoots at points of gall formation.

The Cooley spruce gall adelgid, Ade/ges cooleyi (Gillette), is a native pest that
occurs from coast to coast in the Northern United States and throughout the range
of white spruce in Canada. Its primary hosts in North America are white, blue,
Sitka, and Engelmann spruces. It also has an alternate host, Douglas-fir.

The Cooley spruce gall adelgid overwinters as an immature female under bark
scales near the terminal of twigs of spruce. In early spring the female develops into
a stem-mother and deposits up to 350 eggs under a mass of white, cottony wax. The
eggs hatch in | to 2 weeks. and the nymphs settle down to feed at the bases of young
needles. Elongate, conelike galls (fig. 26) begin to form immediately and develop
rapidly, enclosing the nymphs. Young galls are fleshy and green or purple; older
ones are dry and reddish brown. They vary greatly in size, from about 25 to 75 mm
in length and 12 to 18 mm in diameter. When the nymphs become mature, the galls
open, allowing the nymphs to escape and crawl to the needles. Here they transform
into winged adults and fly to Douglas-fir, if present. Eventually, a winged genera-
tion is produced on this host, and it returns to spruce (27/). Where spruce and
Douglas-fir do not occur close enough together for the aphid to move back and
forth from one to the other, continous generations may be produced independently
on either species.

F-519584

Figure 26.—Galls of Cooley spruce gall adelgid,
Adelges cooleyi, on blue spruce.

82

The Cooley spruce gall adelgid is not usually considered an important pest in the
forest. However, it may be troublesome where spruce and Douglas-fir are growing
close together. Ornamental spruce and young spruce trees in Christmas tree planta-
tions are often seriously damaged by excessive numbers of unsightly galls. No galls
are produced on Douglas-fir but heavy adelgid attacks on this host can cause
abnormal dropping of foliage.

The balsam woolly adelgid, Ade/ges piceae (Ratzeburg), an introduced species
first recorded in North America from Brunswick, Me., in 1908, now occurs in the
Maritime Provinces, Canada, and the Northeastern States. Infestations also occur
over much of Newfoundland, in the Gaspé Peninsula of Quebec, in the southern
Appalachians, in the Pacific Northwest, and in British Columbia. Its hosts are
balsam and Fraser firs in eastern North America. Full-grown adelgids are roughly
spherical, about | mm long, and almost invisible to the naked eye. Because of a
covering of white wax threads, however, they appear conspicuously as dots of white
“wool” (fig. 27).

F-519571
Figure 27.—Infestation of the balsam woolly adelgid,
Adelges piceae, on trunk of Fraser fir.

Clusters of amber eggs are deposited in the late spring and early summer; each
egg is attached to the bark behind the female’s body by a silken thread. Newly
hatched crawlers form the only motile stage in the life cycle of this insect. When
suitable feeding sites are found on the surface of the bark, these crawlers insert their
stylets, become stationary, and turn black except for fringes of white wax plates
around the edges of the body and down the dorsum. After a period of dormancy
lasting from 2 to 8 weeks they develop into the second generation. Adults of this
generation deposit eggs during midsummer. Hatching soon occurs and all stages are
found until late fal!. The winter is spent as stationary first-stage larvae. In the spring
these larvae resume activity and reach maturity by the time the buds begin to swell.
New adults appear by mid-April in the southern Appalachians and in early May in

83

the Northeast. There are two generations per year in the Northeast: in the southern
Appalachians, two and occasionally three generations are produced (/3, 47).

The balsam woolly adelgid feeds at any point on the tree where it can reach the
parenchyma of the cortex with its mouth parts. During feeding, it introduces an
irritating salivary substance into the tissues that causes an abnormal multiplication
of cells and excessive growth in the vicinity of the point of attack. Infested twigs
and small branches become swollen and distorted. Swellings are particularly
noticeable at nodes and around buds. Branchiets may thicken, twist irregularly, and
bend down at the ends. The main stem tapers rapidly toward the top. The tip
becomes bent or flattened and is usually killed. This results in a condition com-
monly known as ““gout.”” The wood of infested trees becomes hard and brittle, and
its surface is usually marked with dark, reddish-brown blotches. This type of wood,
which resembles “compression” wood, is produced in greatest amount on moder-
ately infested fast-growing trees. Trees suffering from heavy stem attack may be
killed in 2 or 3 years (fig. 28).

The balsam woolly adelgid is subject to a considerable amount of control by low
winter temperatures, especially in the northern portions of its range. Tree resistance
is also an important control factor, some trees being less favorable for multiplication
of the insect than others. So far, no insect parasites of the adelgid have been found.
Quite a large number of predators have been recorded and additional ones have been
imported to supplement native species. However, adelgid populations expand so
rapidly and some firs are so sensitive to attack that predators effect little control
before trees are damaged irreversibly.

In a few situations, the spread of infestations may be checked by the prompt
salvage of infested stands during the winter. Short rotation and cutting cycles,
combined with silvicultural methods of reducing the balsam fir content of stands,
have helped in reducing infestations in New Brunswick (50).

Adelges laricis Vallot, an introduced species, occurs in southern Canada and
south and east in the Eastern States to Washington, D.C., and the Lake States. Its
hosts are recorded as European larch, tamarack, and spruces, principally red and
black. Infestations on larch appear as white woolly masses on the needles and as
clusters of adelgids at the bases of needles. Infestations on spruce may be recog-
nized by the presence of small pineapple-shaped galls at the tips of new growth. A
related species, A. lariciatus (Patch), occurs on larch and various species of spruce
(272). It produces galls similar to those produced by the eastern spruce gall adelgid.

The pine bark adelgid, Pineus strobi (Hartig), a widely distributed species in
Europe and North America, occurs over most of the United States wherever white
pines grow. It is small, dark, and covered with flocculent wax. Infestations may be
recognized by the presence of spots and patches of white cottony material on the
smooth bark of the trunks and limbs (fig. 29), at the bases of needles on twigs, or on
buds. Feeding, though, is limited to the bark. The trunks of heavily infested trees
often appear as if whitewashed.

Eggs are laid in the spring by overwintering females, and the eggs hatch into both
winged and wingless females. Winged forms, though, are produced less consis-
tently than wingless ones. The wingless forms remain on the pine host and
reproduce repeatedly. Five generations per year have been recorded as far north as
the Lake States (/0/0). Although winged forms may fly to spruce and lay fertile
eggs, both adults and offspring eventually die.

Trees in parks and recreational areas, ornamentals, and small nursery stock
sometimes become heavily infested by the pine bark adelgid. However, it is

84

doubtful that such attacks produce permanent damage if the trees are healthy.
Similarly, mature trees repeatedly infested in the forest or in eastern white pine
plantations evidently suffer no serious harm.

Sat
F-519567
Figure 28.—Fraser fir killed by the

balsam woolly adelgid, Ade/lges
piceae.

Courtesy Conn. Agric. Exp. Stn.

Figure 29.—Infestation of the pine bark
adelgid, Pineus strobi, on the trunk of
pine.

85

The pine leaf adelgid, P. pinifoliae (Fitch), occurs in both the Eastern and
Western United States. Its range in the East coincides with that of its primary hosts.
red and black spruces. wherever they grow close to its alternate host, white pine.
The offspring produced by aduits migrating from spruce provide the easiest means
of diagnosing an outbreak on pine. The small, scalelike “larvae” are fringed with
white hair and often become numerous enough to cover a shoot completely.

The life cycle takes 2 years to complete. During part of this time, including the
first winter, infestations are found entirely on spruce; during the remaining time,
including the second winter, they are found on pine (752).

Infestations on spruce result in the production of terminal compact galls that have
the appearance of true cones consisting of many chambers, each containing a single
adelgid. These galls are of minor importance except on ornamentals, where they
may be undesirable. Heavily infested white pines, especially young pines in
plantations, may be severely injured. The early symptoms are drooping branches
that turn red by summer, and die. In heavy infestations, growth is reduced and
occasionally trees up to 12 cm in diameter are killed. Some outbreaks spread over
large areas before subsiding.

Pineus floccus (Patch), the red spruce adelgid, feeds on red and black spruces
and eastern white pine, spending | year of its 2-year life cycle on each. It produces
loose, terminal galls on spruce. On pine, its effects are similar to those caused by
the pine leaf adelgid. Heavy infestations on spruce may kill the tips of branches or
they may cause an overproduction of laterals, which leads to bushy, deformed trees.
Damage to pine is usually not serious.

Pineus similis (Gillette), the spruce gall adelgid. produces terminal ragged galls
on various spruces but appears to prefer white spruce in the East. No alternate hosts
have been found for this species. The galls are shorter and thicker than those
produced by the Cooley spruce gall adelgid and the chambers inside are intercom-
municating. Small white spruce growing in the open in Canada has been severely
infested. P. coloradensis (Gillette) has been observed feeding on the needles of red
and pitch pines in Connecticut, although its host range is much wider (774).
Laboratory studies indicate that red pine seedlings readily succumb to heavy

infestations of this species.
Family Phylloxeridae
Phylloxeras

Members of this family res able those of the family Adelgidae except that
winged forms have fewer antermal segments and all species feed exclusively on
dicotyledonous plants.

The genus Phylloxera contains several species that produce galls on hickories
and pecan. These aphids do not produce waxy threads as do many other members of
the family. but some of them may be covered with a waxy powder. Their galls vary
from small disklike or buttonlike swellings, with central openings guarded by
plantlike hairs or processes, to large, hollow, globelike structures up to 18 mm in
diameter.

Phylloxera caryaecaulis (Fitch), the hickory gall phylloxera, is a common
species that produces almost spherical galls, 16 to 18 mm in diameter, on the twigs
and leaf stems of hickory. The galls are green when first formed: later, after the
phylloxeras vacate them, they turn brown or black. The pecan phylloxera, P.
devastatrix Pergande. and the pecan leaf phylloxera, P. notabilis Pergande. pro-
duce galls on pecan and other hickories. P. rileyi Riley is found on white and post
oaks. and P. nyssae Pergande infests blackgum.

86

Family Aleyrodidae
Whiteflies

Members of this family are very small mothlike insects, usually less than 2 or 3
mm in length. The adults all have four wings each. The wings, covered with a
powdery wax, are whitish in appearance. The larvae are very small and scalelike;
they are usually found surrounded or covered with a waxy secretion on the
undersides of the leaves of the host plant. Whiteflies are most abundant in tropical
and subtropical regions, but a few species have been recorded as far north as New
England.

The mulberry whitefly, 7etraieurodes mori (Quaintancé), feeds on mulberry,
dogwood, azalea, hackberry, hoily, mountain-laurel, basswood, maple, and syca-
more. The larvae are less than | mm long, jet black, and ringed around with a white
fringe. Adults are active from June to September. The rhododendron whitefly,
Dialeurodes chittendeni Laing, an introduced species, feeds on rhododendron. The
adult is pale yellow; the larvae and pupae are greenish yellow. Infested leaves have a
yellow, mottled appearance, and their margins curl. The citrus whitefly, D. citri
(Ashmead), feeds on chinaberry and on crapemyrtle in Florida. The azalea white-
fly, Pealius azaleae (Baker & Moles), feeds on azalea as far north as Rhode Island
and Ohio, and sometimes causes severe defoliation. Aleurochiton forbesii (Ash-

mead) feeds on maple.
Superfamily Coccodidea
Scale Insects

Scale insects are among the most destructive pests of shade trees, ornamentals,
and greenhouse vegetation, and may cause serious damage to forest growth. Injury
apparently results from the ingestion of large amounts of plant sap and from the
production of honeydew, which serves as a substrate for the growth of sooty mold.
As with aphid honeydew, many beneficial species feed on this material. Plant
deformation and toxin injury are produced by a few scale insects.

Adult female scales lack wings, may have legs, and are saclike with no definite
head, thorax, or abdomen. Adult males are more insectlike in appearance, usually
with one pair of wings, one pair of “halteres,”’ long legs, and a definite head,
thorax, and abdomen. Adult males are rarely collected because they are small and
normally live only a day or two. Most scale insects produce a waxy secretion that
covers the body either as a shieldlike structure separate from the body or as a
coating on the body surface. The wax varies from a thin, translucent sheet to a
thick, wet mass or to a powdery bloomlike secretion.

Some scale insects are host-specific, but others feed on a wide variety of plants.
Because of the small size and cryptic appearance of most of these insects, many
pest species have been distributed inadvertently by commerce as plant contami-
nants.

Natural dispersal is primarily by highly mobile windblown crawlers or first
instars, but dispersal by birds and mammals has also been suggested. In most
instances, instars other than the crawler are relatively sessile and are unimportant in
natural dispersal.

The life stage most susceptible to chemical control is the first instar, apparently
because during part of this stage there is little or no protective wax covering on the
insects. Attempts to control other life stages are frequently unsuccessful. There-
fore, for effective chemical control, it is important that information on the life cycle
be available so that insecticidal spray applications coincide with crawler
emergence.

87

It is important to remember that egg hatch may occur at different times in
different areas of the United States. Also, some species have short egg-hatch
periods, while others have hatch periods that extend over entire growing seasons.

Natural enemies are frequently effective control agents of scale insects. In
forests, natural enemies are commonly encountered components of the scale insect

ecosystem.
Family Margarodidae
Margarodid Scales

Margarodids are primarily tropical or subtropical in distribution, although sev-
eral groups occur in temperate areas. The family can be divided into two groups—
i.e., the cyst-formers and the noncyst-formers. The cyst-formers produce a heavily
sclerotized, legless stage that develops between the legged crawler and the adult. In
at least one species, this so-called cyst or resting stage is highly resistant to
environmental restraints and has been stored alive for 17 years (S34). Noncyst-
formers are generally similar in appearance throughout their development and do
not form a special “resting” stage. Margarodids may occur anywhere on the host
from the roots to the foliage; many species occur under the bark of their hosts. The
family Margarodidae contains 42 species of 11 genera in the United States. A
comprehensive study of this family is available (887).

The red pine scale, Matsucoccus resinosae Bean & Godwin, was first reported
in western Connecticut on red pine in 1946. It has now spread northward and
eastward in Connecticut and is found in southeastern New York and northern New
Jersey, and an infestation was recently discovered in Morris Arboretum, Phila-
delphia, Pa. (//63). The species apparently is expanding its distribution at a rate of
1 to 3 km per year (77/). Red pine scale occurs only on pines of the sylvestris
subsection of Pinus, which, in North America, is represented by one native
species—red pine (fig. 30). The subsection is diverse in Europe and Asia where
over 17 species are reported. This Eurasian diversity has caused many to suggest
that red pine scale may not be native to the United States but rather 1s introduced
from Eurasia (/7). This idea is supported by its distribution in the United States,
which is entirely outside the natural range of red pine. Red pine scale has been
reported on red, Chinese, Japanese red, and Japanese black pines, and is restricted
to plantations, nurseries, and oraamental plantings.

The species has two generations per year and overwinters as a first-instar crawler
under or in cracks of the bark (79). In April the cyst stage appears, and by the end
of the month the waxy sacks of developing males are obvious on the undersides of
branches. In May and June, adults and eggs are present. Hatching occurs in about
15 days, and crawlers of the summer generation reach the adult stage in August. In
late August the overwintering first instars begin to appear. The red pine scale is the
most destructive insect pest of red pine in the Northeast. Thousands of young to
fully mature trees have been killed. The foliage of infested trees generally changes
progressively from olive green to yellow and then to red. The bark appears swollen
and cracked, and an area of dead tissue is present beneath each feeding scale. It has
been suggested that the death of infested trees might be partially caused by a toxin
produced by the scale (986). Long-range dispersal is principally by wind (772). No
effective natural enemies are known in the United States but biological control
exploration is underway in the Orient. In one experiment, exposure to — 23° C for
two 4-hour periods was lethal to 99 percent of the overwintering crawlers (53/), but
this experiment has never been duplicated. Chemical control is possible, but this is
practical only on ornamental trees.

88

Courtesy Conn. Agric. Exp. Stn.

Figure 30.—Masses of male cocoons and a closeup of
the red pine scale, Matsucoccus resinosae.

Matsucoccus gallicolus Morrison, the pine twig gall scale, is apparently a native
species. It has been recorded in 16 States from New Hampshire to Florida and west
to Misscuri. It occurs on loblolly, pitch, ponderosa, red, shortleaf, spruce, Table
Mountain, and Virginia pines.

The pine twig gall scale has one generation per year and normally overwinters as
an egg. Hatching occurs when the new year’s growth is 2.5 to 7.5 cm long (/49).
The crawlers migrate from the bark to the new green growth where they settle. As
feeding progresses, cells beneath the scale body collapse, eventually leaving the
crawler in a small depression. Molting occurs, and the first-instar crawler trans-
forms into a cyst. The host plant grows around the cyst until the scale is completely
enclosed except for a small hole that apparently is kept open by the trapped shed
skin of the crawler. By the end of July, the cyst molts to the adult female stage; she
squeezes through the hole in the gall, migrates to the trunk or a main branch, and
lays the overwintering eggs under the bark. Adult males have not been reported.

The pine twig gall scale is an important pest of pitch pine and may kill mature
trees, although it 1s most detrimental to young trees (959). This scale frequently
kills limbs and is particularly injurious to ornamental plantings.

Matsucoccus alabamae Morrison, the Alabama pine scale, is reported on the
bark of pines in Alabama, but no major damage has been attributed to this species.
M. macrocicatrices Richards, the Canadian pine scale, occurs in southeastern
Canada and New Hampshire where it is found on eastern white pine. The scale is
unique among the pine scales in that it is associated with the fungus Septobasidium
pinicola Snell in an apparent mutualistic relationship.

Canadian pine scale has one generation every 2 years and overwinters in the cyst
Stage in fungal mats (/255). Eggs are laid in cracks of the bark early in the year.
Crawlers migrate to the edges of fungal mats where they feed and molt to the cyst
stage. The cyst remains in the fungal mat for nearly 2 years before transforming to

89

the adult female early in the third spring. Males have been reported (/025), but
apparently are rare. No major plant damage is attributed to this scale, although
minor host deformation at feeding sites is sometimes apparent when fungal mats are
removed.

Xylococculus betulae (Pergande), the birch margarodid, apparently is a native
species. It occurs from eastern Canada to Tennessee and west to Ohio on birch,
beech, and occasionally on willow and maple. Adult females are bright orange,
about 4 mm long, and are covered with white wax.

The length of a generation is unknown, but a single cycle probably requires more
than | year. The crawler settles in cracks or lenticles on the trunk of the host,
produces a waxy cell, and eventually is covered by the surrounding bark. The
crawler molts to the cyst, which produces a thin, waxy tube that protrudes up to 5
cm above the bark surface and acts as a passage for honeydew (620). The adult
female has legs but normally remains inside the shed skin of the cyst. A hole is
formed near the apex of the cyst-shed skin through which the male mates and the
crawlers emerge to the host surface. The birch margarodid is reported to kill young
birch trees under some circumstances, although damage is normally restricted to
localized necrotic areas on the bark (/079). On beech this scale insect normally
does not kill the tree but causes roughened or swollen spots on the trunk that
eventually dry and form large cracked areas. Damage is usually restricted to narrow
longitudinal strips that start at old branch stubs.

Neosteingelia texana Morrison is native to North America. It occurs in Alabama,
Florida, Georgia, Pennsylvania, South Carolina, Tennessee, Texas, and Virginia. It
is most often collected on pecan and hickory, but material is at hand from
hackberry, sugar maple, and sweetgum. The species occurs 1n native and ornamen-
tal habitats. Adult females and males are unusually large for a scale and are reddish
brown.

This margarodid probably requires more than | year for a generation in Virginia,*
and at least part of its populations overwinter as eggs. Eggs are laid under the bark,
and they hatch in early spring. Adults mate on the bark in late September or early
October. Information is not available on its economic importance or natural en-
emies.

Family Ortheziidae
Ensign Scales

This family is a homogenous group that occurs predominantly in the New World,
although a limited number of species is known from New Zealand, Europe, the
Soviet Union, Japan, and Africa. Ensign scales are easily recognized in the field by
the sharply defined, platelike, waxy tufts over much of their bodies. The ovisac is
constructed of similar wax and is attached to the venter of the scale. This allows the
female to move on the host even while ovipositing. Apparently there are four instars
in the female and five in the male. Males are generally rare. The family contains 32
species in 4 genera in the United States (S86, 888).

In eastern forests several members of this family may be encountered. Orthezia
tillandsiae Morrison, the Spanish moss orthezia, occurs on Spanish moss in
Florida, Georgia, Louisiana, and Virginia, and probably in all areas of the United
States where Spanish moss occurs naturally. O. pseudinsignis Morrison occurs on
many hosts, including walnut, in Louisiana.

+ Kosztarab, M. (personal communication). Virginia Polytechnic Institute and State University, Blacks-
burg, Va.

90

Family Pseudococcidae
Mealybugs

This is a large and diverse family of scale insects that is cosmopolitan in
distribution. Mealybugs are recognized in the field by the filaments around the
body margin and a covering of mealy wax. In general there are four female instars
and five male instars. The family contains 281 species in 45 genera in the United
States. Detailed publications are available for this group (409, 471, 796).

Dysmicoccus wistariae (Green), the taxus mealybug, is apparently an intro-
duced species. In the United States it occurs in 15 Northeastern and Midwestern
States from Maine to Maryland and west to Missouri. It is most abundant on yew
but 1s also reported on maple, rhododendron, dogwood, and Prunus spp. The taxus
mealybug normally occurs in ornamental plantings. Adult females are covered by a
white wax except on the dorsum, where four naked areas allow the red body
pigments to show through as longitudinal lines. The margin of the body is adorned
with 15 to 17 pairs of waxy filaments; the posterior pair is the longest and is about
one-fourth the length of the body. A small, filamentous ovisac is sometimes
constructed beneath the posterior part of the abdomen.

The taxus mealybug has one generation per year in Connecticut (/068) and parts
of New York, and two or three in Massachusetts, New Jersey (5/7), and southern
areas of New York. The species overwinters in the first instar in cracks in the bark;
males are common. Eggs apparently hatch inside the body of the female. Damage 1s
reported on yew where heavily infested plants have sparse foliage, turn yellow, and
are stunted and covered with honeydew and sooty mold (/268). Heavy infestations
can kill young plants.

The striped mealybug, Ferrisia virgata (Cockerell), is probably native to the
New World. In the United States it occurs in 23 States from New York to Florida
west to Louisiana and Missouri; it 1s also known in several Western States. The
striped mealybug is polyphagous; some of its eastern hosts are maple, azalea,
boxwood, catalpa, dogwood, hawthorn, holly, magnolia, apple, mulberry, and
persea. It is found in natural habitats and ornamental plantings. The gray body of
the adult female is covered with a white wax except on the dorsum, where two
longitudinal areas are without wax and form stripes. There 1s a single pair of caudal
filaments extending about one-half the length of the body. Many long, thin,
crystalline rods are present on the dorsum. The adult female produces a small waxy
pad beneath the abdomen on which the eggs are laid. Males may be present.

The striped mealybug has two generations per year in Maryland on azaleas, and
overwinters as a second or third instar (568). Feeding begins in early April in leaf
axils or at the base of flower buds, and adults appear in early May. Eggs are laid and
hatch during the second week of June. Adults of the summer generation are present
in late July, and first instars of the winter generation first appear in early August.
Damage may occur on most hosts of the striped mealybug. Many natural enemies
are reported for this pest, including chalcidoid wasps, lady beetles, flower flies, and
gall midges.

The apple mealybug, Phenacoccus aceris (Signoret), 1s apparently an intro-
duced species from Europe. In the United States it occurs in five States from Maine
south to Massachusetts in the East. This species is polyphagous, although it prefers
rosaceous hosts such as apple and hawthorn. The species is principally a pest of
ornamentals, although it is found occasionally in natural habitats. The green body
of the adult female is covered by white wax. The body margin has 16 to 18 pairs of

91

lateral filaments; the caudal pair is about one-eighth as long as the body. An ovisac
is produced that encloses all but the anterior portion of the adult female.

The apple mealybug has one generation per year and overwinters as a second
instar in crevices in the bark. In early spring, the second instars migrate to the
leaves or new growth of the stems. In late spring and early summer, adults appear,
mating takes place, and egg laying begins. First instars appear in July and feed
primarily on the leaves. Second instars migrate from the leaves to the bark in the
fall. The apple mealybug is an economic pest in apple orchards in Maine, Nova
Scotia, and British Columbia, and is occasionally a pest of ornamentals such as
maple, basswood, and cotoneaster. In many areas this mealybug is held to low
populations by the small wasp Allotropa utilis Muesebeck. Other natural enemies
include chalcidoid wasps and lady beetles.

Phenacoccus acericola King, the maple mealybug or maple phenacoccus, is
probably an introduced species. In the United States it occurs in 16 States from
Maine to Tennessee and west to Illinois. The preferred host is sugar maple, but it is
also reported on other species of maple and on buckeye, basswood, and hornbeam.
The species is normally a pest in ornamental plantings, but is also present in natural
habitats. The adult female is yellow and is lightly dusted with a flocculent white
wax. The body margin has 16 to 18 pairs of broad, waxy filaments which are less
than one-fourth the length of the body. An ovisac encloses all but the anterior end of
the adult female.

In Ohio, the maple mealybug has two or three generations per year on sugar
maple and overwinters in bark crevices as immatures (605). The overwintering
immatures migrate to leaves of the host and feed in the early spring. In June the
adult females migrate to the bark and mate. The adult females return to the leaves,
produce a large ovisac, and lay more than 500 eggs. The ovisacs generally are
produced on the underside of the leaves near the primary veins. Damage caused by
the maple mealybug principally detracts from the appearance of the host. Several
chalcidoid wasp parasites and lady beetle predators have been found associated with
the maple mealybug.

Phenacoccus dearnessi King, the twocirculi mealybug, is of uncertain origin; it
may be a native species. In the East it occurs in Illinois, Indiana, lowa, Kansas,
Maryland, Missouri, New York, Ohio, Pennsylvania, and Wisconsin; it is also
reported in five Western States. The species is restricted to rosaceous hosts:
serviceberry, cotoneaster, hawthorn, quince, and Prunus spp. It is a pest in orna-
mental plantings, although it is found also in natural habitats. The adult female is
red and is covered by a thin layer of white wax. The lateral filaments are short and
inconspicuous or are absent. The ovisac may be absent or restricted to the posterior
part of the venter. Males are abundant.

The twocirculi mealybug has one generation per year and overwinters as second
instars in crevices in the bark. In the spring the females move and begin feeding in
the crotches of twigs or at the bases of bud scales. Adult females produce living
young or eggs that hatch within minutes after being laid. First instars settle on the
leaves and begin feeding. In the summer the crawlers migrate to cracks in the bark
and go through a summer diapause period. During early fall the crawlers return to
the leaves, feed, and develop into second instars. In late fall the second instars
move to the overwintering sites. Heavy populations of the twocirculi mealybug
cause accumulations of large quantities of honeydew and sooty mold and give the
host plants an unsightly appearance. Large infestations may cause early leaf drop.
Natural enemies are not reported for this pest. Normally, chemical control is fairly
successful.

92

Spilococcus juniperi (Ehrhorn), the juniper mealybug, is a native species. In the
East and Midwest it occurs in Florida, Indiana, lowa, Kansas, and Nebraska; it is
also reported from six Western States. The species is restricted to juniper and seems
to prefer redcedar and oneseed juniper. The juniper mealybug is found in native
habitats and ornamental plantings. The adult female is dark purple, is covered with
a smooth gray wax, and has two bare areas forming longitudinal stripes on the
dorsal part of the thorax and abdomen. The body margin has several short, partially
coalesced filaments. The ovisac encloses the venter of the female, forming a
nestlike structure.

The juniper mealybug’s life history is unknown except that it feeds on the foliage
of the host and lays eggs. This mealybug is a serious pest of ornamental plantings of
redcedar and oneseed juniper in Kansas, Oklahoma, Texas, and Indiana. The
foliage of heavily infested trees turns brown and drops, beginning on the lower and
inner branches of the host. Entire trees are sometimes defoliated. Two chalcidoid
wasps have been reared from this mealybug.

The Comstock mealybug, Pseudococcus comstocki (Kuwana), probably was
introduced from the Orient. In the Eastern United States it occurs from New
Hampshire to Florida and west to Missouri; it has also been reported from two
Western States. This mealybug is polyphagous and is frequently collected on
catalpa, mulberry, and yew. Other eastern forest hosts are holly, buckeye, maple,
and poplar. The species is normally restricted to ornamental plantings. The adult
female is reddish brown and is covered with a white wax in which two dorsal bare
areas form longitudinal stripes. The body margin has approximately 17 pairs of
long filaments; the caudal pairs are longest and are about one-third to three-fourths
the length of the body. The filamentous ovisac normally covers all of the adult
female.

The Comstock mealybug has three generations per year, and in Virginia (604)
and California (64) it overwinters as an egg in the ovisac. Adult females usually
move from the feeding areas and oviposit in protected areas on the bark. Ovisacs
contain 200 to 300 eggs; with the exception of overwintering periods, eggs hatch |
to 2 weeks after being laid. Males are common and apparently are required for
reproduction. This mealybug has many natural enemies, including chalcidoid
wasps, platygasterid wasps, lady beetles, green and brown lacewings, and a fly.
Two parasites that have been effective biological control agents are the wasps
Pseudaphycus malinus Gahan and Allotropa convexifrons Muesebeck.

The grape mealybug, Pseudococcus maritimus (Ehrhorn), is a native species.
Because of confusion surrounding the proper identity of P. maritimus and the
closely related species P. obscurus Essig, much of the literature about P. maritimus
actually pertains to P. obscurus. In the United States the grape mealybug is reported
in 21 States; in the East it occurs in 18 States from New York to Florida and west to
Missouri. The species 1s polyphagous, but in the East it often causes damage to yew
and grape. [t occurs in natural habitats and ornamental plantings. The yellow to
orange body of the adult female is covered with a gray wax. The margin of the body
has 17 pairs of thin lateral filaments; the caudal pair is about one-fourth the length
of the body. An ovisac is produced that encloses all but the head of the mealybug.

The grape mealybug has two generations per year in central California (8/2) and
Ohio (9/4) and overwinters as first instars in the ovisac in cracks in the bark. In
early spring the first instars move to the foliage or flower buds of the host and feed.
In early summer adult females move to the bark of the host and oviposit. The eggs
hatch in July, and the first instars feed on the green portions of the stems. In late

93

summer the second generation females mature, move to the bark of the host. and
lay eggs. The eggs hatch and the first instars remain in the ovisac for the winter.
Occasionally. second and third instars may overwinter. Males are known. Many
natural enemies of this mealybug are reported, including chalcidoid wasps, several
lacewings, and five lady beetles. Chemical control apparently is difficult.

Many other mealybug species may be encountered in eastern forests. Because of
this diversity. it is impossible to give detailed information about all species that
might be found. However, additional species include Dysmicoccus morrisoni (Hol-
linger) on pecan and hickory in Alabama, Georgia, Louisiana, Mississippi, Mis-
souri. New Jersey. and New York: D. obesus (Lobdell) on pine in Mississippi: D.
difficilis (Lobdell) on blackgum, ash, and willow in Louisiana, Maryland. Mis-
sissippi, South Carolina, and Virginia: Oracella acuta (Lobdell) on pine, particu-
larly loblolly. in Florida, Georgia, Louisiana, Maryland, Mississippi, North Car-
olina, South Carolina, Texas, and Virginia: and Peliococcus serratus (Ferris) on
beech in Connecticut, District of Columbia, Massachusetts, Maryland, New Jersey.

New Hampshire, New York, Ohio, Pennsylvania, and Virginia.
Family Coccidae
Soft Scales

Soft scales are common pests of ornamentals throughout most of the world,
although they generally are more abundant in tropical and subtropical areas. These
scales are generally without an obvious waxy covering, although members of a few
genera form conspicuous ovisacs or are covered with an ornate, amorphous wax.
There are three or four instars in females and five in males. The family contains 90
species in 22 genera in the United States. Comprehensive studies of this family in
the United States have been published (7/54, 13/0).

Ceroplastes ceriferus (F.), the Indian or Japanese wax scale, was apparently
introduced from the Orient. In the United States it occurs in 16 States from New
York to Florida and west to Texas, although it probably does not survive out of
doors north of Delaware. The host range is extensive, including such plants as
Japanese and Chinese hollies, euonymus, camellia, and eastern hemlock. The
Indian wax scale is normally found in ornamental plantings. Adult females are
covered by a white amorphous wax that is hemispherical in shape and has an
anteriorly projecting horn. The body of the adult female is pinkish purple to dark
reddish-brown.

The Indian wax scale has one generation per year in Virginia and Maryland
(1105): it is possible that more than one generation occurs in southern areas. In
Maryland the species overwinters as adult females. In !ate spring the adults begin to
lay eggs that hatch in about 3 weeks: a single female may lay more than | .000 eggs.
First instars settle on the stems of the host and develop into adult females by late
summer. Males are uncommon. This wax scale does not usually kill its host but
frequently gives the heavily infested plant a scraggly appearance and covers the
host with honeydew, which eventually is contaminated with black sooty mold
(486). Natural enemies include several species of chalcids and a pyralid moth.

Other less common species of Ceroplastes that might be encountered in the East
are the barnacle scale. C. cirripediformis Comstock, which occurs on citrus trees
and gardenia, and the Florida wax scale. C. floridensis Comstock, which is found
on pine, hemlock, and maple. Both species occur in the warm Southern States on
many native and ornamental trees. .

The calico scale, Eulecanium (= Lecanium) cerasorum (Cockerell), is probably
native to the Orient. In the Eastern United States it occurs in Delaware, District of

94

Columbia, Maryland, New Jersey, New York, Pennsylvania, Rhode Island, and
Virginia. The host range is extensive, including many common ornamentals; in
Maryland it is often found on dogwood, maple, the ornamental star magnolia, and
ornamental fruit trees, and is rarely reported in natural habitats. The mature calico
scale is globular and has irregular white patches on a dark-brown background. As
the scale ages, the white patches disappear.

The calico scale in Maryland has one generation per year and overwinters as
second instars on the bark of the host.° In early spring, second instars begin to feed,
molt twice, and develop into globular adults. In late spring and early summer, eggs
are laid. During June, first instars appear on the undersides of leaves. In late
summer and early fall, the crawlers molt to second instars and move from the leaves
to the stems and trunks where they overwinter. Adult males have not been observed
in Maryland. The calico scale normally does not kill trees, but produces large
amounts of honeydew, giving the host an unsightly appearance. Two chalcidoid
wasps have been reported associated with this scale.

Eulecanium (= Lecanium) caryae (Fitch), the large hickory lecanium, is
known primarily in the Northern States from Maine to Virginia and west to Kansas;
the species is also reported from Mississippi. The large hickory lecanium is
polyphagous, being found on most trees in the eastern forests, including hickory,
birch, elm, beech, walnut, oak, and mulberry. This scale is found in native habitats
and ornamental plantings. Adult females are unusually large, sometimes reaching
15 mm in length. The body is normally flat and uniformly brown or orange-brown
and is covered with a white bloom.

The large hickory lecanium has one generation per year in Michigan (/24/) and
overwinters as immatures. Adult females lay over 100 eggs. Hatching occurs in the
summer and first instars feed on the undersides of leaves. In late summer the
immatures move from the leaves to the stems, where they overwinter (/3/0). On
elm this species tends to infest medium-size branches.° The large hickory lecanium
is normally not an economic pest but is a common inhabitant of eastern forests.
Three species of chalcidoid wasps are reported as associated with this scale insect.

The terrapin scale, Mesolecanium (= Lecanium) nigrofasciatum (Pergande), 1s
probably native to North America. In the Eastern United States, it is known from
every State except New Hampshire and Vermont. The species is polyphagous, and
is commonly found on maple, sycamore, and fruit trees in natural habitats and
ornamental plantings. Fully developed adult females are hemispherical and are dark
reddish-brown with many radiating bands extending from the center of the dorsum
to the raised edge of the body.

In Virginia, the terrapin scale has one generation per year and overwinters as
adult females (73/0). In early spring, first instars are produced and remain under
the body of the adult for 1 to 3 days before moving to the undersides of leaves. In
about 18 days, first instars molt to second instars that molt to adult females in a
similar period of time. In the fall, adult females move from leaves to twigs where
they overwinter. Males are usually common in early summer. The terrapin scale has
been reported as a serious pest in fruit tree orchards. It is commonly encountered on
forest and shade trees and produces large amounts of honeydew. Over 30 natural
enemies have been recorded as associated with this scale, including chalcidoid
wasps, lady beetles, green and brown lacewings, a pyralid moth, and fungi.

> Stoetzel, M. B. (personal communication). USDA ARS Syst. Entomol. Lab., Beltsville, Md.
® Weidhaas, J. A. (personal communication). Virginia Polytechnic Institute and State University, Blacks-
burg, Va.

95

The magnolia scale. Neolecanium cornuparvum (Thro), is probably native to
North America. The species occurs in at least 15 Eastern States from New York to
Florida and west to Wisconsin. This scale is restricted to species of magnolia and is
found in natural habitats and ornamental plantings (fig. 31). Newly mature adult
females are covered with a white bloom that is rapidly lost with age. Older females
are brown or yellow, are convex, and are very large, attaining a length of 12 mm.

ce

Ex

Courtesy Conn. Agric. Exp. Stn.

Figure 31.—The magnolia scale, Neo/ecanium
cornuparvum, on magnolia.

The magnolia scale has one generation per year in New York, Connecticut, and
Virginia, and overwinters as first instars on I- or 2-year-old growth. First instars
begin developing in early spring and after at least two molts become adult females
in early August. First instars are born as nymphs in late summer and fall. Heavy
infestations of the magnolia scale can kill branches or entire trees. Large amounts of
honeydew may cover the host. The only natural enemy reported for this scale insect
is a lady beetle.

The European fruit lecanium, Parthenolecanium (= Lecanium) corni
(Bouché), is acommon cosmopolitan pest. Contrary to general opinion, this pest 1s
apparently indigenous to North America, not Europe (426). The European fruit
lecanium occurs throughout the United States on many hosts in both natural and
ornamental habitats. Proper identification of the species is difficult because its
morphology is greatly influenced by the host (363). Until biological studies are
undertaken, it is impossible to sort out host-induced variations from differences
characterizing separate species. The oak lecanium, P. (= Lecanium) quercifex
(Fitch), and the Fletcher scale, P. (= Lecanium) fletcheri (Cockerell), are fre-
quently impossible to separate from P. corni on the basis of microscopic features. P.
quercifex is reported from oaks and P. fletcheri from juniper, northern white-cedar,
and yew. The color patterns and body shape of the European fruit lecanium vary
according to the host. Newly formed adult females are flattened and have a mottled
appearance, but as they mature they become more hemispherical and turn a uniform
brown.

The European fruit lecanium normally has one generation per year and overwin-
ters as second instars. A rapid second generation has been recorded for a few
populations on some hosts (/92). In Pennsylvania a second generation has been

96

reported on peach (28). In late spring, from 50 to more than 2,000 white eggs are
laid beneath the body of the adult female. The number of eggs is generally
proportional to the size of the adult female. In early summer, eggs hatch and first
instars move principally to the undersides of the leaves, although a few settle on the
stems. In late summer or early fall, molting occurs and the resultant second instars
move from the leaves to rough areas of branches, where they overwinter. In early
spring, second-instar females grow rapidly, molt, and develop into adults. Males
may be present or absent, apparently depending on the host or environmental
conditions (426). The European fruit lecanium normally does not kill trees but
produces a large quantity of honeydew which, when contaminated with sooty mold,
covers the plant with an unsightly black film. The species is sometimes a serious
pest of plums, apricots, and other fruit trees. Over 40 natural enemies have been
found associated with the European fruit lecanium, including fungi, lady beetles,
and chalcidoid wasps. Some of the parasitic hymenopterans are effective control
agents.

The cottony maple scale, Pulvinaria innumerabilis (Rathvon), is apparently
native to North America and is known from nearly every State in the United States.
This soft scale prefers maples, particularly silver maple, but is reported on a large
number of important hosts, including basswood, ash, dogwood, hackberry, locust,
sycamore, hawthorn, pear, oak, poplar, elm, willow, apple, peach, plum (fig. 32).
The cottony maple scale is found in natural habitats and ornamental plantings.
Adult females have variable color patterns but are generally reddish brown, and
have a median ridge. The ovisac is one to two times longer than the adult and is
produced by the raised, ventral abdomen.

Courtesy Can. For. Serv.,
Can. Dep. Environ., Sault Ste. Marie, Ont.
Figure 32.—The cottony maple scale, Pulvinaria
innumerabilis, on twigs and leaves of soft maple.

The cottony maple scale has one generation per year (/3/0) and overwinters as
mated, newly mature, adult females on the twigs and branches of the host. The
literature often refers to the overwintering stage as immature females, but slide-
mounted specimens collected during the winter show that the species overwinters as

97

adult females. In early spring, the adult females enlarge rapidly; ovisac develop-
ment and egg laying begins in mid to late spring. A single female may lay 1,000
eggs. First instars begin to appear in early summer and are present until August.
First instars move from the branches to the leaves where they develop into adult
females in late summer and early fall. Adults mate in August and September.
Before leaf drop, adult females move to the stems of the host. Heavy populations of
this scale may cause dieback of branches and under extreme conditions may kill
entire trees. This soft scale produces large amounts of honeydew, which frequently
supports growth of sooty mold. Natural enemies include six species of chalcidoid
wasps, several lady beetles, a green lacewing, a pyralid moth, and the English
sparrow.

Pulvinaria acericola (Walsh & Riley), the cottony maple leaf scale, is appar-
ently indigenous to North America and is common throughout the Eastern United
States. The preferred host is maple, but it is also found on dogwood, persimmon,
holly, blackgum, sassafras, Prunus spp., and andromeda. This soft scale may be
found in natural habitats and ornamental plantings. The adult female is purple with
a longitudinal yellow stripe down the center of the dorsum. An ovisac is produced
with two or three longitudinal ribs.

The cottony maple leaf scale has one generation per year and overwinters as
second instars on the stems and branches. Adults are present in spring when mating
takes place. Adult females move to the leaves and form ovisacs in late spring. or
early summer. The adult female normally falls from the sac soon after it is
completed. Each ovisac may contain nearly 2,500 eggs (644), and hatching begins
several weeks after the eggs are laid. First instars settle on the leaves where they
molt once. In the fall the second instars move back to the twigs and branches where
they overwinter. The cottony maple leaf scale normally does not cause damage, but
heavy infestations may occasionally cause early leaf drop and dieback of branches.
Natural enemies include seven chalcidoid wasps and a lady beetle.

Other species of Pulvinaria that are common in eastern forests are P. floccifera
(Westwood), the cottony camellia scale or cottony taxus scale, known in the East
from Connecticut to Florida and west to Missouri on many hosts, including
camellia, Chinese holly, and yew. The cottony camellia scale is yellow to light
brown with a dark body margin and has a life cycle similar to that of the cottony
maple leaf scale. It can be an important pest of ornamentals.

Pseudophilippia quaintancii Cockerell, the woolly pine scale, is indigenous to
the United States where it occurs on several pine species including Swiss mountain,
loblolly, longleaf, Table Mountain, and pitch pines. This soft scale is reported from
New Jersey to Florida and west to Louisiana and is restricted primarily to natural
habitats. The adult female is elongate oval, yellow, light brown, or greenish yellow,
and is covered with a characteristic white, cottony, waxy secretion. The biology of
this species has not been studied. The woolly pine scale occasionally may be
abundant enough to cause stunting of growth (/0/2).

The spruce bud scale, Physokermes piceae (Schrank), was apparently intro-
duced into North America. It is known in Indiana, Massachusetts, Maine, Mary-
land, Minnesota, Missouri, New Hampshire, New York, Pennsylvania, and
Wisconsin. The taxonomic status of this species in the United States is currently in
question. It is now established that at least some of the reported records of P. piceae
should be treated as P. hemicryphus Dalman. The spruce bud scale is found on
spruce, particularly Norway spruce. Adult females are round, reddish brown, and
closely resemble leaf buds of the host.

98

The spruce bud scale has one generation per year and overwinters as immatures.
Adult females appear in spring, and eggs are retained in the adult female’s body
cavity. First instars appear in late spring and settle on the new growth of the host.
Severe damage may involve the death of lower branches of the host. Chalcidoid
wasps are often found associated with this scale.

The tuliptree scale, Zoumeyella liriodendri (Gmelin), 1s apparently native to
North America. In the Eastern United States, it occurs from New York to Florida
and west to the Mississippi River. This soft scale prefers yellow-poplar and magno-
lias, although it is reported on several diverse hosts, including walnut and
basswood. The tuliptree scale is found in native habitats and ornamental plantings.
Adult females are large, convex, and are usually orange with black mottling near
the body margin. Heavy infestations are common (fig. 33).

F-532855

Figure 33.—The tuliptree scale, Toumeyella liriodendri,
on yellow-poplar.

The tuliptree scale has one generation per year except in southern parts of its
range, where nonsynchronous populations suggest that there might be at least two
generations. The species overwinters as second instars on the twigs of the host. In
early spring the second instars resume feeding. Males appear in early summer and
mating takes place. By late summer the greatly enlarged ovoviviparous adult
females produce as many as 3,000 first instars. These immatures settle on the
stems, feed, and molt to second instars late in the fall. Four kinds of tuliptree scale
damage are reported on yellow-poplar (/76), ranging from greatly weakened trees
with sparse foliage and many dead branches to trees with distorted trunks because
of the death of a previous leader. Seedlings are frequently killed, greatly hindering
future development of economically valuable stands of yellow-poplar. Natural
enemies include five species of chalcidoid wasps, a flower fly, a pyralid moth, two
lady beetles, and a fungus.

The pine tortoise scale, 7. parvicornis (Cockerell), is native in North America.
It is found in nearly every Eastern State and occurs as far west as the Dakotas. This
soft scale seems to prefer Scotch, jack, and Virginia pines, but is also found on

99

Austrian, Swiss mountain, red, loblolly, shortleaf, white, and Chinese pines. The
pine tortoise scale is common in natural habitats and ornamental plantings. Adult
females are convex and are dark brown or black with reddish-brown or cream-
colored mottling. It is possible that specimens identified as 7. parvicornis may
represent more than one species. Specimens collected from needles in the Southeast
may be distinct, but host transfers would be required to establish this fact.

The pine tortoise scale has one generation per year in the Northern States (763),
two generations per year in Maryland (789), and may have more in the Southern
States. In the North, the species apparently overwinters as mated females on the
stems of the host. By late spring, the adult females have greatly enlarged. In early
summer, each female lays about 500 eggs, which hatch soon after being deposited.
First instars develop into adults and mate in mid to late summer. Damage caused by
the pine tortoise scale can be quite severe, particularly in the Northern States.
Injury most frequently is incurred by seedlings and saplings, but heavy infestations
may also damage mature trees. Infested trees may have chlorotic needles or many
dead branches. In some cases, entire trees may be killed. Normally, this soft scale is
held in check by its natural enemies, which include three chalcidoid wasps, nine
lady beetles, and a pyralid moth.

Other Joumeyella species that may be found in eastern forests are 7. pini (King),
the striped pine scale, from Connecticut to Florida and west to Michigan on
shortleaf, Swiss mountain, red, pitch, Scotch, Virginia, and lodgepole pines, and T.
virginiana Williams & Kosztarab, the Virginia pine scale, from Florida, Georgia,
Maryland, and Virginia on longleaf, loblolly, and Virginia pines. The striped pine
scale is reddish brown with a central white stripe on the dorsum. The Virginia pine
scale is uniform salmon pink to reddish brown and is frequently found beneath the

bark.
Family Kermesidae
Gall-Like Scales

This family is unquestionably the least studied of the scale insects in the United
States. Most taxonomic descriptions are based on color patterns of the gall-like
adult females. These patterns have proved to be relatively unreliable. Two studies
that clarify this problem have been completed (34, /7/). Kermesids are Holarctic in
distribution and are almost entirely restricted to oaks (fig. 34). There are approx-
imately 30 species in 2 genera in the United States. Of these species, perhaps only
three occur in the East; biological information is available for only two.

F-519574
Figure 34.—Female scales of Kermes sp. on white
oak twigs and leaves.

100

Kermes pubescens Bogue is known from Kansas and Maryland on white and bur
oaks. Mature adult females are 2 to 3 mm in diameter and are brown with white
bands. There 1s one generation per year, and in Maryland (776) the overwintering
Stage is the first instar. The overwintering first instar sheds its skin about the time
that leaves first appear and then it moves to the new growth. Two more molts occur
by late May or early June, and eggs are laid in late June to late July. The first instars
move from the new growth and settle on main branches or the trunk, where they
overwinter. In the spring, female crawlers move to new growth, including petiole
and main leaf veins, but males remain on trunks and main branches. This species is
reported to cause leaf distortion and flagging terminals of the oaks, especially in
urban areas.

Kermes galliformis Riley, often mentioned as K. kingii Cockerell, is known in
Massachusetts and Virginia on red and black oaks. There is one generation per year,
and in Virginia (522) first instars overwinter in crevices in the bark. In early spring,
the first instars molt and migrate. The second-instar females move to the new
growth of the tree, usually at the base of a leaf petiole. The second-instar males
move down the tree and often settle on the trunk; they also settle on wood chips,
acorns, or stones early in June, and adults appear in mid-June. Mating occurs soon
after the last molt, and egg laying begins in late July. A female may lay as many as
5,800 eggs, which hatch in early September. First-instar females tend to settle on
stems of the host; first-instar males settle primarily on the trunk. Under circum-
stances of host stress, this scale is reported to cause flagging on heavily infested
oaks. Natural enemies include a parasitic wasp, a predacious moth, and a lady
beetle.

Recent studies on first instars’ have revealed that only three of the nine eastern
species can be differentiated morphologically. These are K. pubescens, K. gallifor-
mis, and K. andrei King. The latter is from white and red oaks in Massachusetts.

Family Eriococcidae
Eriococcids

Members cf this family are principally temperate in distribution, although a
small group of species occurs in the tropics. Eriococcids are frequently confused
with mealybugs but are easily separated by the absence of the mealy, waxlike
secretion typical of most mealybugs. There are three female instars and five male
instars. The overwintering stage usually is the adult female or the egg. The family
contains 54 species in I1 genera in the United States (4/2). Species of Cryptococ-
cus are placed in the family Cryptococcidae by some authors.

The beech scale, Cryptococcus fagisuga Lindinger, was apparently introduced
from Europe around 1890 near Halifax, Nova Scotia, on European beech. The first
United States record is from the Arnold Arboretum, Boston, Mass., in 1929, and
there is evidence suggesting the presence of the scale there 10 years before. The
beech scale is reported in Connecticut, Maine, Massachusetts, New Hampshire,
New Jersey, New York, Pennsylvania, Rhode Island, and Vermont. It infests
American, Oriental, and European beeches and is found in natural situations as well
as ornamental plantings.

The life history of the species is variable, depending upon environmental condi-
tions, but the general pattern is as follows: There 1s one generation per year, and
males are unknown even in heavy infestations. The yellow eggs are laid in a

’ Baer, R., and M. Kosztarab (personal communication). Virginia Polytechnic Institute and State
University, Blacksburg, Va.

101

filamentous ovisac in late spring and summer and require a period of at least 20
. days to hatch. First-instar crawlers appear in late summer or early fall and settle in
cracks or lenticels on the trunk or large branches of the host. Soon after settling, the
crawler produces a woolly secretion that encloses the body. The crawler is normally
the overwintering stage, although eggs are occasionally found during this period.
Through the winter the crawlers change in form from oval to pyriform. In the spring
the crawlers molt to the second instar, which apparently is short in duration. This
instar, like the adult female, is enclosed within a woolly, waxlike secretion, 1s
legless, and is yellow. Adult females are present in the spring and most of the
summer.

The first sign of an infestation is the appearance of isolated, minute, white
woolly dots on the bark, usually near the base of the tree. As the infestation
increases, the dots appear in the form of thin vertical lines and then as solid patches.
On heavily infested trees, the trunks (fig. 35) and lower sides of branches may be
completely whitened. Light infestations are not particularly injurious, but when
they increase to about 15 scales per square centimeter, the bark is killed and turns
brown. Depressions or pits in bark tissues around these wounds are frequently
numerous on young trees. The death and shrinkage of groups of cells within feeding
areas Cause ruptures in the bark. Death of the tree does not usually occur until 2 to 5
years after infestation. A fungus, Nectria coccinea Pers. ex Fr. var. faginata
Lohman, Watson and Ayres, gains entry through these ruptures. It penetrates the
cambium and sapwood, killing tissues and interfering with the conduction and
storage processes of the tree. Red fruiting bodies produced by the fungus become so
abundant in some infestations that large areas on the trunk turn red. This infection
leads to death of irregular-shaped areas of bark. Individual fungal lesions also
coalesce and girdle the trunk, leading to crown deterioration and finally to death of
the tree. The fungus is entirely dependent upon the scale for its incidence and
spread.

Figure 35.—Infestations of the beech scale,
Cryptococcus fagisuga, on the trunk of beech.

Enormous quantities of beech have been killed; the only way to prevent such
losses is through cutting and removal of infested trees (/080). This is true despite
the fact that some degree of natural control is provided by several predators,
including Chilocorus stigma (Say). Winter temperatures of — 38° C are highly
effective in the control of infestations exposed above the snow line (45).

Another species of Cryptococcus may be found in eastern forests. C. williamsi
Kosztarab & Hale is found on sugar or, rarely, red maple in Maine, New

102

Hampshire, New York, and Vermont. Probably it will eventually be found in most
areas where there are natural stands of sugar maple.

The European elm scale, Gossyparia spuria (Modeer), apparently was intro-
duced from Europe. In the United States it is reported from 33 States and probably
occurs wherever elms are grown. It feeds on native and introduced species of elm
and is found in natural habitats and ornamental plantings. Adult females are brown
or greenish brown soon after they molt, but as the adults age, the body turns gray. A
felted, waxy sack covers the bottom and lateral areas of the insect, leaving only the
top center of the scale exposed. The first instar is yellow, and the second instar is
reddish brown.

The European elm scale has one generation per year and has long-winged and
short-winged adult males. The second-instar males and females overwinter in
crevices on the bark of the host. In January, February, and March, the second-instar
males molt to the prepupa, pupa, and adult. Mature males are most abundant in
early spring. Molting of second-instar females to the adult stage is coincident with
seed development of the elm host. After molting, the adult females move to the
main branches, where mating takes place. When the elm seeds begin to fall, the
adult females begin to form the characteristic waxy sacks. Egg laying begins 2 or 3
weeks after the first elm leaves are fully formed and continues into late summer. An
individual female may lay as many as 400 eggs. The eggs normally hatch within an
hour of deposition in the ovisac. Crawlers usually move to the undersides of leaves
and feed near the primary veins. A small number of crawlers feed on the stems of
the host. About 6 weeks after hatching, the crawlers molt to the second instar. Most
of the second instars migrate from the leaves to the overwintering sites when the
leaves begin to turn yellow. The European elm scale is frequently injurious,
especially to young or recently transplanted elms. Symptoms include stunting of
new growth, dieback of branches, premature leaf drop, and in small trees, death.
Sooty mold grows on the large quantities of honeydew produced by this insect and
gives the infested tree and surrounding objects an unsightly black, sticky ap-
pearance. Chemical control is frequently effective. Several species of chalcidoid
wasps, lady beetles, and a green lacewing have been reported as natural enemies.

The azalea bark scale, Eriococcus azaleae Comstock, is apparently an Old
World introduction. In the United States, it is reported from 34 States and probably
occurs wherever azaleas are grown. Although normally found on azaleas, this
species 1s also reported on willow, poplar, maple, hackberry, fremontia, huckle-
berry, pieris, rhododendron, gooseberry, and blueberry. It occurs on native and
introduced hosts and is found in natural habitats and ornamental plantings. Adult
females are bright red and are covered with many crystalline rods. A tough, pear-
shaped ovisac is produced that encloses the adult female and the reddish-purple
eggs. The first and second instars are purple or red.

The azalea bark scale has one generation per year in the North and two in the
South. In northern areas the species overwinters as eggs or crawlers in the ovisac.
The crawlers escape through a small hole at the posterior end of the ovisac and
settle in branch cr leaf axils. Second instars appear in early summer, and adults are
prevalent in August. An individual female may lay as many as 250 eggs. In the
South, the overwintering stage is the settled crawler or second instar. Adult females
appear in early spring and lay their eggs in March or April. Adult females of the
second generation mature in late summer and lay their eggs, which hatch in the fall.
The scale causes dieback and gives heavily infested plants a leggy appearance.
Chemical control is possible. A chalcidoid wasp, Coccophagus immaculatus
Howard, often parasitizes this species.

103

Eriococcus quercus (Comstock), the oak eriococcin, is a native species re-
stricted to oak. It is reported from Alabama, the District of Columbia, Florida,
Georgia, Louisiana, Mississippi, New Jersey, Virginia, and several Western States.
This scale is found in natural habitats and ornamental plantings. Adult females are
reddish purple with a faint, yellow stripe down the center of the dorsum. The adult
is covered with many crystalline rods that are noticeably longer along the body
margin. The female produces a tough, pear-shaped ovisac which encloses the adult
and the red eggs.

The oak eriococcin has two generations per year in northern California.* Males
are common, and a single adult female may lay as many as 150 eggs. This scale
normally feeds on the new growth stems of oak hosts. The oak eriococcin occasion-
ally causes economic damage to ornamental plantings of oak. The chalcidoid wasp
parasite, Metaphycus eriococci (Timberlake), has been found on this scale species
in California.

Eriococcus gillettei Tinsley, the Gillette eriococcin, is native to the United
States. It is reported from the District of Columbia, Florida, Maryland, Virginia,
and several Western States, but it is undoubtedly present throughout the Southeast.
This eriococcin is restricted to species of Juniperus, including California juniper,
western juniper, and eastern redcedar. Adult females are yellow before forming an
ovisac, but they eventually turn brownish purple. The body margin has a fringe of
small crystalline rods. The adult female produces a tough, pear-shaped ovisac that
encloses the adult female and the eggs. The first instars are brown when they first
hatch and turn lemon-yellow after feeding. The second instars are yellow with a
brownish mottling.

The Gillette eriococcin has one generation per year in Maryland, and males are
common.” Eggs are the overwintering stage; they hatch in April or May. Second
instars appear in June, and adults become prevalent in late June and early July. Egg
laying begins in the fall and continues into early winter. The eggs are yellow when
first laid, but by February they turn brown. An individual female lays about 50
eggs. Feeding takes place in protected areas on the foliage. This species occasion-
ally builds up to economic proportions in ornamental plantings of juniper.

Family Lecanodiaspididae
Falsepit Scales

This family is a small, homogeneous group that is most abundant in the Southern
Hemisphere. Members of the family form a thick waxy test (or cover) that encloses
the body of the adult female and serves as an ovisac. The shape, color, and pattern
of the test is frequently characteristic of a particular species. There are normally
three instars in the female and five in the male. The family contains five species in
one genus, Lecanodiaspis, in the United States (6/5).

Lecanodiaspis prosopidis Maskell, the common falsepit scale, is native to North
America and occurs from Pennsylvania to Florida and west to Missouri in the
Eastern United States. It is found in ornamental and natural habitats on many trees
and shrubs including: sweetgum, holly, catalpa, boxwood, euonymus, mountain-
laurel, rhododendron, blueberry, buckeye, walnut, yellow-poplar, magnolia,
mulberry, ash, camellia, basswood, azalea, hackberry, persimmon, dogwood, and
elm. The test of the adult female is nearly circular, convex, yellowish white or
brownish red, and has a dorsomedial longitudinal ridge. Eggs are reddish brown.
The male test is smaller, more slender, and is bright yellow.

* Miller, D. R. [n.d.]. Unpublished data on file at USDA ARS Syst. Entomol. Lab., Beltsville, Md.
’ See footnote 8.

104

The common falsepit scale has one generation per year and overwinters as eggs
in the test (6/5). In Virginia, eggs hatch in mid-May, and adults appear in late July
or early August. Egg laying begins in mid-September. Feeding takes place on the
stems and twigs of the host. The common falsepit scale produces large pits and
welts on its host, giving the plant a distorted, unnatural appearance.

Family Cerococcidae
Cerococcids

This family is a small, homogeneous group that is similar in external appearance
to the falsepit scales. Adult female cerococcids form a thick, waxy test that encloses
the body of the female and serves as an ovisac. The ovisac of cerococcids is
generally rougher than the sac of falsepit scales and has a crawler exit hole that
protrudes posteriorly. This family contains eight species in the genus Cerococcus in
North America. Comprehensive studies of the genus have been published (52/7,
616;690;"7 11):

Cerococcus parrotti (Hunter), the Parrott scale, is apparently indigenous to
North America and is reported in Arkansas, Georgia, Louisiana, Maryland, New
Jersey, New York, Ohio, and Virginia in the East. Hosts include maple, bumelia,
pecan, eastern hophornbean, persimmon, sweetgum, hackberry, hawthorn,
sassafras, mahonia, basswood, elm, and buckeye. This species is found in natural
and ornamental habitats. The test of adult females is oval, convex, grayish brown,
brownish red, or yellowish brown, and has medial, mediolateral, and lateral
longitudinal rows of conspicuous projections. The male test is smooth and narrow.

Parrott scale has one generation per year and overwinters as eggs in the tests in
Virginia (6/6). First instars appear in early to mid-May, and adults are present in
late July. This species feeds on twigs, stems, and trunks of its host. It is apparently
not an economic species. Four species of hymenopterous parasites have been
reported as associated with this scale insect.

Cerococcus kalmiae (Cockerell) attacks mountain-laurel, azalea, camellia,
mountain-ash, and persimmon in Georgia, Virginia, Maryland, Massachusetts,
Ohio, and Pennsylvania.

Family Asterolecaniidae
Pit Scales

This family is a diverse aggregation of scale species that occur in all zoo-
geographic regions of the world. The most commonly encountered genus, As-
terolecanium, contains several eastern forest pests. The adult females of this genus
produce a thin, waxy test that covers the body. Many species produce a pit on the
host as they feed. The family contains 19 species in 4 genera in the United States
(1045).

Three species of Asterolecanium occur on oaks in eastern forests. Because these
species are similar in distribution patterns, field characteristics, and life histories,
they are treated together. The species are the golden oak scale, A. variolosum
(Ratzeburg), A. minus Lindinger, and A. quercicola (Bouché). These species may
be found together on the same tree wherever oaks occur in the United States. The
adult female is covered by a translucent greenish-yellow or brownish test that bears
a similarly colored or whitish, waxy fringe around its margin. The test is usually
slightly longer than it is wide and encloses the body of the female. Young adult
females are yellow with a dark-brown streak around the body. As the adults age,
they turn a uniform brown.

These three species of oak scales have one generation per year and overwinter as
adult females. Oviposition occurs in late spring and early summer. During egg

105

laying, the adult female shrivels into the anterior end of the test and eggs fill its
posterior portion. As the eggs hatch in late spring and early summer, crawlers
emerge through a small opening at the posterior end of the test, settle. and within 24
hours form a noticeable pit (/38). One species may be ovoviviparous (958). Second
instars appear in mid-July, and adults are first observed in August. Males are
unknown. These pit scales can be very destructive, especially to white oaks. In
heavy infestations they cause distortion, poor growth, and dieback of the twigs.
Leafing out in the spring may be delayed, and affected hosts may retain many of
their leaves in the winter. Several chalcidoid wasps are associated with these scale
insects.

Asterolecanium puteanum Russell, the holly pit scale, is native to the United
States. It is known from 10 Eastern States from New Jersey to Florida and west to
Pennsylvania and occurs on holly and bumelia. The holly pit scale may be found in
ornamental plantings and natural habitats. Adult females are covered by a translu-
cent, yellow-green test with a whitish-yellow or greenish waxy fringe around its
perimeter.

The life history of the holly pit scale has yet to be worked out, but it probably is
similar to that of the oak-infesting species. The holly pit scale is occasionally
destructive to hollies (293, 294). Heavily infested hosts may be stunted, have
reduced amounts of foliage, and their branches may be distorted with pitted,
roughened bark.

Other species of pit scales may be found occasionally in eastern forests. These
include oleander pit scale, A. pustulans (Cockerell), observed in Florida. Loui-
siana, and Texas on many hosts; A. arabidis (Signoret), found in the District of
Columbia, Massachusetts, New Jersey, New York, Ohio, and Pennsylvania on
many hosts including privet, ash, phlox, and weigela: A. bambusae (Boisduval), in
Alabama, Florida, Georgia, Louisiana, Mississippi, South Carolina, and Texas on
various bamboo hosts: A. miliaris miliaris (Boisduval) and A. miliaris robustum
Green, in Florida on bamboo.

Family Diaspididae
Armored Scales

The Diaspididae is the largest family of scale insects. Most armored scales form
a shieldlike cover that is unattached to the body. The cover is normally composed of
shed skins and wax produced by small glands on the body of the insect (//69). The
terminal segments of the adult female are fused into a strongly sclerotized
pygidium. Scale coverings vary from thick, oystershell-shaped structures to translu-
cent, circular structures. There are three instars in females and five in males. The
family in the United States contains about 297 species in 83 genera (904). The most
comprehensive studies of the family are by Ferris (405, 406, 407, 408).

The hemlock scale, Abgrallaspis ithacae (Ferris), is apparently native to North
America and probably is present in all eastern areas where hemlock occurs.
Distribution records have been compiled from Connecticut, Georgia, Indiana,
Maryland, New York, Ohio, Pennsylvania, Tennessee, and Virginia. This armored
scale is commonly found on hemlock, but is reported on fir and spruce: records
from pine are probably erroneous. The species occurs in natural and ornamental
habitats. The cover of the adult female is slightly convex, circular to oval, and is
gray to black with white margins; the shed skins are subcentral. The body of the
adult female is yellow-green, and the eggs are pale yellow. Petioles and undersides
of needles are the areas inhabited.

106

The hemlock scale in Maryland has two generations per year and overwinters as
second instars (//72). Adults of the winter generation first appear in the middle of
March, and eggs and first instars of the summer generation are present in June and
early July. This generation develops rapidly, and by early to mid-July adults are
present. Eggs and first instars of the winter generation are present from August to
early October, and overwintering second instars are first observed in early Sep-
tember. Adult males are winged. Feeding by this scale produces yellow spots on the
leaves; in heavy infestations, leaves will fall from the tree. An unidentified
hymenopterous parasite has been reported (//72).

Acutaspis morrisonorum Kosztarab, the round conifer scale, is native to North
America and is known from Massachusetts to Florida and west to Michigan. It is
found on many conifers including hemlock, spruce, fir, and juniper and infests
natural and ornamental vegetation. The scale cover of the adult female is oval, flat,
yellow-brown with a light margin; the shed skins are central. The body of the adult
female is orange-yellow.

The round conifer scale is reported overwintering as second instars in Ohio
(689). It apparently does not damage its hosts significantly. Another species of
Acutaspis that might be encountered in eastern forests 1s A. perseae (Comstock),
the redbay scale. This species is most numerous in Southern States where it may
become abundant enough to cause early leaf drop on several ornamentals. The adult
female of the redbay scale has a reddish-brown cover that is flat and circular.
Common hosts are persea and magnolia, but the species is polyphagous.

The oleander scale, Aspidiotus nerii Bouché, is probably introduced into North
America. In the Eastern States, the species is found out of doors in southern areas
and indoors elsewhere. The oleander scale is polyphagous, occurring on numerous
shade trees and woody ornamental shrubs, and can be collected in natural and
ornamental habitats. The cover of the adult female is flat or convex, is transparent
white to opaque tan, and has a central, reddish-brown shed skin. The body is
orange-yellow. Infestations normally occur on the leaves of the host but may be on
the stems, bark, or fruit also. The status of this species is confused; the presence of
biparental and uniparental “forms,” and differences in biology, have led to the
suggestion that there are two sibling species (285, 473).

The oleander scale reproduces almost continuously, and distinct generations are
therefore difficult to discern. Most workers state that the species has three or four
generations per year. Eggs may hatch soon after they are laid, or nymphs may be
deposited directly. Males may be present or absent. This species is sometimes a
serious pest of such diverse hosts as olive, magnolia, and oleander. The oleander
scale has a diverse array of natural enemies, including more than nine chalcidoid
wasps, three lady beetles, and several fungi.

Aspidiotus cryptomeriae Kuwana, the cryptomeria scale, is from Japan; it is
found in Connecticut, Indiana, Maryland, New York, and Pennsylvania in the
United States. This armored scale occurs on an array of coniferous hosts including
balsam fir, spruces, Douglas-fir, eastern hemiock, cryptomeria, white-cedar,
Cypress, yew, and pines and is found in natural and ornamental plantings. The cover
of the adult female is circular or oval and is flat, grayish brown, with yellow, central
shed skins. This species occurs on the leaves of its host.

The cryptomeria scale has two generations per year and, in Maryland, it over-
winters as second instars (//72). Adults of the overwintering population appear in
March and April, and eggs are laid in June. Adults of the summer generation are

107

first present in mid-July, and eggs are laid in late August and September. Overwin-
tering second instars appear in late September. Males are winged. This armored
scale damages the leaves of cryptomeria and other conifers in Japan (898). Natural
enemies in Japan include two species of chalcidoid wasps and two lady beetles.

The juniper scale, Carulaspis juniperi (Bouché), is very closely related to C.
minima (Targioni-Tozzetti), the minute cypress scale, and can be separated on the
basis of microscopic characters only. The major difference is that the juniper scale
has a macroduct between the median lobes and the minute cypress scale lacks this
duct. These species are apparently introduced into North America and occur
throughout most of the United States. They occur on a wide variety of coniferous
hosts including cryptomeria, cypress, incense-cedar, juniper, sequoia, and northern
white-cedar; juniper appears to be the most common host. Infestations are found in
natural and ornamental habitats. The scale cover of the adult female is circular,
convex, white, and has yellow, central shed skins. The body of the adult female and
the eggs are yellow. The scale cover of the male is white with three faint longitudi-
nal ridges and a yellow, terminal shed skin. The species occurs on the foliage of the
host.

The juniper scale and the minute cypress scale apparently have only one genera-
tion per year in northern areas (//62). Because first instars appear in warm southern
and western areas several months in advance of cooler northern areas, and because
only one generation occurs per year in northern Europe whereas two occur in
southern Europe, it is probable that there are two generations per year in warm areas
of the United States. However, only the minute cypress scale is known to have two
generations (33). Both species overwinter as gravid females and begin laying eggs
in early spring in warm areas and late spring in northern regions. Crawlers are
present in spring and early summer. Adult males are common. Heavily infested
plants fail to produce new growth, have dead areas, and eventually die. Natural
enemies include many chalcidoid wasps, several lady beetles, a nitidulid, and a
coniopterygid.

The elm scurfy scale, Chionaspis americana Johnson, is probably native to
North America and is known from almost every Eastern State. This armored scale
prefers elm, but is also found on privet, Prunus spp., hackberry, sycamore, and
basswood. Infestations occur in ornamental and natural vegetation. The scale cover
of the adult female is oystershell-shaped, flat, white or dirty white, and has yellow
or brown terminal shed skins. The body of the adult female without eggs is orange;
with eggs it is red or purple; the eggs are red or reddish orange. The scale cover of
the adult male is elongate, white, has three longitudinal ridges and a beige terminal
shed skin. The adult male is red and may have long wings or short wing stubs.

The elm scurfy scale has two generations per year and overwinters on the bark as
eggs (13/4). In Virginia, first instars appear in late April or early May and molt to
second instars in late May or early June. Adults are present in June and July, and
oviposition begins in July. First instars of the second generation are present in July,
and adults appear in late August and early September. Overwintering eggs are laid
in October and November. Females occur almost exclusively on the bark; males are
predominantly on the undersides of leaves. Damage to elms may include the death
of twigs, branches, or small trees. The feeding of second-instar males on leaves
causes small chlorotic spots. Natural enemies associated with the elm scurfy scale
include 11 chalcidoid wasps, a Leucopis fly, and a lady beetle.

The dogwood scale, C. corni Cooley, is native to North America and is reported
from Massachusetts south to Louisiana and west to Kansas in the Eastern United

108

States. The scale cover of the adult female is white, elongate, expanded posteriorly,
and has orange-yellow terminal shed skins. The scale cover of the adult male is
white with three longitudinal ridges and a yellow, terminal shed skin.

The dogwood scale overwinters as eggs on the bark of the host (605). Males have
been reported. This species is occasionally destructive to ornamental dogwoods in
the Midwest. Two chalcidoid wasps have been associated with this species.

The scurfy scale, C. furfura (Fitch), is probably native to North America and
occurs throughout most of the United States. It is a common pest of many rosaceous
plants, particularly apples; important forest hosts include hawthorn, mountain-ash,
and Prunus spp. The scurfy scale is found in natural and ornamental vegetation.
The scale cover of the adult female is white, oystershell-shaped with an expanded
posterior, and has yellowish-brown terminal shed skins. The body of the adult
female before egg formation is yellowish brown, but it turns red with the internal
development of the eggs, which are also red. The cover of the male is elongate,
white, with three longitudinal ridges. The body of the male is red. Males apparently
have well-developed wings.

The scurfy scale has two generations per year; in Virginia it overwinters as eggs
on the bark (57/). Overwintering eggs hatch in early spring, and first instars settle
on the bark. Adults appear in early June, and eggs are laid in June and July. First
instars of the second generation are found in mid-July, and adult development
begins in mid-August. Overwintering eggs are laid in September and October.
Males are apparently necessary for reproduction in at least some populations.
Scurfy scale may be an important pest in apple and pear orchards but is not usually
a pest in eastern forests. Natural enemies associated with this species include more
than five chalcidoid wasps, a mite, and several lady beetles.

The pine needle scale, C. pinifoliae (Fitch), is closely related to C. hetero-
phyllae Cooley, the pine scale, and can be separated on the basis of microscopic
characters only. The pine needle scale occurs throughout the United States, whereas
the pine scale is restricted to the East, primarily the Southeastern and Gulf States.
Hosts include nearly all needle-bearing conifers including spruce, fir, pine,
hemlock, and Douglas-fir (fig. 36). This scale occurs in natural and ornamental
vegetation. The scale cover of the adult female is oystershell-shaped, white, and
has transparent or light-yellow terminal shed skins. The body of the adult female
and the eggs are purplish red. The cover of the adult male is elongate, white, and
has three longitudinal ridges and a transparent terminal shed skin.

The pine needle scale has one or two generations per year. In the South to as far
north as southern New York and Minnesota, it has two generations per year,
whereas in northern areas, including Canada and in the Western States, there is only
a single generation. Overwintering normally occurs as eggs, but it has been found
that populations on Jeffrey pine from South Lake Tahoe, California, overwinter as
gravid adult females, whereas those on lodgepole pine in the same area overwinter
as eggs (754). In Oregon it was found that adult females laid eggs throughout the
winter into spring (//6/). In the Eastern States only biparental populations have
been reported, but in California and Oregon biparental and uniparental populations
are known. In central New York, the following life cycle is reported (930).
Overwintering eggs hatch in early May, second instars appear in late May, and
adults are present in mid-June. Egg laying begins in early July, with first instars
present from mid-July to early September. Adults appear in mid-August, and
Oviposition begins in early September. The pine needle scale can build to such large
populations that the foliage of infested trees may have a gray appearance. Heavy

109

" ; ee Fe 5

Courtesy Conn. Agric. Exp. Stn.

Figure 36.—Pine needle scale,
Chionaspis pinifoliae, on needles of
red pine.

infestations cause needles to turn yellow and may kill branches or whole trees.
Infestations tend to be heaviest on lower branches. Ornamental plantings in urban
areas are normally most severely damaged. Natural enemies include more than 10
chalcidoid wasps and 5 lady beetles.

Chionaspis nyssae Comstock, the sour-gum scale, is indigenous to North
America and is known from 18 Eastern States from New York to Florida and west to
Missouri. This species occurs almost entirely on black tupelo and water tupelo but
is occasionally found on common sweetleaf, hackberry, hawthorn, and oak. The
cover of the adult female is oystershell-shaped, slightly expanded posteriorly,
white, and has yellow, terminal exuviae. The body of the adult female is yellow but
turns pinkish red shortly before and during oviposition. Eggs are pink when laid but
turn purple before hatching. The cover of the adult male is elongate, white, and has
three longitudinal ridges and a yellow, terminal shed skin.

The sour-gum scale has two generations per year in Maryland and overwinters on
the bark as mated adult females (68/). This scale may be found during the growing
season on leaves and bark; females are most abundant on the bark and males on the
leaves. Egg laying on the bark begins in early April; a single female may lay up to
80 eggs. First instars appear in mid-April on bark and early May on leaves; this
stage 1s present throughout the growing season. Adults of the summer generation
are first observed in mid-June. Winged males are most abundant, but wingless
males are present in small numbers. Adults of the overwintering generation appear
in late August. In the fall eggs are laid by leaf-inhabiting females but normally are

110

not produced by females on the bark. Adult males are predominantly wingless.
Damage caused by this species 1s normally undetected; leaf areas around feeding
sites are chlorotic. Natural enemies of this armored scale have not been reported.

Chionaspis salicisnigrae (Walsh), the willow scurfy scale or black willow scale,
is probably native to North America. In the United States it is known from 32 States
including most of those in the East. Hosts include poplar and willow. The species is
found in natural and ornamental vegetation. The scale cover of the adult female is
white, oystershell-shaped, is broadest centrally, and has translucent or yellow
terminal shed skins. The body of the adult female and the eggs are purplish red. The
scale cover of the adult male is elongate, white, has three small longitudinal ridges,
and a yellow or brown terminal shed skin.

The willow scurfy scale has two generations per year in Colorado and overwin-
ters as eggs on the bark of the host (7/2). Eggs hatch in early spring, and adults are
first present in mid-June. First instars of the second generation appear in early to
mid-July, and adults are first seen in late August. Overwintering eggs are laid in
September and October. The willow scurfy scale can seriously injure willows under
certain circumstances. Natural enemies include two species of chalcidoid wasps
and a lady beetle. Other species of Chionaspis that might be encountered in eastern
forests are: C. acericola Hollinger on maple from Maryland, Missouri, North
Carolina, Ohio, Pennsylvania, and Texas; C. caryae Cooley on hickory and walnut
from 10 States including Connecticut to Florida and west to Missouri; C. kosztarabi
Takagi on ash and American hornbeam from Georgia, Maryland, North Carolina,
Ohio, Pennsylvania, Virginia, and New Jersey; C. /intneri Comstock on many hosts
including viburnum, birch, walnut, alder, and serviceberry from 11 States from
Maine to Florida and west to Indiana; C. parki Hollinger on sycamore from
Indiana, Louisiana, Missouri, Ohio, Texas, and Virginia.

The Florida red scale, Chrysomphalus aonidum (L.), is apparently introduced
into North America. In the United States it occurs out of doors in warm southern
areas and is acommon greenhouse pest in other areas. This species occurs primarily
in ornamental plantings but is occasionally found in natural ones. The Florida red
scale is truly polyphagous and is commonly found on holly, citrus, and palms. The
scale cover of the adult female is circular, convex, dark reddish-brown, and has
reddish central shed skins. The adult female and the eggs are yellow. The cover of
the male is oval and smaller than the female’s and has a subcentral shed skin.

The Florida red scale may have up to five or six generations per year (364),
depending on the climate. Eggs are laid and first instars settle on leaves, fruit, or
rarely on the green stems of the host. Males appear to be necessary for reproduc-
tion. A single female may lay up to 334 eggs. The Florida red scale produces
yellow areas on the leaves of the host. In heavy infestations entire leaves become
chlorotic and drop from the tree. Natural enemies are numerous, including chal-
cidoid wasps, mites, fungi, lady beetles, thrips, and green lacewings. Natural
control is normally effective in Florida.

Other species of Chrysomphalus that might be found in southeastern forests are
C. bifasciculatus Ferris, the bifasciculate scale, and the dictyospermum scale, C.
dictyospermi (Morgan). Both species are polyphagous.

Clavaspis ulmi (Johnson), the elm armored scale, is probably native to North
America and occurs in nine Eastern States from New Jersey to Florida and west to
Ohio. The preferred host is elm, but it is also found on maple, buckeye, catalpa,
and basswood. The adult female cover is circular, convex, yellowish white to gray,

111

and has submarginal shed skins. The body of the adult female is yellow. The male
cover is small and has submarginal shed skins.

The elm armored scale overwinters as second instars on the bark of the host, and
first instars appear in early May in Ohio (689). This species does not seriously
damage its host. The only naturai enemy known Is a parasitic wasp.

The Putnam scale, Diaspidiotus ancylus (Putnam), is probably native to North
America and occurs throughout most of the United States. The species is poly-
phagous but is commonly found on maple, basswood, and elm. The Putnam scale
may be found in natural and ornamental vegetation. The adult female cover is
circular, convex, gray or black, and has yellow or red subcentral shed skins. The
body of newly matured adult females is yellow: on older females the anterior
portion of the body is brown. The male cover is similar to that of the female but is
oval and has subterminal shed skins.

The Putnam scale has two generations per year in Maryland" and Illinois (//46)
and one in Iowa (926) and Ohio (605). The species apparently overwinters as adult
females on stems. In those areas where two generations occur, first instars appear
early in the year and settle on leaves or stems; those that are on leaves are different
morphologically from those on stems (//46). Adult females and winged males are
present in early July. Eggs and first instars appear in July, and adult females and
wingless males are present in late summer and fall. Putnam scale is reported to kill
twigs and branches on heavily infested trees. Natural enemies include more than
three chalcidoid wasps, a lady beetle, a gall midge, and two fungus species.

Diaspidiotus liquidambaris (Kotinsky), the sweetgum scale, is native to North
America and probably occurs in all areas where natural stands of sweetgum are
found. This species 1s reported almost entirely on sweetgum and is found on natural
and ornamental vegetation. This species forms a small gall-like structure on the -
leaves of its host. The cover of the adult female on the leaf has a large, yellow,
central shed skin with white wax around the margin. On stems the adult female
scale is white or gray with subcentral shed skins. The body of the adult female is
yellow. The male cover is oval, white, and has a yellow submarginal shed skin.

The sweetgum scale has two generations per year in Maryland and overwinters as
fertilized adult females on the bark (//72). In Maryland, eggs and first instars
appear in mid-May and are present until the end of June. Settling occurs almost
entirely on the leaves. Adults are first present in mid-June. First instars of the
second generation appear in early July and are present until early October. Settling
occurs on both the leaves and the stems. Adults of the second generation appear in
early to mid-September. Adult males are winged in the first generation and wingless
in the second. The sweetgum scale does not cause serious damage to its host. A
chalcidoid wasp has been reared from this scale.

Diaspidiotus osborni (Newell & Cockerell), the Osborn scale, is apparently
indigenous to North America and is reported in nearly every eastern State. This
scale is reported on several native tree genera but prefers oaks. The cover of the
adult female is circular, flat, gray, and has yellow, subcentral shed skins. The body
of the adult female and the eggs are yellow. The male cover is similar, but is smaller,
oval, and has a submarginal shed skin.

Osborn scale has two generations per year and overwinters as mated adult
females on stems in Maryland (//72). Crawlers are present in Maryland in late May
and June and again in August. Males of the first generation are winged; those of the

'” Stoetzel, M. B. (personal communication). USDA ARS Syst. Entomol. Lab., Beltsville, Md.

LZ

second are apterous. This commonly encountered species is not a pest. Natural
enemies include a chalcidoid wasp and a symbiotic fungus (25/).

Other species of Diaspidiotus that might be collected in eastern forests are D.
mccombi McKenzie, the McComb scale, in 10 States from Pennsylvania to Florida
and west to Louisiana on pine; and D. caryae Kosztarab, the hickory scale, in
Ohio and Georgia on hickory.

Fiorinia externa Ferris, the elongate hemlock scale or fiorinia hemlock scale,
has apparently been introduced from the Orient and is known in || Eastern States
from Connecticut to Georgia and west to Ohio. Hosts include spruce, fir, Douglas-
fir, yew, and hemlock, with the latter being the most common. It occurs in
ornamental and natural vegetation. The external appearance of species of Fiorinia is
unusual because the adult female 1s completely enclosed within the shed skin of the
second instar. The second instar’s shed skin is elongate, yellow to brown, and has
the wax and shed skin of the first instar attached at the terminal end. The body of the
adult female and the eggs are yellow. The cover of the male is elongate, white, has
three longitudinal ridges, and a terminal shed skin. The body of the adult male is
yellow.

The elongate hemlock scale is unusual in that it continues to reproduce and
develop throughout the year in some locations; consequently, all stages are present
at all times (279). In Connecticut there is one complete generation and a second
partial generation. Only about one-third of the population begins the second
generation; the bulk of the population overwinters as adult females or eggs (773).
Adult females produce small batches of eggs which are replaced after hatching.
Adult females may live up to a year. First instars settle on the undersides of needles
and produce two kinds of waxy secretions. The first is a filamentous secretion
produced by a pair of glands between the antennae. This secretion builds into a
mass of tangled strands and becomes so abundant that it gives the undersides of
infested needles a white appearance. The second secretion is produced posteriorly
and eventually forms the wax covering over the body. Winged males are common.
The elongate hemlock scale can cause serious damage to hemlocks in the North-
eastern United States. Severe infestations cause yellowing of the needles, needle
drop, and inhibited growth, giving the hemlock a thin, weak appearance. Natural
enemies include two chalcidoid wasps and a lady beetle.

The tea scale, F. theae Green, was introduced from the Orient and occurs in 14
Southeastern States from Maryland to Florida and west to Louisiana. It is reported
in several Northeastern States but probably is unable to winter out of doors. This
species is truly polyphagous but seems to prefer holly and camellia. It occurs
primarily in ornamental plantings. The second instar’s shed skin is dark brown or
black and has a dorsomedian ridge; the medial area is darkest. The first instar’s shed
skin is yellow and terminal. The body of the adult female and the eggs are yellow.
The adult male’s cover is elongate, white, has three longitudinal ridges, and a
yellow, terminal shed skin.

The tea scale reproduces for an extended period, causing most infestations to
have all stages present at one time. There are apparently many generations per year.
Eggs are laid and crawlers settle on the leaves of the host. Males are common.
Heavy infestations can severely damage the host. This species is generally con-
sidered the most serious scale pest of ornamental shrubs and trees in the Southeast
and is perhaps the most difficult to control. A white powdery host plant is a good
indicator of an infestation of tea scale. Natural enemies include at least one
chalcidoid wasp, a fungus, and several species of lady beetles.

113

Other Fiorinia species that might occur in eastern forests are F. fioriniae
(Targioni-Tozzetti), the fiorinia scale, from most Southeastern States on many
hosts including palms and persea and F. japonica Kuwana, the Japanese fiorinia
scale, from Virginia on hemlock and other conifers.

Hemiberlesia lataniae (Signoret), the latania scale, is so cosmopolitan that it is
difficult to pinpoint an indigenous area. In the Eastern United States it is wide-
spread, feeds on a wide variety of hosts, and occurs on ornamental and natural
vegetation. The cover of the adult female is circular or slightly oval, convex, white
to gray, and has a light-brown submarginal shed skin. The body of the adult female
and the eggs are yellow. The cover of the male is gray, oval, and has a brown
subterminal shed skin.

The latania scale has a variable number of generations per year, depending on the
area; there are two generations per year in Maryland (//72), three in Egypt (379),
and four in Israel (32). In Maryland this species overwinters as second instars, and
adults appear in middle to late April. Eggs are laid late in June, and adults are
present in late July and early August. Eggs of the overwintering generation first
appear in September, and overwintering second instars are found in late September.
The latania scale has sexual and parthenogenetic populations. This species may
build to very heavy populations and may cause dieback of twigs and branches of the
host. Natural enemies include many species of chalcidoid wasps and lady beetles;
green lacewings, fungi, and mites are also known to feed on the latania scale.
Natural enemies usually keep populations of latania scales at low levels.

The greedy scale, H. rapax (Comstock), is of unknown origin. It is reported
from 15 Eastern States from New York to Florida and west to Louisiana, although
in northern areas it may be unable to survive out of doors. The species occurs in
natural and ornamental vegetation and is polyphagous. The cover of the adult
female is circular, convex, gray, and has a reddish-brown submarginal shed skin.
The body of the adult female and the eggs are yellow. The male cover is oval, gray,
and has a reddish-brown submarginal shed skin.

The greedy scale apparently has a variable number of generations per year,
depending on the area, and feeds on all aerial parts of the host. Infestations usually
have all stages present most of the year. Populations may be uniparental or
biparental. Feeding may cause dieback, leaf drop, and unhealthy looking plants.
Natural enemies include several chalcidoid wasps and lady beetles.

The oystershell scale, Lepidosaphes ulmi (L.), was probably introduced from
Europe. It occurs in virtually every State in the United States and is polyphagous.
Commonly infested hosts are lilac, beech, birch, ash, maple, poplar, willow, elm,
boxwood, apple, pear, and Prunus spp. (fig. 37). This species may be found in
natural and ornamental vegetation. The cover of the adult female varies according
to host (466). In Maryland, the cover of populations on poplar is reddish brown
with two transverse yellow bands; on lilac and maple, the cover is dark brown. As
the covers age, they turn grayish, dark brown, or black. The adult female scale is
oystershell-shaped with brown or yellow terminal shed skins. The body of the adult
female is white with a brown pygidium; eggs are white. The cover of the male is
similar to the female cover but is smaller and has one shed skin. The body of the
adult male is light yellow.

The life history of the oystershell scale varies according to host (466). In
Maryland, populations on lilac and maple have two generations per year, those on
poplar and willow have one generation per year, and on boxwood there are one or
two generations; all forms overwinter as eggs, but those from poplar hatch later in

114

F-531245

Figure 37.—Oystershell scale, Lepidosaphes ulmi, on
bark of poplar.

the spring. Bivoltine forms in Maryland have crawlers in May and June and adults
in June and July. Second-generation crawlers appear in mid-July, and adults are first
present in September. Eggs are laid in October and November and subsequently
overwinter. Univoltine forms have crawlers present in late May, with adult females
first appearing in early July. In late July, females begin laying eggs, which
subsequently overwinter. In Maryland, males were found in small numbers on
boxwood, lilac, and maple (bivoltine) but were never found on poplar or maple
(univoltine). The oystershell scale can be very destructive. Branches of trees are
often obscured by scale covers. This scale frequently causes dieback and is often the
apparent cause of the death of ornamental trees. It has been reported that entire
forest stands of ash have been destroyed by this pest (45). Many natural enemies are
known, including parasitic wasps, lady beetles, and mites.

Other species of Lepidosaphes that might be collected in eastern forests are the
camellia scale, L. camelliae Hoke, on camellia, holly, and magnolia in 14 States
from New York to Florida and west to Louisiana; L. pallida (Maskell), Maskell
scale, on many conifers, especially juniper, from 13 States from New York to
Florida and west to Louisiana; and L. yanagicola Kuwana, primarily on euonymus
from 11 States from Massachusetts to Georgia and west to Ohio.

Lopholeucaspis japonica (Cockerell), the Japanese maple scale, is probably
native to the Old World. It occurs in 10 Eastern States along the Atlantic Coast from
New York to Georgia and infests many hosts including maple, ash, and basswood.
The species occurs in ornamental vegetation. Adult females occur within the shed
skin of second instars in a similar manner to species of Fiorinia. The second instar’s
shed skin is dark brown with a thin layer of grayish wax and a yellow, terminal,
first-instar shed skin 1s attached. The male cover is similar to that of the female but
is smaller.

The Japanese maple scale has one generation per year in Rhode Island and in
Tokyo, Japan, and two generations in Oita, Japan (898). The overwintering stage is
mated adult females in Tokyo and immatures in Oita. In Rhode Island first instars
are reported in late June and early July. In Tokyo, winged males occur in mid-
August. This species may kill branches on maples.'' The Japanese maple scale is
attacked by three parasitic wasps and two lady beetles.

'! Davidson, J. A. (personal communication). Dep. Entomol., Univ. Maryland, College Park.

115

The obscure scale, Melanaspis obscura (Comstock), is native to North America
and probably occurs in all Eastern States. The obscure scale is a serious pest of
ornamental oaks. particularly pin oak and pecans. Other hosts reported in the
literature are probably erroneous. The species occurs in natural and ornamental
vegetation. The cover of the adult female is circular, slightly convex, grayish to
black, and has shiny black subcentral shed skins. The body of the adult female is
yellow after molting but changes to dark pink. The eggs are light pink. The male
cover is similar to the female cover but is smaller, oval, with a subterminal shed
skin. The body of the adult male is light brown.

The obscure scale has one generation per year in Maryland (//70) and Louisiana
(40). In Maryland. obscure scale populations on the white oak group of host species
overwinter as crawlers, whereas on the red oak group of hosts, populations over-
winter as second-instar males and females (//7/). In the spring adult females on
red oaks first appear in late April, and egg deposition and hatching begin in late
June. Overwintering second instars are first observed in mid-August. White oak
populations molt to second instars in early May and are adults in mid-June. Egg
laying and crawler emergence begin in early August. In Louisiana, the life history
of populations on pecan is similar to that of red oak populations but is somewhat
advanced chronologically. This scale infests the twigs, branches. and trunk of its
host. Damage involves dieback of small-diameter branches, with the weakening of
heavily infested trees. Pecan nut production may be reduced by this scale. Natural
enemies include a large number of chalcidoid wasps, lady beetles, fungi, and mites.

The gloomy scale, /. tenebricosa (Comstock), is indigenous to North America
and is found in 19 Eastern States from New York to Florida and west to Missouri.
Hosts include a number of important forest trees: the most commonly infested hosts
are species of maples, particularly soft maples. The species is found in natural and
ornamental vegetation. The cover of the adult female is circular, convex, dark gray,
and has black subcentral shed skins. The body of the newly molted adult female is
light yellow but turns pink with age. The cover of the male is similar to that of the
female except that it is smaller and is oval with a subterminal shed skin.

The gloomy scale in Maryland has one generation per year and overwinters as
mated adult females (//68). Eggs and first instars first appear in late June and early
July, and adults first develop in mid-August. North Carolina populations were
found to be somewhat advanced chronologically (S54/). This scale is restricted to
the twigs, stems, and trunks of its host. Natural enemies include several species of
chalcidoid wasps and fungi.

Other species of Melanaspis that might be found in eastern forests are M.
nigropunctata (Cockerell) on many hosts including ash and dogwood from the
District of Columbia, Maryland. and Virginia: and M. smilacis (Comstock) on
many hosts especially smilax from eight States from Maryland to Florida west to
Louisiana.

The black pineleaf scale, Nuculaspis californica (Coleman), is apparently native
to North America and is found throughout most of the United States. Hosts include
Douglas-fir and most species of pine. This species is found in ornamental and
natural vegetation. The cover of the adult female is oval, convex, black with light-
gray margins, and has a yellow central area. Eggs are light yellow. The male scale
cover is similar to the female’s but is smaller and more elongate.

The black pineleaf scale has ane generation per year in northern areas and two in
southern areas. Warm weather during the fall or early winter may allow some
overwintering scales to complete their development and begin a new cycle (1/76).

116

In the northeastern part of Washington State this scale apparently overwinters as
second instars. Adults appear in late May and early June. Eggs and first instars are
present in mid-July to early August. In Spokane, Wash., eggs hatch immediately
after oviposition or hatch within the body of the adult female and are born as first
instars (37/). In southern California, eggs are present in April and mid-May, first
instars are present in May and June, and adults occur in June and July. Second-
generation first instars are present in early August. Irregular abundance of infested
trees in western forests has been hypothesized as a phenomenon of host-scale insect
coevolution (372). This species occurs on the needles of the host and prefers the
basal portion of the needles on the flat surface. Severe infestations of black pineleaf
scale cause the needles to turn yellow or red. Early needle drop and shorter needles
give heavily infested trees a weak, unsightly appearance. Heavy infestations may
kill branches or entire trees. Areas with large deposits of dust seem to be the most
susceptible to attack by the black pineleaf scale. Natural enemies include at least
four species of hymenopterous parasites and two species of lady beetles.

Nuculaspis tsugae (Marlatt), the shortneedle evergreen scale, is indigenous to
Japan and occurs in Connecticut, Maryland, New Jersey, and Rhode Island. Hosts
include fir, cedar, spruce, hemlock (775), and yew (904). The species in combina-
tion with elongate hemlock scale is a serious pest of hemlocks in both natural
habitats and ornamental plantings, and may kill entire trees. The cover of the adult
female is circular to oval, convex, dark gray, and has blackish, subcentral shed
skins. The male cover is similar but is more elongate and has a subterminal shed
skin.

The shortneedle evergreen scale has two generations each year in Connecticut.
Overwintering is in the second instar and crawlers are present from late May to
early July and early August to November (773). This scale prefers the undersides of
host needles and causes leaf chlorosis and premature needle drop. The main parasite
of the shortneedle evergreen scale in Connecticut is a small wasp, Aspidiotiphagus
citrinus (Craw).

A similar species, NV. pseudomeyeri (Kuwana), occurs on cedar, northern white-
cedar, cypress, juniper, and hemlock in New York and Pennsylvania.

Pinnaspis strachani (Cooley), the lesser snow scale, is apparently introduced
and occurs out of doors in Alabama, Florida, Georgia, Louisiana, and Mississipp1.
It is a common greenhouse pest. The species is polyphagous, occurring on more
than 200 hosts, and is normally found in ornamental plantings. The cover of the
adult female is oystershell-shaped, flat, white, and has yellow terminal shed skins.
The body of the adult female is reddish orange. The male cover is slender, white,
has three longitudinal ridges, and a yellow terminal shed skin.

The lesser snow scale reproduces continuously and has many generations per
year. Feeding occurs on all parts of the host, and males are common. Natural
enemies are numerous, including parasitic wasps, lady beetles, and fungi.

Another species of Pinnaspis that might be found in eastern forests is the fern
scale, P. aspidistrae (Signoret), observed on many hosts including palms and °
hollies, from nearly all Eastern States; in northern areas it is found in greenhouses.

Pseudaonidia paeoniae (Cockerell), the peony scale, has apparently been intro-
duced from the Orient and is reported in 14 Eastern States from New York to
Florida west to Louisiana. Common hosts are azalea, rhododendron, camellia, and
holly. This species is most commonly encountered in ornamental plantings but is
sometimes found in natural habitats in southern areas. The cover of the adult female
is circular or oval, convex, brown, and is normally covered by the outer layer of the

117

host; the shed skins are subcentral and are yellowish orange. The body of the adult
female and the eggs are purple. The male cover is smaller than the female cover and
has a submarginal shed skin.

The peony scale has one generation per year in Japan and overwinters as mated
adult females (S98). Overlapping generations are reported in the greenhouse
(1168). In southern areas there are probably many generations per year. Crawlers
are present in Virginia in late May (644). The peony scale occurs on the bark of its
host. This species is a serious pest of azaleas, rhododendrons, and camellias in
southern areas. It is reported to kill twigs and branches of its host. Natural enemies
include a parasitic wasp, several lady beetles, and a fungus.

The camphor scale, P. duplex (Cockerell), is apparently native to the Orient and
is found in Alabama, Florida, Georgia, Louisiana, and Mississippi in the United
States. In Louisiana, nearly 200 hosts are reported, including persimmon, pecan,
camellia, camphor-tree, oak, elm, and maple (266). This species is primarily a pest
of ornamentals but is found in natural habitats. The cover of the adult female is
circular, convex, brown, and has yellow submarginal shed skins. The body of the
adult female is white soon after molting but turns purple with age; eggs are light
purple. The male cover is elongate, flat, brown, and has a yellow terminal shed
skin.

The camphor scale has three generations per year in Louisiana and overwinters
primarily as mated adult females (266). First instars are present in Louisiana in late
March, mid-June, and mid-August. The camphor scale may feed on the leaves or
bark. Adult males are common and are necessary for reproduction in at least some
populations. Damage usually does not involve killing trees, although it has been
stated that camphor-trees may be killed within 6 months of attack (7/275). Normally,
lower branches are killed leaving a weakened host. Natural enemies include at least
nine parasitic wasps, two lady beetles, and Platoeceticus gloveri (Packard), the
orange basketworm.

The white peach scale, Pseudaulacaspis pentagona (Yargioni-Tozzetti) and P.
prunicola (Maskell), the white prunicola scale, have been confused until recently
(250). Morphological and biological characters are useful in separating the species.
The white peach scale has more perivulvar pores, more large macroducts, more
small macroducts on the metathorax and first abdominal segment, and has fewer
gland spines in the third space and these spines are usually bifurcate or trifurcate.
The eggs of a single specimen of the white peach scale may be white or salmon or
both (93), whereas those of the white prunicola scale are all salmon. Both species
have been introduced into North America from the Old World. The white peach
scale is primarily a southern species and is common in the Southeast as far north as
Maryland. The white prunicola scale is abundant in the Northeast but also is
reported from Alabama, Florida, Louisiana, and Mississippi. The white peach
scale occurs on a diverse range of hosts including 115 plant genera in Florida. In the
United States it commonly is taken on peach, mulberry, and persimmon. The white
prunicola scale has a more restricted host range including 21 genera of plants. In the
United States it is commonly found on Prunus spp., particularly Japanese flowering
cherry, and on privet and lilac. Both scales are similar in appearance; the cover of
the adult female is circular, convex, white, and has yellow subcentral shed skins.
The body of the adult female is yellow. The male cover is elongate, white, and has a
yellow terminal shed skin.

The white peach scale has four generations per year in Florida (644) and three in
Maryland (280). The white prunicola scale has two generations each year in

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Pennsylvania (//64) and three in Maryland (280). First instars and eggs of white
peach scale are present in early to mid-May, early to mid-July, and late August to
early September. The life history of white prunicola scale is advanced compared
with that of white peach scale. First instars and eggs are present in late April to
early May, late June to early July, and mid to late August. Males are necessary for
reproduction in at least some populations. These species are very serious pests of
peaches and ornamental cherries. On some hosts they may build to such heavy
infestations that entire branches are white. Natural enemies include parasitic wasps,
lady beetles, a thrips, a mite, and a green lacewing.

Another species of Pseudaulacaspis that might be found in eastern forests is P.
cockerelli (Cooley), which 1s polyphagous and occurs in Alabama, Florida,
Georgia, Louisiana, and South Carolina.

The Forbes scale, Quadraspidiotus forbesi (Johnson), 1s apparently native to
North America and occurs throughout the United States. It is polyphagous and is
often found on hickory, dogwood, apple, and Prunus spp. This species occurs in
ornamental and natural vegetation. The cover of the adult female is circular or oval,
flat or convex, gray, and has orange, subcentral shed skins. The body of the newly
molted adult female is reddish yellow but becomes darker with age. The male cover
is smaller and more slender than the female cover. The adult male may be reddish
brown or yellow, and has wings.

Forbes scale has two generations per year in North Carolina (/2/4) and Ohio
(643), and overwinters as mated adult females. In North Carolina adult females
give birth to first instars. First instars are present from June to early July and again
in August. Wingless males have been reported in mid-April in Ohio (689). The
species feeds on the stems and fruit of its host and can be a serious orchard pest in
cherries, apples, and peaches in Michigan, Ohio, Illinois, Indiana, and North
Carolina. Natural enemies include at least six hymenopterous parasites, several
lady beetles, a mite, and two fung!.

Quadraspidiotus gigas (Thiem & Gerneck), the willow scale, is apparently
introduced into North America and occurs in New York, Ohio, Pennsylvania,
Rhode Island, and Wisconsin in the Eastern States. Hosts are willow and poplar.
The willow scale may be found in ornamental or natural vegetation. The cover of
the adult female is circular, convex, gray, and has orangish-yellow subcentral shed
skins. The body of the adult female is yellow. The male cover is similar to that of
the adult female but is smaller, more elongate, and has a subterminal shed skin.

The willow scale has one generation per year in Czechoslovakia (72/) and
Germany (/066) and overwinters as second instar males and females. First instars
are sometimes present in the winter but are killed before spring. In spring the adults
are present and egg laying occurs from June to September. Males are necessary for
reproduction in at least some populations. First instars are present from June into
winter. Feeding takes place on the bark of the host; this species encrusts the bark in
severe infestations. Damage may include deformation of the bark, death of
branches, or even whole trees. Natural enemies include approximately 15 species of
parasitic wasps and 4 species of lady beetles.

The walnut scale, Q. juglansregiae (Comstock), is apparently native to North
America and has a general distribution in the United States. This species is
polyphagous and has been found on English walnut, holly, ash, elm, hickory,
basswood, maple, sweetgum, yellow-poplar, among others. This scale occurs in
natural and ornamental habitats. The cover of the adult female is circular, convex,
grayish, and has reddish-brown subcentral shed skins. The body of the adult female

119

is orange to reddish orange. The male cover is similar to that of the female but is
smaller, more slender, and has a submarginal shed skin.

The walnut scale has one generation per year and overwinters as second instar
males and females in Maryland (//68); overwintering second instars molt and
mature from mid to late April. Eggs and first instars are present from late June to
late September. Second instars begin to appear in early September. This scale may
kill entire trees in some situations, although infested trees usually have dead lower
branches and few leaves, giving them a weakened appearance (605). Natural
enemies include at least six species of parasitic wasps, two lady beetles, and a green
lacewing.

The European fruit scale, Q. ostreaeformis (Curtis), is apparently native to
Europe and is present in 11 Northeastern States from Maine to Pennsylvania west to
Ohio. Hosts include a variety of plants, but rosaceous trees, especially apple, most
commonly act as hosts. This species is found in natural and ornamental vegetation.
The cover of the adult female is circular, convex, dark gray, and has orange or
yellow subcentral shed skins. The male cover is similar to that of the female but is
smaller, more slender, and has a yellow subterminal shed skin.

The European fruit scale has one generation per year in Germany (/066),
Czechoslovakia (722), and New Zealand (/024). In Europe the species overwinters
as second instar males and females, but in New Zealand it overwinters as first or
second instars. Adults are present in spring or early summer. Eggs and crawlers
appear in late June or July. Males are essential in at least some populations. The
European fruit scale may damage fruit trees in some parts of Europe and the United
States. Natural enemies include at least five chalcidoid wasps, several lady beetles,
and a fungus.

The San Jose scale, Q. perniciosus (Comstock), is Oriental in origin and occurs
throughout the United States (fig. 38). It is polyphagous but is commonly collected
on pyracantha, pears, Prunus spp., and many other rosaceous hosts. The species 1s
found in natural and ornamental vegetation. The cover of the adult female is
circular, slightly convex, gray, and has yellowish subcentral shed skins. The body
of the adult female is yellow. The male cover is similar to the female cover but is
smaller, elongate oval, and has yellow subterminal exuviae.

- Courtesy Conn. Agric. Exp. Stn.
Figure 38.—San Jose scale,

Quadraspidiotus perniciosus. Note
large number of immature scales.

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The San Jose scale may have as many as five generations per year, depending on
the location. The overwintering stage in northern areas is predominantly settled first
instars; in warmer areas all stages may be present, although first instars and gravid
adult females seem to be most common. Parts of some generations are reported to
aestivate during the first instar. It is not surprising that chemical control of this scale
is difficult if one realizes that a single female may produce up to 400 eggs, and
some of the resultant crawlers may develop rapidly while others go through a
resting stage. After one or two generations there is little or no synchrony in first
instar emergence, and nearly all stages are present at any one time (468). First
instars are laid directly on the host, and males are common. This species is most
frequently found on stems, twigs, and fruit, but it also may be collected on leaves.
The San José scale is perhaps the most destructive scale insect in the United States.
If left unchecked it is capable of building up to very large populations and may kill
entire trees. It is a very destructive pest of shade, ornamental, and fruit trees.
Natural enemies include a large number of parasitic wasps, lady beetles, nitidulids,
and fungi.

Other species of Quadraspidiotus that might be found in eastern forests are Q.
socialis (Hoke) on oak from Georgia and Mississippi; Q. taxodii Ferris on bald-
cypress from the District of Columbia, Florida, Georgia, Louisiana, Maryland, and
Pennsylvania; and Q. tillandsiae Takagi & Tippins on Spanish moss from Georgia.

Quernaspis insularis Howell, the island oak scale, and Q. quercicola Tippins &
Beshear, the false oak scale, are native to North America. The latter is known only
from Georgia, whereas the former is recorded from Arkansas, Florida, Georgia,
and Texas. Until recently these species were considered to be the same as Q.
quercus (Comstock), the western oak scale (//55), which apparently occurs only
in the Western States. Because of the confusion, the precise distribution of each
species remains unclear. It 1s likely that the island oak scale and false oak scale will
be found in the Eastern States only, from Ohio to Florida and as far west as Texas.
These scales are reported only on species of oak and are found primarily in natural
situations. The cover of the adult female is oystershell-shaped, gray or white, and
has terminal shed skins. The male cover is elongate, white, with a terminal shed
skin. The male cover of the island oak scale has a conspicuous longitudinal ridge,
whereas the false oak scale lacks this ridge.

The life histories of the eastern species of Quernaspis are unknown, but the
western oak scale overwinters in California as adult females or second instars
(1028). In all three species the adult female occurs on the stems and twigs of the
host, whereas the second instar male covers are found on the leaves. A single
parasitic wasp 1s known on the western oak scale; natural enemies of the other two
species have not been reported.

The euonymus scale, Unaspis euonymi (Comstock), is probably native to the
Old World and will undoubtedly be found wherever euonymus is grown. Although
it is normally found on euonymus, it is also reported on 30 other genera of hosts
including bittersweet, holly, and pachysandra (485). This species normally is found
in ornamental plantings but occasionally is present in natural situations. The cover
of the adult female is oystershell-shaped, convex, dark brown, and has yellow or
brown terminal shed skins. The body of the adult female and the eggs are normally
yellow or yellow-red. The male cover is elongate, white, has three longitudinal
ridges, and a yellow terminal shed skin. The male is pale orange and winged.

The euonymus scale may have two or three generations per year, depending on
the locality. The primary overwintering stage is the mated adult female, although

121

immatures may also overwinter (/94). In Maryland, first instars are present from
late April or early May to late June and early or mid-July to October (485). In
Massachusetts first instars are present in early June and August. In Ohio they are
present from May through June and again in late July through August (689). This
species may be found on the stems, leaves, or fruit of its host. The euonymus scale
can build to such heavy populations that it may cover much of the surface area of a
particular host. Under such circumstances the plant will become white with male
covers (fig. 39). Upper surfaces of leaves become spotted with yellow areas where
scales are feeding on the undersurface. Heavily infested plants may lose many of
their leaves and become leggy in appearance. Natural enemies include at least seven
species of parasitic wasps, one species of lady beetle, a green lacewing, and two
mites. None of these natural enemies has been reported as an effective biological
control agent.

F-519580

Figure 39.—The euonymus scale, Unaspis euonymi, on
euonymus.

122

Velataspis dentata (Hoke), the dentate scale, !s apparently native to North
America and is found in Alabama, Florida, Georgia, Louisiana, and Mississippi in
the Eastern United States. It 1s found on a variety of hosts including bumelia,
hackberry, magnolia, willow, poplar, catalpa, maple, and others. The cover of the
adult female is unusually elongate, flat, white to light green, and has terminal shed
skins. The male cover is similar but is shorter and broader. The life history of this
species has not been studied. It is normally found on the leaves of its host.

Order Lepidoptera—Butterflies, Moths, Skippers

This is the second largest order of insects and one of the most important
economically. More than 5,000 species occur in the Eastern United States alone,
and many are serious pests of forest, shade, and ornamental trees. The adults differ
considerably in appearance from those in all other orders and are not difficult to
recognize. The wings and practically all other parts of the body are typically
covered by a layer of short, flattened hairs, or scales, which rub off like dust when
the insects are handled. The mouth parts, when present, are in the form of a long,
slender, flexible tube that is carried coiled up like a watch spring beneath the head.
The wings are usually very broad and subtriangular in form; the front pair is larger.

Moth, butterfly, and skipper adults usually differ in habits and appearance as
follows: (1) Moths usually fly at night and are frequently attracted to lights;
butterflies and skippers fly in the daytime. (2) Moths usually have the wings
wrapped around the body, folded rooflike on the abdomen, or spread horizontally
while at rest; butterflies usually fold their wings above the back in a vertical
position; skippers usually hold the front and hindwings at a different angie. (3)
Moth antennae are usually threadlike or featherlike; butterfly antennae are
threadlike and clubbed at the tip; the antennae of the skippers are usually recurved
or hooked.

Lepidopterous larvae are all very similar in structure and are known as cater-
pillars. They are usually cylindrical in shape and, besides the head, the body is
composed of 13 segments, 3 thoracic and 10 abdominal. Each thoracic segment
bears a pair of jointed legs, terminating in a single claw, whereas the abdominal
segments bear unjointed fleshy projections of the body called prolegs, typically one
pair each on segments 3 to 6 and 10. Occasionally, some or all of the prolegs are
missing. A distinctive feature of the prolegs of caterpillars is the presence of fine
hooks, known as crochets; these are usually in a circle at the apex, but also may
form bands or rows. Another important characteristic of the Lepidoptera is the
ability of the larvae to produce silk. Many larvae use this material in making
cocoons, and some use it for making shelters. The first instars of certain species
also frequently drop down from the crowns of trees in large numbers when
disturbed, hanging suspended at the ends of long strands of silk. Many of these are
often borne aloft by the wind and transported for considerable distances. Many
nearly full-grown larvae descend the tree in search of better food or pupation sites.

True silk moth larvae typically spin silken cocoons in which to pupate. Some
miscellaneous groups form tough silk or parchmentlike cocoons, which often
include debris. The pupae of many others are naked or are enclosed in slight
cocoons attached to leaves or other surfaces. Depending on the species, cocoons
may be found in the soil, in tunnels in wood, or in other larval habitats. The
caterpillars of butterflies usually do not make cocoons. Their pupae are naked and
are commonly known as chrysalids. They are often attached to leaves or twigs from

123

which they hang head down. Some, such as those of the families Papilionidae and
Pieridae, are girdled with a strand of silk and do not hang head down. Skippers
pupate in cocoons made of leaves fastened together with silk.

The order Lepidoptera contains numerous destructive forest and shade tree
insects. Several species such as the spruce budworm, the forest tent caterpillar, and
the gypsy moth often occur in outbreaks covering tens of thousands of hectares of
woodlands, and losses are very great. Large volumes of timber may be killed, and
larger volumes are lost through reduced growth of surviving trees. Many other
Species cause serious losses by boring into and destroying the buds and shoots of
seedlings and young trees in forest nurseries and plantations, or by mining the
tissues between the upper and lower surfaces of leaves. The attractiveness of shade
trees, parks, and other recreational areas is often reduced or destroyed by these
insects, fire danger is increased, and wildlife habitats are impaired.

The literature on these insects is vast, and the references given here for further
treatment are few, but selected because of their broad coverage (/40, 16/7, 318, 352,
374, 413, 433, 440, 447, 575, 586, 587, 614, 672, 785, 865, 1002, 1003, 1004,
1007, 1204).

Family Eriocraniidae
Eriocraniids

Larvae of the very small species Dyseriocrania auricyanea (Walsingham) mine
the leaves of oak, chestnut, and chinkapin in the Eastern States. The mines are often
blotchlike and may encompass up to one-fourth of the leaf area. Damage is
sometimes a matter of concern to owners of Asiatic chestnut orchards.

Family Hepialidae
Hepialid Moths

Hepialid moths sometimes attract attention late in the day when they are seen
flying swiftly in a zigzag manner, close to the ground. They are medium to large in
size, with rather long, stout abdomens and a wingspread of 25 to 100 mm. The
better known species are yellowish to brown or ashy gray with silvery-white spots
on the wings. The larvae are long-headed and nearly naked, have five pairs of
prolegs, and normally live as root borers. Sthenopis argenteomaculatus (Harris)
breeds in the base of the trunk and roots of alder in the Northeastern States: S. thule
(Strecker) breeds in the roots of willow.

Family Nepticulidae
Nepticulids

This family includes the smallest of the Lepidoptera, the adults of some species
having wingspreads of only 3 mm. Because of their minute size, their retiring
habits, and their irregular flight, they are seldom seen. The larvae of most species
are leafminers: a few construct mines in the bark of their hosts. Many of the more
common deciduous trees in the Eastern United States serve as hosts for one or more
species. The larvae are slightly flattened, and their heads are rather deeply retracted
into the prothorax. When full grown, the majority vacate their mines, drop to the
ground, and spin dense, flattened cocoons in the duff (/47).

The genus Neprticula contains many leafmining species. Eggs are deposited on
either surface of a leaf, usually along the side of a vein. The larvae of certain species
mine only one side of a leaf; some mine both sides; and some mine different sides at
different periods of larval life. The mines may be linear and gradually widen as they
are extended, or, at some point, they may enlarge suddenly into a blotch. Winter is
spent in the larval stage. There are one to four generations per year, depending on
the species.

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The genus Ectodemia contains a number of species, the larvae of which either
form galls on leaf petioles or twigs or mine the bark of various trees. Members of
the genus have only one generation per year. E. populella Busck, the poplar
petiole gall moth, forms pea-size globular galls on the petioles of poplar leaves. E.
heinrichi Busck larvae excavate flattened, oval, spiral mines in the bark of young
branches of pin oak.

The hard maple budminer, Obrussa ochrefasciella (Chambers), bores into and
destroys the buds of hard maple. The larvae spend the winter in axillary buds and
migrate to the main buds in the spring (70/). Heavy outbreaks of O. sericopeza
(Zeller), an introduced species that mines the seed pods and leaf petioles of Norway
maple, have been recorded in Maine. Damaged leaves and seed pods drop pre-
maturely. There are two and perhaps three generations per year.

Family Incurvariidae
Incurvariids

The maple leafcutter, Paraclemensia acerifoliella (Fitch), is the only member of
this family of economic importance in eastern forests. It occurs in southeastern
Canada and in the Northeastern States south to Virginia and west to Illinois. Its
favored host is sugar maple, but the larvae also feed on the leaves of red maple,
beech, birch, elm, and hophornbeam. The adult has long, narrow, pointed wings
and a wingspread of 8.5 to 13 mm. The forewings are steel blue and fringed with
black; the hindwings are pale smoky brown, translucent, and bordered with a pale-
brown fringe of long scales. There is a dense tuft of bright orange-yellow scales on
the top of the head, the thorax is steel blue, the legs whitish, and the abdomen dark
brown. First-stage larvae are flattened, about 1.5 mm long, and they taper from the
front to the rear. Full-grown larvae are slender, flattened, and usually dull white
with amber-brown heads and a broad longitudinal stripe. They are about 6 mm
long. Pupae are light yellowish brown, about 5 to 6 mm long. Abdominal segments
two to eight bear transverse rows of short, stiff, backward-pointed spines on the
back.

In Canada, adults appear in the spring about the time the leaves open, and deposit
their eggs singly in pockets on the undersides of the leaves. When the larvae hatch,
they bore into the leaf tissues and feed as miners for about 10 days to 2 weeks.
Then, each larva cuts a round disk out of the leaf and makes an oval, movable case
in which it resides as a casebearer. As it grows, it cuts out larger oval pieces and
attaches them to its case. In order to feed, it attaches the case to a leaf and reaches
out from it in a circle as far as it can. The uneaten center of this circle often drops
out, leaving a hole up to 12 mm in diameter in the leaf. When the larvae become
full grown, they drop to the ground to pupate. Winter is spent in the pupal stage,
and there is one generation per year (/039).

The maple leafcutter is often a pest in sugar maple stands, especially in sugar
maple orchards. The trees may be severely defoliated, often for several years in

succession.
Family Heliozelidae
Shield Bearers

Shield bearer moths are rather small and have lanceolate wings. The larvae are
strongly flattened and spend most of their lives mining the leaves of their hosts.
The tupelo leafminer, Antispila nysaefoliella Clemens, feeds on tupelo through-
out the eastern part of the United States. The adult is dark brown and has a
wingspread of 7 to 8 mm. The pale-green larvae feed within leaf tissues, forming
blotchlike mines. A full-grown larva spins a cocoon within the mine and then cuts

125

through the upper and lower layers of the leaf around the cocoon, forming a case.
The case, which encloses the larva and cocoon, drops to the ground. Once there,
the larva fastens the case to some object by means of silken threads and then
pupates. Heavily infested trees may turn completely brown by midsummer.

The resplendent shield bearer, Coptodisca splendoriferella (Clemens), also
occurs in the Eastern United States. Larvae mine the leaves of apple, cherry, and
related hosts. They pupate in cases attached to the limbs and trunks of their hosts.
Family Psychidae
Bagworm Moths

Members of this family have the interesting habit of spending the entire larval
stage within silken bags. The bag is usually strong, tough, and camouflaged by an
outer layer containing bits of twigs and leaves. Newly hatched larvae begin to spin
bags about their bodies as soon as they start to feed and they continue to enlarge the
bags as they grow. An opening is maintained at the top of the bag, through which
the head and several segments of the body protrude when the larva is moving,
feeding, or enlarging its case. There is also a smaller opening in the bottom of the
bag, through which excrement drops out. About 24 species have been recorded
from the Eastern United States (282). The species occurring in Texas are discussed
(646).

Oiketicus abbotii Grote occurs over much of the Atlantic and Gulf Coastal Plains
from North Carolina to Texas. It feeds on many species of trees such as bald-
cypress, live oak, bayberry, sycamore, elm, hackberry, sweetgum, and willow. It is
noted for the rather large bag, about 70 mm long, which the larva constructs. Small
twigs used in its construction are placed in a circular pattern around it. Psyche casta
(Pallas) feeds on lichens, mosses, and the beech scale in Massachusetts. The larvae
occasionally climb up on the sides of houses in such large numbers that they are a |
nuisance. Basicladus celibatus (Jones) frequently attaches its bags to the lower
trunks of oaks and pines in coastal areas from North Carolina to Florida. The larvae
are general feeders on low vegetation and may feed on trees.

The bagworm, Thyridopteryx ephemeraeformis (Haworth), is widely distributed
in the Eastern United States and attacks a wide variety of trees. Northern white-
cedar and redcedar appear to be preferred, but many other conifers and hardwoods
such as pine, spruce, black locust, sycamore, willow, maple, elm, basswood,
poplar, oak, baldcypress, and persimmon are also attacked. The male moth is sooty
black, densely hairy, and has a wingspread of about 25 mm. Females are wingless,
have no functional legs, eyes, or antennae, and are almost maggotlike in ap-
pearance. The body is soft, yellowish white, and practically naked except for a
circle of woolly hairs at the posterior end of the abdomen (6/0). Full-grown larvae
are dark brown and about 18 to 25 mm long. The head and thoracic plates are
yellowish and spotted with black.

Male bagworm moths emerge in the fall, fly to the females, and mate. The
female remains in her larval bag and deposits her eggs in the pupal case in the mass
of scales she shed. Winter is spent in the bag in the egg stage and the eggs hatch
during the following May or June. The larvae feed on the surface of leaves at first;
later entire leaves are consumed. Mature larvae attach their bags to twigs (fig. 40)
with silk and pupate in them. There is one generation per year.

The bagworm is most important as a pest on shade trees and ornamental shrubs
growing in yards and hedges, along the streets of cities and towns, and in parks and
other recreational areas. Large numbers of northern white-cedar and other conifers
are lost each year as a result of complete defoliation. Many others only partly

126

defoliated are weakened and rendered unsightly. Damage to forest trees is usually
not very severe (699): however, heavy infestations do occur occasionally in stands
of Atlantic white-cedar and black locust, especially in the Deep South.

Low winter temperatures, bird predation, and parasitism by the ichneumonid,
Itoplectis conquisitor (Say), are often particularly effective in bagworm control.
Handpicking and burning of overwintering bags is also helpful in control, es-
pecially on small trees and ornamentals (/349).

Cryptothelea gloverii (Packard) occurs from South Carolina to Florida and
westward along the Gulf Coast. Its hosts are recorded as two species of scale
insects, and persimmon, oak, hickory, acacia, hawthorn, and sour orange. The
male moth is dark or dark red and has a wingspread of 14 to 18 mm. Females are 9
to 10 mm long. Their cases may be covered with fragments of scale insects, bark,
fruit rinds, or leaves.

A, F-508522
B, Courtesy H. G. Schabel, Univ. Wis.
Figure 40.—Larval cases of the bagworm, Thyridopteryx
ephemeraeformis: A, cases on an infested tree; B,
closeup of cases.

127

Family Lyonetiidae
Lyonetiids

This family contains numerous species of tiny moths, the majority of which
belong to the genus Bucculatrix. At least 18 species are known to feed on trees in
eastern America (/48).

Adults of the genus Bucculatrix have the vertex of the head rough or tufted. and
the face smooth which extends into a point below the eyes. The wings are lanceolate
and broadly fringed with markings of brown, black, or silver. Larvae are cylindrical
and greenish and have well-developed legs and prolegs. Pupation occurs within
longitudinally ridged or ribbed silken cocoons.

Bucculatrix pomifoliella Clemens, the apple bucculatrix. is widely distributed
in southern Canada and the Eastern United States. Its hosts are apple. black cherry.
serviceberry. and hawthorn. Full-grown larvae are dark yellowish-green tinged with
red. have brown heads. and are about 6 mm long. Winter is spent in cocoons spun
on the lower surfaces of twigs. foliage. or fruit. The color of the cocoons differs
according to the larval host—white on apple. pale tannish-ochreous on service-
berry. reddish-brown on black cherry, and dark brown on hawthorn. There is one
generation per year in the North and two in the South.

The birch skeletonizer. 5. canadensisella Chambers. is a common species in
southern Canada and throughout the birch-growing regions of the Eastern States
from Maine to North Carolina and Minnesota. Paper birch appears to be the favored
host. but several other birches and possibly alder are also attacked. The adult has a
brown and white body and a wingspread of about 9 mm. The forewings are marked
with diagonal white bars and the hindwings have broad fringes of scales. Full-
grown larvae are yellowish green, with hairs projecting from white tubercles. and
are about 6 mm long.

Adults are active from late June to late July and lay their eggs singly here and
there over the leaves. Hatching occurs in about 2 weeks and the young larvae enter
the leaves to feed, forming serpentine mines. From 3 to 4 weeks later they emerge
through the lower surface and spin webs within which they molt. After molting,
they leave the webs and feed externally as skeletonizers until they reach maturity.
Full-grown larvae drop to the ground and spin short. brown cocoons in which they
spend the winter on the undersides of fallen leaves or other debris. There is one
generation per year.

Outbreaks occur at frequent intervals. often over large areas, and last for 2 or 3
years. Defoliation may be severe but it seldom causes much tree mortality. The
defoliated trees may be so seriously weakened. however. that they are attacked and
killed by the bronze birch borer.

The oak skeletonizer, B. ainsliella Murtfeldt. occurs from southern Canada and
the Lake States to North Carolina and Mississippi. Its hosts are various species of
oak and chestnut. The adult has a wingspread of about 8 mm. The forewing is
largely blackish. with paler areas outlining an oval, blacker patch on the inner
margin. Full-grown larvae are yellowish green and about 5 mm long.

The winter is spent in the pupal stage in cocoons. The cocoons are white. about 3
mm long. and ridged longitudinally. Adults are active in April and May and again in
July and August. Eggs are laid on the undersides of leaves. First-stage larvae enter
the leaves to feed. forming serpentine or blotch mines. Older larvae feed externally
on the lower surface. often completely skeletonizing the leaves (fig. 41) (475). Two
generations per year have been recorded in Michigan and Massachusetts. Outbreaks
occasionally occur over large areas.

128

F-519521

Figure 41.—Red oak leaves skeletonized by feeding of
larvae of the oak skeletonizer, Bucculatrix ainsliella.

Other widely distributed species of Bucculatrix and their hosts are as follows: B.
packardella Chambers—various oaks and occasionally beech; B. quinquenotella
Chambers—oaks; B. luteella Chambers—white oak; B. recognita Braun—various

white oaks, especially bur; and B. coronatella Clemens—sweet birch.
Family Gracillariidae
Leafblotch Miners

The family Gracillariidae, the largest of the leafmining families of Lepidoptera,
is represented in North America by more than 200 species (380, 9/0). This family
is currently undergoing revision. The adults are tiny and beautifully arrayed in
shining scales and plumes, and their more or less lanceolate wings are overlaid with
glistening scales of silver or burnished gold. While an adult is at rest, the front part
of the body is raised and the wingtips touch the surface on which it sits. Early
instars are very flat and usually feed first within mines in the leaves. Later, some
feed mostly on the leaf tissues from within tentiform mines or they skeletonize the
leaf from shelters made by folding over parts of leaves.

The full-grown larvae of most species spin silken cocoons, usually within
feeding mines or shelters, in which to pupate. Winter 1s spent as larvae, pupae, or
adults, depending on the species.

The solitary oak leafminer, Cameraria hamadryadella (Clemens), occurs
throughout much of the Eastern United States and southeastern Canada. Its hosts
are various species of oak, especially those in the white oak group. The adult has a
wingspread of about 6 mm. The forewings are pale with bronze patches, and the
hindwings are silvery with broad fringes of hairs. Young larvae are tiny, flat, and
taper toward the rear. Full-grown larvae are cylindrical and about 5 mm long. The
larvae feed singly, forming irregular blotchlike mines just below the upper leaf
surface. A single leaf may contain several contiguous mines (fig. 42). Winter is
spent in the larval stage in leaves on the ground and there are several generations per
year. Injury to forest trees is of minor importance, but the beauty of shade trees may
be seriously reduced.

The gregarious oak leafminer, C. cincinnatiella (Chambers), occurs through-
out the same range as that of the solitary oak leafminer, and it feeds on the same

129

F-504082

Figure 42.—QOak leaves with the blotchy mines
characteristic of attack by the solitary oak leafminer,
Cameraria hamaadryadella.

hosts. The adults and larvae of the two species are similar in appearance, but the
feeding habits of the larvae differ. Larvae of this species are gregarious, forming
large brownish-yellow mines, several of which may be found on a single leaf.
Winter is spent in the pupal stage in leaves on the ground, and there are two or more ~
generations per year. Heavy infestations have been reported in oak stands in the
Central States, causing severe browning and premature dropping of infested leaves
over large areas.

Many other species of leafminers attack a wide variety of deciduous trees in
eastern America. C. hamameliella (Busck) mines the upper surface of witch-hazel
leaves. It probably occurs wherever witch-hazel grows. The mines are circular or
somewhat irregular. C. bethunella (Chambers) mines the leaves of oak in Delaware.
C. aceriella (Clemens), the maple leafminer, produces large white mines in the
upper sides of red and sugar maple leaves. C. corylisella (Chambers) mines the
leaves of hazel in Maine. Phyllonorycter salicifoliella (Chambers) occurs through-
out most of the United States and also in Canada. Its preferred host appears to be
quaking aspen (8/7). P. crataegella (Clemens) mines the leaves of apple,
hawthorn, cherry, plum, and quince. Heavily infested trees become ragged and
scorched in appearance. P. /ucetiella (Clemens), the basswood blotchminer. mines
the leaves of basswood. The mine Is nearly square. P. trinotella (Braun) mines the
undersides of the leaves of red and Norway maples. P. robiniella (Clemens)
produces digitate mines in the upper surfaces of black locust leaves. The aspen
blotchminer, P. tremuloidiella (Braun), constructs oval mines in the leaves of
quaking aspen. It is occasionally quite abundant in the Lake States and Maine.
Chrysaster ostensachenella (Fitch) forms blotch mines in black locust leaves.

Marmara fasciella (Chambers) is widely distributed, probably occurring
throughout the range of its host. eastern white pine. Eggs are laid on the bark of
branches of the host. Young larvae bore through the bark and construct linear mines
in the inner bark. Winter is spent as a larva in the mine: pupation occurs during late

130

spring, and adults begin to appear in late May. Several other species of Marmara
have been recorded mining the inner bark of twigs of balsam fir, oak, willow, and
other trees (433).

Phyllocnistis populiella Chambers, the aspen leafminer, is widely distributed in
southern Canada and the Northern States. It feeds on various species of poplar,
especially quaking aspen. In the Eastern States, it has been recorded as far south as
West Virginia. Eggs are laid near the tips of young leaves in the spring. The larvae
form tortuous mines in the leaves (fig. 43), mostly on one side of the midrib (237).
Noticeable infestations have been observed in Maine.

-F-506745

Figure 43.—Small track mines of Phyllocnistis
populiella, the aspen leafminer.

Phyllocnistis liriodendronella Clemens larvae mine the undersides of young
leaves of yellow-poplar, and P. magnoliella Forbes mine the undersides of young
leaves of magnolia. The mines originate near the outer margins of the leaves and
continue tortuously until large portions of the upper surfaces are detached. The
damaged areas have a bluish cast. P. liquidambarisella Chambers mines the leaves
of sweetgum.

The larvae of Parectopa robiniella Clemens feed in the leaves of black locust,
forming triangular, blisterlike mines.

Parornix geminatella (Packard), the unspotted leafminer, occurs throughout the
Northern States, west and south to Colorado, Virginia, and Arkansas. Its hosts are
apple, quince, pear, sour cherry, pin cherry, plum, and hawthorn. Eggs are laid on
the undersides of the leaves, and the larvae devour all of the tissues between the
upper and lower surfaces of the leaves. Four generations per year have been
recorded in Virginia.

The lilac leafminer, Gracillaria syringella (F.), an introduced species, occurs in
many of the Northeastern States and southeastern Canada. Its hosts are listed as
lilac, black ash, privet, and euonymus. The adult has a dark-brown body and a
wingspread of about 10 mm. The forewings are brownish except for six irregular
transverse patches of yellow, and the hindwings are grayish brown. Full-grown
larvae are pale yellowish, translucent, and about 8 mm long. Early instars feed
gregariously inside the leaves and produce blotchlike mines. Later, they abandon
their mines and roll or web together several leaves on which they feed from within
this shelter. There are two generations per year. Injured portions of infested leaves
dry up and become unsightly. Raking and destruction of fallen leaves is a good
control practice.

131

The boxelder leafroller, Caloptilia negundella (Chambers), occurs throughout
eastern America west to the Prairie Provinces of Canada and feeds on boxelder.
Early instars feed within mines in the leaves. A partly grown larva vacates its mine
and moves to the tip of a lobe of the leaf, which it then turns over and attaches to the
lower surface with silk, thus forming a shelter. It then feeds from within the shelter.
There appears to be at least two generations per year as far north as Canada.
Heavily infested trees may be seriously damaged.

Caloptilia sassafrasella Chambers, the sassafras leafminer, feeds on the leaves
of sassafras. The larvae feed first within mines. Later, they vacate their mines,
move to nearby leaves, bend their tips over, and feed from inside the folds. Mature
larvae vacate these folds and form split, capsulelike cases on the undersides of other
leaves in which to pupate. The azalea leafminer, C. azaleella Brants, larva bends
the tips of azalea leaves downward and feeds from inside the folds. Injured leaves
wilt and die. C. packardella (Chambers) larvae mine the undersides of sugar and
Norway maple leaves in the early instars. Older larvae feed on the surface of the
leaves. Other eastern species of the genus and their hosts include C. bimaculatella
(Ely) on maple, C. stigmatella (F.) on poplar and willow, C. quercinigrella (Ely) on
oak, and C. pulchella (Chambers) on yellow birch and alder.

Family Oecophoridae
Oecophorids

This family contains a large number of moderately small moths, only a few of
which are ever very injurious to trees in the Eastern United States. The larvae of
most species roll, tie, or web together the leaves on which they feed. A revision of
the family is available (2/5).

Machimia tentoriferella Clemens, occurs rather commonly in the Northeastern
States and southeastern Canada. Its hosts are various hardwoods such as birch, ash,
maple, oak, honeylocust, mountain-ash, hickory, elm, and pin cherry. The adult is
light ochreous and has a wingspread of 18 to 20 mm. The forewing is dusted with
black and marked with two black discal dots, a spot of black in the fold, a broken
postmedial line parallel to the outer margin, and a series of black terminal dots. The
larva is green and has a large head and tapering body. It lives and feeds inside a
folded leaf.

Psilocorsis cryptolechiella (Chambers) occurs in eastern Canada and the North-
eastern States. The larva feeds principally as a leaftier on American beech. Other
hosts include the birches, maple, and red oak. The biology in Nova Scotia is
discussed (38/).

Full-grown larvae have reddish-brown heads, pale-green bodies, and are about 12
mm long. Eggs are laid singly on the undersides of leaves. During the first three
instars, the larvae feed together and skeletonize the leaves near the veins. Older
larvae usually feed singly from within the silken tubes they spin. Full-grown larvae
drop to the ground and crawl under fallen leaves, where they pupate and overwinter.
P. reflexella Clemens and P. quercicella Clemens occur on oaks in the Eastern
States. P. reflexella has been observed feeding on quaking aspen in eastern Canada.
It probably occurs in the Northeastern States also.

Other tree-infesting species of oecophorids and their hosts in the Eastern United
States are Agonopterix pteleae Barnes & Busck on the common hoptree, A.
robiniella (Packard) on black locust, A. argillacea (Walsingham) on willow, A.
nigrinotella (Busck) on prickly-ash, Bibarrambla allenella (Walsingham) on white
birch and oaks, and Nites betulella (Busck) on white birch and hophornbeam.

132

Family Blastobasidae
Blastobasid Moths

Blastobasid moths are small with long antennae. Long scales on the head often
cover the face and base of the antennae. The scape of the antenna is broad and
armed with a fringe of bristles, and the hindwings are narrower than the forewings.
The larvae feed in the cones, nuts, and seeds of various trees; as scavengers in
hollowed-out nuts or insect galls; or as predators on scale insects.

Valentinia glandulella (Riley), the acorn moth, feeds in acorns, hickory nuts,
and chestnuts in southern Ontario and southward throughout the oak region of the
Eastern and Central States. Larvae are grayish white or yellowish, with blackish
marks on top.

Holcocera lepidophaga Clarke larvae feed mainly in male flower buds and
flowers or among the basal scale leaves of young cones and vegetative buds of slash
and longleaf pines in Florida. It has also been recorded from Massachusetts. The
adult has a wingspread of 11 to 17 mm. The head is ochreous-white, with slight
infuscation posteriorly; the antennae are ochreous-white with narrow, dark annula-
tions; and the forewings are ochreous-buff. The larvae of Zenodochium coccivorella
(Chambers) are internal parasites of female scale insects of the genus Kermes in
Florida.

Family Coleophoridae
Casebearer Moths

About 145 species of casebearer moths occur in North America. The adults are
plain, little moths with markings limited to dustings of lighter or darker colored
scales. The wings are narrow and have a wingspread of 12 mm or less. The larvae of
all species feed within leaf mines in the first instar. At the end of this instar, each
larva constructs a portable case in which it lives thereafter. The larvae of certain
species continue to feed as miners after the first instar. Without leaving the case, the
larva mines in a circle or from each side of the point of entrance into the leaf. The
larvae of other species feed externally in the later instars. Their cases are con-
structed from parts of the mined leaves and are lined with silk. They are enlarged by
the larvae as needed, thus providing shelter at all times. The winter is spent as a
partly grown larva in a case which is usually fastened securely to a twig or branch
with silk.

Coleophora tiliaefoliella Clemens constructs a black case on basswood. C.
atromarginata Braun occurs on southern red and swamp white oaks. Its case is
black with white markings on the underside. C. querciella Clemens occurs on
white and swamp chestnut oaks. Its case is grayish with a black patch on the top
near the rear end. C. a/niella Heinrich has been observed on beech in New York.

The pecan cigar casebearer, C. /aticornella Clemens, occurs from New Eng-
land to Florida and west through Texas. Its hosts are pecan, walnut, and various
hickories. The adult is brownish and has a wingspread of about 10 mm. The larval
case is brown, smooth, cigar-shaped, and about 6 mm long. The winter is spent as a
partly grown larva in the case. Feeding is resumed in the spring on opening buds
and young foliage. Damage is often serious (454).

The elm casebearer, C. u/mifoliella McDunnough, an introduced species first
observed in the vicinity of New York City in 1901, is widely distributed in the
Northeastern States and also occurs in southern Ontario. Its hosts are English,
Scotch, and various native elms. The female adult is buff, covered with gray
markings, and has a wingspread of about 12 mm.

133

Adults appear in late July and lay their eggs on the leaves of their hosts. At first
the larvae feed within mines in the leaves. Then they emerge and construct tiny
cases in which they migrate to the twigs where they spend the winter. Feeding is
resumed in the spring with each larva eating out a tiny circular hole in a leaf and
then, without leaving its case, mining out an area between the larger veins as far as
it can reach. The case is enlarged as needed to accommodate the growing larva,
eventually reaching a length of 9 to 10 mm. This species 1s primarily a pest of shade
and ornamental trees. The mined parts of leaves turn brown and, when numerous,
are unattractive. Fortunately, outbreaks are usually limited.

The larch casebearer, C. Jaricella (Hubner), an introduced species, was first
recorded in North America in Massachusetts in 1886. It now occurs throughout
most of the range of tamarack in North America, west to central Minnesota and
northwestern Ontario. In 1957, an infestation was also discovered on western larch
in Idaho (298). It is now widespread in the region of western larch. The adult is
silvery to grayish brown, has narrow wings fringed with long hairlike scales, and a
wingspread of about 9 mm. Full-grown larvae are about 6 mm long. The pupa is
brown. This species is one of the most serious defoliators of larch in North
America. Outbreaks have been occurring at about 8-year intervals in eastern Can-_
ada. Heaviest losses result from reduced growth, twig mortality, and increasing
numbers of adventitious shoots. Trees completely defoliated for 2 or more con-
secutive years may be killed.

Eggs are deposited singly on needles in early summer and hatch in about 2
weeks. The newly hatched larva bores directly into a needle and continues to mine it
until late summer. Then the larva lines a hollowed section of the needle with silk
and chews the section free at both ends thereby forming a case. The remaining .
larval period is spent in the case, which is enlarged as needed. A single larva may
mine several needles before the needles fall. Before the onset of cold winter
weather, the larva migrates to an outer twig or branch to which it fastens its case,
usually at the base of a bud. Several cases may be found grouped together around
the bases of spurs from which new needles arise in the spring. The most serious
damage is done by the large larvae as they feed on newly developing foliage during
the spring. Adults emerge from late May to early July, depending on locality and
season. There is one generation per year.

The larch casebearer is attacked by more than 50 species of native parasites in
eastern America, but none is particularly effective in control. Two introduced
hymenopterous parasites, Agathis pumila (Ratzeburg) and Chrysocharis
laricinellae (Ratzeburg) are now widely distributed and are believed to be quite
helpful in control. Rearing methods have been devised for these parasites (/046).

The birch casebearer, Coleophora serratella (L.), an introduced species first
observed in North America in Maine in 1927, is now known to occur throughout the
Northeastern States and from Newfoundland and New Brunswick to southern
Ontario. In the past, this insect was also called the cigar casebearer. Its favored
hosts are paper, gray, and European white birches. Eggs are laid in July along the
midribs and larger veins on the undersides of leaves. Young larvae enter the leaves
and feed as miners for several days, then they emerge and construct cases in which
they live and feed thereafter. Winter is spent in cases firmly attached to the bark,
usually in crotches of limbs. In the spring, the larvae feed on buds and young
leaves, mining as far as possible without leaving their cases. Infested leaves tend to
shrivel. This species has been abundant in Quebec, New Brunswick, and northern
Maine. Many stands of paper birch in New Brunswick were completely defoliated

134

in 1968. This insect is also a pest of apple in the Northern States, and attacks
cherry, hawthorn, plum, quince, and pear as well. It is most injurious in the spring
when the larvae feed on expanding foliage, flowers, fruit, and fruit stems.

The palm leaf skeletonizer, Homaledra sabalella (Chambers), feeds on many
varieties of palms, particularly Canary Island date palm and cabbage palmetto, in
the Southern States. It is often quite injurious in Florida. The larvae feed in groups
of 35 to 100 under webs of silk on both the upper and lower surfaces of the leaf
(264). Eggs are usually laid in masses on the interleaf husks. There are no
hibernation stages and there may be up to five generations a year. Cutting out and
burning all interleaf husks and infested fronds is helpful in control.

Family Agonoxenidae
Agonoxenids

Chrysoclista linneella (Clerck), the linden bark borer, an introduced European
species, was first reported in this country in 1928 when it was found infesting
basswood trees near New York City. Other infestations were later found in sur-
rounding parts of New York and New Jersey, and near Boston, Mass. Its current
distribution is not known. Full-grown larvae are whitish with light-brown heads,
and are about 6 mm long. Adults are present from late May to mid-June and are
thought to lay their eggs on the branches of their host. The larvae bore into and
tunnel the bark. Winter is spent in the larval stage, and pupation occurs in the
spring in cells formed in the galleries close to the surface of the bark.

Family Cosmopterigidae
Cosmopterigids

Stilbosis ostryaeella (Chambers), a leafminer of ironwood, has been recorded
from New York, Kentucky, Ohio, and southern Ontario. Its eggs are laid on the
undersurface of leaves, near the midrib. The larvae feed in the tissues of the leaf
between two lateral veins, and form blotch-type mines. Full-grown larvae vacate the
mines in the fall and drop to the ground where they spin thin, loosely woven silken

cocoons in the litter. Winter is spent in the pupal stage (736).
Family Gelechiidae
Gelechiid Moths

Gelechiid moths are small and have narrow forewings. The outer margins of the
hindwings are usually concave. The larvae of some feed in folded or rolled leaves
webbed together; others feed as leafminers; still others feed in buds, seeds and
cones, and roots. A few species are quite destructive.

Coleotechnites apicitripunctella (Clemens) occurs in Quebec and in the North-
eastern States. Its known hosts are eastern hemlock and baldcypress. The adult is
buff yellow to whitish and has a wingspread of 12 mm. The forewings are marked
with blackish spots and dots, and the hindwings are fringed. The larva is greenish,
sometimes with a brownish tinge, and is about 6 mm long. Adults are present from
early June to mid-July, and the larvae feed for the rest of the summer and fall. They
mine the leaves and web them together, forming broad, flat nests. A nest may
contain six or eight mined leaves where the larvae spend the winter. Feeding is
resumed in the spring, and pupation occurs in late spring or early summer. There is
one generation per year. Local outbreaks occur occasionally.

Coleotechnites thujaella (Kearfott) occurs from New Jersey to New Brunswick
and westward to Saskatchewan, in Canada. The larvae are leafminers on northern
white-cedar. The adult is creamy white with heavy dustings of black and brown
scales and has a wingspread of about 9 mm. There are three oblique blackish bands,
a number of costal and terminal dots, and a shaded apical region on each forewing.
Eggs are deposited from late June to early August between scalelike leaves on

135

growing tips. The larvae bore into the tips and mine along the twigs, causing the
foliage to turn brown. Winter 1s spent in the larval stage in a mine, and there is one
generation per year. Ornamental northern white-cedar is subject to serious injury.

Coleotechnites piceaella (Kearfott) occurs commonly from Maine to Colorado
and from the Maritime Provinces to Alberta in Canada. Its hosts are white, blue,
Engelmann, Norway, red, and black spruces. The adult is light gray and has a
wingspread of about 12 mm. The head and thorax are pale yellow to whitish. The
forewings are buff or ochreous near the base, shading to fuscous at the apex, and
are marked with dark-gray, diagonal crossbands and a few conspicuous black spots.
The hindwings are broad and gray with a silvery sheen; the abdomen and legs are
ochreous, sprinkled with gray. Full-grown larvae are reddish to cinnamon brown
and about 8 mm long.

Adults are active from June to late July, depending somewhat on season and
locality. Eggs are laid singly or, rarely, in groups of two or three either between the
axils of current-year needles, in insect-damaged or mechanically damaged needles,
in insect-damaged cones, or in spent staminate flowers. Some also may be depos-
ited at the base of needles or inserted between the scales of sound cones. The larvae
feed as miners in healthy needles, in needles and cones damaged by other insects,
in spent staminate flowers, and in dead needles on shoots damaged by late spring
frosts (802). Winter is spent in the larval stage, and there is one generation per year.
Damage is usually not very injurious, but may be important on ornamentals.

Other eastern species in the genus Coleotechnites include C. juniperella (Kear-
fott) that mine the needles of redcedar and common juniper in the Northeastern
States, C. dorsivittella (Zeller) that feeds on sweetgum, and C. variella (Chambers)
that feeds on baldcypress. Heavy infestations of C. variella have killed the top 60 or
90 cm of baldcypress trees up to 6 m tall in Ohio. C. chillcotti Freeman is a
common needleminer of longleaf pine in Louisiana. The morphology of the pupa is
discussed (94).

The pine needleminer, Exoteleia pinifoliella (Chambers), occurs in southeastern
Canada and south to Georgia and Texas. Jack, pitch, and shortleaf pines are
preferred hosts, but it has also been observed feeding on Virginia, Scotch, longleaf,
loblolly, and red pines. The adult has a wingspread of about 9 mm. The forewing is
reddish to golden brown and is marked by four narrow, grayish bands. The
hindwings are wider than the forewings. Full-grown larvae are pinkish and about 6
mm long.

Females deposit their eggs in recently vacated, mined needles from May to July,
depending on location. Young larvae vacate the old mined needles and bore into the
bases of current-year needles, killing them within 2 or 3 weeks. Older larvae mine
in both old and new needles, killing the apical portions beyond the entrance holes.
Winter is spent in the larval stage, and there may be two or more generations per
year (4/8). Forest-grown trees are occasionally infested heavily, but are seldom
injured seriously. However, heavy infestations on ornamentals or in plantations may
be serious. The mite Pediculoides ventricosus (Newport) appears to be an effective
predator in some areas. It is reported to have destroyed more than 75 percent of late
instars in infestations in North Carolina (74).

Exoteleia dodecella (L.), the pine bud moth, was discovered in the Niagara
Peninsula, Ontario, in 1928, and now occurs throughout southern Ontario. As far
as known, it has not yet crossed the border into the United States. Its preferred hosts
appear to be Scotch and Swiss mountain pines, but several other pines such as
eastern white, red, jack, and Austrian may be attacked in heavily infested areas.

136

Old Scotch pines, particularly those growing along roadsides, are highly subject to
attack. Infestations on such open-grown trees may persist for years. Heavy infesta-
tions cause a thickening or browning of the needles, followed by branch malforma-
tion. Eventually, the needles thin out. A high proportion of the buds on young trees
may be killed.

Eggs are laid on current-year and |-year-old shoots in late July or early August.
The larvae feed on the needles during the first season; the following spring they
feed on the buds. Winter is spent in the third instar in mined needles (5/9).

The pine candle moth, EF. nepheos Freeman, first recorded in southern Canada
in 1958, is also known to occur in Ohio. Its known hosts are red and Scotch pines
and, rarely, Swiss mountain pine. The adult is small and inconspicuous and has a
wingspread of less than 12 mm. Mature larvae are pale yellow with a reddish hue
and are about 6.5 mm long. Winter is spent in the larval stage.

Adults are present from early July to early August in Ontario (in Ohio they may
appear as early as June 1). Eggs are laid singly or, rarely, in clusters of two to four,
on the sheaths of previous year’s needles or, occasionally, under loose bark scales of
twigs.. Newly hatched larvae bore into the tips of the needles. The remainder of the
summer is spent mining the needle, and winter is spent in the tunnel. In the spring,
the larvae vacate their tunnels and reenter the same needles, adjacent needles, or
swelling buds. Larval growth is completed in May, and pupation occurs in the
flowers or shoots of the tree. There is one generation per year (739).

Larval feeding stunts the growth of new shoots, giving the branches of infested
pines a tufted look. Infestations in Ontario have been reported on pines from 3 to
7.6 m tall.

Anacampsis innocuella (Zeller) occurs in southern Canada and from Mas-
sachusetts to Colorado and Texas. The larva is a leafroller on poplars, willow, and
cherry. The adult is ash gray or slightly darker and has a wingspread of 18 to 22
mm. A pale, wavy, transverse line crosses the forewing well beyond the middle.
Behind it, the color darkens and there are three dots in the cells and two in the fold.
The larva is translucent green.

Larval feeding and leafrolling begin in Ohio about the time the leaves of bigtooth
aspen begin to develop. Before pupation in May or June, the larva severs the petiole
of the rolled leaf, causing it and the enclosed larva to drop to the ground. Rolls
containing two or more leaves tend to remain on the tree. Adults appear during the
last half of June. It 1s believed that winter is spent in the egg stage on twigs of the
host (850).

Anacampsis rhoifructella (Clemens) feeds on viburnum and sumac in the North-
eastern States. The adult is grayish brown and has a wingspread of 15 mm.
Markings on the forewings resemble those on A. innocuella. Larvae are pale brown
to dark brownish-red. On sumac, they live in silken galleries within fruit clusters in
the spring and feed in the fruit spikes.

The palmerworm, Dichomeris ligulella Hiibner, occurs in Canada and in the
Northern States from Maine to Minnesota. Its hosts are various hardwoods such as
apple, plum, pear, cherry, hazel, basswood, and oak. Full-grown larvae are green-
ish and translucent, and about 12 to 15 mm long. There is a pair of narrow, whitish
lines down the back, and a wider one runs along each side.

Egys are laid on the undersides of leaves in early spring. The larvae skeletonize
the leaves in exposed positions or within folded or rolled leaves. Pupation occurs in
these rolls or in ground litter. Adults appear during July or August and apparently

137

live until the following spring. Although most important as a pest of apple orchards,
this species is capable of seriously defoliating oak during outbreaks.

The juniper webworm, D. marginella (F.), an introduced species, occurs in
many Eastern and Western States and in southern Canada. Its hosts include various
species of Juniperus, such as Irish juniper, common juniper, and eastern redcedar.
The adult is brownish with white front and rear margins of the forewings, and has a
wingspread of 15 mm. Full-grown larvae are light brown and from 12 to 15 mm
long.

Adults are present during June and July, and deposit their eggs in leaf axils of new
growth. Newly hatched larvae feed first as needleminers, entering the upper surface
of the needle near the axil. Mined needles turn brown and die. Dead needles are
incorporated in webbing constructed between branchlets. Larvae continue to use
mined needles as protective retreats while developing. During July, larvae crawl
from primary mined needles and feed on adjacent needles, eventually constructing
silken tubes leading from holes in the mined needles to other nearby needles. The
webbed area is expanded from July to September. Entire trees up to 2.4 m tall may
be completely webbed (fig. 44). Winter is spent in the larval stage in a silken case.
The larvae resume feeding in the spring, becoming full grown and pupating from
mid-May to early June (939). Trees grown as ornamentals or in permanent plantings
may be seriously damaged by this species. Cutting and removing webbed masses of
foliage is a helpful control practice.

The redbud leaffolder, Fascista cercerisella (Chambers), feeds on redbud from
Delaware and Maryland to Illinois and southward. The adult is velvety black except
for its white head and a white collar. The forewings are slightly bronzed and marked
with three costal spots and several white terminal points. The larvae feed on leaves
which they web together. There are at least two generations per year.

Battaristis vittella (Busck) is widely distributed throughout the Eastern United
States and southern Canada. It has been reared from the buds of Swiss mountain
pine; from the cones of Scotch, Austrian, and longleaf pines; and from galls of
midges on loblolly pine. The adult has bright-red eyes, the forewings are cinnamon
brown and traversed by gray bands, and the wingspread is 8 to 10 mm. Mature
larvae are slender and from 4 to 6 mm long. Winter is spent in the larval stage in a
tunnel in a bud or cone. Pupation occurs in the spring, and adults appear by May.
Superfamily Yponomeutoidae—Family Plutellidae
Plutellids

The plutellids loosely web leaves together. The mimosa webworm, Homadaula
anisocentra Meyrick, an introduced species first reported in the United States at
Washington, D.C., in 1942, is now widely distributed from New Jersey and
Pennsylvania southward to Florida, Alabama, and Mississippi, and westward to
Kansas and Nebraska. Its host trees are silktree and honeylocust. The adult has a
wingspread of about 12 mm. The forewings are mouse gray, except for a silvery
luster and a stippling of black. Full-grown larvae are pale green to dark brown, are
marked with five longitudinal white lines, and are about 12 mm long. Pupae are
yellowish brown and about 6 mm long. They are found in whitish, silken cocoons.

The pupae generally overwinter in the soil or in other protected places. Adults
appear by June, and the female deposits her eggs on flowers or foliage, or on the
bark of small branches and twigs. Egg laying continues throughout the season. First
and second instars spin webs around flowers and leaves, within which they live and
feed (fig. 45). Adjacent surfaces of webbed leaves may be skeletonized, turn brown,
and die. Older larvae feed on the tender, terminal leaves. Pupae of this generation

138

Osea oe ie
Courtesy Can. For. Serv., Can.
Dep. Environ., Sault Ste. Marie, Ont.

Figure 45.—Webs formed by the
mimosa webworm, Homadaula
anisocentra.

F-519583

Figure 44.—Defoliation by the juniper
webworm, Dichomeris marginella.

are formed in the webbing of infested trees. Second-generation adults appear during
late July and early August and deposit their eggs on webs formed by first-
generation larvae. In heavily infested areas, the larvae of this generation may
completely defoliate their hosts. Some of the second-generation larvae pupate in
cocoons spun in webs on trees, and the adults appear during the fall. Other larvae
move to the soil or other protected places, pupate, and spend the winter (/276).

The mimosa webworm 1s a serious defoliator of ornamental plantings of silktree
and honeylocust. The thornless variety of honeylocust is heavily attacked 1n nurser-
ies. Season-long protection against attack has been obtained by the use of systemic
insecticides (3/9).

139

Family Yponomeutidae
Ermine Moths

The ermine moths web foliage together. The ailanthus webworm, Afteva punc-
tella (Cramer), feeds on the foliage of ailanthus throughout the Southern States and
north to New York and the Lake States. The adult is orange to brownish and has a
wingspread of 25 to 30 mm. The forewings are bright yellow, and each one has four
rows of round yellow spots on a blue background. The larvae feed on leaves
enclosed in frail, silken webs. The pine needle sheathminer, Zelleria haimbachi
Busck, feeds rather commonly on jack pine in southern Ontario. It may occur in the

Lake States.
Family Argyresthiidae
Argyresthiids

Moths of this family are small, usually brightly patterned, and have rather broad
wings.

The genus Argyresthia contains a number of leafmining species. The adults have
wingspreads of about 8 to 12 mm. While at rest, their wings are folded close to the
body, their front legs are extended forward, and their hindlegs are slanted upward at
45° angles.

Argyresthia freyella Walsingham is generally distributed in eastern Canada and
south and west to the Middle Atlantic States and Missouri. Its hosts are redcedar
and northern white-cedar. The adult is whitish and has a wingspread of about 8 mm.
The forewings are golden with silvery spots and bands and a black dot at the apex of
each. The larvae feed in mines that they extend along main branches and
branchlets. Winter is apparently spent in the mine as a full-grown, yellowish-green
to green larva, and there is one generation per year. Pupation takes place in the
spring in cocoons spun on leaves or branches. Damage to ornamentals and nursery
seedlings appears to be less than that caused by the related species, A. thuiella. A.
aureoargentella Brower attacks northern white-cedar. It resembles A. freyella ex-
cept that its cocoons are whiter and larger.

The arborvitae leafminer, A. thuiella (Packard), feeds on northern white-cedar
throughout much of the same area as that occupied by A. freyella. The adult is light
gray to white and has a wingspread of about 8 mm. The forewings are marked with
brown and there is a black spot in the middle edge of the distal end of each. The
larva is about 3 mm long. The head and cervical shield are shiny black, the body is
green with a reddish tinge, and the legs and anal plate are black. Adults appear from
late May to mid-July, and the female deposits her eggs in the axils of branchlets or
along the edges of leaves. Newly hatched larvae bore into the leaves and feed in
them as miners for the rest of the season. Winter is spent in the larval stage in the
mine and pupation and adult emergence occur in the spring. Outbreaks in Maine
have severely damaged forest stands of northern white-cedar. Damage to ornamen-
tals and nursery seedlings is often serious. Heavily defoliated trees may be killed
(160). The habits of the species of arborvitae leafminers have been discussed
(1081).

Larvae of the apple fruit moth, A. conjugella Zeller, bore in the fruit of
mountain-ash, serviceberry, hawthorn, apple, plum, and cherry in the Northeast.
The adult is dark gray and has a wingspread of 10 to 12 mm. The forewings are dark
gray, very slender, crossed by black and silver-white bands, and each bears a
yellowish-white spot on the outer margin. A. laricella Kearfott, the larch shoot
borer, larvae bore in the shoots of larch in southern Canada and the Lake States.

140

Family Sesiidae
Clearwing Moths

Members of this family are known as clearwing moths because the greater part of
one or both pairs of wings are without scales, thus leaving them clear or trans-
parent. The forewings are long and narrow, with the outer margins short and the
anal veins reduced. The hindwings are somewhat broader than the forewings, and
the anal areas are well developed. In some species, the two sexes are colored
differently. Many species bear a striking resemblance to bees or wasps. The adults
are swift fliers and are most often seen around flowers. The larvae are ivory-white
and mostly unmarked. They bore in the roots and basal stalks, the trunks, or
branches of trees, shrubs, and vines, or in the stems and roots of herbaceous plants.
A few form galls, others are inquilines in galls, and some inhabit injured areas on
the trunks or branches of their hosts. A number of species are important pests of
forest and shade trees and ornamentals. The family is discussed (345, 354, 795).

The hornet moth, Sesia apiformis (Clerck), an introduced species first observed
in North America around 1880, is now known to occur in Massachusetts, Connecti-
cut, New York, New Jersey, Pennsylvania, and California. [ts hosts are poplar and
willow. The larvae bore in the roots, trunks, or large limbs. The adult is brownish
black except for yellow markings on the head and sides of the thorax; it has a
wingspread of 34 to 44 mm. It also has black and yellow bands on the abdomen,
and brown legs. Because of its close resemblance to the giant hornet, it is known as
the hornet moth. Full-grown larvae are white with reddish heads and are about 30 to
50 mm long. They excavate extensive tunnels in their hosts, causing swellings to
occur. Young trees are often killed. Two years are required to complete the life
cycle. The first winter is spent in the larval stage in the wood; the second, as a larva
in a cocoon in wood borings in or close to the base or roots.

Sesia tibialis (Harris), the poplar clearwing moth, attacks poplar and willow
from New York to Nova Scotia westward across the Northern States and southern
Canada to the Pacific Coast and southward to California. Adults are distinguished
_ by their black abdominal segments, of which all but the second and fourth have
narrow posterior yellow margins. Full-grown larvae are 40 mm or more long.
Infestations are located well down on the trunk or in the roots.

The dogwood borer, Synanthedon scitula (Harris), occurs in southeastern Can-
ada and throughout the eastern half of the United States. Although normally a bark
borer in oaks, it also attacks a wide variety of other deciduous hardwood trees,
shrubs, and sometimes pine. It is often an important pest of flowering dogwood and
pecan. In the South, it is commonly called the pecan borer. Abnormal growths such
as woody galls, excrescences due to fungi, rusts, blight, and bruises and healing
wounds are attractive as points of attack. Galls, such as those produced by the
cynipid Callirhytis cornigera (Osten Sacken) on black and pin oaks, sometimes
occur in the thousands on a single tree, and nearly every one will be infested by this
borer (354).

The adult is a small, blue-black moth with yellow-banded legs and yellow stripes
on segments two and four of the abdomen. The wings are transparent with blue-
black margins, and the wingspread is from 14 to 20 mm. The larvae are whitish
with brown heads and are up to 14 mm long.

Eggs are laid from late spring to midsummer on rough bark or around wounds.
The larvae enter the bark through openings and feed in the cambial area. Infested
areas are sometimes up to 61 cm or more in length and may contain up to SO larvae
each on the larger trees. A single larva can kill a dogwood 10 cm in diameter in |

141

year. There are one and possibly two generations per year, depending on locality.
Keeping the bark smooth, especially at the base of branches, has been recom-
mended for control on dogwood (/239).

The rhododendron borer, S. rhododendri (Beutenmiller), attacks rhodo-
dendron and sometimes laurel and azalea in the Atlantic Coastal States. The adult is
blackish, except for various white and yellow markings, and has a wingspread of 10
to 15 mm. Mature larvae are up to 15 mm long. The winter 1s spent as a full-grown
larva in the stem of its host. Pupation occurs in the spring, and the adults appear
during May and June. Eggs are deposited on the bark, and the larvae bore into the
stems, Causing ugly scars and sometimes killing large branches and small plants.
Cutting out and burning infested parts of plants and the use of two to three well-
timed insecticide applications are recommended control practices (908).

The maple callus borer, S. acerni (Clemens), is widely distributed in southern
Canada and throughout the Eastern United States south to Florida. Its hosts are
sugar and red maples. The adult is amber colored and has a wingspread of 20 to 25
mm. The head is orange-yellow and there is a red tuft of hairs at the posterior end of
the yellow-banded abdomen. Full-grown larvae are white and from 14 to 19 mm
long.

Adults appear in May and June and deposit their eggs in roughened places on the
trunk, preferably on or near wounds. The larvae bore through the bark and into the
sapwood. Slightly wounded trees are often severely damaged. Winter is spent in the
larval stage and pupation occurs in the spring. Just before it changes to an adult, the
pupa wriggles part way out of its burrow. When the adult emerges, the cast pupal
skin is left sticking out of the bark. There is one generation per year. Smoothing
roughened bark areas, removing borers from under the bark in the spring, and
painting wounds are recommended control practices. The southern form attacks
silver maple in Florida and Georgia.

The pitch mass borer, S. pini (Kellicott), occurs in eastern Canada and the
Eastern United States southward to Georgia and Tennessee and westward through
the Great Lakes region. Its preferred host apppears to be eastern white pine but it
also attacks Austrian and Scotch pines and spruce. The forewings are blue-black,
with a metallic-green luster, and have a wingspread of 25 to 30 mm. There is an
orange band on the fourth abdominal segment and a tuft of orange scales at the tip
of the abdomen. Mature larvae are white to pink and 25 mm long.

Eggs are deposited on the bark during June and July, usually near a wound, on
old scars, or just below a branch. The larvae bore in the inner bark and sapwood,
excavating more or less transverse tunnels, and cause copious flows of pitch. This
pitch accumulates in masses 8 to 10 cm in diameter at the entrance hole. Pupation
occurs in the pitch mass, and when the moth emerges the empty pupal case is left
sticking out of the mass. The life cycle requires 2 to 3 years. Although not a killer
of trees, this species causes some lumber degrade as a result of its activities. No
practical control methods are known for use on forest-grown trees. Borers in shade,
ornamental, or park trees may be removed with a knife.

Several other species of Svnanthedon also occur in eastern North America. S.
geliformis (Walker) 1s a bark borer in dogwood and pecan. It also infests cynipid
galls on the trunks and branches of oaks, and diseased and injured tissues of various
other plants. The apple bark borer, S. pyri (Harris), attacks apple, hawthorn,
serviceberry, and mountain-ash. The larvae excavate shallow, tortuous tunnels in
and beneath the bark of trees usually injured by storms and disease, causing the

142

bark to blister and peel. The lesser peachtree borer, S. pictipes (Grote & Robin-
son), breeds in peach, American plum, and in fungus growths on American plum. It
is a major pest of peach. S$. viburni Engelhardt larvae bore under the bark of
viburnum, preferably in injured areas or galls. S. acerrubri Engelhardt is a bark
borer in maple, preferably in branches and often in wounds and scars caused by
other insects. S. sapygaeformis (Walker) has been recorded infesting woody galls in
oaks in Florida. S. castaneae (Busck) bores in the trunks of American chestnut,
preferably in bruised areas. S. rubrofascia (Henry Edwards) larvae excavate long,
sinuous tunnels in the bark of tupelo from Maryland to Florida. Injuries and healing
wounds on well-matured trees are preferred. S. sigmoidea (Beutenmiuller) attacks
low-growing willows in bays, along streams, and in depressions in sand dunes of
coastal and lake regions. Infestations are often heavy. S. proxima (Henry Edwards)
larvae bore in the branches and exposed roots of low-growing willows in moist or
shady locations. S$. bolteri (Henry Edwards) also attacks low-growing willows.

The persimmon borer, Sannina uroceriformis Walker, occurs from southern
Maryland to Kansas, Texas, and the Gulf Coast and feeds as a larva in the solid
wood of the base and taproots of persimmon. The adult is mostly bluish black and
has a wingspread of 28 to 32 mm. Full-grown larvae are up to 30 mm long. Trees
growing in cutover areas and in hedgerows are particularly subject to infestation.
Young trees in nurseries are also damaged seriously at times. The life cycle requires
2 or 3 years.

The lilac borer, Podosesia syringae (Harris) (fig. 46), occurs from Texas and
Saskatchewan east throughout the United States and southern Canada. It attacks
lilac, ash, privet, and other trees and shrubs of Oleaceae. The adult has a wing-
spread of 26 to 28 mm. The species is polymorphic with several distinct adult
morphs varying from dark brown sometimes with chestnut-red markings to light
brown and yellow to distinct yellow banding of the abdomen (345, 1006). The
larvae bore into trunks and branches, excavating galleries up to 15 cm long and
causing nursery cull, decline and mortality in shelterbelts, and degrade in timber
stands. Winter is spent in the larval stage, and there is one generation per year.
Woodpeckers and parasites are the major natural controls. Cultural practices that
avoid bark wounds and maintain good tree vigor help to minimize losses. Chemical
control is also effective.

a

is 7 Mt £% :
Courtesy D. G. Nielson, Ohio State Univ.
Figure 46.—Adult of the lilac borer, Podosesia syringae.

Be a CR

143

The banded ash clearwing,. P. aureocincta Purrington & Nielson (1006) (fig.
47), is known from New York south to Florida and west to Oklahoma. It has been
reared from green and white ashes and probably attacks other Fraxinus spp. The
adult is mostly brownish black but can be distinguished from the lilac borer by a
narrow, orange-yellow, partial band on the fourth abdominal segment. Also, the
eggs are darker, slightly larger, and have a smoother surface than those of the lilac
borer. It is particularly destructive to ornamental plantings and to trees grown for
timber. However, populations are more scattered than those of the lilac borer.

Courtesy D. G. Nielson, Ohio State Univ.

Figure 47.—Adult of the banded ash clearwing,
Podosesia aureocincta.

Paranthrene simulans (Grote) occurs throughout the Eastern United States west
to Texas and in eastern Canada. It attacks trees in both the red and white oak
groups, but host preference varies with region. In the Gulf States it shows a decided
preference for trees more than 30 cm d.b.h. in the red oak group, especially Nuttall
and cherrybark oaks (//33, 1/135). In the Northeast, injury is greatest in nursery
stock, saplings, and young trees in both the white and red oak groups. Moths
emerge in June and July and deposit eggs in bark crevices on the lower trunk (fig.
48). The larvae bore through the bark and into the wood. Here they excavate
galleries up to 9 mm in diameter and 10 cm long. For the first 3 to 5 cm the gallery
slopes upward, then it continues straight up. The life cycle requires 2 years.
Damage includes nursery cull, entryways for decay fungi, and degraded butt logs.
Open-grown trees are most susceptible, thus populations can be minimized by
maintaining well-stocked stands, preventing bark injuries, and removing or treating
brood trees.

Paranthrene dollii (Neumoegen) bores in the base and root-collar area of young
cottonwoods and black willow in the Deep South. The adult is brown with dark
wings and yellow crossbands on its abdomen and has a wingspread of about 37 mm.
Eggs are laid in bark crevices. The larvae bore in the wood and pith, constructing
open tunnels up to 15 cm long. Heavily infested trees are weakened and subject to
windbreakage. Damage is often severe in cottonwood nurseries and plantations in
the Deep South where losses average 12 percent cull in nursery stock (//32). There
is One generation per year.

144

F-049923; 519922
Figure 48.—Adult and larva of Paranthrene simulans.

Paranthrene tabaniformis (Rottemberg) larvae bore into the branches and termi-
nals of young cottonwoods, weakening them and causing some to break. There are

one or two generations per year.
Family Choreutidae
Choreutids

The apple-and-thorn skeletonizer, Eutromula pariana (Clerck), an introduced
species first recorded in New York State, is now known to occur in Virginia west to
Indiana, and north to southern Canada. The larva feeds mostly on the leaves of
apple, pear, and hawthorn but mountain-ash, birch, willow, plum, and sour cherry
are also attacked. The adult is dark reddish-brown with a purplish tinge and has a
wingspread of 12 mm. The forewings are often marked with faint, pale bands and
wavy, black lines. Three or four white spots are usually along the costal margin.
Full-grown larvae are yellowish green with prominent black tubercles and are about
12 mm long.

Young larvae skeletonize the undersurfaces of the leaves under loose webs. Older
larvae move to the upper surfaces and draw the opposite sides of leaves together
with silk. They feed inside the fold, consuming everything but the lower epidermis
and larger veins. Damaged leaves curl, turn brown, and fall by early September.
Pupation occurs in cocoons spun in the angles or folds of leaves, on weeds or other
objects, and even in cracks of buildings. There are three and possibly four genera-

tions per year.
Family Cossidae
Carpenterworm Moths

Carpenterworm moths have fairly heavy, spindle-shaped bodies, and narrow,
pointed wings. The larvae excavate large galleries in the wood of trees, often
causing serious injury.

The leopard moth, Zeuzera pyrina (L.), an introduced species, is known to
occur from the Philadelphia area to northern Massachusetts. Its favored hosts are

145

elm, maple, ash, beech, walnut, oak, chestnut, poplar, willow, apple, pear, and
plum, but it will attack scores of other species. The female is heavy-bodied and has
a wingspread of 62 to 75 mm. Full-grown larvae are about 50 mm long. The body is
usually pale yellow but may have a pinkish tinge. It is aiso sparsely hairy and dotted
with brown or biack tubercles.

Adults are present from late spring until fall. Eggs are deposited either singly or
in small groups in bark crevices. Young larvae bore directly into twigs, branches, or
the trunk and feed in the heartwood. When a larva becomes too large for a twig or
branch, it vacates it and migrates to a larger one. As the larva feeds, it pushes chips,
matted excrement, and frass to the outside through the entrance hole. Pupation
takes place within the tunnel, and the life cycle requires 2 years (6/2).

The leopard moth may cause considerable damage to its host. Damaged twigs
wilt and break off, small branches break and hang down, larger branches are girdled
(fig. 49) and may break in the wind, and small seedlings are killed. Ugly scars
appear on the trunks of large trees where the bark dies and splits over wounds. The
removal and destruction of infested twigs, branches, and heavily infested trees is a
recommended control practice. Borers in valuable shade trees can be killed by
probing their tunnels with flexible wires.

Courtesy Conn. Agric. Exp. Stn.

Figure 49.—Elm branch nearly severed by larvae of the
leopard moth, Zeuzera pyrina.

The pecan carpenterworm, Cossula magnifica (Strecker), occurs throughout
the Southern States. Its hosts are pecan, oak, and hickory. The adult is grayish with
brown markings and has a wingspread of 37 mm. Full-grown larvae have pinkish
bodies sparsely clothed with short, fine hairs and are about 37 mm long.

Adults appear in May and June and deposit their eggs on the bark of small twigs.
Newly hatched larvae bore into the twigs and tunnel in the pith. Later, they emerge
and migrate to larger branches or the trunk where they enter and excavate tunnels up
to 8 cm long. Frass is extruded through holes that are about 6 mm in diameter in the
trunk; it can usually be found in small heaps at the base of infested trees. Winter is
spent in the larval stage, and pupation occurs in the tunnel in the spring. There
appears to be one generation per year (/35).

Acossus centerensis (Lintner), the poplar carpenterworm, and A. populi
(Walker), the aspen carpenterworm., also occur in the Eastern United States. A.
centerensis bores in poplars from New Jersey to North Dakota. A. populi is found
sparsely in poplars and cottonwoods from coast to coast.

The carpenterworm, Prionoxystus robiniae (Peck) (fig. 50), is widely dis-
tributed in the United States and southern Canada where it breeds in various

146

hardwoods. In the Eastern and Southern States, the oaks, particularly those of the
red oak group, are most heavily damaged. In the Prairie States, green ash is the
chief host. Other hosts are black locust, elm, maple, willow, cottonwood, and
occasionally fruit trees and ornamental shrubs (540). The adult is dark, slightly
mottled, stout-bodied, and has grayish black hindwings. The female is lighter
colored and considerably larger than the male and has a wingspread of 75 mm. The
posterior half of the hindwing of the male bears a large yellowish to orange spot
with a black border. Full-grown larvae are greenish white, nearly naked, bear
simple setae, and are 50 to 70 mm long. The head is shiny brown and armed with
powerful, nearly black mandibles. The thoracic legs are yellowish and _three-
jointed, and each bears a curved, pointed tarsal claw.

F-532850-54

Figure 50.—The carpenterworm, Prionoxystus robiniae:
A, adult male; B, adult female; C, eggs; D, larva; E,
pupal cases.

147

Carpenterworm moths begin to appear in early April in the South, the last week
of May in the Central States, and the first week of June in more northerly regions.
The female deposits as many as 450 to 800 eggs in groups in bark crevices, near
wounds, or under vines, lichens, or moss. Young larvae bore directly to the inner
bark or enter it through openings, then bore into the wood, their tunnels angling
upward in the sapwood and straight upward in the heartwood. Feeding may be
finished in | year in the Deep South but may continue for 3 or 4 years in the North.
Tunnels are kept open and enlarged as needed by the growing larvae; eventually
tunnels may reach a diameter of 18 mm and a length of 300 mm. Mature larvae line
their tunnels with loose, silky, yellowish-brown webs. Pupation occurs at the upper
end of the tunnel. Before completing its transformation to an adult, the pupa
wriggles to the mouth of the tunnel and continues until its head and thorax are
protruding. Even after the adult has formed and departed, the pupal case usually
remains in place, sticking out of the opening.

Carpenterworms seldom kill trees outright, although heavily riddled small trees
may be broken off by the wind. Open-grown trees, or trees growing on poor sites
such as dry ridgetops or ridge slopes, are especially subject to attack and damage.
The greatest damage results from the degrade of lumber cut from infested trees. The
overall value of rough-cut, oak lumber may be reduced by as much as 15 percent.
Borer-caused losses, mostly carpenterworm, in oaks cut in the Ozark National
Forest, Ark., and updated to 1980 lumber prices amounted to $42 per 5.7 cubic
meters (//3/).

Woodpeckers, other birds, and spiders destroy large numbers of carpenterworms
and are among the most important natural controls. Insect parasitism is inconse-
quential. Fungi and other disease pathogens kill some larvae. Cultural practices that
promote tree vigor, recognize and remove brood trees, and minimize bark injuries
will help to reduce losses in the forest. Shade trees can be protected by painting
injuries with wound dressing. Larvae in small trees can be killed by probing their
tunnels with a wire. Insecticides applied at the time of oviposition and egg hatch
can be used to protect high-value trees.

The little carpenterworm, P. macmurtrei (Guérin), a species very similar to the
carpenterworm, is widely distributed in eastern Canada and the Eastern United
States. The larvae are borers in oaks, and their habits are similar to those of the
carpenterworm. They spend their first summer in the outer layers of bark and the
second in the sapwood. During the third summer, they bore in the woody part of the
tree, making a labyrinth of crossing and recrossing tunnels. The third winter is
spent in the pupal stage in the tunnel. This species attacks all parts of the tree more
than 25 mm in diameter, and trees infested continuously become badly hon-

eycombed. Fortunately, infestations are usually local.
Family Tortricidae—Subfamily Olethreutinae
Olethreutine and Leafroller Moths

This subfamily contains a large number of economically important species of
forest insects (1004). Several are particularly important as pests in nurseries and
plantations. The larvae differ widely in their feeding habits. Some feed by boring
into the buds, twigs, stems, roots, seed, or fruit of their hosts; others, as leafminers,
feed on the foliage from within folded leaves or on the exposed surface. Many
adults and larvae of North American species are described (55/, 552, 792, 793).

Episimus argutanus (Clemens) is widely distributed and feeds as a leafroller on
sumac, poison-ivy, witch-hazel, and various other shrubs. The adult has a wing-
spread of about 13 mm and is dull reddish or grayish brown, mottled with darker

148

colors. Each larva lives in a rolled leaf or between two leaves fastened together with
silk. There are two or more generations per year.

The spruce needleminer, Endothenia albolineana (Kearfott), occurs from coast
to coast in southern Canada and from Maine to North Carolina, Colorado, and
Idaho in the United States. Its hosts are white, Norway, Engelmann, Colorado blue,
and black spruces. The adult is dark brown and has a wingspread of about 13 mm.
The forewings have three irregular, transverse, broken, grayish-white bands. Its
biology has been studied (//88).

Adults are present from mid-May to mid-June. Eggs are deposited so that they
overlap in a single row on the undersides of needles in groups of 2 to 12 eggs each.
Young larvae are gregarious and bore into the bases of old needles, hollowing them
out. Older larvae feed singly. Shortly after beginning to feed, the larvae construct
nests composed of dead needles and frass which are held together by fine silk
strands. As the larvae develop, their nests are continually enlarged. Winter 1s spent
in the larval stage in the nest. Feeding is resumed on adjacent needles in the spring.
Pupation takes place in silken cocoons within the nest during late spring or early
summer.

The spruce needleminer is most important as a pest of ornamental spruce, but it
also occasionally causes serious defoliation in forest stands. The presence of webs
on ornamentals reduces their esthetic value. This can be largely prevented by
handpicking or washing the webs from the trees before the buds break in the spring.
Webs washed to the ground should be picked up and burned.

Evora hemidesma (Zeller) feeds as a leafroller on alder, poplar, and willow in
southern Canada, and in the Eastern States from Maine to Virginia and Kentucky.
The adult is reddish brown with a darker median band on the forewing and has a
wingspread of about 15 mm. Full-grown larvae are dark green to almost black,
sparsely hairy, with light-colored tubercles and light-brown heads, and are about 18
mm long. Larvae are present from May to July.

The European pine shoot moth, Rhyacionia buoliana (Denis & Schiffermiiller)
(fig. 51), an introduced species first recorded in North America on Long Island,
N.Y., in 1914 (178), is now widely distributed in the Northeastern States and
southern Canada. Outlying infestations also occur in Washington, Oregon, and
British Columbia. Its hosts include Scotch, red, Austrian, Swiss mountain, Jap-
anese red, Japanese black, ponderosa, eastern white, jack, pitch, longleaf, and
Virginia pines (853, 998). Red, Swiss mountain, Scotch, and Austrian pines are
most heavily attacked in the Eastern United States, especially red and Swiss
mountain. The adult moth is rusty orange-red and has a wingspread of about 20
mm. The forewings are marked with several irregular, forked, silvery crosslines;
the hindwings are dark gray, and the legs are whitish.

Adults appear in late spring and fly at dusk. They lay eggs singly or in groups of 2
to 10 on the bases of buds, on needle fascicles and twig tips, or on the bark of new
and old shoots. Newly hatched larvae spin resin-coated, tentlike webs between
needle sheaths and the stems of current-year growth, then they bore through the
sheaths and mine the bases of the needles. About midsummer, the larvae move to
buds and construct new resin-coated webs. At first these webs glisten brightly, then
they solidify into yellow-white masses. Feeding ceases in August. Winter is spent
in the larval stage in a feeding tunnel in or near a bud. When activity is resumed in
April, the larvae move to undamaged buds and new shoots, construct new tents, and
resume feeding. During this period, a single larva may feed on more than one bud

149

if
ei

- F-493841, 493467

Figure 51.—European pine shoot moth, Rhyacionia
buoliana: A, red pine tip with solidified resin mass
broken open to show partly grown larvae; B, red pine
stand badly damaged by the shoot moth. Note spiked
and bushy tops.

or shoot. Larvae reach maturity in May, and pupation occurs inside the burrow in
late May or June. From 2 to 3 weeks later, the larva works its way out of its
chamber. There is one generation every 12 months, but occurring in 2 consecutive
years.

The most important and permanent damage to trees results from spring feeding.
The killing of terminal and lateral buds results in dead, spike tops. The develop-
ment of adventitious buds below this dead portion often causes the formation of
dense, bushy growth the following season. The killing of the terminal bud and the
development of several lateral buds into competing leaders results in forked stems.
Shoots weakened by larval tunneling may fall over, but continue to grow. This.
results in the formation of crooked trunks and branches called “post horns.” All
open-grown young trees of susceptible species below a height of 6 to 8 m are subject
to attack and damage. Taller trees or trees growing in closed stands are usually not
seriously damaged. Also young trees having adequate soil moisture during early
summer are not damaged; apparently the needlemining larvae are “pitched out.” In
Ontario this insect often destroys second-year pine cones on young trees and on the
lower branches of larger trees (760).

Climate is an important factor affecting the distribution and abundance of the
European pine shoot moth. Overwintering larvae are killed by temperatures colder
than about — 29° C. Warm, dry summers followed by mild winters permit max-
imum survival. Rate of tree growth is also important as damage is usually most
severe on slow-growing trees. Introduced and native species of parasites normally
destroy about 10 percent of the population. Several species of parasites introduced
to North America have become established, and two of these species, Orgilus
obscurator (Nees) and Lypha dubia Fallén, appear to be most successful. Refer-
ences are available for more complete information on arthropods attacking Rhya-
cionia spp. (526, 788, 1184, 1363).

The planting of susceptible pines on good sites, ““snow-depth pruning,” and
Christmas tree shearing are helpful in reducing damage (552). Fumigation in the
spring has provided a degree of control on seedlings and larger ornamental spec-
imens (197).

Many reports have been issued on the biology, ecology, and control of the
European pine shoot moth. In addition to those cited in preceding paragraphs, there

150

are several others that are of special interest (452, 500, 541, 545, 856, 858, 992,
1206).

Rhyacionia adana Heinrich has been recorded from Massachusetts, Pennsylva-
nia, Virginia, Michigan, Wisconsin, Ontario, and elsewhere (995). Red, jack, and
Scotch pine seedlings usually under | m tall in nurseries, plantations, and natural
stands and the lower half of trees up to 8 m tall are attacked. The adult has a
wingspread of about 16 mm. The forewing has the outer third red and the remainder
gray with four pairs of grayish-white vertical bars.

In Ontario, eggs are laid between needles just above the needle sheath of the
needle fascicle. Young larvae spin silken cases between two old needles just above
the sheath. Then they enter the needles and mine toward the tip. Later, after new
needle growth has begun, they enter and mine the developing shoots. Several larvae
may inhabit a single shoot and riddle it with their tunnels. Then they vacate the
shoot and move to and destroy the buds. Full-grown larvae crawl down the stem and
pupate in cocoons cemented to the stem below the soil surface. There is one
generation per year (S20). Serious damage has been recorded in red and Scotch pine
plantations in Canada and the Lake States. Pole-volume yields have not been found
to be seriously affected, however (S60).

The pitch pine tip moth, R. rigidana (Fernald), occurs from Florida to Texas
and north to Missouri, New York, and Maine (998). It feeds on pitch, Corsican,
Virginia, red, Scotch, loblolly, and slash pines. The adult is similar to the adult of
the Nantucket pine tip moth, but is slightly larger and more colorful. The forewings
are silver-white with crossbands of silver-white scales, and the hindwings are silver-
gray. Damage 1s also similar to that caused by the Nantucket pine tip moth. Pitch
pines up to 40 cm in diameter may be infested.

The Nantucket pine tip moth, R. frustrana (Comstock) (fig. 52), 1s widely
distributed in the Eastern, Central, and Southern States, and elsewhere (998). Its
hosts include nearly all species of pines growing within its range; the only excep-
tions are longleaf and eastern white pines (/366). Slash pine is also somewhat
resistant, but is occasionally attacked. In the South and Southeast, loblolly and
shortleaf pines are preferred; in the Northeast and Middle Atlantic States, pitch,
Virginia, and Scotch appear to be favored; and in the Central States, shortleaf is
attacked most heavily. The adult has the head, body, and appendages covered with
gray scales. The forewings are marked with irregular brick-red and coffee-colored
patches, the patches being separated by irregular bands of gray scales; the wing-
spread is about 12 mm.

Winter is spent as a pupa within the injured tips of the host. Adults begin to
appear on warm, sunny days in early spring—as early as January in the Deep
South. Egg laying begins in a few days, during dusk and darkness. The eggs are
deposited on new or old-growth needles, in the axils of needles and stems, on
developing tips, or on buds. Newly hatched larvae wander about the shoots looking
for suitable feeding sites. Soon, they construct delicate webs in axils formed by
developing needles and stems. Then the larva bores into a needle sheath and feeds
on the needle, which is then severed. Second instars spin new and larger webs
between buds, or between buds and needles; they feed in the buds. When a bud is
consumed, the larva moves to another bud on the same or a different shoot.
Eventually, the connective tissue of the tip is severed, and the damaged portion
turns brown. The larva continues to feed within the shoot and bud. Once having
consumed the bud, it bores down the center of the stem. The larval period lasts for 2

151

A, F-531246; B, F-532848
Figure 52.—Nantucket pine tip moth, Rhyacionia
frustrana: A, adult; B, pupa in loblolly pine tip.

to 4 weeks. Toward the end of this period, the larva constructs a webbed cell within
the shoot in which it pupates. In the Deep South, there are three or more generations
per year (97). In Pennsylvania and Ohio, there are only two per year.

The Nantucket pine tip moth is a major pest of young pines in the Eastern United
States. During recent years, it has become increasingly abundant and destructive as
a result of the establishment of large areas of pine plantations and seed orchards.
Damage by this insect retards height growth, causes crooks or forks in main stems,
reduces cone crops, and, occasionally, results in the death of the tree. Attacks are
generally restricted to trees less than 4.5 m in height and are most severe in young
plantations, but severe attacks on commercial-size trees have also been reported.
This species causes serious losses of flowers and conelets of shortleaf pine in seed
orchards (362).

At least 60 species of parasites of the Nantucket pine tip moth have been recorded
(1363), but they rarely provide satisfactory control. Damage can be reduced by
limiting the planting of susceptible pines to sites to which they are well adapted.
Close spacing and planting under an overstory may also be helpful.

Rhyacionia bushnelli (Busck), the western pine tip moth, occurs from the
Dakotas and Nebraska south to Kansas and New Mexico east to Missouri. Its
preferred hosts are ponderosa, red, jack, and Scotch pines. Infestations have been
particularly severe in pine plantations in Nebraska (307, 497). The adult is indis-
tinguishable from those of the Nantucket pine tip moth, but it is believed that R.
bushnelli is a distinct species because of its overwintering habit and size (998).
There is one generation per year in the Dakotas, one to two per year in Nebraska,
and an additional one southward (798).

In 1925, Campoplex frustranae Cushman, a common parasite of the Nantucket
pine tip moth in the East, was liberated in infested plantations in the Nebraska
National Forest. It became established immediately and increased rapidly. By 1930,
parasitism at the original liberation point had reached 80 percent. Unfortunately, the

152

southwestern pine tip moth, Rk. neomexicana (Dyar), which the parasite also attacks
but in which it is unable to develop, increased very rapidly in the area about this
time, and parasitism of R. bushnelli declined. This may have happened because the
parasite deposited too many of its eggs in R. neomexicana.

The subtropical pine tip moth, R. subtropica Miller, occurs throughout the
range of slash pine in the South (998). Slash pine is the preferred host, but longleaf,
loblolly, and Caribbean pines are also attacked. South Florida slash pine is fairly
resistant. Heavy infestations have been recorded in Florida where typical slash pine
has been planted south of its range, whereas naturally regenerated slash pine
seedlings have rarely been infested. Serious losses of grafted slash pine scions in
tree improvement programs have been incurred.

Rhyacionia sonia Miller has been recorded in Ontario, Michigan, and Manitoba.
Its host is sack pine. There is one generation per year and it overwinters as a pupa
beneath trees. Superficially, the adult is similar to that of R. aktita Miller, a species
occurring on the Coastal Plain from Maine to Texas (998). The hosts of R. aktita are
pitch, loblolly, and slash pines. It overwinters as a pupa in infested tips. Both
species have been mistaken for R. frustrana trom which they differ structurally in
the adult stage.

Extensive studies of Rhyacionia pheromones indicate the utility of such com-
pounds for surveying for Rhyacionia and other purposes (99).

The pitch twig moth, Petrova comstockiana (Fernald), occurs from Maine to
North Carolina and west into the Central States. Its hosts include the hard pines, but
pitch pine is preferred in the Midwest. The adult is reddish brown with gray
mottlings and has a wingspread of about 19 mm.

In Ohio, eggs are deposited on twigs during May and June. The larvae bore into
and downward in the twigs for distances from 8 to 10 cm, and pitch masses form
over the entry holes (fig. 53). Winter is spent as a larva under the pitch mass.
Development is resumed in May, and pupation occurs under the mass. There is one
generation per year. Damaged twigs break off, leading to deformation of infested
trees. The braconid Agathis pini (Muesebeck) has been a major factor in control
(S59). A similar moth, P. taedana Miller, attacks loblolly pine throughout the South
(S54).

SAUER

pSirasess

ea

EATEN

F-519519

Figure 53.—Larva of the pitch twig moth, Petrova
comstockiana, in a red pine twig. Pitch mass covers
the opening to the larval gallery.

153

Petrova pallipennis McDunnough feeds on jack pine in the Lake States and
southern Canada. The larvae feed in and hollow out the terminals and adjacent
lateral buds. Winter is spent as a larva in the large central bud. Feeding is resumed
about mid-May and pupation occurs in the bud in early June. Damage to terminals
and lateral buds results in the death of an entire whorl, causing crooked stems and
“stag-headedness.”

The northern pitch twig moth, P. albicapitana (Busck), occurs in all parts of
North America where jack pine grows naturally. Scotch and lodgepole pines are
also attacked. The adult is reddish brown with grayish patches on the forewings,
and has a wingspread of about 18 mm.

Young pines from 0.3 to 1.5 m tall are most heavily infested. Smaller ones are
not attacked and taller ones are rarely injured. although signs of attack may be seen
in trees up to 9 m tall. Larvae feed singly under masses of pitch, about 2 cm in
diameter, generally at an internode or fork. As they develop, their feeding may be
extended almost to the pith. Two years are required to complete the life cycie, and
winter is spent in the larval stage. Pupation occurs under the pitch mass.

When an attack occurs at the base of a growing terminal, the shoot may be
girdled and killed, or the terminal may survive as a weakened. crooked trunk.
Damage in jack, Scotch, and lodgepole pine plantations may be severe. Areas
planted entirely during a 1- or 2-year period suffer much less damage than areas
planted in small blocks over a period of several years (/2/5).

Petrova houseri Miller attacks shortleaf pine from Ohio south to Florida (854).
The adult is dark gray with light- and dark-brown areas and has a wingspread of
about 15 mm. Before its description as a new species (S5/), its damage had been
attributed to the pitch twig moth. The larvae feed in the inner bark of current shoots,
usually girdling them. Pitch blisters, which form over the entrance holes, average
about 1.3 cm in diameter. Usually, there is only one blister per shoot, and it is
situated away from a branch node. Toward the end of summer, the larva bores down
to the pith where it spends the winter. Activity is resumed in the spring, and the
larva tunnels toward the base of the shoot for a distance of about 2.5 cm. Pupation
occurs beneath the blister. There is one generation per year. Infested shoots usually
turn reddish brown, die, and eventually break off. The braconid Agathis pini
(Muesebeck) is an important parasite of the species.

The eyespotted bud moth. Spilonota ocellana (Denis & Schiffermiller), an
introduced species, occurs from coast to coast in the Northern States and southern
Canada and south to North Carolina. Its hosts are hawthorn, larch, laurel. oak, and
several species of fruit trees. The adult is dark, ashy gray, with a large irregular
whitish median band on the forewing and has a wingspread of about 14 mm.

Adults appear in June and early July and lay their eggs on the undersides of
leaves. Young larvae feed for a short time on the leaves and then migrate to young
twigs where they spin tiny, silken hibernacula in which to spend the winter. Feeding
is resumed in the spring on opening buds and unfolding leaves. which are bound
together with silk. Pupation occurs in June in silk-lined cocoons. There is one
generation per year (942). This species is most important as an orchard pest.

The eastern pine shoot borer, Eucosma gloriola Heinrich, is widely distributed
in the Northeastern and Lake States and southern Canada. Its hosts are eastern
white. jack, red. Scotch, Austrian, and Swiss mountain pines, and Douglas-fir. The
adult is coppery red, with two shining, gray transverse bands on the forewings, and
has a wingspread of about 15 mm.

154

Eggs apparently are laid on needle sheaths on new shoots or laterals during May
or early June. Young larvae bore into the pith a few centimeters above the node of
the new shoot and construct tunnels toward the tip several centimeters long.
Infested shoots are eventually girdled from the inside (333). The full-grown larva
bores a hole to the outside through which it escapes and drops to the ground. There
it spins a cocoon in the litter or just below the soil surface in which it pupates.
Winter is spent in the pupal stage (29/).

The eastern pine shoot borer is a serious pest of young jack pines in the Lake
States and Ontario and in eastern white, red, and Scotch pine plantations in New
York and Pennsylvania. Infested terminals bend over or break off. Damaged laterals
turn yellow, then red, and gradually die as the season advances. Infested trees
become bushy after repeated attacks (fig. 54).

Via, a ;
a 24 GAbiad of E g ee a2 ®

F-492898

Figure 54.—Damage to pine terminal by the eastern
pine shoot borer, Eucosma gloriola.

Eucosma monitorana Heinrich, the red pine cone borer, attacks the cones of
various pines in the Northeastern and Lake States and southern Canada. The adult is
reddish brown with tan and silver markings on the forewings and has a wingspread
of about 14 mm.

Young larvae enter the upper parts of second-year cones in June and bore down
the cone axis. They feed on immature seeds until the supply is exhausted and
sometimes sever the cone at the base. A single cone may contain up to 25 larvae.
Final instars sometimes migrate to and enter relatively fresh cones. At first they
bore around the cone for a short distance, then they tunnel toward and into the axis
where they feed on surrounding seed. Mature larvae vacate the cones during July
and drop to the ground to pupate, the stage in which the species overwinters. While
on the ground, the larvae are vulnerable to prescribed burning (855). There is one
generation per year (58).

Eucosma cocana Kearfott, the shortleaf pine cone borer, bores in cones mainly
of shortleaf pine on the Eastern Seaboard (360). The adult is rust colored with

155

shining white markings on the forewings and the wingspread is about 21 mm.
Larvae feed from April to July, dropping to the ground for pupation and overwinter-
ing. There is one generation per year. E. tocullionana Heinrich, the white pine
cone borer, has been recorded from Ontario and New Brunswick, Canada, and
from Wisconsin, Massachusetts, New York, Connecticut, Pennsylvania, Virginia,
North Carolina, and Tennessee. The larvae feed on the cones of various conifers,
such as eastern white pine, spruce, balsam fir, and eastern hemlock (996).

The cottonwood twig borer, Gypsonoma haimbachiana (Kearfott), has been
recorded from Ontario, from the northern tier of States from New York to Michigan
and south through the Midwestern States of Missouri, Oklahoma, Arkansas to
Mississippi, Louisiana, and Texas. Its hosts are listed as various poplars. The adult
is ash gray and has a wingspread of about 15 mm. The basal portion of the forewing
is darker than the apical portion. Full-grown larvae are pale and about 15 mm long.
The head is brownish yellow; the thoracic shield, brownish yellow edged with
brown; and the anal shield, brown or grayish.

Winter is spent mostly as young larvae in silk-covered, shallow pits excavated in
healed-over borer entrance holes, in the margins of corky bark ridges below leaf
bases, or in depressions of leaf scars. Lesser numbers of late instars overwinter in
hollowed-out terminal buds. When the younger larvae resume activity in the spring,
they enter the tender, new shoots to feed and complete their development. When the
older overwintering larvae resume feeding in the spring, they frequently kill the bud
and up to 25 cm of the terminal. There appear to be four or five generations per year
in the Mississippi Delta Region (877).

This is one of the most destructive of the insects that damage young eastern
cottonwood trees. Damaged trees are stunted, have crooked trunks, and produce too
many limbs. This leads to a great reduction in the quality and quantity of merchant-
able pulpwood, saw logs, or veneer from these trees.

Proteoteras moffatiana Fernald feeds on terminal buds of maples. The resulting
disruption in apical dominance causes forking and branching which seriously
affects tree quality of sugar maple in the Lake States (S557). This insect occurs in the
Northern United States and southern Canada from the Atlantic Coast to Wisconsin.
The adult is bright green with irregular, brown forewing markings and has a
wingspread of about 17 mm. There is one generation per year and adults are present
in July and August. In late summer, larvae enter and mine terminal buds in which
they pass the winter. In spring they mine additional buds before shoot elongation is
completed.

Proteoteras aesculana Riley occurs throughout southern Canada and the North-
ern States, south to Tennessee. The adult is olive green with yellow, gray, and black
markings, and has a wingspread of about 14 mm. The larvae bore in the seeds, leaf
stalks, and terminal twigs of buckeye and maple and sometimes cause serious
injury. Seedlings in nurseries have been heavily attacked. Adults emerge in July and
August.

The boxelder twig borer, P. willingana (Kearfott), attacks boxelder and maple
in many of the Northern and Midwestern States and southern Canada. The adult is
white to brownish, marked with streaks, rings, and clusters of yellowish-tan to
black scales; it has a wingspread of about 17 mm. The larva destroys dormant leaf
buds in the fall and early spring. Later in the spring, it burrows in succulent twigs
(fig. 55), causing the formation of spindle-shaped galls (982). These galls become
woody when they dry out. This usually prevents further terminal growth. Severe
damage has been recorded in shelterbelt plantings in the Prairie Provinces of
Canada.

156

F-500805

Figure 55.—Larva of the boxelder twig borer, Proteoteras
willingana, in a boxelder stem.

The spruce bud moth, Zeiraphera canadensis Mutuura & Freeman, occurs
throughout the range of spruce in the Northern United States and Canada (984). In
addition to spruce, it feeds on several other conifers, especially fir. The adult is
light brown and has a wingspread of about 14 mm. The forewings have darker
diagonal markings, and the outer margins are straight. Full-grown larvae are
yellowish or grayish green and about 18 mm long.

Young larvae feed singly in opening buds in the spring or on tender terminal
needles which they web together. Some pupae are found on the foliage of the trees,
but the majority occur immediately below the ground surface. Winter is passed in
the egg stage, and there is one generation per year. Z. improbana (Walker) occurs
all the way across southern Canada and possibly in the northern tier of States. It
appears to prefer larch and spruce, but several other conifers are also infested. The
foregoing two Zeiraphera species were formerly thought to be European but now
they are shown to be specifically distinct from their European counterparts (900).
Z. unfortunana Powell occurs commonly from coast to coast in southern Canada.
Infestations also have been found in Maine. It apparently prefers white spruce.

The maple trumpet skeletonizer, Epinotia aceriella (Clemens), occurs from
southern Canada to North Carolina. Its hosts are principally red and sugar maples,
but it has also been collected on hawthorn and beech. The adult is white with
dustings of gray or brown, and has a wingspread of about 15 mm. Each larva spins a
long trumpetlike tube of silk and frass on the underside of a leaf, causing the leaf to

157

fold around it. It feeds from within this tube and skeletonizes the part of the leaf
covered by the web, causing it to crumple. Damage is usually not very serious
(248). E. solandriana (L.) feeds as a leafroller on quaking aspen, paper birch, and
various other hardwoods. It is sometimes abundant in the Northeast and Ontario.

Epinotia nanana (Treitschke), an introduced species first recorded in North
America in Massachusetts in the early days of this century, now occurs from Maine
to Ohio and Michigan and in Ontario, Quebec, and British Columbia. Its hosts are
various species of spruce, especially white, Norway, and Colorado blue. The adult
is dark, smoky brown and has a wingspread of about 11 mm. The forewing has a
rather blunt apex, which is black and diffused below by a white dash. A blackish
band crosses from the middle of the costa to before the anal angle. There are five
distinct white spots on the costa of fresh specimens, and the wing is also flecked
with whitish scales.

Adults are present during June in Canada and deposit their eggs on spruce
needles. Newly hatched larvae attack the old needles, boring into them and
hollowing them out completely. From the third instar on, they feed in about equal
numbers on both old and new needles. A single larva may feed on several needles .
which it ties together with silk. The winter is spent in the larval stage inside a mined
needle. In the spring, the larva moves to a new needle and continues its feeding.
Before reaching maturity, it may destroy several other needles, all of which are tied
together in bunches appressed to the twig. Pupation occurs in silken cocoons in
hollowed-out needles, in old staminate flowers, on the bark, or in litter on the
ground. There is one generation per year (27S). Washing webs from the trees in the
spring with a strong stream of water before the buds break has been suggested for
control on ornamentals.

Ancylis platanana Clemens occurs commonly on sycamore wherever it grows in
this country. The adult is whitish with pale, reddish forewings and has a wingspread
of about 12 mm. Eggs are deposited along the midribs or larger veins on the
undersurface of leaves during early spring. Young larvae feed on both sides of the
midrib near the base, and spin fine, silken webs over the leaf surface. Older larvae
feed beneath these webs and skeletonize the leaf. The winter is spent in the larval
stage (297). Ancylis spp. feed inside folded leaves of birch in the East, and
populations occasionally become very heavy. Two common species are A. disci-
gerana Walker and A. logiana L.

The spruce seed moth, Cydia strobilella (L.) (formerly youngana (Kearfott)), is
widely distributed in the United States, Canada, and Eurasia. It attacks the cones of
white, red, black, Colorado, Sitka, and Engelmann spruces. The larvae make
tortuous mines near the cone axis, destroying both scales and seeds. White spruce
cones are especially susceptible. The adult is smoky brown with four crossbars of
shining silver and four shining costal spots on the forewings and has a wingspread
of about 9 mm. Full-grown larvae are creamy white and about 10 mm long.

Winter is spent as a full-grown larva within a cone. Most of the larvae pupate in
the spring; some may remain in diapause for | to 2 or more years (/2/0). Female
moths rarely fly; as a result, populations tend to build up on old cone-bearing trees.

The hickory shuckworm, C. caryana (Fitch), occurs in southern Canada and
from the East Coast to Missouri and Texas in the United States. The larvae feed on
hickory nuts and pecans. The adult is smoky black and has a wingspread of about
12 mm. Full-grown larvae are creamy white and about 9 mm long.

Adults appear in early spring (as early as mid-February in Florida) and lay their
eggs on the nuts or foliage. Young larvae bore into and feed inside the nuts, thus

158

preventing future nut development. Winter is spent as a larva in the shucks of fallen
nuts, and there are from one to four generations per year, depending on locality.
Heavy infestations may seriously reduce hickory and pecan nut crops. Gathering
and destroying infested nuts during the winter is helpful in control.

The eastern pine seedworm, C. toreuta (Grote), occurs over almost all the
eastern half of the United States and southern Canada (547). It attacks the second-
year cones of various pines. The adult is gray brown with two prominent black-
bordered silver bands across each forewing and has a wingspread of about 14 mm.
The mature larva is creamy white and about 10 mm long (360).

Adults emerge during late May and early June in the mid-South and during late
June in Ontario and deposit their eggs in crevices on the surface of the cone. Young
larvae bore into the upper part of second-year cones and feed from seed to seed as
they spiral around the cone scales. Infestations appear to be heaviest on open-grown
trees with branches to the ground or on trees in low-density stands (693). Winter is
spent as a full-grown larva in the woody axis of the cone, and pupation occurs in the
spring. A braconid parasite, Phanerotoma sp., may destroy a high percentage of the
full-grown larvae (760).

The slash pine seedworm, C. anaranjada (Miller), occurs throughout the range
of typical slash pine and South Florida slash pine in the South (547). The adult has
a wingspread of about 15 mm. The abdomen is pearl white and the forewings are
yellowish orange to rusty orange, with four more or less equally spaced, mostly
pearl-white crossbands. The species has been reared principally from mature cones
of slash pine, occasionally from longleaf cones, and rarely from loblolly cones
(360).

Eggs are usually laid either singly or in small clusters on second-year cones.
Young larvae bore into the cones and feed on the seed, moving from seed to seed,
consuming their contents, and lining the tunnels between seeds with silk. A single
larva consumes from five to seven seeds. Eventually, the full-grown larva bores into
the woody cone axis where it spends the winter. Pupation occurs in the spring, and
the adults emerge during April and May. There is one generation per year. Accord-
ing to estimates, this species destroys from 2 to 10 percent of the seed in open-
grown slash pine stands in northeast Florida each year (836).

The longleaf pine seedworm, C. ingens (Heinrich), has been recorded from
South Carolina, Georgia, Florida, Alabama, and Mississippi and probably occurs
throughout the range of its favored host, longleaf pine. It also infests slash and
loblolly pines to a minor extent and may occur occasionally on other southern pines
(360). The adult is grayish brown and has a wingspread of about 18 mm. Eggs are
laid in rows of two to nine eggs each on scale apophyses of 1-year-old cones. Young
larvae bore downward through the scales and enter seeds through the micropyle.
Larvae feed during the first two instars, each in a seed, vacating it and tunneling
through the cone in search of another seed. Before it reaches maturity the larva may
consume two to five additional seeds. A full-grown larva bores into the rachis and
usually tunnels toward the base for a distance of 2.5 to 5 cm. Winter is spent in the
larval stage in this tunnel. Pupation occurs in the spring, usually between mid-
March and mid-May, and the adult appears about | to 2 weeks later. There is one
generation per year, except for occasional individuals that enter diapause (255).
This species causes an estimated loss of 21 percent of longleaf pine seed in the Gulf
States region.

The filbertworm, Melissopus latiferreanus (Walsingham), occurs throughout
most of the United States and in southern Canada. The larvae feed in hazelnuts,

159

filberts, acorns, beechnuts, chestnuts, and oak galls. The adult is reddish brown
and has a wingspread of 11 to 20 mm. High percentages of acorn crops may be
destroyed during poor seed years.

Endopiza liriodendrana (Keartott) has been recorded feeding on yellow-poplar in
New Jersey and on magnolia in Florida. The adult is brownish or blackish, and
there is a tuft or mass of erect, dark-colored scales at the rear of the thorax. Eggs are
laid on the leaves from May to September. Young larvae bore into the midvein near
the petiole and feed in mines during the first instar. Older larvae feed under webs
spun on the lower surface of the leaves or between two leaves webbed tightly
together. Pupation occurs under the web (979). E. palliolana (McDunnough)
attacks terminals of eastern larch (892).

The locust twig borer, Ecdytolopha insiticiana Zeller, attacks black locust
throughout the Eastern United States and in parts of southern Canada. It also occurs
in Colorado, Arizona, and California. The adult has a wingspread of about 22 mm.
The forewings are dark, ashy brown with large, dull, pinkish-white patches on their
outer parts with several small, blackish spots near the middle of each of the patches.
Full-grown larvae are reddish to straw yellow with a darker dorsal line, and are
about 16 mm long.

Adults are present from early May to the end of June and again from July to
October. The larvae are twig or stem borers, and cause the formation of elongate
galls up to 7.5 cm long. Winter is passed as a mature larva in a cocoon among the
leaves on the ground. In heavily infested areas, seedling mortality may be high. A
high percentage of the twigs on larger trees also may be damaged.

Other olethreutine moths likely to be encountered in eastern forests and their
hosts are as follows: Hedya chionosema (Zeller)—hawthorn, occasionally red oak
and mountain-ash; the raspberry leafroller, Olethreutes permundana (Clem-
ens)—hickory; O. quadrifidum Zeller—cherry; Pseudosciaphila duplex (Wal-
singham) and Pseudexentera oregonana (Walsingham)—poplar; Epinotia stroe-
miana (F.)—gray birch; E. lindana (Fernald)—alternate-leaved and flowering
dogwood: Griselda radicana Heinrich—various conifers, principally white spruce
and balsam fir: the pecan bud moth, Gretchena bolliana (Slingerland), and G.
concitatricana (Heinrich)—black walnut.

Subfamily Tortricinae

Members of this subfamily eat the foliage of a wide variety of coniferous and
deciduous trees of all sizes and ages. Many species are important pests, some
extremely so. The larvae either fold or roll individual leaves or parts of leaves or tie
several leaves or shoots together forming enclosures in which to rest and feed or
from which they move out to feed. The adults are usually small and have wide,
oblong, fringed wings that appear bell-shaped while at rest. The wingspread is
about 25 mm. The larvae are usually some shade of green, seldom more than 25
mm long, and they pupate in flimsy silken cocoons. Just before the pupa transforms
to the adult, it works its way partly out of its cocoon. Information about many
species has been summarized (446, 794).

Adoxophyes furcatana (Walker), a leafroller on sycamore, occurs from New
England to Pennsylvania and the Mississippi River Valley. The adult is straw yellow
and has a wingspread of 20 mm. The forewings are marked with five golden-brown
lines and two irregular light-brown bands. The hindwings and their fringes are
shiny white. Full-grown larvae are light green, taper toward each end, and are about
18 mm long.

160

Amorbia humerosana Clemens occurs from coast to coast in southern Canada
and in the Northeastern States south to Pennsylvania. Its hosts include a wide
variety of tree species, both coniferous and deciduous. Adults are light gray and
have wingspreads of about 25 mm. Larvae are light green except for light-brown
heads.

Sparganothis acerivorana (MacKay) occurs in southern Canada and the Lake
States. In Canada it feeds on several hardwoods, mostly sugar and red maples, and
also on young Scotch and red pines. In the Lake States, it usually feeds on sugar
maple. The adult is reddish yellow with reddish-brown spots and has a wingspread
of about 25 mm. Full-grown larvae are yellowish green and about 21 mm long.
Winter is apparently spent in the egg stage, and hatching occurs during May of the
following spring. Each larva rolls a leaf in which to rest and from which it moves
out to feed. Pupation occurs inside the roll.

Defoliation by this species was an important factor leading to the development of
maple blight, a condition responsible for the deterioration and death of large
numbers of sugar maple trees in Wisconsin (483).

Sparganothis pettitana (Robinson) occurs in southeastern Canada and south and
west to Florida and the Mississippi River Valley. The larvae are solitary leafrollers
on various hardwoods, especially basswood and maples. The adult is lemon yellow
and has a wingspread of about 22 mm. The forewings are sometimes marked with
two oblique lines of light-brown scales, and the hindwings are white. Full-grown
larvae are dull, yellowish green, with reddish-brown heads and brown to blackish
cervical shields.

Sparganothis sulfureana (Clemens) has been observed feeding in the tips and
shoots of small red pines in Canada and in the terminals of loblolly pine seedlings in
Georgia. Willow, honeylocust, apple, and cherry are also attacked. S. reticulatana
(Clemens) occurs over about the same range as S. sulfureana. It feeds on white ash,
maple, bigtooth aspen, and paper birch. S$. diluticostana (Walsingham) occurs
commonly on oak in New Jersey and Maine. It has also been recorded feeding on
white ash and paper birch in southern Canada. S. fristriata (Kearfott) occurs fairly
commonly on jack pine in southern Ontario. It probably occurs in the Lake States
also.

The genus Pandemis contains several species that attack a wide variety of
hardwoods in eastern North America. P. /Jamprosana (Robinson) feeds on beech,
paper birch, red and sugar maples, oak, elm, basswood, ironwood, and sassafras
from Maine to New Jersey and in southern Ontario and Quebec. The threelined
leafroller, P. /imitata (Robinson), occurs from coast to coast in Canada. It has also
been recorded from Maine, Pennsylvania, Georgia, Illinois, and Arkansas. Its hosts
include paper birch, willow, oak, basswood, elm, boxelder, alder, and cherry.

Archips rosanus (L.), an introduced species, occurs in southeastern Canada,
British Columbia, and from New England to the Lake States. The larvae are general
feeders on a number of deciduous species. Privet appears to be especially attractive.
Adults are dull light-brown to olive-brown and have wingspreads of about 20 mm.
Full-grown larvae are dull green and about 18 mm long. The larvae tie together two
or more leaves at the tips of twigs or branches and feed on them during May and
June.

The uglynest caterpillar, A. cerasivoranus (Fitch), occurs from coast to coast in
the Northern States and southern Canada. Its preferred hosts are common
chokecherry and black cherry, but it may also be found on a wide variety of other
hardwoods. The adult is dull orange and has a wingspread of about 21 mm. The

161

forewing is irregularly speckled with dark reddish-brown and has three patches of
the same color. The hindwing is bright orange. Full-grown larvae are yellowish or
greenish yellow except for black heads and cervical shields, and are about 20 mm
long.

Winter is spent in the egg stage. Hatching begins in May, and larvae are present
until September, depending on location. They live together in dense nests they
construct by webbing twigs and leaves together (fig. 56). In heavily infested areas,
these nests are often numerous. Some may be large enough to entirely enclose small
trees.

The uglynest caterpillar is usually of little or no economic importance because
the trees attacked are usually of low value. However. the presence of its webs on
roadside vegetation may be objectionable because of their unsightliness. Cutting
and removing nests provides adequate control in most situations.

The oak webworm, A. fervidanus (Clemens), occurs throughout the oak regions
of the Northeastern and Lake States and in various parts of southern Canada. Its
favored hosts appear to be scrub and bur oaks, but it also feeds on the seedlings and
sprouts of several other oaks. The adult is brownish and has a wingspread of about
21 mm. The forewing is yellowish brown with dark patches. The hindwing is
uniformly smoky except for a light-colored fringe marked with a fine, basal line.
Full-grown larvae are grayish green and about 20 mm long. Larvae live together in
webs. some of which may be 8 cm wide and up to 0.5 m long (fig. 57).
Occasionally, they are large enough to enclose all of the leaves at the top of a tree.
Winter is spent in the egg stage. Larvae appear in midsummer, and pupation occurs
within the nest. The species is of little economic importance.

F-519520

Figure 56.—Nest of the uglynest caterpillar, Archips
cerasivoranus, on wild cherry.

162

The fruittree leafroller, A. argyrospilus (Walker), occurs throughout the United
States and from coast to coast in southern Canada. Its hosts include many fruit trees
and many forest and shade trees such as ash, hickory, elm, oak, maple, walnut,
poplar, birch, basswood, and buckeye. The adult is pale yellow to orange-red and
has a wingspread of about 21 mm. The forewings are mottled with golden scales,
their tips triangular. The hindwings are fuscous and have dirty-white fringes. Full-
grown larvae are light green and about 22 mm long.

Adults are present from June to August, depending on location. Eggs are
deposited in small, round or convex masses containing about 100 to 150 eggs each,
usually on twigs or small branches. Winter is spent in the egg stage, and hatching
occurs in early spring. Young larvae feed on opening buds, blossoms, young fruit,
and unfolding leaves which they web together with silk. Later, several leaves may
be webbed together, forming a nest in which the larvae live and from which they
move out to feed. Pupation occurs in flimsy cocoons spun inside the nest or on the
branches or trunk of the tree. A number of outbreaks of this species, some of which
covered tens of thousands of hectares, have occurred in oak stands in Eastern and
Lake States.

Archips semiferanus (Walker) defoliated oaks on more than 400,000 hectares in
midcontinent areas (375, 1/330). Northern red, scarlet, and northern pin oaks of all
ages are affected. The adult is light olive brown with a diagonal rusty band on the
forewing; the wingspread is about 22 mm. Mature larvae are greenish yellow and
about 21 mm long. Masses of I5 to 125 eggs covered with tan scales from the
female’s abdomen are deposited on the bark of tree crowns during July and early
August. Hatching occurs the following spring, in mid-May, about the time of

Courtesy Conn. Agric. Exp. Stn.

Figure 57.—Nest of larvae of the oak webworm, Archips
fervidanus.

163

budbreak. At first, larvae feed within the protective folds of young leaf clusters and
later roll or fold one or more entire leaves or leaf parts. Pupation takes place in leaf
rolls in June and adults appear in July.

Many other species of Archips are also encountered in eastern forests (446). A
few of these and some of their hosts are as follows: A. infumatanus (Zeller) —
hickory and pecan, A. negundanus (Dyar)—boxelder, A. rileyanus (Grote)—
hickory and walnut, A. georgianus (Walker)—oak, A. griseus (Robinson)—oak
and hickory, A. magnolianus (Fernald)—cucumbertree, and A. purpuranus (Clem-
ens)—basswood, paper birch, quaking aspen, willow, and black cherry.

The spruce budworm, Choristoneura fumiferana (Clemens) (fig. 58). occurs in
North America from Virginia to Labrador and westward to the MacKenzie River
Valley, Yukon Territory (//50). At one time, it had been considered present
throughout the ranges of spruce and fir in North America. Forms occurring in
western Canada and the Western United States are now considered different species
(448, 997): however, the question of speciation does not appear to be settled (997,
1165). Balsam fir is the preferred host of the spruce budworm. To a lesser degree, it
also feeds on white, red, and black spruces, and on larches, pines, and eastern
hemlock (698).

Spruce budworm adults are mostly gray, sometimes ochreous gray. The male has
a wingspread of about 24 mm; the female, about 26 mm. Full-grown larvae are
about 20 to 23 mm long. The head is usually almost entirely dark brown, the
prothoracic shield is brownish yellow with some diffusion of brown pigment or is
entirely dark brown, and the anal shield is brownish yellow. Pupae are light to
reddish brown except for darker bands and spots.

Spruce budworm adults are active from late June to early August, depending on
location, and the females deposit their eggs in elongate masses of 2 to 60 eggs each,
the eggs overlapping like shingles on a roof. A large proportion of the masses are —
usually found on needles near the periphery of the crown. The eggs hatch in about
10 days under normal conditions. After a period of | to 2 days, during which larvae
are dispersed by wind throughout the tree and stand, the larvae spin hibernacula in
suitable sites and molt to the second instar. While the majority of hibernacula are
found on the branches of host trees, some are also found in mined buds, in flower
scars, under bark scales, or under lichens. Many larvae become dislodged while
searching for overwintering sites and drop down on silken threads. At this time,
they may be blown considerable distances by the wind, often into uninfested stands.

In the spring, after several days of warm weather but before balsam fir buds
expand, the overwintering larvae emerge and start to feed. The new buds of
staminate flowers are attacked first if present; otherwise, the larvae bore into old
needles. After a short period of feeding here, the larvae move to the ends of
branches and bore into expanding, vegetative buds. Later, they feed on the new
foliage of developing shoots. When about half grown, they begin tying the tips of
two or more twigs together with silk, forming a small nest. During this period, old
needles are avoided until all of the new ones are eaten or cut through. Feeding is
usually completed during late June or early July. Pupation usually takes place
within the last-formed nest but may occur at twig axils. Adults appear in about 10
days and are subject to considerable dispersal by the wind. The female does not fly
until she has deposited one or two egg masses. Wind dispersal may be the most
important factor influencing population trends in any given area (503, 582).

In light or moderate infestations, spruce budworm injury is restricted to the
partial loss of new foliage, especially in the upper portion of the crown. Damaged

164

A,B,C,E, courtesy Can. For. Serv., Can. Dep.
: Environ., Sault Ste. Marie, Ont.
D, courtesy S. J. Krieg, Emporium, Pa.

Figure 58.—Spruce budworm, Choristoneura fumiferana:

A, moth; B, egg masses: left—unhatched; center—

shows hatched and unhatched as well as hatching,

with completely hatched mass to the right; right—egg

mass parasitized by Irichogramma minutum, C, instar

ll; D, final instar; E, pupae.

165

needles on webbed branch tips turn reddish brown by midsummer. In heavy,
persistent infestations, all of the new foliage may be consumed (fig. 59) for several
successive years, and opening vegetative buds and developing shoots may be killed
in their formative stages. Top-kill usually occurs after about 3 years of severe
infestation, and tree mortality, after about 5 years. In sustained outbreaks, nearly
complete mortality of the merchantable volume of balsam fir may occur by the 8th

year (89, 799). During widespread outbreaks, the magnitude of these losses is
great.

F-492901

Figure 59.—Balsam fir defoliated by the spruce
budworm, Choristoneura fumiferana.

Much has been learned about the factors or conditions conducive to the develop-
ment and decline of spruce budworm outbreaks. For example, it was determined
many years ago (//82) that outbreaks generally begin in extensive and continuous
areas of mature and overmature balsam fir. Later, it was learned that even when
these conditions prevail, at least three or four summers of clear, dry weather are
necessary for populations to explode (502, 985, 1271). Much has also been learned

166

about factors tending to hold populations in check during intervals between out-
breaks or which assist in bringing them under control once they are underway (/27,
882).

The spruce budworm has many natural enemies, including parasites and preda-
tors, birds, mites, spiders, and several pathogenic organisms. Their effectiveness in
control is always important, but it is greatest only after another agent, such as
weather or starvation, has reduced the budworm population considerably (548). A
number of authors have evaluated the control effectiveness of natural enemies in
specific outbreaks (/24, 126, 329, 330, 636, 787).

Other important natural control factors affecting the spruce budworm are: (1) the
loss of young larvae through competition for new foliage in heavy infestations; (2)
loss of young larvae as a result of cooler than normal temperatures and late frosts in
the spring; (3) loss of large larvae through starvation following complete defolia-
tion; and (4) loss of adults through dispersal.

A number of management practices have been suggested for reducing the
chances of spruce budworm outbreaks. These include the utilization of balsam fir
(125, 1182); the regulation of age classes to prevent the occurrence of large areas of
overmature fir (125, 498), and the favoring of less susceptible species, such as red
spruce. Discussions that favor forest management oversimplify the special rela-
tionships between the budworm and balsam fir (66).

The removal of overstory mature balsam fir was suggested for preventing out-
breaks (806, 1277); management plans and risk-rating systems for selective cutting
were presented (807, SO8, 1278); and risk-rating systems for the Lake States were
devised (69, 70, 78, 495). Publications on the budworm and forest management in
the Maritime Provinces of Canada are available (46, /95).

For further information on the spruce budworm, the reader is referred to reviews
of the literature (66, 635, 797).

The jack pine budworm, C. pinus pinus Freeman, is known to occur in Nova
Scotia, Ontario, Manitoba, and from Michigan to Minnesota. Its hosts include
various species of pine, especially jack and red. The adult has a wingspread of
about 22 mm. The head, thorax, and forewings are ochreous-tawny; the forewings,
distinctly maculate; and the hindwings smoky with dark basal lines through their
white fringes. The full-grown larva has a shiny light-brown to black head, a dark-
brown “‘collar” separated from the head by a narrow white band, a reddish-brown
body with yellowish sides, and two rows of white dots along the back; it is about 21
mm long (fig. 60). Pupae are pale green when just formed; later, they become dark
reddish-brown, and they are about 12 mm long.

Courtesy Can. For. Serv., Can. Dep. Environ.,
Sault Ste. Marie, Ont.
Figure 60.—Larva of the jack pine budworm,
Choristoneura pinus pinus, feeding on needles of jack
pine.

167

Jack pine budworm adults are present from early July to early August and lay
their eggs in clusters of about 40 in two or three rows on the flat side of a pine
needle. Hatching occurs in about 10 days. A few days later, the young larvae,
without feeding, spin hibernacula under bark scales on the trunk or larger limbs. or
between needles. Then they molt to the second instar, the stage in which they
remain throughout the rest of the summer, fall, and winter. In the spring, about the
time the staminate flowers are shedding their pollen, they emerge and begin feeding
on the pollen. Some usually remain in the flower clusters throughout the entire
feeding period, but the majority migrate to new foliage on which they feed. once it
is well developed. The needles are not consumed entirely, but are usually clipped
off at the base and webbed together. Pupation occurs among the needles or between
webbed shoots.

The jack pine budworm usually does not cause heavy mortality of merchantable
jack pine, but it may cause top-kill and “‘stag-headedness.** During outbreaks,
however, heavy losses in pole-size trees, saplings, and reproduction may result. In
heavily infested stands, young understory red and eastern white pines also are often
severely defoliated and killed. Fortunately, outbreaks usually last only 2 to 4 years.

Cutting practices designed to remove the jack pines that most commonly produce
staminate flowers are helpful in preventing outbreaks (580). These trees are usually
coarsely branched and large-crowned, or suppressed and slow growing. Growing
hard pines in fully stocked stands or in groups, eliminating large-crowned “wolf”
trees, utilizing trees before they become mature, and encouraging species suited to
the site are also recommended practices (674). Parasites and a polyhedral virus
disease aid in the control of infestations following periods of heavy defoliation.
Rapid declines in populations have also been attributed to decreases in staminate
flower production.

A subspecies, C. pinus maritima Freeman, has been found feeding on Virginia
and pitch pines in Pennsylvania, Massachusetts, New Jersey, and Kentucky. Adults
are larger and redder than those of the jack pine budworm (448, 997).

The large aspen tortrix, C. conflictana (Walker), occurs throughout much of the
range of quaking aspen in Canada and the United States (/288). Several other
hardwoods also serve as hosts such as balsam poplar, bigtooth aspen, paper birch,
willow, and alder. The adult is dull, light gray and has a wingspread of about 28
mm. Full-grown larvae are usually dark green, sometimes almost black, and are
about 16 mm long. The prothoracic shield is reddish brown to black: the thoracic
legs, black.

In Canada, eggs are laid in flat clusters usually on the upper surfaces of leaves in
June or July. The first instars feed gregariously on leaf surfaces during July,
spinning much silk and webbing the surfaces together (fig. 61). Later they move to
the trunk in search of hibernation sites in rough bark or under moss. Here they molt
and spend the winter as second instars. The following spring, they climb the trees
and mine the swelling buds. Later, they roll leaves and feed within the enclosures.
Pupation occurs within the rolls, and adults begin to emerge in June (/00/). Many
outbreaks have occurred in aspen stands in Canada. Serious defoliation has also
been recorded in New England, New York, and Michigan.

The obliquebanded leafroller, C. rosaceana (Harris), occurs in southern Can-
ada and throughout most of the United States. It is recorded as a general feeder on
the foliage of deciduous trees and shrubs. In Canada, it occurs most frequently on
quaking aspen, paper birch, and willow but has also been observed feeding on
Scotch pine. In New York, it has seriously damaged coniferous seedlings in

168

F-50674

Figure 61.—Pupae of the large aspen tortrix,
Choristoneura conflictana, in webbed leaves.

nurseries (/088). Seedlings most seriously affected were those of eastern white,
Scotch, and red pines. The infestation is believed to have resulted from an invasion
of the nursery by larvae developing on weeds or other vegetation surrounding the
nursery. The adult is reddish brown and has a wingspread of about 25 mm. The
forewings are marked with three dark-brown, oblique bands. Full-grown larvae are
greenish and about 9 mm long.

Winter is spent as young larvae in tightly woven cases under bud scales or loose
bark, or between leaves. The following spring the larvae feed first on the surface of
unfolding leaves. Later, each larva ties two or more leaves together with silk and
feeds from within the case. Adults may appear as early as June, and there may be
two generations per year, depending on location.

Choristoneura fractivittana (Clemens) feeds on sugar maple, beech, paper birch,
red maple, elm, and red oak in southern Canada and from Massachusetts to
Wisconsin and Colorado. It 1s sometimes mistaken for C. rosaceana.

Cudonigera houstonana (Grote) attacks various species of Juniperus, especially
eastern redcedar, and is a pest in windbreak and ornamental plantings in western
Kansas (554).

The redbanded leafroller, Argyrotaenia velutinana (Walker), occurs in southern
Canada, mostly in the southeastern part, and throughout the Eastern United States,
westward to lowa, Missouri, and Texas. In Canada and Maine it occurs commonly
on various conifers, especially white, red, and black spruces, balsam fir, and larch.
In the United States, !t occurs on a wide variety of deciduous trees. Damage is often
serious in apple orchards. The adult has a wingspread of about 14 mm. The
forewing is marked with a band that widens as it runs from the middle of the costa
to the outer third of the inner margin. Full-grown larvae are pale green and about 16
mm long. Winter is spent in the pupal stage among leaves and debris on the ground,
and there may be three or four generations per year. A granulosis virus disease that
tends to retard larval development has been reported in Virginia. Eggs are some-
times heavily parasitized by Trichogramma minutum Riley.

The pine tube moth, A. pinatubana (Kearfott), feeds on various pines, mostly
eastern white pine, in southern Canada, the Northeastern States, Florida, and
Louisiana. The adult is small, slender, and grayish; it has a wingspread of about 14
mm. The forewings have broad, orange to reddish-ochreous patches and are crossed
by two whitish, oblique lines; the hindwings are smoky; and the abdomen is gray,
blackish, or mouse colored with ochreous, apical tufts. Full-grown larvae are 12
mm long.

169

The larva lives within the tube it constructs by drawing from 5 to 20 needles
together and fastening them with silk. It feeds on the tips of these needles (fig. 62).
Winter is spent in the pupal stage within the tube. There may be two generations per
year. Heavily infested pines often have a ragged appearance. This may be objec-
tionable where valuable ornamentals are involved.

Courtesy Conn. Agric. Exp. Stn.

Figure 62.—Larval tube and larval feeding damage by
the pine tube moth, Argyrotaenia pinatubana.

The hickory leafroller, A. jug/andana (Fernald), occurs in southern Canada and
throughout the Eastern States. Its principal host is hickory, but it will feed on plum
and viburnum. The adult is dark brown and has a wingspread of about 21 mm. The
front wings are marked by two parallel, oblique, blackish bands. Full-grown larvae
are pale to translucent, with pale green heads tinged with brown; larvae are about 20
mm long. They feed from within longitudinally rolled leaves and pupate beneath
the bark on the trunk of the tree.

Argyrotaenia quercifoliana (Fitch) occurs in southern Canada and south to
Florida and Texas. The larvae feed on the foliage of red, black, white, scrub, and
pin oaks. The adult is cream yellow marked with light-brown dots and has a
wingspread of about 20 mm. The forewings are marked with two oblique brown
bands, and the hindwings are white. Full-grown larvae are light green except for
amber-yellow heads and are about 20 mm long. This species sometimes causes
serious defoliation locally.

170

Several other species of Argyrotaenia are recorded as feeding on eastern trees
(446); A. occultana Freeman—on spruce and occasionally balsam fir and larch in
Canada and New York; the graybanded leafroller, A. mariana (Fernald)—on
paper birch, willow, chokecherry, elm, pear, Vaccinium, and possibly oak from
eastern Canada to Florida; A. quadrifasciana (Fernald)—on hawthorn, service-
berry, plum, and pear in southeastern Canada and from Maine to Missouri; and A.
alisellana (Robinson)—on oak from southern Canada to Florida.

The oak leaftier, Croesia semipurpurana (Kearfott), feeds on various oaks from
southeastern Canada and Massachusetts to Minnesota and Texas. Adults have
wingspreads of about 12 mm. The forewings vary in color from almost solid yellow
to yellow with dark-brown markings. The full-grown larva is dirty white to light
green except for a pale head and brown to black thoracic legs, and is about 12 mm
long.

Eggs are laid individually on the bark of second-year wood of branches in late
June and early July and hatch in April of the following year. Newly hatched larvae
enter unopened buds and feed on the young leaves. Older ones fold together
sections of leaves and feed inside the folds. When they reach maturity in May, they
spin down to the ground and pupate in the litter. Adults appear | to 2 weeks later.

Serious outbreaks have occurred in the Northeastern and Middle Atlantic States
(210). During 1964 and 1965, approximately one-quarter million hectares of red
oaks were severely defoliated in Pennsylvania alone, resulting in considerable tree
mortality.

The eastern blackheaded budworm, Acleris variana (Fernald), occurs from
Cape Breton Island and the northeastern corner of the United States across the
coniferous forest region of Canada to Saskatchewan and eastern Alberta. (The
western blackheaded budworm occurring farther west 1s now considered to be a
different species, A. gloverana (Walsingham) (995).) Balsam fir is its preferred
host, but during epidemics, white and sometimes red and black spruces and
hemlock may also be defoliated (770). The adult is predominantly mottled gray
with various brown, white, or gray ragged bands across the wings. Some indi-
viduals have a white, yellow, or orange stripe down each wing. The wingspread ts
about 19 mm. Full-grown larvae are bright green and about 14 mm long.

Adults appear during August and September and deposit their eggs singly on the
undersides of needles, mostly on the upper branches. The winter is spent in the egg
stage. Hatching occurs in the spring, and the young larvae burrow into the expand-
ing buds. As the new needles grow, the larvae web together a few of them and feed
within. Once the new needles are devoured, the larvae feed on the old ones.
Pupation occurs within webbed masses of partially eaten and damaged needles.

Extensive outbreaks tend to occur at intervals of 10 to 15 years in stands where
maturing balsam fir is dominant, but they usually subside before many trees are
killed.

Acleris chalybeana (Fernald) has been recorded from Ontario, Quebec, Wiscon-
sin, Maine, New York, and Pennsylvania. The larvae feed on the foliage of sugar,
red, and mountain maples, yellow birch, beech, and eastern hophornbeam. The
adult is grayish and has a wingspread of about 21 mm. Full-grown larvae are light
green and about 19 mm long. In Wisconsin, apparently they winter as first or
second instars in hibernacula on the twigs. During June, the majority of them were
found in rolled leaves previously occupied by larvae of another tortricid, Spar-
ganothis acerivorana; the remainder rolled their own leaves. Pupation occurs inside
the roll. Damage by this species appears to have been one of the factors contributing

171

to the development of maple blight, a condition leading to the deterioration and
death of sugar maples and saplings (483). A. logiana (L.) skeletonizes the leaves of
paper and river birches. Larvae are dull green, with the head, cervical shield, and
front legs black; they have black warts on the prothorax. They usually feed singly
inside folded leaves. Sometimes they are found between two leaves folded together.
A. tripunctana (Hubner) feeds on paper birch.

Aphelia alleniana (Fernald) normally feeds on weeds and clovers, but may also
attack and injure small seedlings in coniferous plantations. Seedling losses have
been severe in the Lake States and southern Canada. The larvae tie the shoots of the
seedlings together and feed from within the sheath on the stems and new needles
during May and June. Injured seedlings become twisted and deformed. Winter is
spent in the larval stage, and there is one generation per year.

Xenotemna pallorana (Robinson) also normally feeds on weeds and clover, but it
also occasionally damages seedlings in young coniferous plantations in the Lake
States and southern Canada. It is widely distributed in the Eastern United States,
occurring from New England to the Lake States, Missouri, and Texas. The larvae
pull the young shoots of seedlings together, fastening them with loose silk. Then,
they feed on the young needles and tunnel into the shoot. Heavy damage to young
eastern white, red, jack, and Scotch pines has been recorded during May and June
in Michigan. There are two generations per year in Ontario. Winter is spent as

larvae in hibernacula spun within folded leaves (8/8).
Family Cochylidae
Cochylids

The moths of this family resemble those of the subfamilies Olethreutinae and
Tortricinae. Only one eastern species 1s worthy of mention.

Aethes rutilana (Hubner) is an introduced species first recorded in this country in
1878. Its present distribution seems to be limited to southern Canada and from New
England to New Jersey and Indiana. Its host plants are various junipers, especially
common juniper on which it is often abundant. The adult is yellowish and has a
wingspread of about 10 mm. There are red markings on the head, thorax, and
forewings, those on the wings occurring as four broad crossbands. The larvae spin
webs on the foliage, tying the needles together and forming tubes in which they live
and feed. Pupation takes place on the tree in the webbing. The foliage of heavily
infested trees may turn brown.

Family Hesperiidae
Skippers

Members of this family are commonly known as skippers because of the way the
adults flit or dart from place to place. They are distinguished by the head that is
nearly as wide or wider than the thorax, and the antennal club that usually ends in a
recurved hooklike apiculus. The larvae usually have large heads and strongly
constricted necks. They are also usually solitary, each one concealing itself under
part of a leaf that it cuts and folds over.

The silverspotted skipper, Epargyreus clarus (Cramer), one of the largest
species in the family, is widely distributed throughout the United States and
southern Canada, and the larvae feed on black locust and wisteria. Adults are brown
except for yellow and white triangular spots on the forewings. The forewings are
elongate, and the hindwings have rounded tips. Full-grown larvae are nearly 50 mm
long. The body is leaf-green, the head dull red except for two yellow spots on the
lower part of the face, the neck and sides of the first thoracic segment are red, and
the cervical shield is black. The body is also marked with dark rings.

172

The larvae feed from within nests made by tying several leaves together with
silk. Sometimes they cause heavy defoliation locally. Pupation takes place in loose
cocoons spun among the leaves, usually on the ground. In the South there are two
generations per year, farther north there may be only one generation or one and a
partial second.

Family Papilionidae
Swallowtail Butterflies

Swallowtail butterflies are of considerable interest to many people because of
their large size and striking appearance; otherwise, they are of minor importance.
The adults are distinguished by the wavy margins and taillike prolongations of the
hindwings; the larvae, by the protrusile, bright-colored, forked processes rising
from the first thoracic segment. These processes also emit a disagreeable odor
when the caterpillar is disturbed.

The tiger swallowtail, Papilio glaucus L., commonly occurs in eastern North
America. The caterpillars feed on various deciduous trees such as ash, birch,
basswood, cherry, and poplar. Full-grown larvae are dark green and about 37 mm
long. The third thoracic segment is enlarged and marked on each side by a large
yellow spot. This spot is edged with black and encloses a small purple spot that is
also edged with black. The distal part of the first abdominal segment bears a
transverse, yellowish ridge, edged posteriorly with black. The caterpillar spins a
silken mat upon the surface of the leaf which usually causes the leaf to fold
lengthwise. Resting caterpillars are found inside this fold. Transformation to the
chrysalis usually takes place on some object above the ground.

The spicebush swallowtail, P. troilus L., occurs throughout the eastern part of
the United States and its principal food plants are spicebush and sassafras. Full-
grown larvae are about 37 mm long. The body is widest at the third thoracic
segment. The head and venter are pink, the dorsum pea-green, the sides yellowish,
and there is a transverse black line on the prothorax. The third thoracic and first
abdominal segment each bears two orange spots. Those on the thorax have black
centers. Six small spots are on second and seventh abdominal segments and four on
the eighth. P. cresphontes Cramer feeds on prickly-ash in the Northern States and
on citrus in the South.

Graphium marcellus (Cramer), the zebra swallowtail, feeds on pawpaw.
Family Nymphalidae
Brushfooted Butterflies

This family contains some of our most common butterflies. The adults are
medium to large and are distinguished by having the forelegs much reduced and
without claws. Only the middle and hind pairs are used for walking. The head of the
caterpillar is usually bilobed, the tips of the lobes often supporting branched spines,
and the body is spiny or bears fleshy, hair-covered warts. The chrysalids are naked
and are usually suspended by the cremaster.

Polygonia interrogationis (F.), the question-mark, feeds on elm, basswood, and
hackberry in eastern America, especially on sprout growth along roadsides. Full-
grown caterpillars (fig. 63) are brownish with yellow mottlings and are about 37
mm long. Each body segment bears a transverse row of light-colored branched
spines. There are two generations per year. P. comma (Harris), the comma, occurs
on elm, nettle, and hops from Canada to the Carolinas and Texas. Full-grown
caterpillars are yellowish white, and each body segment bears a transverse row of
branched spines.

173

F-519524

Figure 63.—Larvae and chrysalis of the butterfly,
Polygonia interrogationis, the question-mark.

Nymphalis vau-album (Denis & Schiffermiller), the Compton tortoiseshell,
feeds principally on gray and paper birches, but also on poplar and willow in
Canada and south to Pennsylvania in the Eastern States. Caterpillars have black
heads, the body is reddish to blackish on the dorsum with dots of light green, and
each segment bears a transverse row of branched spines.

The mourningcloak butterfly, NV. antiopa (L.), is a widespread species, occur-
ring throughout the subarctic regions of North America. The larvae, commonly
known as spiny-elm caterpillars, feed on elm, willow, poplar, and hackberry, and
are sometimes abundant locally, especially on shade and ornamental trees and along
fence rows. Adults are black-bodied and have wingspreads of 60 to 80 mm. The
upper wing surface is dark reddish-brown except for a broad, creamy-yellow border
that contains a row of blue spots. Full-grown caterpillars (fig. 64) are black, with a
scattering of white dots and a red dot on the dorsum of abdominal segments one to
seven. The head is covered with tubercles: the body, with many large, branched
spines.

Courtesy Rocky Mt. For. & Range Exp. Stn.
Figure 64.—Larva of the mourningcloak buttertly,
Nymphalis antiopa.

174

Winter is spent in the adult stage, and the adults appear in early spring. Eggs are
deposited in clusters around small twigs. The larvae feed gregariously until almost
full grown and usually defoliate one branch before moving to another. Chrysalids
are formed in June or early July. Adults soon appear and lay the eggs for a second
brood. Larvae of this generation are present until September. They then pupate and
the adults emerge to hibernate. There are one or two generations per year, depend-
ing on location.

The mourningcloak butterfly 1s of minor importance in the forest, but is some-
times injurious to shade and ornamental trees. Infestations can be controlled by
cutting and burning infested twigs and small branches.

The viceroy, Basilarchia archippus (Cramer), occurs Over most of the United
States and feeds on poplar and willow. The adult resembles the well-known
monarch butterfly, Danaus plexippus (L.), but differs by being slightly smaller,
having a narrow black line across the hindwings and only a single row of white
spots in the black marginal band of the wings. The full-grown caterpillar is about 37
mm long. The head is large, pale green, and bilobed. Body segments one and two
are pinkish to brownish; segments three to six and the sides of seven are brownish or
greenish; the tops of segments seven and nine and nearly all of eight are pale
pinkish or whitish; and the top of nine and nearly all of the last three are brownish or
greenish. There are two barbed, club-shaped, brown tubercles on top of the second
thoracic segment and two smaller ones armed with spines on the top of the other
segments. There are two generations per year. The two related species, B. arthemis
(Drury), the white admiral, and B. astyanax (F.), the red spotted purple, are
found on poplar, willow, birch, black cherry, apple, and basswood. The caterpillars
are similar in appearance to those of B. archippus.

Asterocampa clyton (Boisduval & LeConte), the tawny emperor, occasionally
seriously defoliates hackberry in the Lake States (7/4). Full-grown caterpillars are
25 to 37 mm long. The body is greenish except for a yellow stripe down the back
and a deep biue, yellow-bordered stripe on each side. The head is armed with
branches, antlerlike spines, and there are two projections at the posterior end of the
body. A. celtis (Boisduval & LeConte), the hackberry butterfly, is also occasion-
ally abundant on hackberry in the Lake States. The caterpillar is greenish with a
row of yellow dots down the back and three yellow lines along each side. Other-
wise, it resembles the caterpillar of A. clyton.

Family Megalopygidae
Flannel Moths

The bodies of flannel moths are covered with dense coats of scales and long
crinkly hairs. The larvae are also densely covered with long soft hairs, with an
intermingling of venomous spines. Females deposit their eggs in small batches,
usually on leaves, and cover them with hairs from the abdomen.

The puss caterpillar, Megalopyge opercularis (J. E. Smith), occurs throughout
the Southern States where it feeds on various deciduous trees and shrubs. Forest and
shade trees commonly infested include oak, elm, hackberry, maple, and sycamore.
The adult moth is yellowish brown, with brownish spots on the wings, and has a
wingspread of about 25 mm. The wings bear long, wavy, white hairs, especially
along the veins. The larvae are densely clothed with long yellow and reddish-brown
or mouse-gray hairs. The hairs on the rear end are taillike.

Young larvae feed gregariously on the surface of the leaf and skeletonize it. Older
larvae devour the entire leaf. Serious infestations have been recorded in Florida and
Texas. Several thousand hectares of turkey oak were defoliated during an outbreak

175

in Florida in 1966. Generally speaking, however, the species is most important as a
pest of people, because of its poisonous spines (//4). There may be two generations
per year in the more southerly portions of its range. Winter is spent as a pupa in a
cocoon spun some place on the host tree. Handpicking larvae, wearing gloves, is a
common control practice. The tachinid parasite, Carcelia lagoae (Townsend), is
sometimes abundant in Texas infestations.

The crinkled flannel moth, Lagoa crispata (Packard), occurs throughout the
eastern half of the United States and feeds on a wide variety of plants, including
oak, locust, birch, cherry, and apple. According to some reports, it occurs most
commonly in the northern parts of its range; however, it has completely defoliated
shin oak over several hundreds of hectares of rangeland in Texas. The adult is cream
colored, with black wavy lines and brownish, crinkled hairs on the forewings. Full-
grown larvae are oval and about 25 mm long. The body is covered with long, silky
brown hairs that meet in the form of a ridge along the back and then slope off,
rooflike, on each side. Winter is spent in the pupal stage in a cocoon. The cocoon is
unique in being urn-shaped and having a flat, hinged, circular lid that is lifted as the
moth emerges. The stings produced by spines on the larvae of this species appar-
ently are less severe than those produced by related species (//4).

Norape ovina Sepp feeds on redbud, silktree, and beech from New Jersey and
southern Pennsylvania southward. The adult is a pure white moth with a small

amount of crinkly hair. The larvae are spotted and sparsely clothed in tufts of hair.
Family Limacodidae
Slug Caterpillar Moths

Limacodid larvae are sluglike in appearance. The head is concealed in the
thorax, the thoracic legs are small, and the prolegs are replaced by sucking discs.
Pupation occurs in dense, brownish, oval, silken cocoons spun between leaves or
attached to twigs. Each cocoon has a hole covered by a lid at one end through which -
the adult emerges.

The saddleback caterpillar, Sibine stimulea (Clemens), is widely distributed in
the Eastern and Southern United States and feeds on a wide variety of trees and
ornamental plants. The larva is brownish except for a green patch, which resembles
a saddlecloth, on the middle of the back. In the middle of this patch is an oval,
purplish-brown, saddlelike spot. The body is armed along the sides with fascicles of
poisonous spines, and has a pair of spiny tubercles at each end (fig. 65).

Courtesy Conn. Agric. Exp. Stn.
Figure 65.—Larvae of the saddleback caterpillar,
Sibine stimulea.

176

The hag moth, Phobetron pithecium (J. E. Smith), feeds on various deciduous
trees and shrubs. The larvae are brown, about 10 mm long, and each bears nine
pairs of lateral brown processes. The third, fifth, and seventh pairs are long, curved
and twisted, and are suggestive of the disheveled locks of a “hag.”’ These processes
are clothed with stinging hairs.

The oriental moth, Cnidocampa flavescens (Walker) (fig. 66), an introduced
species first recorded in this country near Boston in 1906 (223), is still confined to
eastern Massachusetts. The adult has a wingspread of 30 to 42 mm. Full-grown
larvae are about 22 mm long and marked with yellow, blue, green, and purple. The
larvae feed on a large number of tree species, including Norway and planetree
maples, sweet birch, cherry, apple, pear, plum, oak, aspen, willow, honeylocust,
hickory, and hackberry.

F-519528
Figure 66.—The oriental moth, Cnidocampa flavescens:

Upper, adults and cocoons; middle, a defoliated

ole), maple; lower, full-grown and newly hatched

arvae.

177

Adults deposit their eggs either singly or in groups in the undersides of leaves.
Young larvae feed on the lower epidermis of the leaves; older ones consume all but
the larger veins. A full-grown larva forms a cocoon by spinning a network of
threads around itself and attaching them to the bark in the forks of limbs or twigs.
Later it secretes a fluid that fills the spaces between the threads and hardens. Winter
is Spent as a prepupa in the cocoon. People suffer severe skin irritation when they
come into contact with the larva’s venomous spines. The tachinid parasite,
Chaetexorista javana Bauer & Bergenstamm, was imported from Japan against this
species in 1929 and 1930 and has exerted a considerable degree of control.

Prolimacodes badia Hubner feeds on various hardwoods such as‘oak, beech, and
black cherry in the Northeastern States and southern Canada. It has also been
observed feeding on maple in North Carolina. P. scapha Harris feeds on pin cherry
and blackgum in Massachusetts and New Jersey. /sa textula Herrich-Schaffer feeds
on Norway maple and oak, Packardia geminata Packard on pin cherry, and
Tortricidia flexuosa Grote on oak, gray birch, and pin cherry in the New England

States.
Family Pyralidae

Several subfamilies, sometimes listed as separate families, of the Pyralidae
contain species of interest in forestry. These include the Pyraustinae, Epi-
paschiinae, and Phycitinae.

Subfamily Pyraustinae
Pyraustine Moths

The grape leaffolder, Desmia funeralis (Hubner), an important pest of grape,
both cultivated and wild, occurs in eastern North America and along the West
Coast. It also feeds on Virginia creeper. The body of the adult is black except for a
white band on the abdomen of the male and two white bands on the abdomen of the
female. The forewings and hindwings are black with white spots and have coppery .
reflections. The wingspread is about 25 mm. The larvae live in tubes formed by
rolling over the edges of leaves and tying them with silk. When they are fully
grown, larvae are covered with sparse, fine, yellow hairs, are translucent yellow-
green, and are 18 to 25 mm long. The winter is spent in the pupal stage inside the
larval tube. There are two generations per year in the South.

The basswood leafroller, Pantographa limata Grote & Robinson, occurs on
basswood in southern Canada and throughout the Eastern United States. The adult
is yellowish white, with an abundance of olive or dull-brown markings, and has a
wingspread of about 37 mm. Full-grown larvae are bright green, except for black
heads and black cervical shields, and are about 25 mm long. Adults appear during
June and July; larvae are present from July to September. Each larva rolls the apical
half or more of a leaf into the form of a tube in which it lives (fig. 67). Full-grown
larvae spend the winter in cocoons constructed by folding a part of a leaf. The
folded leaf drops to the ground with other leaves. Although frequently abundant,
this species does not seem to cause serious damage.

Phlyctaenia coronata (Hufnagel), the elder leaftier, larvae feed as leafrollers on
the leaves of American elder in the Northeastern States. The adult is brown except
for the presence of creamy-white spots and streaks, and has a wingspread of 22 mm.
Full-grown larvae are translucent, whitish or pinkish, and about 18 mm long.
Winter is spent as a prepupa in a hibernaculum usually spun in the hollow stems or
pith of elder. There may be two generations per year. This species occasionally
causes serious defoliation.

178

Subfamily Epipaschiinae

The pine webworm, Tetralopha robustella Zeller, occurs in southern Canada and
throughout most of the eastern half of the United States. Its food plants include
several species of pines: jack, red, eastern white, Scotch, pitch, Virginia, shortleaf,
longleaf, loblolly, and slash. Jack pine is preferred in the Lake States and adjacent
parts of Canada. In the Northeast, pitch pine is preferred. The adult has a wing-
spread of about 25 mm. The basal part of the forewing is purple-black, the central
part grayish, and the outer part blackish. Full-grown larvae are yellowish brown,
with two dark-brown longitudinal stripes on each side, and are about 18 mm long.

Adults are present from June to August and deposit their eggs on pine needles.
Young larvae mine the needles; older ones live in silken tubes that extend through
globular masses of brown, coarse frass webbed together by strands of silk (fig. 68).
These masses, which are found on the twigs, enclose the needles upon which the
larvae feed, and range in length from about 8 to 13 cm. Pupation occurs in a cell in
the soil. In the northern part of its range, there is usually one generation per year; in
the South there may be two (/240).

The pine webworm is often troublesome in pine plantations. Young seedlings up
to 0.6 m tall are sometimes completely defoliated and killed by the larvae in a

NN

F-519529 F-532810

Figure 67.—Apical portion of leaf rolled Figure 68.—Web nest of the pine
into a tube by a larva of the webworm, Jetralopha robustella,
basswood leafroller, Pantographa on slash pine.

limata.

179

single nest. Ugly nests on the twigs and branches of young pines being grown for
the Christmas tree trade sometimes make it impossible to sell them.

Tetralopha asperatella Clemens, the maple webworm, occurs in southeastern
Canada and throughout the Eastern United States. The larvae feed on the foliage of
various hardwoods such as sugar, red, and mountain maples, oak, elm, beech,
quaking aspen, and willow. The moth is powdery gray, with the outer half of the
forewing somewhat lighter. Full-grown larvae range from pale yellow through
shades of green to brown or black and are about 25 mm long. On sugar maple in
Wisconsin, eggs are laid on leaves partly rolled by other insects. When the larvae
hatch they feed on these leaves as skeletonizers; older larvae web together groups of
leaves, sometimes including all of the leaves on a branch in a web (fig. 69).
Heaviest infestations apparently occur in the more open portions of the crowns of
trees growing in the most exposed positions in the stand. When the larvae become
full grown, they leave the nest and drop to the ground on strands of silk. The winter
iS spent as a prepupa in a cocoon spun in the duff on the ground. There appears to
be only one generation per year (483).

F-519532

Figure 69.—Characteristic nest of Tetralopha asperatella
on oak.

This species had never been considered economically important until it was
shown to be a major factor leading to the development of “maple blight,” a
condition responsible for killing thousands of valuable sugar maples in Wisconsin
during the late 1950's.

Other species of Tetralopha that occur in the Eastern United States are T. militella
Zeller feeding on sycamore; and 7. melanogrammos Zeller, on sweetgum. Another
species identified only as being near T. asperatella is sometimes abundant locally
on beech in New England. The larva is yellowish green and has two pale, brownish

stripes running down the back.
Subfamily Phycitinae

The subfamily Phycitinae contains many important tree-infesting species. The
larvae differ considerably in their habits. Some feed in rolled or folded leaves: some

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construct cases of silk, or frass and silk, and feed from inside them; at least one is a
predator of scale insects; and many feed as borers in shoots, bark, roots, cones,
nuts, or fruits. Members of the subfamily are rather difficult to identify because of
the variability of many of their distinguishing characters.

The pecan leaf casebearer, Acrobasis juglandis (LeBaron), occurs from south-
ern Canada to Florida and Texas, and the larvae feed on the buds, flowers, and
leaves of walnut, butternut, and pecan. The adult is a grayish-brown moth with the
forewings black at the middle of the costa and reddish-brown near the middle of the
inner margin, a black raised-scale ridge is also present on the forewings, and the
head and thorax of the male are distinctly white above. Wingspread is 14 to 17 mm.
Full-grown larvae are mostly olive green to dark olive green and about 16 mm long.
The larvae feed during two growing seasons. The first summer, they feed on the
lower surfaces of leaves. They spend the winter in small cases attached to buds or
twigs. The following spring, they resume their feeding by eating into bud after bud
and constructing new cases as necessary. When the larvae become about half
grown, they move onto the underside of the rachis. The rachis and base of the
adjacent leaflets are notched and a tube of silk is formed and fastened to the notch in
the rachis. The leaflets are pulled down about the silk tube and eaten. This, plus the
injury to the buds, often results in serious damage to infested trees (454, 927).

Acrobasis caryivorella Ragonot occurs in southern Canada and throughout the
Eastern United States and the larvae feed on the foliage of hickory, walnut, and
pecan. The forewings of the adult are black with indistinct patches of gray; a black
raised-scale ridge is also present on the forewings. The wingspread is about 20 mm.
The larvae are purplish green and about 19 mm long. In the spring, they feed by
boring into new shoots that they tie together with silken threads. Full-grown larvae
construct oval cocoons in which they pupate. Several generations occur each year.
Generations occurring after the spring generation feed on silked-together leaflets.
In Florida and Texas, there are probably three or four generations per year. This
species has seriously damaged pecan seedlings in nurseries in Florida and Texas.

The pecan nut casebearer, A. nuxvorella Neunzig, occurs from North Carolina
and southern Illinois to Florida and Texas. Its host tree is pecan (920). The adult is
mostly dark brown with a dark-brown raised-scale ridge on the forewings, and has a
wingspread of 12 to 19 mm. Mature larvae are mostly purplish brown with greenish
undertones and are about !2 mm long. Winter is spent in the larval stage in a small
case near the base of a bud. Later, larvae bore into tender shoots, causing them to
become stunted and distorted. Larvae of the second generation bore into newly set
nuts, destroying from two to five nuts each. Larvae of later generations feed in the
large nuts or on the shucks. There are three or four generations per year in the
South.

Many other species of Acrobasis occur on forest and shade trees in the Eastern
United States. The birch tubemaker, A. betule/la Hulst, is common, feeding on
the foliage of various species of birch, particularly paper birch in the Northern
United States and southern Canada; A. betulivorella Neunzig occurs on river birch
in the Southeastern United States; A. rubrifasciella Packard, the alder tubemaker,
feeds on foliage of alder; A. indigenella (Zeller), the leaf crumpler, is associated
with hawthorn foliage; A. demotella Grote, the walnut shoot moth, bores in shoots
of walnut, hickory, and pecan; A. exsulella (Zeller) occurs in buds and between
leaves of hickory and pecan; A. e/yi Neunzig bores into, and galls, the rachis of
hickory leaves; A. carpinivorella Neunzig feeds on the leaves of American

181

hornbeam. Details on the immature forms and biology of these and other species of
Acrobasis are available (921, 922).

The genus Dioryctria has Holarctic distribution, and the larvae of all species are
borers in conifers. Many are highly injurious to seeds and cones (360, 547). The
larval habits of some of the more widely distributed species are quite variable. This
suggests that more than one species is involved in certain cases. This can be
established only through further taxonomic and biological investigations. The
genus has been discussed (357, 360, 553, 897, 901, 902, 924, 925).

The Zimmermann pine moth, D. zimmermanni (Grote), is widely distributed in
the Northern United States and southern Canada. The larvae feed in the cambial
region and outer xylem under the bark of limbs and trunks of all commonly grown
species of pines. The adult has forewings of gray blended with red-brown and
marked with transverse zigzag whitish and black lines: the hindwings are yellowish
white. Mature larvae are variably dirty-white, pink to greenish, with prominent
small black setal bases, with black muscle attachments: they are about 18 mm long.

Adults emerge from mid-July to mid-August and deposit their eggs at the edges
of wounds, on resin masses, in bark crevices. or on terminal buds. Hatching occurs
in about 8 to 10 days and the young larvae, without feeding. enter bark recesses and
spin hibernacula. Here they remain until the following spring. Feeding begins in
May or June, first in the bark, then by tunneling in the cambium area of new growth
on terminals or laterals. Damaged terminals usually become “‘fish-hooked™ and
turn yellowish green. Toward the end of June, the larvae leave the new growth and
tunnel beneath the bark in the whorl area, girdling branches and leaders. Pupation
occurs in resin masses or in the tunnel. Dead tops, dead branches hanging on trees
(fig. 70), and burllike growths on trunks above girdled whorls and pitch masses
(fig. 71) are evidence of attack. Infested trees tend to be attacked again and again,
thus becoming so-called brood trees (/0/7).

F-49295 F-492960
Figure 70.—Damage to young pine by Figure 71.—Burllike growths on trunk
the Zimmermann pine moth, above girdled whorl, a sign of attack
Dioryctria Zimmermanni. by Zimmermann pine moth.

182

Zimmermann pine moth damage has been especially serious in Christmas tree
plantations of Scotch, red, and Austrian pines in the North Central States and
southern Canada.

The remaining eastern Dioryctria species are known primarily for their injury to
cones although other tree parts may be infested (360).

The southern pine coneworm, D. amatella (Hulst), occurs throughout the
Southeast from Virginia to Florida and westward to Texas. In addition to cones, it
infests male flowers, shoots, fusiform rust cankers, Cronartium quercuum (Berk. )
Miyabe ex Shirai f. sp. fusiforme, and southern cone rust, C. strobilinum Hedgc. &
Hahn, infested cones and tree wounds of southern pines, particularly of longleaf,
slash, and loblolly pines. The adult (fig. 72) has a wingspread of 27 to 32 mm; the
forewing is dark brown, nearly black, with contrasting white patches and zigzag
lines. Mature larvae are brownish to purplish above, pale whitish to greenish below,
with black muscle attachments; they are about 25 mm long.

F-532840

Figure 72.—Adult of the southern pine coneworm,
Dioryctria amatella.

First-stage larvae overwinter under bud or bark scales; occasional larger larvae
overwinter in damaged cones. In longleaf pine, overwintered larvae usually feed
first upon flowers, both male and female, then enter shoots or second-year cones.
The larvae in shoots may mature there or migrate to second-year cones. In slash and
loblolly pines, overwintered larvae more frequently occur in fusiform rust cankers
where they complete development. Later overlapping generations occur mainly in
second-year cones of all host pines. Pupation occurs within host materials. This
species is frequently reported as the cause of heavy cone losses of southern pines.

Dioryctria abietivorella (Grote), the fir coneworm, ranges across Canada and
the adjoining Eastern and Western United States. It is a coneworm that also infests
buds and twigs, and occasionally feeds under the bark. It occurs on a variety of
conifers including the eastern spruces and firs and red, eastern white, and jack
pines. The moth has blue-gray forewings crossed by diffuse, pale zigzag lines; the
hindwings are pale gray. Wingspread is from 20 to 30 mm. The mature larva is
reddish carneous to amber- or purplish-brown with indistinct darker dorsal and
subdorsal lines, and is from 15 to 25 mm long (791, 897).

Damage is similar to that of the spruce coneworm. Prepupal larvae or pupae
overwinter and produce early season moths in June and July. However, some larvae
apparently mature and pupate and the moths emerge in late summer and fall. These

183

give rise to a group of overwintering larvae. Essentially, however, a single genera-
tion occurs each year (759).

The spruce coneworm, D. reniculelloides Mutuura & Munroe, occurs through-
out the forested regions of Canada and in the Northern and Western United States.
It feeds primarily upon cones and associated vegetative parts of white spruce and
various Other spruces and firs but also has been reported from other conifers within
its range. The moth has dark-gray forewings strongly shaded blackish and crossed
by distinct white zigzag lines. The hindwings are dusky with an obvious paler,
diffuse postmedial (outer) crossband. The wingspread is 22 to 27 mm (90/).
Mature larvae are pale yellow with distinct brown longitudinal stripes and are about
17 mm long (797).

In Canada adults are present from late June to August. Eggs are laid in bark
crevices and similar niches. The first instars diapause over winter. In spring they
first mine needles then bore buds, staminate flowers, cones, or shoots. Masses of
webbed frass enclose the food material. During certain years this coneworm takes a
heavy toll of white spruce cones (S05).

The blister coneworm, D. clarioralis (Walker), occurs throughout the South and
northward into coastal Massachusetts. It infests vegetative buds, male bud clusters,
conelets, and cones of pines including longleaf, slash, loblolly, and shortleaf.
Adults have blended brown and black forewings marked with wide, nearly black
transverse bands near the base; wingspread is from 22 to 29 mm (fig. 73). Larvae
are yellowish brown to brownish orange, often heavily suffused with gray; they are
about 18 mm long. This coneworm was the most common coneworm found in
cones of shortleaf and loblolly pines in Arkansas in 1960 (/367), but southwide
seems to be of lesser importance.

Winter is spent in the developing larval stage in buds or conelets. Flowers, cones,
and buds are bored into during the spring, then succeeding generations infest buds,
shoots, conelets, and occasional cones. A characteristic of damage by this species
is the presence of a resin-coated silk blister over the entrance hole where ejected
frass accumulates. Pupation occurs outside the hollowed-out food material on a
twig or cone stalk in silken cocoons covered with bits of twig scales. Normally
three generations per year occur in the South.

The webbing coneworm, D. disclusa Heinrich, occurs in southern Canada and
throughout the Eastern United States. It infests cones of red, jack, and Scotch pines
in the Northern States and cones of Virginia, loblolly, shortleaf, and longleaf pines

F-494452
Figure 73.—Adult of the blister coneworm, Dioryctria
clarioralis.
184

in the South. The adult has forewings of bright yellow to orange, shaded red-brown
with sharp white crosslines and markings, and has a wingspread of 17 to 29 mm.
The larvae vary from olive green to buff. Northern larvae are small, slender, and
from 14 to 18 mm; southern larvae are robust and from 20 to 25 mm long.

In the North, winter is spent as first-stage larvae in hibernacula beneath bark
scales of branches. Young larvae infest staminate flowers in the spring then enter
and tunnel in second-year cones as third to fourth instars (759). In the South partly
grown larvae have been reported to feed and overwinter in conelets (925), but in
northern Georgia no cone damage was observed until spring when third-stage
larvae infested young second-year cones without evidence of prior conelet boring.

Infested cones are hollowed and have characteristic masses of tightly webbed
frass over the entry holes. Pupation occurs in the damaged cones, and adults
emerge from late May (South) through July (North). A single generation occurs
each year. Damage by the webbing coneworm is generally sporadic; however,
serious infestations and cone losses were experienced in loblolly pine seed orchards
from 1978 to 1982 throughout the South. Pheromone-baited traps have been
successful in trapping adults to determine population levels and periods of activity
(288).

The baldcypress coneworm, Dioryctria pygmaeella Ragonot, occurs in the
Eastern United States. It infests the cones of baldcypress and pondcypress from
Maryland to Florida. In northern Florida it has destroyed more than 75 percent of a
single year’s crop. There are three complete generations in its southern range (839).

Several Dioryctria species of importance in the Southeast have only recently
been described (547, 902); previously these were often either misidentified or
referred to species groups in the literature. These include: the south coastal
coneworm, D. ebeli Mutuura & Munroe, limited to the lower Southeast from
southern Georgia and Florida westward along the Gulf area. It infests slash,
longleaf, and loblolly pines and also occurs on pondcypress. The moth (fig. 74) 1s
similar to D. abietivorella and has a wingspread of about 20 mm. Mature larvae are
brown to purplish and about 15 to 20 mm long. In the spring this coneworm infests
conelets infected by cone rust, Cronartium strobilinum Hedge. & Hahn. Through-
out the summer repeated generations attack second-year cones, both intact and
damaged. Fall infestations occur in fusiform cankers or terminal buds where larvae
mature over winter. Larvae drop from feeding sites to pupate, apparently in soil
litter. This species is periodically common, but its affinity to diseased or damaged
cones reduces its impact on cone crops (357).

F-519534
Figure 74.—Adult of the south coastal coneworm,
Dioryctria ebeli.

185

Dioryctria merkeli Mutuura & Munroe, the loblolly pine coneworm, occurs
throughout the Southeast and northward to Maryland. It infests flowers, shoots, and
cones of loblolly, slash, and longleaf pines, and occasionally other southern pines.
The moth resembles D. zimmermanni but is duller in color and the hindwings are a
light gray-brown. Wingspread is about 25 to 30 mm. The mature larva is purplish
black above, deep blue-green below, with black muscle attachments, and is 18 to 25
mm long. In early spring, small larvae infest male and female flowers, then soon
migrate to shoots or second-year cones. In slash and longleaf pines such larvae may
mature in shoots; in loblolly they migrate to second-year cones. Mature larvae
aestivate in dead shoots and cones where they pupate and emerge as adults in
August and September. Larvae from this generation overwinter- under bark or bud
scales as nonfeeding first instars. A single generation occurs yearly. This species
periodically causes extensive cone loss in loblolly pine (/365).

The mountain pine coneworm, D. yatesi Mutuura & Munroe, attacks second-
year cones of Table Mountain pine. It ranges throughout the Appalachian Mountain
region from Pennsylvania to Georgia (547, 902).

Dioryctria taedae Schaber & Wood, the Atlantic pine coneworm, attacks
loblolly pine cones in Maryland and occurs sparingly in the Southeast (/057).

Dioryctria resinosella Mutuura resembles D. zimmermanni but feeds on red pine
shoots and cones in the Northeast, Lake States, and southern Ontario (759, 899).
Mature larvae are gray to green with black muscle attachments, and about 15 mm
long.

The locust leafroller, Nephopteryx subcaesiella (Clemens), occurs from south-
ern Canada and Maine to North Carolina and westward to Colorado. The larvae
feed inside tied-together leaves of black locust, other locusts, and wisteria. The
adult is dark gray with some reddish scales at the wing base, and has a wingspread
of about 25 mm. Mature larvae are about 20 mm long. The head is dark brown and
the body light green, except for a darker green line from the middle of the dorsum
and for three faint lines on each side. The prothorax is usually distinctly darker than
the mesothorax and metathorax. The winter is spent as pupae in silken cocoons in
the soil. This species is quite common at times, but is seldom injurious. N.
virgatella (Clemens) 1s a leafroller on black locust. The larvae resemble those of the
locust leafroller except that they are much more lightly pigmented. N. subfuscella
(Ragonot) is a leafroller on sumac from Maine to Texas. The adult is ash gray and
has a wingspread of about 22 mm. Mature larvae are yellowish green except for
dark-green lines on the dorsum and sides, and are 13 to 20 mm long. Additional!
information on the species of Nephopteryx is available (244, 3/5, 316, 923).

The lesser cornstalk borer, Elasmopalpus lignosellus (Zeller), a widely dis-
tributed species, has damaged black locust seedlings in nurseries in several south-
ern states. Loblolly pine seedlings in nursery beds in Georgia, Virginia, and Rhode
Island have also been damaged. Gall-like growths occur at points of injury on the
lower stems of locust seedlings, causing them to die or break off at the ground line.
The male adult is mostly ochre-yellow to light brown but the female is heavily
shaded black. The wingspread is 16 to 24 mm. The forewings are long and narrow
and marked with several black spots. Mature larvae are greenish white with
interrupted longitudinal stripes of dark brown or purple. The winter is spent in the
larval or pupal stage and there are up to four generations per year in the Deep South
(923, 1205).

The American plum borer, Euzophera semifuneralis (Walker), is widespread
and attacks a wide variety of hardwoods, including plum, pin cherry. apple.

186

London plane, mountain-ash, walnut, persimmon, mulberry, basswood, poplar,
sweetgum, and ginkgo. Serious infestations have occurred on young London plane
trees in newly developed areas on Long Island. The larvae bored in the trunks and
larger branches, and were associated with a canker condition (644).

Euzophera ostricolorella Hulst has been recorded from Long Island to Georgia
and westward to Ohio, Kentucky, Tennessee, and Louisiana. Its preferred host
appears to be yellow-poplar, but it also attacks magnolia. Heaviest infestations on
yellow-poplar are found at the base of trees over 25 cm in diameter (537). Attacks
above a height of 60 cm on the trunk are rare. The larvae feed in the inner bark,
excavating tunnels which extend both above and below the ground line. Winter is
spent in the pupal stage in the tunnel. There appears to be one generation per year in
the North and two per year in the South. Trees of all sizes may be killed by heavy
infestations.

Euzophera magnolialis Capps has been observed damaging or killing magnolia
seedlings in Florida nurseries. The adult has a wingspread of about 25 mm and the
forewing is purplish brown with black terminal dots. Full-grown larvae are white
and flattened, they taper toward the rear, and are about 28 mm long. Attacks occur
at the base, the larvae tunneling in the lower centimeters of the trunk and in the
larger roots entering the crown. Heavily infested trees may be girdled and killed
(196).

Canarsia ulmiarrosorella (Clemens) occurs in southern Canada and throughout
the Eastern United States. The larvae feed on the leaves of elm that they web
together with silk. The adult has a wingspread of 15 to 20 mm. The forewings are
gray-dusted white and crossed by dark gray or blackish wavy lines. Mature larvae
are green, sparsely hairy, and about 18 mm long. Winter is spent in the pupal stage,
and there are two generations per year. This species is sometimes abundant enough
to attract attention, but is seldom injurious.

Moodna ostrinella Clemens is rather widely distributed in southeastern Canada
and the Eastern United States. In Canada, the larvae have been found in second-
year pine cones damaged by other insects. They have also been found in damaged
pine cones in Arkansas; in sumac heads in New York, Pennsylvania, and Texas; and
in fusiform rust cankers and in dead and diseased cones on pines in Florida. Full-
grown larvae vary from yellow with a greenish tinge to purplish brown with bright
orange-brown heads, and are about 8 to 10 mm long.

Actrix nyssaecolella (Dyar) is a leaffolder of tupelo from western Pennsylvania to
New Jersey and Massachusetts and south to North Carolina. The adult is powdery
gray and has a wingspread of 18 mm. Full-grown larvae are black with yellow
heads. Winter is spent in the pupal stage, and larvae are present from May to
August.

Laetilia coccidivora (Comstock) larvae feed on various scale insects. According
to some reports, it has aided materially in the control of outbreaks of the pine
tortoise scale. It has been recorded from Pennsylvania and Ohio to Florida and

Texas.
Family Geometridae
Geometrid Moths

This is one of the largest families in the order Lepidoptera, and approximately
1,200 species occur in the United States and Canada alone. The larvae are easily
distinguished by their peculiar method of locomotion, which consists of bringing
the rear end up to the thoracic legs, forming a loop of the body, and then extending
the whole body forward. This characteristic, which results from the absence of

187

prolegs near the center of the body, has given rise to a number of common names
for the group such as geometers, measuring worms, inch worms, loopers, canker-
worms, and spanworms. The larvae of many species have the interesting habit of
standing nearly erect on the posterior prolegs when disturbed. Motionless in this
position, they are almost indistinguishable from small twigs or spurs. With the
exception of several Hawaiian species, which are predators of fruit flies, all
members of the family are foliage feeders.

Archiearis infans (Moschler) occurs from coast to coast in southern Canada and
in the Northern States south to Pennsylvania. Its hosts are paper and gray birches.
The adult has cryptic forewings and orange and black hindwings, and has a
wingspread of about 25 mm. Mature larvae are green to reddish brown and about 25
mm long. Two fine yellowish lines run the length of the dorsum, and there are two
similar lines plus a broad subspiracular stripe on each side. Larvae are present from
May to July, and winter is spent in the pupal stage.

The fall cankerworm, Alsophila pometaria (Harris) (fig. 75), occurs from the
Maritime Provinces to Alberta in southern Canada, and throughout the Eastern
States south to Georgia and west to Missouri and Montana (397). Its hosts are elm,
apple, hickory, maple, ash, beech, boxelder, basswood, cherry, and the oaks (399).
The male moth is brownish gray and has a wingspread of 25 to 35 mm. The
forewings are rather glossy with purplish reflections and are crossed by two faint,
jagged, whitish bands. The hindwings are grayish brown, and each has a faint
discal dot. Female adults are wingless and about 12 mm long. Full-grown larvae
vary from very light green to very dark brownish green and are about 25 mm long.
The head and anal segment vary from pale green to almost black, and are some-
times mottled. A median, longitudinal, black stripe extends down the back of the
darker larvae. Light-green larvae have longitudinal white lines. These color forms
are associated with population density, with the darker larvae occurring in denser
populations (334). There are three pairs of prolegs—a very small pair on the fifth
abdominal segment and larger pairs on the sixth and anal segments.

The fall cankerworm spends the winter in the egg stage, and hatching occurs
from mid-April to early May. Young larvae skeletonize the young leaves at the tips

Figure 75.—The fall cankerworm, Alsophila pometaria:
A, adult female ovipositing; B, egg mass.

188

of branches; older ones devour all but the midribs or larger veins of the leaves. They
become mature in 5 to 6 weeks at which time they drop to the ground and enter the
soil where they pupate in tough cocoons made from silk and soil particles. Adults
emerge from November to January and in some localities into the springtime,
usually following some freezing weather. Reproduction is by sexual mating and a
type of parthenogenesis tentatively described as gynogenesis (the genotypes are
only of the female) (866, 1067). Females climb the trees, mate, and deposit about
100 brownish eggs in compact, uniform rows of single layers on the trunk, smaller
twigs, and branches (397). There is one generation per year.

The fall cankerworm is an important pest of forest and shade trees. Outbreaks
occur periodically and sometimes cover large, forested areas. Shade trees in urban
areas are subject to heavy infestation and may be seriously damaged. Severe
infestations in recreational areas are particularly annoying. Natural control by the
egg parasite, Telenomus alsophilae Viereck, 1s a major factor in the decline of fall
cankerworm populations (398).

The Bruce spanworm, Operophtera bruceata (Hulst), occurs from coast to
coast in Canada and from New England to the Lake States. In eastern America its
preferred hosts are sugar maple, quaking aspen, willow, and beech, but many other
hardwoods such as paper birch, red oak, pin cherry, common chokecherry, alder,
and serviceberry are also attacked. Serious outbreaks have occurred in both the
United States and Canada.

Female moths are light brownish-gray and practically wingless. Males are light
brown and have fully developed wings; their forewings are semitransparent, banded
with brown or gray, and have a wingspread of 25 to 30 mm. Full-grown larvae may
vary from bright green to dark brown and have three narrow yellow stripes on each
side; larvae are about 18 mm long.

In Canada, winter is spent in the egg stage and hatching occurs in early spring.
The larvae feed either openly on leaves or within the shelter of leaves loosely rolled
and webbed together with silk. In heavily infested stands, trees are sometimes
literally festooned with this silk. Pupation occurs in thin, silk cocoons in the soil or
duff. Adults appear in the fall. The female climbs trees and deposits eggs singly in
bark crevices, under loose bark, or in other sheltered locations on the tree (/62). A
polyhedrosis virus disease has occurred commonly in New Brunswick infestations.

The winter moth, O. brumata (L.), an introduced species known to have been
present in Nova Scotia since about 1930, now occurs also on Cape Breton and
Prince Edward Islands, and in the Pacific Northwest. Although it was originally
found in the United States at Portland, Oreg., in 1958, the male specimens were
previously misidentified and not correctly determined until 1978. The female moth
has a dusky-brown to silver-gray body with occasional irregular black spots and is
practically wingless. Males have fully developed wings with a wingspread of 27 to
30 mm. The body is dusky brown with black spots and the forewings are dusky
brown with obscure markings.

The most common hosts are apple, northern red oak, American elm, red maple,
basswood, and eastern hophornbeam, but a number of other hardwoods are also
attacked, such as poplar and willow. Persistent severe attacks result in thin tops,
dead twigs and branches, and ultimately the death of trees. Severe defoliation of
principal host trees often occurs in Nova Scotia.

Winter is spent in the egg stage and hatching occurs from late April to late May.
Young larvae feed first on opening buds and on the undersides of developing leaves.
Older larvae feed inside loose leaf rolls. When larvae become mature, they drop and

189

enter the ground where they pupate in earthen cells at a soil depth of 5 to 12 cm.
Pupation occurs during late June and early July, and the pupae remain in their cells
during the remainder of the summer and fall. Adults appear from late October to
mid-December. The females climb tree trunks or other objects near their emergence
sites and deposit their light-green eggs in bark crevices, under lichens on the trunks
or branches, or in other places that afford suitable shelter (270). The population
dynamics of the winter moth has been investigated (382), and a key is available to
distinguish this defoliator from associated species of Operophtera, Erannis, and
Alsophila (403).

Two imported parasites, the tachinid, Cyzenis albicans (Fallén), and the ich-
neumonid, Argypon flaveolatum (Gravenhorst), have provided successful biological
control of the pest in Nova Scotia (383).

The genus Eupithecia contains a large number of species, a number of which
occur in forests of the Eastern States and eastern Canada. E. palpata Packard, the
small pine looper, occurs on various species of pine, especially eastern white and
jack; E. luteata Packard, the fir needle inchworm, feeds on various spruces,
particularly white, and other conifers, especially balsam fir and larch; FE. filmata
Pearsall, the early brown looper, is found on a wide variety of conifers, especially
white spruce and balsam fir; and E. transcanadata MacKay, the small conifer
looper, that feeds on a wide variety of conifers, especially white spruce, black
spruce, and balsam fir (738).

Hydria prunivorata Ferguson, the cherry scallop shell moth, occurs on black
and occasionally other wild cherries throughout much of eastern North America.
The adult has a wingspread of about 37 mm. The forewings are marked with 12 to
15 whitish, scalloped, parallel lines and a black discal dot; the hindwings are
marked with 6 to 8 whitish lines. Full-grown larvae are about 20 mm long. The
body is blackish above with four longitudinal yellow lines, and is straw yellow
beneath. The larvae live in nests that they construct by webbing together the leaves
toward the end of a branch. Heavily infested trees occasionally are completely
defoliated. An egg parasite, Telenomus sp., is the principal mortality factor in
populations that have remained at epidemic levels for 2 years or more (/070).

Rheumaptera hastata (L.), the spear-marked black moth, feeds on paper birch,
willow, and alder from Alaska east to Labrador and Newfoundland. It occurs as far
south as New Mexico and North Carolina. Major areas of defoliation have been
limited to interior Alaska (/274). Full-grown larvae are about 25 mm long. The
head is brown to shiny black; the body is dark brown to black with a row of small
black dots and a few white to brick-red spots on each side. In the Northeastern
States, larvae are present from June to September, and winter is spent in the pupal
stage.

Lomgarapha semiclarata (Walker), the wildcherry looper, occurs on pin cherry
in southern Canada and the Atlantic Coast States. Full-grown larvae are light green
and about 18 mm long. The related species, L. vestaliata, (Guenée), is known to
occur on cherry, mountain-ash, and birch in Maine.

Cabera erythemaria (Guenée) feeds on willow in the Northeastern States. Full-
grown larvae are about 27 mm long. They are light green with a reddish stripe on
each side of the head, reddish patches on the back, and a red stripe on each side of
the body. Winter is spent in the larval stage. The related species, C. variolaria
Guenée, is found on willow and quaking aspen in the Northeastern States and
southwestern Canada.

190

Itame pustularia Guenee occurs on red maple from eastern Canada to Florida. It
is occasionally abundant. It has been reported from widely scattered localities in
southern Canada, and high populations have occurred in the areas of central New
Brunswick that were repeatedly sprayed for control of spruce budworm (/234). The
adult has a wingspread of about 25 mm and Is pure white except for four brownish
spots on the margin of the forewing. The light-green eggs are laid singly on bark,
lichens, or in litter. Eggs overwinter and larvae hatch in early spring. Young larvae
are light green and closely resemble other geometrids, especially cankerworms and
Bruce spanworm. Older larvae have two whitish lines down the back, bordered by
yellowish-white lines; the integument is quite wrinkled. Before pupating, larvae
change color from green to pink. Larvae are present during May and June.

Semiothisa signaria dispuncta (Walker), the spruce-fir looper, feeds on the
foliage of various conifers such as eastern white pine, spruce, balsam fir, and
hemlock. It has a transcontinental range. Full-grown larvae are about 20 mm long.
The body is light green with brownish tinges on top, and there are two light
longitudinal lines on the back. S. granitata (Guenée) feeds on Virginia pine and
probably pitch pine in the Eastern States from southern Maine and Ohio to the
Carolinas. S. ocellinata (Guenée) occurs in southern Ontario and the Northeastern
and Lake States and feeds on honeylocust and black locust. Full-grown larvae are
light yellow-green to green except for faint, wavy red lines and are about 25 mm
long. S. bisignata (Walker) occurs in southern Canada and the Northeastern States
and feeds principally on eastern white pine. S. sexmaculata (Packard), the green
larch looper, a widely distributed species, feeds on larch. These species of
Semiothisa overwinter as pupae in the duff. Eggs are laid singly on host foliage
throughout the growing season (782).

Eufidonia notataria (Walker) occurs on eastern white pine in the Atlantic Coast
States. In southern Canada, it has been observed mainly on tamarack, eastern
hemlock, balsam fir, and spruce. Full-grown larvae are dark green and about 25 mm
long. They have a light-colored line on the dorsum, two stripes along each side, and
fine brown markings in herringbone pattern on lobes. Larvae are present from July
to September, and winter is spent in the pupal stage. The russet-brown eggs are laid
singly or in clusters in leaf axils and at forks of branches of recent growth (783).

Melanolophia canadaria (Guenée) feeds on the foliage of various deciduous
plants such as basswood, birch, ash, willow, larch, and a few shrubs in the United
States and Canada. Full-grown larvae are 20 to 25 mm long. The body is yellow-
green and has a broken, purplish line on the top and yellowish stripes suffused with
purplish red on each side. Larvae are present from June to early August, and they
overwinter in the ground.

Anacamptodes ephyraria (Walker) has been observed feeding on hemlock in the
Northeastern States and on several hardwoods in Canada and Northeastern States
south to Texas and Florida (753). Full-grown larvae are greenish or light gray to
reddish-brown with a brown herringbone pattern on lobes, and are about 25 mm
long. The head is bilobed and wider than the thorax, and the second abdominal
segment is usually swollen at the sides. Winter is spent as a pupa. A. pergracilis
(Hulst) feeds on baldcypress in the South (fig. 76). During a 1980-81 outbreak in
Florida, trees on 24,000 hectares were defoliated. This species may have as many
as Six generations a year. In winter, when normal baldcypress 1s leafless, larvae feed
on previously defoliated trees which have reflushed. The overwintering form is
unknown. Light-green eggs are deposited under the bark of the stem primarily in
the upper branches. Pupation occurs under bark or in epiphytic bromeliads (fig.

19]

77). A larval parasite, /chneumon navus Say, appears to be the major natural
controlling agent for this looper (338).

Iridopsis larvaria (Guenée) feeds on aspen, paper birch, alder, and other decid-
uous plants in the Northeastern States and southern Canada. Full-grown larvae are
yellowish green to light brown and 20 to 25 mm long. There is a reddish band on
the second abdominal segment, several reddish diamond-shaped formations on the
dorsum, and a pair of blunt tubercles on the eighth abdominal segment (783).

F-532013
Figure 76.—Adult female moth of Anacamptodes
pergracilis. Note extended ovipositor, which is used to
insert eggs in bark of baldcypress.

tit tin 5 HE Al

F-532014

Figure 77.—Anacamptodes pergracilis pupae in cells in
bark of baldcypress.

Ectropis crepuscularia (Denis & Schiffermuller), the saddleback looper, feeds
on the foliage of a wide variety of trees, especially conifers, from coast to coast in
southern Canada. South of Canada, it is usually found on hardwoods such as birch,
maple, oak, poplar, and walnut. Full-grown larvae are reddish to chocolate brown
and 25 to 30 mm long. The head is somewhat mottled, and there is a pair of blunt
tubercles on the eighth abdominal segment. Larvae are present from May to
September in the Northeastern States and the winter is spent in the pupal stage in
the ground.

Epimecis hortaria (F.) occurs on sassafras and yellow-poplar in the Atlantic
Coast States. The adult is either dark colored to almost black or light gray with a
dusting of brown. The forewings are marked with lines and the wingspread is about

192

50 mm. Full-grown larvae are yellowish to dark brown with five pale, yellowish
longitudinal lines and are about 30 mm long. Larvae are present during June and
July, and they feed at night. There is one, and possibly a second, generation per
year.

Phigalia titea (Cramer) occurs in southern Canada westward to Saskatchewan
and the Eastern States. Its hosts include various hardwoods, especially northern red
oak, red and sugar maples, basswood, hickory, and elm. Numerous outbreaks have
been recorded in mixed oak and maple stands in the Northeastern States and
Virginia. The male moth has a wingspread of 37 mm. The thorax is whitish, and the
abdomen is marked with two rows of black dots on the dorsum. The forewings are
dotted with dark-brown specks and are marked with three blackish lines and a row
of black spots along the outer margin. The wings of the female are vestigial and
functionless. Full-grown larvae are pinkish with many blackish longitudinal lines
and are about 37 mm long. The thoracic segments are thick, and there are hairy
tubercles on all body segments. Pinkish-red eggs are laid in crevices in the bark of
dead branches. Larvae are present from May to July, and winter is spent in the
pupal stage beneath litter (//87).

The spring cankerworm, Paleacrita vernata (Peck) (fig. 78), occurs throughout
the same general range as the fall cankerworm, also farther southwest and west to
Texas, Colorado, and California. Its hosts are about the same as those of the fall
cankerworm. Apple and elm are especially favored. Male moths have a wingspread
of about 21 to 30 mm. The forewings are silky with loosely attached, brownish
scales, and are crossed by three jagged, dark lines. The hindwings are pale, ashy
gray, and each bears a dusky discal spot. Each of the first seven joints of the
abdomen bears two transverse dorsal rows of stiff, reddish spines, pointed posteri-
orly. Female moths are wingless and generally whitish or brown or black. There is a
darker stripe down the back and two transverse rows of reddish spines on each of the
first seven joints of the abdomen. These spines are often so prominent as to give the
dorsum a reddish aspect.

Full-grown larvae are reddish to yellowish brown, yellowish green, or blackish
and are about 18 to 30 mm long. The head is light and mottled with brown. The
body is usually marked with a yellow stripe just below the spiracles, and a broad
greenish-yellow stripe down the middle of the venter. There are two pairs of
prolegs, one on each of the eighth and anal segments.

Spring cankerworms do not spin cocoons, but spend most of the winter as larvae
in earthen cells. Pupation occurs in late winter, and the adults appear in early spring
about the time frost leaves the ground. Female moths crawl up the trunks of trees,
mate, and deposit loose clusters of 100 or more eggs in bark crevices or under bark
scales on the trunk or branches. Hatching occurs by early May and the larvae
become full grown by early June, at which time they drop and enter the soil. Like
the fall cankerworm, this species may be an important pest of shade trees and
forested areas. _

Fall and spring cankerworm populations are normally kept under control by
natural control factors. Outbreaks often occur, however, over large areas. Isolated
trees can be protected by placing sticky bands around the trunks to prevent female
moths from climbing to lay their eggs (/224).

The linden looper, Erannis tiliaria (Harris), occurs from Nova Scotia to central
Alberta and south to Missouri, Kansas, and Utah (753). The larvae feed on the
foliage of a wide variety of hardwoods, especially basswood, elm, hickory, maple,
oak, birch, and apple. The female adult is light gray to brownish, wingless, about

193

A,B,D, F-531247
C, courtesy Can. For. Serv., Can. Dep. Environ.,
Sault Ste. Marie, Ont.

Figure 78.—Spring cankerworm, Paleacrita vernata: A,
females and eggs; B, male; C, fully grown larva; D,
pupae.

12 mm long, and is marked with black spots on the sides and back. Male moths
have fully developed wings with a wingspread of about 42 mm. The forewings are
buff and marked with two transverse, wavy brown bands and a sprinkling of
brownish dots. Full-grown larvae (fig. 79) may be bright yellow, with rusty-brown
heads, and 10 dark wavy lines running down the dorsum, or they may be entirely
light yellow. They are about 37 mm long.

Winter is spent in the egg stage and hatching occurs in April or May. Larvae are
present until July, then pupate in cells in the ground. Adults are present from
October to December. The females crawl up the trunks of trees and deposit their
eggs singly or in small groups in bark crevices. There is one generation per year.
Several outbreaks have been recorded in the Northeastern States and Canada. At
such times, male moths are attracted to street lights in nearby towns in considerable
numbers. Two dipterous parasites, Pseudotachinomyia slossonae (Townsend) and
Phryxe vulgaris Fallén, commonly attack the larvae. A virus epizootic occurred in
an outbreak in Quebec in 1961.

194

Courtesy Can. For. Serv., Can. Dep.
Environ., Sault Ste. Marie, Ont.

Figure 79.—Larva of the linden looper, Erannis tiliaria.

Larvae of Lycia ursaria (Walker), the stout looper, feed on the foliage of a
number of hardwoods such as paper birch, willow, elm, maple, poplar, basswood,
ash, pin cherry, and alder in eastern Canada and the Atlantic Coast States. Willow
and paper birch appear to be preferred. The adult is gray, stout-bodied, and has a
wingspread of about 45 mm. Both pairs of wings are crossed by diffused, blackish
lines. Full-grown larvae are about 35 mm long. There are four large whitish spots
on the front of the prothorax, and the gray body is marked with numerous
longitudinal, wavy red-brown lines and creamy-white spots. Larvae are present
from May to July, and winter is spent as pupae in the ground. This species is
occasionally quite common in New England.

Nacophora quernaria (J. E. Smith), the oak beauty, occurs on oaks in the
Atlantic Coast States and Pennsylvania, and on paper birch from New Brunswick
west to central Saskatchewan in Canada. Full-grown larvae are about 50 mm long.
The head is bilobed and angular; the body slate gray, stoutish, and armed with
brownish tubercles. The tops of the first two thoracic segments are also marked
with reddish brown and black. Larvae are found from June to September; winter 1s
spent in the pupal stage. The yellowish-green egg is iridescent and acorn-shaped
with a circle of white spots on the top (434).

Plagodis serinaria Herrich-Schaffer feeds on various hardwoods such as red and
sugar maples, yellow and paper birches, beech, and aspen in the Northeastern
States and eastern Canada. Full-grown larvae are about 37 mm long. The head is
bilobed and angular; the body, dull brown with blotches of lighter and darker shades
and a prominent swollen area on the back of the sixth abdominal segment. Winter is
spent in the pupal stage, and adults are present during May and June. P. kuetzingi
(Grote) feeds on ash in the Northeastern States, Nova Scotia west to Ontario and the
Lake States (434).

Probole amicaria (Herrich-Schaffer) and P. alienaria Herrich-Schaffer feed on
various hardwoods throughout much of the Eastern United States and from coast to
coast in southern Canada. Full-grown larvae are green to brownish and about 30
mm long. The head is rather small and flattened in front, and there are two whitish
spots on the tops of all body segments except the prothorax. Larvae are present from
July to early September, and winter is spent in the pupal stage.

The filament bearer, Nematocampa limbata (Haworth), feeds on the foliage of
hemlock, fir, and such hardwoods as maple, oak, gray birch, pin cherry, buckeye,

195

and apple in the Eastern United States and eastern Canada. Full-grown larvae are
greenish brown with reddish-brown heads and are about 18 mm long. There is a pair
of tubercles on the first abdominal segment; a pair of long, slender, brownish
filaments on each side of the second and third segments; a pair of small, rusty-red
tubercles on the eighth segment; and a broad stripe that extends from the dorsum of
the prothorax to the first pair of filaments. Larvae are active from May to July;
adults, from late June to August.

The elm spanworm, Ennomos subsignarius (Hubner) (fig. 80), occurs in south-
ern Ontario and throughout the Eastern United States, west to Wisconsin, Colo-
rado, and Texas. Its preferred hosts are hickory, oak, red maple, and ash but it
attacks a large number of other hardwoods (208). About 100 years ago, it was best
known as a shade tree pest in the larger cities of the Eastern United States (56/). It
also has been a predominant forest pest with widespread severe outbreaks occurring
in the southern Appalachians and Connecticut (395, 653).

The adult is a powdery white moth with a wingspread of 30 to 37 mm. Larval
color is related to population density (334). Full-grown larvae in outbreak areas are
usually dull, slate black except for rusty head capsules; a small proportion of the
larvae may be light green with yellow head capsules. When populations are low,
there is a higher proportion of the lighter colored larvae. They are about 50 mm
long.

Winter is spent in the egg stage. Hatching begins in late April in the South.
Farther north, it may not start until late May or early June. Young larvae feed on the
lower surfaces of leaves and produce a typical shothole effect. Older ones eat the
entire leaf with the exception of the midrib and petiole. To pupate, mature larvae
spin coarse, netlike cocoons of silken threads, often on partly eaten leaves. In
completely defoliated stands, cocoons may be spun on exposed branch tips, in leaf
axils, in bark crevices, or stumps of undergrowth.

The elm spanworm is capable of completely defoliating large areas of mixed
hardwood forests during outbreaks. Between outbreaks, populations often exist at
very low levels. A number of insect parasites and predators were found in fairly
large numbers during the latter stages of some of the outbreaks and probably helped
in bringing them to an end. The hymenopteran 7elenomus droozi Muesebeck
parasitized and destroyed over 80 percent of the eggs in certain areas (209, 336,
337, S95). In Connecticut, Ooencyrtus ennomophagus Yoshimoto, a par-
thenogenetic egg parasite with more than one generation per year, ended an
outbreak in about 2 years (654). The predacious beetle Calosoma scrutator (F.), a
voracious feeder on lepidopterous larvae, was also abundant in outbreak areas.

Ennomos magnarius Guenee, the notched-wing geometer, is widely distributed
in southern Canada and the Northern States. It feeds on many species of hardwoods
such as white ash, basswood, maple, quaking aspen, paper birch, beech, willow,
and elm. The adult is yellowish and has a wingspread of about 60 mm. The
forewing has a reddish tinge, is thickly flecked with brown dots, has a conspicuous
lobe near the middle, and is shaded with brown on the outer margin. Mature larvae
are about 50 mm long and yellowish green to dark brown or twiglike. Reddish areas
occur on the tops of segments two and five and on the venter of segment three.
Adults deposit oblong eggs in long, single strings during August and September.
and winter is spent in the egg stage. Hatching begins in May, and larvae are present
from May to July or August. This species is sometimes fairly abundant but is
seldom of economic importance.

196

SS <oY

D

A,B,C,D, F-531248
E, F-500619

Figure 80.—Elm spanworm, Ennomos subsignarius: A,
adults mating (male distinguished by feathery
antennae); B, eggs; C, light-colored larva; D, dark-
colored larva; E, pupae (light larvae and pupae
develop in low-density populations, darker from dense
populations).

Pero honestaria (Walker) occurs on black locust, pin cherry, and larch in the
Eastern States. Full-grown larvae are about 37 mm long and dark brown, with paler,
longitudinal markings. The head is bilobed and the body increases in girth toward
the rear. There appears to be two generations per year. P. morrisonaria (Henry
Edwards) occurs on a wide variety of trees, including fir, spruce, larch, pine,
willow, and aspen. The species is single-brooded and occurs from Nova Scotia
south to Virginia and west to the Pacific Coast (434).

197

The pine conelet looper, Nepytia semiclusaria (Walker), attacks loblolly, sand,
shortleaf, and slash pines from southeastern Virginia south and west to eastern
Texas. The mature larva is from 25 to 50 mm long. It is vividly marked with a
broad, brick-red dorsal stripe flanked by paired bright-yellow lateral stripes sepa-
rated by a series of fine black and white lines. Head, thoracic legs, and prolegs are
bright orange. The adult is grayish tan with a wingspread of from 25 to 30 mm.
Each front wing is crossed by a pair of brown scalloped lines while a single similar
line occurs on each hindwing. This species overwinters as an egg in masses under
bark scales. Hatching occurs early in the growing season and the young larvae feed
on the female flowers and conelets of slash, loblolly, and shortleaf pines and
possibly also sand pine. In sand pine, old-growth foliage is the principal food
(359). Pupae are found among the needles. N. pellucidaria (Packard) occurs on fir,
larch, pine, spruce, and hemlock in the Northeastern States.

The false hemlock looper, NV. canosaria (Walker), occurs on hemlock, fir, white
and black spruces, northern white-cedar, and larch in the Northeastern States and
southern Canada, often in conjunction wiin hemlock looper (/036). Full-grown
larvae are about 25 mm long. The head is whitish or reddish brown and bears a few
black dots; the body is whitish to green with tinges of yellow or red, black dotted,
and has a yellowish stripe on each side below which are four or five dark, wavy
lines. Larvae are present from June to August. The ellipsoidal eggs are truncated at
one end and laid on bark in small clusters (434).

The hemlock looper, Lambdina fiscellaria (Guenée), occurs from New-
foundland to Alberta in Canada and south to Georgia in the Eastern States. Its
preferred hosts are balsam fir, white spruce, and hemlock. It also feeds on many
other species during outbreaks such as larch, red and black spruces, jack pine,
basswood, maple, paper and yellow birches, elm, and cherry. In the more southerly
portions of its range, infestations develop mainly on hemlock. Outbreaks were
recorded in Ohio, Wisconsin, Michigan, New York, and Maine during the 1920's
and in Massachusetts, New Hampshire, and Vermont from 1949 to 1952. However,
severe, prolonged outbreaks have been recorded only in eastern Canada (/036).
The adult is creamy tan to grayish brown with a tinge of purple and has a
wingspread of about 62 mm. Two irregular purplish-brown lines cross the fore-
wings, and a dot of the same color lies between them near the costal margin. Full-
grown larvae are usually grayish green to grayish and are about 30 mm long. The
head and body are marked with distinct black dots.

Adults are present from around mid-August to early October and deposit their
eggs singly or in small groups on moss, lichens, or bark on limbs and trunks.
Winter is spent in the egg stage and hatching occurs in June. Young larvae feed on
opening buds or on old needles. Needles are often chewed off at the base and drop;
others that are partially chewed off, usually dry out, turn brown, then drop.
Pupation takes place in bark crevices, or in masses of lichens on or near the bark
(198).

Hemlock looper outbreaks may develop very suddenly, the most serious ones
occurring in mature and overmature hemlock and balsam fir stands.

The eastern pine looper, L. pellucidaria (Grote & Robinson), occurs in several
Atlantic Coast States and feeds on pitch, red, shortleaf, loblolly, pond, and possibly
other pines. Several widespread outbreaks have been recorded during the past 70-
odd years. The adult is ash gray to smoky in color and has a wingspread of about 37
mm. The forewing is crossed by two irregular diffuse dusky lines and bears a
slightly sinuate discal dot. The hindwing is also crossed by a dusky line. Full-grown

198

larvae are pale straw to yellow with black markings and are 27 to 37 mm long. The
body bears faint rows of blackish dots and short wavy lines on the top and sides, and
the head is densely marked with light and dark spots.

Adults emerge in May and June and lay their eggs on the needles. The larvae feed
on the needles until late September. Winter is spent in the pupal stage in the duff
beneath the trees.

Lambdina fervidaria athasaria (Walker) occurs on hemlock, maple, oak, and
beech in several Eastern States. Local outbreaks have occurred in Massachusetts,
Connecticut, and Ohio. The adult resembles that of L. pellucidaria except for a
slightly shorter wingspread. Full-grown larvae are yellowish and about 30 mm long.
The head is marked with irregular brown to blackish spots. The top of the body is
lighter than the sides. The sides are marked with wavy lines of dark or reddish
brown, interrupted with dashes of white. Winter is spent in the pupal stage on the
ground, just beneath the top crust of the leaf mold. Larvae are present from July to
late September.

The chainspotted geometer, Cingilia catenaria (Drury), occurs in eastern
Canada and the Northeastern States. It has many hosts including gray and paper
birches, oak, poplar, willow, cherry, balsam fir, larch, and white spruce. Young
eastern white and red pines growing in mixture with the above hosts are also subject
to heavy defoliation. Blueberries, huckleberries, and small trees growing in pas-
tures and cutover areas are especially subject to infestation. Local outbreaks have
been recorded frequently in the Northeastern States.

The adult has a wingspread of 30 to 42 mm. The head and part of the thorax are
orange-yellow; the body, white with black markings. The wings are smoky white
and black spotted. Full-grown larvae are straw colored and about 50 mm long. The
head and body are dotted with black spots, those on the sides of the body produce
chainlike effects. There also are three or four thin, black lines below these rows of
dots (/037). Winter is spent in the egg stage. Larvae are present from June to
August.

Tetracis cachexiata Guenée occurs on cherry in the Northeastern States. Full-
grown larvae are about 37 mm long. The head is grayish, flattened, and square in
front; the body, reddish brown to black with white markings and a black line
running down the middie of the back after the fifth segment. Tubercles are also
prominent. Larvae are present from July to September, and winter is spent in the
pupal stage.

Eutrapela clemataria (J. E. Smith), the purplish-brown looper, is multivoltine
and occurs on basswood, maple, quaking aspen, paper birch, pin cherry, sweet-
gum, and hemlock in eastern Canada and the Eastern United States, westward to
the Mississippi River Valley. Full-grown larvae are purplish brown and up to 60 mm
long. The head is rounded and bilobed, the second and fourth abdominal segments
are swollen on top, and there are prominent tubercles on the fifth and ninth
abdominal segments. Larvae feed from June to August in the North, and from April
to October in the South. Winter is spent as pupae in cocoons in leaves on the
ground. The truncate eggs are laid erect in large masses, much like those of
Alsophila (434).

Prochoerodes transversata (Drury), the large maple spanworm, occurs on
various hardwoods such as maple, oak, willow, quaking aspen, paper birch, and
mulberry in southern Canada and the Eastern United States. Full-grown larvae are
light to purplish brown and about 50 mm long. The head is rounded and flattened in
front, the second thoracic segment is swollen and streaked with red, and the eighth

199

abdominal segment bears a pair of prominent tubercles. Larvae feed from June to
October, winter is spent in the pupal stage, and there are two generations per year.

The pepper-and-salt moth, Biston betularia cognataria (Guenée), occurs in
eastern Canada and the Northeastern States where it feeds on elm, willow, poplar,
black locust, cherry, and apple. The adult is dark brown to black and has a
wingspread of about 55 mm. The light-gray forewings are suffused with gray-brown
and have dark-brown to black crosslines (783). Full-grown larvae, commonly called
cleft-headed spanworms, are 50 to 75 mm long. The head is deeply cleft,
granulated, and flat in front; there are tubercles on each side of the prothorax and on
the fifth and eighth abdominal segments (/037). Larvae feed from July to October,
and winter is spent in the pupal stage in the litter.

Campaea perlata (Guenée), the fringed looper, is a common species in the
Northeastern United States and Canada, and the larvae feed on balsam fir, hemlock,
and many species of deciduous trees. Adults have whitish wings; the forewings are
pointed at the apex and have two gray crosslines; the hindwings are scalloped on the
outer margin (754). The full-grown larva is 20 to 25 mm long and light brown to
grayish with dark-brown markings (754, 1/036). Protoboarmia porcelaria indica-
taria (Walker) is also a common species in northeastern forests. Adults are gray.
Larvae feed on a wide variety of trees including balsam fir, white spruce, larch, jack
pine, birch, quaking aspen, and willow.

Family Mimallonidae
Mimallonid Moths

Adults of this family are stout-bodied moths with pectinate antennae. The
forewings are falcate, bent at the middle, and heavy-veined. The humeral angle of
the hindwing is much enlarged, and the frenulum is rudimentary. Two species are
known to occur in the Eastern United States, neither of which is of economic
importance. However, they both often attract attention.

Cicinnus melsheimeri (Harris), Melsheimer’s sackbearer, is fairly common on
oak, especially bur oak, from New England to the Lake States and southward. The
adult is reddish gray and has a wingspread of 37 to 50 mm. The body is sprinkled
with minute black dots; each wing 1s crossed by a narrow blackish band and marked
by a black discal spot or bar. A newly hatched larva makes a shelter for itself by
drawing two leaves together with strands of silk. Eventually, the larva constructs an
ellipsoidal portable case from pieces of leaves and silk, leaving a circular hole at
each end. It lives in the case but can leave it at will. To move the case, it bites off
the strand of silk that anchors it and transports it to the new location. When it 1s at
rest, the larva anchors the case with silk and plugs the openings of the case with its
head and rear end. Winter is spent in the pupal stage, and adults appear during May
and June.

Lacosoma chiridota Grote feeds on oak throughout much the same range as
Melsheimer’s sackbearer. Adults are dark yellowish brown, with deeply scalloped
forewings, and have wingspreads of 25 to 30 mm.

Family Apatelodidae
Apatelodids

This family is represented in North America by only five species, two of which
are encountered fairly often in eastern forests (442).

Apatelodes torrefacta (J. E. Smith) occurs throughout the eastern part of the
United States and feeds on maple, black cherry, and various other trees and shrubs.
Full-grown larvae have rounded yellowish heads and are about 50 mm long. The
body is whitish to yellow on the dorsum, except for a row or line of more or less

200

connected black spots. It has a row of black spots on each side, is blackish beneath,
and is densely clothed with long, fine, white or yellow hairs. Long pencils of hairs,
pale at the base and black at the tip, arise from the second and third thoracic and
eighth abdominal segments. Larvae are present from June to September, and winter
is spent in the pupal stage on the ground.

Olcerlostera angelica (Grote) larvae feed on the leaves of ash and lilac from New
England and southern Canada to Florida and westward into the Ohio River Valley.
Full-grown larvae are 50 to 62 mm long. The head is rounded, brown, and mottled
with light and dark shades. The body is gray and covered with a network of fine
wavy black lines, except for the top of the thorax that bears two broad black
transverse bands. The tops of segments one and seven bear yellowish-green spots,
long brown and white hairs project forward over the head from the prothorax, and
whitish hairs project forward from the middle of the second and third thoracic
segments. Most of the rest of the body is sparsely clothed with short white and
black hairs. Larvae are found in August and September. Winter is spent in the pupal
stage on the ground.

Family Lasiocampidae
Tent Caterpillar Moths and Allies

The family Lasiocampidae is represented in North America by about 37 species,
several of which are important economic pests of trees (442). The moths are
medium-size and stout-bodied; the body, legs, and eyes are hairy; and the antennae
are somewhat feathery. The larvae vary in form from nearly cylindrical to very
much flattened; they are very hairy.

The majority of important species belong to the genus Malacosoma, and are
commonly known as tent caterpillars. Four and a possible fifth species occur in
the Eastern United States and eastern Canada. The remaining 11 North American
species have a western distribution. A revision of the genus 1s available (//49). This
includes keys to the adults, mature larvae, and egg masses of all species and a
discussion of their distribution, their hosts, and some of their habits, and parasites
recovered. A complete listing of the various species is also available (/344).

Eggs of tent caterpillar moths are laid either in flattened masses on the bark of
limbs or trunks of trees or in masses that may encircle small twigs. The number of
eggs per mass ranges from 100 to 400, usually from 150 to 250. As the eggs are
deposited, they are held in place by a frothy substance called spumaline (579). The
majority of species cover their eggs with this material. Hatching occurs in the
spring, about the time the new leaves of the host tree appear. Young larvae feed first
on egg shells, then on the buds and young leaves. Those of tent-building species
also immediately begin to construct a tent on a branch or in a nearby tree crotch.
They do not feed from within the tent but on the leaves of neighboring branches. As
they crawl to these branches, they spin strands of silk that they usually follow in
returning to the tent, where they remain during periods between feedings. Species
that do not construct tents assemble in clusters on branches and trunks during
periods between feedings. Toward the end of an instar they spin nests of silk on
branches and trunks on which they congregate to molt.

The larvae usually pass through five or six instars. By the time the last instar is
reached, they are no longer gregarious, and they travel extensively in search of
food. At this time, they are not very selective in their food requirements and will
feed on a wide variety of hosts. When their food supply becomes scarce, they may
migrate in search of other food, often for considerable distances. When they reach
maturity, they spin cocoons 1n which to pupate. The cocoons are about 25 mm long,

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fairly loosely constructed, and white or yellow because of a powdery material
dispersed between strands of silk. Cocoons may be found within the old tents,
inside logs, beneath loose bark, and between folded leaves.

The eastern tent caterpillar, M. americanum (F.) (fig. 81), 1s generally dis-
tributed throughout the eastern half of the United States and southern Canada. Its
preferred hosts are cherry and apple, but it also attacks a wide variety of other
forest, shade, and fruit trees. The adults are light to dark chocolate brown, the
wings are lightly dusted with white scales, and the wingspread varies from about 37
to 50 mm. Each forewing is crossed by two oblique white or yellowish-white lines.
The hindwing is uniformly chocolate brown and crossed by a faint white area. Full-
grown larvae have black heads, sparsely clothed with long, fine, light-brown hairs,
and are marked with an apparently continuous middorsal light stripe, bordered on
each side with longitudinal reddish-brown and black wavy lines. The subdorsal area
is marked with a central black area on each segment, crossed by a vertical blue
mark posteriorly.

A, courtesy Can. For. Serv., Can. Dep. Environ.,
Sault Ste. Marie, Ont.

B, F-532847
C, D, courtesy Conn. Agric. Exp. Stn.

Figure 81.—Eastern tent caterpillar, Malacosoma
americanum: A, adult male and female; B, egg
masses encircling twig; C, larvae; D, a typical tent.

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Winter is spent in the egg stage, and hatching occurs about the time the buds of
the host tree begin to unfold in the spring. In Florida, adults may appear and lay
their eggs even before hatching occurs in the northern parts of the insect’s range.
The larvae are gregarious. As soon as they hatch, they begin the construction of a
tent in a nearby trunk or branch crotch, and continue to enlarge the tent as they
grow. From this tent, the larvae crawl out to the foliage to feed. After feeding, they
return to the tent to rest. When they become full grown, they leave the nest and
wander in search of places to pupate. Pupation occurs in tough silken cocoons,
dusted with a yellowish powder, on the bark of trees, on fences, on brush and
weeds, among dead leaves and other debris on the ground, and even on the sides of
buildings. When the adults appear, they lay eggs in essentially a clasping mass on
small twigs or branches, or on the trunks of small trees. In the Lake States, eggs are
often found on the trunks of very small trees about 15 cm above the ground. There
is One generation per year (/56, 302).

Most of the hosts of the eastern tent caterpillar have little value, thus it usually
does not cause economic losses. It may be of some importance, however, when it
defoliates commercial-size black cherry whose wood is of value for furniture (700).
The species is primarily a nuisance pest. Infested trees in parks, recreational areas,
along roadsides, and in the vicinity of homes may be disfigured.

During most years, the eastern tent caterpillar is controlled satisfactorily by its
natural enemies. Egg parasites may account for one-quarter of the exposed eggs,
but only | percent of the spumaline-covered eggs (276). Periodically, however,
populations reach outbreak proportions. An effective method of control on isolated
trees is to prune off and burn twigs containing egg masses. Destroying larval tents,
preferably when the tents are still small, is also effective. Bacillus thuringiensis and
a number of chemical insecticides are effective in controlling the larvae.

Malacosoma californicum lutescens (Neumoegen & Dyar), the prairie tent
caterpillar, occurs throughout the Great Plains area east of the Rocky Mountains to
central Texas. Its hosts are recorded as common chokecherry, willow, American
plum, and gooseberry. Male adults range from dark reddish-brown to very light
yellow; females are yellowish to medium reddish-orange brown. The forewings are
crossed by light yellowish lines, and the wingspread is about 37 to 50 mm. Full-
grown larvae are about 50 mm long. The head is blue, mottled with black, and
sparsely covered with fine whitish to orange setae. The middorsal area of each
abdominal segment is marked with an elongate somewhat pointed, blue-white dash.
These dashes combine to form a broken middorsal stripe.

Eggs are laid in flattish, clasping masses on twigs and branches and are covered
with light-brown or grayish spumaline. The larvae construct relatively large tents
that look like those of the eastern tent caterpillar. Defoliation is usually confined to
branches, but during outbreaks trees may be completely defoliated. Because of the
low value of its hosts, the species is of minor economic importance.

The Sonoran tent caterpillar, M. tigris (Dyar), occurs in the southern Great
Plains, southern Rocky Mountains, the Southwest, and Mexico. So far, it is not
known to occur farther eastward than central Texas. Its host plants are various oaks
and possibly other species. Eggs are laid in encircling bands on very small twigs,
occasionally dead twigs. The masses differ from those of other members of the
genus in not being covered with spumaline. The larvae construct relatively small
tents on which they congregate to molt. Tents are usually formed near the end of
each instar. This species does not appear to be of much economic importance,

203

unless it occurs in association with other defoliating species. It has combined with
the forest tent caterpillar in heavily defoliating oaks in south-central Texas.

The western tent caterpillar, M. californicum (Packard), occurs primarily in
western America. However, spotted infestations also occur eastward across central
Canada to Quebec. It has also been collected in New Hampshire, New York, and
northwestern Minnesota (//49). Alder, serviceberry, willow, common chokecherry,
birch, apple, plum, cherry, and quaking aspen are known to be attacked. Full-grown
larvae are predominantly black and yellow or yellow-orange. The head is mottled
blue-black and sparsely covered with fine yellow-orange setae. The dorsum of the
abdomen is marked by a stripe formed by a series of elongate bluish-white,
somewhat pointed dashes, one per segment.

Eggs are laid in flat, clasping masses covered with brown or dark-brown
spumaline. The tents are similar to those of the eastern tent caterpillar. The species
is of little or no economic importance in eastern America. Larvae may be controlled
with Bacillus thuringiensis (S63).

The forest tent caterpillar, M. disstria Hubner, occurs throughout most of the
United States and Canada and feeds on a wide variety of hardwoods (7/). In the
North and West, quaking aspen is preferred; in the South, water tupelo, blackgum,
sweetgum, and various species of oak are most heavily defoliated. The adult is
stout-bodied, light buff-brown, and has a wingspread of about 25 to 37 mm. The
forewings have two darker oblique bands near the middle (fig. 82A). Full-grown
larvae have light-blue heads mottled with black and sparsely covered with fine,
whitish setae. Each abdominal segment is marked dorsally with a yellowish-buff,
keyhole-shaped spot which may be divided to form an anterior spot and a smaller |
posterior spot (fig. 82B). The venter is blue-gray to dark gray, usually with a
median spot on each segment, and often with a dark-gray area running the full
length of the body between the bases of the legs.

Winter is spent as a complete embryo within the eggshell and hatching occurs in
the spring, about the time the buds on the host tree begin to swell. Young larvae
feed on expanding buds; older ones devour the foliage, often completely defoliating
the tree. During the first four to five instars, the larvae are gregarious. At first, all of
those from one egg mass cluster on one leaf or one group of small expanding
leaves. Later, they become more widely dispersed on surrounding foliage. Al-
though this species is referred to as a tent caterpillar, it does not construct tents.
However, larvae do lay down strands of silk along which they travel. They also form
silken mats on the trunks or branches on which they congregate in masses to rest or
to molt (fig. 82C). As they approach maturity, they tend to wander individually over
the trees and other vegetation in search of food or places to pupate. Pupation occurs
in pale-yellow cocoons spun in folded leaves, in bark crevices, on shrubs or other
vegetation, and occasionally on buildings. Adults appear from late May in the
South to late June and July in the North. Eggs are laid in masses of 100 to 350 in
bands 25 to 37 mm wide that encircle twigs up to 37 mm in diameter. The eggs are
cemented together and are coated with dark-brown spumaline. There is one genera-
tion per year.

The forest tent caterpillar has been an important enemy of forest, orchard, and
shade trees for many years. During the period from 1886 to 1962, several general
outbreaks, some of which covered thousands of square kilometers and lasted for 2
to 6 years, occurred in the Northern United States and eastern and western Canada.
Since the 1930’s several outbreaks have also occurred in Virginia, South Carolina,
Mississippi, and Louisiana (577). Outbreaks occurred repeatedly in Alabama

204

A and B, Courtesy Conn. Agric. Exp. Stn.;
C, F-506692
Figure 82.—Forest tent caterpillar, Malacosoma disstria:
A, adults; B, full-grown caterpillar, showing keyhole
spots along the dorsum; C, caterpillars at rest on trunk
and branches of an aspen.

during the 1960’s, mostly defoliating water tupelo on a gross area of 17,000
hectares (2/3, 1/47). Damaging populations arose again in 1981, including Loui-
siana, North Carolina, Kentucky, and Alabama (58/). In general, tree mortality has
not been severe, but losses in reduced growth following defoliation have been great.
Studies in Minnesota indicate that there 1s about a 70 percent reduction in basal area
growth of aspen during the first year of heavy defoliation. About 90 percent
reduction also occurs during the second year of heavy defoliation, plus about 15
percent reduction during the year of recovery. Total reduction for the 3-year period
averages about 58 percent (349). Heavy defoliation of bottom-land gums in the
South results in substantial mortality and dieback, as well as severe reduction in
annual growth. In the Northeast, heavy defoliation in sugar maple orchards not only
causes serious injury to the trees, but also a reduction in the quantity and quality of
the sap. Outbreaks in recreational areas adversely affect business because of the
nuisance created by migrating caterpillars and the midwinter appearance of defoli-
ated trees during the tourist season. Discussions of the effects of defoliation on
quaking aspen in Canada are published (570, /035). Outbreaks have been associ-
ated in eastern and central Canada with a cool winter and warm spring 2 to 4 years
earlier (628).

Outbreaks usually subside after 3 or 4 consecutive years of defoliation. Several
adverse environmental factors are responsible for population declines. Mortality of
pharate larvae, possibly caused by low winter or spring temperatures, may be quite
high (1/279, 1346, 1347). Harsh weather conditions may cause death of large
numbers of the early instars. Excessively high temperatures later in the spring may
kill large numbers of adults and seriously reduce the viability of newly laid eggs.
Mortality of the late instars may be severe or complete as a result of starvation in
heavily or completely defoliated stands. A polyhedrosis virus disease sometimes
kills enormous numbers of larvae in the late stages of outbreaks (//45). The
sarcophagid parasite, Sarcophaga aldrichi Parker, often becomes extremely abun-
dant during the late stages of outbreaks in the Lake States and greatly aids in
termination of outbreaks (576). Other natural enemies include many species of
insect parasites (1/345). Bacillus thuringiensis and various chemical insecticides
may be used to control larvae (527).

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Tolype velleda (Stoll), the velleda lappet moth, occurs from southern Canada
south through the Atlantic States to Florida and west to the Lake States. The larvae
feed on the foliage of various hardwoods including ash, quaking aspen, basswood,
cherry, elm, maple, holly, oak, and apple. The adult has a white head and thorax, a
gray abdomen, and a large blackish spot in the middle of the dorsum. The wings are
usually gray, sometimes dusky, are crossed by white lines, and have a wingspread
of 37 to 62 mm. Full-grown larvae are gray with faint longitudinal lines and are
about 62 mm long. The body is flattened and has lateral lappets, each of which has
many long hairs, thus forming a fringe along each side of the body. There is a pair
of warts, bordered posteriorly by a velvety black band, on the metathorax. Adults
appear in September and October. Females lay their eggs in rows and cover them
with hairs from the abdomen. Larvae are present from June to August. Pupation
takes place in tough, flattened, parchmentlike cocoons on the bark. There is one
generation per year. This species seldom causes serious injury. The related but
smaller species, 7. /aricis (Fitch), feeds on larch. Its range and life cycle are similar
to those of T. velleda.

The lappet moth, Phyllodesma americana (Harris), 1s generally distributed, but
not often common, through the Eastern States and southern Canada. Its hosts are
quaking aspen, cherry, hickory, birch, maple, oak, and various other hardwoods.
Adults are reddish brown and have wingspreads of 30 to 50 mm. The inner angle of
the forewing and the costal margin of the hindwing are deeply notched, and there is
a pale band edged with irregular dark-brown lines beyond the middle of each. Full-
grown larvae are bluish gray, somewhat mottled above, have lateral lappets, and are
about 62 mm long. There are transverse scarlet bands on the second and third
thoracic segments, in each of which are three black dots. Winter is spent in the
pupal stage in a tough, flattened cocoon, usually on the bark of the tree. There is

one generation per year in the North and a partial second one in the South.
Family Saturniidae
Giant Silkworm Moths

This family contains some of the largest and most colorful moths in the Eastern
United States (226, 403). The larvae, also large and conspicuous, are more or less
armed with tubercles and spines. Because of their habit of spinning large, dense
silken cocoons in which to pupate, they are known as giant silkworms. The larvae
feed on a wide variety of trees and shrubs; however, because they usually occur
singly and in low numbers, they are seldom injurious. Some of the most common
Species are discussed briefly below.

The cynthia moth, Samia cynthia (Drury), was introduced into this country from
Asia more than 100 years ago and now occurs in urban areas from Massachusetts to
Georgia and west to Indiana. Its preferred host is ailanthus, but cherry, plum, and
other hosts are also infested. The adult is olive brown with rows of tufts of white
hairs on the abdomen, and has a wingspread of 150 to 200 mm. Full-grown larvae
are about 75 mm long and have yellowish-green heads and a whitish bloom dorsally
on the body. The rest of the body is light bluish green to yellowish, spotted with
black, and there are long, bluish tubercles with short bristles on each body
segment. Moths are present from June to August; larvae from July to October.
Winter is spent as pupae in pendulous cocoons on the tree or on the ground.

The cecropia moth, Hyalophora cecropia (L.), occurs throughout the Eastern
United States and in southern Canada. The larvae feed on a wide variety of
hardwoods such as apple, ash, birch, cherry, hawthorn, walnut, maple, sassafras,
willow, elm, poplar, and basswood. Feeding damage can be severe in shelterbelt

206

plantings in the northern Great Plains. The moth has a wingspread of 125 to 150
mm. There is a white, crescent-shaped spot near the center of each wing, a red-
bordered crossband on each wing, and a dark spot near the end of each forewing.
They fly from April until June, depending on location. Full-grown larvae are 75 to
100 mm long. The head is green with two black spots on each side, and the body is
pea green. There are pairs of large, coral-red tubercles dorsally on the second and
third thoracic segments, 15 yellow tubercles on top of the first to eighth abdominal
segments, and blue tubercles on each side. All tubercles bear stiff, black bristles.
Winter is spent in large, thick, tough, gray-brown silken cocoons firmly fastened
lengthwise to bare branches. Larvae may be found from June to October, depending
on location. The tachinids Lespesia samiae (Webber) and Winthemia cecropia
(Riley) are common parasites in the Northeast.

The promethea moth, Callosamia promethea (Drury), occurs throughout most
of the Eastern United States and southern Canada, and feeds on a variety of hosts
such as ash, cherry, lilac, sassafras, spicebush, yellow-poplar, maple, and birch.
The adult has a wingspread of about 75 mm. The wings of the female are reddish
purple to brown with light-brown borders, and are crossed near the middle with a
wavy white line; each bears an angular white spot near the middle. Each forewing
also bears an eyelike spot near the apex. A full-grown larva is about 50 to 62 mm
long. The head 1s small and yellow; the body bluish or greenish white. The second
and third thoracic segments each bear a pair of large, coral-red tubercles or horns;
the eighth abdominal segment bears a large yellow one. Each segment is also
ornamented by a series of black buttons. Winter is spent in a tough, light-colored
cocoon enclosed in a leaf. The petiole of the leaf is attached to the twig by a very
strong band of silken threads.

The luna moth, Actias luna (L.), occurs from southern Canada to Florida and
Texas and has many hosts such as beech, birch, persimmon, sweetgum, willow,
oak, hickory, black walnut, eastern hophornbeam, and butternut. The adult moth, a
very beautiful insect, has delicate green wings that may spread to well over 100
mm. Each forewing bears a conspicuous eyelike spot and is edged with a purplish-
brown band. The hindwings extend into long, curved swallowtails. The full-grown
larva is about 75 mm long. The head is bluish green with brown on the sides; the
body, pale green with six pinkish or greenish tubercles armed with bristles arising
from each segment. A pale-yellow line runs along each side of the body. The tips of
the dorsal tubercles on the second and third thoracic segments are red; the re-
mainder, yellow. Winter is passed in a cocoon, usually on the ground. Moths are
present from April to September, depending on location. There may be two or even
three generations per year in the South.

The polyphemus moth, Antheraea polyphemus (Cramer), is widely distributed
throughout the United States and Canada and feeds on many species of trees
including basswood, beech, birch, elm, hawthorn, hickory, maple, oak, yellow-
poplar, willow, ash, butternut, walnut, sassafras, and sycamore. The moth is
yellowish brown and has a wingspread of 100 to 150 mm. There is an eyelike
transparent spot and a sooty, transverse stripe outwardly edged with light pink near
the outer margin of each wing. Full-grown larvae are apple green and about 75 mm
long. The head is reddish brown, and the thoracic shield is sometimes margined
with yellowish green along the front. Body segments are angular on the back, and
each bears six orange or golden tubercles from each of which arise one to three
bristles. The last segment bears a purplish-brown V-shaped design. Winter is spent
in a tough, thick cocoon usually enclosed in a leaf attached to a twig or on the

207

ground. Larvae may be found from June to October. The tachinid Lespesia
sabroskyi Beneway is a common parasite everywhere.

The io moth, Automeris io (F.), occurs throughout the Eastern United States and
attacks a wide variety of trees including paper birch, cherry, black locust, quaking
aspen, willow, beech, apple, maple, oak, hickory, elm, mulberry, dogwood, and
sycamore. Female moths are purplish red and have a wingspread of about 75 mm;
males are yellowish and slightly smaller. There is a large, circular, black eyespot
with a tiny white center on the upper surface of each hindwing and a smaller but
similar one on the lower surface of each forewing. Small larvae are gregarious. Full-
grown larvae are about 65 mm long. The body is pale green with a broad, reddish-
brown stripe on each side, margined with white and reddish lilac. Whorls of
branched, black-tipped, green, poisonous spines rising from small conical tubercles
on each body segment cause a severe nettling effect when they come into contact
with the skin. Moths are present from May to September; larvae, from July to
October, with two broods in the South. Winter is spent in the pupal stage in a thin,
brown, oval cocoon, often covered with bits of dead leaves and other debris, and
usually on the ground.

The buck moth, Hemileuca maia (Drury), occurs from Maine and Wisconsin to
Florida, Louisiana, and Texas, and it apparently feeds exclusively on certain
species of oak. Adults fly mostly in October, and have black, thin-scaled, some-
times semitransparent wings. A common white band crosses both the forewing and
hindwing near the middle. Small larvae are gregarious. Full-grown larvae are about
62 mm long. The head is deep reddish-brown; the body, dull brownish to black and
covered with small, yellowish dots. Each body segment has tufts of bristles or
compound spines arising from tubercles. The spines cause a nettling effect when in
contact with the skin. Winter is passed in the egg stage, and larvae are present from
May to August. A common parasite is the tachinid Leschenaultia fulvipes (Bigot).
H. lucina Henry Edwards feeds principally on spirea in the Northeast. H. nevaden-
sis Stretch, the Nevada buck moth, is mainly a western species that feeds on
poplar and willow. It occurs as far east as Wisconsin, Illinois, Nebraska, and Texas.

Royal moths constitute a subfamily of the Saturniidae and are medium-size to
large, with stout bodies and large, strong wings (/029). The head is generally
sunken in the prothorax, and the male antennae are feathery for only a little more
than half their length. The larvae are armed with horns or spines, and some are
sparsely hairy. The horns or spines on the second and sometimes third thoracic
segments are long and usually curved. The larvae feed on the foliage of various
trees, and they pupate in the ground without forming cocoons.

The spiny oakworm, Anisota stigma (F.), feeds on oak and perhaps hazel from
southern Ontario and Massachusetts to Florida and Texas. The adult has a wing-
spread of 45 to 62 mm. The wings are yellowish to rusty brown, sometimes tinged
with pink, and heavily speckled in both sexes. Each forewing has a white spot. Full-
grown larvae are about 37 to 50 mm long. The body is tawny brown, often tinged
with rose or pink. It is covered with tiny ivory-white specks, denticles, or granules,
and is marked indistinctly with single dorsal and lateral stripes. There are two long,
curved spines on the second thoracic segment. The remaining segments bear
backward-pointing black spines.

In the North, adults are present in June and July and the larvae from July to
September. The winter is passed as a pupa in the ground, and there is one
generation per year. This species is normally not very abundant, but infestations
have occasionally covered several hectares.

208

The orangestriped oakworm, A. senatoria (J. E. Smith), feeds on various oaks
from eastern Canada to Georgia and westward to Minnesota, lowa, and Texas. It is
rare in the Southeast. The adult has a wingspread of 37 to 50 mm, and its thick body
is covered with yellowish-red hairs. The female (fig. 83A) looks like a small,
faded, spiny oakworm, and the smaller, darker male has translucent areas in the
forewings. The hindwings of the male are distinctly angular and only about two-
thirds as long as the body. Full-grown caterpillars are black, with eight longitudinal
orange-yellow stripes on the dorsum and sides; the larvae are about 40 mm long
(fig. 83B). There is a pair of black, slender, stiff, erect, blunt, recurved spines on
the second thoracic segment. Each succeeding segment bears a number of small,
sharp, black spines.

ea | Courtesy Conn. Agric. Exp. Stn.
Figure 83.—The orangestriped oakworm, Anisota
senatoria: A, adult and eggs; B, larva.

In the North, adults appear during June or July and deposit their eggs in clusters
of several hundred each on the undersides of leaves. Young larvae feed in groups on
each side of the leaf, consuming everything but the veins. Older larvae are less
gregarious and are often seen crawling around on lawns or the sides of houses, or
feeding singly on the foliage of their host. During September or October, they crawl
to the ground and often do considerable wandering in search of suitable places to
pupate. Pupation takes place in the soil at a depth of 8 to 10 cm. There is one
generation per year, possibly two in the South. This species has been responsible
for a considerable amount of defoliation in oak stands in Connecticut, Missouri,
New Jersey, and Pennsylvania. A similar species, A. peigleri Riotte, occurs com-
monly in the Piedmont of North Carolina, South Carolina, and Georgia (fig. 84).

The pinkstriped oakworm, A. virginiensis (Drury), occurs from southern Can-
ada southward to Arkansas and Virginia. The larvae feed principally on various
oaks, but also, perhaps, on many other hardwoods such as chestnut, hazel, maple,
and birch. There is one generation per year. Adults (fig. 85) are yellow to reddish

209

brown, often with a purplish cast. Females have a wingspread of about 57 mm,
males are considerably smaller. The forewings are thinner, less speckled, and more
transparent beyond the discal dot than those of the orangestriped oakworm or the
spiny oakworm. Full-grown larvae are about 50 mm long. The body is greenish
gray with two dorsolateral, rose-colored stripes and a similarly colored stripe along
each side. The entire body is covered with minute, white granules. Two spines on
the second thoracic segment are slightly curved and are conspicuously longer than
the others. Two similar species, A. pellucida (J. E. Smith) and A. discolor Fer-
guson, have two generations per year and occur on oaks throughout the Southeast
and in Texas and Oklahoma, respectively.

F-532020

Figure 84.—Larva of Anisota peig/eri found on chestnut
in North Carolina.

Courtesy Duke Univ. Sch. For.

Figure 85.—Adult of the pinkstriped
oakworm, Anisota virginiensis.

The greenstriped mapleworm, Dryocampa rubicunda (F.), 1s found throughout
most of the Eastern United States and in adjacent areas of Canada. Its preferred
hosts are maples, but it also feeds on various oaks and boxelder, especially where
they are growing in mixture with maple. Populations may become heavy enough to
cause serious defoliation anywhere within its range, but this is most likely to occur
in the South. The moth (fig. 86A) has a woolly body and a wingspread of 37 to 50
mm. The body is yellow on top and rose-pink beneath. The forewing is rose-pink
on the inner and outer borders with a yellow band between. The hindwings are
either pure yellow, or yellow with rose-pink streaks. Pale, whitish forms are also
known. Full-grown larvae (fig. 86B) have cherry-red heads, pale yellow-green

210

bodies, and are about 37 mm long. The body also has seven dark-green lines
running its entire length, two prominent, slender horns on the second thoracic
segment, two rows of short spines on each side of the body, and four larger spines
on the terminal abdominal segments.

Courtesy D. C. Allen,
SUNY, Coll. Environ. Sci. & For.
Figure 86.—The greenstriped mapleworm, Dryocampa
rubicunda: A, male and female adults; B, fifth instar
larva; C, egg cluster.

There is one extended generation per year in the North and two in the South.
Eggs (fig. 86C) are laid on the undersides of leaves during May and June and hatch
in about 10 days. Larvae feed singly, devouring entire leaves, and become full
grown in about a month. With two generations in the South, trees may be defoliated
twice in the same season. Full-grown larvae crawl to the ground and form cells in
the soil or duff where they pupate and spend the winter.

Several species of parasites of the greenstriped mapleworm have been recorded,
but never in any great abundance. Birds devour some of the larvae but are probably
not very effective in population suppression (9, /327/).

The regal moth, Citheronia regalis (F.), occurs throughout the Southern States,
northward to Hlinois and Massachusetts. The larva, known as the hickory horned
devil, feeds on a wide variety of plants, including many species of trees such as
hickory, walnut, butternut, persimmon, sweetgum, sycamore, ash, and sourwood.
Adults are quite large; males have wingspreads of 100 to 125 mm, and females, 125
to 150 mm. The head and body are orange with pale-yellow markings; the fore-
wings are olive-gray with reddish-brown veins and yellow spots and the hindwings
are orange-red with somewhat redder veins. Full-grown larvae are 100 to 125 mm
long and are startling to behold (fig. 87).

Adults appear in June and larvae are present from July to September. Winter is
spent in the pupal stage in the ground. In most of its range there is only one
generation per year, but in the Deep South there may be a partial second. This

211

Courtesy Conn. Agric. Exp. Stn.
Figure 87.—Hickory horned devil, larva of Citheronia
regalis.

species is of little or no economic importance. It is of interest mostly because of the
frightening appearance of the larvae.

Citheronia sepulcralis Grote & Robinson is occasionally found feeding on the
needles of pines from Maine to Florida. Moths are dark brownish gray with a lilac
tinge. There is a dusky discal spot on each forewing and a reddish base on the
hindwing. The wingspread is about 75 to 100 mm. Full-grown larvae are dull
brown, armed with short, orange horns, and are about 100 mm long.

The imperial moth, Eacles imperialis (Drury), occurs in the Eastern United
States and southern Canada. The larvae feed on the foliage of a wide variety of trees
including pines, redcedar, oaks, sweetgum, elm, persimmon, hickory, maple,
beech, honeylocust, and baldcypress. The moth is sulfur yellow with brown and
lilac markings and has a wingspread of 100 to 150 mm. Full-grown larvae are
heavy-bodied and about 100 mm long. The head is orange-yellow with green sides;
the body is pale green to dark green or reddish brown and sparsely covered with
long, whitish hairs; it has short tubercles on the second and third thoracic and the
last two abdominal segments. Adults appear during June and July and larvae are
present from July to October. Winter is spent in the pupal stage in the ground; there
is One generation per year in the North, two in the South.

Eacles imperialis pini Michener feeds on eastern white and jack pines in New
York, Michigan, and southern Canada. The adults are smaller than those of the
imperial moth and have dark markings of pink to pinkish brown. Spots on the wings
are also heavier than those of the imperial moth. Adults are present from mid-June
to mid-July.

Family Sphingidae
Sphinx Moths

The adults of this family are distinctive in appearance and are known by such
common names as sphinx moths, hawk moths, and hummingbird moths (574,
1075). They have stout, spindle-shaped bodies and long, narrow, very strong
wings. The antennae are more or less thickened at the middle or toward the tip and
are usually pointed or curved back in the form of a hook. The mouth parts are
usually very long, and when not in use are held coiled beneath the head like a watch
spring. Mature larvae are long, usually naked, and each bears a horn, an eyelike
spot, or a low tubercle on top of the eighth abdominal segment. The pose of the
larva while at rest is distinctive—it clings to its support with its prolegs, holds the
front part of the body aloft, and bends its head downward.

Z1z

Sphinx moths are strong fliers. Some fly only at night, others at twilight or
during the day. They are usually seen hovering like hummingbirds over flowers and
feeding on nectar while in flight.

The elm sphinx, Ceratomia amyntor (Geyer), feeds on basswood, birch, and
elm throughout the Eastern United States. Full-grown larvae are pale green to
reddish brown; are marked on each side with seven oblique, whitish stripes; bear
pairs of horns on the tops of the second and third thoracic segments; and have one
caudal horn. They are about 75 mm long. C. undulosa (Walker), the waved sphinx,
feeds on ash and lilac from eastern Canada and Maine to central Florida and
westward to Texas and eastern Alberta. Full-grown larvae are about 62 mm long.
The head is bluish green marked laterally by a broad, pale band. The pea-green
body tapers toward the head and is marked by seven oblique, yellow stripes on each
side; the spiracles are orange; and the caudal horn is reddish and curved toward the
tip.

The catalpa sphinx, C. catalpae (Boisduval), occurs from New York to Florida
and westward to Michigan, Kansas, Iowa, and Texas, but appears to be most
abundant in the Southeastern States. It feeds exclusively on catalpa trees, often
completely defoliating them. The injury is of relatively minor significance to the
tree (1157). The adult is heavy-bodied and has a wingspread of about 75 mm. The
forewings and body are brownish with irregular dark and light bands and markings;
the hindwings are almost uniformly brownish gray. Full-grown larvae are about 75
mm long and armed with a stout, black horn near the posterior (fig. 88). There are
two highly variable color forms of large larvae—dark and light (6//). The dark
form is black on top and pale yellow underneath; the light form is pale yellow with
markings and patches of black on top.

Winter is spent as a pupa in the soil. Adults begin to appear as early as March in
the South, but much later farther north. Eggs are deposited in large masses on the
undersides of leaves or in smaller masses on twigs and branches. Young larvae feed

A
_ Zz Z j
Courtesy OS
& Dev. Cent., Dep. Entomol.

Figure 88.—Larvae of the catalpa sphinx, Ceratomia
catalpae.

213

gregariously; older ones, singly. In the Deep South, all life stages may be present
during the summer, and there may be three or four generations per year. In the
North there is only one generation per year. Larvae are often heavily parasitized by
the hymenopteran Cotesia congregata (Say) (fig. 89).

Courtesy OSU, Ohio Agric. Res.
& Dev. Cent., Dep. Entomol.

Figure 89.—Larvae of the catalpa sphinx
parasitized by Cotesia congregata.

The great ash sphinx, Sphinx chersis (Hubner), feeds on lilac and ash through-
out the United States. Full-grown larvae are usually light green with yellow-banded,
bluish heads and are about 75 mm long. There are seven light-yellowish stripes,
edged above with bluish green, on each side of the body, and the caudal horn is pale
blue and curved downward. The larvae are found most commonly on young trees
and sprout growth in the open or along roadsides. Populations are sometimes heavy
enough to cause noticeable defoliation in such areas.

Sphinx kalmiae J. E. Smith, the laurel sphinx, feeds on white ash, fringetree,
mountain-laurel, rhododendron, and lilac from southern Canada throughout the
Atlantic States to Georgia and west to Saskatchewan and Mississippi. Full-grown
larvae are yellowish green and about 75 mm long. The body is marked by seven
oblique, yellow stripes edged above with black; the caudal horn is arcuate and blue
with black, raised markings. S. /uscitiosa Clemens feeds on poplar and willow in
the Atlantic Coast States and west through the northern Plains. S. drupiferarum J.
E. Smith, a widely distributed species, feeds on apple, cherry, hackberry, plum,
and peach.

Eastern white, pitch, red, and jack pines are fed on by the larvae of Lapara
bombycoides Walker, the pine tree sphinx, in eastern Canada and from the Atlantic
Coast through the northern Plains. The larvae are present from July to September.
Full-grown specimens are green except for a broad, brick-red, median dorsal stripe;

214

a reddish ventral stripe; and three longitudinal white stripes on each side. They are
about 50 mm long. Brick-red patches sometimes enclose the spiracles, and there is
no caudal horn. L. coniferarum (J. E. Smith) feeds on pines, especially longleaf and
loblolly, in more southern areas.

Aspen, willow, and many other trees are fed on in the whole eastern area by
Paonias excaecatus (J. E. Smith), the blinded sphinx. Full-grown larvae are light
green, studded with pointed granulations, and about 62 mm long. There are seven
oblique, yellowish stripes running backward on each side of the body; the head is
conical, granulated, and has a white or pale-yellow stripe on each side, meeting at
the apex. The spiracles are deep lilac or black; the caudal horn is usually green and
nearly straight; and the thoracic legs are lilac or reddish. Larvae of P. myops (J. E.
Smith), the smalleyed sphinx, feed on various kinds of cherry in the Eastern United
States. They have rose-colored spiracles; otherwise, they closely resemble the
larvae of P. excaecatus. The larvae of Smerinthus jamaicensis (Drury), the twin-
spot sphinx, feed mostly on poplar and willow, but also on apple, birch, elm, and
plum. They also resemble the larvae of P. excaecatus. They differ mainly in having
a bluish-purple caudal horn and violet thoracic legs. Subdorsal rows of reddish
spots may occur on each side of the body and around the spiracles in any of these
three species.

The walnut sphinx, Cressonia juglandis (J. E. Smith), occurs from eastern
Canada to Florida and westward to the eastern boundary of the Great Plains. The
larvae feed on black walnut, butternut, the hickories, beech, and pecan. Full-grown
larvae are light green to reddish, coarsely granulated with white, and about 50 mm
long. The head bears a yellowish stripe on each side and two rough, brownish
projections on the apex. There are seven light-yellowish oblique stripes, sometimes
reddish bordered above, on each side of the body. The caudal horn is brownish and

very granulated. There are two generations per year in the South.
Family Notodontidae
Notodontid Moths

More than 100 species of notodontid moths occur in the United States and
Canada, and many of the larvae feed on the foliage of a wide variety of deciduous
trees and shrubs. The family name refers to the fact that in some species there are
backward-projecting tufts on the hind margin of the wings that protrude when the
wings are folded. The larvae may be solitary, but many are nest builders. When
disturbed, they often elevate each end of the body and remain attached by the four
pairs of prolegs at the middle of the body. Larvae in some species spray a mixture of
formic acid and other chemicals in response to attacks by predators (377). These
chemicals can burn human skin (656). Many feed exposed on the foliage; others
feed from within folded leaves or tents.

The poplar tentmaker, Clostera inclusa (Hiibner), occurs in southern Canada
and from New England to Georgia and Colorado, and feeds on various species of
poplar and willow. The adult is brownish-gray with three whitish lines crossing each
forewing, and it has a wingspread of about 25 mm. There is a crest of dark-brown
hairs on the front of the thorax, and the hindwing is crossed by a wavy band. Full-
grown larvae are brownish to nearly black and up to 42 mm in length. There are four
lines of light yellow on top, and one bright and several indistinct lines and yellow
marks are on the sides. Also, there are black tubercles on the tops of the first and
eighth abdominal segments (878).

Adults appear from March to April and from July to August, depending on
location. Eggs are laid in clusters on the undersides of leaves. The larvae are

215

gregarious and live in tents or webs that they construct by pulling together the edges
of one or more leaves and lining them with silk (fig. 90). They feed from May to
October, then craw] to the ground and pupate in loose cocoons to spend the winter.
Old, abandoned nests often remain on trees throughout the winter. There is one
generation and a partial second or two per year in the North, and up to five in the
South. This species often seriously defoliates small groups of trees, especially trees
growing more or less in the open.

F-519526

Figure 90.—Nest of the poplar tentmaker, Clostera
inclusa, torn open to show the larvae.

Other species of Clostera frequently encountered are C. albosigma (Fitch) on
aspen; and C. apicalis (Walker), C. brucei (Henry Edwards), and C. strigosa
(Grote) on aspen and willow. None is very important.

The yellownecked caterpillar, Datana ministra (Drury), occurs in southern
Canada and throughout most of the Eastern United States. Its food plants include
many species of fruit, shade, and forest trees. Important forest and shade tree hosts
include paper and yellow birches, basswood, elm, oak, maple, butternut, walnut,
mountain-ash, hophornbeam, and honeylocust. The adult has a wingspread of about
50 mm; its forewings are cinnamon brown and marked with irregular dark lines.
Full-grown larvae are about 50 mm long and are moderately clothed with long, soft,
white hairs. The head is jet black; the prothorax, bright orange-yellow; and the
body, marked longitudinally with alternate black and yellow or whitish stripes.

Adults appear during June and July. Eggs are laid in masses of 100 or more on the
undersides of the leaves. The larvae feed in colonies (fig. 91) near the ends of twigs
and branches. When disturbed, they elevate both ends of the body. At maturity, they
drop to and enter the soil to depths of 5 to 10 cm where they pupate and spend the
winter. There is one generation per year (/222).

Damage is seldom serious in the forest, although heavily infested trees may be
completely defoliated. Fruit, shade, and ornamental trees are injured most severely.

216

Courtesy Ill. Nat. Hist. Surv.

Figure 91.—Cluster of larvae of the yellownecked
caterpillar, Datana ministra.

Two species of tachinid parasites, Compsilura concinnata Meigen and Winthemia
datanae Townsend, are important natural enemies.

Datana angusi Grote & Robinson feeds on hickory, oak, beech, gray birch, and
butternut and occurs throughout the Eastern United States, west to Illinois, and
along the north shore of Lake Erie in Ontario, Canada. Full-grown larvae resemble
those of D. ministra, but differ in having an entirely black cervical shield (644).

Datana perspicua Grote & Robinson, the sumac datana, feeds on sumac
throughout most of the Eastern United States and in southern Canada. Full-grown
larvae are moderately hairy and about 50 mm long. The head 1s dark reddish to
black; the cervical shield, reddish brown; and the anal plate, blackish. The body 1s
deep straw or lemon yellow, with 11 longitudinal, dark reddish-brown to blackish
stripes.

Datana contracta Walker feeds on oak and sycamore in the Eastern United States
west to the Lake States and Arkansas. Full-grown larvae are about 50 mm long and
clothed with long, white hairs. The body is black with 11 longitudinal, yellowish-
white stripes.

Datana drexelii Henry Edwards feeds on basswood, walnut, sassafras, and
witch-hazel from the Atlantic Coast to Ohio. Full-grown larvae are moderately
hairy and about 50 mm long. The head and body are black; the cervical shield and
front of the thorax, honey-yellow; and the body bears 11 longitudinal stripes.

The walnut caterpillar, D. integerrima Grote & Robinson, occurs commonly in
southern Ontario and throughout the Eastern United States where it feeds on a wide
variety of deciduous trees, preferably walnut, butternut, pecan, and hickory. Adults
are stout-bodied, have wingspreads of about 50 mm, and are clothed with dull-
brown to chestnut-brown scales. The forewings are brownish and crossed by dark,
irregular lines. Full-grown larvae may attain 50 mm in length. The body is black
with longitudinal, yellowish stripes and is covered with long, white or dirty-gray
hairs.

PAG

Adults are present during the spring and summer. Egg laying begins in early
June, and eggs are deposited in masses on the undersides of leaves. The larvae feed
in colonies until almost full grown (fig. 92). They are often found in masses on the
trunk and larger limbs where they congregate to molt. Later, they return to the
foliage to continue their feeding. Full-grown larvae drop to the ground and wander
about searching for pupation sites. At this time they are often found in large
numbers along the foundation walls of houses. Pupation occurs in the soil and there
are one or two generations per year, depending on locality (532).

Courtesy Ill. Nat. Hist. Surv.

Figure 92.—Colony of larvae of the walnut caterpillar,
Datana integerrima.

The walnut caterpillar is frequently a serious pest of walnut. Trees heavily
defoliated 2 or more years in succession are seriously injured or killed. Isolated
trees or trees growing in small groups are especially subject to heavy attack. Losses
have been particularly severe in the Central States (39/).

Datana major Grote & Robinson feeds on azalea and apple and various shrubs
from the East Coast to Illinois. Full-grown larvae have mahogany-red heads,
cervical shields, and legs. Their bodies are marked with longitudinal, yellow lines
broken with black, giving them a finely spotted appearance.

Hyperaeschra stragula (Grote) has been recorded from southern Canada and the
Northern States, where it feeds on willow and quaking aspen. Full-grown larvae are
about 37 mm long. The head 1s flattened in front and slightly bilobed. The body is
mostly pearly gray with a reddish-brown dorsal line between the head and second
tubercle. The second and third abdominal segments each bears a conical tubercle
directed backward, and there is a prominent, pale-rust hump on the eighth abdomi-
nal segment. Larvae are found from June to October, and the winter is spent as
pupae in the ground. There are one and sometimes two generations per year.

Pheosia rimosa Packard, the false-sphinx, occurs rarely in eastern Canada and
from coast to coast in the Northern United States. Its hosts are recorded as poplars,
especially quaking aspen, and willow. Full-grown larvae are lead-colored with a

218

purplish tinge and about 43 mm long. The body segments are slightly smaller at the
middle, and the eighth segment bears a well-developed horn. Larvae are present
from July to October, depending on locality, and the winter is spent as a pupa in the
ground. There may be one or two generations per year.

Lophodonta angulosa (J. E. Smith) occurs from southeastern Canada to Florida
and Texas and feeds on various species of oaks. Full-grown larvae are pea green and
about 37 mm long. The body is marked with a faint, double, whitish line down the
middle of the back, and a distinct reddish stripe down each side. Larvae may be
found from May to October, and winter is spent in silken cocoons on the ground.
There are two generations per year in the South, but only one in the North. L.
ferruginea Packard occurs on paper birch in the Northeastern States and south-
eastern Canada.

Nadata gibbosa (J. E. Smith), the green oak caterpillar, occurs from coast to
coast in southern Canada and throughout the United States. The larvae feed on the
foliage of a wide variety of deciduous trees such as various species of oaks, red and
sugar maples, beech, paper birch, and willow. Full-grown larvae are pale pea green,
have large, rounded heads and tapering bodies, and are about 43 mm long. The
spiracles are deep red, and there is a yellowish stripe along each side. Larvae may
be found from May to October, and winter is spent as a pupa beneath the litter, but
not in the ground. There may be two generations per year as far north as New
England.

Nerice bidentata Walker feeds on elm in southern Ontario and from New
England to the Lake States and Kansas. Full-grown larvae are bluish green and
about 30 mm long. There are four white bands on the front and sides of the head,
and there is a large forward-pointing tubercle on each of the first eight abdominal
segments, and a pair of small ones on the ninth. Larvae are present from June to
September, and the winter is spent in silken cocoons on the ground. There is one or
one and a partial second generation per year.

Symmerista canicosta Franclemont, the redhumped oakworm, occurs in south-
eastern Canada and throughout much of the Northeastern United States. Its hosts
are various oaks, preferably white and bur, and several other deciduous trees such
as basswood, sugar maple, paper birch, beech, and elm. Outbreaks covering
several thousand hectares of oak type have occurred in Michigan and New England
(1S). The adult is ash gray, with a long, white area near the outer two-thirds of the
costal margin of each forewing, and it has a wingspread of 37 to 50 mm. The full-
grown larva has a rounded, orange-red head and a yellowish body that increases in
width back to an orange-red enlargement on the eighth abdominal segment. The
body is also marked with five fine, black dorsal lines (fig. 93).

Adults appear from May to July and the female deposits her eggs in masses on
the undersides of leaves. The larvae feed gregariously at first and skeletonize the
foliage. Later, they scatter out and feed singly, devouring entire leaves except the
larger veins. Mature larvae move to the ground and spin cocoons in rolled leaves
where they pupate and spend the winter. There appears to be one generation per
year.

Populations are usually too light to cause serious injury. Sometimes, though, they
are heavy enough to cause severe defoliation in isolated spots.

Symmerista albifrons (J. E. Smith) occurs in the Eastern United States. Its hosts
are much the same as those of the redhumped oakworm, a species with which it is
easily confused both in appearance and habits. At least 5,000 hectares of northern
hardwoods were defoliated in Upper Michigan and Wisconsin during an outbreak in

219

F-519531

Figure 93.—Larvae of Symmerista canicosta, the
redhumped oakworm.

1961. There are two generations per year. The orangehumped mapleworm, S.
leucitys Franclemont, occurs in the Northern States and southern Canada where it
feeds principally on sugar maple. Its life history and habits are similar to those of
the redhumped oakworm. It is distinguished from the latter by an orange headcap-
sule and hump and three fine, black dorsal stripes (/0).

The variable oakleaf caterpillar, Heterocampa manteo (Doubleday), occurs in
nearly all of the States and Canadian provinces east of a line drawn from western .
Ontario through eastern Texas. Its hosts include a wide variety of deciduous trees.
All species of oaks are attacked, but white oak is preferred. Other important species
attacked include beech, basswood, paper birch, American elm, walnut, boxelder,
persimmon, and apple (/3/8). The adult is ashy gray and has a wingspread of 37 to
42 mm (fig. 94A). The full-grown larva is about 38 mm long with variable
coloration. The head is large with two broad, lateral bands, the inner brown or
black and the outer creamy white. The body is smooth, yellowish green with a pale,
middorsal longitudinal line, and with more or less reddish-brown coloration on each
side bordered laterally by a creamy-yellow stripe, below which is a yellowish,
stigmatal stripe (fig. 94B).

Pupation occurs in early spring, and the adults appear from early May in the
South to late May or early June in the North. Eggs are deposited singly on the
leaves, each female laying up to 500. Young larvae skeletonize the lower surfaces of
the leaves and older ones eat entire leaves, except the larger veins. Mature larvae
move to the ground and spin cocoons in the litter or top soil. The larvae spray a
chemical mixture containing formic acid, when disturbed. The mixture is capable
of blistering the skin of humans (656). Winter is spent in the prepupal stage. There
are two generations per year in the South and one in the North.

Trees of all sizes are attacked, and heavy defoliation may occur anywhere in the
insect’s range, especially in the South. Some outbreaks have been extensive,
covering thousands of square kilometers and extending for hundreds of kilometers
(1179). Tree mortality has usually not been serious, however, since most trees of
sapling size or larger are able to withstand several years of extensive defoliation.

The saddled prominent, H. guttivitta (Walker), occurs in southeastern Canada
and throughout the Eastern United States. Beech, yellow birch, and sugar maple are

220

its preferred hosts, but many other species of deciduous trees are fed upon in
heavily infested stands. The adult is greenish gray or brownish gray with splotches
of creamy white, and has a wingspread of about 50 mm. Full-grown larvae are
usually gréen with reddish to purple or brown markings on the back, and they are
about 30 mm long (fig. 95).

Adults emerge in late May or early July and the female deposits from 200 to 300
eggs singly on the lower surfaces of the leaves. Young larvae skeletonize the upper
surface of leaves; older ones eat the entire leaf except the principal veins. They
often migrate from tree to tree and, where abundant, may collect in large numbers
about the bases of defoliated trees. Pupation occurs in the leaf mold from mid-July
until late August. Winter is spent in the pupal stage. There is one generation per
year in the northern parts of the insect’s range.

Many outbreaks have been recorded since the turn of the century (222, 96/)
during which heavy defoliation occurred over large forested areas. In areas suffer-
ing two consecutive years of defoliation, considerable tree mortality occurred.
Many of the trees that survived lost some of their large branches as well as large
portions of their tops. The most damaging effects of defoliation of sugar maple
occur to sapling and pole-size trees on poor sites (504, 707).

Figure 94.—The variable oakleaf caterpillar,
Heterocampa manteo: A, adults; B, larvae.

F-519533

Figure 95.—Dorsal and lateral views of larvae of the
saddled prominent, Heterocampa guttivitta.

22\

Several species of parasites have been reported from H. guttivitta (8), but the
most important are the egg parasite Jelenomus coelodasidis Ashmead and the pupal
parasite Cratichneuman sublatus (Cresson).

Other species of Heterocampa likely to be encountered in eastern forests include:
H. umbrata Walker—on oak and maple: H. biundata Walker—on beech, paper
birch, black cherry, and maple; and H. bilineata (Packard), the elm prominent—
on elm and birch.

Macrarocampa marthesia (Cramer) occurs from Maine to Florida and Texas and
feeds on the leaves of beech, maple, quaking aspen, oak, and sycamore. Full-grown
larvae are pale green and up to 50 mm long. The head is flat in front: there is a
small, double, reddish tubercle on the prothorax; and the body is marked with a
longitudinal, yellowish-white stripe and occasional pink spots on the back. Larvae
feed from July to October, then spin cocoons between leaves on the ground in which
they pupate and spend the winter.

Oligocentria lignicolor (Walker) feeds on various species of oaks and beech
throughout the Eastern United States. The larva is about 37 mm long and resembles
larvae of the genus Schizura. Its prominent characteristics include a pale, bilobed
head, with dark branched bands on each side of the face meeting on the vertex.
There is also a large, slightly cleft tubercle on the first abdominal segment and a
smaller brownish one on the eighth abdominal segment. The sides of the thorax are
pea green. Adults appear in July and August in the Northeastern States; the larvae,
from August to October. Winter is spent in a tough, parchmentlike cocoon on the
ground.

The redhumped caterpillar, Schizura concinna (J. E. Smith), occurs throughout
the United States and in most of the Canadian provinces. Its hosts include fruit trees
and a long list of forest and shade trees, such as elm, quaking aspen, willow,
hickory, black locust, dogwood, sweetgum, persimmon, and paper birch. The adult
is grayish brown and has a wingspread of about 50 mm. Full-grown larvae are about
25 mm long. The head and a hump on the eighth abdominal segment are red; the
body is marked with black and yellowish lines and bears a double row of short,
stout, black lines on top (fig. 96). When at rest, the larva holds the rear end in an
elevated position and, when handled, it gives off a pungent, disagreeable odor.

Courtesy Conn. Agric. Exp. Stn.
Figure 96.—Larva of the redhumped caterpillar, Schizura

concinna.

222

Adults appear from late May to August, depending on locality. Eggs are laid in
masses containing up to 100 each on the undersides of leaves. The larvae are
gregarious. At first, they skeletonize the undersurfaces of the leaves. Later, they
devour entire leaves except the midrib. During their feeding, they completely
defoliate one branch before moving on to another. Mature larvae move to the
ground and construct parchmentlike cocoons in the duff in which to spend the
winter. There is one generation per year.

Populations are often heavy in unsprayed appie orchards, along roadsides and
fence rows, and on ornamentals. In light infestations or on ornamentals, it is often
practicable to collect and destroy the colonies as soon as they are discovered.

Three other eastern species of Schizura are sometimes common locally: the
unicorn caterpillar, S. unicornis (J. E. Smith), S. leptinoides (Grote), and S.
ipomoeae Doubleday. Each species feeds on a wide variety of hosts, such as apple,
pin cherry, elm, quaking aspen, hickory, beech, paper birch, and willow.

Gluphisia septentrionalis Walker occurs in southeastern Canada and the northern
portions of the Eastern United States. It feeds on various poplars, especially
quaking aspen. Full-grown larvae are pale green and about 37 mm long. The head
has blackish stripes on each side; the body is largest in the middle, and is marked
with pinkish to reddish blotches on the back and a yellow line along each side.
Larvae are found from June to September, and winter is spent in cocoons on the
ground. As arule, there is one generation per year, but in some localities there may
be two.

The genus Cerura 1s represented in eastern forests by C. borealis (Boisduval), C.
occidentalis Lintner, C. cinerea Walker, and C. multiscripta canadensis McDun-
nough. The first feeds on quaking aspen, willow, and black cherry; the last three on
quaking aspen and willow. Notodonta simplaria Graef also feeds on quaking aspen
and willow. Misogada unicolor (Packard) occurs on sycamore, and Hyparpax

aurora (J. E. Smith) and H. perophoroides (Stecker) are found on oak.
Family Arctiidae
Tiger Moths and Allies

This is a large family of stout-bodied moths with moderately broad wings. In
general they are moderate in size and have broad heads and pectinate or ciliate
antennae. Many species are marked with brightly colored spots and stripes. All are
night fliers and are attracted to lights. They usually fold their wings rooflike upon
the abdomen while at rest. The larvae of most species are clothed with dense
clusters of hairs. In some species, certain of these clusters are larger and longer than
others, causing the larvae to resemble those of the tussock moths in the family
Lymantriidae. The hairs of certain species are irritating to humans. The majority of
species prefer the foliage of low-growing plants, but a few feed on the foliage of
trees and shrubs.

The hickory tussock moth, Lophocampa caryae (Harris), occurs in southern
Canada and south in the Eastern States to North Carolina. The larvae feed on the
foliage of a wide variety of deciduous trees and shrubs including walnut, butternut,
hickory, birch, elm, black locust, basswood, and quaking aspen. Walnut, butternut,
and hickory appear to be preferred. Adults are light brown or buff, with numerous
silvery-white spots on the forewings, and they have wingspreads of about 50 mm.
Full-grown larvae are about 37 mm long. The body is clothed with short, spreading
tufts of grayish-white hairs. There is a row of black tufts on the first eight
abdominal segments and pairs of long, black, hair pencils on the first and seventh
abdominal segments (fig. 97).

228

CHEN \\.
Courtesy Can. For. Serv., Can. Dep. Environ.,
Sault Ste. Marie, Ont.

Figure 97.—Larvae and cocoon of the hickory tussock
moth, Lophocampa caryae.

Adults appear from late May to early July. The female deposits her eggs in
batches of 50 to 400 each in a single layer on the undersides of leaves. The larvae
feed gregariously until nearly mature. Winter is spent in the pupal stage in gray,
hairy cocoons under rubbish and stones on the ground. There is one generation per
year. This species is often abundant locally, but it seldom, if ever, causes serious
defoliation.

The spotted tussock moth, 1. maculata (Harris), occurs in the Northern States
and southern Canada. The larvae feed on the foliage of various deciduous trees such
as oak, poplar, birch, beech, black locust, boxelder, black cherry, maple, and
willow. The oaks, willow, and poplar are particularly favored. The adult is pale
yellow with long, somewhat pointed, brown-spotted forewings, and has a wing-
spread of 37 to 50 mm. Full-grown larvae are about 30 mm long, and dull black
above. They are thickly clothed with tufts of black and bright-yellow to whitish
hairs, and have a row of short tufts down the middle of the dorsum. The tufts on the
third thoracic and eighth abdominal segments are longest and bear an intermixture
of white, yellowish, and black hairs; those on the thorax overhang the head. The
larvae are solitary feeders except during outbreaks and are found from July to
October. Winter is spent as a pupa in a hairy cocoon, and there is one generation per
year. This species is occasionally abundant enough to damage shade trees.

The pale tussock moth, Halysidota tessellaris (J. E. Smith), occurs in southern
Canada and throughout the eastern part of the United States. The larvae feed on
practically all common deciduous trees and shrubs. The adult is pale yellow and has
a wingspread of about 50 mm. The forewings are translucent and crossed by five
broad, dark bands. Full-grown larvae are dingy and densely clothed with compact
tufts of light-yellow or dirty-white fine hairs. Hair pencils rise in pairs from the
second and third thoracic and eighth abdominal segments (fig. 98). Adults appear in
June and July and the larvae, which usually feed singly, are present from August to
October. The winter is spent as a pupa in a brownish, hairy cocoon, and there is one
generation per year. This species is frequently abundant in the forest and along
roadsides in the South Central States, but is usually of minor importance.

224

Courtesy Conn. Agric. Exp. Stn.

Figure 98.—Larva of the pale tussock
moth, Halysidota tessellaris.

The sycamore tussock moth, H. harrisii Walsh, feeds on sycamore and London
plane trees, and it probably occurs wherever sycamore grows in this country. Adults
are indistinguishable from those of the pale tussock moth, and the larvae of the two
species differ only in color. Larvae of this species have yellowish bodies clothed in
whitish to yellow hairs, and the long hair pencils are orange. Infestations are often
heavy on shade and ornamental sycamore in the Northeast.

The fall webworm, Hyphantria cunea (Drury), occurs throughout the United
States and southern Canada. Its hosts include more than 100 species of forest and
shade trees. The adult has a wingspread of 30 to 42 mm, and the bases of the front
legs are orange or bright yellow. In the southern part of its range, the moth is white,
usually with dark spots on the wings. In the North, particularly in eastern Canada,
it is nearly always pure white and was formerly referred to as H. textor Harris. Full-
grown larvae are usually pale yellowish or greenish, with a broad, dusky stripe
down the back and a yellowish stripe down each side. They are about 25 mm long.
Their bodies are covered with long, silky, gray hairs arising in tufts from orange-
yellow or black tubercles. Head color varies from red to black. The larvae of H.
textor are dark.

Adults appear mostly from May to July and deposit their eggs in hair-covered
masses of several hundred each, usually on the undersides of leaves. Newly
emerged larvae immediately begin to spin a silken web over the foliage on which
they feed; as they grow, they enlarge the web to enclose more and more foliage (fig.
99). On heavily infested trees several branches may be enclosed in webs. Small
trees are often enclosed entirely. The larvae are gregarious until the last instar.
During the early stages, they feed on the upper surface of the leaves; later they
devour entire leaves except the larger veins and midribs. As they approach maturity,
some larvae leave the web and feed individually. Pupation occurs in thin cocoons
usually spun in the duff or just beneath the surface of the soil. There are one to four
generations per year, depending on location.

The fall webworm is ordinarily of no great importance as a forest pest since it
usually attacks understory weed species of no economic value. Outbreaks may

225

Bast *

F-531249
Figure 99.—Defoliation and webbing caused
by the fall webworm, Hyphantria cunea.

occur, however, sometimes encompassing tracts of several square kilometers. The.
fall webworm is often a serious pest of shade trees and ornamentals. These trees
may be heavily or completely defoliated, and the presence of numerous, unsightly
webs is esthetically detracting. Persistent infestations on individual trees may cause
branch and top-kill. There is considerable literature on this insect in the United
States and Canada (/55, SS/, 853, 854, 946, 947, 1207).

Seirarctia echo (J. E. Smith) larvae feed on the foliage of persimmon, runner
oak, and cabbage palmetto from Florida to Mississippi. The adult is white and has a
wingspread of about 55 mm. The wing veins are edged with dark brown or black.
Full-grown larvae are clothed with coarse, black-tipped hairs and are about 50 mm
long. The body is black on top except for a pair of yellowish stripes and a row of
orange warts that cross each segment.

Other species of arctiids likely to be encountered in eastern forests include:
Haploa clymene (Brown)—on maple, hickory, and apple; H. lecontei (Guérin-
Méneville)—on maple, birch, oak, and cherry; Apantesis radians (Walker)—on
slash pine seedlings in Georgia; and Lexis bicolor Grote—on balsam fir and

spruce.
Family Ctenuchidae
Ctenuchids

Lymire edwardsii (Grote) has been recorded feeding as larvae on the foliage of
banyan and fig trees in Florida. The adult is a sluggish, bluish-gray to purplish-gray
moth, with plumose blue antennae. The thorax 1s orange-red beneath, the abdomen
white beneath and blue above. Larvae are somewhat whitish with a dark tuft of hairs
on the thorax. When touched, they usually flip from the leaf and drop to the ground
on silken threads. This species is occasionally destructive on shade or ornamental
trees (469).

226

Family Lymantriidae
Tussock Moths

This family includes some of the insects that most seriously defoliate trees in the
United States. The female moths of certain species are wingless; others, though
winged, are so heavy-bodied that they are either unable to fly, or can fly for only
short distances. The remainder are strong fliers. The females of some species pack
and cover their eggs with their abdominal hairs; others coat their eggs with a viscid
secretion that hardens and forms a protective covering. The urticating hairs of
certain species in all stages may cause allergic reactions in people when they come
in contact with skin. The larvae of our native species also bear conspicuous tufts of
hairs on top of certain body segments.

The rusty tussock moth, Orgyia antiqua (L.), occurs throughout southern
Canada and in the northern part of the United States. Its hosts include scores of
species of both deciduous and coniferous trees (404). The male adult is rusty
colored. The forewings are crossed by two darker bands and each bears a conspic-
uous white spot near the border. Females are gray and wingless. Full-grown larvae
are about 28 mm long and have black heads and dark-gray bodies. The second
abdominal segment bears a black hair pencil on each side, and there are reddish-
orange tubercles bearing hairs. The tufts on the prothorax and abdomen are similar
to those on the larvae of the whitemarked tussock moth. The female deposits her
eggs in a single-layered, naked mass on the cocoon from which she emerges. There
are one and possibly two generations per year. This species is sometimes abundant
locally. Conspicuous defoliation has been recorded in Canada.

The whitemarked tussock moth, O. leucostigma (J. E. Smith), occurs com-
monly throughout the Eastern United States and eastern Canada and feeds on a
wide variety of deciduous and coniferous trees. Preferred species appear to include
the following: apple, basswood, elm, and poplar; Norway, silver, and planetree
maples; sycamore; paper and yellow birches; larch; and balsam fir. The female adult
is wingless, grayish to light brown, hairy, and about 12 mm long. Males are ashy
gray, and have fully developed wings with a wingspread of about 30 mm. The
forewing has a conspicuous white spot near the anal angle, and is marked with dark
wavy bands. Full-grown larvae (fig. 100) have coral-red heads and thoracic shields,
yellow to cream-colored bodies, and are about 25 to 37 mm long. There is a pair of
upright pencils of black hairs on the prothorax and another black tuft on the eighth
abdominal segment. There are also brushlike tufts of white or yellowish hairs on
each of the first four abdominal segments and reddish dots on the sixth and seventh
segments. The sides of the body are clothed in white and blackish hairs radiating
from rows of small yellow tubercles.

Winter is spent in the egg stage and hatching occurs between April and June.
Young larvae feed on the surface of the leaves, skeletonizing them. Later, they chew
holes in other leaves and finally consume all but the larger veins. Young larvae
often spin down on silken threads and are sometimes transported considerable
distances by the wind. The larvae become full grown in 5 or 6 weeks, then, under
branches or in bark crevices, they spin grayish cocoons consisting of silk and hairs
from their bodies. The pupal stage lasts about 2 weeks. Usually there are two
generations per year.

The whitemarked tussock moth may be a pest of shade trees in cities and towns.
It also occurs in forested areas but usually causes minor damage there. The life
history, habits, and parasites of the species are discussed (404, 609).

DoT

Courtesy Conn. Agric. Exp. Stn.

Figure 100.—Larvae of the whitemarked tussock moth,
Orgyia leucostigma.

Oregyia definita Packard, the definite-marked tussock moth, occurs in southern
Ontario and the Eastern States and feeds on quite a wide variety of deciduous trees
such as willow, apple, pin cherry, elm, paper birch, red oak, red maple, and ash
(404). The adults and larvae closely resemble those of the whitemarked tussock
moth in form and size, and in the arrangement of the tufts of hairs on the larvae.
Wingless females are clothed in golden-brown hairs. They lay their eggs in masses
on the cocoons from which they emerge, covering them with hairs from their.
bodies. In addition to the conspicuous pencils and tufts of hair, the larva is yellow,
with a faint dorsal stripe and a black spot behind each of the second and third tufts
of hair on the abdomen. This species is seldom of economic importance.

The genus Dasychira is represented in eastern forests by a number of species, a
few of which are economically important. Adults of the different species are very
similar in appearance and are difficult to distinguish. In both sexes they are winged,
and the females are heavy-bodied. The larvae have tufts of hairs characteristic of the
tussock moth group and their bodies are densely clothed with hairs. In some species
there is a feathery black hair in each lateral tuft.

Dasychira basiflava (Packard), the dark tussock moth, is sometimes locally
common in the Eastern States, where it feeds on various deciduous trees such as
slippery elm, white oak, beech, flowering dogwood, and hickory. The head of the
larva is hidden by yellowish clusters of barbed spines, and there are tufts of hairs on
the first and fourth abdominal segments. Heavy infestations on valuable shade and
ornamental trees may cause serious damage. It has several natural enemies includ-
ing parasites and pathogens (655).

The pine tussock moth, D. pinicola (Dyar), occurs in the Northeastern States
west to the Lake States and in southeastern Canada. The larvae feed on various
conifers such as jack, red, and eastern white pines, spruce, and balsam fir. Jack pine
is especially favored. The moth is gray-brown with lighter and darker stripes across
the forewings, and has a wingspread of 25 to 37 mm. Full-grown larvae are gray-
brown and about 37 mm long. There are four tufts of grayish or brownish hairs on
the dorsum. The first tuft has two black hair pencils on the front and three similar
ones on the rear.

228

Adults are present from early July to early August in the Lake States. Eggs are
usually deposited in small, irregular clusters on or near the female pupal case,
mostly on the needles near midcrown but also on the trunk, dead twigs, and similar
vegetation. Young larvae feed on the flat surfaces of needles. During August, the
second or third instars spin a few silken threads about themselves and go into
hibernation. They may be found beneath rough bark on large trees or between the
bases of needles on young, smooth-bark trees. Feeding is resumed in the spring on
staminate flowers and young needles. Later, old needles are also attacked, with
everything being consumed down to the needle sheath. In heavy infestations, the
entire tree may be completely defoliated. Full-grown larvae spin silken cocoons on
twigs or among needles. There is one generation per year (/237).

Several extensive outbreaks have occurred in the Lake States since the turn of the
century, and losses, especially of jack pine, have been serious. During 1961-62,
outbreaks covering several hundred square kilometers of jack and red pines and
white spruce were recorded in the region.

Other eastern species of Dasychira and their hosts are as follows: D. plagiata
(Walker)—on conifers at high elevations in the Appalachians, northern Mas-
sachusetts into Canada and west to the Rocky Mountains (//43); D. obliquata
(Grote & Robinson) and D. vagans (Barnes & McDunnough)—a willow and poplar
defoliator; D. meridionalis (Barnes & McDunnough)—on oak in the Southeast; D.
cinnamomea (Grote & Robinson)—on elm from New England to the Lake States;
D. tephra Hubner—on oak in the Southeastern States; D. dorsipennata (Barnes &
McDunnough)—on oak in Maine and southern Canada.

The gypsy moth, Lymantria dispar (L.), was introduced into the United States in
1869 when a French scientist brought a number of egg clusters from France for the
purpose of crossing the species with the silkworm. During the course of his work,
larvae escaped, and the species became established. About 20 years later, cater-
pillars had spread over an area of about 900 square kilometers around Boston, and
shade and fruit trees were being completely defoliated. The gypsy moth has
continued to spread and now occurs throughout the New England States, except
northern New Hampshire and Maine, and northeastern Vermont. It also occurs in
New Jersey, New York, Pennsylvania, Delaware, Maryland, Michigan, Virginia,
and southern parts of Ontario and Quebec. Relatively small, isolated infestations
have occurred in Arkansas, California, Illinois, Nebraska, North Carolina, Oregon,
and Wisconsin. Adult males have been trapped in numerous other States. [ts hosts
include most species of hardwoods, the oaks, apple, basswood, willows, birches
(except yellow and sweet), and poplars being most highly favored. Several conifers
are also attacked, usually when growing in mixture with the more highly favored
hardwoods (42). A list of food plants, divided into groups by host preference, has
been published (593).

The general biology of the gypsy moth has been reported by several authors (/6,
727, 928). The male of the gypsy moth (fig. 101A) is dark brown, with blackish
bands across the forewings, and has a wingspread of about 37 mm. Females (fig.
101B) are almost white and have wingspreads of about 50 mm. The abdomen of the
female is clothed in yellowish hairs and is so large and heavy that she is unable to
fly. Full-grown larvae (fig. 101C) are about 37 to 60 mm long. The head has yellow
markings, the body is dusky or sooty-colored and hairy, and there is a double row of
five pairs of blue spots followed by a double row of six pairs of red spots on the
dorsum. The pupa (fig. 1O1E) is reddish brown with a sprinkling of reddish hairs.

229

D x.

F-489191-95

Figure 101.—Life stages of the gypsy moth, Lymantria
dispar: A, adult male moth; B, adult female moth; C,
mature larva; D, egg mass; E, female pupa with cast
larval skin.

Adults appear in late July and August, and mate. The males fly vigorously in a
zigzag manner, often within about | m of the ground. The females, being unable to
fly, crawl a short distance from the empty pupal case and mate. Immediately after
mating, they lay their eggs in oval masses (fig. 101D), about 100 to 1,000 eggs
each, then cover them with buff-colored hairs from their abdomens. While the
majority of the masses are deposited on the trunks and limbs of trees, many are also
laid in various other places such as under stones, inside hollow stumps and trees, on
leaves, and even on buildings. Some masses are deposited on vehicles, and this
accounts for the long-distance dispersion of the gypsy moth. The winter is spent in
the egg stage, and hatching occurs about the first of May, usually about the time oak
leaves unfold.

Young larvae crawl from the egg mass and move toward the tops of trees. During
this period they often spin down on silken threads, especially when disturbed, and
some may be transported by the wind (22/, 824). Newly hatched larvae feed first
on leaf bases, then on the leaf surfaces, chewing small holes in the leaves. Older
larvae feed almost entirely from the edge of the leaf. During this part of their lives,
they feed mostly at night and tend to congregate in sheltered places during the day
(436). Large larvae consume entire leaves except the larger veins and midribs. In
heavily infested stands the entire crop of foliage may be consumed before the larvae

230

reach maturity. When this happens, the larvae usually vacate the trees and often
migrate in search of food. When the food supply is depleted but not completely
consumed before the larvae mature, many succeed in pupating but they are smaller
than normal. This results in smaller than normal adults, and the number of eggs
deposited per female is greatly reduced. Larvae seek sheltered places in which to
pupate. Pupae may be found attached by silken threads to limbs and trunks of trees,
stones, picnic tables, forest debris, buildings, vehicles, and the like. In heavy
infestations, they are often found massed together in large numbers. The pupal
stage lasts about 10 days or 2 weeks.

The gypsy moth has long been considered one of the most important forest
insects in the United States. Since the turn of the century, enormous sums of money
have been expended by the Federal Government and the affected States to eradicate
it, to suppress it, or to prevent its further spread (8/0). Despite these efforts,
outbreaks have continued to occur in the older infested portions of New England
often at 8- to 10-year intervals (/9), and have expanded to newer infested areas to
the South and West (table 1).

Defoliated trees show reduced growth and are more susceptible to attack by
wood-boring insects and fungi than are undefoliated trees (249, 348, 608, 702,
1245, 1246). Mortality of defoliated trees has been variable (/9/, 660, 702, 1156).
Further losses result from the reduction in quality of forest sites in stands suffering
repeated defoliations. Esthetic and recreational values are also reduced in outbreak
areas.

When the gypsy moth was introduced, the insect parasites, predators, and
pathogens affecting it in its native habitats abroad were left behind, and it encoun-
tered none in the Northeast that were capable of holding it in check. To remedy this
situation, a program of importation of foreign parasites and predators was initiated
in 1905 and has been continued intermittently to the present (252, 328, 613, 6/8,
1014). Collections representing 45 different species of the parasites and predators
were imported, colonies of which were released in infested stands in this country.
Ten species of parasites and two predators became established (6/8, 1090). Two are
hymenopterous egg parasites—Ooencyrtus kuvanae (Howard) and Anastatus dis-
paris Ruschka; four are tachinid larval parasites—Compsilura concinnata
(Meigen), Exorista larvarum (L.), Parasetigena silvestris (Robineau-Desvoidy),
and Blepharipa pratensis (Meigen); two are hymenopterous larval parasites—
Phobocampe disparis (Viereck) and Cotesia melanoscelus (Ratzeburg); two are
hymenopterous pupal parasites—Monodontomerus aereus Walker and
Brachymeria intermedia Nees; and two are coleopterous predators—Calosoma
sycophanta (L.) and Carabus auratus L.

One of the most important factors affecting heavy gypsy moth populations is the
so-called wilt disease, caused by a nuclear-polyhedrosis virus. This pathogen 1s also
a native of Europe. It may have entered this country in gypsy moth material
imported for the recovery of parasites. During moth epidemics, it increases tremen-
dously and decimates populations over large areas (3/2, 731, 990). Other natural
control factors include low winter temperatures—exposed eggs begin dying at a
temperature near — 28° C (48/, 1178). Small mammals devour large larvae and
pupae found on the forest floor. Also, many larvae die of starvation in woodlands
entirely stripped of foliage before the larvae reach maturity. Discussions of the
population dynamics and ecology are available (/04, 187, 189, 726, 1106).

Silvicultural practices designed to promote the health and vigor of stands are
helpful in increasing tree resistance to gypsy moth damage (/05). A reduction in the

231

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232

proportion of favored host species also decreases the chance of infestation and
limits the degree of population buildup (4).

Insecticidal spraying by aircraft has been used widely in gypsy moth control.
Various chemical insecticides are effective. Attempts to secure control by spraying
infested stands with the nuclear-polyhedrosis virus continue with some success
(733, 989, 1033). Spraying with the microbial insecticide, Bacillus thuringiensis
Berliner (25, 343), has given foliage protection. Another possible method of
control is the use of synthetic sex attractants. The natural female attractant has been
isolated, identified, and synthesized (/06, 626); the synthetic pheromone is known
as disparlure. It has been particularly useful in survey and detection work (/072).
The use of the sterile-male technique for controlling gypsy moth continues to be
investigated (827, 1073).

The satin moth, Leucoma salicis (L.) (fig. 102), an introduced species first
discovered in North America near Boston, Mass., in 1920 and in British Columbia
during the same year, now occurs throughout most of New England and in the
Maritime Provinces of eastern North America. In the West, it has spread southward
through western Washington into northwestern Oregon. The larvae feed on most
species of poplar and willow. Adults of both sexes are pure white with a satiny sheen
and have a wingspread of 37 to 50 mm. The head, thorax, and abdomen are black
but are so densely clothed with long, satiny-white hairs they appear white. Full-
grown larvae are about 34 mm long. The head is black with a bluish tinge. The
body is blackish on top with a row of large white blotches down the middle and a
narrow broken line along each side. There 1s also a transverse row of reddish-brown
tubercles on the top of each body segment, each bearing a tuft of yellowish-brown
hairs.

Winter is spent in the larval stage. Feeding is resumed in April in New England.
Young larvae feed only on the leaf surface, most often on the underside. Partly
grown larvae eat small, irregular holes in the leaf, and full-grown larvae devour the
entire leaf except the large veins. Feeding is completed by June. Pupation occurs in
loosely woven cocoons of silk spun in the leaves or on twigs or other objects. Adults
appear in late June and early July. Eggs are deposited on leaves, branches, and
trunks of trees or on other surfaces in masses of 100 to 400. Each mass is oval,
about 9 mm long, and covered with a glistening white secretion. Hatching occurs in
about 2 weeks. Newly hatched larvae feed for 5 to 6 days and then spin small webs
in which they molt to the second instar. These larvae then feed again for 5 to 6 days.
Then they crawl to limbs or the trunk and spin hibernacula in bark crevices or under
loose bark where they remain until the following spring (/74).

The satin moth is not very important as a forest insect in the Eastern States,
although heavy infestations are occasionally reported. From time to time, it se-
riously defoliates poplars in ornamental plantings. A number of natural enemies
(dipterous and hymenopterous parasites, fungi, insect and vertebrate predators)
cause mortality of eggs, larvae, pupae, and adults (328, /236).

The browntail moth, Euproctis chrysorrhoea (L.) (fig. 103), an introduced
species, was first recorded in North America in Somerville, Mass., in 1897. During
the next few years, it increased enormously and spread rapidly. By 1905, it was
extremely abundant throughout Rhode Island, eastern Massachusetts, southern
New Hampshire, and southwestern Maine, and had been recorded as far north as
New Brunswick and Nova Scotia. Ten years later, most of the area east of the
Connecticut River, with the exception of northern New Hampshire and Maine, was
heavily infested. Infestations also occurred in Vermont and west of the Connecticut

239

A, B, and D courtesy of L. Nef, For.
Biol. Res. Cent., Genck, Belgium:
C courtesy of Can. For. Serv.,
Marit. For. Res. Cent.

Figure 102.—The satin moth, Leucoma salicis: A,
mating pair; B, hibernating webs on bark opened to
show overwintering second instars; C, mature larva on
aspen leaf; D, pupae (females are larger than males).

River in Massachusetts and Connecticut. Since then, the area and intensity of
infestations have declined greatly. At present. infestations occur almost entirely in
old, abandoned apple orchards and on beach plum on Cape Cod and in coastal
counties in Maine. During the first several years following its introduction, its hosts
were recorded as several species of deciduous trees, principally apple. pear, plum,
oak, willow, elm, and maple.

234

ware

A courtesy of Ziegler; B, C, and D courtesy o
Shrdder; Landwirtschaftskammer, Rheinland.

Figure 103.—The browntail moth, Euproctis
chrysorrhoea: A, female ovipositing; B, egg mass on
leaves; C, larvae; D, silken cocoons opened to show
pupae.

Browntail moth adults are pure white except for the tip of the abdomen, which is
covered with brown hairs. The female is rather heavy-bodied and has a wingspread
of about 37 mm. Males are more slender and somewhat smaller. Full-grown larvae
are about 37 mm long. The head is light brown; the body is dark brown to almost
black with a broken white line on either side and two conspicuous red spots near the
posterior end. There are also numerous tubercles with long, barbed hairs and many
short, brown hairs on the dorsum and sides. These urticating hairs are allergenic to
humans and cause a severe rash when they come in contact with skin.

Adults appear in early July, and the female deposits her eggs in elongate oval
masses from 12 to 18 mm long on the undersides of leaves. Each mass contains
about 300 eggs and is covered with brown hairs from the female’s abdomen. Young
larvae feed gregariously on the surface of the leaves. Later, they tie two or more
leaves together with silk near the tip of a branch. Then they spin a web over the
outside of these leaves and fasten them securely to the twig, thereby forming a
tough, grayish web from 5 to 15 cm long where they spend the winter. Feeding 1s
resumed in the spring, and the larvae become full grown by mid-June. Pupation
occurs in silken cocoons spun usually among the leaves at the tips of twigs.

The browntail moth, although one of the better known forest and shade tree
insects in the Eastern United States, is no longer of economic importance. For some
undetermined reason, it has almost disappeared. Introduced parasites, many of
which became established (328), and a fungus disease probably helped toward its
control, but it seems unlikely that they were primarily responsible for its decline.

235

Family Noctuidae
Owlet Moths and Underwings

This is the largest of all the families of Lepidoptera in North America. There are
more than 2,900 species in the Noctuidae in North America. The adults are mostly
nocturnal; many of the moths attracted to lights at night belong to this group. The
larvae are usually dull colored and naked. Many are foliage feeders, some are
borers, and others gnaw in fruits. Others are found on the foliage of forest and
shade trees and shrubs, but they usually do not cause serious damage. A number of
species attack seedlings in nurseries and young trees in plantations, and occasion-
ally cause serious injury. This family has received extensive treatment (268, 435,
684, 1002, 1032, 1108).

The genus Acronicta contains upwards of 100 species, many of which feed on the
foliage of forest and shade trees. Some are known as “‘dagger moths” because of the
presence of a daggerlike mark near the anal angle of the forewing.

The American dagger moth, A. americana (Harris), occurs throughout the
Eastern United States and from Newfoundland to Alberta, Canada. Its hosts include
a wide variety of hardwoods such as boxelder, basswood, red and sugar maples,
paper and yellow birches, elm, ash, oak, willow, hickory, and sycamore. Full-
grown larvae are about 50 mm long. They are clothed with fine, yellowish hairs and
there are long, black hair pencils on the backs of abdominal segments one, three,
and eight. Larvae are present from June to October, and overwinter in the pupal
stage in dense silken cocoons. During an outbreak in Maine, larvae were so
abundant that they literally swarmed all over buildings and gardens.

The cottonwood dagger moth, A. lepusculina Guenée, occurs from coast to
coast in southern Canada and the Northern States. Its favorite hosts appear to be |
willow and quaking aspen, but it also occasionally infests other species of poplar.
Full-grown larvae are clothed with long, soft, yellowish hairs and are about 37 mm
long. There are single, long, black hair pencils on the backs of abdominal segments
one, three, four, five, and eight. Larvae are present from July to October. Winter is
spent in a cocoon composed of silk and bits of wood.

Many other species of Acronicta are also found on trees in eastern forests. Some
of these with some of their more common tree hosts are as follows: A. innotata
Guenée—hickory and birch; A. morula Grote & Robinson, the elm dagger
moth—elm and basswood; A. interrupta Guenée—elm, cherry, birch, and apple;
A. lithospila Grote—hickory and oak; A. funeralis Grote & Robinson, the paddle
caterpillar—hickory, elm, and birch; A. modica Walker—oak; A. retardata
(Walker)—maple; A. leporina (L.), the poplar dagger moth—poplar, willow, and
birch; A. distans (Grote)—birch, cherry, poplar, willow, apple, and alder; and the
smeared dagger moth, A. oblinita (J. E. Smith)—usually herbaceous vegetation
but also occasionally poplar, willow, boxelder, pin cherry, alder, and apple.

The genus Catocala contains many large, conspicuous species, the larvae of
which feed on the foliage of various forest trees and shrubs. As a group, they are of
little economic importance. The forewings are usually grayish and mottled with
lighter and darker spots. This makes them very inconspicuous on the trunks of trees
when they are resting with their wings folded. The hindwings, in contrast, are
beautifully marked with bright red, yellow, white, or brown. This has given rise to
the common name “underwing moths.” More than 50 species have been recorded
from New York State alone, mostly on oak and hickory. The life histories of many
species have been discussed (56, 1/055).

236

Many other species of noctuids are commonly known as “‘cutworms.” The larvae
are usually stout, naked, and dull grayish or brownish. They are seldom seen,
however, because of their habit of feeding at night and hiding on the ground or under
bark during the daytime. A number of species feed on the roots, or on portions of
stems or foliage near the ground; many others climb the stems or trunks of their
hosts and feed on buds, flowers, fruit, and other succulent parts.

The more important species of cutworms in nurseries and forests in eastern
America and some of their hosts are as follows: Feralia jocosa (Guenée), the
joker—hemlock, spruce, larch; /dia aemula (Hubner), the spruce epizeuxis—
spruce (particularly in ornamental plantings; the larvae commonly are found in
webbed masses of dried needles and frass on the branches); Panthea furcilla
(Packard), the tufted white pine caterpillar—pines, P. acronyctoides (Walker),
the tufted spruce caterpillar—spruce; Papaipema furcata (Smith)—ash seedlings
in nurseries in the Lake States; Colocasia propinquilinea (Grote)—birch, beech,
maple, walnut; C. flavicornis (Smith)—hickory; Charadra deridens (Guenée)—
oak, maple, elm, birch; Raphia frater Grote, the yellow-marked caterpillar—
poplar, willow; Euxoa scandens (Riley), the white cutworm—young white and
overcup oaks (serious defoliation reported in Minnesota); the zebra caterpillar,
Melanchra picta (Harris)—usually most abundant on herbaceous vegetation, but
occasionally damages pine seedlings in nurseries in the northern Great Plains;
Orthosia hibisci (Guenée)—the opening buds and fruit of many deciduous trees
and shrubs and a few conifers; Lithophane laticinerea Grote—many species of
deciduous trees (severe defoliation of red maples on islands in Penobscot River,
Maine); the green fruitworm, L. antennata (Walker)—ash, boxelder (has occured
in larger numbers locally in New York and Vermont); Anomogyna elimata
(Guenée), the chameleon caterpillar—various conifers, especially balsam fir and
jack pine (one of the most common of the cutworm larvae on conifers in the
Northeast); and Palthis angulalis (Hubner), the spruce harlequin—a wide variety
of conifers, especially balsam fir and white spruce, also occasionally on various
deciduous trees.

Clean cultivation is recommended for preventing the development of damaging
populations of cutworms in forest nurseries.

Order Coleoptera—Beetles

The order Coleoptera contains more named insects than any other; in fact, over a
fourth of the named animals are beetles. The estimated number of named beetle
species is about 350,000 (27), and about | ,000 new species are named each year.
Over 26,000 species occur in the United States (980). Beetles can be distinguished
from all other insects, except earwigs, by their hard and horny or leathery forewings
that meet in a straight line along the middle of the back. The forewings, or elytra,
cover the membranous hindwings used for flying; some species are flightless and
lack hindwings.

All beetles have chewing mouth parts and complete metamorphosis, 1.e., eggs
hatch to produce larvae that feed to maturity; then the larvae transform to inactive,
nonfeeding pupae which become adults that mate and lay eggs to repeat the life
cycle. The life cycle varies greatly in length, from 10 or more generations per year
to | generation in many years. For most beetles the larval stage is long and the egg,
pupal, and adult stages are of short duration.

oon

Beetle larvae and adults have a wide range of habits. The majority of the beetles
are terrestrial but some species, even entire families, are aquatic both as larvae and
as adults. Some feed on vegetable matter; others feed on animal matter. Many are
phytophagous, many are predacious, some are scavengers, some feed on mold and
fungi, and a few are parasitic. Beetles that feed on plants are of primary interest.

The order Coleoptera contains many of the most destructive forest insects. Adults
and larvae of the phytophagous species have certain modifications in form and
habits that enable them to feed on different portions of plants. A beetle species may
cause damage to tree flowers. fruits, and seeds; roots, stems, or foliage of living
trees; dead or dying trees: drying logs and lumber: or wood products many years
after wood is placed into use. Considering the invaluable resources provided by
forests—wood and wood products, clean air and water, recreation and wildlife as
well as esthetic wilderness values—humanity is inadequately appreciative of the
fact that parasites and predators help to limit the damage beetles might cause in
most situations.

Beetles annually cause enormous losses to valuable timber or shade trees; to
forest products by degradation, often through the introduction of blue-stain fungi or
other pathogens: to seedlings in nurseries, plantations, seed orchards, or natural
stands by growth loss or mortality; and to seed orchards by reducing harvest of
viable seed. Most of the destruction results from feeding by larvae. Charac-
teristically, initial attack by adults of many species is restricted to wood that is in a
certain condition—either living, recently dead or dying, or decaying. Larvae of the
Cerambycidae, Buprestidae, and some other families may continue to survive
within wood that is too dry for initial attack. Some other beetles, e.g., the Lyctidae
and some of the Anobiidae, are capable of repeatedly infesting seasoned wood in
storage or service.

Identification.— Adult beetles in many families feed as defoliators or predators,
but the most serious damage to forests and forest products results during the
extended feeding period of the larvae of most families. Frequently the adults are
short-lived or nocturnal and the feeding larvae are hidden, either beneath bark or
deeply within the wood, and not readily available for identification. Although
larvae are found more often than adults, specific or generic identification of larvae
is often a difficult task, even for a taxonomic specialist. The family, group, and
sometimes even the species can often be quickly and easily identified from careful
attention to the general appearance of larvae and to what is being damaged, 1.e.,
seeds, twigs, bark, wood of living, dying, or dead trees, logs, and wood products,
ec.

Beetle larvae from various families can be grouped by general appearance
according to their body form or shape. For a few families, larvae have a different
body form with each molt. Larvae characterized by the absence of legs on the
thorax are the weevil or curculionid type. They have thick crescent-shaped bodies
and a well-developed head (families Curculionidae, Scolytidae, and late-stage
Bruchidae). Larvae with thoracic legs are either very active with flattened bodies
(thysanuriform) or slow moving with thick caterpillarlike bodies (eruciform). The
thysanuriform type is further subdivided into the caraboid and triunguloid types.
The caraboid type larvae have strongly developed mandibles and are mostly
predacious (families Carabidae, Cicindelidae, Cleridae, Staphylinidae, Histeridae,
etc.). The first instars of the Meloidae and Rhipiphoridae are about the only
triunguloid examples.

238

Many wood-infesting beetle larvae are eruciform, but again there is a subdivi-
sion. Larvae of the scarabaeoid subtype are typical grubs with wrinkled, crescent-
shaped, and often hairy bodies (families Scarabaeidae, Lucanidae, Anobiidae,
Bostrichidae, and Lyctidae). Larvae of the cerambycoid subtype have straight
rather than crescent-shaped bodies and are more active than larvae of the
scarabaeoid subtype. The larvae of the Buprestidae have somewhat flattened
bodies, but Cerambycidae and Elateridae larvae typically have rounded bodies.

There are extensive references to the taxonomy of coleopteran adults (27, /28,
145, 635, 723, 724, 725) and larvae (1, 142, 243, 980). Keys to identification and
descriptions of habits of species occurring in eastern North America are available
(304). Extensive checklists and manuals of the major families of beetles are being
prepared by the North American Beetle Fauna Project of the Beetle Research
Institute of America (27).

The families described here are discussed in phylogenetic order from primitive to
advanced. Some families do not contain forest-destroying insects but are included

because they contain beetles frequently encountered in the forest.
Family Carabidae
Ground Beetles

The Carabidae, one of the largest families of beetles, contains hundreds of
species. Both as adults and as larvae, nearly all ground beetles are predacious on
other insects. Adults of most species are black or dull brown, but adults of a few
species are yellow and may be marked with black. Some species are brightly
colored, either metallic blue, green, or purple. Many have an orange prothorax or
orange markings on the elytra. Size also varies greatly among species, but the body
is flat to oval in cross section. Adults are unusually long lived—1 to 4 years—and
are active even in winter. Ground beetles occur, sometimes abundantly, in most
habitats, e.g., under stones, logs, forest litter, bark, or running about on the ground
or branches.

Larvae are elongate, fusiform, and usually active. The integument is darkly
sclerotized in the free-living species, but it is white in those species living in the
ground or bark. The larvae have five-jointed legs, frequently with spines and single
or paired movable claws on the fifth joint for grasping prey. Their mandibles are
sickle-shaped for the same purpose.

Many carabids are very beneficial, preying on many of the forest pests that cause
much damage. The prey include lepidopterous and hymenopterous defoliating
pests, such as the gypsy moth and sawflies, and the immature stages of bark
beetles. A number of species in the genus Calosoma are important predators of tree-
defoliating pests and other insects. Adults are large, about 20 to 30 mm long, and
some are brilliantly colored. Like many other carabids, these conspicuous predators
emit a disagreeable odor when handled or disturbed.

A species introduced from Europe in 1906, Calosoma sycophanta (L.), 18 now
widespread and well established over most of the area infested by gypsy moths and
even beyond. Adults have a bluish-black pronotum with golden-green elytra, and
they are from 24 to 30 mm long (fig. 104). Both the adults and larvae readily climb
trees and feed voraciously on gypsy moth larvae and pupae. Many other insects are
also preyed upon. Adults live up to 4 years, overwinter and lay eggs in the ground,
and emerge about June. After climbing trees to feed to maturity, larvae return to the
ground to pupate. The effectiveness of this predator has been investigated (/88).

Native species widely distributed in the Eastern States include C. frigidum Kirby
(fig. 105), C. willcoxi LeConte, C. scrutator (F.), and the fiery hunter, C. calidum
(F.) (fig. 106).

239

F-532841 F-531250

Figure 104.—Adult of Calosoma Figure 105.—Calosoma frigidum on a red
sycophanta. maple twig in the field.

8
=

Courtesy Conn. Agric. Exp. Stn.
Figure 106.—Larva of the fiery hunter, Calosoma
calidum, feeding on a gypsy moth larva.

Sa

Carabids frequently occurring in bark beetle galleries include Mioptachys and
Tachyta spp. Several species, Geopinus incrassatus (Dejean), Harpalus spp., and
Scarites subterraneus F., reportedly cause feeding injuries to tree seedlings: how-

ever, members of the latter genus are strictly predacious (304).
Family Cicindelidae
Tiger Beetles

Tiger beetles are not forest pests but their appearance and habits arouse interest
and curiosity. Both adults and larvae are predacious on other insects. Most species
encountered will probably be in the largest genus Cicindela. Unlike other tiger
beetles, most members are active in the daytime. Adults of Cicindela spp. are 9 to
18 mm long; they are mostly metallic green or bluish gray with yellowish to whitish
bands or spots on the elytra. They are strong flyers and most often are seen swiftly

240

running along sandy stream banks or along woodland trails on bright, sunny days
from early spring to fall. They will often stop, turn, and assume an aggressive
posture toward intruders. This habit and the way they pounce on their prey has led
to their common name. In striking contrast to the active adults, the larvae wait for
prey to fall into conical, vertical burrows in soil.

Family Cupedidae
Cupedids

Beetles in this family are not of economic importance. Both adults and larvae
occur in moist, badly decayed wood or under bark. Cupes concolor Westwood is a
common, widely distributed species, sometimes found in structures of pine or oak.
Adults have flattened bodies, 7 to 11 mm long. They are grayish brown and densely
covered with scales. The larvae have a peculiar elongate, fleshy appearance and are
25 mm long. Each fleshy five-jointed leg is armed with a movable claw, a
characteristic common to predacious forms. These are considered to be the most
primitive of all beetles.

Family Silphidae
Carrion Beetles

Carrion beetles are often seen on animal excrement or around the bodies of dead
animals in the forest. Many of the more commonly observed species are in the
largest genera, Si/pha and Nicrophorus. Adults of various species range from 10 to
35mm in length. Their black bodies are often ornamented with yellow or red on the
prothorax or elytra. Their bodies are soft and robust, usually elongated but some-
times circular. Several of the species of small size, like Agathidium oniscoides
Palisot de Beauvois, apparently act as scavengers under the bark of logs and dead
trees in galleries of bark beetle-infested hardwoods.

Family Histeridae
Hister Beetles

Hister beetles have widely varied habits and habitats. Adults as well as larvae are
mostly carnivorous. Many are predacious on other adult insects and their larvae.
Habitats include excrement, carrion, decaying fungi, fermenting sap, beneath bark,
in ant and termite colonies, or in nests of small mammals and birds. Adults are
generally small (2 to i0 mm), sluggish, hard-shelled, and often with shortened
elytra. Most can feign death, drawing in their appendages when disturbed. The
species with flat bodies live under bark, often of only certain tree species, appar-
ently because they are predacious on a pest of the particular tree, e.g., Hololepta
spp. under bark of poplar and yellow-poplar trees. The species with cylindrical
bodies are found in galleries of bark beetles and wood borers, e.g., Plegaderus spp.
prey on eggs and larvae.

Family Byrrhidae
Byrrhids

These beetles are oval, convex, and usually black, | to 10 mm long. The head is
deflexed, retracted, and usually concealed from above. The hard-bodied, slow-
moving adults retract their appendages when disturbed to form a compact ball and
thus are referred to as pill beetles. Adults and larvae are herbivorous.

Although several species of the genus Byrrhus are abundant and cause much
damage to tree seedlings, specimens are difficult to find because the adults are
nocturnal. Adults are 5 to 10 mm long, black, and densely covered with grayish
hairs. The distribution of this genus apparently 1s limited to Alaska, Canada, and
the contiguous Northern States.

241

Family Staphylinidae
Rove Beetles

This very large family has over 300 genera, several of which contain numerous
species. Rove beetles occur commonly in eastern forests on, in or near the forest
floor, or under bark of dead trees. Many species are conspicuous and easily
recognized, though the family is difficult to define. Adults are usually elongate,
slender beetles, small to medium in size, and can run with considerable speed. The
wing covers of nearly all species are decidedly shortened, exposing a long and
slender abdomen. When disturbed, many of the larger species turn the tip of their
abdomen up and over the body, as if threatening to sting, as they run away. Most
species are uniformly black or brown, but many have red or yellow body segments
interspersed among the somber colors. A large species often: encountered, Phi-
lonthus cyanipennis F., is iridescent blue, purple, or green. Rove beetle larvae are
highly variable, and several distinct types occur from soft, white larvae to heavily
sclerotized, black larvae.

Most members of this family of beetles are predacious on insects or other similar
soft-bodied small animals. A few species feed on fungi, a few on pollen, and some
members of the genus Aleochara are known to be parasitic on fly pupae. Staph-
ylinids are not economically important though many species occur in any given
forest type. Common and conspicuous predacious genera that occur in a variety of
forests are Staphylinus and Philonthus, which contain some of the largest members
of this family. Other general predators are species of Quedius, Tachyporus, Medon,
Lathrobium, and Astenus.

Members of at least 20 different genera in the eastern regions are known to be
predacious on bark beetle eggs and first instars, and small wood borer larvae. |
Among those, the species of Nudobius, Quedius, Phloeopora, Atrecus,
Olisthaerus, Charhyphus, Placusa, and Phloeonomus are common in eastern
forests. Other genera contain species that can be predacious on insects under bark,
such as Tachinus, Sipalia, Leptacinus, and Homaeotarsus. A few genera often
occur in bark beetle galleries, but apparently do not feed on bark beetle eggs or
larvae. Of these, members of the genus Afhefa are common, and sometimes also
Siagonium.

None of the staphylinid species is known to be a major factor in the control of any
harmful forest pest. However, they are a consistent and abundant component of the

complex forest ecosystems.
Family Passalidae
Passalid Beetles

These beetles are discussed because of their interesting appearance and biological
habits. Most species occur in the tropics. Of the three species recorded from the
United States, only the horned passalus, Odontotaenius disjunctus (Illiger), has a
very wide distribution in the Eastern United States. Adults are large, somewhat
flattened, parallel-sided, shiny black beetles from 30 to 40 mm long. The thorax
and abdomen are separated with a narrowed “waist.” The elytra are deeply grooved
and a short, forward curved hook occurs on the top of the head. The three rigid,
terminal antennal segments are widened into a loose club that cannot be rolled
together.

Larvae and adults live together in colonies in damp, decayed logs and stumps.
The cream or dull-white larvae are fed wood that has been reduced to a pulp and
treated with digestive secretions by their parents (499). Both adults and larvae have
stridulatory organs. When disturbed, or for communication, adults rub together

242

roughened areas located on the underside of the wings and on top of the abdomen;
on the larvae, these roughened areas are on the third pair of legs that are very greatly
reduced.

Family Trogositidae

Trogositid Beetles

The family Trogositidae is made up of beetles formerly in the family Ostomidae.
Species in several genera are important predators of some of the most destructive
forest insects. These beetles are closely related to clerid beetles. Larvae of the two
groups are very similar in appearance; however, most of the adults are different.
Unlike the hairy bodies and contrasting colors of clerids, trogositid adults rarely
have hairs and are entirely brown, blue, green, or black. Their bodies are usually
flattened and elongate or oval. The terminal 3 segments of the 11-segmented
antennae form a loose club.

Temnochila virescens (F.) is one of the most important insect predators of the
southern pine beetle. Adults are iridescent, bluish green, and 10 to 18 mm long.
They feed on bark beetle adults on the surface of infested wood, though not as
actively as do the clerids. Larvae are elongate and cylindrical with a sclerotized
plate bearing a pair of unbranched recurved hooks. They burrow through bark
beetle galleries as they prey on eggs and larvae of the host.

Other predacious trogositids occurring in eastern forests and their insect hosts
include: Tenebroides corticalis (Melsheimer)—bark beetles; 7. bimaculatus
(Melsheimer)—Agrilus larvae in oaks; Corticotomus cylindricus (LeConte) and

Airora cylindrica (Serville)—ambrosia beetles and cossonid weevils.
Family Meloidae
Blister Beetles

Blister beetle adults are most commonly seen defoliating forest vegetation.
Distinctive characteristics are: green, gray, brown, or black color; usually 8 to 18
mm long; slender, soft-textured, and usually elongated body and elytra; sub-
cylindrical; head and elytra wider than pronotum; head narrowed abruptly into a
neck; often large and swollen abdomen; long legs; and cleft tarsal claws each with a
lower appendage. The larvae of the genus Epicauta prey on grasshopper eggs, all
others being predators in nests of wild bees.

Species that defoliate seedlings in nurseries and ornamental plantings in the
Midwest include: The threestriped blister beetle, Epicauta lemniscata (F.); the
margined blister beetle, E. pestifera Werner; the ashgray blister beetle, £.
fabricii (LeConte); Lytta aenea (Say); and Say blister beetle, L. sayi (LeConte).
The caragana blister beetle, £. subg/abra (Fall), the Nuttall blister beetle, L. nut-
talli Say, and the ashgray blister beetle defoliate caragana in the Great Plains; E.
torsa (LeConte) defoliates Kentucky coffeetree, honeylocust, and mimosa in Flor-
ida and Oklahoma. L. polita Say feeds on male strobili of shortleaf and pond pines
in the South,consuming the pollen sacs of mature male flowers, leaving only the
flower stalks or axes. The biology and control of seven common species are
discussed (602) and plant host records of Meloidae associated with bees have been
compiled (385).

Family Nitidulidae

Sap Beetles

Sap beetles are usually oval, sometimes elongate and flattened, and 1.5 to 12 mm in
length. They are usually black and often marked with red or yellow. Antennae are
11-segmented, with a variable shaped but always 3-segmented club. Elytra are
usually short and truncated at the apices, exposing the tip of the abdomen. The
larvae are nearly white, subcylindrical, moderately elongate, and usually 7 to 12

243

mm in length. Paired and forked horny projections are often found on the ninth
abdominal segment. Keys to the species occurring in Delaware are available (232).

Both adults and larvae of many species feed on decaying fruit, fermenting sap
under bark, and sap exudations from freshly cut logs, stumps, or wounds on living
trees. Bark-infesting species often kill patches of cambium (fig. 107) that heal over
to form pocket defects causing lumber degrade losses in valuable species like oak.
A number of species apparently can penetrate the bark of oak where there are no
previous injuries. Numerous studies have shown that nitidulid beetles are vectors of
oak wilt fungus, Ceratocystis fagacearum (Bretz) Hunt.

F-519925, 519926

Figure 107.—Evidence of nitidulid attacks on oak: A,
scars overgrown after attacks; B, pockets in the wood
beneath scars shown on A.

244

|

Family Coccinellidae
Lady Beetles

Lady beetles are small, usually less than 5 mm long. Their bodies are round,
usually convex or hemispherical, and shiny, often marked with red, yellow, black,
or white spots. The larvae have a large, protruding head with a fleshy, elongate,
tapering body that is covered with tubercles or spines, and have well-developed,
five-jointed legs. They are black or gray and are frequently marked with bright
colors.

About 400 species occur in the United States. There are keys to the identification
of species occurring in Minnesota (//5/). As a group, lady beetles are among the
most beneficial and most widely recognized of all insects; one review covers 206
references on predacious species (5/2). Both adults and larvae of most species are
predacious on scales, aphids, and spider mites or on the eggs and larvae of larger
insects. Both stages usually occur on the host plants of their prey, and eggs are laid
in clusters on leaves or bark. Adults overwinter, sometimes in enormous congrega-
tions. However, species in the subfamily Epilachninae are herbivorous and some,
such as the Mexican bean beetle, Epilachna varivestis Mulsant, are notorious pests
of food plants. Some other species feed on spores of mildew or rust fungi. A few of
the most important predators of forest insects will be discussed briefly.

An introduced species, the twospotted lady beetle, Adalia bipunctata (L.), is an
important predator of pine needle, juniper, beech, and terrapin scales in the
Northern States. Adults are 3 to 5 mm long. The head is black with two yellow
spots between the eyes; the thorax is black with yellow margins; and the wing
covers are red, each with a black central spot.

The twicestabbed lady beetle, Chilocorus stigma (Say), occurs in the Northern
States and feeds commonly on pine needle, beech, terrapin, and juniper scales. The
adult is 4 to 5 mm long, shiny black with a round red spot on the disk of each

elytron. The venter of the abdominal segments is red.

The convergent lady beetle, Hippodamia convergens Guérin-Méneville, feeds
on aphids and scales throughout most of the United States. Adults are 4 to 6 mm
long. The pronotum is black except for pale margins and is marked with two discal
bars; the elytra are reddish with a black scutellar spot and 12 more or less distinct
black spots; and the venter and legs are black. In some parts of the country this
species hibernates in clusters of tens of thousands. In very hot summers it aestivates
in a similar manner.

Hyperaspis congressis Watson is widely distributed in the Eastern United States
and is an important enemy of the pine tortoise scale. Adults are shiny black,
strongly punctate, and 2.5 mm long. A rather large yellowish spot occurs on the
middle of the elytra and often a fainter one is present on the wing tips. Anatis mali
(Say), the fifteen-spotted lady beetle, feeds on a wide variety of forest insects,
including young larvae of the gypsy moth and cankerworms, in the Eastern United
States. Adults are yellowish and 8 to 10 mm long. There are two pale spots enclosed
in a black disk on the thorax, a single spot on the scuiellum, and seven black spots
on each elytron.

Coccinella novemnotata Herbst, the nine-spotted lady beetle, is a common and
widely distributed species that feeds on aphids and scales on both conifers and
hardwoods. Adults are 5 to 7 mm long. The body is pale yellow, and black beneath.
Each elytron bears four black spots, and one black spot is on the elytral suture.

The transverse lady beetle, C. transversoguttata richardsoni Brown, feeds on
scale insects infesting pines in the more northern States. Adults are reddish, 6 to 7.5

245

mm long. The wing covers may be without spots or each one may be marked with a
long, transverse subbasal spot, a shorter transverse spot near the middle, and a third
near the tip.

Several species of coccinellids have been imported against the balsam woolly
adelgid in recent years. The European species, Aphidecta obliterata (L.), initially
became established in infested Fraser fir stands in North Carolina but its present
fate is unknown (396). Eggs are laid on the needles or trunks of infested trees, and
the larvae and adults feed on all stages of the adelgid except the first instar.
Attempts to establish this species in New Brunswick were unsuccessful (/63).
Scymnus (Pullus) impexus Mulsant, one of the most important enemies of the
adelgid in Central Europe, has been released in infested stands of fir in New
England and North Carolina. The adults and larvae feed on second and third instars
of the adelgid. It does not appear to be established (396).

Other important species of native coccinellids include S$. /acustris LeConte, an
enemy of the pine tortoise scale in the Lake States; Microwesia misella (LeConte), a
minute species less than 1 mm long that feeds on various species of scale insects;
Cleis picta (Randall), which often occurs in large numbers on pines infested with
the red pine scale in the Northeast; and the spider mite destroyer, Stethorus
punctum Casey, which is the mainstay of integrated spider mite control in apple
orchards in Pennsylvania.

Family Derodontidae
Derodontids

This little-known family contains about five species in the Eastern United States.
Adults are brown or motley brown and are 2 to 6 mm long. Larvae and adults of
some species occur together on or in slime molds and shelf fungi. These habitats
and the sharp projections on the margin of the prothorax have led to the name
“tooth-neck fungus beetles.”” Some native species are general, though not impor-
tant, predators of adelgids on conifers.

Laricobius erichsonii Rosenhauer, a European predator of the balsam woolly
adelgid, has been established in adelgid-infested stands of fir in Canada, New
England, North Carolina, and the Pacific Northwest. It appears to be the only
foreign predator of the adelgid established in North Carolina (396). The adult is an
elongate oval beetle, 2.2 to 2.4 mm long, with a deflexed head that is usually
hidden from view dorsally. The body is covered with yellowish to brownish to black
hairs; the central portions of the elytra, the antennae, and the legs are reddish
brown. Adults deposit eggs within adelgid egg clusters or under lichens on the bark.
The larvae, often covered with bits of adelgid’s wool and bark, feed to maturity on
adelgid eggs, drop to the ground, and pupate in cocoons of compacted soil particles
(164).

Family Cucujidae
Cucujid Beetles or Flat Bark Beetles

Adults of this family are yellow, red, brown, or black. They are extremely
flattened since they are adapted to living under close-fitting bark of unhealthy trees
or recently felled trees and logs. The larvae are depressed in shape. The majority
are scavengers but a few are predacious on mites or insects.

Cucujus clavipes F. is conspicuously bright red, extremely flat, and 10 to 14 mm
long. It is common in the Northern States, especially under bark of recently dead
ash and poplar. Catogenus rufus (F.) larvae are usually found parasitizing the pupae
of wood-boring Cerambycidae or larvae of Braconidae. The adults are reddish
brown and are up to 12 mm long.

246

Family Colydiidae
Colydiids

Members of this family vary in form and habit. They are hard-bodied, shiny,
often beautifully sculptured, and reddish brown to nearly black. Many species are
elongate, cylindrical, and rarely more than 5 mm long, but some larger species are
strongly flattened and oval. Larvae of the latter are more robust in form and usually
are found as scavengers in bark beetle galleries. Larvae of other species—
Nematidium filiforme LeConte, Bitoma carinata (LeConte), and Colydium lineola
Say—are elongate, slender forms that prey on larvae of ambrosia beetles, cossonid
weevils, and some other wood borers. Other species are phytophagous and live on
fungi or decaying vegetation.
Family Melandryidae
Melandryids

Adults of this family are 2 to 15 mm in length, and they are usually elongate to
oval, loosely jointed beetles. Their antennae are 11-segmented and filiform; their
color is dark brown to black. Adults are seldom seen, but the larvae occur
commonly under the bark on dry, decaying logs or in dry fungi.

Some of the more common species and places to expect them are: Melandrya
striata Say—many different hardwoods in association with black lines of decay;
Orchesia castanea Melsheimer—oak, maple, yellow-poplar, sycamore, and
hickory; Hypulus concolor (LeConte)—rotting pine sapwood; Serropalpus sub-
striatus Haldeman and Eustrophus tomentosus Say—various conifers; and Dircaea

quadrimaculata (Say) and Holostrophus bifasciatus (Say)—various hardwoods.
Family Oedemeridae
Oedemerid Beetles

Adults are usually elongate and subcylindrical with elytra soft in texture and
sometimes covered with fine silken hair. Their color varies from pale to black, often
with red, yellow, or orange markings. They range in length from 5 to 20 mm. The
next-to-last tarsal segment is dilated and bears a dense, hairy pad beneath. Not
much is known of the habits or habitats of the 68 species in the United States.
Adults commonly visit flowers to feed on nectar, sometimes pollen, and they are
attracted to lights. The larvae feed in damp, decaying wood.

The wharf borer, Nacerdes melanura (L.), 1s cosmopolitan and occurs on most
coastlines and around the Great Lakes (33/). Adults are reddish yellow with a dark
patch on the head, two dark patches on the thorax, and blackened tips on the elytra.
Larvae of this species, the only economically important member of the family,
occasionally are very destructive to moist wood poles, pilings, boardwalks, or
basement timbers (885). Adults are present from April to July in outdoor infesta-

tions, but they may emerge at other times within buildings.
Family Tenebrionidae
Darkling Beetles

Many species of this large family occur in the forest. The adults are hard-shelled,
usually dark brown or black, 3 to 20 mm in length. Many are clumsy and slow-
moving. The larvae are long and slender, and are often covered with tough, horny
skins. The majority of forest species feed as scavengers on dead vegetable matter
and fungi. They are commonly found under the bark of dead and dying trees or
decaying logs, and occasionally in the galleries of bark beetles. A few species in the

genus Strongylium bore into the wood of living trees at stubs or wounds.
Family Lymexylonidae
Timber Beetles

Two species of timber beetles are known from the Eastern United States. Adults
are slender and elongate; the head is deflexed and narrowed behind the large eyes to

247

form a neck. Larvae are also elongate and slender, with globular heads. They bore
deeply into timbers, causing pinhole defects—small tunnels across the grain having
dark, stained walls.

The chestnut timberworm, Melittomma sericeum (Harris), once a destructive
pest of chestnut, 1s now found chiefly in white oak. The nocturnal adults are brown,
clothed with fine silky hair, and are 11 to 15 mm long. Eggs are laid in cracks in the
wood surface.

The sapwood timberworm, Hylecoetus lugubris Say, 1s a reddish-brown or
blackish beetle 10 to 12 mm long. Eggs are laid from April to July on bark of dying
trees or unseasoned logs. Larvae bore across the grain of the sapwood of various
hardwoods, such as poplar, birch, yellow-poplar, basswood, buckeye, and black
walnut. The larvae have a distinguishing feature, a conical, slender, barbed spine on

the ninth segment.
Family Mordellidae
Tumbling Flower Beetles

Adults of this family can be recognized by their habit of jumping or tumbling off
flowers and by their characteristic strongly arched, wedge-shaped bodies that
usually end in a long, conical process. They are usually black or dark brown, often
spotted or banded with yellow or silver, and from 3 to 7 mm long. The head is bent
downward, and the hindlegs are long, flattened, and spiny. The larvae of many
species feed in rotting wood; some species are reportedly predacious.

Beetles That Damage Seasoned Wood

Numerous beetle species may damage seasoned wood in storage or use. Beetle
larvae bore meandering, powder-filled feeding tunnels within wood, and the adults
chew small round or oval holes in the surface of wood as they emerge. Adults of a
few species bore short tunnels into wood for egg laying. When heavily infested or
repeatedly attacked over an extended period, wood is often reduced to a mass of
powder surrounded by a thin shell of sound wood perforated with holes. Damage of
this type has commonly been termed “powderpost.”’

Beetles that cause a powderpost type of damage to wood are, as a group, ignored
and misunderstood by many entomologists, foresters, and wood users. Damage,
superficially similar in appearance, may be caused by numerous species within the
families Anobiidae, Lyctidae, Bostrichidae, Ptinidae, Curculionidae (subfamily
Cossoninae), and a few species within the families Cerambycidae and
Oedemeridae. Many of these beetles can reinfest wood after it is dry, although some
require wood in a very moist condition (Oedemeridae, Cossoninae weevils, and
Ptinidae) and with bark attached (most Bostrichidae).

Seasoned wood may also contain damage from various other beetles within the
families Buprestidae, Cerambycidae, Platypodidae, and Scolytidae. Although
adults initiate the attacks in the moist wood of trees recently dead or dying or in
drying logs and lumber, larvae may complete their development in wood after it is
dry. These beetles, but not necessarily their damage, are usually removed during
wood processing; damage is frequently misidentified as that of beetles capable of
reinfesting dry, seasoned wood. Occasionally, adults may emerge from air-dried
wood after it is in use. Additionally, larvae of the families Dermestidae, Ten-
ebrionidae, and nonwood feeding Anobiidae occasionally use wood as a pupation
site.

Fortunately, the numerous beetles that damage seasoned wood may be identified
to group by characteristics of habits and damage. The incidence and location of

248

damage are dependent on the habits and food requirements of the various beetles
and on people’s utilization of wood (/299, /302). Infested wood products result
from susceptible wood being placed in favorable conditions for beetle attack. Some
of the many beetles that can reinfest seasoned wood have less restrictive require-
ments than others. Therefore, some beetles occur in structures much more fre-
quently than others, e.g., Anobiidae, Lyctidae, and Cerambycidae (old house
borer, Hylotrupes bajulus (L.)) (fig. 108). These three groups of beetles can be
distinguished easily by characteristics of damage. Group, not species, identification
is necessary for the proper selection of control measures. Failure to make distinctive
identifications, or failure to use distinctive common names may lead to inadequate
or unnecessary control operations (/300). Habit and damage characteristics are
summarized for the beetles most frequently attacking seasoned wood (table 2).
Many different beetles with different requirements may occur in the same situation.
Therefore, key damage identification characteristics are given to distinguish in-
festations of beetles that cannot reinfest seasoned wood from those that can. In
addition, guides for inspection of new and existing structures are available to aid
wood users in damage identification by providing color illustrations of damage by
various beetles, other insects, and decay organisms, and by giving the incidence of
each organism both within structures and within the United States (729, 730). For
the most important beetles infesting seasoned wood, conditions affecting their
attack and changes in wood processing and use that influence the likelihood of
infestation are reviewed to suggest the outlook for prevention and control (/302).

Family Anobiidae
Deathwatch and Drugstore Beetles

This family contains about 52 genera and 318 species in the United States. The
common names “‘deathwatch beetles” and “drugstore beetles” are not useful terms.
A more acceptable name would be “shield beetles’ because of the way they shield
themselves, as described below. Adults are of widely diverse sizes and colors,

fj
RS

F-521295

Figure 108.—These are adults—seldom seen by the
homeowner—of the three most destructive groups of
wood-destroying beetles. Left: One of the Lyctus, or
true powderpost beetles, which attack recently
seasoned hardwood sapwood. Center: One of the
anobiid beetles, which are among the most common
feeders on both hardwoods and softwoods within
buildings. Right: The old house borer, sometimes
called a longhorn borer from its antennae, primarily
attacks softwood framing lumber.

249

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ranging from 1.5 to 8 mm and from bright reddish-brown with orange patches on
the wing tips to a shiny ebony black. Adults of many species appear yellowish or
grayish from the dense pubescence covering their bodies. To varying degrees, all
species share common characteristics of a hoodlike pronotum that conceals the head
when viewed from above, convex body shape, and the contractile ability to shield
their appendages by retraction. Many genera and species can retract their appen-
dages into specialized grooves when at rest; thus, they resemble the seeds of their
host plant.

Members of this family have widely diverse habits, food hosts, and habitats.
Anobiid beetles occur in most forest habitats, but many are not well known or
frequently collected. Apparently, no species causes damage of economic signifi-
cance to living plant material although many species in the genus Ernobius are
frequent secondary invaders of aborted pine cones. Many species smaller in size
feed on puffballs and other fungi or on plant seeds, but the majority of the larger
species have wood-infesting habits. Many feed on dead twigs and branches or
aborted cones, others in or beneath the bark of decadent trees. Many other species
require bark-free dead wood such as branch stubs, lightning scars, or fallen trees.
These species perform a function similar to that of subterranean termites and help
reduce dead wood to nutrients for new plant growth. Some species with this
function are restricted to wood of deciduous trees, others to only coniferous wood,
and still others can attack wood of both types. Some of the species that attack
seasoned wood have become frequent pests of wood in fabricated structures,
especially when conditions surrounding the wood are favorable to their develop-
ment.

Two nonwood-feeding species have become so adapted to human activities that
they are now cosmopolitan pests of stored products. These are the drugstore
beetle, Sregobium paniceum (L.), and the cigarette beetle, Lasioderma serricorne
(F.), and they will feed in a wide variety of products including cereal grains, spices,
tobacco, etc. Occasionally, they will excavate wood near food products when
pupating, thus suggesting that they originaliy had wood-infesting habits.

Only a few species that most commonly damage seasoned wood in storage or use
will be discussed. Adults of most species attacking seasoned wood are reddish
brown to brownish black and range from 2.8 to 7.5 mm in length. With the
exception of a few species, eggs are usually laid on the wood surface in cracks or
crevices or in the holes made by emerging adults. The larvae are whitish with
yellowish-brown heads, grublike, and may be 10 mm in length at maturity. The size
of the adults and their exit holes are variable within a species. Exit holes are
circular, typically 3 to 4 mm in diameter. The duration of the life cycie varies with
the temperature and relative humidity conditions surrounding the wood as well as
with its nutritive content. The cycle may require | to 5 years or more.

Many of the wood-infesting species can be identified with keys (/253, 1285).
Xyletinus peltatus (Harris) is the most widely distributed species in the Eastern
United States, with a range from southern Canada to Texas to Florida. X. peltatus 1s
probably the most common species infesting structures in the southern half of its
range (869, 1/42, 1308) and the third most common species in the northern half
(1086). Adult beetles are seldom seen because they are nocturnal and may live for
only a few weeks. On the Mississippi Gulf Coast, adult emergence may begin in
mid-April, peak in late May or early June, and continue through September, but 80
to 90 percent of the adults emerge within a 4- to 6-week period surrounding the
peak of emergence (/309). These beetles cause the greatest damage in the warm,

22

humid areas of their range. Nevertheless, infestations usually occur only when
unfinished, untreated wood is exposed in a humid environment such as improperly
stored lumber, in crawl spaces of buildings, or in outbuildings (/298, /299).
Damage may occur to heartwood if sapwood is attacked, but usually only the
sapwood is damaged. Wood from both coniferous (softwood) and deciduous (hard-
wood) trees are attacked; in fact, yellow-poplar sapwood has proved to be a
preferred species for egg laying, and best for larval growth and survival over seven
other woods tested (/306, /307). Most structural infestations originate in soft-
woods that are exposed in crawl spaces (fig. 109), as softwoods are most commonly
used for building construction. With time and favorable conditions, infestations
may spread upward to the hardwood flooring, molding, and furniture. Populations
within infestations increase slowly, and wood has usually been in service 10 or
more years before attacks are discovered (/304, 1308). Several factors explain this:
(1) relatively few eggs are laid per female, (2) the life cycle is usually at least 2
years in favorable woods and may be 3 to 5 years in unfavorable woods, and (3)
damage is not easily detected until adults emerge and make holes on the exterior
surface. These factors just discussed, larval feeding primarily in springwood layers
of softwoods (/30/), and load sharing capacities of complete floor systems indicate
that control of infestations in crawl spaces can be achieved after attack and before
structural weakening occurs if done before infestations spread upward (/303).

F-531251
Figure 109.—Damage to softwood lumber in crawl

space of a house. Note the piles of frass as the result
of feeding by anobiid powderpost beetle larvae.

Hemicoelus carinatus (Say) is the most common species in structures in the
Northeastern United States (/086). Its distribution is from Maine to Manitoba
southward to Kansas and eastward to North Carolina. Both hardwoods and soft-
woods are attacked, with the preponderance of attacks paralleling the prevalence of
hardwoods in the region. Host woods include: silver and sugar maples, yellow and
paper birches, white and northern red oaks, white ash, American beech, and
American basswood (/084). The frass is loosely packed within tunnels and will fall

out when damaged wood is gently tapped. The frass contains elongated pellets
(1087).

253

Common predators and parasites associated with H. carinatus include: Pel-
ecotoma flavipes Melsheimer (Coleoptera: Rhipiphoridae), Heterospilus flavicollis
(Ashmead), and Histeromerus canadensis Ashmead (Hymenoptera: Braconidae).

Ptilinus ruficornis Say is the second most common species in structures in the
Northeast (/086), but is the most common species in northeastern hardwood forests
(3). This species attacks only hardwoods, and this habit may possibly explain why
it is found in forests more than in structures. Its distribution is from Newfoundland
to Alberta and extends southward to Kansas, southeastward to Alabama, and
northeastward to Virginia. Host woods include the same hardwood species given
for H. carinatus except the oaks and ash (/084). P. ruficornis females bore short
tunnels into bark-free wood for egg laying. Apparently, females have a mating or
aggregating pheromone (3). Unlike other anobiids, the frass is very tightly packed
in larval tunnels and does not contain discrete pellets.

Other native and introduced species may occasionally be found damaging wood
in use. Their occurrence in structures is of academic rather than practical signifi-
cance because species identification is not needed for controlling infestations.
Descriptions and distributions of these species are available (/086, 1283, 1284).
Additional information (3) is available for the native species which include:
Platybregmus canadensis Fisher, the Canadian powderpost beetle: Priobium
sericeum (Say); Oligomerus obtusus LeConte; O. alternans LeConte; and Xyletinus
harrisii Fall.

A number of species have been introduced that are well-known wood-destroying
pests in other parts of the world: Nicobium hirtum Uliger (//23); the furniture
beetle, Anobium punctatum (De Geer) (132, 567, 740); Xestobium rufovillosum
(De Geer), the deathwatch beetle (/32, 567); and Ptilinus pectinicornis (L.)
(1085). Although these species have been reported from various locations, probably
only the first species has a very wide distribution.

Many anobiid beetle species can infest only dead wood in branch stubs or
lightning scars of living trees. Various species within the genus Ernobius com-
monly occur in bark, dead twigs, or aborted cones on coniferous trees. E. mollis
(L.), an introduced species, has been reported from southeastern Canada southward
to Florida and Texas. The larvae feed under the bark but may bore into the wood
where the bark is thin or when infested wood with bark is used. Thus, adults may
emerge from wood in structures but they cannot reinfest dry wood. Bark-beetle
weakened trees are particularly susceptible to attack. E. granulatus LeConte, E.
tenuicornis LeConte, and E. filicornis LeConte are other species of rather wide-
spread but not well-known distribution. Adults of these Ernobius species are
yellowish to reddish brown to brownish black and they have a similar range in size
of 2.3 to 4.3 mm. More specific descriptions are available (/253, 1254). Their
hosts are not well known.

Many other anobiids are apparently indigenous to hardwood forests and reported
only from hardwood hosts (3, 1/283). Many of the species are in the genus Petalium
(437).

Petalium bistriatum (Say) occurs from Massachusetts and New York to Ohio and
south to the Gulf of Mexico. It breeds in the twigs of various hardwoods such as
oak, dogwood, walnut, and buckeye (74). Adults are 1.8 to 3.0 mm long: the head,
pronotum, and undersurfaces are reddish black to black, the elytra black, the legs
reddish, and the antennae yellow. P. seriatum Fall has been observed breeding in the
dead twigs of pine, oak, and bittersweet. Adults are reddish brown to nearly black
and are 1.5 to 2.5 mm long.

254

Hadrobregmus notatus (Say) occurs in southeastern Canada and southward to
North Carolina and Mississippi. It has been recorded feeding in dead oak branches
in Ohio. Adults are dark reddish-brown and 3.2 to 4.3 mm long. Eucrada humeraiis
(Melsheimer) larvae feed beneath the bark of dead oak and beech trees. It occurs
from Quebec to Michigan in the North and southwestward and southeastward to
Iowa and South Carolina.

Many of the smaller anobiid species within the genera—Caenocara, Dor-
catoma, and Tricorynus—feed on various plant seeds and fungi. Some species have
been reported attacking stored pine cones or small tree branches.

Family Lyctidae
Powderpost Beetles

This family contains 12 genera and more than 60 species worldwide. Each region
has indigenous species plus established introduced species. In fact, many species
have been widely distributed through commerce and they are considered cosmopol-
itan. The taxonomy of the family has been revised and keys to identification are
available to all 12 genera and to all 35 species reported from North America (47/).

Adults are elongate, flattened, reddish brown to black, and 5 to 6 mm long.
Unlike anobiids and bostrichids, the head is easily seen in dorsal view. The head is
slightly deflected and constricted behind the eyes. The antennal club is two-
segmented. Larvae are white and grublike; the body 1s C-shaped and enlarged at the
thorax. The eighth spiracle on the abdomen is much larger than all the others; this
characteristic distinguishes lyctid larvae from anobiid or bostrichid larvae.

Beetles in the family Lyctidae probably have a greater capacity for destroying
portions of seasoned wood in a shorter period of time than any other native beetle.
In comparison with other beetles, females lay many eggs, larval populations are
high, and the life cycle is short. Other biological characteristics, however, severely
restrict the frequency of infestations and extent of damage. For wood to be
susceptible to lyctid attack, it must have at least 3 percent starch and pores (vessels)
large enough for the insertion of eggs; eggs are not laid on the surface of wood.
These requirements usually restrict damage by lyctid beetles to the unfinished
sapwood portions of hardwoods, although heartwood of some wood species may be
attacked also. Lyctid beetles are primarily problems of firms that produce, process,
or sell hardwood products. Wood loses starch through respiration after tree felling,
and infestations are rare in wood that has been in service more than 4 to 5 years
(131, 471). The frequency of infestations and the sum of damages are largely
dependent upon hardwood species utilization and storage and processing pro-
cedures. In general, the likelihood of attack and the extent of damage are propor-
tional to the amount of sapwood, the starch content, and the length of storage for
unfinished hardwoods.

The wood of ring-porous species is more susceptible to attack than that of
diffuse-porous species. Highly susceptible native woods include ash, white oak,
pecan, and hickory. Other susceptible native hardwoods include cherry, elm,
persimmon, sycamore, walnut, and many others. Many of the lighter colored, low
density tropical hardwoods are highly susceptible to lyctid attack, e.g., banak
(Virola spp.), luaun/meranti (Shorea spp.), and obeche (Triplochiton scleroxylon K.
Schum. ). Information on the identification, uses, and insect resistance of native and
imported woods is available (/00, 136, 300).

Lyctid beetles do not infest domestic hardwoods as frequently as they once did;
since the demand for these woods is greater now, prolonged storage is not required
and there has been much improvement in transporting, processing, and drying.

255

Proper kiln drying eliminates infestations within raw hardwood materials, and use
of water repellents seals pores of processed materials to prevent egg laying. Because
of limited supplies, many of the most susceptible hardwoods such as ash, oak,
pecan, and walnut are used as veneers, especially in furniture and paneling.
Processes of drying and gluing veneers minimize lyctid beetle attacks.

Lyctid infestations are frequently associated with the use and movement of
tropical hardwoods (/3/, 275). This same association should occur in the United
States because imported hardwoods are now frequently used as millwork, molding,
picture framing, or as core material for plywood and paneling (7/0). These and
other solid wood products are exposed to lyctid attack during the long period from
tree felling in the tropics to final use in this country.

Adults may emerge from infested wood at any season in heated buildings. Wood
may be successfully attacked when its moisture content is between 6 and 30
percent. Larval development is better when the moisture content is near the
midpoint of this range rather than at either extreme (/038). Thus, wood in any
centrally heated or airconditioned building contains enough moisture for suc-
cessful attack, and the same wood may be infested repeatedly until it is rendered
useless. Infestations may also spread to other hardwoods nearby. In this manner and
through the sale of infested products, lyctid beetles may become widely dispersed
from a single source. Nevertheless, lyctid beetle damage is more common in the
warmer Southern States than in the Northern States.

About 14 species in the genus Lyctus were reported from the United States. The
southern lyctus beetle, L. planicollis LeConte, is a common native species. Adults
are usually black but may be reddish brown; they are 4 to 6 mm long. Usually there
is only one generation per year, but under favorable conditions a generation may be
completed in 3 months or less (206, 207, /36/). Information on rearing techniques —
has been published (833, //13, 1360).

Lyctus brunneus (Stephens), an introduced species, is the species most fre-
quently associated with imported woods. Adults are 4 to 6 mm long; most of them
are reddish brown, but some are black. L. brunneus is widely distributed and
studied throughout the world. Rearing methods (529, /038) have been developed
and information on its biology summarized (/3/, 567).

Lyctus africanus Lesne and Minthea rugicollis (Walker) are also often inter-
cepted in imported hardwood products. Both species appear to be of tropical origin
and apparently have not become established in this country.

Trogoxylon parallelopipedum (Melsheimer) is another common native species.
Adults are smaller in average size than adults of L. planicollis and usually reddish in

color. Studies of the biology of this species have been published (206, 207).
Family Bostrichidae
False Powderpost Beetles

The family Bostrichidae contains about 455 species, most of which are indige-
nous to the tropics. Like the anobiids, members of the family vary considerably in
habits—most species attack dead portions of trees, some breed in fungi, some are
cosmopolitan pests of stored products, many species attack a wide variety of plants,
and a few species reportedly attack weakened living trees. Like lyctid beetles,
starch is an essential nutrient, and the sapwood portions of hardwoods are fre-
quently attacked. Unlike lyctids, bostrichids do not reinfest wood after it is dry.
Many of the larger bostrichid species in tropical regions cause extensive damage to
unseasoned hardwoods used as packing crates and to bamboo products (/32).

256

Female beetles lay eggs in cracks or crevices on the wood surface or bore short
tunnels in bark-free wood.

None of the more than 60 native species attacks bark-free wood except possibly
the leadcable borer, Scobicia declivis (LeConte), in the West (367). Seasoned
wood suffers no damage of major economic importance caused by any species
native to eastern North America. Attacks are initiated in partially seasoned wood
containing bark. Females bore short tunnels into the outer sapwood and deposit
eggs. Once attacks are initiated, larvae of many species may continue to feed and
develop at a reduced rate. Adults may emerge from wood in use, usually within a
year, but possibly even as long as 5 years after tree felling. Because bostrichids
differ from lyctids in adult appearance, size of exit holes, and frass, bostrichids are
referred to as false powderpost beetles. Many other species attacking trees are often
called branch or twig girdlers, a name more appropriate to their habits.

Adults of native species are reddish brown to black and 3 to 6 mm long. They
resemble adults of the family Scolytidae but differ in that the pronotum is tubercu-
late and rasplike, the tarsi are five-segmented, and the antennae are straight rather
than elbowed. The antennal club is three- or four-segmented; the thorax is usually
hoodlike, covering the head; and the posterior portion of the elytra is frequently
concave and tuberculate. The femora and tibiae are broad and the latter are
frequently toothed on one margin (425).

Xylobiops basilaris (Say), the redshouldered shothole borer, is a common
species, widely distributed over much of the Eastern United States. The adults are
basically black but the basal part of the elytra is dull reddish-orange, hence the
common name (fig. 110). Hickory, pecan, and persimmon are most frequently
infested, but several other hardwoods are also attacked. Larvae feed parallel with
the grain, mostly within the sapwood, but heartwood may be damaged to some
extent. Populations may be high, thus extensive damage may result. Feeding may
continue until the wood is very dry. Adults often emerge from wood within
structures such as firewood.

The apple twig borer, Amphicerus bicaudatus (Say), is another common species
in most of the Eastern United States. It breeds in dying wood such as large
prunings, exposed roois, and injured branches of most shade and fruit trees. Some
of the many other species and the woods they attack are: Lichenophanes arminger
(LeConte)—various hardwoods, chiefly oak; L. bicornis (Weber)—sycamore,
hackberry, oak, pecan, hickory, apple, beech, elm, black locust; Scobicia bidentata
(Horn)—hickory, elm, oak, chestnut, hackberry, and sassafras.

Several native bostrichids commonly attack softwood timbers. Adults may
emerge within structures from bark-covered wood that is used as structural timbers
or rustic furniture. Stephanopachys rugosus (Oliver) is widely distributed in the
Eastern United States. It and several other species in the same genus—S. densus
(LeConte), S. cribratus (LeConte), and S. hispidulus (Casey)—breed in pine. S.
substriatus (Paykull), found in most of the Northern States, attacks Douglas-fir and
hemlock, as well as all species of pines and true firs.

Many of the tropical species are frequently intercepted during quarantine inspec-
tions at ports of entry to this country. Even though tropical in origin, at least two
species are established in some Southern States.

The bamboo powderpost beetle, Dinoderus minutus (F.), occurs in Florida and
Louisiana. It is frequently intercepted in bamboo products at ports of entry. Adults
are reddish brown to brownish black and are 2.5 to 3.5 mm long. The wing covers

Zo

F-519941, 480481

Figure 110.—xylobiops basilaris, a false powderpost
beetle: A, adult; B, larval damage (note openings to
frass-filled larval tunnels in cross section of a
persimmon log).

are often redder than the rest of the upper surface. The female bores tunnels across
the grain into bamboo culms at breaks in the rind and then deposits eggs in exposed
vessels. The larvae often reduce products to masses of powder surrounded by easily
shredded fibers.

In 1967, a large infestation of Heterobostrychus aequalis (Waterhouse), the
oriental wood borer, was found in oak and mahogany lumber in a mill yard in
southern Florida (/2/8). This is one of the most common and most serious wood-
boring beetles in India. It is widely distributed in Southeast Asia and it has been
recorded attacking the wood from more than 35 species of trees. The adults are
reddish brown to black and are from 6 to 13 mm long. The larvae become large, up
to 13 mm in length. Usually the life cycle is 1 year, but it may take about 6 years.

258

Several species of western origin may be shipped to the Eastern States in infested
wood. Polycaon stouti (LeConte) is commonly encountered. Its appearance 1s
unlike other bostrichids in that the head is visible and the pronotum is not hoodlike.
The pronotum and elytra lack tuberculate projections. The subcylindrical body is

coal black and from !2 to 25 mm long.
Family Ptinidae
Spider Beetles

The common name for this family appropriately describes the appearance of
these beetles. Adults are 2 to 4.5 mm in length. The head and pronotum are much
narrower than the abdomen, the legs are long, the antennae are long and filiform,
and the elytra are usually very convex and shiny. The majority of species breed in
old grass roots or in dead bark. The brown spider beetle, Ptinus clavipes Panzer,
and the whitemarked spider beetle, P. fur (L.), occasionally damage pine boards
in old buildings. Both are cosmopolitan species that frequently damage dried
vegetable and animal products in warehouses and museums. Gibbium psylloides
(Czenpinski) occasionally is a pest in attics of houses where organic matter has

accumulated.
Family Bruchidae
Seed Beetles

Members of the family Bruchidae are distinguished by their compact and usually
oval bodies. Their small heads are prolonged into beaks, and their short wing
covers leave the top of the abdomen exposed. The larvae are quite small and feed
almost entirely in the seeds of plants.

Amblycerus robiniae (F.) is occasionally a pest in the Eastern United States. The
female deposits her eggs on the pods of honeylocust and the larvae feed on the seeds
within the pod. Adults are reddish brown and about 7 mm long. The body is clothed
with grayish-yellow hairs and there are five rows of black spots across the elytra.
Gibbobruchus mimus (Say) breeds in the seeds of the redbud and Caryobruchus

gleditsiae (L.), in palmetto.
Family Chrysomelidae
Leaf Beetles

Leaf beetles are one of the larger families of Coleoptera, with about 2,000
species occurring in North America. The adults usually are medium size or small,
short-bodied, and more or less oval. The legs are generally short, but in some
species the femora of the hind pair are enlarged. There is great variation in coloring
and markings, occasionally even within a species. In some species the entire body,
elytra, and legs may have a bright metallic sheen. Some are hairless; others are
pubescent or covered with scales or scalelike hairs.

Leaf beetle larvae are usually soft-bodied and frequently have highly pigmented
sclerites. They vary greatly in shape, depending on their feeding habits. The head
usually protrudes, except in the leafminers, and is bent downward for feeding. The
body varies greatly in shape, from short and compact to depressed cuneiform,
depending on whether the larva is free-living or a leafminer, casebearer, or root
feeder.

All members of the family feed on the foliage of plants as adults or larvae, or
both. The adults are diurnal and are usually slow moving. Species that are exposed
in the larval stage feed gregariously, whereas leafmining larvae usually feed singly,
each within its own mine. Adult feeding is characterized by the presence of holes
eaten all the way through the leaf, or by skeletonization usually of the lower surface
of the leaf. Free-living larvae either fasten themselves to the surface of the leaf to
pupate, or move to the ground to do so. Leafmining larvae pupate either within the

259

mine, or they vacate it and pupate in the ground. There is considerable literature on
this family (53, 723, 724, 725, 1289).

The cottonwood leaf beetle, Chrysomela scripta F., occurs throughout the
United States and Canada and feeds on the leaves of poplar, willow, and alder.
Adults are about 6 mm long. The head and thorax are black and the margins of the
thorax are yellow or red. The wing covers usually are yellowish with black stripes,
but are sometimes almost pure golden to black. Mature larvae are blackish and
about 12 mm long. There are two whitish spots on each side. They are located at the
site of the scent glands, one on each side of each segment except those on the
prothorax and the last two on the abdomen. The scent glands emit a pungent odor
when the larvae are disturbed.

Winter is spent in the adult stage under loose bark or debris or in clumps of grass.
Emergence begins in early spring and the adults feed on unfolding leaves or on
tender bark at the tips of twigs (528). Eggs are deposited in groups of 15 to 75 on
the undersides of leaves. The young larvae are gregarious and, feeding side by side,
skeletonize the leaves. Later, they feed separately and consume the entire leaf,
except the larger veins. At maturity they attach themselves to leaf surfaces, the
bark, or to weeds and grass beneath the trees to pupate. There are several genera-
tions per year. Severe infestations occur occasionally and cause considerable
damage.

The aspen leaf beetle, C. crotchi Brown, and C. interrupta F. occur commonly
throughout the Eastern States. The adults and larvae of C. crotchi feed on poplar
(1093); those of C. interrupta feed on the leaves and at times on the tender bark of
alder. C. knabi Brown feeds on poplar in the Northeastern States. It has also been
observed feeding on willow in Tennessee.

The elm leaf beetle, Pyrrhalta luteola (Miller), an introduced species first:
recorded in North America at Baltimore, Md., well over a century ago, now occurs
throughout most of the United States (fig. 111). Its hosts are all species of elm. In
the Northeastern States, American elm is often severely attacked and seriously
damaged. Farther west, Siberian elm is also frequently heavily attacked. European
elms are especially subject to injury. Adults are about 6 mm long and yellowish to
dull green, with a black stripe along the sides of the wing covers. There is also a
short, dark spot at the base of each wing cover. Full-grown larvae are dull yellow,
with two rows of black spots on the dorsum, and are about 12 mm long. The head,
legs, and tubercles are black and there is a broad, yellow stripe down the middle of
the dorsum. Pupae are bright orange-yellow with a few black bristles and they are
about 5 mm long.

Elm leaf beetles spend the winter in sheltered dry places. In the Northeastern
States most of them hibernate in house attics, barns, and sheds. During periods of
warm weather in the winter many of these beetles become active and cause
considerable annoyance by crawling into living quarters or onto windows. Spring
emergence begins about the time the buds of elm begin to swell, and the adults fly
to nearby elms and feed by chewing holes in the unfolding leaves. Egg laying
begins in late May or early June, each female laying from 400 to 800 eggs (/223,
1264). Hatching begins in about | week and the larvae feed for 2 or 3 weeks on the
undersurfaces of the leaves. Only the veins and upper surfaces are left, and the
leaves soon dry out and turn brown. Full-grown larvae crawl down the tree and
pupate in bark crevices or at the base of the tree. In the Eastern United States there
may be one or two complete generations and a partial third. Usually the first

260

Courtesy Agric. Res. Serv.

Figure 111.—Elm leaf beetle, Pyrrhalta luteola. A,
undamaged elm leaf; B, elm leaf showing typical
feeding damage, skeletonization, and perforation; C,
egg mass; D, larvae; E, pupa; F, adult.

261

generation is the most injurious. Beetles maturing in the summer begin entering
hibernation quarters as early as August, on or near the tree on which they fed.

Shade tree elms are often heavily defoliated, whereas those growing in the forest
are usually not seriously infested. The native parasite, Tetrastichus brevistigma
Gahan, and the coccinelid predator, Coleomegilla maculata (De Geer), occasion-
ally exert a considerable degree of control in the Northeastern States (/02, 1263).

The cherry leaf beetle, P. cavicollis (LeConte) occurs in southern Canada and in
the Eastern States southward through the Allegheny and Appalachian Mountains to
North Carolina and westward to the Rocky Mountains. Its preferred host is pin
cherry but it also feeds on plum, other cherries, and peach in heavily infested areas.
The adult is red, shining, coarsely punctured, and about 5 mm long. Larvae are
dark brown, with black and yellow spots, and are about 6 mm long.

The winter is spent in the adult stage. In the spring, eggs are deposited in the soil
at the base of the tree or on the trunk near the base. The larvae climb the tree and
feed on the foliage. Full-grown larvae return to the ground and pupate in the leaf
mold or soil. There is one generation per year. The gray willow leaf beetie, P.
decora decora (Say), and P. tuberculata (Say) feed on willows.

The elm calligrapha, Calligrapha scalaris (LeConte), has been reported from
eastern Canada and from several widely distributed points in the Eastern United
States. Its preferred and probably only host is elm. Adults are elongate-oval,
creamy-white beetles from 8 to 10 mm long. The head and thorax are dark, metallic
green. Each elytron bears from 10 to 14 metallic green spots, a dark-green, boot-
shaped spot at the base, and a dark, metallic-green, irregular stripe along the inner
edge. Full-grown larvae are hump-shaped and have yellow heads. The abdomen is
light yellow or cream with a black line down the middle of the dorsum.

Adults overwinter in bark crevices, in sheltered places around the base of the:
tree, or in the top 2.5 to 5 cm of soil. When the adults emerge in the spring, they
chew oval or circular holes in the leaves. The larvae devour entire leaves except the
veins. When they reach maturity, they craw] down the trunk in search of places to
pupate and spend the winter. In heavy infestations, they frequently occur by the
thousands on the trunk or under the larger limbs. American elm is sometimes
heavily defoliated and damaged in the Midwest (283).

Calligrapha multipunctata bigsbyana (Kirby) sometimes completely defoliates
willows over large areas in the Northeastern States. It is also a common species in
the Midwest. The adult is a somewhat bronzed, metallic-green beetle about 6 to 8
mm long. The antennae and legs are reddish, and the margins of the pronotum and
most of the elytra are pale yellow.

The larger elm leaf beetle, Monocesta coryli (Say), occurs throughout most of
the Eastern United States from Georgia and Alabama northward to Pennsylvania,
Ohio, Indiana, and Illinois and westward to the Plains States (20). Its hosts are
recorded as native and Japanese elms, river birch, pecan, hawthorn, and hazel.
Slippery elm is especially favored. The adult is about 12 mm long. Its color is dull
yellow to dark brown, with large, greenish patches at the ends of each elytron. Full-
grown larvae are reddish brown, metallic lustered, and about 13 mm long.

Winter is spent as a full-grown larva in a cell 6 to 10 cm below the surface of the
soil. Pupation occurs in early spring, and adult emergence begins in late May. The
adults fly to the tops of their hosts, and feed on the leaves for several days. Eggs are
deposited in masses on the undersurfaces of leaves; the larvae are gregarious and
skeletonize the foliage; and there is one generation per year. Outbreaks have
occurred in river bottoms of the Piedmont in the Carolinas and Alabama.

262

The basswood leafminer, Baliosus ruber (Weber), occurs in Canada and
throughout most of the Eastern United States. Basswood is the favored host, but
oak, maple, willow, birch, hophornbeam, apple, and cherry are also subject to
attack. The adult is broad, flat, wedge-shaped, reddish yellow, and from 4.5 to 7
mm long. There are indistinct markings on its sides and on the apical half of the
elytra. Full-grown larvae are about 6 mm long. The head and thorax are brownish
red; the rest of the body is white.

The winter is spent in the adult stage under leaves and trash on the ground
beneath infested trees (578). In the Lake States, the adults emerge in the spring
about the time basswood leaves begin to unfold. They feed on these leaves,
skeletonizing them. Eggs are deposited singly at the edges of skeletonized areas on
these leaves about mid-June, and the larvae feed by mining the leaves (fig. 112).
Trees heavily infested for 2 to 3 years are characterized by thin crowns and the
presence of dead branches; some trees may be killed. There is one generation per
year.

F-506746

Figure 112.—Leaf of basswood mined
by the basswood leafminer, Baliosus
ruber.

The alder flea beetle, Al/tica ambiens alni Harris, occurs in southern Canada and
from Maine to Minnesota and New Mexico. It is sometimes a pest of alders growing
along roadsides and in parks and other recreational areas. The adult is cobalt blue to
greenish blue above, bluish black beneath, and about 6 mm long. Full-grown larvae
are dark brown to almost black above, dark yellow beneath, and a little longer than
the adult. In Maine, overwintering adults emerge in early spring and feed for a
short time by eating small holes in the leaves before laying their eggs on the lower
surface of the leaves. The larvae feed on both leaf surfaces and become mature in
about 5 weeks (1357). In the North there is one generation per year; in the South
there may be two. The elm flea beetle, A. carinata Germar, a related species, feeds
on elm; another, A. subplicata (LeConte), feeds on willow (30/).

The pine colaspis, Colaspis pini Barber, occurs from Maryland to central Florida
and westward to east Texas. It feeds mostly on southern pines but also occasionally

263

on baldcypress and ornamental spruce. The adult is an elongate-oval convex, rusty-
yellow or brown beetle with green reflections, and it is about 4.5 mm long. Fuil-
grown larvae are sparsely covered with short hairs. Small clusters of longer hairs
occur at the lower, outer edges of each body segment.

The winter is spent in the larval stage in cells in the soil. Pupation occurs in the
spring and adults begin to emerge by early May. They feed on the needles of the
host, chewing from the edges into the midrib. In light infestations, feeding is
generally limited to the needles on new growth; whereas in heavy infestations
needles over the entire crown may be attacked and entirely consumed. Where this
occurs, infested stands appear as if scorched by fire. The larvae feed on the roots of
grasses and herbaceous vegetation until fall, and then move deeper into the soil
where they spend the winter. Infestations tend to occur on pines growing along the
edges of stands bordering on grassland, or on isolated groups of pines growing in
fields or yards. Severe infestations have been recorded in pine plantations in
Florida, Georgia, and the Gulf Coast States. The biology of the species in Loui-
siana has been reported (368).

The locust leafminer, Odontota dorsalis (Thunberg), occurs in southern Canada
and throughout most of the Eastern United States. Its favored host is black locust,
but several other tree species such as apple, birch, beech, cherry, elm, oak, and
hawthorn are also attacked occasionally. The adult is an elongate, flattish beetle
about 5 to 6 mm long. The head is black and the thorax and most of the wing covers
are bright orange. The inner edge of each elytron is black, with the blackened area
widening posteriorly. The elytra are also deeply pitted, and each elytron bears three
longitudinal ridges. A full-grown larva 1s yellowish white, somewhat flattened, and
a little longer than the adult.

The winter is spent in the adult stage in bark crevices or under debris on the
ground. Overwintering adults emerge in the spring about the time the leaves begin
to unfold, and feed for a short time on the leaves, skeletonizing the lower surfaces
and eating holes in them. Eggs are deposited on the lower surfaces of leaves in
groups of three to five. They overlap like shingles on a roof, are glued together, and
are covered with excrement. All the larvae from a given group of eggs bore into a
leaf and feed in a common mine. Later, they separate, and each larva feeds in its
own mine. Before reaching maturity, a larva may mine several leaves. Pupation
occurs in the mine, and there is one generation per year.

Outbreaks of the locust leafminer occur practically every year somewhere within
its range, and black locust trees on tens of thousands of hectares are often defoli-
ated. The defoliated trees are seldom killed, however, unless the damage is incurred
during poor growing seasons. At such times trees may be killed in large numbers
(605). The eulophid parasite, Closterocerus tricinctus (Ashmead), 1s reported to
have destroyed over 50 percent of the pupae in West Virginia infestations (/258).
Direct control is seldom attempted in the forest, but is sometimes desirable in parks
and other recreational areas.

The imported willow leaf beetle, Plagiodera versicolora (Laicharting) (fig.
113), an introduced species, was first reported in this country from Staten Island,
N.Y., in 1915 (589). It is now widely distributed in the Eastern States and southern
Canada (it has also been reported from Alaska) where it feeds on several varieties of
willow and poplar. The adult is moderately stout, oval, and about 3.5 to 4.5 mm
long. It is metallic blue or greenish blue, and sometimes tinged with red or bronze.
Full-grown larvae are almost jet black and about 5 mm long. Rows of protuberances
run both across and along the body.

264

Courtesy Can. For. Serv., Can. Dep. Environ.,
Sault Ste. Marie, Ont.

Figure 113.—The imported willow leaf beetle,
Plagiodera versicolora. Note skeletonization of leaves
by larvae.

The winter is spent in the adult stage under the bark, or in debris or tufts of grass
around or near the base of trees. The beetles emerge in April or May and feed for a
short period by skeletonizing the leaves or by cutting holes through them. Then the
female lays irregular masses of eggs on the leaves. Hatching occurs in about a
week. The larvae are gregarious and feed in groups or in rows on the leaf surfaces,
which they skeletonize. Three generations per year, and a partial fourth, have been
recorded in Massachusetts. Additional generations probably occur farther south-
ward. Heavily infested trees may become entirely brown as early as mid-June. A
considerable degree of natural control is exerted by the imported pupal parasite,
Schizonotus latus (Walker) (326). Extremely cold winters are also fatal to poorly
protected adults.

Systena marginalis (liliger) feeds on oak in the Midwest and on baldcypress in
northern Florida and southern Georgia. The adult is dull, pale yellow except for two
black lines along the front margin of the elytra and a single black line along the hind
margin. The wing covers are densely and coarsely punctate. Adults are present from
mid-June to late August and feed by gouging out linear-shaped punctures in the
leaves. This usually causes part or all of a baldcypress leaflet to turn red and die.
The beetles occur in large swarms that tend to move about, spending only | to 3
days in any one place. A single swarm may encompass more than a dozen trees.

Zengophora scutellaris Suffrian, an introduced species, feeds on cottonwood
and other poplars from New York and New Jersey to Montana and New Mexico.
The adult is about 4 mm long. The head, prothorax, and legs are yellow; the tarsal
claws are toothed; there is a prominent tubercle on each side of the prothorax; the
elytra are coarsely punctate; and the abdomen is black. The remainder of the body
is yellow. Adults feed by skeletonizing the lower surfaces of leaves. The larvae feed
singly in the soft inner tissues, chiefly against the upper surface of the leaf, making

265

large black blotch mines. When they become full grown they vacate their mines and
drop to and enter the ground. Here they construct cells 6 to 10 cm below the surface
in which to pupate. Trees heavily fed on by both larvae and adults may be
completely defoliated.

Glyptoscelis pubescens (F.), the hairy leaf beetle, occurs primarily east of the
Mississippi River from Canada to Georgia and feeds on various species of pines
and, reportedly, spruce and hemlock (67/). The adult is elongate-oval, robust,
broadly rounded posteriorly, dark brown with a brassy or golden sheen, and is from
7 to 10 mm long. It is also sparsely clothed with a mixture of white and brownish
hairs (fig. 114). Adults feed on the edges of pine needles, causing them to turn
brown. Defoliation has been reported in seed orchards of Virginia, shortleaf, and
eastern white pines in North Carolina. G. barbata (Say) feeds on hickory and
related trees from Connecticut to Pennsylvania. The adult resembles the adult of G.
pubescens except that it is smaller and its upper surface is shining brown. The genus
Glyptoscelis in the United States and Canada has been reviewed (694).

Courtesy H. C. Coppel, Univ. Wis.

Figure 114.—Glyptoscelis pubescens, the hairy leaf
beetle, ovipositing into a cocoon of the introduced pine
sawfly on eastern white pine.

Many other chrysomelids also feed on various species of trees in the Eastern
United States. A few of these and their hosts are as follows: Pachybrachis peccans
Suffrian—hickory and birch: P. tridens (Melsheimer)—willow: P. othonus (Say )—
ash and elm: and P. carbonarius Haldeman—oak; the claycolored leaf beetie,
Anomoea laticlavia (Forster)—honeylocust, black locust, elm, live oak, and silk-
tree: Tymnes tricolor (F.)—oak, walnut, and eastern hophornbeam:; Bassareus
literatus (F.)—hickory: Derocrepis aesculi (Drury )—buckeye: Plagiometriona
clavata (F.)—sycamore, basswood, and oak; Neochlamisus platani Brown—syc-
amore; Xanthonia decemnotata (Say)—oak, beech, and elm: Paria sexnotata
(Say)—redcedar: P. guadrinotata (Say)—walnut and mountain-ash; and Syneta
ferruginea (Germar)—birch and oak.

266

Family Lucanidae
Stag Beetles

Stag beetles are distinguished by very large mandibles, which in the males of
certain species are branched like the antlers of a stag, and by the plates of the
antennal club, which are rigid and cannot be opened or closed. They are usually
found in or beneath rotting logs or stumps. The larvae feed on the juices of rotting
wood.

The giant stag beetle, Lucanus elaphus F., the most familiar species, infests
dead stumps in the South. Adults are large, fearsome insects, up to 60 mm long.
Male mandibles are branched and are more than half as long as the body.
Pseudolucanus capreolus (L.) is also a common species. It breeds in the trunks of
old, partly decayed trees such as apple, cherry, willow, and oak. The adults fly at
night and are frequently attracted to lights. Other eastern species include Platycerus
quercus Weber, Ceruchus piceus (Weber), Dorcus parallelus Say, and Sinodendron
rugosum Mannerheim. The first three breed in moist, almost completely decayed

logs. S. rugosum breeds in decayed alder, willow, and poplars.
Family Scarabaeidae
Scarabs

The family Scarabaeidae is represented in the United States by more than 1,400
species, the majority occurring in the eastern half of the country. Depending on
their feeding habits, members of the family fall into two distinct groups. One group
comprises the so-called dung beetles, the larvae and adults of which are
saprophytic, feeding on such materials as dung, carrion, and decomposing plants.
The second group consists of species whose larvae feed on the roots or juices of
living plants, decaying vegetable matter, rotten wood, leaf mold, and sometimes
manure. The adults feed chiefly on the foliage of plants. They are commonly
referred to as lamellicorn leaf chafers. Many of the plant-feeding species are
important pests of nursery, plantation, woodlot, shade, and forest trees. The adults
of most of these are nocturnal and are strongly attracted to lights.

Scarab beetles have stout bodies, the last three to six or seven segments of the
antennae are leaflike and capable of being opened or closed. Their front legs are
fitted for digging. The larvae or grubs are usually thick, white or yellow, enlarged
posteriorly, bent in the shape of a crescent, and have well-developed legs.

The genus Phyllophaga is represented in the United States and Canada by more
than 100 species, the majority of which occur in the East (756). The adults,
commonly called May or June beetles, are robust, oval, light straw to very dark
brown, and from about 12 to 25 mm long (fig. 115A). The wing covers are smooth
and shiny or are covered with short hairs. The antennae are lamellate and end in
three-jointed clubs; the tarsal claws are armed with a small tooth near the middle.
The larvae, commonly called white grubs, are milky white, strongly curved, and
about 25 mm long at maturity (fig. 115B). The head is brownish, all of the hind
parts are shiny, and body contents are visible through the skin.

The adults of certain species are most abundant in the spring, usually in May;
others reach peaks of abundance in June or July (/030). They tend to stay out of
sight under stones, leaves, or trash, or in the soil during the day; and to fly, mate,
and feed at night. Eggs are laid in masses in the soil at depths of 8 to 18 cm, each
egg being placed in a cavity in the center of a ball of dirt. Newly hatched larvae feed
on organic matter; then they move to tender roots of seedlings and other plants to
feed. The winter is spent in the larval stage at depths determined by temperatures
and frost levels. Pupation takes place in the soil at depths of a few centimeters to 30

267

A, F-532842: B, F-494654
Figure 115.—Phyllophaga spp. A, adult; B, white grubs.

or more. In the South, the life cycle is completed in 1, 2, or 3 years: in the Central
States, 2 to 3 years are required; farther north, from 3 to 4 years are needed (//4/).

Phyllophaga larvae, or white grubs, have caused heavy losses in forest nurseries
and plantations in the South, the East, and Central and Lake States. Also, here and
there throughout the region, many trees are lightly to heavily defoliated by the adult
beetles every year, especially in woodlots and around the edges of forest stands. A
few of the more common and important species are discussed briefly below.

Phyllophaga drakei (Kirby) occurs throughout most of the Eastern United States
and southern Canada. Adults are dark brown, shiny, and about 25 mm long. They
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 luctuosa (Horn) occurs primarily along the Atlantic and Gulf
Coasts in sandy, oak-pine regions, but also farther north and inland to Tennessee,
Oklahoma, and Iowa. Adults are dark brown to black, moderately shiny, and about
21 mm long. They 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 tristis (F.) occurs throughout the Eastern United States and in
southern Canada. Adults are light or dark yellowish-brown or slightly reddish, and
about 12 mm long. They seem to prefer the foliage of oaks but also feed on maple,
persimmon, hickory, elm, and willow. The larvae have caused serious losses in
nurseries in the Lake States.

Phyllophaga prununculina (Burmeister) occurs in the South Atlantic and Gulf
Coast States. It is especially common in the Sand Hills of South Carolina. Adults
are reddish brown to black, with the surface either shining and slightly pruinose, or
dull smoky, and are 12 to 18 mm long. They 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 rugosa (Melsheimer) occurs mostly in the Northern States and
southern Canada. Adults are reddish brown to black, shiny, and from 18 to 25 mm
long. They feed on a wide variety of hardwoods. The larvae are often destructive in
coniferous nurseries in the Lake States.

268

Phyllophaga crenulata (Froelich) occurs throughout the Eastern United States.
Adults are brown, with a covering of short, recumbent hairs, and are about 17 to 20
mm long. They 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 forsteri (Burmeister) occurs generally throughout the Eastern
United States. Adults are reddish brown and shiny, have dusky heads, and are about
16 mm long. They 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 South.

Phyllophaga prunina (LeConte) occurs throughout the central part of the United
States east of the Rocky Mountains. Adults are chestnut brown to black and about
18 mm long. They feed on various hardwoods, such as beech, elm, walnut,
basswood, and willow. Feeding on pine has also been observed. The larvae are
sometimes injurious in nurseries in the Lake States.

Phyllophaga implicita Horn occurs mostly in the Mississippi and Ohio River
Valleys. Adults are orange-brown to brown (with the head and thorax darker), shiny,
and about 14 to 18 mm long. They 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 thirties.

Phyllophaga micans (Knoch) is found over most of Eastern United States. Adults
are brownish black, and from 15 to 17 mm long. They feed at night on female
flowers and needles of pines. Although damage is not common, one report from
Louisiana cites that 14 percent of the pine conelets were killed by this scarab (547).

The genus Polyphylla is represented by a number of species in the Eastern United
States, a few of which are sometimes injurious. The beetles are somewhat larger
than those of the genus Phyllophaga. A few species are entirely brown while the
remainder are brown- or white-striped. They are distinguished further by their
massive antennal clubs, which consist of six or seven extremely long, thin, flat,
parallel, leaflike plates.

The larvae of Polyphylla variolosa Hentz have caused heavy losses in coniferous
nurseries in the Northeast. The larvae of P. occidentalis (L.) have been observed
feeding on the roots of pine seedlings, but they prefer the roots of grasses. They
would appear to be potentially harmful in nurseries and plantations. P. hammondi
LeConte occurs in the western part of the Central States and has the interesting
habit of depositing its eggs in rotten wood.

The genus Serica contains a number of species, the adults of which closely
resemble those of the genus Phyllophaga except for their much smaller size and
their regularly spaced, elytral striae. They are usually less than one-fourth as large
as May beetles. The adults are sometimes abundant enough to cause noticeable
defoliation in hardwood stands. The larvae have also been known to cause damage
in heavily infested nurseries, but they are usually not very destructive. Adults
emerge from mid-May to mid-August but are usually most abundant in June. The
life cycle requires 2 to 3 years. Common species include S. sericae (Iliger), which
is often abundant; S. tristis LeConte, which has been known to defoliate spruce in
plantations in southern Canada; and S. vespertina (Gyllenhal) and S. intermixta
Blatchley.

The genus Dichelonyx is represented in eastern North America by many species.
The beetles are small, about 6 to 12 mm long, and are often brightly colored. A
distinguishing characteristic is the presence of two spurs on each middle and hind

269

tibia. The larvae are grubs, which never exceed 18 or 20 mm in length. Adults are
most common in June and July, and the life cycle requires 2 to 3 years (1056).

Dichelonyx albicollis (Burmeister) is a well-known species. It has been recorded
from New Jersey, Michigan, and Ontario and feeds during both the day and night on
the needles of pine, especially jack pine. The adult is greenish, shiny, and about 12
mm long. D. elongata (F.) occurs from New England and New Jersey to Oklahoma
and Kansas. The adults feed at night on the leaves of various hardwoods, especially
sweet birch and alder. The adult is smaller and somewhat darker than the adult of D.
albicollis. A third species, D. subvittata LeConte, has been recorded feeding on
oak, hazel, and pine from New England to the Lake States and in southern Canada.

The genus Diplotaxis contains a number of species that feed mostly on conifers,
especially pines. The beetles are usually brown or reddish brown and, except for
having five visible ventral abdominal sternites, resemble members of the genus
Phyllophaga. The exoskeleton is also quite hard and rigid. D. sordida (Say) occurs
commonly on red and jack pines in the Lake States, and the larvae have caused
serious damage in nurseries and plantations in New York. The adult is slate colored
with yellowish hairs on the pronotum, and it is about 10 to 12 mm long. D. liberata
(Germar) also occurs commonly on pines in the Lake States. The adult is blackish,
hairless, and about 12 mm long.

The Asiatic garden beetle, Maladera castanea (Arrow), an introduced species
first recorded in North America in New Jersey in 1921, is now widely distributed in
the Eastern States south to South Carolina. Adults are usually cinnamon brown and
about 6 to 12 mm long. They fly at night and feed on more than 100 species of
plants, including forest and shade trees such as maple, willow, boxelder, buckeye,
and ailanthus. Young pines, hemlocks, and yews in nurseries are defoliated occa-
sionally and seriously injured. The roots of rhododendron and azalea are damaged
occasionally by the larvae.

The rose chafer, Macrodactylus subspinosus (F.) is widely distributed in the
Eastern United States. Adults are tan to reddish brown, densely covered with dull,
yellow scales or hairs, and have long reddish-brown legs. They skeletonize the
leaves of a wide variety of hosts, including many species of forest and shade trees
(fig. 116). In heavily infested areas, they appear in swarms in late May or early June
and feed first on the opening buds. Later, they attack the flowers, fruit, and foliage.
The larvae feed mostly on the roots of grasses but may also attack the roots of tree
seedlings.

The European chafer, Rhizotrogus (= Amphimallon) majalis (Razoumowsky),
an introduced species first recorded in New York in 1940, now occurs in several
Eastern States and southern Ontario (228). The adult is oval-shaped, light brown or
tan, and about 14 mm long. A distinguishing characteristic is its toothless, uncleft,
hind tarsal claw. During the peak of the flight season they are often seen swarming
around various trees and tall shrubs. The larvae feed on the roots of a wide variety
of plants, including the seedlings of such tree species as spruce and Douglas-fir
(1189).

The pine chafer, Anomala oblivia Horn, occurs from New York to the Lake
States and south to Georgia. Male beetles have the head and pronotum greenish
bronze and the elytra dark tan. They are about 6.5 mm long. Females are light tan
and about 9 mm long. They feed mostly on the new needles of various pines, eating
notches in them just above the sheath and causing the ends to die. Heavily infested
trees become brownish or scorched in appearance. Needle browning has occurred

270

Courtesy Conn. Agric. Exp. Stn.

Figure 116.—Birch leaf skeletonized by the rose
chafer, Macrodactylus subspinosus.

on several thousand hectares of young loblolly pines in southeastern North Car-
olina. The base of the damaged needle usually survives, however, and the needle
grows to about one-half its normal length. The winter is spent in the larval stage 6
to 10 cm deep in the soil. Pupation occurs in the spring, and the adults emerge in
June or earlier. Eggs are deposited in the soil near their pine hosts. Hatching occurs
in 10 to 15 days and the larvae feed on the roots of various plants, including trees,
until the onset of cold weather. There is one generation per year.

Infestations of the pine chafer tend to occur in open pine stands and plantations.
Numerous outbreaks have occurred in the Lake States and the South. The majority
were short-lived and limited in size, but some outbreaks were quite extensive and
lasted for several years. A. lucicola (F.) occurs from New England to the Lake
States and Kentucky. It severely damaged larch seedlings in a nursery in New York.

The Japanese beetle, Popillia japonica Newman, an introduced species first
recorded in North America near Riverton, N.J., in 1916, now occurs in all or parts
of at least 15 States from New Hampshire and Vermont to Georgia and Ohio (427).
Spot infestations have also been recorded in many other States and in Ontario and
Nova Scotia, Canada. The adult is broadly oval and nearly 12 mm long. The body
is a bright, metallic green; the legs, a darker green; and the elytra, a coppery brown.
There are two small tufts of white hairs just behind the wing covers and five patches
of white hairs on each side. The wing covers are shorter than the abdomen. Full-
grown larvae are about 25 mm long, typically grub-shaped, and have the last two
rows of spines on the underside of the last abdominal segment arranged in the shape
Oral Veue

The Japanese beetle feeds on the foliage, flowers, and fruits of a wide variety of
plants. It is present north of Florida, in most States east of the Mississippi River and

271

south of Wisconsin and Minnesota, and in California. Many species of forest and
shade trees are subject to defoliation, especially Japanese and Norway maples,
buckeye, sycamore, gray birch, walnut, Lombardy poplar, basswood, mountain-
ash, and American, English, and Siberian elms. Feeding is usually confined to
young, tender leaves. Damaged leaves may be skeletonized or they may also have
large, irrregular holes chewed out. In heavily infested areas, the trees may be
almost entirely defoliated. The larvae may also seriously damage the roots of
ornamental nursery stock.

The winter is spent in the larval stage in the soil. Pupation occurs in early spring,
and the adults emerge from late May to early July. They are gregarious and are often
found feeding in masses on certain plants, while nearby plants are uninfested.
Female beetles enter the soil to depths of 3 to 10 cm to deposit their eggs. Moist,
loamy soil covered with closely cropped grass is a favorite site. There may be two
generations per year in the southern parts of the species’ range. Farther north, the
life cycle may require 2 years.

Disease pathogens, especially the bacterium, Bacillus popilliae Dutky, often
destroy large numbers of the larvae. Several introduced parasites, Tiphia vernalis
Rohwer and 7. popilliavora Rohwer in particular, also exert a considerable degree
of control (427).

Cotalpa lanigera L., the goldsmith beetle, occurs throughout the Eastern United
States and feeds on the foliage of various hardwoods, such as aspen, oak, and
willow. The adult is broadly oval, convex, and from 20 to 26 mm long. It is a
brightly colored beetle; the elytron is lemon yellow, and the head and thorax
burnished golden. The venter is greenish to copper and covered with whitish wool.
The legs are reddish yellow. Full-grown larvae are about 43 mm long. The head is
tan and the underside of the last abdominal segment is thickly covered with hooked
spines. Adults emerge in late spring and the females deposit their eggs in the soil.
The larvae feed on the roots of various plants, probably including young conifers.
Adults have been observed fairly commonly in nurseries in the Lake States. The life
cycle requires from 2 to 3 years for completion.

The green June beetle, Cotinis nitida (L.), one of the most widely recognized
members of the family, occurs throughout much of the Eastern United States, most
commonly in the Atlantic and Gulf Coast States and in the Mississippi River Valley.
The adult is usually velvety green above, with the margins orange-yellow, and is
about 10 to 25 mm long. The undersurface is shining green and orange-yellow, and
the head is armed with a hornlike process. Full-grown larvae are up to 50 mm long
and have the interesting habit of crawling on their backs.

Adults are most numerous during June and July, and the females deposit their
eggs in soil rich with organic matter. The larvae feed on the organic matter during
the remainder of the season and then hibernate. In the spring, they move close to the
surface and feed on both dead vegetation and the roots of living plants. Lawns and
golf courses are often damaged severely. Seedlings in forest nurseries are some-
times injured.

The genus Dynastes, which contains the largest known beetles, is represented in
the Eastern United States by only one species, the eastern Hercules beetle, D.
tityus (L.). It has been recorded from New York, Indiana, and Arkansas south to the
Gulf of Mexico. D. tityus breeds in decayed hardwood stumps and logs, and is also
found occasionally in cavities in the bases of living trees (488). The adult is very
large, from 40 to 60 mm long, and is usually greenish gray or tan except for
mottlings or blotchlike areas of black. The male is armed with a large pronotal horn

oP

that projects forward and almost meets another horn that projects upward from the
front of the head. Females are somewhat smaller than the males and are without
horns. The habits of the adults are not well understood; however, they have been
observed feeding on sap oozing from wounds on ash trees.

The genus Xy/oryctes 1s represented in eastern forests by three or four species,
one of which, the rhinoceros beetle, X. jamaicensis (Drury), 1s common in
hardwood stands in the South. The adults are robust, shiny beetles, about 28 mm
long. They are dark chestnut to blackish brown above, and paler and thickly clothed
with reddish hairs below. The male has a single, large curved horn on top of the
head. In the female, a large tubercle replaces the horn. The larvae are usually found
in leaf mold on the forest floor. Adults are usually found in the vicinity of ash trees.

At least four species of the genus Pelidnota occur in eastern forests, one of
which, P. punctata (L.), 1s fairly well known. The larvae are usually found in
decayed hardwood stumps but sometimes in decaying roots and logs. The adult is
about 20 mm long and reddish brown above, with three black spots on each wing
cover and one on each side of the pronotum. The base of the head, the scutellum,
and the entire underside of the body are deep bronzed green. The species is not
injurious.

The genus Parastasia is represented in eastern forests by several species. The
most common and widely distributed one is P. brevipes LeConte. The larvae are
found most commonly in decayed hardwood stumps, roots, and logs. Adults are
blunt, convex, dark chestnut-brown, and about 16 mm long. The species is not
considered to be injurious.

The genus Osmoderma is represented in eastern forests by at least four species,
two of which, O. eremicola Knoch and O. scabra (Beauvais), are fairly common.
The larvae feed in decayed cavities of dead or dying trees or logs. The adults are
broadly oval and depressed dorsally and have heavy leathery elytra; O. eremicola is
dark chestnut-brown, smooth and shiny, whereas O. scabra is bronzy purple-black
and rough in texture. They are approximately 25 mm long. When handled, they
emit a strong leatherlike odor.

The genus Trichiotinus contains several species, the adults of which are fre-
quently seen around flowering trees and shrubs, cherry in particular. They are
variegated, the body is densely pubescent, the elytra are almost as wide as long, the
legs are long and slender, and they range in length from 9 to 15 mm. The larval

stage is spent in old logs and stumps.
Family Elateridae
Click Beetles, Wireworms

Click beetles are so-called because of the presence of a spine on the prothorax
that snaps into a groove on the mesosternum with an audible click. This mechanism
enables an adult lying on its back to throw itself into the air and land on its feet. The
larvae are known as wireworms because of their long, narrow, fusiform, tough-
skinned bodies. Forest-inhabiting species are predominantly phytophagous, but a
few are predacious on other insects, such as wood borers and sawflies in cocoons in
the soil. Species that feed on forest vegetation usually confine their attacks to dead
and often well-decayed wood and are of little economic importance. The wire-
worms of New York State have been reported (303).

The eyed click beetle, Alaus oculatus (L.), is a voracious feeder on various
species of borers in hardwoods and the related species, A. myops (F.), on borers in
pines. Adults of A. oculatus are grayish black, with two large eyelike spots on the
prothorax, and are 25 to 50 mm long. Adults of A. myops are more slender and
darker, and are only about 20 to 40 mm long. They also have eye spots.

2S

Other predacious species include Lacon discoidea (Weber) and L. avita Say that
are found in pines, and Hemicrepidus bilobatus (Say) that occurs in hickory.
Crenicera triundulata (Randall) and C. nitidula (LeConte) have been observed
feeding on cocoons of the European spruce sawfly in Canada (879), and an
undetermined species of Crenicera was found feeding on the larch sawfly in

Minnesota (332).
Family Cleridae
Checkered Beetles

This is one of the most important families of insect predators attacking injurious
forest insects. The adults are active, antlike, brightly colored, hairy beetles about 3
to 13 mm long. They feed on adult beetles. The larvae live in the galleries and
tunnels of bark beetles and wood borers and destroy the immature stages of these
insects.

Adults are distinguished by their I 1-jointed and generally serrate antennae, the
outer joints of which are longer and form open or compact clubs. The tarsi are five-
jointed and the first four joints bear membranous appendages. Larvae are soft-
bodied, elongate and parallel-sided, frequently highly colored though often white
and thin-textured, and are from 9 to 13 mm long.

Most species spend the winter in the larval stage. Others overwinter as pupae or
as adults in pupal cells in the bark. The larvae travel down the tunnels of their hosts
eating one larva after another. Some are capable of consuming several times their
own weight of these larvae (/4/). A few of the more important predators of forest
insects are discussed below.

Chariessa pilosa (Forster) is one of the most common species in eastern forests.
Its known hosts include several species of borers in hardwoods, and the smaller
European elm bark beetle. The adult (fig. 117) is a wedge-shaped, flattened beetle
about 6 to 13 mm long. The thorax is red with two black stripes and the wing covers
are black with dense, fine punctures. The larva is fairly robust, widest at the
middle, and of a bluish tinge. Adults are often observed feeding on insects attracted
to freshly cut logs during the summer.

F-519949
Figure 117.—Adult of Chariessa pilosa,
a predator of wood-boring larvae.

274

Thanasimus dubius (F.) is one of the most important predators of the destructive
bark beetles in the Eastern United States (95, 445, 568). The adult is a brightly
colored, hairy beetle about 7 to 10 mm long. The head, thorax, and base of the
wing covers are dull red; the antennae and legs are red to pitch black; and the wing
covers are mostly black with crossbands of whitish hairs. The larva is elongate,
fusiform, and purplish with brown sclerotized areas.

Winter is spent in the larval, pupal, or adult stages. In early spring, the adults
emerge and fly to beetle-infested trees or logs and feed on bark beetles as they
emerge from hibernation. Eggs are deposited in entrances to bark beetle galleries.
Young larvae feed on bark beetle eggs; older ones feed on beetle larvae, pupae, and
adults. Pupation occurs in cells in the outer bark (590).

Enoclerus nigripes (Say) larvae feed on bark beetles in conifers, and on wood
borers in hardwoods. Adults are brightly colored and about 8 to 12 mm long. The
head, thorax, base of the wing covers, and the undersides are dull red; the
remainder is black except for two yellowish crossbars on the wing covers. The
larvae are similar to those of 7. dubius. The blackbellied clerid, EF. lecontei
(Wolcott), has many hosts including bark beetles and weevils in pine, spruce, and
juniper. It also feeds on bark beetles, weevils, and small borers in hardwoods.

Monophylla terminata (Say) feeds on borers and bark beetles in hardwoods.
There also are reports of its feeding on white pine weevil larvae in white pine.
Adults are about 4 to 8.5 mm long. The eyes are deeply notched in front, the last
joint of the antenna 1s as large or larger than all of the others combined, the thorax is
yellow with a black disk, and the sides of the wing covers are yellow. The larvae are
white, soft-textured, and bear two well-separated hooks on the ninth abdominal
segment.

Cymatodera bicolor (Say) 1s an important enemy of roundheaded and flatheaded
borers in hardwoods in the Eastern United States. Adults are about 5 to 10 mm
long. The color is dull blackish except for the legs, thorax, and basal joints of the
antennae which are reddish yellow mixed with black. The larvae are purplish.

Tarsostenus univittatus (Rossi) is an important predator of powderpost beetles
and other borers in dry, seasoned wood. The adult is small, slender, and shiny black
except for a white mark across the middle of the elytra. The larva is very small and
a light violet, except for brown or yellow markings. There are two recurved hooks
on the ninth abdominal segment.

Neichnea laticornis (Say) feeds on various species of bark beetles in the Middle
Atlantic States. The adults are small, slender, and black except for a spot on the
head and the sides of the thorax, which are golden yellow.

Many other clerids are also predacious on various forest insects in the Eastern
States. Priocera castenae Newman feeds on bark beetles in conifers; Phlogistoster-
nus dislocatus (Say) and Orthopleura damicornis (F.) feed on the larvae of borers
and bark beetles in the twigs of hardwoods; Cregya oculata (Say) feeds on larvae of
borers. and bark beetles in both hardwoods and conifers. Descriptions of clerids
occurring in Ohio are available (683).

Family Dermestidae
Dermestid Beetles

Dermestid beetles are compact, oval to convex, and usually extremely hairy.
Some are spotted with gray, brown, or orange hairs which rub off easily. The larvae

are cylindrical and covered with long hairs. Some larvae are soft; others are hard-
shelled.

215

A few species are found under the bark of trees where they feed on dead insects.
Some are troublesome pests in collections of insects or stuffed animals, on which
they feed. The hide beetle, Dermestes maculatus De Geer, and the larder beetle,
D. lardarius L., have damaged cargoes of lumber in ship holds where hides were
stored previously. In efforts to construct pupal chambers, D. maculatus has also
been known to damage seriously the surface of lumber in warehouses. The majority

of dermestids feed on skins, dried meats, furs, and carpets.
Family Buprestidae
Flatheaded Borers

The flatheaded borers include several destructive pests of forest and shade trees.
More than 150 species and varieties have been recorded east of the Mississippi
River (443). The larvae of all species are borers, and they feed in ail parts of the
tree. Some mine the leaves, and some construct tunnels in the inner bark and outer
wood of the trunk, branches, and roots. The majority of species, however, excavate
winding tunnels through sound and decaying sapwood. Many bark-boring species
are capable of girdling and killing both healthy and injured trees. Wood-boring
species are often highly destructive of recently felled saw logs, often seriously
reducing or destroying their usefulness as lumber.

Buprestid beetles are usually somewhat flattened or oval-shaped and are beau-
tifully marked or metallic colored. The head is strongly deflexed and is inserted
into the prothorax to the eyes. The antennae are serrate, | 1-jointed, and inserted on
the front; the prosternum is prolonged behind and fits into the mesosternum; the
elytra usually cover the abdomen; and the first two of the abdominal sternites are
fused.

Buprestid larvae are distinguished primarily by well-developed ambulatory plates
on the upper and lower surfaces of the first segment behind the head, by the
presence of a central line, groove, or V on the upper plate, and by the absence of
legs. The larvae of all bark- and wood-boring species are typically “‘flatheaded,”’ a
condition caused by the greatly enlarged first and sometimes second and third
thoracic segments (fig. 118). Leafmining larvae are flattened, rather oval-shaped,
deeply notched at the sides, and gradually taper toward the rear. The true head in all
larvae is comparatively small, more or less retracted into the first thoracic segment,
and scarcely visible.

eS

Courtesy R. T. Frankiin, Univ. Ga.

Figure 118.—Typical larva of a buprestid or flatheaded
borer.

Bark- and wood-boring buprestids deposit their eggs singly or in masses either
on the bark, in crevices in the bark or wood, or under the bark at the edges of
wounds. Weakened, injured, dead, or dying trees and stumps are usually attacked.
Occasionally, green trees are also infested. The larvae feed either under the bark, in
the sapwood or heartwood, or in two or more of these places. Their mines are

VALS

winding and usually oval in cross section. Eventually, they terminate in elongated
pupal cells that are connected to the surface by short, oval exit holes. Charac-
teristically, the mines are usually packed tightly with layers of sawdustlike borings
and pellets and their walls are scarred with fine, transverse lines. Many wood-
boring species spend the winter as adults in pupal cells. A few overwinter in the
larval stage. The life cycle usually requires | to 2 years, but in certain species it
takes many years.

Acmaeodera pulchella (Herbst), the flatheaded baldcypress sapwood borer,
breeds in baldcypress in Eastern and Southern States. The adult is blue-black to
blackish and about 6 to 10 mm long. In some individuals the thorax is dull bronze
and the wing covers and outer angles of the thorax are marked with spots and
patches of waxy yellow. Full-grown larvae are about 13 mm long and the pro-
thoracic plates are marked by brownish median grooves or lines. The sapwood of
dead and dying baldcypress trees and recently cut baldcypress logs is subject to
severe attack and damage. The removal of unseasoned logs from the woods before
the adults fly in the spring, or girdling trees in the fall that are to be felled in the
spring, should aid in the reduction of damage.

The twolined chestnut borer, Agrilus bilineatus (Weber), occurs in southern
Canada and throughout the Eastern United States. It breeds in various hardwoods,
preferably chestnut and several of the oaks (348). Trees weakened by drought or
defoliation, and trees with low root starch (/244) are usually attacked. Adults are 6
to 12 mm long, subcylindrical, and black with a more or less greenish tinge. The
sides of the thorax and elytra are clothed with light golden-yellow pubescence, and
each elytron is marked with a stripe of the same color. The larva is slender,
considerably flattened, about 25 mm long, and has two spines at the posterior end.

Winter is spent in the final (fourth) instar in pupal cells constructed in the outer
layers of the sapwood and sometimes in the bark (249). In the spring, the adults
emerge through characteristic D-shaped holes in the bark. Eggs are deposited on
the bark in late spring or early summer. Young larvae bore directly through the bark
to the phloem. There they excavate winding mines in the inner bark and outer wood
of the main trunk and larger branches. These mines run back and forth in all
directions (fig. 119), and in the event of heavy attack, they girdle and kill the trees.
Attacks usually begin in the tops of trees and are extended downward as the trees
continue to weaken (5/0).

Control under forest conditions is usually impractical. A chalcid wasp, Pha-
sgonophora sulcata Westwood, is the beetle’s primary parasite, accounting for 10
percent of the host in Wisconsin (5//). Except for management practices that
maintain or promote tree vigor, such as watering and fertilizing valuable shade
trees, there is little that can be done to protect trees from attack.

The bronze birch borer, A. anxius Gory, apparently occurs throughout most of
the range of birch in Canada and the United States. Various birches, especially
paper and yellow, are preferred hosts. The adult is deep green-bronze and about the
Same size and shape as the adult of the twolined chestnut borer. There are coppery
reflections on the front of the pronotum; the front of the head is greenish in the male
and copper-bronze in the female. Full-grown larvae are slender, flattened, about 25
mm long, and have two spines at the posterior end.

Adults begin to emerge in late May or early June and continue until August,
depending on locality, and they feed on leaves for about 3 weeks before egg laying
begins. Eggs are deposited singly or in small groups beneath loose curls of bark and
in cracks and crevices in the bark, mostly on unshaded parts of the tree. Young

PAT

Courtesy Pa. Bur. For., For. Pest
Manage., Middletown

Figure 119.—Galleries of the twolined chestnut borer,
Agrilus bilineatus.

larvae bore directly through the bark to the cambium area. There they excavate
galleries between the bark and wood, with occasional side trips into the xylem to
molt and then to spend the winter. The galleries wind back and forth, usually across
the grain of the wood. Mature larvae construct oblong cells in the wood or thick
bark in which they spend the winter and pupate in the spring. Larvae of all sizes and
ages have been found in infested trees during the winter, but the only ones capable
of developing to adults were those which became mature before winter set in and
were later subjected to subfreezing temperatures. In the North, 2 years are required
to complete the life cycle; in the South there 1s one generation per year. The biology
of the bronze birch borer in New Brunswick has been reported (62).

The bronze birch borer prefers weakened or injured trees (765). Damage may be
extremely severe in stands of such trees. Weakened residual trees following log-
ging, and individual shade and ornamental trees weakened by drought or other
factors, are often seriously injured. Enormous volumes of birch suffering from
dieback have been killed in the Northeastern States and Canada.

278

The bronze poplar borer, A. liragus Barter & Brown, breeds in poplar in
Canada and southward in the United States to Pennsylvania and Arizona. Adults are
blackish with deep-green reflections and are about 7 to 12 mm long. Overmature or
defective trees and young trees suddenly released by the removal of dominant trees
are most often attacked; however, felled, topped, or girdled trees and trees damaged
by Saperda calcarata Say and infected by the fungus Hypoxylon mammatum
(Wahl) Miller are also attractive. Infestations in standing trees usually begin in the
crown and move downward.

Adults are present during a period of several weeks in the summer and feed on the
leaves of poplar. Eggs are deposited in bark crevices and the larvae feed in the
cambial region, excavating long tunnels that zigzag back and forth in a compact
manner. The larvae feed for two seasons in Ontario before becoming mature (63).
Woodpeckers took heavy tolls in mature larvae, pupae, and, presumably, adults in
standing trees in Ontario.

Agrilus horni Kerremans occurs in Ontario and the northern tier of Eastern
States, and breeds in young, apparently healthy aspen suckers, especially quaking
aspen, some of which it girdles and kills. Adults are almost identical to those of the
bronze birch borer except for shorter ovipositors in the females.

Eggs are deposited on the smooth bark at the base of aspen suckers. The larvae
bore into the bark and tunnel downward and out along large roots, gradually
working their way through the bark to the cambium region. Once here, they turn
around and tunnel back toward the main stem, making spiral galleries that encircle
the roots and which may be continued up the trunk for 6 to 10 cm. Pupation takes
place in a cell in the center of the stem, and the life cycle requires 2 years. Damage
appears to be most severe in sparsely stocked stands or in stands growing on poor
sites (937).

Agrilus arcuatus torquatus LeConte, the hickory spiral borer, has been re-
corded from New York, Ohio, North Carolina, and Mississippi, apparently favor-
ing hickory and pecan, but also infesting many other hardwoods. It, or a closely
related species, has been observed attacking young oaks and other hardwoods in the
Piedmont of the South (74). Adults are about 8 mm long. Males have a greenish-
bronze head and thorax, purplish-black elytra, and brassy underparts. Females are
completely bronze.

Adults appear from May to July and feed on the leaves of their host, making large
irregular holes. Eggs are deposited singly on the bark surface or on terminals or
twigs, usually near the base of a small shoot of the current season’s growth. The
larvae feed downward beneath the bark during the summer, and during the fall they
sever the wood by constructing a spiral burrow. The following spring, they continue
to feed beneath the bark, constructing long, irregular tunnels that deeply engrave
the wood. When they are full grown, they make a second transverse spiral cut
around the wood, working first toward the pith and then out again to the phloem,
leaving the bark intact. Then, they mine upward in the phloem for 2.5 to 5 cm and
construct pupal cells entirely within the pith. Two years are required to complete the
life cycle (159). This species is often very destructive of hickory seedlings in the
South.

Agrilus acutipennis Mannerheim, the spotworm borer, has been observed
attacking overcup oak in Louisiana and Arkansas (876). Eggs are deposited on the
bark. The larvae bore through the bark and then excavate patches of inner bark up to
12 mm in diameter. Later, they enter the wood and tunnel spirally upward in the
outermost growth ring. Pupation occurs within the tunnel and the life cycle requires
2 years (1133).

279

Damage by A. acutipennis results in a defect known as “grease spot.”’ This is
caused by a fungus that spreads through the wood from the tunnels. In cross
section, grease spots are oval to diamond- or spindle-shaped and about 19 mm
wide. Their presence in lumber greatly reduces its value. Infestations have been
found frequently in river bottoms where the trees were subject to backwater
flooding during the winter and spring. In such situations, entire stands of trees over
12 mm in diameter are usually infested.

Many other species of Agrilus also occur in eastern forests. Some of these and
their more important hosts are as follows: A. juglandis Knull—butternut: A.
difficilis Gory—honeylocust; A. /econtei Saunders and A. ce/ti Knull—hackberry;
A. betulae Fisher—river birch; A. cephalicus LeConte—dogwood; A. fuscipennis
Gory—persimmon; A. egenus Gory—black locust; A. otiosus Say—hickory; and
A. masculinus Horn—maple (423).

The genus Buprestis is represented by a fairly large number of wood-boring
species. Many seem to prefer dead and decayed wood; others are found in either
weakened or perfectly healthy trees. The larvae construct tunnels in the sapwood
and often the heartwood and frequently cause serious damage. The adults come in
many different colors: metallic green, blue, gold, red, yellow, or orange. Often,
there are many color variations and patterns within a species (555).

The turpentine borer, B. apricans Herbst, long considered the most important
eastern member of the genus, occurs throughout the southern coastal regions from
North Carolina to Texas, and breeds in longleaf, slash, loblolly, shortleaf, and pitch
pines. The adult (fig. 120) is grayish bronze with a greenish, metallic luster and is
about 25 mm long. It is elliptical, somewhat flattened, and each elytron bears eight
rows of large punctures. Full-grown larvae have the prothoracic plates roughened
and marked above by a dark-brown Y, and they are up to 40 mm long.

Courtesy Fla. Dep. Agric. & Consum. Serv.,
Div. Plant Ind., photo by V. Jane Windsor

Figure 120.—Adult of the turpentine borer,
Buprestis apricans.

280

Turpentine borer beetles emerge in February or March and feed for a short time
on the needles in the tops of their hosts. Eggs are deposited in exposed wood
containing season checks, especially at the edges of turpentine faces and on fire-
scarred surfaces. The larvae tunnel in the sapwood and heartwood, construct long,
narrowly oval, tortuous mines, and fill them with solidly packed, fine, granular,
pitchy frass. At maturity, they form cells in which to pupate near the surface. There
the adult spends the winter. About 3 years are spent in the larval stage.

The turpentine borer used to be the most destructive insect in the turpentine
orchards of the South (72, 74). Borer-riddled trees were weakened so severely they
became subject to windbreakage. The lumber value of such trees was virtually
destroyed, and gum production was seriously reduced. Attacks can be prevented or
reduced by preventing the exposure of dead, dry wood to fire, logging, or other
forest operations. Acid treatment to increase gum flow in naval stores operations
has virtually eliminated dry faces, thereby greatly reducing the damage caused by
the species.

A number of other species of Buprestis are also encountered in eastern forests. A
few of the more common ones and their hosts are as follows: B. striata (F.)—adult
brown with greenish reflections; it breeds in dead branches of pine, hemlock, and
baldcypress. B. lineata (F.)—adult medium-size and dark, usually with brick-red to
yellow longitudinal markings on the elytra; it attacks longleaf, loblolly, pitch, and
Virginia pines. B. rufipes (Olivier)—adult slender and dark green, with a long,
yellow basal patch and two crossbars of yellow on the elytra; infestations occur
around scars on elm, beech, hickory, oak, maple, yellow-poplar, and tupelo. B.
salisburyensis (Herbst)—adult short, oval, and green with a tooth on the inner
margin of each elytron; it breeds in pitch pine. B. maculipennis Gory—adult rather
small and blackish, with a brassy tinge and scattered yellow spots or patches on the
elytra; it attacks pine and hemlock.

The genus Chrysobothris contains some of the most common and injurious
members of the family. The larvae are all borers in the wood of both deciduous and
coniferous trees. All parts of the tree are attacked, from the roots to the twigs in the
crown, but the majority of attacks occur on the main trunk. The adults are small to
medium size and are usually not conspicuously colored. The pronotum is usually
wider than long and the scutellum is small and triangular. The elytra are rounded or
angulate at the base and they strongly converge posteriorly. The legs are robust,
with the femora swollen at the middle. There usually is a large tooth on each front
leg. Several species are very injurious, especially to young trees weakened by
drought, defoliation, or other adverse factors. The genus has been revised (424).

The flatheaded appletree borer, C. femorata (Olivier), one of the commonest
and best known of the flatheaded borers in North America, occurs throughout most
of Canada and the United States. It attacks a wide variety of deciduous trees such as
sycamore, silver maple, boxelder, black walnut, willow, white and black oaks,
yellow-poplar, elm, beech, hickory, hackberry, apple, and pear. The adult is oval,
flattened, dark green-bronze above, bright brassy beneath, and about 7 to 16 mm
long. The elytra are marked with two wavy, depressed, light bands. Full-grown
larvae are about 25 mm long.

Adults appear throughout the summer and feed on the foliage of their hosts,
occasionally causing serious defoliation. Eggs are deposited under bark scales or in
bark crevices on the main trunk or larger branches. The larvae bore into the bark
and feed in the phloem and outer sapwood. Their tunnels are sometimes 8 cm or
more long, especially in young trees. Tunnels in older trees are confined mostly to

281

the thick inner bark. Mature larvae construct cells in the outer wood during late
summer where they spend the winter and pupate the following spring. There is one
generation per year (402).

The flatheaded appletree borer is especially destructive to newly planted trees
and trees weakened by drought, defoliation, or other adverse factors. Young trees
are often girdled and killed; larger trees are often seriously injured through the loss
of large patches of bark over mined areas. Maintenance of tree vigor, wrapping the
trunk with high-grade wrapping paper or burlap when trees are planted or pruned,
and shading the south side of newly planted trees are recommended control
practices.

The Australianpine borer, C. tranquebarica (Gmelin), also commonly known
as the mangrove borer, occurs in southern Florida and breeds in living mangrove
and horsetail casuarina trees. The adult is greenish bronze and about 12 to 17 mm
long. Eggs are deposited in the spring under roughened areas of bark. The larvae
feed beneath the bark until nearly full grown, and then enter the wood to construct
pupal cells. Damage to ornamental or windbreak trees is often severe. The removal
and destruction of infested wood during fall and winter is helpful in control.

Chrysobothris orono Frost attacks living red and jack pines in the Lake States.
Eggs are laid singly on the trunk. The larvae feed in the bark during the first three
instars, excavating cells and causing pitch to coagulate into large masses (fig. 121).
Older jiarvae feed in the wood. The winter is spent in the larval stage. In the spring
of the third year, the larva constructs an L-shaped cell, plugged with frass and wood
chips, where pupation occurs. This species is seldom injurious, although its injury
may remain in the bole as a defect after the wounds have healed.

Several other species of Chrysobothris also occur fairly commonly in eastern
forests. Some of these and some of their more important hosts are as follows: C.

F-519916

Figure 121.—Pitch mass on red pine caused by feeding
of the larvae of Chrysobothris orono.

282

pusilla Castelnau & Gory—pitch, shortleaf, and eastern white pines, white spruce,
and hemlock; C. dentipes (Germar)—eastern white, shortleaf, longleaf, and Vir-
ginia pines, and larch (this species is reported to be strongly attracted to sawmills);
C. floricola Gory—probably all species of pines; C. scabripennis Castelnau &
Gory—pine, spruce, hemlock, and balsam fir; C. trinervia (Kirby)—white pine
and spruce; C. harrisi (Hentz)— Virginia, eastern white, and pitch pines; C. azurea
LeConte—white oak, dogwood, maple, basswood, birch, and willow; C. sexsig-
nata (Say)—ash, red maple, walnut, hickory, beech, yellow birch, white oak,
hemlock, baldcypress, and pitch pine; C. texana LeConte—redcedar; C. adelpha
Gemminger & Harold—hickory and pecan; C. viridiceps Melsheimer—red maple,
red oaks, and cherry; C. blanchardi Horn—eastern white, Virginia, and pitch
pines, and larch; and C. neopusilla Fisher—balsam fir.

Chalcophora virginiensis (Drury), the large flatheaded pine heartwood borer,
breeds in injured, dying, and dead pines and in pine stumps throughout the Eastern
United States. Adults (fig. 122) are dull-black or dark-bronze beetles about 23 to 33
mm long. The thorax is broader than long, and the elytra are marked with dark or
shiny elevations and rough, grayish or brassy depressions. Full-grown larvae are up
to 50 mm long and the dorsal thoracic plate is marked with a Y. Eggs are deposited
around scars on living areas and in bark crevices or holes in the bark of logs and
stumps. Living trees may be severely damaged by larval tunnels in the wood. Pine
logs left too long in the woods are also subject to severe damage. C. liberta
(Germar) and C. georgiana LeConte are often found in association with C.
virginiensis. Adults of C. liberta are copper or brass colored, while those of C.
georgiana are golden bronze.

Chalcophorella campestris (Say), the flatheaded sycamore-heartwood borer,
breeds in injured, dying, or dead sycamore, beech, oak, maple, yellow-poplar, and

~~

F-532843

Figure 122.—Adult of Chalcophora virginiensis, the
large flatheaded pine heartwood borer.

283

basswood, preferably in moist, rotting logs. The larvae resemble those of Chal-
cophora virginiensis except that they are longer, and the dorsal thoracic plate is
marked with a V or U instead of a Y. This species often attacks at ax blazes and
wounds and frequently riddles the wood beneath with its tunnels.

The genus Dicerca contains many species that breed in dead and dying trees and
logs. The adults are dark-gray to brown, medium-size, metallic beetles. The head is
flat, the pronotum is wider than long and grooved or ridged down the middle, the
scutellum is very small, the hind coxae are strongly dilated, and the elytra are
extended into taillike appendages (9/5). A few of the more common eastern species
and some of their hosts are as follows: D. divaricata (Say), the divergent beech
beetle— various dead, dying, or injured hardwoods; D. obscura (F.)—reared from
persimmon and sumac, collected on oak and hickory; D. punctulata (Sch6nherr)—
various pines (often common around sawmills); D. /urida (F.)—hickory, American
hornbeam, basswood, willow, and alder; D. tenebrosa (Kirby), the gloomy
borer—pines and spruce; and D. tenebrica (Kirby)—poplar.

The genus Melanophila consists of small- to medium-size beetles, the larvae of
which bore in the inner bark and outer wood of their hosts. The majority of species
attack very slow-growing, dying, or recently felled trees and are of no economic
importance. In the West, certain species are strongly attracted to fires and are
known as “fire bugs.”’ A key to the North American species has been published
(1092).

The hemlock borer, M. fulvoguttata (Harris), occurs throughout the Eastern
United States and eastern Canada. Hemlock appears to be the preferred host, but it
also occasionally attacks several other conifers such as eastern white pine, larch,
balsam fir, and red, white, and black spruces. The adult is black with a metallic
sheen and about 10 mm long. Each elytron usually bears three orange or yellow
spots of equal size.

Adults appear from late spring to late summer and deposit their eggs in groups
deep in bark crevices on weakened, dead, and dying trees or on logs and wind-
thrown trees in which the cambium is still moist. The larvae bore into the inner
phloem and, if conditions are not favorable for their development, they remain there
until they die. Under more favorable growth conditions they penetrate to the
cambium region and construct tortuous, frass-filled galleries (fig. 123). Before
becoming full grown, they construct cells in the outer bark in which they spend the
winter. The life cycle may be completed in | year in dead trees and logs. In living
trees, several years may be required.

Management practices designed to promote rapid growth and good health of
hemlock trees should be helpful in preventing damage by the hemlock borer. A
study indicates that hemlock borer attacks are not successful until 60 percent of the
roots are dead (9/6). Rapid salvage or cutting heavily infested trees is helpful in
preventing population buildup in stands weakened by windthrow or defoliation
(764).

Several other species of Melanophila also occur in eastern forests. A few
common ones and their hosts are as follows: The flatheaded fir borer, M.
drummondi (Kirby)—fir, larch, spruce, and hemlock; M. notata (Castelnau)—
pines; M. acuminata (De Geer)—spruce, fir, pine, and northern white-cedar
(adults are common around forest fires and scorched timber); and M. aeneola
Melsheimer—pines.

The genus Brachys contains many species of leafminers. The larvae differ from
the larvae of wood- or bark-boring species in having the prothorax only slightly

284

F-514869

Figure 123.—Mature larvae and galleries of the hemlock
borer, Melanophila fulvoguttata, on the surface of the
sapwood.

broader, if any, than the first abdominal segment. The adults are small and oval, and
they feed on the leaves of hardwoods, sometimes riddling them with holes. The
larvae mine the tissues of the leaves. B. tesselatus F. is very common on scrub oak
in the sandhills of the southeastern Coastal Plain. Heavy annual defoliation is not
unusual (/2/2). B. ovatus (Weber) also mines the leaves of oak. B. aeruginosus
Gory mines the leaves of elms.

Numerous other species of bark- or wood-boring buprestids are also encountered
in eastern trees. Ptosima gibbicollis (Say) breeds in living redbuds. The adult is
dark blue, about 6 mm long, and has two yellow spots on each wing cover. Damage
is sometimes severe. Trachykele lecontei (Gory) breeds in dead baldcypress in
Southern States. The adult is dark, ashy bronze, with black, velvety spots, and is
about 9 mm long. The larvae feed in both the sapwood and heartwood, often
causing a serious degrade of lumber. Actenodes acornis (Say) breeds in the dry
heartwood of red maple, birch, beech, oak, and hickory; Poecilonota cyanipes
(Say) breeds beneath the bark at wounds on living poplars; P. thureura (Say) breeds
beneath the bark at wounds on willow; Agrilaxia flavimana (Gory) breeds in the
small branches of white oak; and Cinyra gracilipes (Melsheimer) breeds in the dead

branches of oak and eastern hophornbeam.
Family Cerambycidae
Longhorned Beetles or Roundheaded Borers

The family Cerambycidae is one of the largest and most important of the families
of wood-boring beetles. More than 1,400 species have been recorded from the
United States, about 450 of which occur east of the Mississippi River (373). A list
of 262 species either known or believed to occur in Ohio alone was published (682);

285

the species occurring in Georgia have been discussed (394). The majority of species
breed in the dead wood of trees and shrubs but many, including some of the most
destructive ones, attack either slightly weakened or healthy trees and other plants.
Some species are vectors of the pine wood nematode, Bursaphelenchus xyloplilus
(Steiner & Buhrer) Nickle.

The larvae of all but a few members of the family live as borers in the tissues of
trees and other woody plants. Almost no part of a tree of any age or size is immune
to infestation by some species. The twigs, branches, and stems of sprouts and
seedlings and the twigs of branches of mature trees are girdled and severed. The
sapwood and the heartwood of large limbs and trunks of living trees are often
riddled and weakened, leading to windbreakage or death. Species that feed under
the bark of living trees may weaken and kill their hosts, or cause defects and stains
which seriously degrade lumber value. Species that attack recently felled trees,
logs, or seasoned timber also cause heavy losses. Not all species are harmful,
however; many of those that attack slash, stumps, and dead and dying trees are
actually beneficial because they aid in the quick removal of such waste material
from the forest floor and aid in its incorporation into humus.

Cerambycid beetles are distinguished by their oblong, often cylindrical bodies,
their long, usually 11-segmented antennae, and their long legs. The tarsi are five-
segmented, with the fourth segment small and partly concealed by the bilobed third
segment. The beetles move rapidly when disturbed and are strong fliers. Some
species make squeaking noises when captured. .

Cerambycid larvae (fig. 124) are distinguished by a few prominent characters.
They are always fleshy, thin-skinned, white or yellowish, and more or less cylin-
drical or depressed. In some species, the body tapers somewhat posteriorly, but the
anterior segments are never suddenly and conspicuously larger than the following
ones. The body is never curved and there are no prolegs or gripping processes on
the last abdominal segment. There are two overlapping, circular bands of skin
between each pair of body segments, the ventral mouth parts are always about on a
line with the base of the mandibles, and the tenth abdominal segment is modified
into two or three small retractile lobes. These borers are the subject of numerous
publications (204, 257, 259, 742, 743, 744, 745, 746, 747, 748).

F-531252
Figure 124.—Typical larva of a roundheaded wood borer.

The locust borer, Megacyllene robiniae (Forster), one of the most important of
the cerambycids, occurs in eastern Canada and throughout most of the United
States wherever its host, black locust, grows. The adult ranges from 11.5 to 28 mm
long. The jet-black background is marked with bright-yellow bands extending

286

across the thorax and wing covers, the third band on the wings being W-shaped.
The legs and antennae are moderately long and yellow. Full-grown larvae are robust
and about 25 mm long.

Adults are present in late summer or early fall, but are most abundant during
September. They are commonly seen feeding on the pollen of goldenrod blossoms
during morning hours. Later in the day, sometimes well after sunset, they are
usually seen running up and down the trunks of black locust trees in search of
oviposition sites. Eggs are usually deposited in rough bark crevices and around
wounds on the trunks of living trees. Newly hatched larvae bore into the inner bark
and construct small hibernation cells in which they spend the winter. Activity 1s
resumed in the spring when the leaf buds begin to swell. At this time, oozing sap
may be seen around larval entry holes in the trunk. The larvae soon bore into the
wood where they continue to feed until mature, around mid-July. During this period
they construct extensive tunnels throughout the heartwood (fig. 125A). As the larva
grows, it enlarges its tunnel to the exterior, through which it pushes its granular
frass to the outside and through which the adult eventually emerges. There is one
generation per year (464, 1352).

The locust borer has destroyed thousands of hectares of natural regeneration and
plantations of black locust. Enormous numbers of older trees, especially slow-
growing overtopped trees, have also been badly damaged or killed. However,
vigorous dominant trees over 10 years old have seldom been killed. Infested trees
are physically weakened by the larval tunnels, the smaller ones often being re-
stricted to shrub form by repeated attacks (fig. 125B). Trees growing on poor sites
are especially susceptible to attack and suffer serious damage during periods of
prolonged drought.

A number of practices have been suggested for the prevention or reduction of
damage by the locust borer. These include planting superior varieties of black
locust, the use of mixed species in planting, the removal of old stag-headed brood
trees, selection of good sites for planting, thinning and mulching of stands, and
protection of young trees from fire or livestock grazing (5/5, 830, 1351).

The painted hickory borer, M. caryae (Gahan), is widely distributed in the
Eastern United States. Freshly cut hickory logs are normally preferred for breeding,
but dead trees of several other hardwoods such as black locust, honeylocust, oak,
hackberry, mulberry, walnut, butternut, and ash also are attacked occasionally.
Hickory wood cut during the winter may be completely riddled by midsummer. The
adult closely resembles the adult of the locust borer, but is slightly shorter on the
average, ranging from 10 to 20 mm long. The elytra also are more tapering, the
prosternum is wider than long, and the antennae of the male are longer than the
body.

Adults emerge in early spring and deposit their eggs beneath bark scales on logs
cut the previous winter. The larvae feed for several weeks under the bark and then
bore into the sapwood and later the heartwood. Pupation occurs in the fall at the end
of the larval mine behind a wad of fibrous frass. The adult, like the adult of the
locust borer, emerges through the larval gallery and entrance hole. Winter is spent
in the pupal stage and there is one generation per year.

Megacyllene antennatus (White), commonly known as the mesquite borer,
breeds in mesquite and acacia in Texas and other Southwestern States. The adult is
robust, brownish black, and from 12 to 30 mm long. It is marked with fine white or
gray hairs and a dark spot in the center of the thorax. Eggs are deposited in crevices
of the bark of recently cut wood. The larvae feed first beneath the bark and then in

287

: Bins:
F-501514, 501515
Figure 125.—Damage caused by the locust borer,

Megacyllene robiniae: A, larval damage in heartwood;
B, damage to small trees.

288

the wood, excavating extensive mines and pushing quantities of frass to the outside
through a hole in the bark. There are two generations a year. This species 1s
especially injurious to mesquite cordwood. Fenceposts also are greatly weakened or
destroyed. Information on control has been published (258).

The banded hickory borer, Knulliana cincta (Drury), occurs throughout much
of the Eastern United States where it breeds in the dead branches and trunks of a
wide variety of hardwoods, such as hickory, walnut, oak, eastern hophornbeam,
plum, and apple. Hickory appears to be preferred. The adult (fig. 126) is dark
brown and from 16 to 30 mm long. The body is clothed with fine grayish hairs, and
there is a short sharp spine on each lateral margin of the thorax. Each elytron has an
oblique yellow spot near the base and two slender spines at the tip. Eggs are
deposited during the summer beneath the bark or directly on the wood of recently
felled, dying, or dead trees. The larvae feed beneath the bark during the remainder
of the summer, deeply scarring the wood and pushing out huge quantities of
granular frass through small openings in the bark. During the fall and following
summer they bore into the wood and mine it extensively. Pupation occurs in the fall
or spring between wads of fibrous frass at the end of the tunnel. The life cycle
probably requires 2 years for completion. Cordwood, logs, posts, and rustic work
are frequently seriously damaged by the species. The prompt milling and seasoning
of summer-cut wood are recommended control practices.

yey

F-480484
Figure 126.—Adult of the banded hickory borer,
Knulliana cincta.

The cottonwood borer, Plectrodera scalator (F.), breeds in the bases and roots
of living cottonwood, poplars, and willows mainly in the Southern States, but is
also found from New York and Michigan to Montana and Texas. The adult is 25 to
38 mm long and beautifully marked. The ground color is black, but this is obscured
by patches and cross stripes of fine, pure white hairs that surround black, hairless
areas (fig. 127A).

Adults appear in late spring or early summer and feed on the tender shoots of
young trees. These shoots often break, shrivel, and turn black. Eggs are deposited
in pits chewed in the bark at the base of the tree, at just above, or below the ground
line. Trees of all sizes are subject to attack. Larvae (fig. 127B) begin boring at the

289

f f

F-519918, 519919

Figure 127.—The cottonwood borer, Plectrodera
scalator: A, adults; B, larva.

root collar and tunnel downward into the roots. Coarse, fibrous frass may be ejected
from galleries at the root collar during early larval development. Part of a brood
develops in | year and the remainder in 2 years (878, 1/30). Young plants may be
hollowed, partially severed, or girdled at or slightly below the root collar, causing
breakage (875). Damage has been greatest in nurseries, young plantations, and
young natural stands growing on sandy soils. Infestation counts have run as high as
27 percent in I- and 2-year-old nurseries (//30). Borer populations and losses can
be reduced by using borer-free planting stock and by plowing out and destroying
nursery rootstock at 3-year intervals. Insecticides can be used to control the adults
before oviposition.

The sugar maple borer, Glycobius speciosus (Say), breeds in living sugar
maples in southern Canada and throughout the Northeastern States, westward to the
Lake States and southward through the Appalachians. The adult is robust, velvety
black, and from 19 to 28 mm long. The head is clothed with fine yellow hairs; the
pronotum is much wider than long, constricted at the base, and marked with two
parallel yellow bands on each side. Each elytron bears five yellow bands, with
those at the front forming a W-shaped design. Full-grown larvae reach a length of
50 mm (583).

Eggs are deposited in bark crevices, under bark scales, or around wounds,
usually during July and August. The larvae feed beneath the bark. Their tunnels run
more or less across the grain and cut deep channels in the wood. The winter is spent
as a larva in a chamber formed in the sapwood. The following spring, feeding is
resumed, with the larva cutting a larger gallery in the sapwood. The mature larva
bores deep into the wood and constructs a pupal cell at the end of its tunnel. Before
entering the cell, it cuts an exit hole through which the adult emerges. During this
activity, the larva pushes considerable quantities of sawdust to the outside. Pupation
occurs in the spring, and the life cycle requires 2 years (766, 1/56).

The presence of transverse ridges or elevations on the large limbs or trunks of
sugar maple, or of sawdustlike frass and moisture on the bark, are evidence of
attack by the sugar maple borer. The bark over ridges is pushed outward at an angle
or is broken up in the form of cracks, some of which may completely girdle the tree.

290

These cracked, swollen areas often resemble cankers or galls. Damage is generally
most severe to shade trees or to trees growing in open stands or along streams. The
larval galleries and associated stain, decay, and twisted grain cause defect and
degrade in lumber sawed from infected trees (583). Growing sugar maple in well-
stocked groups, avoiding overgrazing, and removing and burning infested dead
limbs and trees before adult emergence in the spring should be helpful in reducing
losses (1078). Borers in shade trees can be killed by injecting a fumigant into their
tunnels or by piercing their bodies with a wire pushed into their tunnels.

The red oak borer, Enaphalodes rufulus (Haldeman), breeds in the trunks of
living oaks in southern Canada and throughout the Eastern United States, west to
Minnesota, Iowa, and Texas. In the Central States, red, scarlet, and black oaks are
especially subject to attack. Adults are light brown with spots of lighter fine hairs,
and are 14 to 28 mm in length (fig. 128). There are two small tubercles on the disk
of the thorax and one triangular spot at its rear. The elytra are notched at the apex,
and the sutural angles are produced into spines.

Eggs are deposited singly on living trees over 5 cm in diameter in bark crevices
or beneath lichen patches during June and July. Young larvae bore directly into the
phloem, and they feed there during the remainder of the summer, excavating cave-
type burrows 10 to 15 mm square. The larvae spend the winter in these burrows and
continue their phloem feeding in the spring. In early summer, they bore into the
wood and direct their tunnels obliquely upward in the sapwood and straight upward
in the heartwood for distances of 15 to 25 cm. The second winter is also spent in the
larval stage with pupation occurring in the spring. The life cycle requires 2 years in
the Central States with adult emergence in the odd-numbered years. Granular frass
pushed out from points of attack and wet spots caused by sap leakage are evidence
of attack (539).

A high percentage of the large oaks in the Eastern, Southern, and Central States
are attacked by this species, resulting in serious defects and serious degrade in the
timber. The loss in grade can amount to 40 percent of the current tree value, which,
at 1980 prices, is about $80 per 5.7 cubic meters for factory-grade lumber in terms

Courtesy J. D. Solomon, South.
Hardwood Lab; Stoneville, Miss.

Figure 128.—Adult of the red oak borer, Enaphalodes
rufulus.

291

of reduced quality caused by larval tunnels. About 38 percent of the oak wood
used for lumber, cooperage, and veneer in the Eastern United States is affected
(321). Damage to upland oaks in the Central States can be reduced by poisoning
borer-infested trees with a herbicide after egg laying is completed in August. Larval
mortality occurs in a short time if the tree is deadened while they are still feeding in
the phloem. This can be accomplished during ordinary stand-improvement opera-
tions, because the trees selected for removal in this work are the very ones most
likely to be heavily infested (538).

Enaphalodes cortiphagus (Craighead), the oak-bark scarrer, breeds in the bark
of living, mature oaks throughout the Eastern United States and westward through
the Ozark Mountains. White and chestnut oaks are especially subject to attack.
Adults are dark brown with patches of fine, short, gray hairs on the head, pronotum,
and elytra and are 16 to 27 mm in length. Eggs are deposited in bark crevices in the
spring, and the larvae feed in the bark for nearly 3 years. At the end of this period
they bore deeper into the bark and excavate a large pupal cell in the sapwood. This
excavation usually damages several layers of annual growth of the wood, causing a
large black defect and the formation of scars on the outer surface of the bark. The
presence of this defect results in considerable degrade of the lumber. The related
species E. atomarius (Drury), breeds under the bark at the bases of dead trees and in
the stumps of oaks, chestnut, walnut, hickory, and hackberry in southern Canada
and throughout the Eastern United States.

The genus Goes contains a few species that attack living hardwoods. Large
tunnels constructed in the heartwood result in defects in lumber cut from infested
wood. Eggs are deposited singly in oval pits chewed through the bark. Tunnels in
the wood are excavated inward and upward and are kept open. Fibrous, granular
frass is expelled through openings maintained at the egg scar. Life cycles vary from
2 to 5 years, depending on species.

The white oak borer, G. tigrinus (De Geer), the largest species in the genus
occurs throughout the Eastern United States. White oak is its preferred host
throughout most of its range, but various other oaks, and hickory and walnut are
attacked occasionally. The adult is large, robust, dark brown, and from 25 to 30 mm
long. It is irregularly covered with a dense coat of fine white hair, giving it a white
and brown mottled appearance (fig. 129). The basal part of the elytra is roughened
with small, black, elevated points: and there is a strong spine on each side of the
thorax. Full-grown larvae are up to 37 mm long.

Adults emerge in May and June and feed for 1 or 2 weeks on the bark of tender
twigs and leaves of oaks. Eggs are deposited in niches gnawed in the trunks of
young trees, usually from 5 to 30 cm in diameter, or in the branches of larger trees.
The young larvae bore directly into the sapwood. Then they tunnel upward and
penetrate deep into the heartwood, excavating tunnels up to 25 mm in diameter and
to 25 cm in length. Each borer makes two separate holes in the tree—a small
elongate entrance hole that it keeps open for the expulsion of frass and a round hole
made for the emergence of the adult. The life cycle requires from 3 to 5 years,
depending on locality (//33).

The white oak borer is a major pest of overcup oak in the bottom lands of
Mississippi (//34). Small trees down to 25 mm in diameter are attacked and
seriously damaged. Trees growing on heavy clay soil with poor drainage. or where
flooding is prolonged into the growing season, are frequently infested. It has been
ranked as the primary pest responsible for defect in rejected staves in production of
white oak cooperage in Ohio (322).

292

Courtesy J. D. Solomon, South.
Hardwood Lab., Stoneville, Miss.
Figure 129.—Adult of the white oak borer,
Goes tigrinus.

Goes pulcher (Haldeman), the hickory borer, occurs in southern Canada and
throughout the Eastern United States and breeds in the sapwood and heartwood of
the trunk and branches of hickory and pecan (//28). The adult is from 17 to 25 mm
long. Its body is dark brown, covered with fine yellowish hair. The elytra are clay
yellow with dark bands across the base and the middle. In central Mississippi,
adults emerge during May and June and lay eggs singly in niches between the bark
and wood on trunks 2 to 14 cm in diameter, and at heights up to 4.8 m. The young
larvae bore irregularly shaped mines | to 2 cm wide under the bark, then extend
tunnels obliquely and vertically upward in the wood 8 to 17 cm, and then turn
horizontally back to near the surface. At maturity larvae reach a length of 18 to 28
mm. The adult chews a circular emergence hole through the bark at the terminal
point of the larval gallery. The life cycle requires 3 to 5 years.

The borer is common in the South, but populations vary greatly from one locality
to another. Hickories growing in the Piedmont area of North Carolina are particu-
larly prone to attack by this species. Open-grown trees and those near openings
within young stands are most heavily attacked.

The oak sapling borer, G. fesselatus (Haldeman), occurs from Pennsylvania
southward and in the Midwest where it usually breeds in the base and roots of small
oak saplings, preferably white and chestnut oaks (/58). Young chestnuts and
serviceberry are also attacked occasionally. The adult is dull brown, 20 to 27 mm
long, and its body is covered with small patches of prostrate, fine gray or yellowish
hairs occasionally arranged in rows. The larvae require 3 to 5 years to complete
their development. During this period they may completely hollow out the base of
the tree and cause its death. Less seriously damaged trees often produce bulblike
swellings around the wounds at the ground line. These trees are subject to wind-
breakage.

Goes pulverulentus (Haldeman), the beech borer, breeds in the small trunks and
branches of various hardwoods such as beech, oak, elm, and sycamore throughout
the Eastern United States and southern Canada (//27). The adult is brown, 18 to 28
mm long, and its body is clothed with short, brownish-gray hairs. The thorax has a
sharp spine on each side and the elytra are faintly barred at the middle and base with
fine, pale-brown hairs. In central Mississippi, adults emerge during May and June.
Eggs are laid singly in niches between the bark and wood. The niches usually are

293

clustered, averaging 17 per cluster. The larval boring habit is similar to G. tigrinus.
At maturity, larvae reach a length of 20 to 30 mm. The life cycle requires 3 to 5
years.

There is a tendency for the borer to favor the less vigorous, slower growing trees.
Since attack occurs only on trunks less than 12 cm in diameter and branches of
larger trees, the borer cannot be termed a major pest. Infested trees are easily
recognized by egg-niche clusters, sap-stained bark, dark-colored frass, and later by
ragged, longitudinal, overgrown scars.

Goes debilis LeConte, the oak branch borer, breeds in small, living branches
and terminals of oak in the Eastern United States, often causing gall-like swellings.
Adults are brownish and from 11 to 19 mm long. The head, thorax, and apical half
of the elytra are clothed with fine reddish-yellow hairs. The elytra are crossed by
two irregular brown bands, and each basal half is mottled with grayish hairs. The
life cycle requires 3 to 4 years. Infested stems become swollen and gall-like and
often break or die back (//29).

Many species of the genus Neoclytus occur in eastern forests and attack both
coniferous and deciduous trees. The larvae feed first beneath the bark, then bore
into the sapwood and heartwood, often completely riddling it with long mines
tightly packed with frass. The adults are slender-bodied beetles of medium size with
quite long legs and short and more or less clubbed antennae. The pronotum bears
transverse ridges and the body is marked with transverse yellow lines.

The redheaded ash borer, N. acuminatus (F.), one of our most common wood
borers, occurs generally throughout the Eastern United States and in southeastern
Canada. Its hosts include nearly all dying and dead hardwoods, but chiefly ash,
oak, hickory, persimmon, and hackberry. Unseasoned logs of ash, oak, and hickory
with the bark intact are especially subject to heavy attack. The adult is from 6 to 18
mm long. The head and thorax are reddish; the body is light brown with the apical
part of the elytra sometimes much darker. The elytra are also marked with four
transverse bands of fine yellow hairs and the middle and hind legs are long and
reddish.

Adults become active by mid-February in the Deep South and progressively later
until May or June in the North. Eggs are deposited beneath the bark of dead,
unseasoned wood. The larvae feed first beneath the bark, then they tunnel into the
sapwood and often reduce it to powder. In the South there are several generations
per year; in the North, only one (/252).

Neoclytus caprea (Say), the banded ash borer, occurs in eastern Canada and
throughout much of the United States. Its hosts are recorded as ash, hickory, elm,
mesquite, and, rarely, white oak. In the Eastern United States, it commonly breeds
in ash logs. Adults are dark brown to almost black and from 8 to 17 mm long. There
is a line of fine, white or yellowish hairs on the thorax and four bands of the same
material and color across the elytra. The first two bands meet, almost forming
circles.

Adults emerge in early spring and fly to host material where they deposit their
eggs in crevices in the bark. Ash logs cut during the winter are especially subject to
attack. The larvae feed for awhile under the bark and then bore into the sapwood
where they feed for the remainder of the summer. Pupation occurs in the fall, but the
adult does not emerge from the wood until the following spring. There is usually
one generation per year; however, if the infested material is sawed, stored, and dried
out, the life cycle may require several years. Ash logs left in the woods or stored

294

with the bark on literally may be honeycombed with tunnels tightly packed with
frass (fig. 130).

Other species of Neoclytus likely to be encountered in eastern forests and some of
their known hosts are as follows: N. mucronatus mucronatus (F.) (fig. 131)—the
dead branches and trunks of hickory; N. jouteli jouteli Davis—dead oak twigs; N.
fulguratus Casey—dead branches of oak; N. scutellaris (Olivier)—elm, oak,
hickory; and N. muricatulus muricatulus (Kirby)—larch, spruce, pine (northern
coniferous forests of North America, in the East and down the Rocky Mountains to
New Mexico and down the Cascades to Oregon).

The genus Saperda is represented in eastern forests by several important species
(401). Depending on the species, the larvae bore in (1) large branches or trunks of
living trees; (2) small branches and produce galls; or (3) living or dead wood of
dying or recently killed trees. Adults are medium size and cylindrical in form. The
head is quadrate in front and very flat; the antennae are about as long as the body;
the first joint of the hind tarsus is elongated; and the body is rather densely clothed
with a hairy covering.

The poplar borer, S. calcarata Say, occurs throughout the United States and
Canada, wherever poplars grow (878). Willows are also subject to attack. In the
southern latitudes, cottonwoods are the primary host; in the northern area, aspens

: Sy

PN eS

Courtesy Duke Univ. Sch. For.

Figure 130.—Galleries of Neoclytus caprea, the banded
ash borer. Note that tunnels are tightly packed with
granular frass.

295

Courtesy Duke Univ. Sch. For.

Figure 131.—Adults of Neoclytus mucronatus
mucronatus.

are most severely damaged. Living trees, healthy or injured, are attacked. A strong
preference is shown for trees that have partially succumbed to former attacks. The
adult is from 20 to 28 mm long, grayish black or reddish brown, and densely
clothed with fine, gray and yellow hairs (fig. 132). There are also yellowish stripes
on the thorax and orange-yellow markings on the wing covers. Full-grown larvae
are creamy white and about 30 mm long. Adults appear during the summer, feed on
the bark of young twigs, and deposit their eggs in small slits cut in the bark, usually
in the middle third of the tree. The larvae bore into the inner bark and sapwood
where they later spend the winter. In the spring, they bore into the sapwood and
heartwood and feed there until they are mature. Attacked trees are characterized by
the presence of swollen scars and holes in the trunk and larger branches. Each larva
bores an opening out to and through the bark through which frass is expelled and
sap exudes. Wet areas around these holes blacken and appear varnished. The life
cycle requires 3 to 5 years in the North and 2 years in the Deep South. The usual
time is 3 years for most of its range. In the North, the second winter is spent as a
mature larva in a cell at the end of its tunnel (98/).

Small trees are occasionally killed by larvae girdling beneath the bark. Larger
trees are seldom killed outright, but the large larval tunnels make them susceptible
to windbreakage. They literally may be riddled with tunnels. Larval openings and
tunnels serve as openings for various rots, decays, and other injurious insects which
may kill the tree or degrade the lumber (878). Poplar plantations may be ruined and
valuable shade trees severely damaged. Damage in forest stands also may be
severe. In the Lake States, successful attacks appear to be concentrated in indi-
vidual trees or small groups of trees unevenly distributed throughout the stand.
These trees, commonly known as brood trees, are usually the larger, faster growing

296

F-519920

Figure 132.—Adults of the poplar borer,
Saperda calcarata.

trees in stands averaging between 7.5 and 28 cm d.b.h. Lake States infestations also
tend to increase with a decrease in stand density.

The best practice in the management of poplar in the Lake States apparently is to
maintain well-shaded stands and then clearcut them at maturity. The removal of
“brood trees” should also be helpful (388).

The roundheaded appletree borer, S. candida F., occurs in Canada and
throughout the Eastern United States. It is most serious as a pest of apple orchards,
but it also breeds in mountain-ash, hawthorn, and serviceberry. The adult is
brilliantly white except for three broad, brown, longitudinal stripes extending the
full length of the back, and is from 15 to 20 mm long. Full-grown larvae are creamy
white and about 30 mm long (562).

Adults are present from June to September and deposit their eggs in slits cut in
the bark at the base of living trees. They feed on ‘the foliage and sometimes on
tender bark. The larvae feed beneath the bark for | year and then bore into the
wood, making large excavations and riddling it. The presence of tendrils of frass on
the bark or at the base of the tree 1s evidence of attack. Heavily infested trees may be
killed in a single season. Two or more years are required to complete the life cycle.
Keeping trees healthy is an effective preventive measure. Borers in high-value trees
can be killed by injecting a fumigant into borer burrows.

The linden borer, S. vestita Say, occurs in the Northeastern States and Canada.
Its preferred host is basswood but it also attacks poplars. The adult is reddish
brown, densely covered with prostrate, fine, olive-yellow hair, and is from 12 to 21
mm long. The tips of the antennae are brown, and there are three small black spots
on each wing cover. Adults feed on leaf petioles, the larger veins of leaves, and the
bark of growing shoots, often killing the tips of infested branches. The larvae feed

297

beneath the bark and often bore deep into the wood. Unhealthy and weakened trees
are most susceptible to attack.

Saperda discoidea F. breeds in dead and dying hickories and butternut from New
York to Louisiana and Nebraska. Hickory trees infested with the hickory bark
beetle are particularly subject to attack. Adults are 10 to 17 mm long. The male is
blackish with reddish legs and lines of fine, grayish hair on the thorax. The female
is reddish brown or nearly black, clothed with fine, yellowish hairs, and has two
spots separated by a curved bar on the elytra. Eggs are deposited in bark crevices or
in holes in the galleries of bark beetles during the late spring and summer. The
larvae feed gregariously beneath the bark, making extensive meandering mines.
Most of the larvae mature in | year.

Saperda inornata Say, the poplar-gall saperda, breeds in the living branches
and main stems of poplars and willows in the Northern and Central States (936).
Adults are black with fine, gray hairs and are about 12 mm long. Eggs are laid in U-
shaped slits in the bark. Callous tissue forming over these slits causes the formation
ot globose galls. The larvae mine around the stem and then bore into wood, making
galleries about 25 mm long that run parallel to the axis. The life cycle requires | or
2 years. Usually the infested twigs are not killed. Branches or small stems bearing
numerous galls may stop growing, however, and become subject to windbreakage.
Growers should avoid establishing new plantings near heavily infested stands
(1336).

Saperda obliqua Say, the alder borer, breeds in the bases of living alders and
sometimes birch in the Northern and Central States. The adult is reddish brown and
from 10 to 20 mm long. There are two dark bands on the thorax and four oblique
bands on the wing covers. Larvae feed first beneath the bark near the base, often
girdling the stem. Later they bore into the stem and tunnel upward for 8 to 15 cm. ©
Swollen areas appear at points of attack on stems that survive, and large amounts of
frass are pushed out through the openings at the egg scars. Alders in ornamental
plantings are frequently killed.

Saperda fayi Bland, the thorn-limb borer, occurs in the Northeast and Midwest
and breeds in the twigs of hawthorn causing gall-like, gnarly swellings. Adults are
reddish brown and about 12 mm long. There is a white stripe on each side of the
thorax that extends up to the base of the elytra. The elytra bear two white spots at
the base and tip, and a single large one in between. Infested twigs are subject to
windbreakage. S. moesta LeConte causes the formation of globose galls on the
stems and branches of poplar and willow in the northern parts of the United States.
Adults are dark brown to black and about 10 mm long. S. /ateralis F. breeds in dead
hickory, elm, basswood, oak, cherry, and in hickory sprouts. Adults are black or
brownish black with the head, thorax, and elytra margined by broad, red lines and
are about 12 mm long. The male has a tooth on each of its claws. S. imitans Felt &
Joutel breeds in dead hickory, willow, and basswood; S. cretata Newman, in living
apple and the limbs of living hawthorn; S. mutica Say, in dead willow.

The elm borer, S. tridentata Olivier, occurs in southeastern Canada and
throughout the Eastern United States and breeds in dead and dying American and
slippery elms. The adult is grayish or fawn colored and from 9 to 17 mm long (fig.
133). An orange-yellow or yellowish band extends from the head to the tip of each
wing cover, twin black spots occur below these bands on the thorax, and there is a
crossband at the base of the wing covers followed by two oblique bands. Full-grown
larvae are about 12 to 15 mm long.

298

WE
NS
N

SRO}

Courtesy lil. Nat. Hist. Surv.

Figure 133.—Adult of the elm borer,
Saperda tridentata.

Adults are present from late spring to late summer, and feed on young leaves and
young twigs. Eggs are deposited in small holes chewed in bark crevices, usually on
freshly cut logs or weakened trees. The larvae bore beneath the bark, filling their
mines with fibrous frass, and completely destroying the phloem and cambium
(973). When they reach maturity they bore into the wood and construct cells in
which to pupate. There 1s usually one generation per year; however, in rapidly dried
wood 3 years may be needed to complete the life cycle.

Park and shade tree elms, especially the older ones and those in a weakened
condition, are severely injured by this species. Affected trees tend to die very
slowly, a branch at a time. The removal of infested branches is sometimes helpful in
control.

Smodicum cucujiforme (Say), the flat powderpost beetle, occurs throughout the
Eastern United States. The larvae excavate extensive meandering galleries in dry
heartwood of oak and hickory. Stored lumber is frequently infested, the larvae
continuing to feed in it until the wood is thoroughly riddled. The adult is small,
elongate, very depressed, dull yellowish, shiny, and 7 to 11 mm long. Adults
appear from June to August. Eggs are laid in crevices of exposed wood. The larvae
excavate tunnels about 3 mm in diameter in the wood, tightly packing them with
frass. Pupation occurs in an enlarged portion of the mine near the surface of the
wood. There is normally one generation per year. In dry wood, several years may
be required to complete the life cycle.

The genus Oberea is represented in the Eastern States by a number of slender,
cylindrical beetles, the larvae of which bore in the twigs, branches, or stems of
various forest, shade, and ornamental trees. These beetles are distinguished by the
presence of a broad tooth on each tarsal claw.

The dogwood twig borer, O. tripunctata (Swederus), breeds in dogwood, elm,
sourwood, laurel, azalea, viburnum, and various fruit trees in the Eastern United
States. The adult is yellowish and about 14 mm long. The head is reddish or dark
brown and there is a black spot on the scutellum. Adults appear in early spring and,
after girdling the tip, the female deposits her eggs in living twigs of the host. The
larva bores down the center of the twig, making a long series of holes for the
expulsion of frass and cutting off portions of the twig as it bores into the green

299

wood. The winter is spent in the larval stage in the twig. Pupation occurs in the
spring between two wads of fibrous frass. The portion of the twig containing the
cell may have been girdled previously. Occasionally dogwoods and elms are
seriously damaged. Cutting off and burning infested twigs might be helpful in
control (220).

Oberea schaumii LeConte occurs in southern Canada and south in the Eastern
States to the Gulf Coast. It breeds in poplar. Adults are about 14 mm long; the
thorax is yellowish or yellowish orange to black and is marked by four round,
smooth spots (935). The elytra are either yellowish or black. Eggs are laid in the
stems and branches of suckers and seedlings; also in the twigs of saplings, poles,
and large trees. The larvae feed near the pith, excavating tunnels up to 15 cm long,
and boring holes to the outside for the expulsion of frass. Black, necrotic areas
develop around these holes. The life cycle usually takes 3 years in northern
Michigan, but a few take 2 or 4 years. Infested twigs do not die and are usually not
sufficiently weakened to permit windbreakage.

Oberea myops Haldeman, the rhododendron stem borer, breeds in rhodo-
dendron, azalea, and mountain-laurel. The adult is pale yellow and about 12 to 15
mm long. There are two black spots on the thorax and the elytral margins are dark.
Eggs are deposited in the bark of twigs between two girdles about | cm-apart. The
larva bores down the twig, into the stem, and on down to the ground. Here, it cuts
off the stem, and then bores into the roots. Frass is expelled through holes along the
stem and at the ground line. This species is frequently a serious pest in ornamental
plantings of rhododendron. Cutting and burning infested twigs as soon as they are
noticed is recommended.

Oberea ocellata Haldeman, the sumac stem borer, occasionally causes serious
injury in sumac plantings. Adults are about 13 to 15 mm long. The head and
underside of the body are red; the thorax is red with two black spots on the disk; and
the elytra are black. The female girdles the tip of the plant and then lays an egg just
below the girdle. The larva bores down the stem through the pith to the roots. Here,
it constructs long tunnels and feeds for two seasons. During the fall of the second
year, it cuts the plant off near the ground and plugs the stub with a wad of frass.
Below this plug, pupation occurs the following spring.

Oberea ruficollis (F.) breeds in the stems and roots of sassafras. Adults are pale,
reddish yellow and about 18 to 20 mm long. The antennae, tibiae, and tarsi are
black and the elytra are densely clothed with fine gray hairs. The feeding habits of
the larvae are similar to those of O. ocellata except that they seldom cut off stems
near the ground. Infested plants usually survive but may be badly deformed.

Other species of Oberea sometimes encountered are: O. pallida Casey—in alder
in Pennsylvania; O. ulmicola Chittenden—in the branches of oak, elm, hickory,
dogwood, and black cherry in Ohio and Pennsylvania; and O. ferruginea Casey—in
willow canes.

The genus Oncideres contains several species, the females of which deposit their
eggs in previously girdled terminals, twigs, or small branches of large trees, or in
the stems of seedlings or sprouts. The girdled portions soon die and most of them
fall to the ground. The larvae feed in the wood of this fallen material until the
middle of the following summer, loosely filling their mines with frass. Pupation
occurs in a cell formed by walling off a portion of the mine with fibrous frass. The
adults appear in late summer or early fall and feed on the thin bark of twigs or
stems. There is one generation per year in the South. In the North, many larvae do
not pupate until the second year.

300

The twig girdler, O. cingulata (Say), is the most important eastern member of
the genus. Its hosts include hickory, pecan, persimmon, elm, oak, honeylocust,
hackberry, poplar, basswood, dogwood, sourwood, and various fruit trees. As a
pest of pecan in the South it has been incorrectly referred to as Oncideres texana
Horn, the pecan twig girdler (74/). Adults (fig. 134A) are grayish-brown and
about 15 mm long. The front of the head is clothed with fine golden hairs; the elytra
are clothed with fine gray hairs and are marked with scattered yellow spots. Full-
grown larvae are up to 18 mm long.

; Courtesy Duke Univ. Sch. For.

Figure 134.—The twig girdler, Oncideres cingulata: A,
adult; B, young hickory tree deformed by repeated
attacks.

Adults emerge in late summer and feed on the tender bark and tips of twigs of
their host. The egg laying and larval habits are as described earlier for the genus.
On heavily infested large trees dozens of branches may be girdled and severed;
often, many of them hang on for long periods before dropping to the ground. Such
trees are not only damaged severely but also are ragged and unattractive. Hickory
seedlings are especially prone to attack and damage, often becoming distorted
following attack (fig. 134B). Young pecan trees grown in plantations for timber
purposes have had up to 22 percent of the terminals girdled in a single year (662).
Honeylocust seedlings in nurseries are also subject to serious injury. Collecting and
burning infested twigs and branches during the fall or winter is an effective method
of control, provided that the trees to be protected are located at some distance from
heavily infested stands.

Oncideres pustulatus LeConte attacks sweet acacia, acacia, and mesquite in the
Rio Grande Valley and other parts of the Southwest.

301

Eburia quadrigeminata (Say), the ivory-marked beetle, occurs throughout the
Eastern United States and breeds in the dry heartwood of various hardwoods,
including oak, hickory, ash, maple, honeylocust, elm, chestnut, and baldcypress.
Adults are light brown, with pairs of ivory spots at the base and middle of each
elytron, and are from 12 to 24 mm long. The larvae are wedge shaped, have tough,
shiny skins, and are sparsely clothed with golden hairs. Mature oaks having
catfaces or scars through which the larvae can gain access to the heartwood are
often badly damaged. Seasoned lumber is also subject to occasional attack. The
normal life cycle appears to be 2 years, but it may be considerably longer. There are
records of adults emerging from flooring, doorsills, and furniture 25 years after
they were placed in use.

The twig pruner, Elaphidionoides villosus (F.) occurs throughout the Eastern
United States. It breeds in the twigs and branches of living hardwoods such as the
oaks, hickory, maple, locust, hackberry, walnut, elm, sweetgum, and pecan. Adults
are slender, elongate, brown, and are from 11 to 18 mm long. The dorsal surface is
clothed with irregular patches of fine gray hairs. There are spines on the first few
joints of the antennae, and the tips of the elytra are notched and bispinose.

Eggs are deposited in slits in the bark at leaf axils near the tips of twigs and small
branches in late spring. Young larvae feed beneath the bark, often consuming much
of the wood toward the base of the twig. Older larvae bore down the center of the
stem toward the base until late in the summer. Then they sever the branch by
making several concentric circular cuts from the center outward to, but not includ-
ing, the thin bark (fig. 135). These branches, from 0.5 to 5 cm in diameter, break
and fall to the ground with the larvae in them. The ground under heavily infested
trees may be literally covered with these fallen twigs and branches. The larva
returns up through the fallen branch and forms a cell between wads of fibrous frass
where it pupates in the spring, or in the fall. There appears to be one generation per
year. Heavily infested trees may be seriously damaged but are seldom killed. Shade
and park trees may be so severely pruned that they lose much of their esthetic value.
The presence of numerous dead twigs and branches hanging in the crown also
detracts from their appearance. Collecting and burning infested twigs and branches
in the spring before the adults emerge should be helpful in control.

F-504092

Figure 135.—Larva of the twig pruner, Elaphidionoides
villosus, in an oak twig. Note ends of severed twigs.

302

Elaphidionoides incertus (Newman), the mulberry bark borer, a species that
closely resembles the twig pruner, breeds in the outer bark of living mulberry trees
in the Eastern United States. Infestations have also been recorded in chestnut cak
and pignut hickory. It does not cause serious damage.

Elaphidion mucronatum (Say), the spined bark borer, breeds in the dead
branches of various hardwoods in the Eastern United States. Adults are dark brown,
irregularly clothed with fine brown hair and are from 13 to 19 mm long. Eggs are
deposited beneath bark scales, and the larvae feed beneath the bark during the first
year of their lives. During the second year, they bore in the sapwood. Wood used in
making rustic furniture is often damaged.

Desmocerus palliatus (Forster), the elder borer, breeds in elder wherever it
grows in the Eastern United States. Adults are bright, metallic blue with nearly all
of the basal half of the elytra yellow, and are from 18 to 26 mm long. The pronotum
is much wider than long and is constricted at the apex; the wing surfaces are densely
and coarsely punctured. Eggs are deposited in crevices of the bark at the base of the
stem. The larvae feed in the roots and base of their host, eating out the pith and
filling their mines with coarse, rather fibrous frass. The larval period lasts 2 to 3
years. Heavily infested plants may be seriously injured or killed.

Dorcaschema wildii Uhler, the mulberry borer, breeds in the living branches of
mulberry and Osage-orange in the Southern and Central States. The adult is dark
brown and from 16 to 22 mm long (fig. 136). The body is covered with fine gray
hairs except for small bare punctures scattered over the surface. Each wing cover
has a light-brown stripe along its outer margin.

In the Deep South, adults appear as early as mid-May. Eggs are deposited in
niches chewed in the bark. Young larvae feed between the bark and wood, destroy-
ing irregular 2- to 8-cm? patches of cambium. After about | to 3 months they bore
into the wood, constructing galleries that angle upward and inward for about 5 cm
and then bend back toward the surface. These galleries are close together and often
overlap. The winter is spent in the larval stage, and the life cycle varies from | to 2
years. Suppressed and fire-damaged trees are preferred, but vigorous, healthy trees
are also attacked occasionally. Individual branches and even entire trees may be
completely girdled and killed. Trees that recover usually have large scars on the
trunk (//26). The smaller but related species, D. alternatum (Say), is commonly
found in trees infested with the mulberry borer. The larvae feed in dead and dying
branches of mulberry, sometimes attacking green limbs that have been slightly
injured.

Aneflomorpha subpubescens (LeConte), the oak-stem borer, occurs in the
Eastern, Southeastern, and Central States. It breeds in small living oak and chestnut
seedlings from | to 3 cm in diameter, and occasionally in the branches of larger
trees. The adult is narrow, elongate, light brown, clothed with semierect fine brown
hairs, and 17 mm long (fig. 137). There is a stout spine on each of the third and
fourth segments of the female antennae, and the tips of the elytra are notched and
bispinose.

Eggs are deposited at leaf bases near the tops of seedlings and sprouts. The larvae
bore into the center of the stem and tunnel downward, mining out the wood as they
feed. Section after section of the stem is cut off as the larvae proceed toward the
base. Frass is extruded through a single row of small holes cut through the bark to
the outside. During late summer the full-grown larva burrows to the base of the
main stem and often into a root. Here it constructs a pupal cell between two wads of
fibrous frass. The stem is usually cut off at the ground line. There appears to be one

303

generation per year. During certain years a high proportion of the oak seedlings in
the Southeast are killed by this species.

Callidium antennatum antennatum Newman occurs throughout the Eastern
United States and breeds in dead or recently felled conifers, or in lumber with the
bark on. It is especially common in dead pines in the South. The adult is flattened,
bright metallic blue or bluish black and from 9 to 14 mm long (fig. 138A). Male
antennae are somewhat shorter than the body; the thorax is rounded, with depres-
sions on each side of the middle; and the legs are black with large femora.

Adults appear early in the spring and deposit their eggs beneath bark scales on
dead trees or on cut wood that has seasoned over winter. The larvae feed in the
phloem and outer sapwood making broad, wavy tunnels in the wood (fig. 138B)
and pushing large quantities of frass through small holes in the bark. Pupation
occurs in the wood in long cells plugged with wads of fibrous frass. There is usually
one generation per year.

This species frequently causes serious losses to improperly edged lumber in
lumber yards. Rustic work and houses built of pine logs are also subject to serious
damage, the wood being badly riddled and weakened and the bark so loosened that
it falls away. Prompt utilization of logs, the removal of wane on sawed lumber, and
kiln drying are recommended methods of control.

Callidium texanum Schaeffer, the blackhorned juniper borer, breeds in various
conifers, preferably juniper, throughout the United States. Its habits are similar to
those of C. antennatum antennatum. Other eastern species include: C. schotti
Schaeffer, which breeds in the dead branches of eastern redcedar in the Midwest,
and C. violaceum (L.), which occurs in pine, larch, and spruce in the Northeastern
States.

The genera Tylocerina and Neacanthocinus contain a number of species com- —
monly known as pine bark borers (305). The adults are elongate, rather flattened,
and from 7 to 28 mm long. They are also usually black and mottled or striped with
grayish-white or brown pubescence. The basal joints of male antennae are fringed
beneath with hairs, and the ovipositor of the female is characteristically extended.

Tylocerina nodosa (F.) is a common species in the South, but is found as far
north as Pennsylvania. It breeds in the thick bark of pine logs and in dying and
recently killed pines. The adult (fig. 139) is gray with velvety black markings and is
about 25 mm long. Male antennae are sometimes at least three times the length of
the body. Eggs are deposited in pits chewed in the bark or in the exit holes of bark
beetles. The larvae feed in the bark where they often compete with and destroy bark
beetle broods. Pupation occurs in nestlike chambers in the bark, near the surface.
There is one generation per year.

Neacanthocinus obsoletus (Olivier) breeds in recently cut pine and balsam fir in
eastern North America. The adults are 7 to 14 mm long. N. pusillus (Kirby) is
limited to the northern tier of States from Maine to Minnesota and into Alaska. The
adults are 7 to 10 mm long. It has been reared from windthrown and fire-killed red
and jack pines, balsam fir, and spruce.

Strophiona nitens (Forster), the chestnut bark borer, breeds in thick, moist bark
in crotches and at the base of living chestnut and oak trees. The adult is velvety
black and from 10 to 15 mm long. There are golden-yellow bands on the margins of
the thorax and on the elytra. Damage is seldom serious although large patches of
bark are sometimes killed.

Stictoleptura canadensis (Olivier) breeds in dead pines, spruces, hemlock, and
sometimes balsam fir in southern Canada and the Northern States. Living trees are

304

Courtesy Duke Univ. Sch. For.

Figure 137.—Adult of Aneflomorpha
subpubescens, the oak-stem borer.

Bie RE!

A, courtesy Duke Univ. Sch. For.
B, F-519924

Figure 136.—Dorcaschema wildii, the
mulberry borer: A, adult; B, damage

by larvae.

"tip tl

}

A Bs \ — —
A, F-519946; B, courtesy Duke Univ. Sch. For.

Figure 138.—Callidium antennatum antennatum: A,
adult; B, larval galleries under the bark and in the

wood.
305

Courtesy Duke Univ. Sch. For.

Figure 139.—Adult of Tylocerina
nodosa, a pine bark borer.

attacked occasionally at wounds, the larvae boring into the heartwood. The adult is
dull black, with the base of the elytra bright red. The upper surface is coarsely
punctured, and the antennae are generally ringed with yellow. Additional eastern
species of borers and their hosts include the following: Brachyleptura vagans
(Olivier)—butternut, hickory, birch, and pine logs; Lepturopsis biforis (New-
man)—white pine; Pygoleptura nigrella (Say)—pines and spruce; Strangalepta
vittata (Swederus) and Trachysida mutabilis (Newman)—red spruce; Trigonarthris
minnesotana (Casey)—hickory, elm, black cherry, and red spruce; and T. proxima
(Say)—maple, hickory, and basswood.

Phymatodes testaceus (L.), the tanbark borer, breeds in the bark of dead oak
trees and occasionally in stored hemlock bark in the Eastern and Central States.
Adults are elongated, flattened, and are from 8 to 17 mm long. Some are brownish
yellow or dark brown with lighter elytra and in some the thorax, abdomen, tibiae,
and tarsi are reddish yellow and the elytra, blue. The remainder are intermediate in
color. The larvae feed within or beneath the bark and pupation occurs in the
sapwood. Several years may be required to complete the life cycle. Bark stored for
tanning purposes is subject to heavy damage. P. dimidiatus (Kirby) breeds in fir,
spruce, and larch in the Northern States. Adults are from 5 to 11 mm long, and dark
brown, with a light-brown band across the base of the elytra. P. varius (F.) breeds in
or beneath the dead bark of hickory and oak in Eastern and Southwestern States.
The adult is light brown or dark brown to black and shiny. There are white bands on
the elytra, and the adult borer is 6 to 10 mm long.

Physocnemum andreae (Haldeman), the cypress bark borer, breeds under the
bark of girdled, felled, or dead baldcypress trees. The adult is reddish brown and
ranges in length from 11 to 21 mm. There is a curved white mark on each elytron, a
small tubercle near the base of the thorax, and club-shaped femora. Eggs are
deposited beneath bark scales, and the larvae feed beneath the bark, excavating
large mines that deeply scar the wood. Winter is spent in the larval stage, and
pupation occurs in the spring. The species often causes serious losses to bald-
cypress trees felled or girdled during lumbering operations. Rustic work con-

306

structed from this wood is also subject to serious damage. Rapid utilization of
girdled or felled trees and the storage of logs in ponds are effective control
practices.

Physocnemum brevilineum (Say), the elm bark borer, breeds in the corky bark
of living elm trees in southern Canada and the Eastern and Central States (5/4).
Adults are dark brown to black and from 9 to 20 mm long. The elytra are frequently
bluish with three longitudinal white marks. Eggs are deposited beneath bark scales.
The larvae feed in the phloem, constructing meandering, frass-packed galleries.
The bark over these galleries dies and falls off. P. violaceipenne Hamilton breeds in
the small branches of white oak in eastern Canada and the Northeastern States.
Adults are 8 to 17 mm long.

Parandra brunnea brunnea (F.), the pole borer, occurs in central and eastern
North America, and attacks a wide variety of hardwoods and conifers. Logs, poles,
and other wood products in contact with the ground, such as untreated crossties and
structural timbers, are also infested. The adult is flat, shiny, mahogany-brown, and
is from 8 to 21 mm long. Full-grown larvae taper slightly toward the rear and are
about 30 mm long.

Adults appear from July to October and deposit their eggs singly but close
together, deep in either solid or decayed wood. Attacks on living trees are usually
made at places where the wood is exposed such as at scars, wounds, or broken
branches. The larvae feed in the wood for 3 or 4 years. Although the wood may be
completely honeycombed, a covering shell of sapwood 1s always left intact. Pupa-
tion occurs in a cell in the wood. Many of the adults do not emerge from the wood
but mate and lay eggs in the cavities in which they are working. In living wood, the
wounds where the larvae gain entry will often heal over, leaving no external signs of
attack. Shade trees, telephone and telegraph poles, and structural wood in moist
locations or in contact with the ground are subject to severe damage. A consider-
able degree of protection of valuable shade trees can be provided by keeping them
healthy, by the removal or treatment of exposed dead and decaying wood, and by
covering pruning scars with paint.

The cedartree borer, Semanotus ligneus (F.), occurs throughout the United
States. Practically all species of conifers are subject to attack, but dying and
recently felled thujas and junipers are preferred. Adults are dark brown to black and
from 7 to 16 mm long. The thorax is rounded and hairy, except for several bare
spots on the disk. The elytra are sometimes black, but are usually dark blue with
yellow or orange markings.

Eggs are deposited beneath bark scales in the spring. The larvae feed first
beneath the bark, scarring the wood deeply. Then, they bore into the sapwood and
occasionally the heartwood. There is one generation per year. A related species, the
firtree borer, S. litigiosus (Casey), has been recorded from the Eastern United
States, but is primarily western in distribution. Its hosts include several species of
true firs, Douglas-fir, larch, plus several spruces. Male adults are usually all black,
whereas females are black, marked with orange.

The genera Asemum and Arhopalus contain a number of species that breed in the
sapwood and heartwood of the stumps of felled trees and in the lower portions of
dying trees. When abundant, the larvae may destroy large portions of the sapwood.
Asemum striatum (L.) and Arhopalus rusticus obsoletus (Randall) are common
eastern species.

Tragosoma depsarius (L.), the hairy pine borer, occurs from coast to coast in
southern Canada and the Northern States. It also occurs southward through the

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Appalachians in the Eastern States where it breeds in various dead pines. The adult
is a heavy, shiny dark-brown beetle from 20 to 40 mm long. The underside of the
body is very hairy, and the elytra are ridged. Larvae are tough-skinned and have
four teeth on the front of the head. Occasionally untreated crossties and poles, and
timbers in contact with the ground, are seriously damaged.

Atimia confusa confusa (Say), the small cedar-bark borer, breeds in dying
thujas, junipers, and related trees throughout the Eastern and Central States. The
adult is small, stoutish, and 6 to 9 mm long. The head, pronotum, elytra, and
venter are black. The dorsum is clothed with recumbent, fine gray hairs, and the
elytra are notched at the apices. Adults appear in early spring and again in early fall
and deposit their eggs beneath bark scales. The larvae feed entirely between the
bark and wood, packing fibrous frass behind them. Pupation occurs in cells in the
sapwood. The winter is spent as larvae or as adults. Rustic work constructed from
improperly seasoned thuja and juniper is subject to attack and serious damage. The
bark over damaged areas dries and peels off.

Aneflus protensus (LeConte) attacks living mesquite in Texas and other South-
western States. The adult is light or dark brown, has spines on the basal joints of the
antennae and the tips of the elytra, and is from 25 to 32 mm long. Eggs are
deposited in bark crevices at the forks of small branches. The larvae bore into the
branch and hollow it out. Frass is extruded through small holes in the bark. Black,
watery liquid drips from these holes and stains the foliage and ground beneath. The
interior of the larval mine is always stained black. Two years are required to
complete the life cycle. Infested branches and small trees are sometimes killed.

Tetropium cinnamopterum Kirby breeds beneath the bark of dead and recently
felled coniferous trees, and occurs in eastern Canada and the Northeastern States
(1009). The adult is oblong, somewhat flattened, brown to blackish, and about 12
mm long. The eyes are completely divided into upper and lower lobes. The larvae
feed gregariously beneath the bark, packing frass behind them in the mines. The
shallow-bored pupal cells in the sapwood are largely removed by slabbing. There 1s
one generation per year in the United States. This was the most common ceram-
bycid attacking windthrown spruce in the Adirondacks following a severe wind-
storm in November 1950.

Neoptychodes trilineatus (L.), the fig tree borer, breeds in the branches and
trunks of living and dying fig trees in the Southern States. Alder is also attacked
occasionally. The adult is gray except for small reddish-yellow spots on the body
and white stripes on the elytra, and is 18 to 25 mm long. Eggs are deposited in
small holes chewed through the bark. The larvae feed at first beneath the bark and
then bore into the heartwood, where they construct long tunnels. The life cycle
usually requires several years. Infested branches and small trees are often killed.

Tylonotus bimaculatus Haldeman, the ash and privet borer, breeds in a wide
variety of deciduous trees and shrubs, preferably ash and privet, in the Eastern and
Central States. Adults are dark brown and from 10 to 18 mm long. There are light
spots on the wing covers and a median line and two small, shiny spots on the
thorax.

Eggs are deposited at the base of privet plants and beneath bark scales on living
and dying ash trees during early summer. Young larvae feed in the phloem; later
they penetrate deeper and scar the wood. Large branches of ash are usually attacked
and killed before the trunk is attacked. Old, mature, and drought-ridden trees,
especially those growing in parks and windbreaks, are killed gradually, branch by

308

branch. Privet hedges are subject to severe damage. The larval stage extends over a
period of 2 years.

Encyclops caerulea (Say), the oak-bark scaler, bores in the outer bark of living
white oaks, yellow-poplar, maple, tupelo, black ash, and pignut hickory in the
Eastern States. Dry scales peel off over damaged areas of bark. The adult is slender,
metallic blue or green, has light-brown legs, and is 7 to 11 mm long.

Oeme rigida rigida (Say) breeds in dead and dying baldcypress and juniper
throughout the Eastern United States. The adult is light to dark brown and between
16 to 20 mm in length. Eggs are deposited beneath bark scales and the larva feeds
beneath the bark and in the wood, producing large quantities of granular frass.
Deadened baldcypress and rustic work constructed from improperly seasoned wood
are often severely damaged, causing the bark to peel off and sometimes causing
destruction of pieces of wood up to 8 cm in diameter.

The genus Prionus consists of a number of species of robust, black or brownish-
biack, shiny beetles from 25 to 45 mm long. The antennae have from 12 to 27
segments and are heavy and imbricated in the male. There are three spines, or teeth,
on each side of the prothorax. Several species are known to feed in the roots of a
wide variety of living fruit, forest, and shade trees in the Eastern United States.
Root rots, such as Armillaria mellea (Vahl ex Fr.) Kummer, and various species of
secondary boring insects often attack the damaged roots and kill the trees. Mature
trees growing in open stands, in well-drained gravelly soil and hillsides, in pastures,
and in heavily used recreational areas are especially subject to attack. Small trees
and shrubbery are occasionally killed by borers that cut off their roots at the ground
line.

Prionus species have been found to be associated with declining pecan trees in
Georgia and Alabama. Whether the larvae are primary or secondary invaders on
pecan is unknown. There is no known adequate method for controlling the larvae or
beetles (1/37).

The broadnecked root borer, P. /aticollis (Drury), breeds in the roots of a wide
variety of trees and shrubs in the Eastern United States. Mostly hardwoods are
attacked, including oak, pecan, poplar, chestnut, and basswood. Infestations have
also been recorded in the logs and stumps of all the above species. The adult is dark
brown, shining, and 20 to 45 mm long. The head is depressed between the eyes, the
antennae of the male are shorter than the body, and the base of the pronotum is as
wide as the elytra. Full-grown larvae may reach a length of 75 mm. Eggs are
deposited in groups on the ground and the young larvae crawl to the roots to feed.
At first they feed in the bark, but soon enter the root which is completely hollowed
out and occasionally severed. They move from root to root through the soil, feeding
on the surfaces of smaller roots as they go and causing many injuries or wounds.
Mature larvae come to within 8 to 13 cm of the soil surface in the spring and form
oval, compact cells in which to pupate. The life cycle requires 3 to 5 years. This
species seldom causes substantial damage in the forest.

The tilehorned prionus, P. imbricornis (L.), occurs throughout much of the
Eastern United States and breeds in the living roots of oak, pecan, chestnut, pear,
and various herbaceous plants. Adults (fig. 140) are dark brown, shiny, and 24 to 50
mm long. Male antennae have 18 to 22 jointed segments that overlap. Eggs are
deposited in groups in the soil around the base of the tree. Larval habits are similar
to those of the broadnecked root borer. The life cycle requires 3 to 5 years.

Prionus pocularis Dalman breeds in decaying pine logs and stumps throughout

309

aa

Courtesy Duke Univ. Sch. For.
Figure 140.—Adult of the tilehorned
prionus, Prionus imbricornis.

the Eastern United States. Adults are light brown, shiny, and 25 to 45 mm long. The
elytra are densely punctured.

Several species of Xylotrechus occur in eastern forests. Adults are characterized
by their moderate size, their short, filiform antennae, and a V-shaped callosity on
the front of the head.

Xylotrechus quadrimaculatus (Haldeman), the birch and beech girdler, breeds
in the branches of birch, beech, American hornbeam, maple, and alder in eastern
Canada and Northeastern United States. Adults are 7.5 to 14 mm long. The thorax
is black with four yellow spots; the wing covers, pale brown with faint white marks.
Eggs are deposited in the axils of twigs, in bark crevices, and in healed-over
injuries on branches. Young larvae feed beneath the bark, often girdling and killing
the branch quickly. Later they bore toward the pith in concentric circles until the
branch is almost severed. At this point? they turn and bore toward the tip, packing
their tunnels with granular frass. Before reaching the tip they construct pupal cells
in which they spend the winter and in which they pupate in the spring. Damaged
branches up to 5 cm in diameter frequently fall to the ground during midsummer
with the larvae in them. There is one generation per year.

The gallmaking maple borer, X. aceris Fisher, breeds in the trunks and
branches of various maples, especially small red maples, causing the formation of
galls. The adults resemble those of the birch and beech girdler, although they have
fainter spots on the thorax and stronger markings on the elytra. The adults are 10 to
14 mm long. Eggs are deposited during midsummer in wounds or at the bases of
small dead twigs along the trunk. The larvae bore directly into the sapwood. Later
they construct tunnels in the heartwood, often completely destroying the center of
the trees and causing the formation of galls or swellings about the wounds. During

310

the second summer of their lives, they bore either directly upward or downward and
pupate in cells at the end of their tunnels. Infested trees are never killed outright but
are seriously weakened, becoming subject to windbreakage.

Xylotrechus obliteratus LeConte, the poplar-butt borer, a serious pest of quak-
ing aspen and other poplars in the Rocky Mountain region, also occurs in many
parts of the Eastern United States. The adult is dark colored and about 10 to 18 mm
long. The thorax is crossed by yellow bands at the front and rear margins; the elytra
are crossed by three yellow bands—the first one oblique, the middle one curved,
and the last one transverse.

Adults are present during late summer. Eggs are deposited in irregularities of the
bark or exposed wood, and the larvae feed beneath the bark until fall. The following
year they bore into the wood where they feed for several years, much of the time in
parts of the tree below the ground line. Females continue to deposit eggs in the butts
of infested trees until the wood is completely honeycombed and the tree dies or is
broken by wind or ice.

Xylotrechus sagittatus sagittatus (Germar) breeds in dead conifers from eastern
Canada and the Northeastern States to Florida and New Mexico. In areas where
pines predominate, it is especially common. Logs, slash, and trees killed by fire or
bark beetles are particularly attractive. Adults (fig. 141) are dark brown and from
12 to 21 mm long. Each elytron is marked with variable stripes of fine gray hairs—
one stripe along the sutural margin, one along the outer margin, and one at the
apex. The larvae feed first beneath the bark, then they tunnel deep into the wood.

Courtesy Duke Univ. Sch. For.

Figure 141.—Adult of Xy/otrechus sagittatus
Sagittatus.

311

The rustic borer, X. colonus (F.), one of the most common of all cerambycids in
the Eastern United States, feeds under the bark of almost all dead hardwoods. It
occurs also in southern Canada. The adult (fig. 142) is light to dark brown and 8 to
15 mm long. Each elytron is marked with an irregular line of fine yellow hairs back
of the base and three transverse bands of gray hairs—one band just behind the
yellow line at the base, one back of the middle, and one at the apex. The larvae feed
almost exclusively in the bark without scarring the wood (463). Recently killed
trees are preferred.

Courtesy Duke Univ. Sch. For.

Figure 142.—Adult of the rustic borer,
Xylotrechus colonus.

Xylotrechus annosus annosus (Say) breeds in poplar and willow in eastern and
central Canada and in the Northeastern and Lake States. X. undulatus (Say) breeds
in recently cut spruce, Douglas-fir, and lodgepole pine from eastern Canada and the
Lake States to Yukon Territory and central Alaska.

The genus Monochamus is represented by several important wood-boring species
in eastern forests, all of which breed in various conifers. The larvae are commonly
known as “‘sawyers” because of the loud noise they make while feeding. Freshly
cut, feiled, dying or recently dead trees are preferred. Young larvae feed on the
inner bark, cambium, and outer sapwood, forming shallow excavations called
surface galleries which they fill with coarse, fibrous borings and frass. As they
grow older, they bore deep into the heartwood, and then turn around and bore back
toward the surface, thereby forming a characteristic U-shaped tunnel. A pupal cell
is formed at the outer end of the tunnel. from which the adult emerges by chewing a
hole out through the remaining wood and bark. Full-grown larvae are often more

312

than 50 mm long. Members of this genus often cause heavy losses in windthrown or
fire-killed timber, in saw logs left too long in the woods before milling, and in
improperly handled pulpwood. The adults are vectors of the pine wood nematode
(1340). A key to the adults of the genus 1s available (60/).

The southern pine sawyer, M. sitillator (F.), occurs throughout the Eastern and
Southern States and breeds in recently cut, windthrown, fire-killed, insect-killed,
and dying pines. The adult is mottled gray and brown, and is from 18 to 30 mm
long (fig. 143). Male antennae are often 2 to 3 times as long as the body, there is a
strong spine on each side of the thorax, and the elytral sutures are prolonged into
sharp spines. Full-grown larvae are up to 60 mm long.

Figure 143.—Adult of the southern pine sawyer,
Monochamus titillator, on shortleaf pine.

In the Piedmont area of the South, adult emergence reaches a peak in April and
May. However, adult activity continues until late fall and probably to some extent
throughout the winter. Larval habits are as described for the genus. There are at
least two generations per year in the South, with overlapping broods (/26/).
Prompt salvage and utilization of windthrown and dead and dying trees, debarking
recently dead trees, and water storage of logs will prevent attacks by this species.

The whitespotted sawyer, M. scutellatus (Say), occurs from Newfoundland to
North Carolina, Minnesota, and Alaska. Eastern white pine appears to be its
favorite host, but it also attacks many other conifers such as red and jack pines;
balsam fir; white, black, and red spruces; and larch. The adult (fig. 144) is about 18
to 25 mm long. The male 1s completely shiny black except for a small, rounded
white spot at the base of the elytra; females are of the same color or have the elytra
mottled with white spots. Two years are required to complete the life cycle in the
Lake States and Canada. Farther south there is one generation per year. Adults
emerge through circular holes cut in the bark and feed for short periods on needles
and tender bark of various conifers. Eggs are then deposited in slits or niches
chewed in the bark, preferably near old branch scars or in wrinkled areas on logs,
pulpwood, and recently killed trees (/034, 1319).

The whitespotted sawyer causes heavy losses to saw logs and pulpwood in the
Northern States and southern Canada. Damage to saw logs can be prevented by

313

Courtesy H. F. Cerezke, Can. For. Serv.

Figure 144.—Whitespotted sawyer, Monochamus
scutellatus: A, adult; B, larvae and damage.

cutting the trees between September and early June and removing them from the
woods before late June. Damage to pulpwood can be reduced by piling it in the
shade of standing trees or covering the piles with layers of slash 0.3 to 0.6 m thick.

The northeastern sawyer, M. notatus (Drury), occurs in eastern Canada and in
the Northeastern States, westward to the Great Lakes region, and breeds in dead
and dying eastern white pine and balsam fir and in windthrown red spruce. Adults
are dark brown and up to 30 mm long. The head and pronotum are irregularly
clothed with fine white hairs; the elytra are covered with fine gray and white hairs
arranged in the form of interrupted stripes. The female head 1s greatly flattened and
elongated.

The balsam fir sawyer, M. marmorator Kirby, breeds in true firs in eastern
Canada and the Northeastern States, west to the Great Lakes region and south to

314

North Carolina. Adults are dark brown, marbled with irregular bands of white and
yellow, and from 18 to 25 mm long. Recently felled trees are particularly attractive
to them. M. carolinensis (Oliver) breeds in dead and dying pines in the South-
eastern States. Adults are only about 17 mm long, otherwise they are very similar in
appearance to those of the southern pine sawyer. The spotted pine sawyer, M.
maculosus Haldeman, a common species in the Far West, also occurs in eastern
forests. It breeds in dead and dying pines. Adults are dark reddish to blackish with
patches of fine yellowish hair, and are about 16 mm long. The tip of each elytron is
prolonged into a tooth.

Orthosoma brunneum (Forster), the brown prionid, breeds in decaying co-
niferous and hardwood logs in southeastern Canada and throughout much of the
Eastern United States. The adult is light brown, flattened, and from 25 to 50 mm
long. The pronotum is narrower than the elytra and has three sharp spines on each
side. There are three fine, raised longitudinal lines on each elytron. The body of the
larva is slightly tapering and shines with a lemon or yellowish tinge.

Eggs are deposited from June to late summer in wood that has been dead for
several years, especially in wood with very high moisture content and containing
decay fungi. The larvae feed for 2 to 3 years, packing their tunnels with coarse,
fibrous frass. Crossties, structural timbers, poles, or other wood in contact with the
ground may be severely damaged or destroyed. Damage can be prevented by
keeping the wood dry or by treating it with a preservative before placing it in
contact with the ground.

Stenodontes dasytomus dasytomus (Say), the hardwood stump borer, occurs
from Virginia southward and westward and breeds in the heartwood of living
hardwood trees such as various oaks, sycamore, willow, and boxelder. Wood in
contact with the ground is also attacked. The adult is a large, somewhat flattened,
reddish-brown beetle from 23 to 47 mm long. The head 1s large and the sides of the
prothorax are armed with small, fine teeth. Eggs are deposited around wounds,
particularly near the base of the tree, and the larvae bore into the heartwood. They
feed there gregariously for 3 or 4 years, completely honeycombing the wood.
Shade trees are sometimes weakened so badly that they break and fall during
storms. Crossties and other wood products in contact with the ground are also
subject to serious damage.

Archodontes melanopus melanopus (L.) breeds in the roots of live and water
oaks, boxelder, persimmon, and hackberry from Virginia to Florida and westward
along the Gulf Coast. The adult is broad, rather flat, dark brown, and from 33 to 50
mm long. The head is distinct and rather large, and the edges of the prothorax are
finely toothed. Full-grown larvae are almost 90 mm long.

Eggs are deposited at the base of young trees, just below the ground line. The
larvae bore into the roots and excavate large, flattened galleries in the wood. Huge
galls form on infested roots and interfere with the growth of the tree. Heavily
infested trees may die and be replaced by clumps of bushlike suckers. It has been
suggested that this species was largely responsible for the creation of large,
comparatively barren areas of scrub oak in parts of southern Georgia and Florida.

Rhagium inquisitor (L.), the ribbed pine borer, occurs throughout the United
States and in southern Canada, and it breeds in the inner bark of various species of
dying conifers. The adult is black except for mottlings of reddish-brown or gray and
is about 9 to 21 mm long. The thorax is slender and bears a spine on each side. The
larvae are distinguished by their very thin, flat heads. Eggs are deposited in early
spring in crevices of the bark of trees that died or were cut during the preceding

315

winter. Trees dead for only a short time and containing considerable amounts of
moisture are preferred by the larvae. They feed entirely beneath the bark, excavat-
ing irregular galleries and packing them with fibrous frass. When they become full
grown they construct oval, fibrous-edged cells in which they pupate and spend the
winter as adults.

The old house borer, Hylotrupes bajulus (L.), is native to the Atlas Mountains
of northern Africa (/93). It was introduced into North America more than a century
ago. Serious infestations have occurred in the Atlantic and Gulf Coast States from
Massachusetts to Texas. Spot infestations have been found as far north as Wisconsin
and Minnesota and as far west as Nebraska. It breeds in dry, seasoned, coniferous
sapwood. Pine and spruce appear to be preferred, but hemlock, true fir, and
Douglas-fir are also attacked. The adult (fig. 145A) is a slightly flattened,
brownish-black beetle from 8 to 20 mm long. The head and forward part of the
body are clothed with gray hairs; the thorax has several small tubercles at the side
and a black line and two black spots on the disk; and each wing cover bears either
patches of gray that fuse to form two crossbands, or two whitish spots. Full-grown
larvae are wedge shaped, deeply segmented, and up to 30 mm long.

F-494430, 494432

Figure 145.—The old house borer, Hy/otrupes bajulus:
A, adults; B, damage.

Eggs are deposited in fan-shaped clusters or in rows and layers in holes or tight
crevices. Stacked lumber and cracks and natural checks in the wood of houses are
especially subject to attack. Young larvae feed near the surface of the wood; older
ones bore into the sapwood and seriously damage it with their frass-packed tunnels.
The larvae seldom break through the surface of the wood. Thus, timbers so severely
damaged as to be near collapse may appear from the outside to be perfectly sound.
The length of the life cycle is not exactly known. In the Southern States, from 3 to 5
years may be required. In the northern parts of its range, an additional 2 to 3 years
may be necessary. The length of the cycle may also vary considerably in a given
building. In attics, where generally warmer temperatures prevail, adults may
emerge up to 2 years earlier than those in wood in the basement.

The old house borer causes severe damage in houses and other buildings.
Structural timbers (fig. 145B), framing members, and other wood parts are se-
riously weakened by larval mining and tunneling. Air-dried pine floor joists, plates
or sills, and subflooring are apt to be damaged most severely. Other framing such as
studs, stair carriages, furring strips, and roof rafters are sometimes attacked.

316

When an old house borer infestation is discovered, two types of remedial action
are necessary: (1) repairing or correcting serious structural defects, and (2) control-
ling the remaining insects present in wood left in place (S70, 87/, 1305).

Chlorophorus annularis (F.) has been recovered from infested, imported bamboo
at several locations in this country, and it may be established here. The adult is
blackish, with green markings on the thorax and a yellow, X-shaped mark on the
elytra, and is about 10 mm long. The first two pairs of legs and the inner parts of the
antennae are red. Well-seasoned bamboo is mined extensively, and the mines are
tightly packed with fine, powdery frass. This species is a serious pest of bamboo in
India and Japan.

Superfamily Curculidnoidea
Snout Beetles

Snout beetles are a morphologically distinct group in the order Coleoptera that
were formerly placed in a suborder, Rhyncophora. Adults of this superfamily have
the area in front of the eyes elongated to form a rostrum or snout. But there 1s much
variation in the extension of the area. In one family, the Scolytidae, the area is only
slightly extended and might go unrecognized. The mandibles are at the end of the
snout. In addition to the snout, these beetles, with a few exceptions, are dis-
tinguished by having the gular sutures united on the median line; the palpi are short,
conical, and rigid. The labrum is absent.

The larvae have large sclerotized heads. The body 1s light colored and soft with
conspicuous transverse folds or ridges in both the thoracic and abdominal seg-
ments. They are legless. When removed from their galleries (the larvae of most of
the species in the superfamily are borers), they assume a characteristic C-shaped
posture. Many of our most destructive forest insects belong to this superfamily. In
the United States, 90 percent of all tree mortality is caused by insects; more than 60
percent of this is caused by scolytids (1356). Flowers, fruit, and seeds are also
destroyed by various species of snout beetles. Those species that do not kill trees
often severely reduce the quality of the wood because they bore into the stem, thus
producing crooked logs, or holes, or pitch pockets. Other species reduce the
quantity of wood either directly or by reducing the vigor of the tree by eating roots,
stem, branches, or leaves.

Most of the species in this superfamily, both in terms of numbers and the amount
of economic damage they produce, are found in two families: Curculionidae—the
weevils—medium-size beetles with elongate snouts, no tibial spines, and the
antennae inserted somewhere along the snout; and Scolytidae—the bark beetles—
small, cylindrical beetles without a prominent snout, tibiae with spines, and
antennae not much longer than the head.

But there are many interesting weevils that belong to a number of much smaller
families. Some of these families, such as the Anthribidae and Brentidae, have been
recognized as families for a long time. Other families such as the Nemonychidae,
Rhynchitidae, Attelabidae, and Rhynchophoridae, have been considered, until

recently, subfamilies of the family Curculionidae.
Family Anthribidae
Fungus Weevils

The fungus weevils are a relatively small family of about 90 species in North
America. Few are economically important. Adults of this family have a clearly
visible labrum that is not fused with the rostrum, which sets them apart from other
families. Other distinguishing features are: visible and movable palpi, the pygidium
is exposed, the third tarsal segment is bilobed and set into the apex of segment two,
the antennae are not elbowed.

317

The larvae are found in dead and dying twigs and branches, and frequently in
polypore fungi. Tropideres fasciatus (Olivier) larvae have been found in fungi
associated with dead sycamore. The adults have been found on sassafras and birch.
T. dorsalis (Thunberg) occurs in dead hemlock. Choragus zimmermanni LeConte
has been found in fungi on dead maple and oak. The larvae of this species

overwinter in the wood (23, 1/227).
Family Brentidae
Brentid Beetles

This family is composed of mostly tropical species. The snout of these weevils is
straight and projects directly forward. The antennae are not elbowed. The thorax is
longer than wide. The oak timberworm, Arrhenodes minutus (Drury), 1s the only
tree-infesting species that occurs in the Eastern United States. It breeds chiefly in
oak, beech, and poplar, and its life cycle usually requires 3 years.

Adults are dark reddish-brown with yellowish spots or bars on the elytra. They
vary greatly in length: males may be from 7 to 35 mm long and females may be up
to 25 mm long. The snout of the female is straight, narrow, and much longer than
the head; that of the male is short and broad with large mandibles (fig. 146). The
larva of this species is an exception to the legless condition usually found among
weevils. Timberworm larvae have vestigial two-jointed legs.

F-519917
Figure 146.—Adults of the oak timberworm, Arrhenodes
minutus. Male on left; female on right.

Adults are active from May through August. Both sexes are attracted to fresh
wounds, and females lay their eggs in the tissue around wounds. Although most
females lay their eggs on living trees, occasionally eggs are laid on dead trees, but
always in wounds made before the trees died. The larvae bore through the wood,
constructing galleries in all directions. The larval galleries cause serious degrade of
infested trees. Occasionally much of the timber in a stand is so badly damaged that

it is unfit for cooperage (/69).
Family Nemonychidae
Pine-Flower Weevils

This small family is represented in the West by the genus Diodyrhynchus and in
the East by the genus Cimberis. These weevils are characterized by having a

318

functional labrum, labium, and palpi. The gular suture is double, and the antennae
are straight. On the apex of the tibia there are two spines (/227). Adults are usually
dull red, less than 5 mm long, and have flat snouts that are about as long as the
thorax. They feed on the staminate flowers of various conifers. The eggs are laid in
staminate flowers and under bud scales on dead shoots where the larvae develop. C.
pilosus (LeConte) is found in Virginia pine and C. elongatus (LeConte) in jack pine
(547, 1200).

Family Rhynchitidae

Rhynchitids

Members of this family can be distinguished by the absence of the labrum; the
palpi are short and rigid; the antennae are straight, |1-segmented including a
distinct 3-segmented club. The prothorax is not margined, and the elytra generally
cover the abdomen. The mandibles are flattened and toothed on both the inner and
outer margins. There is a single small spur on the apex of the tibiae.

There are two genera with species commonly found in the forest:
Pselaphorhynchites with larvae that develop in dead twigs, and Eugnamptus with
larvae that mine dead leaves. The adults of P. aeratus (Say) are black with a brassy
or coppery cast. They are 2.1 mm to 2.8 mm long. It has been reported from most
States and southern Canada in the east. Oak and willow are the major hosts
reported. P. cyanellus (LeConte) is a larger species (2.3 mm to 3.5 mm) with a
more northern distribution: southern Canada west to Alberta and Saskatchewan and
south to Pennsylvania, Ohio, and Illinois. This species is also black, but has a
distinct bluish cast. The antennae and tips of the legs may be paler. Adults have
been collected on willow, birch, and oak (519). Species of Eugnamptus differ from
Pselaphorhynchites by having the pygmidium covered by the elytra. They are more
slender, longer weevils. Adults of EF. collaris (F.) are 3.5 mm to 4.7 mm long. The
weevils are predominantly bluish black and red, but an entire array of color varieties
have been described and named. Hickory and butternut have been reported as hosts.
E.. striatus LeConte eats hickory and walnut. An adult may be from 4.5 mm to 5.0
mm long, have a red head and thorax, and black elytra. Other species attack oak,

sumac, and dogwood (670, 983).
Family Attelabidae
Leaf-Rolling Weevils

Four or five species of this small family can be found on eastern trees and shrubs.
This family is closely related to the Rhynchitidae but can be distinguished from it
by the absence of teeth on the outer margins of the mandibles, and also by the two
large spurs on the inner margin of the front tibiae.

The species most likely to be encountered are Homoeolabus analis (Iliger) and
Attelabus bipustulatus F. on oak, hickory, and walnut; A. nigripes LeConte on
sumac; and Himatolabus pubescens (Say) on alder and hazelnut.

The interesting thing about this group of weevils is the preparation the females
make for the larvae. When an egg is to be laid, the female chews a slit in a leaf
blade on both sides near the petiole. She then lays an egg near the tip of the leaf.
The slits at the base permit her to fold the leaf along the midrib and then to roll it
toward the petiole, enclosing the egg in a neat leaf cylinder. The petiole is then
partially chewed through so that the leaf wilts and eventually falls to the ground.
The larva develops in the leaf roll, and pupates either there or in the ground. There

may be more than one generation a year (454, 520, 670).
Family Curculionidae
Weevils

This family is reported to contain more species than any other in the animal

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kingdom (625). More than 35,000 have been described worldwide, 2,500 in North
America (670). With a few exceptions, the weevils are vegetarian; some are gall
makers. Many are wondrously disguised, with cryptic color and form to mimic
other insects, bird droppings, or buds. In many cases, the technique of deception is
enhanced when disturbed weevils remain motionless for a long time or feign death
and fall to the ground.

Some of our most destructive agricultural pests belong to the Curculionidae: the
cotton boll weevil, the alfalfa weevil, and the grain weevils. Foresters have prac-
tically stopped planting eastern white pine in the Northeast because of the white
pine weevil.

With such a large number of entities involved, the interpretation of the systemat-
ics of the family presents great difficulties, and authorities are not always in
agreement. The sequence of species used here closely follows that of ‘*Cur-
culionidae of America North of Mexico” (670).

There are five subfamilies of ““broadnose” weevils that are characterized by
short, broad snouts, and an oval depression on the face of the mandibles that marks
the position of a pupal cusp. Many species of this group have larvae and pupae that
are subterranean. The cusp is thought to assist adult emergence. These larvae eat
roots. Consequently, those species that feed on the roots of woody plants are of
concern to foresters and nurserymen. A disproportionately large number of the
most common and economically important of these weevils have been introduced
from Eurasia.

The arborvitae weevil, Phyllobius intrusus Kono, is a Japanese species first
recorded on nursery stock in Rhode Island in 1947. [t is now known to occur in the
other New England States as well as New York, New Jersey, and Pennsylvania. It
feeds on many varieties of arborvitae, northern white-cedar and eastern redcedar.
Damage in nurseries as a result of root pruning may be severe when populations
build up over a number of years.

Adult weevils are 5.1 to 6.3 mm long, black and covered with minute, light
metallic-green scales and hairs. The legs and antennae are more sparsely covered.

Eggs are laid in the soil during May and June. The larval period usually extends
through the winter, but some individuals overwinter as pupae. The weevil pupates
among the roots in an earthern cell that may be as deep as 25 cm. Adults begin to
emerge about mid-May (663).

The arborvitae weevil is destructive in both the larval and adult stages but
primarily in the larval stage. Damage in nurseries may be severe where infestations
occur over a period of many years. In severely infested areas, well over 200 larvae
may be found feeding on the roots of a single plant. This results in severe root
pruning.

Phyllobius oblongus (L.), the European snout beetle, was first recorded in the
United States at Rochester, N.Y., in 1923. It has been reported in New Jersey,
Pennsylvania, and Michigan. The adults are brown and 4 to 7 mm long. They seem
to prefer the leaves of sugar maple reproduction, but also eat leaves of elm, birch,
cherry, and alder. Adults are present in June and July. There is one generation a
year. Mature larvae overwinter in the ground.

Another introduced species (Nova Scotia, in 1884), one often associated with P.
oblongus, is Sciaphillus asperatus Bonsdorff. This 5- to 6-mm-long ash gray
weevil has been found as far south as Maryland, and west to Michigan. It feeds at
night, primarily on sugar maple, but also on leaves of mountain and red maples,
yellow birch, hazel, and hophornbeam. Because of its habit of feeding on buds, this

320

species has been implicated in excessive forking of sugar maple. The adults also
feed on petioles and shoots. Females are active from June through September.

No males have been reported. The larvae of both these species feed on roots of
plants on which the adults are found (/089, /343).

All the species of the genus Otiorhynchus in North America have been intro-
duced. The two species of greatest importance to eastern trees and shrubs, the
black vine weevil, O. sulcatus (F.), and the strawberry root weevil, O. ovatus (L.),
are thought to have arrived early in the 19th century. Both species are par-
thenogenetic and flightless (/249).

Adults of the black vine weevil are brownish black and 9 to 12 mm long. The
prothorax is rough with rounded tubercles, each of which has a curved yellow hair.
The elytra are also tuberculate with scattered tufts of metallic-yellow scales. It feeds
on more than 100 plant species including yew, spruce, rhododendron, hemlock, and
grape (/249). The adults eat foliage, buds, and shoots, but the greatest damage is
done by larvae in nursery and container plantings. In summer and fall young larvae
feed on rootlets. The following spring older larvae eat the cortex of larger roots
causing severe injury or death to the plant. Often the injury is not apparent until the
plant is transplanted.

Adult emergence usually begins about mid-June. They are nocturnal feeders so
spend the day in secluded places, frequently in the litter beneath the host plant. The
eggs are laid on the ground. Larvae develop through the summer and following
spring and pupate in cells down to 20 cm below the surface (929). There is one
generation a year (//04).

The strawberry root weevil is a shiny dark-brown or black weevil. The elytra are
covered with fine yellow setae. It resembles O. sulcatus but is smaller, about 5.5
mm long. More widely distributed than O. sulcatus, it has been collected in most
States in the United States, including Alaska, and the provinces of Canada.
Arborvitae is the preferred host of the adults, and twig girdling may cause severe
injury. Hemlock is less often attacked. White, red, Scotch, and Swiss mountain
pines, juniper, and Norway, white, and Colorado blue spruces have been reported as
hosts. The larvae prefer the roots of hemlock (462). In some places this species may
have two generations a year. Occasionally, during the summer, large numbers of
adults invade dwellings causing much distress, but no damage.

Because higher populations of these two weevils can cause heavy losses in
nurseries and to ornamental plants, some control may have to be employed. Some
insecticides, for foliar application, have been registered for these weevils.

Polydrusus impressifrons (Gyllenhal), first recorded in New York State in 1906,
feeds on various hardwoods, chiefly willow, poplar, birch, apple, pear, and plum in
New York and Connecticut, and possibly adjoining states. The adult is rather
slender, 4 to 5.5 mm long, and uniformly light metallic-green. Eggs are deposited
around scars and beneath loose flakes of bark in roughened areas. The cut ends of
pruned branches and twigs of young trees are especially attractive oviposition sites.
After hatch, larvae drop to the soil and feed on roots. Pupation occurs in the soil,
and adults appear from mid-May to early June. They feed on developing buds,
foliage, and succulent shoots. In heavy infestations, defoliation may be severe, and
large numbers of stems may be girdled and killed. Damage is usually most serious
in nurseries (960).

Brachyderes incanus (L.), an introduced species, has been recorded from Mas-
sachusetts and Long Island. Its hosts are various species of pine. The adults feed on
the needles; and the larvae, on the roots. Adults are brownish, with metallic

321

reflections, and are from 8 to 11 mm long. This species is often a serious pest of
pine and spruce in Europe. So far, it has not been very injurious in this country.

Four species of the genus Graphognathus, whitefringed beetles, have been
introduced into the Southern States (/248). First observed in Florida in 1936, they
now range north to Virginia and Missouri (524). The adults feed on the foliage of
hundreds of species of plants, including such tree species as pecan, hackberry,
black tupelo, yellow-poplar, blackjack oak, hawthorn, and sassafras, but their
damage is of minor importance. Most of the injury caused by this group results
from larval feeding on the roots of plants. There are records of damage to tree
seedlings in nurseries and fields.

The female adult (no males have been found) is dark gray and about 21 mm long.
The body is densely covered with short pale hairs, with those on the elytra being
somewhat longer. The forewings are fused together on the inner margins and the
hindwings are rudimentary, thus the beetle cannot fly. Eggs are deposited in masses
covered and held together by a sticky, gelatinous substance that hardens upon
drying. They may be attached to plants or other objects at the ground line or in the
soil, just below the surface. Winter is usually spent in the larval stage, and there is
one generation per year (/368).

The twobanded Japanese weevil, Callirhopalus bifasciatus (Roelofs), although
first reported from Connecticut in 1920, was first collected in Philadelphia in 1914
(166). It is now widely distributed in Eastern United States. Only females have
been recorded here. They are stout weevils, 4.5 to 7 mm long, with the snout
almost as wide as long. The elytra are fused, brownish gray, with two dark bands.
The eggs are laid on freshly fallen leaves or leaf fragments that the female then
folds and seals to make a pod. An average of three eggs is laid in each pod. They
hatch in about 14 days; the larvae then burrow into the soil and begin to feed on the
roots. Adults begin to appear in September.

All developmental stages can overwinter, but usually it is the adult. The adults
become active in April (//). The larvae and pupae have been described (8/4).
Adults eat at the leaf margin of a great variety of trees and shrubs. Reported eaten
are such diverse plants as flowering dogwood, hemlock, rhododendron, oak, aralia,
maple, forsythia, viburnum, and poplar.

The Asiatic oak weevil, Cyrtepistomus castaneus (Roelofs), an introduced
species first recorded in North America at Montclair, N.J., in 1933, is now known
to occur from New Jersey to Georgia and Missouri. Oaks and Asiatic chestnuts
appear to be its preferred hosts but it also attacks hickory, beech, dogwood,
hazelnut, and azalea (/209). Adults are black to reddish brown, irregularly clothed
with minute green scales, and about 6 mm long.

The Asiatic oak weevil is parthenogenetic and has a l-year life cycle. Appar-
ently, eggs are laid in the soil from early July to mid-September. Larvae have been
found around the roots of oak seedlings at depths of 15 cm in the soil. Winter is
spent mostly in the larval stage. Adults appear in the spring and feed on the leaves
of sapling oaks and chestnut. They chew in from the margins toward the midribs
and devour everything but the larger veins. Later, they fly to larger trees to feed.
During the fall, they sometimes invade houses in large numbers, presumably in
search of hibernation quarters.

Species of the genus Hylobius are another group of weevils with larvae that feed
primarily on or near roots. These attack conifers predominantly. Often a complex of
Hylobius spp., Pissodes spp., and Pachylobius picivorus (Germar) will attack the

322

same tree. Seven species of Hylobius occur in North America. Keys to the adults
have been published (4/6, 86/).

The pales weevil, H. pales (Herbst), is the most serious insect pest of pine
reproduction in cutover pine land (75, 938). It is also a problem in young reforesta-
tion areas and Christmas tree plantations. Pales weevil occurs in North America
east of the Great Plains and north to Ontario. The adult is dark reddish-brown to
black, 7 to 12 mm long (fig. 147). There is a patch or line of yellow-white scales on
the head, and irregular patches of light scales on the elytra.

F-532346
Figure 147.—Adult pales weevil, Hylobius pales, feeding
on loblolly pine seedling.

Pales weevil breeds in all species of pine within its range. Pitch, white, loblolly,
and shortleaf pines are favored species. It has been reported from spruce, fir,
juniper, larch, hemlock, northern white-cedar, and Douglas-fir.

In the North, winter is spent as adults beneath litter or as larvae in roots. In the
South, adults may be active throughout the winter but are in reproductive diapause
(214). Depending on location, those adults in hibernation emerge from March to
June. The adults are active at night and hide during the day in the soil and litter
around seedlings and saplings on which they have fed. After a brief period of
feeding they fly to cut, damaged, or recently dead pines (572). Here they feed and
mate and the females lay their eggs in the roots. Sometimes they may burrow as
much as 30 cm in search of roots. The larvae feed downward in long tunnels under
the bark and pupate in cells in the outer sapwood. Pupation and emergence may
take place in the late summer or fall; or the larvae may overwinter, and pupation and
emergence occur in early summer the following year (133, 317, 415, 1139). In the
North there is usually one generation a year, although some adults may live for 2
years. In the South there may be a second generation if weevils emerging in late
summer or fall oviposit before winter. Adult populations peak in March through
May and again in July and August (4/5, 975, 1190).

The most serious damage pales weevil does is in natural and planted seedlings in
recently cutover pine areas. Weevil-caused mortality among first-year seedlings of
30 to 60 percent is not uncommon, and mortality exceeding 90 percent has been
recorded (938). First evidence of attack is a series of small holes chewed in the bark
by adults. On lightly infested trees these may fill with oleoresin and heal over. With

323

heavy feeding, the damaged areas merge and may girdle and kill seedlings up to 1
cm in diameter. The seedling may be completely stripped, leaving a bare, curled
stem surrounded at the base by a pile of detached needles. Adults may feed on
terminals and twigs on large trees. Damage is not serious although the ends of
branches may be killed. In Christmas tree plantations, twig feeding may deform.
trees thus lowering their value to the extent of the damage (242).

Damage to young pines in cutover areas can be minimized by cultural practices.
In the South, land cutover and site prepared before July can be planted the
following winter without weevil damage. Weevils leave these areas before the
planting season. In areas cut or site prepared in July or later, planting should be
delayed | year. In the North and in the southern Appalachians, the waiting period
should be | to 2 years. When delay of planting is undesirable, seedlings should be
chemically treated (934, 938, 1140, 1242). In the South where stands are to be
established by direct seeding, no delay is necessary because weevils will have left
the area before seedlings are large enough to become suitable food. In the North, a
l-year delay of direct seeding may be necessary.

There are six ways to prevent or control pales weevils in Christmas tree planta-
tions: (1) delay planting | to 2 years if land was previously used for Christmas
trees, (2) treat seedlings with insecticide, (3) remove stumps in spring, (4) treat
stumps in spring with insecticide, (5) spray trees with insecticide from mid-August
to mid-September to control weevil feeding on branches, and (6) leave a whorl of
live branches on the stump to keep the stump alive and unattractive to weevils (242,
938).

The pine root collar weevil, H. radicis Buchanan, breeds in the root collars of
healthy pines in southern Canada from Newfoundland to Manitoba and in all of the
Northeastern States south to Virginia and west to Minnesota. Of its hosts, Scotch
pine seems to be the most severely damaged; but jack, red, Austrian, eastern white,
pitch, and Swiss mountain pines are also attacked. The adult (fig. 148) is dark
reddish-brown to black. It lacks the line or spot of pale scales on the head that H.
pales has. The body is marked with irregular patches of white to yellow hairlike
scales, and the elytra have longitudinal rows of elongated indentations. It is a larger
weevil than H. pales—its body length ranges from 9.5 to 13.0 mm (4/6).

-F-488105

Figure 148.—Adult of the pine root collar weevil,
Hylobius radicis.

324

In most localities, eggs are deposited from May to September in cavities in the
inner bark at the base of the tree, or in the soil nearby (/064). Peak egg laying
occurs in June (/332). The larvae feed downward in the inner bark of the base of the
trunk and in the base of large roots below the ground line, widening their galleries
as they develop. Galleries are also formed in the soil around the base. In the Lake
States, the winter is spent in the larval stage. In southern Ontario, it 1s spent in the
larval and adult stages and occasionally in the pupal stage (4/7). Pupation occurs
from June to September in cells constructed in the soil. Adults appear from August
to October but do very little egg laying before entering the soil or bark crevices to
spend the winter. When they emerge in the spring, they feed during the day on the
bark of duff-covered branches, mate, and lay eggs. During warm evenings, they
feed on the bark of the upper branches and fly to other trees. Most of these adults
overwinter a second time and then are active for part of the following season
(1338).

Heavily infested trees may be severely injured (fig. 149). Small trees, from 2.5 to
10 cm in diameter, are most severely damaged; smaller ones are seldom attacked.
The presence of pitch flows at the root collar and layers of pitch-infiltrated soil near
damaged areas are evidences of infestation. A number of silvicultural practices have
been suggested for reducing populations (/327). These include avoidance of sus-
ceptible species such as Scotch and red pines, particularly in mixed plantings;
shallow planting and maintenance of fully stocked stands. Weevils in plantations
can also be reduced by pruning the lower whorls of branches and removing the litter
from around the hole. This treatment was effective for up to 5 years (/33/). In
high-value plantations, insecticide should be applied to the base of the trees in late
May and June before egg laying begins (5/6).

ETA.
F-488104
Figure 149.—Damage by the pine root collar weevil,

Hylobius radicis, at the base of a pine tree.

325

The pine root tip weevil, H. rhizophagus Millers, Benjamin, & Warner, is a
species attacking jack pine in the Lake States (86/7). Red, Scotch, and eastern white
pines can be affected. The adult is shiny black and between 8.0 and 11.7 mm long.
Dense patches of coarse scales occur on the dorsum, and the elytra bear longitudi-
nal rows of pits containing fine setae. The adults are nocturnal. Eggs are laid mostly
in June near root tips down to 75 mm. The larvae feed in roots less than 12.5 mm in
diameter, tunneling from the smaller end toward the base. They complete develop-
ment in June and July of the second year. Pupation occurs in cells in the roots and
lasts about 20 days. Adults live 2 years. Pole-size pines in closed plantations on
formerly cultivated land are most frequently infested; reproduction in well-estab-
lished infestations, and red pine mixed with jack pine are attacked occasionally
(658).

Hylobius warreni Wood, Warren’s collar weevil, attacks most species of con-
ifers growing on moist to wet sites in southern Canada and south to North Carolina
in the Eastern United States. The adult is a large, robust, reddish-brown to black
weevil, from 12 to 15 mm long. The wing covers are thick, tough, and veinless,
and each bears 10 rows of longitudinal punctures. The hindwings are vestigial.
Eggs are deposited around the root collars of healthy trees, and the larvae feed in
the inner bark and cambium of roots and root collars for | to 2 years. Infested trees
bleed heavily at the ground line and severely injured ones may be completely
girdled and killed. Up to 40 percent of the trees in a 40-year-old Scotch pine stand
in Quebec is reported to have been killed by the species. The species has been
described in detail (/250).

Hylobius pinicola (Couper), Couper’s collar weevil, occurs throughout the
range of H. warreni and apparently breeds in the same species of trees. The adults
of the two species are similar in appearance, but the hindwings of H. pinicola are
fully developed.

Hylobius congener Dalla Torre, Schenkling, & Marshall occurs from the north-
eastern coast to Alaska, and in the East breeds in the inner bark of logs and stumps
of red, Scotch, and eastern white pines. It will also attack larch and spruce. Adults
range from 5.8 to 9.0 mm in length. The body is brown to black, the thorax without
scales, but the foretibia of the male has a fringe of white hair.

Adults emerge from hibernation in May and eggs are laid in logs and stumps.
Logs seem to be preferred. Larvae become prepupae in chip cocoons in August and
September and overwinter in the prepupal stage. The adult stage occurs in July and
August. They then feed on slash and logs until hibernation (822). Recently this
species has been found to attack seedlings in plantations causing up to 40 percent
mortality. Removal of litter from around seedlings has reduced damage (606).

The southern pine root weevil, H. aliradicis Warner, was discovered attacking
the roots of healthy young pines with root-collar diameters of 0.5 to 5 cm in
southern Georgia (36/, 1247). The larvae hollow out the smaller roots and bore
extensively in the root collar. Heavily infested trees are killed. The most serious
infestations are in plantations up to 4 years old on disturbed sites.

The pitcheating weevil, Pachylobius picivorus (Germar), occurs throughout the
Eastern United States, but is most common in the South. It attacks various species
of pine, including shortleaf, loblolly, and slash pines. The adults feed on the inner
bark of small twigs of residual pines and on stems of small pine seedlings (fig. 150).
Larvae bore in roots of recently cut or killed pine trees. Damage is often serious in
recently cut pine areas, especially where a new crop of seedlings is established
during the first year after cutting the old stand. Adults are dark brown, robust, and

326

clothed with patches of short, flattened, yellowish or reddish-brown hairs. The
tibiae are thick with the outer part enlarged; the tarsi are densely hairy underneath
with the third segment bilobed.

F-532347
Figure 150.—Adult pitcheating weevil, Pachylobius
picivorus.

Adults are attracted to and breed beneath the bark of dying pine stumps or roots.
To lay eggs, the female bores directly down through the soil to green roots,
sometimes to a depth of | m. Eggs are deposited in the cambial region of roots as
small as 5 mm in diameter. Burrowing may occur as far as 8 m from the stump.
Small piles of large soil particles surround the entrance holes into the soil. The
newly hatched larvae feed beneath the bark, packing their mines with fine, red,
boring dust and frass. The surface of the sapwood may be etched, depending upon
the number and size of the larvae and the size of the root. Mature larvae excavate
cells and construct chip cocoons in which to pupate.

When there are many larvae, these cells are formed in the wood. Adults emerge
in 6 to 11 months, depending on the season in which the tree was cut. Then they fly
to newly cut areas where they feed and repeat the cycle. Most adult feeding occurs
at night on seedlings near fresh stumps, but some feeding also occurs on cloudy
days (444, 1193).

Damage varies with the season, and with the size and vigor of the tree. All of the
buds, bark, and foliage may be removed to the ground. Roots as deep as 20 cm may
be damaged. Attacks on newly established seedlings are usually severe. Mortality
is particularly severe among seedlings planted within 3 months after overstory
pines are cut. Heaviest attack with resulting seedling mortality occurred in eastern
Texas between March and June. Later, in July and August, very little feeding occurs
and damage mainly consists of puncture wounds and the removal of patches of bark.
Feeding in autumn is very limited.

Pine seedlings may be planted safely during the winter in eastern Texas on areas
cut earlier than the previous July. Farther north, a longer wait may be necessary.
Seedlings should be chemically treated before planting on areas cut within 6
months. Stands may be cut during seed years without danger of excessive losses to
regeneration (934, 1193).

The bark weevils of the genus Pissodes attack both living and dead conifers.
There are about 30 species in North America; their taxonomy is not altogether clear.

32]

The white pine weevil, P. strobi (Peck), is able to feed and breed in a large
number of spruce and pine species, both native and exotic. As a consequence, its
range extends from coast to coast (//20). In eastern North America, eastern white
pine is the most common host, but Norway spruce, wherever it has been planted, is
attacked with almost equal severity.

The adult (fig. 151A) is a small brown weevil 4 to 6 mm long. The elytra and
body are covered with irregularly shaped patches of brown and white scales. Near
the apex of each elytron there is a large white patch and a brown patch. In P. strobi
these two patches are close together, and brown and white scales are often found
intermixed.

DA

F-482574, 482573, 482572

Figure 151.—White pine weevil, Pissodes strobi: A,
adult; B, infested terminal of a young eastern white
pine; C, eastern white pines deformed by white pine
weevil attack.

328

The adult weevils spend the winter in the litter. They emerge from hibernation
during March, April, and May in response to temperature, and feed on the cambial
tissue of the main stems of the host. Most of the feeding is done within 25 cm of the
terminal buds. The eggs are laid in some of the cavities made'in the bark by the
females when they feed. Eggs hatch in 7 to 10 days and the young larvae bore
downward in the cambium under the bark. When there are many larvae they feed
side by side in a ring around the stem. Toward the end of July the larvae begin to
make chambers in the woody part of the stem. The shreds of wood removed are
placed on top of the chamber to form a chip cocoon. The larvae pupate in the
chambers and 10 to 15 days later emerge as adults. The adults disperse over all
parts of the tree and feed until late fall; this differs from their spring-feeding habit.
Sometimes, this feeding alone is severe enough to kill entire shoots.

The white pine weevil is the most serious insect pest of eastern white pine in
North America. Its damage results in two types of loss: (1) reduction in volume,
and (2) lumber degrade in the remaining volume. Studies in New Hampshire
showed an estimated volume loss of 40 percent in the saw-log portion of sawtimber
trees and 70 percent loss in the portion above saw-log limits of merchantability. The
average loss in pole-size trees was 13 percent (/254).

The first evidence of attack in the spring is pitch flow from feeding punctures on
the terminal shoots (534). Later, the new growth appears stunted, and finally, the
needles wilt (fig. 151B). Trees up to | m tall may be killed. Killed terminals on
taller trees are replaced by one or more branches of the topmost living whorl
assuming vertical growth. This results in crooked or forked stems (fig. 151C). Trees
suffering this type of damage for several years become multiple-stemmed, cabbage-
shaped, and worthless.

The white pine weevil has been studied intensively, and much has been learned
about its biology, ecology, and control. As a result of these investigations, the
following management practices have been recommended for reducing losses to
white pines: (1) planting white pines with hardwoods or under a hardwood cover;
(2) planting white pine on medium soils only, where soil matting or hardpan does
not occur within | m of the surface and where the trees will not suffer from
competition with hardwoods, or jack or red pines; (3) selecting and pruning the
least injured pines for an improved final crop in heavily infested stands; and (4)
removing less desirable trees from white pine stands (236, 309, 762, 523, 1362).
Other types of indirect control, such as that exerted by insect parasites, predators,
and birds, are helpful in preventing excessively high weevil populations but are
incapable of preventing intolerable levels of loss (525, //91). Key factor analysis
suggests that larval, pupal, and winte’ submortalities are the major influences on
population change (308). Drench-spr.y of terminals at the time adults emerge from
hibernation provides protection (5/6).

The northern pine weevil, P. approximatus Hopkins, a close relative of the
white pine weevil, occurs from the Atlantic Coast to Manitoba in Canada and
southward to Minnesota and North Carolina in the United States. Its preferred hosts
appear to be red and Scotch pines, but it also attacks eastern white, pitch, jack,
shortleaf, Virginia, Table Mountain, and Austrian pines, and red, black, and white
spruces.

The adult is similar to the white pine weevil. It is larger, and the snout is
proportionately longer. The distal brown and white spots on the elytra rarely overlap
in P. approximatus. This is the single most reliable morphological feature by which
these two species can be distinguished from one another (4/4). The taxonomic

329

position of P. approximatus and P. nemorensis (Germar), discussed later, remains
unclear. With the use of multiple discriminant analysis, these two species and P.
strobi can be separated on the basis of external morphology (490).

Winter may be spent in the larval, pupal, or adult stage, but most overwinter as
adults in the duff and topsoil beneath infested trees. Occasionally, they may be
found under scales or in the crevices of the bark of these trees. In the spring the
weevils leave their hibernation sites and feed for several weeks on the inner bark of
pine branches and on the stems of seedlings and small trees. Individuals that
overwinter as larvae and pupae complete their development and emerge as adults in
July and August.

The insect breeds beneath the surface of the bark on recently cut stumps and logs,
and on the main stems or branches of dead or dying trees. Attacks occur on the tree
from the roots up to branches as small as | cm in diameter. Attacks also occur at the
root collar and on the lower stems of apparently healthy young trees. Eggs are
deposited in pockets chewed through the bark by the female.

A pocket usually contains only one egg, but as many as five can be found. A
larva may feed in any direction from the site of the egg, but usually follows the
grain of the wood. Larvae pupate in chip cocoons usually on the surface of the wood
but the chamber can be | to 2 cm below the surface.

Damage is often severe in regions where there are large quantities of breeding
material, such as fresh stumps in Christmas tree plantations and in stands under
intensive management. Because of the scarcity of breeding material in natural pine
stands, damage there is of little consequence. Damage in nurseries and plantations
can be reduced by destroying breeding sites or rendering them unsuitable for egg
laying (420).

The deodar weevil, P. nemorensis Germar, occurs in the Southern States north to
Pennsylvania, and breeds in the true cedars, deodar cedar, Atlas cedar, cedar of
Lebanon, and some pines, notably loblolly, shortleaf, and longleaf. The adult (fig.
152) is similar to P. approximatus and can be distinguished by discriminant
analysis (490). Damage is of three principal kinds—feeding on young shoots of the
crown in sapling and pole-size trees, terminal killing by both adults and larvae, and
branch-end flagging on pole-size and small saw-log-size trees. Attacks have been
made directly on boles of trees 6 to 12 m tall, which killed the trees (948, 952).

4

Courtesy Duke Univ. Sch. For.

Figure 152.—Adult of the deodar weevil, Pissodes
nemorensis.

330

The adults are active all winter and lay eggs during this time. Eggs are deposited
in the inner bark through holes chewed in the bark. The larvae feed beneath the bark
in a manner somewhat similar to that of white pine weevil larvae, girdling and often
killing the stem. Evidence of their presence is the swelling of the bark over feeding
areas. Pupation occurs in chip cocoons in the wood. Adults apparently aestivate
during the summer months. They again appear in the fall to feed on the twigs and
leading shoots. Apparently, depending on location, all stages can be found through-
out the year. May is the month of greatest adult emergence.

Several other species of Pissodes also attack various coniferous species in eastern
forests. P. affinis Randall breeds in the stumps and logs of eastern white, red, jack,
and Scotch pines from New England to the Lake States. The adults feed on the
inner bark of branches of living trees up to 17 m tall. They are dark brown or black,
from 5 to 8 mm long, and are marked with white spots. P. dubius Randall, balsam
bark weevil, breeds in windthrown, dying, or recently dead balsam fir and red
spruce. It is probably the most important insect attacking dead and dying balsam fir
following spruce budworm defoliation (//82). It also commonly attacks balsam fir
weakened or killed by the balsam woolly adelgid. P. rotundatus LeConte, small
spruce weevil, and P. fiskei Hopkins breed in windthrown red spruce.

The New York weevil, /thycerus noveboracensis (Forster), 1s the only species of
the subfamily Ithycerinae known in the East. It is sometimes placed in a separate
family, Belidae. The adult is one of our largest weevils, 12 to 18 mm long. The
body and elytra are covered with prostrate ash-gray and pale-brown hairs that give
the insect a mottled appearance. It is widely distributed in the Eastern and Central
States, but is most abundant in the Mississippi River Valley (304). Adults have been
collected from oak, hickory, beech, and a number of fruit trees. The larvae are
thought to breed in twigs and under branches of oak (/027).

The pine gall weevil, Podapion gallicola Riley, occurs throughout the Eastern
United States and southeastern Canada. It breeds in red, pitch, and Virginia pines.
The adult is black and about 5 mm long. During June, eggs are laid in niches
chewed into the bark of 1-year-old twigs. Young larvae feed first on the sides and
floor of the egg niche and then bore into the cambium. Here they separate and
tunnel outward from the niche. They continue to feed through three seasons.
Pupation occurs in funnel-shaped cells in the bark during May of the fourth season.
Galls are formed by hypertrophy of the xylem tissue surrounding each larva. The
galls first appear as slight swellings on one side of the stem (/333). By the time the
adults emerge, these swellings are larger, generally ovoid, and taper gradually
toward the distal end (fig. 153). Old galls continue to enlarge, even after the insects
leave them, some reaching a length of 37 mm. When several galls are formed on a
small branch, the branch may be killed (/323).

The members of the genus Magdalis are small cylindrical weevils frequently with
toothlike spines at the anterior corners of the pronotum.

The black elm bark weevil, M. barbita (Say), breeds in the trunks and branches
of unhealthy elms from North Carolina to southern Canada. Adults are jet black,
have long slender beaks, and are about 6 mm long. They emerge in May or June
and deposit their eggs in the bark. The larvae feed in the inner bark and sapwood,
constructing galleries up to 4 cm long. Pupation occurs in oval cells just beneath the
bark. There appears to be one generation per year. The bark of heavily infested trees
may be literally peppered with small, circular, emergence holes.

The red elm bark weevil, MM. armicollis (Say), occurs in the Eastern States and
southern Canada and breeds in dying or recently dead elms or occasionally in dead

331

F-501808

Figure 153.—Galls in pine produced by the pine gall
weevil, Podapion gallicola.

branches of living, suppressed elms. The adult (fig. 154) is reddish and somewhat
smaller than the adult of the black elm bark beetle. Eggs are deposited in punctures
in the bark, often in groups around knots or at the bases of twigs. Larval tunnels
usually radiate away from the egg-laying site and follow the grain of the wood,
scarring both the inner bark and the wood. Pupation occurs in cells at the end of the
tunnels. There is one generation per year (554).

Magdalis perforatus Horn breeds in the dead and dying branches of pines from
Canada to Florida. The adult is bluish black, wedge shaped, shiny, and from 4 to 6
mm long. Eggs are deposited singly in pits chewed in the base of needle fascicles at
or near the tips of branches. The larvae bore to the center of the stem and then
tunnel through the pith toward its base. Trees under 4 m tall killed by insects or
disease appear to be preferred; however, slash left during thinning or pruning
operations and the dead, lower branches of old trees in closed stands are also
attacked. Adults feed on the new shoots of pines, usually on trees growing in young,
open stands. Prepupae overwinter in the pith. Pupation occurs in May, and adults
emerge in June (S22).

F-519947
Figure 154.—Adult of the red elm bark weevil, Magdalis
armicollis.

o32

Several other species of Magdalis are also encountered in eastern forests: M.
austera Fall and M. hispoides LeConte are found on the needles of eastern white
pine in the Northeast (987). The adults bore through the scales of succulent, young
needles, causing the distal portions of the injured needles to turn yellow and break
off. M. austera substriga Fall is found on young Scotch pines; M. olvra (Herbst), on
weakened hickories; M. salicis Horn, on willow and chestnut; and M. barbicornis
(Latreille) and M. pandura (Say), on elms.

The genus Curculio is represented in eastern North America by 15 species (478).
These attack the seeds of nut-bearing trees. Most attack acorn, but several species
are important pests on hickory, chestnut, and pecan. Injury is caused by the larvae
eating the contents of the nuts (/50). Damage can be severe; the entire seed crop of
some tree species is destroyed in some years. C. caryae is considered the major
insect problem in commercial pecan culture (/83).

The life history of curculios is essentially the same for all species. From 2 to 3
weeks before the nuts ripen, the adults appear. The female lays her eggs in
chambers chewed in the nut near the inner surface of the shell. Usually one egg is
laid in each chamber. After hatching, the larvae feed for several weeks. When
mature, they leave the nut to form pupal cells underground. However, pupation is
usually delayed | or 2 years and sometimes as long as 5 years. The pupal stage lasts
up to 3 weeks and maturation of callow adults may take another 20 days.

In the adult stage, species of the genus can be differentiated by snout shape and
length, characteristics of the mesosternal intercoxal process, antennal insertion, leg
structure, and hair and scale pattern (478).

The pecan weevil, C. caryae (Horn), breeds exclusively in the nuts of the genus
Carya. Except for the more northerly and eastern portion, this weevil is found
throughout the range of hickories. Adults are dark reddish-brown and densely
covered with golden to dark-brown hairs and scales. The body length varies from
7.5 to 12 mm. The female snout is longer than the body and down-curved in the
apical quarter of its length. The male snout also curves down but 1s rarely longer
than one-half the body length. Adults emerge in July and August. Eggs are laid in
maturing nuts and the larvae leave the nuts in September and October, sometimes
when the nuts still cling to the tree. Most larvae spend two winters in the pupal cell
before pupation. C. caryae is a serious problem in commercial pecan orchards.
Early maturing cultivars offer some promise of reducing damage; however, weevil
emergence, dependent on rainfall, is erratic, so in some years the crop is heavily
infested (/84). Most orchardists depend on insecticidal control. A number of
compounds are registered for use (966).

The large chestnut weevil, C. caryatrypes (Boheman) (the largest Curculio in
the East), and the small chestnut weevil, C. sayi (Gyllenhall), breed exclusively in
chestnut. At one time they were common, but since the passing of the American
chestnut they have become rare. They have been reduced to remnant populations in
orchards and isolated trees of introduced chestnut species, but they can be a serious
problem that requires insecticidal control in these instances (968).

The hazelnut weevil, C. neocorylus Gibson, breeds exclusively in hazel species.
It is a dark reddish-black weevil with somewhat lighter legs and antennae. The body
is covered with yellow to gray scales and hair that may have bands of darker scales
on prothorax and elytra. They are 6.0 to 7.5 mm long.

All other species of Curculio found in the East feed on oak. Some, like C. fulvus
Chittenden, utilize only one species, live oak; others such as C. nasicus (Say) and
C. iowensis (Casey), use three or four species. At the other extreme are species like

333

C. sulcatulus (Casey), which feeds on almost all oak species but prefers the red oak
group, and C. proboscideus F., reported from more than 20 species of oak (/230).

The willow flea weevil, Rhynchaenus rufipes (LeConte), occurs in eastern
Canada and south and west through the Eastern States to New Mexico. Willow is its
preferred host, but it also feeds on many other hardwoods such as elm, red maple,
aspen, red oak, gray and paper birches, cherry, serviceberry, and apple. The adult
is black, broadly elliptical, and about 2 mm long. The eyes are large and almost
meet in front, the antennae are reddish yellow and elbowed, the scutellum is white,
the legs are reddish yellow, the hind femora are thick and fitted for jumping, and
there is a small white spot at the base of the wing covers.

In Maine, winter is spent in the adult stage beneath loose bark, under stone walls,
in debris, or in the soil. Overwintering adults emerge in early spring, fly to their
hosts, and feed by eating circular holes in opening buds and new leaves. Eggs are
laid in pits on the underside of leaves, and the larvae feed almost entirely within the
tissues of the leaf, forming large blotch mines. Pupation occurs within the mine and
new adults appear in August. In heavily infested areas, they may crawl over
buildings in such large numbers as to be a nuisance (905).

The apple flea weevil, R. pallicornis (Say), feeds on apple, hawthorn, winged
elm, hazelnut, and quince from New York to Hlinois and Missouri. Adults are black
and about 2.5 mm long.

Odontopus calceatus (Say) has been reported from most Eastern States wherever
its host plants—yellow-poplar, magnolia, and sassafrass—are found. Adult weevils
are 2.5 to 3.9 mm long, and from 1.4 to 2.4 mm wide. Most are black with dark-
brown antennae, mouth parts, and tarsi. Southern specimens may have yellow tarsi
and antennal segments.

In the Gulf States, adults emerge in early February, but not until late April or
early May in the northern reaches of its range. At first they feed on buds and
stipules. Later, they feed on expanding leaves producing typical oval or rice-shaped
holes (fig. 155). The females lay their eggs in the leaf midrib on the underside of
leaves. Upon hatching, the larvae feed in the leaf blade; as many as 19 larvae have
been reported in a single mine. When feeding is done, the larvae weave spherical
cocoons in the mine and pupate. New adults emerge during the summer months.
After a brief period of feeding, mostly on the under surface of leaves, the adults
enter aestivation. Then, apparently with no further activity, they hibernate in the
leaf litter. Late spring frosts can cause high mortality in the northern parts of the
weevil’s range (177).

This species has been particularly abundant on yellow-poplar in eastern Ken-
tucky since 1960. Heavy infestations have also occurred in Ohio, West Virginia,
Virginia, and Tennessee. Prior to 1960, it seems to have occurred most commonly
on sassafras and magnolia. Parasites have destroyed up to 50 percent of the pupae in
certain areas of Kentucky.

The poplar-and-willow borer, Cryptorhynchus lapathi (L.), is a native of
Europe and Asia. It was first reported in North America in New York in 1882.
Since then it has spread across the continent in southern Canada. In the East, it is
found as far south as South Carolina. Willow is its primary host, but poplar, except
quaking aspen, is attacked and species of birch and alder are occasionally infested.
Infestations in plantations and ornamentals may require insecticidal control to
prevent serious damage.

The adult is 8 to 10 mm long (fig. 156). The snout is as long as the head and
thorax combined, and the elytra, at the base, are a third wider than the thorax. The

334

. >

7

F-519311

Figure 155.—Adult feeding damage to yellow-poplar leaf
by Odontopus calceatus.

F-519950

Figure 156.—Aduit of the poplar-and-willow borer,
Cryptorhynchus lapathi.

antennae and tarsi are reddish brown, the body black. The body is covered with
black and yellow or white scales intermixed with erect black bristles. The white
scales predominate on the apical third of the elytra.

Adults appear in late July and August and feed on the inner bark of young shoots.
Eggs are deposited singly or in groups of two to four in slits cut in corky bark, often
in lenticels and scar tissue. Young larvae feed in the cambial region and outer layers
of sapwood. They tunnel in all directions and push their borings out through small
holes. In late fall, they hollow out small chambers in the inner bark, and there spend

335

the winter. Feeding is resumed in the spring. The larvae usually bore around the
branch or stem. When ready to pupate, the larvae bore upward and construct cells in
the center of the stem. There is one generation per year.

The first evidence of attack is the occurrence of dead patches or cracks in the bark
on the trunks of small trees or on the branches of larger ones, and the presence of
small holes chewed in the bark. The wood under these patches eventually becomes
honeycombed with larval tunnels. Branches and small trees may be completely
girdled, or so badly weakened by tunnels that they break. All poplars and willows
over 2.5 cm in diameter are subject to attack, especially recently planted trees and
nursery stock (530, 1096).

The genus Conotrachelus, like the genus Curculio, contains a number of species
important as pests of acorns, nuts, and fruit. The adults of Conotrachelus resemble
those of Curculio, but tend to be smaller. They have shorter and less curved beaks,
the body is scaled on the upper surface, and the prosternum is grooved for reception
of the beak.

Probably the most economically important species is the plum curculio. C.
nenuphar (Herbst). It breeds in plums, cherries, peaches, apples, and other fruits.
This weevil is a serious pest in commercial orchards, where repeated applications of
insecticides are required during the first growing season for control. A number of
insecticides are registered for use. The quince curculio, C. crataegi Walsh, is also
a pest in orchards. It attacks quince, pear. peach, and hawthorn fruits. Some races
of this species that attack apple have been found (8/3).

There are three acorn-infesting species in the United States. C. naso LeConte can
be distinguished from the other two by the absence of femoral teeth: C. carinifer
Casey has prothoracic punctures larger than elytral punctures: and C. posticatus
Boheman has prothoracic punctures that are not larger than those on the elytra
(477).

Conotrachelus naso occurs from Maine to Florida and west to Minnesota in the
Eastern United States. It has been bred from acorns of 16 species of oaks. The adult
is shiny black to light reddish-brown. Light and dark scales make varied patterns on
the elytra and thorax. White scales usually make four spots on the thorax and base
of the elytra and a band on the distal third. Adults are from 4.8 to 6.6 mm long. The
snout is moderately stout and curved, and longer than the prothorax. It is slightly
longer and less stout in the female than in the male. Adults emerge from April to
August, depending on locality. Eggs apparently are deposited in damaged. cracked,
sprouted. or previously infested acorns. The larvae mature in 2 or 3 weeks. Then
they vacate the nuts and enter the soil to pupate. New adults emerge about 1” to 3
months later.

Conotrachelus carinifer occurs in the Coastal States from New Jersey to Texas
and west to Arizona. It has been reared from the acorns of eight species of oaks.
The wing covers are dark reddish-brown with a few black areas. These are covered
with brown to white setae in patches and bands. The prothorax is black and clothed
in pale-brown to white setae. The beak is curved and longer than the prothorax in
both sexes. Adults emerge in late summer or early fall and begin laying eggs within
4 days. The larvae become mature and vacate the acorns within 12 to 18 days.

Conotrachelus posticatus occurs throughout much the same range as C. naso. It
is known to breed in acorns from nine species of oaks. The adults have dark reddish-
brown wings with black splotches, the black sometimes predominating. The elytra
are sparsely covered with pale tan, yellow, and white setae with some patches.
Usually there is a narrow, curved posterior band. Their bodies are 4 to 5 mm long.

336

The beak is feebly curved and longer than the prothorax. Adults emerge from June
to August. The eggs, like those of C. naso, are deposited in damaged, cracked,
sprouted, or previously infested acorns. The larval stage lasts 10 to 30 days. The
first winter is spent as a larva in the soil; the second, as an adult beneath leaves on
the ground (476).

Other common eastern species of Conotrachelus include: C. juglandis LeConte,
which breeds in the nuts, stems, and leaf petioles of a number of species of Juglans;
C. retentus (Say), which attacks the nuts of black walnut and butternut; C. affinis
Boheman and C. hicoriae Schoof, two closely related species that are difficult to
separate and which breed in hickory nuts; C. aratus (Germar), which attacks young
shoots and leaf petioles of hickory; C. anaglyptius (Say), which breeds in the
tissues around fresh wounds on various deciduous trees including hickory, birch,
maple, and oak; and C. elegans (Say), which breeds in Phylloxera galls on hickory
leaves. Another weevil, Anthonomus suturalis LeConte, also develops in Phyllox-
era galls on hickory and pecan (467).

The subfamilies Rhynchophorinae and Cossoninae are placed in the family
Rhynchophoridae by some authors.

Rhynchophorus cruentatus (F.), the palmetto billbug, occurs from North Car-
olina to Florida and Louisiana and breeds in the trunks of weakened palm trees and
cabbage palmettos. The adult has a red thorax with black margins, shiny black wing
covers, fringes of long yeilowish hairs on the legs, and is about 30 mm long. It
feeds on bruised terminal buds or on sap exuding from wounded or recently felled
trees.

The subfamily Cossoninae, the broad-nosed bark weevils, contains several
genera and many species. A number of species breed in the sapwood of hardwoods
and conifers killed by bark beetles; others, some of which are important pests,
breed in the woodwork of buildings. The larvae of all species cut meandering
galleries across the grain of the wood and pack them tightly with granular frass
except for that portion immediately behind their bodies. Adults may be found in the
wood, but they usually occur under the bark about a year after the tree 1s killed. The
larvae of a number of species have been described (24).

Several species in the genus Cossonus breed in the sapwood of bark-beetle killed
trees. C. corticola Say, a shiny black species—under the bark of dying pines; C.
platalea Say, a flat, shiny black species—under the bark of hardwoods; C. im-
pressus Boheman, a dull black species—in the sapwood of both conifers and
hardwood (reported from Florida); and C. concinnus Boheman—under the bark of
hardwoods.

Species of Cossoninae that attack and damage wood in buildings deposit their
eggs in crevices in the wood. Adults of certain species reattack the wood from
which they emerge, thus intensifying the damage they already caused. The more
important eastern species are as follows: Hexarthrum ulkei Horn—damages wood-
work in old buildings, often reducing the wood to powder; Jomolips quercicola
Boheman—damages seasoned coniferous wood such as pine flooring and pecky-
cypress paneling; and Pselactus spadix (Herbst)— occasionally damages damp
wood beneath buildings and saltwater pilings above the high watermark.

Dryophthorus americanus Bedel occurs very commonly in hickory killed by the
hickory bark beetle. Stenoscelis brevis Boheman is common in dead, dying, and
rotting hardwoods.

Family Scolytidae
The family Scolytidae comprises a large, diverse, and important group of

beetles. A small segment of the family is actually responsible for its bad reputation;

337

a somewhat larger group occasionally causes losses in forest products, while most
species have never been economically important.

Two subfamilies are recognized (/354): the Hylesininae and the Scolytinae. The
Hylesininae are generally rounded behind, have a series of teeth or granules on the
anterior margin of each elytron, and have the head visible from the dorsal aspect.
The Scolytinae usually have a declivity at the posterior end, have no armament on
the anterior margin of the elytra, and have the head either partly or completely
concealed from above by the pronotum. Another mode of dividing the family
Scolytidae is by feeding habit. The scolytid beetles of concern in the forest feed on
the inner bark (phloeophagy), seeds and cones (spermophagy), wood (xylophagy),
or on mutualistic fungi that they culture in wood (xylomycetophagy or ambrosia
habit). The taxonomic divisions of the family do not necessarily correspond with
the feeding groupings because the various modes of feeding have repeatedly
evolved independently. This discussion of Scolytidae lists the genera and species by
feeding habit and by approximate phylogenetic order within the feeding groups.
Identification of some of the more important genera and species can be made by
using illustrations and the descriptions of beetles’ morphology, gallery systems. and
hosts. For the most part, taxonomic treatises on the Scolytidae of North America
(7356) and Canada (/53) should be consulted when a reliable identification of a
species of litthe economic importance is necessary.

Scolytid adults are typically soft and yellowish when first formed but soon
harden and turn reddish to dark brown or black. They are cylindrical beetles, from
about 0.9 to 9.5 mm long. The larvae are white, curved, legless grubs with
enlarged thoracic segments. The heads and mandibles are usually strongly scle-
rotized and dark red-brown. Larvae of the Scolytidae are very similar to those of the
Curculionidae (weevils).

When scolytids attack, the adults construct entrance tunnels into the inner bark
(phloem), pith, wood, or cone axil, depending on the species. The entrance tunnel
empties into a nuptial chamber or is simply elongated into an egg gallery. In
conifers, pitch and sap may exude from these holes and harden at the bark surface
to form a pitch or resin tube. Ambrosia beetles and most bark beetles push boring
dust out through the holes. Egg tunnels of bark beetles may be entirely in the
phloem but more often engrave the wood. Egg tunnels of wood-feeding (xy-
lophagus) species may be deeply cut into the wood surface or may be completely in
the wood. Tunnels of ambrosia beetles are always in the sapwood and may be
simple, branched, or compound. The females of most species deposit their eggs
individually in niches cut into the sides of the tunnel and cover them with fragments
of bark or wood. Others deposit groups of eggs in larger niches or grooves and
cover them with boring dust. Still others, particularly some of the pith beetles and
ambrosia beetles, place their eggs free in the tunnels.

When bark beetle larvae hatch they feed away from the egg tunnel, usually at
right angles to it. Many species can be recognized by their host and gallery pattern.
Ambrosia beetle larvae remain in the egg tunnel and feed on fungi growing on its
walls. Most Scolytidae pupate in cells at the ends of larval galleries, in the wood, in
the outer bark, or between the wood and bark. The adults remain in those cells until
their exoskeletons harden, sometimes even longer. While there, they usually feed
on whatever phloem or xylem remains after the larvae finish feeding. When ready
to emerge, the beetles gnaw holes through the bark to escape. The majority of a
particular colony will emerge within a few days. The ambrosia beetle adults emerge
through their parents’ entrance tunnels.

338
Se ee

Most Scolytidae fly and attack a new host as soon as they mature and some
species congregate under the bark of the new host to hibernate or mature sexually. A
few species feed on the twigs, buds, or bark of other trees before attacking a tree for
breeding purposes. Generally speaking, the adults spend only enough time outside
the bark to find new host material. Exceptions are species (Ips, Dendrocranulus,
Trypodendron, etc.) that overwinter in litter on the forest floor.

Beetles usually overwinter in the tree in the larval or adult stage, or in the forest
litter as adults. In warm regions, eggs and pupae may also be present in winter.
Activity is resumed with the advent of warm weather in the spring. Parent adults
continue their egg laying in the extended portion of galleries started the year before
or in newly constructed tunnels or freshly attacked trees. In the South, there usually
are several generations per year. In the North and at high elevations, there may be
only one or a partial generation per year.

Bark Beetles
Phloeophagus Scolytidae

In the United States, bark beetles kill an average of 25.5 million cubic meters of
sawtimber and pulpwood annually. This accounts for 90 percent of all insect-caused
tree mortality and 60 percent of the annual loss of timber to all causes (/22/). The
most destructive eastern species, the southern pine beetle, inflicted an annual
economic loss of $225 million by killing 9 million cords of pulpwood and 3 billion
board feet of sawtimber from 1960 to 1978 (969).

Most tree mortality 1s caused by species in the genera Dendroctonus, Ips, and
Scolytus. Scolytids in other genera occasionally may kill or damage trees or
indirectly cause their death by transporting pathogens from diseased trees where
they breed to healthy trees in which they feed. Rapid death of mass-attacked trees is
actually caused by fungi that the beetles carry in mycetangia (fungal pockets). The
fungal inoculum is able to penetrate the resinous (or other chemical) resistance of
the host; then the fungi colonize (and usually stain) the sapwood and disrupt the
flow of water to the tree crown. Even the tree-killing bark beetle species may most
often be restricted to breeding in broken or moribund material because healthy trees
are able to resist attacks of beetles and fungi in the vital phloem-cambial region.
However, bark beetles can overcome host resistance by mass attacks orchestrated by
aggregation pheromones produced when the pioneer beetles attack a tree (///). A
sufficient population mass attack can kill any host, regardless of its vigor. When the
attacking population is insufficient to exhaust the natural defense mechanisms of a
tree, the host will survive and the attackers will be repelled or killed (/03). Success
or failure of an attack can be visualized as a seesaw on which host resistance and
beetle numbers are balanced on a fulcrum—time. The balance will be affected by
an increase or decrease in either attack density or resistance, or a shift in the
position of the fulcrum (duration of attack). This concept explains why bark beetle
outbreaks generally follow a predisposing event (logging, flooding, drought, or
fire) that reduces host resistance or allows for a dramatic increase in the bark beetle
population. Once the population exceeds the threshold beyond which host resistance
is a factor, tree killing will continue until the supply of hosts is exhausted or natural
events (winterkill, disease, predation) reduce the population.

Major outbreaks of bark beetles have probably never been controlled by human
intervention. However, timber losses in localized eruptions have been curtailed by
direct action such as logging infested trees or felling them for treatment with
insecticides, solar heat, or fire. Synthetic pheromones have been able to disrupt

339

local “‘spot” populations of southern pine beetles (/026), and using pheromone
lures for mass trapping European elm bark beetles has contributed to control of
Dutch elm disease in urban areas (7/6). Parasites and predators (especially wood-
peckers) may keep endemic-level bark beetle populations from increasing, but
probably do not cause the collapse of outbreaks. Foresters can assist in biological
control of bark beetles by leaving a few dead and defective trees for nesting of
woodpeckers rather than felling all cull material during thinning and logging
operations.

A trap-log technique can be used beneficially against bark beetles that preferen-
tially attack cut material. Trees are felled shortly before beetles are expected to fly.
Infestation may occur naturally or be assured by baiting the trees with synthetic
aggregation pheromone. Before beetle broods mature, logs are debarked, milled,
etc. Although this method has been applied effectively against [ps typographus (L.)
in Europe for three centuries (/074), it has not been widely applied in North
America. A modified trap technique in which trees are killed with cacodylic acid
and baited with pheromone is more efficient because herbicide-induced drying of
the bark inhibits brood development and renders follow-up action unnecessary
(943, 1233). The most satisfactory means of reducing losses to bark beetles is to
prevent the occurrence of outbreaks by maintaining sanitation of logging debris,
lightning-struck or windthrown trees, etc., and by promoting stand vigor by
thinning and other cultural practices (85). Loss in quality of softwood logs due to
bark beetle-carried stain fungi can be avoided by prompt milling, ponding, or
sprinkling of logs.

Subfamily Hylesininae

The genus Hy/urgops is represented in eastern forests by only one species, H.
pinifex (Fitch). This species is widely distributed in the Eastern United States and |
Canada where it breeds in logs, stumps, and basal portions of dead and dying pines,
spruces, and larch. The adult is reddish brown to nearly black and from 4.5 to 5 mm
long. The undersurface is black and the declivity is covered with small, ash-gray
scales and a few long, erect hairs.

The genus Hylastes is represented by 15 species in North America, several of
which occur commonly in the Eastern United States (/2/). They usually breed in
the bases or roots of dying pines and spruces or in stumps or the bottom sides of
logs in contact with the ground. The adults occasionally kill pine transplants or
young plantation trees by chewing the bases of the stems; otherwise, members of
the genus are of minor economic importance.

Hylastes porculus Erichson occurs from Maine to the Carolinas and west to the
Lake States. It breeds in stumps and roots of dying pines. The adult is dark reddish-
brown to black and from 4 to 5.3 mm long. Adults are strongly attracted to freshly
cut lumber or to ongoing construction. In the Southeast, they fly during the period
April to November.

Hylastes salebrosus Eichhoff occurs commonly in the south Atlantic States. It
has been recorded breeding in loblolly, longleaf, and shortleaf pines, and in
spruces. It is also strongly attracted to freshly sawn lumber. The adult is dark
reddish-brown and about 4.5 mm long.

Hylastes tenuis Eichhoff occurs over most of the United States. Its hosts include
pines, spruce, and fir. The adult is dark reddish-brown to black and from 2 to 3.5
mm long.

Hylastes exilis Chapuis occurs in the Southern States from the District of
Columbia to Florida and Texas where it infests various species of pine. The adult 1s
dark reddish-brown and about 2.9 mm long.

340

rr eee te eee ee ae ee eee crepes ee

The genus Hylesinus contains a number of species that breed by preference in
various species of ash, and are commonly known as ash bark beetles. The adults
differ from most other bark beetles in having variegated coloration produced by
bands or spots of light-colored scales alternating with areas of dark scales. The
body is rather stout, and the elytra gradually descend behind; the antenna has a
seven-jointed funicle, with the club elongate, fusiform, and compressed. Favorite
breeding material consists of recently cut or broken trees; however, living trees
weakened by mechanical injury, disease, or fire may be attacked. Parasites and
predators, especially the braconid, Coeloides scolytivorus (Cresson), and the clerid,
Enoclerus nigripes (Say), provide a considerable degree of control at times.

Hylesinus aculeatus (Say), the eastern ash bark beetle, is the most common
eastern species. It breeds in ash species throughout the eastern part of the United
States and southeastern Canada. The adult is grayish brown with nearly black
markings and is 2 to 3 mm long. Winter is spent in the adult stage in tunnels in the
bark of living or felled trees. The adults emerge in the spring and fly to the trunks or
limbs of recently felled, dying, or seriously weakened trees to breed. Egg galleries
are constructed between the bark and wood, both of which are deeply engraved.
The galleries are biramous and transverse, with the two arms connected by a short
tunnel or nuptial chamber just below the entrance hole (fig. 157). Eggs are laid in
niches along the sides of the gallery. The larvae feed away from the gallery,
following the grain. Pupation occurs in deep oval cells between the bark and wood.
There are one to three generations per year, depending on location. This species and
the slightly larger H. pruinosus (Eichhoff) are economic pests to producers of rustic
ash products; otherwise, these beetles cause little or no economic damage.

7
Courtesy Duke Univ. Sch. For.

Figure 157.—Transverse egg galleries and vertical larval
tunnels of Hylesinus aculeatus, the eastern ash bark
beetle, in white ash.

34]

The genus Hylurgopinus is distinguished by its seven-segmented antennal funi-
cle, the strongly chitinized first two sutures of the antennal club, and the widely
separated forecoxae. One species occurs in North America.

The native elm bark beetle, H. rufipes (Eichhoff), breeds in various species of
elm in southern Canada and throughout the Eastern United States north of Alabama
and Mississippi. The adult is brownish black, thinly clothed with short, stiff, yellow
hairs, and from 2 to 3.5 mm long (fig. 158A). The head is convex, thickly
punctured, and nearly invisible from above; the antennal club is almost twice as
long as wide; the pronotum is narrow in the front, densely punctured, and reddish at
the rear; the elytral striae are composed of deep punctures; and the legs and
abdominal sternites are red.

A, F-532844
B, courtesy G. N. Lanier, SUNY Coll. Environ.
Sci. & For. photo by E. M. Gallagher

Figure 158.—The native elm bark beetle, Hy/urgopinus
rufipes: A, adult; B, gallery pattern.

Beetles overwinter as larvae or adults. Adults that mature during the late summer
or fall fly to healthy elms in which they bore overwintering tunnels in the bark at the
root collar or lower bole. In April and May overwintering adults emerge from
hibernation chambers, crawl up the tree and burrow into the bark of small (1 to 5 cm
in diameter) branches (/20/). Later they fly to dead and dying trees, broken limbs,
or recently cut logs or limbs to breed. Usually, dying or fairly moist recently dead
limbs at least 5 cm in diameter are selected. Entrance holes are made in bark
crevices or under overhanging bark flakes and they penetrate directly to the surface
of the wood. A biramous egg gallery is constructed with the arms extending away
from the entrance hole at various angles (fig. 158B). The gallery may be con-
structed horizontally, but is most often inclined from the horizontal. Galleries may
be constructed entirely in the bark, or they may scar the wood slightly. Eggs are laid
on both sides of the gallery, sometimes very close together. Young larvae feed away
from the gallery, usually following the grain. Pupation occurs in cells at the end of
the larval tunnels in the bark. There appears to be two generations per year in the
southern portions of the insect’s range and one and one-half in the northern portions
(O51; 1207).

342

Adults emerging from elms dying from Dutch elm disease often carry spores of
the disease-causing fungus on their bodies. When they bore into the bark of healthy
elms to feed or hibernate, some of the spores rub off onto the walls of their tunnels,
inoculating the tree with the disease. In most of the United States, the native elm
bark beetle has been displaced by the more aggressive smaller European elm bark
beetle. In northern New York, New England, and Canada, where the latter species
is limited by low winter temperatures, the native elm bark beetle is the primary
vector of Dutch elm disease.

The genus Dendroctonus contains some of the most destructive insects affecting
conifers in North America. The adults are reddish brown to black and from 2 to 9
mm in length. The body is cylindrical and rather stout; the head is broadly rounded
and visible from above; the antennal funicle is five-segmented; and the short
antennal club is sutured toward the tip. Important papers on the biology and
taxonomy of the genus are available (59/, 1353).

The southern pine beetle, D. frontalis (Zimmermann), the most destructive of
the eastern species of bark beetles, occurs throughout the Southeastern and South-
ern States. From 1948 to 1975 more than 14 million cubic meters of pine timber
were killed during a series of outbreaks in the Southern States (fig. 159). The insect
also occurs in Mexico, Guatemala, and Honduras. During a catastrophic outbreak
in Honduras in the early 1960’s, more than 55 million cubic meters of pine were
killed. It breeds in all species of yellow pines in its range; also in eastern white, red,
and spruce pines, and red and Norway spruce. Shortleaf, loblolly, Virginia, and
pitch pines are the most highly favored among the yellow pines. Infestations in
slash, longleaf, and eastern white pines are usually unsuccessful because of heavy
resin exudations. Attacks on red spruce are also unsuccessful, the beetles dying in
them after constructing short tunnels (692).

Figure 159.—Deteriorated loblolly pines 5 years after
they were killed by the southern pine beetle,
Dendroctonus frontalis.

343

The southern pine beetle adult (fig. 160A) is reddish brown to black and from 2.2
to 4.2 mm long. The front of the head has a distinct longitudinal median groove
bordered by a narrow elevation of tubercles on each side; the pronotum is slightly
narrowed at front, broadest at the middle, and about as long as wide; the elytra are
as wide as and over twice as long as the pronotum; and the declivity is convex. The
full-grown larva is a whitish, legless grub with a glossy reddish-brown head, and it
is about 5 mm long.

A B

F=Dol2or

Figure 160.—The southern pine beetle, Dendroctonus
frontalis: A, adult; B, pitch tubes on shortleaf pine; C,
winding egg galleries.

Winter is spent in the bark in all stages—egg, larval, pupal, and adult.
Emergence in the spring varies with the overwintering stage, geographic location,
and climatic conditions. In the southern Appalachians, the overwintering adults
emerge about mid-April, while those that develop from overwintering eggs may not
appear until late June. The bark of trees through which the adults have emerged is
peppered with small, round holes. Hibernating beetles do not attack healthy, living
trees after they emerge. Instead, they seek out and invade trees attacked but not
killed the previous fall (590). Generally, beetles attack the middle and upper trunk
first, especially in the Middle Atlantic States. Later they continue their attacks
down the trunk to within 1.5 m or less of the ground. In the Deep South,
overwintering adults emerge during warm periods in the winter and may attack the
upper and lower portions of the tree from which they emerged. However, most of
these beetles emerge in late February or early March and first attack the mid to
lower trunk of new host trees. Continuing attacks extend the zone of attack down to
the ground line and up into the base of the live crown.

The female bores directly into the cambium and constructs a nuptial chamber.
Points of attack are characterized by the presence of distinct pitch tubes and fine
reddish boring dust or white resinous boring particles in the bark crevices or on

344

upper leaf surfaces of understory vegetation (fig. 160B). The female is joined in the
chamber by a male and after mating she begins construction of a gallery diagonally
across the grain of the wood, etching the surface of the wood faintly. The direction
of the gallery eventually is reversed, thus creating a typical S-shaped or serpentine
pattern (fig. 160C). Eggs are deposited at intervals of 3 to 25 mm or more in niches
in each side of the gallery, one egg per niche. Hatching occurs in 3 to 9 days, and
the larvae tunnel away at right angles to the gallery. Young larvae initally produce
threadlike mines which are visible when the inner bark is exposed. As the larvae
continue to develop, they mine outward into the corky bark where they construct
individual cells in which to pupate. Each adult bores its own emergence hole. All
the adults in a given brood may emerge during a period of 10 to 32 days. Three to
five generations develop each year in western North Carolina. In Virginia and West
Virginia, four generations and a partial fifth occur; in the Deep South there are as
many as seven per year.

Newly emerged beetles may attack adjacent trees or they may fly to stands some
distance away. Trees under 15 years of age and less than 5 cm in diameter are rarely
attacked. When outbreaks develop, scattered groups of pines growing under dense
stand conditions, disturbed by recent logging or lightning damage, or on sites
subjected to moisture stress and disease are attacked. Once an epidemic is under-
way, stands of all age classes and densities are vulnerable. Spot infestations, which
may range from a few trees to stands occupying hundreds of hectares, are charac-
terized by a central zone of defoliated trees, one or more adjacent areas of red-
topped trees, and one or more peripheral fingers of infested trees with yellowish or
green foliage extending out into the surrounding forest. The needles of pines
infested during midsummer turn yellow in 2 or 3 weeks and reddish brown in 4 to 6
weeks. Death results either from the girdling of the main stem by the beetle or from
the effects of the blue stain fungus, Ceratocystis minor (Hedgc.) Hunt, which the
beetles introduce into the tree.

Southern pine beetle outbreaks appear to be caused by conditions that reduce tree
or stand vigor, whether by natural phenomena or human intervention. Drought or
flooding, overstocking, stand disturbance and site depletion are most often associ-
ated with initial infestations. Beetle populations increase rapidly in response to
favorable host and environmental conditions and cause extensive damage. Only in
the more northerly or high elevation areas of the insect’s range do low winter
temperatures serve as a check in terminating outbreaks (73). A return to more
favorable growing conditions tends to be associated with reduced beetle activity
over large areas.

Management practices designed to improve and maintain the vigor of stands
(e.g., timely thinning), and the removal of high-risk trees, such as those injured by
recent logging, struck by lightning, or attacked and weakened by other insects, are
helpful in reducing losses. Once an outbreak is underway, the salvage or chemical
treatment of infested trees may be helpful in suppressing populations. To be most
effective, these practices must be applied on a timely basis with first priority given
_to larger spots that have a greater volume of actively infested trees.

Several reviews of the life history and habits, natural enemies and associated
organisms, sampling and population dynamics, impacts, utilization, site, stand,
and climatic factors affecting stand susceptibility and vulnerability, silvicultural
and direct control methods and materials have been issued (56, 87, 107, 246, 247,
Dope SLO FOLD 35, 40, :/00, 1155, 41803. 1195).

345

The black turpentine beetle, D. terebrans (Olivier), occurs from coastal Mas-
sachusetts, New York, and New Hampshire to Florida, Missouri, and Texas. The
northern part of its range is coincident with stands of pitch pine near the Atlantic
Coast. All species of southern pines and red spruce are attacked, but loblolly and
slash pines apparently are the most frequently injured. The adult (fig. 161) is dark
reddish-brown to black, and from 5 to 10 mm long. The head is densely granulate,
roughly punctate, and convex in front. The pronotum and elytra are coarsely and
shallowly punctate. Full-grown larvae are creamy white, legless, about 12 mm
long, and have glossy reddish-brown heads.

Winter is spent in the adult stage in the northern parts of the insect’s range. In the
Deep South all life stages are present throughout the year. Eggs are principally laid
in the basal 90 cm of the trunk and large roots of weakened and dying trees and
stumps of recently cut trees. Green logs may also be attacked occasionally. The
female bores a hole through the bark to the cambium. Here she is joined by a male
and, working together, they excavate an egg gallery up to 25 cm wide and 50 cm
long on the face of the sapwood, usually in a downward direction. Eggs are
deposited in a long group on one side of the gallery. The larvae feed away from the
gallery in the phloem. They feed together in groups and excavate large cavelike
galleries, usually somewhat fan-shaped, and occasionally up to 30 cm across.
When fully grown they construct pupal cells either in the corky bark or between the
bark and wood. The adults emerge through holes chewed through the bark and fly
to trees or stumps to start a new generation. Several may emerge through a single
hole. There are two to three generations per year in the Deep South.

Beetles are attracted by terpenes released by stumps and injured trees. Trees
weakened by fire, logging, adverse climatic conditions, or naval stores operations
are also highly prone to attack. Occasionally uninjured, apparently healthy trees are
infested. Attacks are usually confined to a height of less than 2 m on the trunks of
standing trees. Sometimes, though, they occur to a height of 3.5 m. Attacked trees
have large reddish to whitish pitch tubes on the bark surface (fig. 162) and whitish
pitch and bark pellets in the bark crevices or at the base of the tree. Infested trees are
almost always secondarily attacked by ambrosia beetles, which produce piles of
fine white sawdust around the base of the tree.

F-532017

Figure 161.—Adult of the black turpentine beetle,
Dendroctonus terebrans.

346

F-519568

Figure 162.—Pine tree attacked by the black turpentine
beetle, Dendroctonus terebrans. Pitch tubes are
reddish to white at first but soon assume a grayish
hue.

Before 1949, the black turpentine beetle was thought of as a stump-infesting
scavenger or, at most, as a species that killed patches of bark on apparently healthy
pines. The first evidence of its widespread killing of trees occurred during an
outbreak from 1949 to 1951 in Louisiana, when tens of thousands of cubic meters
of lumber and thousands of cubic meters of pulpwood were killed. Since then,
severe infestations on slash, longleaf, and loblolly pines have been reported from
Florida, Georgia, Alabama, Mississippi, and eastern Texas. Losses in turpentine
orchards have been severe. Heavily infested trees yield little resin and usually die
within a few months. Losses of unprotected seed trees and losses in seed orchards
also may be severe. Pitch and Austrian pines of landscape value have been killed on
Cape Cod, Massachusetts.

Some degree of natural control may result from the feeding activity of engraver
beetles, wood borers, weevils, and termites which compete for the food of the
beetle larvae. In low-lying areas, considerable brood mortality also occurs during
periods of flooding. Losses may be prevented or reduced by minimizing damage to
- residual trees during logging operations, particularly in wet weather, or by rapidly
salvaging recently infested trees. Where the salvage of infested trees is not practica-
ble, they can be sprayed to kill the broods (///9).

The red turpentine beetle, D. valens LeConte, the largest species of the genus,
occurs in southern Canada, all the coniferous forests of the continental United
States except in the south Atlantic and Gulf Coast States, and Mexico. It attacks all

347

species of pine within its range, and occasionally spruce, fir, Douglas-fir, and larch.
The adult is light reddish-brown to dark brown and from 5.5 to 9 mm long (fig.
163A). Full-grown larvae are grublike, legless, 10 to 12 mm long, and white except
for a brown head and small brown area at the rear. Mature larvae have a row of
small, pale-brown tubercles along each side of the body. The adult is frequently
confused with lighter specimens of the black turpentine beetle, especially where the
ranges of the two species overlap.

The habits of the red turpentine beetle are very similar to those of the black
turpentine beetle. It, too, usually attacks trees of reduced vigor, but can attack
apparently healthy trees. Individual trees or groups of trees and fresh stumps are
attacked most frequently. However, destructive populations may also develop in
trees disturbed by logging, fire, or land clearing. Injured trees around construction
sites or adjacent to piles of fresh lumber are infested frequently. Trees of pole size or
larger are most susceptible (///5).

In the colder parts of the insect’s range, the winter is spent chiefly in the adult
stage but, to some extent, in the larval stage. In the warmer parts of its range, adults
fly intermittently during the warmer winter months. Attacks on standing trees are
initiated by females boring through the bark to the wood, usually in the basal 2 m of
the tree, but sometimes to a height above 3.5 m. Like those of the black turpentine
beetle, these attacks are characterized by the presence of pitch tubes on the bark
(fig. 163C). The gallery first runs horizontally or slightly upward, then it turns
downward. When the attack is made just above the ground line, the gallery may be
continued below ground line along a large root. A fully developed gallery may be 1

F-494425, 494422, 494420

Figure 163.—Red turpentine beetle, Dendroctonus
valens: A, adult; B, gallery with mass of eggs along
the side; C, pitch tubes at base of pine.

348

cm to more than 3 cm wide and up to | m or more in length. Eggs are laid in one or
more elongate masses along the sides of the gallery (fig. 163B). The larvae feed
gregariously away from the gallery in the phloem. Their feeding may kill a patch of
inner bark ranging from a few centimeters to more than 30 cm wide. Pupation
occurs in separate cells located between the bark and wood, either in the gallery ora
short distance forward into fresh inner bark. Adults emerge through holes chewed
through the bark; sometimes several use the same hole. In southern latitudes and at
lower elevations, there may be two or three generations per year. Farther north and
at higher elevations, there may be only one generation per year or every 2 years.

In areas where lumbering is continued for several years, the red turpentine beetle
often becomes very abundant. The sudden discontinuance of these operations,
therefore, may lead to attacks on healthy trees, causing catfaces and the killing of
decadent trees in the stand. Shade tree pines in areas of new construction are also
attacked and may be weakened or killed. Damage can be reduced or prevented by
avoiding damage to trees or stands, deep earth fills over roots, or piling lumber of
green logs near trees. It is also helpful to debark or spray freshly cut stumps with
insecticide and to cut and remove pines dying from other causes. Watering or
fertilizing individual trees also increases their resistance.

The spruce beetle, D. rufipennis (Kirby), occurs throughout the spruce forests of
America, and breeds in native spruces. The adult is bicolored (black with red-brown
elytra) or uniformly black and from 4.5 to 6.2 mm long. Normally, windfalls,
prostrate dying green trees, and overmature or weakened standing trees over 20 cm
d.b.h. are attacked. During epidemics, however, almost all trees are attacked
regardless of size or vigor. Attacks usually begin on the lower third of the bole
except for the first 60 to 90 cm above the ground. Later in the season, they are
continued upward and downward, exclusive of limbs and parts of the trunk less than
20 cm in diameter.

Females construct vertical, almost straight egg galleries in the phloem, engraving
the wood. Eggs are deposited in groups along the side of the gallery. After the eggs
are laid, the attacking adults may vacate their galleries and construct new ones in
the same tree or in nearby trees. The larvae may feed gregariously for the first two
instars and individually for the last three instars. Pupation occurs in cells at the end
of larval tunnels and the winter is spent in the bark in either the larval or adult stage.
Signs of attack are red boring dust and pitch tubes on the bark, fading and dropping
needles, and the reddish appearance of the twigs after the needles drop.

Several major outbreaks have occurred in eastern forests. One, from 1897 to
1901, killed more than 5.7 million cubic meters of valuable spruce in northern New
England and eastern Canada (59/). The underlying causes of outbreaks are not well
understood, but piles of slash in mature stands are believed to trigger outbreaks
(1181). Some degree of control is possible by cutting infested trees in the fall and
removing them from the woods before spring, or by storing the logs in water.

The eastern larch beetle, D. simplex LeConte, occurs in the Northeastern States
south to West Virginia and west to Minnesota. It also occurs from coast to coast in
Canada and northwestward to Alaska. Its preferred host is tamarack but it has also

~been recorded from red spruce. Adults are dark brown, the elytra often having a
reddish cast, and are from 3.4 to 5 mm long.

Winter is spent as young adults or larvae in the brood galleries. Adults are active
from May until late August. Eggs are deposited in alternate groups of three to six
each in niches arranged along the sides of longitudinal, sinuate galleries. The larval
mines are in the inner bark and are quite short. Adults may reemerge and construct

349

several additional galleries during the season. Up to three generations may be
produced annually. The first of these reaches maturity by midsummer and the
second, by mid-September. The third brood spends the winter as larvae or young
adults.

The eastern larch beetle generally infests dying or injured trees. However, during
the 1970's, thousands of apparently healthy tamarack were killed in the Adirondack
and Green Mountains. In many stands almost all larches more than 10 cm d.b.h.
were killed. Plantations of European larch at Wanakena, N.Y., also suffered losses.

The lodgepole pine beetle, D. murrayanae Hopkins, is known from the Great
Lakes region where it attacks stumps and the root collar of injured, windthrown,
decadent or dying jack pine. Occasionally it infests red or eastern white pines. In
the Rocky Mountain region of the West, this species has periodically been a serious
killer of apparently vigorous lodgepole pine. Tree killing by D. murrayanae is often
mistakenly attributed to the better known spruce beetle, which this species very
closely resembles (/353). Like some spruce beetles, adults are distinctly bicolored,
having reddish-brown elytra and black bodies. It is most easily differentiated from
the spruce beetle by its host and by the habit of the larvae to continuously feed in
contact with one another rather than construct isolated feeding tunnels.

The Allegheny spruce beetle, D. punctatus LeConte, attacks stumps and the
root-collar region of weakened or dying white and red spruces. This species is
infrequently collected and apparently rare. It can be distinguished from the closely
related D. rufipennis and D. murrayanae by its uniformly brown or dark-brown
body covered with orangish, rather than pale-yellow, setae. Little is known about
the biology of this species.

The genus Phloeotribus is represented by a number of eastern species, all but one
of which breed in deciduous trees. The adults are distinguished from other bark
beetles by the loosely jointed antennal club, all three parts of which are extended on
the inner side into a leaflike process. They breed in dead or cut material or in
weakened or dying trees. Young adults burrow into the bark of living trees during
the fall where they spend the winter. Their burrows often extend into the outer part
of the living bark, causing irritations which result in abnormal growths. These may
show up as swellings on the trunks of a badly infested tree. Trees that harbor
overwintering aggregations are not killed, but they may be seriously weakened.

Phloeotribus frontalis (Olivier) breeds in mulberry and is believed to occur
wherever its host grows in the Eastern United States. Adults are brown and about 2
mm long. The branches and trunks of living trees or the trunks and stumps of killed
trees are preferred for breeding. The gallery system consists of two short and deeply
engraved branches extending transversely from the entrance hole. Aggregations of
adults overwintering in bark of healthy trees may kill patches of bark, which are
later sloughed.

Phloeotribus dentifrons (Blackman) breeds in hackberry in the South and in the
Midwest. Adults are dark brown to black and 1.5 mm long. Injured or dying limbs
are preferred as breeding material. Girdled or weakened trees or green logs are also
attacked if the bark is fairly smooth and not too thick. Many of the adults of the fall
generation spend the winter in their burrows. Others may emerge, then bore into the
bark of living trees to hibernate. Adults overwintering in their burrows often
obliterate the gallery patterns during their prolonged periods of feeding under the
bark.

The peach bark beetle, P. liminaris (Harris), occurs in southern Canada and
from New Hampshire to Michigan and south to the Gulf Coast in the Eastern United

350

States where it attacks cherry and other stone fruit trees. Breeding attacks some-
times kill defoliated forest trees (/07/) and aggregations of overwintering adults
may damage orchard trees by killing patches of bark. Adults are light brown to
nearly black, feebly shiny, sparsely clothed with long, fine, whitish hairs, and from
1.5 to 2.2 mm in length. The habits of the species are similar to those of P. frontalis
except that the galleries tend to be somewhat more irregular.

The genus Phloeosinus is represented in North America by 27 species, 5 of
which occur in eastern forests (/22). They breed preferably in cut, broken, or
decadent conifers. The adults construct short, longitudinal egg galleries between
the bark and wood. Newly emerged adults feed briefly before attacking a new host.
Sometimes they clip off and eat young leaflets on healthy trees. Generally, however,
they bore into the twigs. This occasionally causes the twigs to wilt, die, break, and
drop to the ground.

Phloeosinus dentatus (Say), the eastern juniper bark beetle, occurs from New
Hampshire to Georgia and westward to Texas and Nebraska. Its most common host
is eastern redcedar, but it also attacks northern white-cedar. The adult, piceous-
brown to black, is clothed with rather abundant short, gray hairs, and is 2.2 to 2.8
mm long. Eggs are laid in short galleries that extend upward from the entrance
hole. The larvae mine for short distances across the grain, then upward with the
grain (fig. 164). Infestations are usually found in cut, broken, or fire-damaged
trees. Attacks have been reported on living, overtopped redcedars infested with the
root rot fungus, Fomes annosus (Fr.) Karst, in North Carolina. Neither the insect
nor the fungus working alone usually kills trees. Working together, however, they
kill trees of all sizes (76). Cutting and burning infested branches and keeping the
trees in a healthy condition should be helpful in control.

Courtesy Duke Univ. Sch. For.
Figure 164.—Egg galleries and larval mines of
Phloeosinus dentatus, the eastern juniper bark beetle,
on trunk of eastern redcedar.

Phloeosinus taxodii Blackman, the southern cypress beetle, breeds in bald-
cypress and probably occurs wherever its host grows in the South. The adult is
brownish black, has reddish-brown elytra, and is from 2.1 to 3 mm long. Logging
slash and the larger limbs and trunks are especially attractive for breeding purposes.
The galleries, and the larval and adult feeding habits are similar to those of the
eastern juniper bark beetle. Because it apparently does not attack and kill living
trees, the species is of minor importance.

351

The other three species of Phloeosinus occurring in eastern North America are:
P. canadensis Swaine—on eastern redcedar and northern white-cedar in eastern
Canada and from Maine to the Lake States: P. pini Swaine—on various pines and
spruces in the Lake States and southern Canada: and P. scopulorum neomexicanus
Blackman—on juniper and cypress in Texas.

The genus Chramesus is represented in the East by four species of stout, strongly
convex, “humpbacked” beetles, less than 2 mm long. They are further dis-
tinguished by their large, elongate-oval, unsegmented antennal clubs and the five-
segmented antennal funicle attached to the side of the club. They breed in broken or
dying twigs and small limbs. The adults construct longitudinal or transverse
unbranched egg galleries, partly in the bark and partly in the sapwood. C. hicoriae
attacks various species of hickory throughout the Eastern States and in eastern
Canada. C. chapuisi LeConte attacks hackberry from Pennsylvania to Florida and
Texas (/20), and C. subopacus Schaeffer attacks hackberry in the Southern United
States to Honduras. The fourth species, C. wisteriae Wood, attacks wisteria in
Mississippi.

The genus Carphoborus is represented in eastern forests by at least two species.
The adults are dark brown to black, more or less covered with short scalelike hairs,
and are less than 2 mm in length. C. bifurcus Eichhoff is a fairly common species in
the South where it breeds in dying, broken, and cut limbs of pines.

Polygraphus rufipennis (Kirby), the four-eyed spruce bark beetle, occurs in the
spruce forests of the United States south in the Eastern States through the Ap-
palachians. This dull dark-brown beetle, 2 to 3 mm long, can be identified easily by
its completely divided eyes. Its hosts, in addition to the spruces, are larch, pine,
and balsam fir. Infestations are usually found in slash and in dead and dying trees.
However, when heavy populations develop in such material, nearby living trees are
also subject to attack. The adult is dark brown to black and about 2.3 mm long.
Eggs are laid in the sides of three to five irregular, short galleries that radiate away

from a central nuptial chamber. The bark, but not the wood, is slightly engraved.
Subfamily Scolytinae

The genus Scolytus is represented in eastern North America by a number of
species of true bark beetles, several of which are of economic importance (//9).
Most species feed in twigs or buds of a host species while in the process of seeking
a weakened host in which they can breed. All species except one breed in deciduous
trees, and two species have been implicated in the transmission of Dutch elm
disease. The adults differ from other eastern bark beetles in having the outer angle
of the foretibia produced into a curved hook and the ventral surface of the abdomen
ascend abruptly to the rear. In some cases the abdominal declivity is concave and
ornamented by spines, tubercles, etc.

The smaller European elm bark beetle, S. mu/tistriatus (Marsham), one of the
two principal American vectors of the Dutch elm disease fungus, Ceratocystis ulmi
(Buisman) C. Moreau, was first observed in North America in 1909 at Boston,
Mass. (202). Since then, it has spread over most of the United States and into
southern Canada. Its hosts include all species of elm (U/mus) and the related
Japanese zelkova, Zelkova serrata. Adults are 2 to 3 mm long and distinctly two-
toned with the elytra red-brown and the other parts black. Males have a bright
yellow brush on the front of the head. The underside of the posterior is concave and
armed with a stout spine projecting from the anterior margin of the second
abdominal sternite (fig. 165A). The larvae are typical legless grubs about 3 mm
long.

352
tet. Os | irae een See.

F-531255

Figure 165.—The smaller European elm bark beetle,
Scolytus multistriatus: A, adults (male on left, female
on right); B, gallery pattern.

Larvae overwinter in pupal chambers in the bark. Pupation occurs at the onset of
warm weather in the spring and adults emerge through “shot holes” they chew
through the bark at about the time the first elm leaves are fully expanded (late
March to early June, depending on the latitude). Emerging adults fly directly to
-weakened or diseased elms to breed in the inner bark or to healthy elms where they
feed in twig crotches. It is during twig feeding by contaminated beetles that healthy
trees are inoculated with spores of the Dutch elm disease fungus. Chances of
infection are greatest in the spring and early summer when the long vessels of the
tree are functioning and near enough to the surface for the beetles to cut them while
feeding.

353

Breeding attacks begin in material that is weakened by drought, disease, .or
injury. The bark of any wood recently cut from a living elm may be colonized.
Attacks may quickly spread to undiseased parts of the tree and to adjacent healthy
elms. Rapid death of entire trees or sections of trees and the retention of shriveled
brown leaves indicate mass attack and successful colonization by the beetles,
although beetles may also colonize material after leaves have wilted and been shed.
Trees that are vigorously attacked emanate a sticky liquid that attracts ants, yellow
jackets, and certain moths. Bark that has been removed or dried to the point of
cracking will not be infested. Females initiate attack and release an aggregating
pheromone that attracts both sexes to the breeding site. After mating near the
entrance of their attacking tunnels, females bore egg galleries 2.5 to 8 cm long.
engraving the surface of the wood, parallel with the grain (fig. 165B). Eggs are
deposited in niches along the sides of the gallery. The larvae feed in the inner bark
and the surface of the wood, angling away from the gallery. When fully grown, they
form cells in which to pupate in the bark. During the spring and summer, the life
cycle may be completed in 35 to 40 days. The spring-flying adults produce a
generation that emerges from mid-June to mid-August, depending upon the lati-
tude. Most of the progeny of the summer-flying adults enter a developmental
diapause and overwinter, but a portion continues to develop and emerges in the fall.

The vertical orientation, the unbranched condition, and the engraving of the
sapwood by both adults and larvae distinguish the gallery system of this species
from the branched horizontal galleries of the native elm bark beetle.

Because of its role in the transmission of Dutch elm disease, the smaller
European elm bark beetle has been the subject of considerable research, and huge
expenditures have been made to eradicate or control it. Eradication efforts proved
futile, but progress has been made on its control. The fundamental mode of
controlling the size of the elm bark beetle population is limiting their supply of
breeding material (/287). Elm firewood, broken limbs, and severely stressed trees
as well as elms that have Dutch elm disease should be eliminated or treated with an
insecticide. In addition to the physical removal and burning or burying of infested
or potential beetle breeding material (sanitation), diseased or unwanted elms can be
rendered unsuitable for beetle breeding by the trap tree technique based upon
injection of certain herbicides that cause the bark to dry before larvae are fully
developed (943).

Beetles that escape sanitation can be lured away from elms and killed on sticky
traps baited with a synthetic copy of their aggregation pheromone (7/6, 977).

An integrated approach that employs a medley of techniques to manage the elm,
the bark beetles, and the fungus is most effective in reducing losses to Dutch elm
disease.

Further information on control of elm bark beetles and Dutch elm disease is
available (688, 7/5, 717, 718).

The larger shothole borer, S. mali (Bechstein), also introduced from Europe,
has been recorded from southern Ontario and Quebec in Canada, and from Con-
necticut, Maryland, New York, New Jersey, Ohio, Indiana, and Pennsylvania. It
breeds in moribund apple, cherry, and elm. During its dispersal phase it sometimes
feeds in twig crotches of healthy trees and may be an occasional vector of Dutch
elm disease. The very dark-brown adult is from 3.4 to 4.4 mm long, and is about
one-half as wide as long. The elytra have the punctures arranged in regular strial
and interstrial rows of nearly equal size. The abdomen is weakly concave ventrally.
The fifth sternite is longer than the third and fourth combined and the elevation of

354
ee ae ae

the posterior margin of the abdomen is lacking in the male. This species differs
from other eastern Scolytus by its oblique second abdominal sternite. Dying and
weakened limbs and freshly cut wood are preferred for breeding purposes. The
gallery system is slightly larger but similar to that of the smaller European elm bark
beetle. When fully grown, larvae bore cells into the sapwood where they overwinter
and pupate in the spring. There is one generation per year (972).

The hickory bark beetle, S. guadrispinosus Say, occurs from southern Quebec
to Georgia, Alabama, and Mississippi and west to Minnesota, Kansas, Oklahoma,
and Texas. Its range is probably coincident with the natural range of hickories. This
species is a serious pest of hickories and has also been recorded feeding on
butternut and pecan. The adult is stout, black, almost hairless, and 4 to 5 mm long.
The venter of the male is deeply excavated. The third abdominal segment is armed
with three spines; the fourth with one large median spine. The venter of the female
is without spines.

Adults appear in early summer and feed for a short time at the bases of leaf
petioles and on the twigs of hickory before flying to the trunks and branches of
living trees where they construct rather short (ca. 3 cm), longitudinal egg galleries
between the bark and wood (fig. 166). In thick-barked trees, the gallery may
scarcely touch the wood; in thin-barked limbs it may occur almost entirely in the
wood. Eggs are deposited in niches at each side of the gallery. The larvae feed
across the grain until nearly fully grown, then they abruptly turn and mine parallel
to the grain. Before reaching maturity they leave the phloem and bore into the bark
where they construct cells in which to pupate. The winter is spent in the larval stage
and pupation occurs in the spring. There is one generation per year in northern
areas. In the South, there are normally two broods per year.

Courtesy Duke Univ. Sch. For.

Figure 166.—Galleries of the hickory bark beetle,
Scolytus quadrispinosus, in phloem of hickory. Note
short vertical egg galleries and fan-shaped larval
galleries.

355

The hickory bark beetle is one of the most important insect pests of hardwoods in
the Eastern United States (76). During drought periods. outbreaks in the Southeast
have killed large tracts of hickory timber. At other times, damage is generally
confined to the killing of single trees or to portions of their tops. The foliage of
infested trees or tree limbs turns red within a few weeks of attack. Control practices
include felling infested trees and destroying the bark during the winter months.
treating infested bark with insecticides, or storing infested logs in ponds. To be
effective. this type of control should be conducted over large. natural units.

The shothole borer, S. rugulosus (Muller), an introduced species known to have
been in the United States since 1878. now occurs throughout temperate North
America. It breeds in most fruit trees, and uncommonly in mountain-ash,
hawthorn, and elm. The adult is grayish black, and from 1.7 to 2.9 mm long. The
elytra are covered with short hairs and are reddish brown at the apex. The venter is
unarmed and more gradually ascending than in other Sco/ytus that occur in North
America. Because of its preference for broken, cut, or dying material, this species
is of minor economic importance in forestry. However, the shothole borer is an
important horticultural pest because populations that build up in pruned limbs and
culled trees kill buds, limbs, and sometimes entire trees. Twig and bud damage
results from adults feeding: branches are killed when they are mass attacked and
colonized. Egg galleries are somewhat irregular but generally are parallel with the
grain. Larvae mine the cambial region and pupate there during summer months.
Larvae that mature in the fall generally bore cells into the sapwood where they
overwinter and then pupate in the spring. There are one to four generations per year,
depending on locality. A parthenogenic race occurs in Israel (508).

Scolytus fagi Walsh breeds in beech and hackberry from Illinois to Texas. Adults
are 4.5 to 5 mm long. The elytral striae are distinctly impressed, and the strial
punctures are much coarser than those of the interstrial rows. The species is of slight
economic importance.

The hackberry engraver. S. muticus Say. occurs from North Dakota and
Pennsylvania to Texas and Florida, and breeds in dying and dead limbs of hack-
berry. The adult is reddish brown to black, and from 4.5 to 7.5 mm long. There are
long, ashen hairs on the elytra and sides of the pronotum. Egg galleries are similar
to those of the hickory bark beetle. The larvae feed first between the bark and wood:
later they burrow into the wood where they pupate. There are two generations per
year in the Deep South and one per year in the Lake States.

Scolytus piceae (Swaine). the spruce scolytus. occurs from Alaska and Nova
Scotia to California and New York approximately coincident with the range of
white and Englemann spruces. It breeds in all spruce species within its range:
reports of spruce scolytus in larch and balsam fir represent aberrant occurrences or
errors. Adults are readily distinguished from other eastern members of the genus by
a ventral spine that arises from the center rather than the anterior margins of the
second abdominal sternite. The unique burrows of this species typically consist of
two and sometimes three egg galleries extending across the grain from a central
nuptial chamber. Broken limbs and tops are preferred breeding material.

Members of the genus Pseudothysanoes are closely related to those of the genus
Thysanoes. They differ. however. by breeding in the bark instead of the wood of
their hosts. P. lecontei Blackman attacks various oaks, hackberry. hophornbeam.,
chestnut, and walnut from New York to North Carolina. The adult is dark brown.
shiny, and about 1.2 mm long. P. rigidus (LeConte) breeds in basswood from
Canada to Michigan, Ohio, and West Virginia.

356

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Pseudothysanoes dislocatus (Blackman) is brown and about 1.3 mm long. The
head is slightly concave at the middle, with a row of long hairs at the lower margin
of the concavity. The antennal funicle is six-segmented, the club unsegmented, and
the scape armed with long hairs. It constructs its galleries just beneath the bark of
twigs of its hosts. It has been recorded from hickory from West Virginia to Florida.

The genus Pityogenes contains a number of species, all of which breed in the
twigs, limbs, and thin-barked portions of the boles of pines. Some species prefer to
breed in slash, whereas others most commonly attack decadent lower limbs of living
trees. Vigorous trees are usually not attacked except in heavily infested areas. Trees
weakened by drought or transplanting, or by ground fires or mechanical means, are
frequently attacked and killed. Adults are usually stout and sparsely pubescent. The
antennal funicle is five-segmented, and the antennal club is flat and sutured on both
sides. The elytra are marked with rows of punctures, excavated, and ornamented
with teeth at the posterior end. Females of all species except P. meridianus
Blackman have conspicuous cavities in the front of their heads. Males have three
slightly enlarged to very enlarged spines on each side of the elytral declivity.
Attacks are initiated by the males which are joined by two to seven females. The
gallery system is usually engraved on the surface of the wood and consists of
several egg tunnels radiating from a central nuptial chamber.

Pityogenes hopkinsi Swaine is a common species wherever its host, eastern white
pine, occurs in eastern North America. Rarely other pines and spruces serve as
hosts. In early spring, overwintering adults infest winter-pruned limbs and other
white pine slash. A summer generation attacks trees weakened by basal canker or
blister rust or joins the pine engraver or the red turpentine beetle in attacking
stressed trees. Logging slash may be infested any time when adults are active. P.
hopkinsi concentrates its attacks in the smooth-barked portions of the tree, thereby
limiting competition with the pine engraver, which infests areas with fissured bark.
The insects are 1.8 to 2.1 mm long. The male has three similar teeth on the elytral
declivity; the female has large frontal fossa.

Pityogenes meridianus Blackman is known to occur in North Carolina and
Mississippi. It breeds in slash and the dead and dying lower branches of pines
weakened by shading or injured by ground fires. The adult is dark reddish-brown
and from 2.7 to 3 mm long. Males have a large, blunt, downward curving spine at
the upper margins of the elytral declivity and two smaller, upward curved spines at
the lower margin of the declivity. Females have no frontal fossa. Burrows consist of
two to five galleries originating at and radiating away from a central nuptial
chamber. The related species, P. plagiatus (LeConte), breeds in pines in the
Atlantic States. The elytral declivity is similar to that of P. meridianus but the
species are easily distinguished by the female’s large frontal fossa that is divided by
a vertical partition.

The genus Pityokteines is represented in the United States by several species,
only one of which occurs in eastern forests. The beetles breed primarily in dying or
felled trees, particularly firs and spruces. The eastern species, P. sparsus (Le-
Conte), the balsam fir bark beetle, is frequently injurious to balsam fir, killing
large groups of trees. Pine, spruce, and larch are also attacked. Infestations are
found in logging slash in the limbs and tops of trees dying suddenly, in windthrows,
and in weakened and perfectly healthy trees. The adult is about 2 to 3 mm long and
is distinguished by long yellow hairs arising from the front of the head and from the
apical margin of the pronotum. Eggs are deposited in large niches along the sides of
several galleries that radiate away from a central nuptial chamber and scar the wood
deeply. Larval tunnels are longitudinal and follow the grain of the wood.

357

The genus Orthotomicus is represented by one species in eastern forests. It is
closely related to the genus /ps, but the aduits differ in having obliquely truncate
rather than flattened antennal clubs and feeble rather than well-developed teeth on
the margin of a shallowly concave declivity.

Orthotomicus caelatus (Eichhoff) occurs throughout the East, Lake States, and
eastern Canada. It commonly breeds in thick bark on stumps and logs or at the
bases of weakened or dying pines, spruce, larch, and balsam fir. The adult is dark
reddish-brown to nearly black and from 2 to 2.3 mm long. Short, radiating egg
galleries originate at central nuptial chambers, and from one to six eggs are laid in
large niches or pockets along their sides. Specimens of a morphologically closely
related species, possibly a variety, have been collected from the twigs of fire-killed
young loblolly pines in North Carolina, where they were apparently breeding as
well as mining out the pith and wood. Adults were also reared from dry twigs and
the tips of longleaf pine logging slash (76).

Among the bark beetles, the genus /ps ranks next in importance to the genus
Dendroctonus in its destructiveness to pines and spruces. Infestations normally
occur in lightning-struck trees and those recently felled by windfalls, snow-
breakage, and logging, and in road slash. However, when heavy populations
develop in this material, the adults emerge and attack and kill adjacent groups of
young healthy pines and the tops of older trees. Infestations in green timber are
usually of short duration unless the trees have been weakened by drought, fire, or
other disturbances. Spot- or group-killing in pulpwood or pole-size trees or, less
often, in mature stands is characteristic of outbreaks. Widespread outbreaks occur
frequently, and losses may be extremely severe. More than 3 million cubic meters
of commercial timber and more than 1.8 billion cubic meters of pulpwood were
killed in the South Atlantic and Coastal States during the period of 1952-55. It is
estimated that annual losses of 765 million cubic meters of pulpwood are incurred
in Florida alone. The North American species of /ps have been arranged into a
number of taxonomic groups (592, 593, 594, 595, 596, 597, 598, 599, 600).

The male initiates the attack by boring through the bark to the wood and
constructing a nuptial chamber. Here he 1s joined by one or more females (typically
three), each of which excavates an egg gallery in the phloem. These galleries radiate
in all directions from the chamber through the phloem, but eventually tend to run
parallel with the grain of the wood. The resu!tant pattern tends to form a rough Y or
| shape. Eggs are deposited in small niches at irregular intervals along the sides of
the gallery, and the larvae tunnel in the phloem until fully grown. Pupation occurs
in cells hollowed out in the inner bark. Young aduits feed for a short time beneath
the bark and then emerge, several often using the same exit hole. In northern areas,
the winter is spent in the adult stage under the bark of the brood tree or in the litter
below the infested tree. In the South, winter may be spent in the bark in all life
Stages.

Ips calligraphus (Germar), the six-spined engraver, occurs through eastern
America. Although it breeds in any species of pine available, its distribution in the
Northeast coincides with that of pitch pine. Trunks, stumps, and large limbs of
recently felled trees appear to be favored for breeding purposes, but the trunks of
apparently healthy pines are also attacked, especially in the Southeastern States
where it frequently attacks in concert with other /ps and Dendroctonus species.
Attacks on living trees usually occur on the lower portions of trunks with diameters
of 15 cm or more. In the South, this is one of the first species to attack drought-
stressed trees.

358

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The adult is dark reddish-brown to black and from 3.5 to 6.5 mm long. The
declivity is deeply excavated and coarsely punctured. Each side is armed with six
teeth, and the apical margin is strongly produced. The egg galleries, usually three
to five, radiate from a central nuptial chamber and run longitudinally, grooving both
the bark and wood (fig. 167). The larval mines are broad, tortuous, often long, and
transverse. In the South, the life cycle may be completed in 25 days, and there may
be 6 or more generations per year.

Ips grandicollis (Eichhoff), the southern pine engraver, occurs in eastern
Canada and in the Eastern United States from Massachusetts to Minnesota,
Nebraska, and Texas and south to Florida and Mississippi. Like J. calligraphus, it
attacks any pine species available but its distribution north of the southern pine
region is coincident with the distribution of pitch pine. Recently felled trees and
slash are preferred, but the trunks and limbs of apparently healthy trees are also
infested when attack occurs in concert with other bark beetle species. Heaviest
infestations in large living trees are found on limbs and the upper portions of trunks.
Spot- or group-killing of pines is characteristic of the species. During periods of
extreme drought, these groups increase in size and abundance. Populations nor-
mally develop in areas of recent logging operations.

The adult is dark reddish-brown to black and from 2.8 to 4.7 mm long. The
declivity is deeply excavated, coarsely punctured, armed with five teeth at each
side, and the apical margin is strongly produced. The egg galleries, three to five,
radiate from a central nuptial chamber and run longitudinally, grooving both the
bark and wood (fig. 168). The larval mines are more or less transverse. In the
South, the life cycle requires from 20 to 25 days, and there are 6 or more
generations per year.

The small southern pine engraver, /. avulsus (Eichhoff), the smallest species of
Ips, breeds in all species of pines from southern Pennsylvania to Florida and Texas.
Thin-barked slash, such as the limbs and tops, is preferred but groups of young,
vigorous trees and the tops of large, living trees are also attacked frequently and
killed. Attacks on large trees are usually associated with attacks on the lower
portions of the trunks by other species of ps or Dendroctonus frontalis. Adults are
attracted to freshly cut and injured trees. Any disturbance that causes pitch flow
may induce attack. Spot-killing occurs at times among pines showing no evidence
of previous injury or decreased vitality.

The adult is reddish brown to black and about 2.3 to 2.8 mm long. The declivity
is shallowly excavated and deeply punctured. Each side is armed with four small
teeth and the apical margin is slightly produced. Adults make one to several long
winding egg galleries that originate from a central nuptial chamber (fig. 169).
Larval galleries are short, transverse, and each ends in a pupal cell in the phloem.
In the South, the life cycle may be completed in 18 to 25 days, and there may be 10
or more generations per year.

The pine engraver, /. pini (Say), occurs throughout most of the coniferous
forests of North America except the Pacific Coastal forests, the southern pine
forests, and Mexico. In the Eastern States it is ubiquitous in the north and extends
. south in the Appalachian Mountains to Georgia. It breeds in all species of pine and
spruce within its range. Infestations usually develop in logging slash and windfalls
or in trees dying of other causes. When heavy populations build up in this type of
material, nearby healthy trees may be attacked and killed. Heavy infestations have
occurred in cutover and burned-over areas in Canada.

359

Courtesy Duke Univ. Sch. For. Courtesy Duke Univ. Sch. For.
Figure 167.—Gallery of /ps calligraphus Figure 168.—Gallery of the /ps
on limb of a_ shortleaf pine. grandicollis, southern pine engraver,

on limb of loblolly pine. Note egg
niches and incompleted larval mines.

The adult is brown to black, is from 3.5 to 4.5 mm long, and has four teeth on
each side of the declivity. Egg galleries, from three to six, radiate away from a
central nuptial chamber in the phloem, deeply scarring the sapwood. Larval tunnels
extend a short distance in the inner bark and end in pupal cells. Adults remain under
the bark for a short period before emerging. While there, they make irregular,

360

Cc r.
Figure 169.—Galleries of the small southern pine
engraver, /ps avulsus, in bark of shortleaf pine. Note
radiating tunnels of adults, short larval mines, and
pupal chambers.

meandering food tunnels, deeply engraving the wood. Winter is spent in the adult
stage on the ground. There appears to be three generations per year as far north as
Wisconsin (1065).

Other less common eastern species of /ps include: /. perturbatus (Eichhoff), the
northern spruce engraver— breeds in white spruce in the Lake States and Canada;
I. perroti Swaine—breeds in red and jack pines in Minnesota; /. borealis Swaine,
the northern engraver—breeds in white spruce in Maine, Canada, and the Lake
States; /. latidens (LeConte)—breeds in eastern white, red, jack, and Scotch pines;
rarely in white spruce; and hemlock in New York, the Lake States, and adjacent
parts of Canada.

Lymantor decipiens (LeConte) occurs from Michigan and Quebec to Kansas and
Mississippi. The adult is reddish brown and about 1.7 mm long. The antennal
funicle is four-segmented, the club sutured on both sides and slightly longer than
wide. It breeds in dead dry limbs, sprouts, and seedlings of living hickory, maple,
and apple. Its burrows are constructed in the wood, usually just beneath the bark
but sometimes deeper. The adults and larvae reportedly feed on certain black wood
fungi that are always present in the dead wood.

The genus Dryocoetes is represented in North America by seven species, five of
which occur in eastern forests (/5/). They usually breed in the upper portions of
trunks, in the roots of injured or dying trees, or in windfalls. Both coniferous and
deciduous trees are attacked.

Dryocoetes affaber (Mannerheim), the most common North American species,
occurs throughout the spruce forests of the continent north of a line from North
Carolina to New Mexico. Spruces are preferred hosts, but pines and larches are also
attacked. Infestations occur in felled trees, stumps, and the trunks of standing trees.

' The female adult is reddish brown to black, has the frons pubescent, and is from 2.5
to 3.3 mm long.

Dryocoetes autographus (Ratzeburg) is widely distributed in the coniferous
forests of North America. Infestations are usually found at the base and in the roots
of dying or injured standing trees, or in stumps or felled trees. A wide variety of

361

trees are attacked including spruce, hemlock, Fraser fir, and pine. There have also
been reports of infestations in yellow-poplar. The adult is from 3.5 to 5 mm long. It
differs from the adults of all other species in the genus in having a distinctly
punctured pronotal disk. a convex declivity with the sutural interspace only slightly
raised, and in the absence of a dense mat of hair on the female frons.

The birch bark beetle, D. betulae Hopkins, occurs from coast to coast in
Canada and south to Florida and Mississippi in the Eastern States. Its hosts are
recorded as birch, beech, sweetgum, cherry, and pear. Attacks are generally
confined to dead and dying trees, logs, and stumps. The adult female is reddish
brown, has a dense mat of hairs on the frons, and is from 2.8 to 4.5 mm long.

Dryocoetes caryi Hopkins occurs throughout the northern coniferous forests and
south to North Carolina in the Eastern States. Its hosts are white. red. and
Engelmann spruces. Infestations are usually confined to the trunks of small,
weakened. shaded-out, suppressed trees. The female adult is reddish brown and
from 2.1 to 2.7 mm long.

Dryocoetes granicollis (LeConte), a rather rare species, breeds in spruce from
Quebec to North Carolina. The female adult is reddish brown and from 2.3 to 3 mm
long.

The genus Crypturgus is represented in eastern forests by three common species,
none of which is economically important. The adults are brown or black, about |
mm long, and are found in the inner bark of dead or dying conifers. Their short.
irregular burrows usually originate from the burrows of larger bark beetles, but
sometimes from ventilation holes made by Monochamus spp. C. alutaceus Sch-
warz, the smallest of all North American bark beetles, occurs from New Jersey to
Florida. It breeds in species of pine, and in black and Norway spruces. C. borealis
Swaine breeds in various conifers from Maine to Pennsylvania. C. pusillus
(Gyllenhal), a species introduced from Europe, attacks pine, spruce, balsam fir, and
larch in eastern Canada and from Maine and New York to West Virginia and the
Lake States.

The genus Pseudopitvophthorus is closely allied to Pityophthorus, but differs in
that the adults have a longer and more acute prosternal process (//8). The first
segment of the antennal club also is longer than those in Pityophthorus, and the
males rather than the females have long, yellowish hairs on the front of the head.
The majority of species prefer to breed in the inner bark of recently cut or dying
limbs of various species of oaks. A few species attack other tree species, and some
attack and kill perfectly healthy limbs.

Pseudopityophthorus minutissimus (Zimmermann), the oak bark beetle. a com-
mon and widely distributed species from Quebec and Massachusetts to Georgia and
westward to Mississippi and Colorado, breeds in various species of oaks and
occasionally in many other hardwoods. The adult is dark reddish-brown, from 1.5
to 1.9 mm long. and can be distinguished from other species in the genus by the
reticulate frons and pronotal disk in both sexes. In the southern portions of its
range, it is active throughout the year, and there are at least two generations per year
as far north as the Lake States. Eggs are laid in circumferential galleries in branches
from 12 to 100 mm in diameter and the larvae tunnel away from the gallery.
following the grain of the wood. When the adults emerge. they fly to the tops of the
trees and feed on the buds, in twig crotches, in the axils of leaves, and in immature
acorns. Because of these habits and because of the abundance of the species in
stands suffering from oak wilt disease, this species is strongly suspected of playing
a primary role in the transmission of the fungus (/68).

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Other common eastern species of Pseudopityophthorus are: P. pruinosus
(Eichhoff)—breeds primarily in dead and dying oaks; also in beech, American
hornbeam, hickory, maple, and hophornbeam. Smooth-barked branches in the
upper parts of standing trees are preferred, but slash and other recently felled
material also are attacked. The adult is dark brown and about 1.5 to 1.8 mm long.
This species also is suspected of transmitting oak wilt fungus (/023). P. pubescens
Blackman—breeds in various oaks, American hornbeam, and chestnut in the
Southeast. The adult is dark brown, about 1.8 mm long, and the male has rows of
very long hairs on the elytra. P. asperulus (LeConte)—breeds in various species of
oaks, chestnut, and gray birch from Maine to Florida and Texas. The adult is dark
reddish-brown and from | to 1.5 mm long.

The genus Cryphalus consists of small, dull, dark-brown to black beetles about 2
mm or less in length. They usually breed in small, suppressed trees but may also be
in twigs, branches, and seedlings. C. fraseri Hopkins attacks Fraser fir in the
Southern Appalachians and balsam fir throughout the Northeastern States and in
eastern Canada. C. ruficollis Hopkins is found commonly in red and white spruces
in Maine and New York. C. rubentis Hopkins has been collected from red spruce in
West Virginia.

The genus Pityophthorus is represented by more than 100 species in North
America, many of which occur in eastern forests. The majority of species breed in
the inner bark and central pith of twigs or small branches, but a few may be found
in larger material. Most species utilize conifers, especially pines; a few species
attack broadleaf trees. 3

Gallery systems usually consist of 2 to 4 egg tunnels radiating from a central
nuptial chamber, although the pith-feeding species construct an unbranched system.
All species deposit eggs individually in niches. Material colonized by
Pityophthorus is usually unthrifty, cut or broken, except that certain pith-feeding
species may bore into healthy twigs.

Despite its pervasiveness, this group of insects causes little economic injury
except killing of twigs on drought-stressed saplings, and infesting scions on special
grafted planting stock.

Adults are very small (1.3 to 2.0 mm long), shiny, and have a distinct elytral
declivity that may be armed with tiny spines. Females usually have a frontal brush
of yellow hairs while males do rot (//5, 1/6). Identification of species in this
difficult genus is usually a task for experts. Detailed information on Pityophthorus
can be found in a generic revision (/54) or in a monograph on the North American
Scolytidae (1356).

Pityophthorus lautus Eichhoff breeds in redbud, sumac, and a variety of other
hosts from Minnesota and Quebec to Missouri and North Carolina. P. liqguidam-
barus Blackman breeds in sweetgum, probably throughout the natural range of this
tree. P. crinalis Blackman breeds in poison-sumac and P. scriptor Blackman breeds
in staghorn sumac.

Pityophthorus pulicarius (Zimmermann) occurs throughout the Eastern United
States and in eastern Canada. It attacks all species of pines in its range and has also
been recorded from deodar cedar. Infestations occur in the wood and pith of twigs
of dead and dying trees, in small trees scorched by fire, in slash, and in |-year-old
cones of felled pines. Scions of grafted slash pines being prepared for seed orchard
establishment in Florida have been seriously injured.

Additional eastern species of Pityophthorus and their hosts are as follows: P.
opaculus LeConte—eastern white pine, tamarack, balsam fir, and various spruces

363

from Maine to West Virginia and west to California; P. biovalis Blackman—various
spruces and pines in New York, Maine, and Michigan; P. balsameus Blackman—
balsam fir, spruces, and pines from Michigan to Maine and south to North Carolina;
P. dentifrons Blackman—red and white spruces from Minnesota to Maine and
North Carolina; P. cariniceps LeConte—balsam fir, pines, and spruces in the
northern tier of States and southern Canada; P. annectens LeConte—living trees
and slash of various pines from West Virginia to Florida and Texas; P. intextus
Swaine—white and red spruces from British Columbia to Newfoundland in Canada
south to West Virginia; P. pulchellus Eichhoff—probably all species of pines from
Maine to North Carolina and Texas, also red spruce and balsam fir; P. puberulus
(LeConte)—all species of conifers from British Columbia to New Brunswick in
Canada to Kansas, Texas, and Florida; and P. ramiperda Swaine—tred and eastern
white pines in eastern Canada and from Maine to the Lake States.

The genus Pityoborus contains only one eastern species, P. comatus (Zimmer-
mann). It is widely distributed in the Southeastern States, but appears to be most
common in the Mississippi area. Its known hosts are shortleaf, loblolly, longleaf,
and slash pines. It breeds beneath the bark on the undersides of living but weakened
branches of its host and is of little or no importance. Egg galleries radiate in any
direction, deeply engraving the wood. The larvae construct wide, short, connecting
galleries. The adult is brown to black with yellowish appendages and is about 1.8
mm long. The female is distinguished by the presence of a patch of fine, dense,
silky hair on each side of the pronotum.

Cone and Seed Beetles
Spermophagous Scolytidae

The genus Conophthorus, which best exemplifies spermophagy (cone and seed
feeding) in eastern North America, is very destructive to many species of pines
(547). The close relationship of Conophthorus with twig beetles of the genus
Pityophthorus is indicated by their occasional infestation of new shoots.

Several tropical species that infest seeds of hardwoods occasionally are imported
into North America. For example, Hypothenemus obscurus (F.) is frequently
intercepted in shipments of Brazil nuts.

The adults of Conophthorus are stout, dark brown to black, and about 1.2 to 4
mm long. The female adult bores into a second-year cone at the petiole or base into
the axis, where she constructs a small tunnel and deposits eggs at intervals in niches
along its sides. The larvae feed on the scales, seeds, and tissues of the cone, often
completely honeycombing the interior. Infested cones wither and die before reach-
ing maturity. Conophthorus species usually attack one or a few pine species. If
second-year cones are scarce, first-year cones and current-year shoots may be
attacked. Another way in which these insects survive years in which few cones are
available is that one portion of the population emerges from the infested cone after
the first winter, while other brood adults remain dormant in the cones or in the litter
until the second or possibly the third year. By periodically destroying a majority of
the annual cone production of a species, cone beetles can delay restocking in logged
or burned areas.

Cone beetles and other cone insects perennially destroy most of the seed pro-
duced. For satisfactory yields from seed orchards it is usually necessary to treat the
trees with topical or systemic insecticides. Controlled burning or cleaning the
ground in the fall or early spring should provide substantial relief from infestation
since Conophthorus and most other cone-feeding insects overwinter in fallen cones
or in the ground litter.

364
a ne i A a ed

The white pine cone beetle, C. coniperda (Schwarz), occurs throughout the
natural range of its host, eastern white pine, in eastern America. Apparently it
rarely attacks cones of loblolly pine in Virginia (7354). The adult is shiny black,
from 2.8 to 4.2 mm long, and covered with moderately long, erect hairs.

Winter is spent in the adult stage in infested cones on the ground. These adults
begin to emerge in late April and fly to the tops of nearby pines. When a female
finds a suitable cone, she bores into the petiole or the base of the cone. A dark-
reddish pitch tube marks the point of attack. Once into the cone axis, she bores
toward the cone tip. A male joins her at this time and the tunnel is extended the full
length of the cone (fig. 170). The larvae feed on the seed and tissues until full
grown and then pupate in cells at the ends of the tunnels. Infested cones die and fall
to the ground in mid to late summer. Some adults emerge during the fall but the
majority remain in the cone until the following spring. Some of the fall-emerging
individuals fly to the tops of pines and attack first-year conelets; the remainder stay
on the ground where they feed on other infested cones. There is one generation per
year.

F-505549

Figure 170.—Cone damage by the white pine cone
beetle, Conophthorus coniperda.

_ This is one of the most destructive insect pests of eastern white pine seed. Entire
seed crops in many stands in New England have been completely destroyed (944).
Most of the damage results from the killing of second-year cones; however,
considerable damage also results from attacks on first-year conelets, shoots, and
occasionally buds and male flowers.

The red pine cone beetle, C. resinosae Hopkins, occurs in southeastern Canada
and the northern tier of States from Maine to Minnesota. It breeds by preference in

365

second-year red pine cones, but also attacks current-year red pine shoots and
occasionally second-year jack pine cones. It is also found in cones of Virginia pine
in West Virginia (/356). The adult is shiny black, with sparse, short fine hairs and
is about 3 to 3.5 mm long (758).

Seasonal activity begins in May when the overwintering adults emerge and attack
current-year shoots and second-year cones of red pine. The adults feed for a few
weeks and then attack cones for oviposition purposes. Cones are entered by females
near the petiole on the underside, often forming an open groove at the cone base.
The tunnel is extended in the pith to the end of the cone. Eggs are deposited singly
in niches along the sides of the tunnel. After oviposition is completed, the female
returns to the base, fills the base of the tunnel with a plug of resin and debris, then
vacates the cone. Infested cones soon wither, harden, turn brown, and occasionally
drop from the tree. The larvae feed on seeds and scales in the cone and pupate in
frass-lined cells which often are near the base. During late summer, new adults
emerge either through the plugs in the bases of the tunnels or through the top or
sides of the cones. Soon after emergence they bore into current-year red pine shoots
and tunnel forward through the pith into vegetative buds where they spend the
winter. There is one generation per year. Weakened at point of beetle entry, pine
shoots soon break off and fall to the ground. Damage by C. resinosae is often
severe enough to make the commercial collection of red pine seed impractical or
impossible. Prescribed burning is recommended as a control technique for the red
pine cone beetle (855).

Conophthorus banksianae McPherson, the jack pine tip beetle, a species
closely related to C. resinosae, also occurs in the Lake States and southern Canada.
It breeds in the shoots of jack pine. The adults bore into the shoots about 25 mm
below the bud then tunnel toward the bud, either to feed or to deposit their eggs.
Ovipositing lasts from late May into mid-July. Winter is spent in infested buds, and
there are two generations per year. Infested terminals are often killed, leading to
multiple branching and flat-topping of infested trees. Damage to natural jack pine
reproduction and in jack pine plantations is often severe.

Wood-Boring Beetles
Xylophagus Scolytidae

Genera of Scolytidae that feed on wood occur in both subfamilies (Hylesininae
and Scolytinae). Xylophagus scolytids in eastern North America bore into mori-
bund to dry dead twigs on various hardwood species. Mining by adults and larvae
deeply scores the surface of the wood or is entirely within the wood. A rapid change
in color and texture of the wood near the tunnels suggests that micro-organisms
introduced by the beetles are important in nutrition of the beetles. Species with this
feeding habit cause no economic loss.

Genera included here under this feeding habit include Hypothenemus,
Hylocurus, Micracis, Micracisella, and Thysanoes. A few species of Pityophthorus
may be truly xylophagus but this genus is grouped with the bark beetles because
tunneling in the wood of twigs by some species appears to be incidental to their
feeding in the bark and pith.

The genus Micracis contains a number of species that breed in the wood or pith
of their hosts. Adults are similar to those of the genus Hylocurus but differ in having
the first joint of the antennae flattened and ornamented with long hairs.

Micracis swainei Blackman, a widely distributed species in the South and
Southwest, breeds in poplar shoots and in dead and dying twigs of redbud and

366

willow. The adult is brown and about 2.7 mm long. The terminal hook of the
anterior tibia is strongly curved. M. suturalis LeConte breeds in the twigs and small
branches of redbud, walnut, and willow.

The genus Micracisella includes certain species formerly included in the genus
Micracis (123). Female adults have the antennal scape flattened, somewhat ex-
tended laterally, and clothed with long hairs.

Micracisella opacicollis (LeConte) occurs from Minnesota and New York to
North Carolina. It breeds in the pith of dead twigs, in sprouts, and in shoots of oak,
maple, redbud, and baldcypress. The adult is dark brown to black and from 1.7 to
1.9 mm long. M. nanula (LeConte) breeds in oaks and redbud from South Carolina
to Florida and Texas.

The genus Thysanoes is closely allied to the genera Hylocurus, Micracis, and
Micracisella, but differs in having the apex of the elytra broadly rounded. 7.
fimbricornis LeConte breeds in the twigs of red and black oaks, maple, redbud,
hornbeam, hackberry, hickory, and acacia from Pennsylvania to Florida and Texas.
The adult is yellowish brown and from 1.6 to 1.9 mm long. Egg galleries are
constructed almost entirely in the sapwood, nearly encircling the twig in a diagonal
direction just beneath the bark. Larval galleries run parallel with the grain. The
related species, 7. lobdelli Blackman, has been observed breeding in oaks and
maple in Mississippi, Florida, and Georgia, and 7. berchemiae Blackman, in elm,
oaks, and rattan-vine from Virginia to Florida and Texas.

The genus Hylocurus contains a number of species that rear their brood in the
wood of their hosts. In attacking the host, the adults bore directly into the sapwood
or pith, where they construct nuptial chambers. From each chamber, one to several
egg galleries are then extended obliquely through the wood or in several directions
through the pith. The majority of species are of minor importance since they
normally breed in recently cut, dying, or dead limbs of their hosts. Adults are quite
small, 3 mm or less in length. The head is concealed from above by the pronotum,
which is strongly roughened in front; the first joint of the antennae is club-shaped;
and the elytra are elongated and pointed at the apex.

Aylocurus rudis (LeConte) breeds in the twigs and branches of hickory, walnut,
maple, and hackberry throughout much of the Eastern United States. The adult is
dark brown to black and 2.2 to 3.0 mm long. H. spadix Blackman, H. biorbis
(Blackman), H. bicornus (Blackman), and H. harnedi (Blackman) breed in hickory.

Hylocurus langstoni (Blackman) occurs in the South from the East Coast to
Texas. It breeds in the limbs and trunks of dying or recently killed honeylocust,
hackberry, mulberry, and slippery elm. Green poles and posts are also frequently
attacked and damaged. The damage resembles that caused by powderpost beetles.

The genus Hypothenemus contains numerous species that breed in dying and
dead twigs, dead bark, seeds, and hulls. Included in the genus are species formerly
assigned to the genus Stephanoderes. A few of the seed-infesting species may be
injurious at times.

Hypothenemus dissimilis (Zimmermann) 1s widely distributed in eastern America
from Quebec to Florida and west to Michigan. It breeds in dying branches and dead
twigs of various hardwoods such as the hickories, oaks, honeylocust, hornbeam,
and redbud. The adult is dark brown to black and from 1.2 to 1.8 mm long. They
bore deep entrance holes, and one or more elongate tunnels are constructed more or
less parallel to the grain. They may be found in the sapwood just beneath the bark,
deep in the sapwood, or in the pith. Where numerous, they may honeycomb the
wood.

367

Hypothenemus rotundicollis (Eichhoff) occurs in the Southeastern States and
breeds in the limbs of oaks and hickories. The adult is dark brown to black and
about 1.6 to 1.8 mm long. H. quercus (Hopkins) attacks various oaks,
hophornbeam, honeylocust, and hickories in the Southeastern States. The adult is
dark brown and from 1.5 to 1.9 mm long. H. chapuisi (Eichhoff) breeds in
sassatras and redbud. It has been recorded from Texas, Mississippi, Georgia, and
North Carolina. The adult is dark brown to black and about 1.9 mm long. H.
interstitialis (Hopkins) attacks oaks, hickory, walnut, and many other hardwoods.
It has been recorded from Connecticut and Kansas to Florida. The adult is dark
brown to black and from 0.9 to 1.6 mm long. There is a slightly reddish rugose area
on the pronotum. The adult is dark brown and from 1!.1 to 1.6 mm long. H.
obscurus (F.) 1s frequently intercepted in shipments of Brazil nuts. H. eruditus
Westwood attacks dogwood, redbud, hickory, and black cherry from Michigan and
New Jersey south to Argentina. The adult is dark brown to black and from 1.1 to
1.3 mm long.

Ambrosia Beetles
Xylomycetephagus Scolytidae

Numerous species of beetles in the families Scolytidae and Platypodidae are
known as ambrosia beetles because, in all cases, both the adults and larvae feed on a
mold-type of fungus known as “‘ambrosia.”’ The beetles introduce this fungus into
tunnels bored into the sapwood and sometimes heartwood of trees and logs, where
it grows on the walls and is propagated. Female ambrosia beetles possess spe-
cialized structures called mycetangia (479), which are variously located in and on
the body of the insect. In a few species these organs are found in the male and, in at
least one species, Xyloterinus politus (Say), 1n both sexes (2). Since the discovery
that these specialized organs are possessed by ambrosia beetles, much important
knowledge about the relationship of beetles to their specific microsymbiotic com-
plexes has been gained (4/, 67, 392, 393, 441).

More than 54 genera of ambrosia beetles, some of which include up to 1,200
species, have been recorded throughout the world. A number of species breed in
living trees, but decadent, dying, or recently cut trees, logs, and pulpwood or
stumps are usually preferred. All species require a considerable amount of moisture
for development. In the Southern States, timber is not attacked unless the moisture
content of the wood is at least 48 percent. Seasoned timber is never infested (205).

Ambrosia beetles are important chiefly because of the degrade of sawed lumber
that results from their invasion of trees or logs. This degrade is caused both by holes
bored into the wood and by the presence of black stains caused by the fungus
inhabiting the tunnels. Trees cut during the summer in the South and left for more
than 2 weeks in the woods are often severely damaged. This is especially true of
sweetgum, baldcypress, and oak logs.

There are four general types of ambrosia beetle tunnels: simple, branched,
compound, and cave. Simple tunnels are unbranched, often penetrating deeply into
the wood. Branched tunnels penetrate deeply into the wood and then break up into
several branches that extend in various directions on the same plane. Compound
tunnels also branch off from a single entrance gallery but have egg niches extending
upward and downward from the tunnel. In the cave-type gallery, a simple tunnel
extends into the wood where it is expanded vertically upward and downward into a
cavelike chamber in which the eggs are laid. One of the most common is the cave-
type cavity made by Xyleborinus saxeseni. As these tunnels are excavated, the

368

beetles push the boring dust to the outside. The larvae of certain species live and
feed in the egg niches which they expand into so-called cradles that branch off from
the main galleries. Others live in the main galleries. The larvae of some species
extend the galleries and cradles made by the parent beetles. Ambrosia beetle
galleries differ from those of other wood-boring insects in that they are of uniform
diameter throughout, are free of borings or other refuse, and have their walls stained
black or brown.

Ambrosia beetles are discussed at greater length in several publications (67, 76,
2014214221019, 800, 1232, 1356).

The Columbian timber beetle, Corthylus columbianus Hopkins, occurs from
Kansas to Massachusetts south to Florida and Arkansas. It breeds in various living
deciduous trees such as sugar, silver, and red maples, sycamore, yellow-poplar,
boxelder, basswood, beech, elm, yellow birch, and several species of oaks. Adults
are robust, very dark brown to black, and about 3.6 to 3.8 mm long (fig. 171A).
The front of the head of the female is broadly concave and covered with short, stiff
hairs. That of the male is convex and almost hairless. The pronotum is broadly
rounded and asperate in front. The elytra are shiny, striate, and coarsely and
shallowly punctured. The declivity is armed with small tubercles.

erseensee

SS
SENG

B ‘ | ue
F-531258

Figure 171.—The Columbian timber beetle, Corthylus
columbianus: A, adult; B, callow adults in tunnels in
the wood of a living red maple.

369

Adults become active during May and June in Indiana and West Virginia (648).
They tend to reattack the tree in which they develop, but some dispersal occurs
(933). The wood is entered through bark crevices, usually on the main trunk near
the base. In oak, a tunnel is bored straight into the sapwood until it nears the
heartwood, then it turns right or left (fig. 171B), whereas in diffuse porous wood
such as maple and yellow-poplar, the tunnel branches but then extends toward the
pith. Entrance holes are clean-cut and about 2 mm in diameter, with boring dust and
sap exuding from them. Short tunnels or chambers leading from the upper and
lower surfaces of the main tunnels are excavated at intervals. Eggs are laid in the
chambers and the larvae live and develop in them. The larval food is a yeast of the
genus Pichia (649) and another fungus, Ambrosiella xylebori Brader (68). Food is
stored and transmitted by prothoracic mycetangia possessed by the male beetle
(450). Winter is spent in the adult stage in short tunnels under bark scales or vines
at the base of the tree (932). There are two to three generations per year.

The Columbian timber beetle seems to prefer vigorous trees, and it attacks trees
of practically all sizes. Damage is conspicuous in cross sections of the trunk of
infested trees. Streaks of stain originating from the tunnels extend, often for
considerable distances, above and below them. These and the black-stained tunnels
cause defects known variously as “‘grease spots,” “‘steamboats,”’ “spot worm,”
‘“flag-worms,”’ and “black holes.’ Damaged wood is rendered unfit for such uses
as face veneer, cooperage, or furniture. In southern Indiana, red and silver maple
woods, which are highly valued in the furniture industry, are reduced in value by 38
percent (SOQ).

Corthylus punctatissimus (Zimmermann), the pitted ambrosia beetle, occurs
from southern Canada to Georgia and westward to Colorado. It breeds in a variety
of trees and shrubs, such as maple, dogwood, American hornbeam, eastern
hophornbeam, sassafras, rhododendron, and azalea. Young sugar maples are es-
pecially subject to damage, and destructive infestations in them have been reported
both in North Carolina and southern Canada. Cultivated rhododendrons and azaleas
are also frequently attacked and killed. The adult is rather stout, cylindrical, dark
brown or black, and 3.0 to 3.3 mm long. The antennae and legs are rusty red-
brown. The prothorax is longer than wide, roughly tuberculate in front, finely and
sparsely punctured, shiny behind, and extends hoodlike over the head. The elytra
are strongly punctured but not in rows, are rounded behind, and are without furrows
or teeth.

The adult bores into its host near the ground line, then excavates a tunnel which
may encircle the stem one or more times, girdling it. Small stems, from 3 to 12 mm
in diameter, are usually attacked. Nearly all attacked stems die and annual mortality
of 8 percent has been recorded in southeastern Canada (4/9).

Two species of the genus Monarthrum, M. fasciatum (Say) and M. mali (Fitch),
occur in eastern North America. Adults of M. fasciatum are yellowish brown with
the anterior half of the prothorax and the posterior third of the elytra usually dark
brown, and are about 2.3 to 2.8 mm long. In this species, mycetangia are possessed
by the female (753). This species is most common in the South but occurs as far
north as Massachusetts and Wisconsin. It breeds in many species of hardwoods,
and has also been observed in pine. Adults of M. mali resemble those of M.
fasciatum except for their slightly smaller size and their uniform yellowish to light
reddish-brown color. This species breeds in injured and dying trees or recently cut
logs and stumps of many species of hardwoods throughout the Eastern States. The
galleries of both species of Monarthrum are branched with larval chambers extend-

370

ing upward and downward from the tunnels (20/). Both species are highly destruc-
tive of green lumber and fresh logs of gum in the Gulf States.

Xyloterinus politus (Say), a widely distributed species in eastern North America,
breeds in injured, dying, and recently cut trees and limbs of a variety of trees such
as beech, birch, hard and soft maples, hickory, oak, ash, magnolia, black cherry,
red spruce, pine, and hemlock. It is rare in coniferous hosts. The adult is dark
brown to black and about 2.7 to 3.7 mm long. The pronotum is almost square,
rugose in front, and has the anterior margin armed with four teeth. The galleries of
this species differ from those of many other ambrosia beetles in that they often fork
and branch secondarily, and they also possess four rows of larval cradles, two above
and two below the gallery (800, SO/). Lumber cut from infested wood may be
severely degraded by adult entrance holes and by associated stains.

Five species of the genus Trypodendron occur in eastern North America. Adults
are distinguished by having completely divided eyes and by the absence of distinct
sutures in the antennal club. The frons of the male is broadly excavated; that of the
female is convex. The oral region of the head is visible from above. The gallery
penetrates the sapwood and branches one or more times. The larvae are reared in
cradles extending upward and downward from the tunnels. The cradles are enlarged
by the larvae and serve as pupation chambers (/356).

The striped ambrosia beetle, 7. lineatum (Olivier), occurs over much of
Canada, and the Western and Northeastern United States. It also occurs in the
mountains of western North Carolina. It breeds in a wide variety of conifers and
occurs rarely in hardwoods. The adult is from 2.7 to 3.5 mm long. They are dark
brown to black, with each elytron usually marked with two light yellowish-brown
stripes parallel to the suture. These stripes extend to the base of the pronotum. The
main gallery of the species extends straight into the wood for 2.5 to 5 cm and then
divides into two or more branches. Larval cradles are situated at the upper and lower
surfaces of these branches. Damage to felled timber, and to damaged, injured, or
fire-scorched trees is often severe.

Trypodendron scabricollis (LeConte) occurs from Minnesota and Arkansas east-
ward, and breeds in various species of pine and hemlock. The trunks (fig. 172) and
larger limbs of weakened and dying pines are preferred, but freshly cut lumber is
also subject to attack and serious damage. Adults are from 3.5 to 4.1 mm long, and
brown with a faint, light stripe extending from the basal one-fourth of the pronotum
down each elytron. The elytra are smooth and finely striate. The gallery system has
an entrance tunnel extending 2.5 to 5.0 cm into the wood. The gallery branches left
and right, following an annual ring. Larval cradles extend above and below these
secondary tunnels (76).

Trypodendron retusum (LeConte), the largest of the eastern species, attacks
poplars across the Northern United States and southern Canada. It has also been
recorded from West Virginia and several Western States from Alaska to Arizona.
Adults are 3.6 to 4.6 mm long and uniformly black when fully mature except for
pale yellowish-brown areas at the base of the pronotum and on the elytra from the
base to the declivital margin continuing on the sides of the declivity to the apex. T.
betulae Swaine occurs throughout the Northern States and across Canada, and
breeds in unthrifty standing birch. Adults are dark brown to black and from 2.7 to
3.5 mm long. There is a faint yellowish-brown area of variable size on each elytron
from the base to the declivital margin (/356).

Trypodendron rufitarsis (Kirby) attacks injured or dying pines and spruces,
evidently preferring standing trees over logs. It occurs over much of western North

371

F-532334, F-532335
Figure 172.—Two examples of damage from tunnels of
the ambrosia beetle, Trypodendron scabricollis, in
loblolly pine.

America but it has been found in northern Minnesota and eastern Canada (1/356).

The genus Gnathotrichus contains several species of true ambrosia beetles (//7).
The adults are small, cylindrical, and dark brown or black. The head is invisible
from above, and the body surface is finely punctured, smooth, and sparsely covered
with hairs near the elytral declivity.

Gnathotrichus materiarius (Fitch) occurs from eastern Canada south to Florida
and westward to South Dakota and eastern Texas. It breeds in the lower portions of
the trunks of dead and dying pines, spruce, balsam fir, larch, and perhaps other
conifers. Adults are dark brown and about 1.7 to 3.1 mm long. The elytra are
glabrous except for the declivity and sides which have short, sparse hairs. The
declivity of the elytra is grooved at the suture.

Adults bore directly into the wood and their main galleries may have several
branches (fig. 173). Larval cradles are extended both upward and downward from
these galleries, and run parallel to the grain of the wood. Pines killed by /ps bark
beetles in the South are often attacked by this species (76).

The genus Xyleborus is represented by 19 native and introduced species in the
Eastern United States (752, 1356). All species are ambrosia beetles. Many breed in
both conifers and deciduous trees and shrubs of all sizes. Dying, unhealthy, felled,
or weakened trees or wounds and dead areas in living trees are preferred for attack.
Males are very rare, quite different from females, and flightless.

342

Courtesy Duke Univ. Sch. For.
Figure 173.—Tunnels of the ambrosia beetle,

Gnathotrichus materiarius, in the wood of a shortleaf

pine.

Xyleborus celsus Eichhoff, the largest member of the genus occurring in the
United States, breeds in dead, dying, and recently felled trees and stumps of
hickory in the Eastern United States west to lowa and Texas. Females are reddish
brown and about 3.6 to 4.5 mm long. The declivity of the female is steep, almost
flat. The sides and upper margin of the declivity are armed with several acute
tubercles, and the area between the suture and first stria is broad, smooth, and
armed with two large teeth. The galleries of this species extend directly into the
wood for | to 3 cm or more, then branch one to six times in a fan shape. Eggs are
laid near the ends of the galleries. There are two generations per year in southern
Missouri (459). Hickories killed by the hickory bark beetle are especially subject to
attack in the Southeast (76).

Xyleborus dispar (F.) is widely distributed in southern Canada and the Eastern
States south to North Carolina. It attacks all the common fruit trees as well as a
great many other hardwood trees. It has also been reported from pine and hemlock.
Females are black and 2.8 to 3.5 mm long. In large trunks, the galleries extend
straight into the wood from | to 3 cm and branch on the same plane following an
annual ring. In small trunks and branches, the galleries spiral upward from the
point of attack around the stem or limb, branching upward or downward from the
spiral gallery (20/).

Xyleborus ferrugineus (F.) occurs from Massachusetts and Michigan south to
Florida and Texas. It breeds in dead, dying, or felled trees of a wide range of
species including oak, hickory, ash, baldcypress, walnut, pine, beech, and sweet-
gum. Females are reddish brown and from 2 to 3.3 mm long. The elytral declivity
of the female is flat. The third interspace of the declivity is armed with one to three
acute granules on the basal half and one coarse denticle at the middle; the fourth
and fifth interspaces are armed with two to four granules each. Galleries resemble
those formed by X. ce/sus except that they are smaller in diameter; they also may be
longer and more winding and branch less frequently in the same plane. Side
branches are formed which lead to other sets of galleries at different levels. The
galleries may penetrate all of the sapwood but they are less common in the
heartwood. X. ferrugineus may form shallow galleries that are visible on the surface
of the wood when the bark is peeled (/356).

373

Xyleborus affinis Eichhoff occurs in the Eastern United States east of a line from
Missouri to Texas and south of Michigan and Massachusetts. It breeds in dying
trees and in green logs and lumber of various hardwood trees such as oak, hickory,
sweetgum, river birch, hackberry, silktree, persimmon, baldcypress, and black
locust. Adults are yellowish to reddish brown and are from 2.0 to 2.7 mm long. The
elytral declivity of the female is dull, opaque, and broadly sloping, and the
interspaces are armed with a few minute granules. Galleries are similar to those of
X. ferrugineus except surface galleries are more commonly found and usually more
extensive. Adjacent tunnels are often interconnected, and there are no cradles (76,
1356):

Other eastern members of the genus and their hosts are as follows: X. xy-
lographus (Say)—oaks; X. pubescens Zimmermann—pines; X. intrusus
Blandford—pines; X. obliquus (LeConte)—birch, hickory, and chestnut; X.
rubricollis Eichhoff—oak, hickory, walnut, dogwood; X. /econtei (Hopkins)—
hickory and palm in Florida; X. tachygraphus Zimmermann, X. obesus LeConte
(the stout ambrosia beetle), and X. sayi (Hopkins)—a wide variety of deciduous
trees; X. volvulus (F.)—probably various deciduous trees and shrubs in southern
Florida; X. devexulus Wood—hickory; X. validus Eichhoff—beech and black oak;
and X. planicollis Zimmermann, X. viduus Eichhoff, and X. opimus Wood—for
which the hosts are unknown.

Xyleborinus saxeseni (Ratzeburg) occurs commonly throughout southern Canada
and the United States and breeds in a wide variety of trees. Some of its more
important eastern hosts are pecan, hickory, honeylocust, walnut, sweetgum,
yellow-poplar, oak, American beech, maple, birch, dogwood, persimmon, holly,
hemlock, baldcypress, and shortleaf and loblolly pines. The adult is dark brown
and from 1.6 to 2.4 mm long. It is distinguished from Xyleborus spp. by its conical
scutellum. The elytral declivity of the female is steep, convex, shallowly bisulcate,
and armed with two rows of five to seven acute tubercles on each side of the suture.
The male is smaller with all characters poorly formed. The gallery of this species
consists of a single tunnel bored | to 7 cm directly into the wood. The innermost
section of the tunnel is widened vertically upward and downward into a cavelike
chamber in which eggs are laid and in which the larvae live and feed. The larvae
feed on ambrosial fungi and wood, enlarging the tabular tunnel of the female. The
life cycle 1s completed in 2 months. This species can cause severe economic
damage (/356).

The genus Xy/osandrus is represented in the United States by five species, of
which at least four are introduced. They usually breed in twigs, branches, and small
stems of trees and other plants (/356). The female excavates a tunnel into the pith of
small stems or into the wood for | to 3 cm. Here a small cavity is made where the
eggs are laid. The larvae feed on ambrosial fungi growing on the walls of the tunnel
and also apparently on the host tissue as they enlarge the gallery.

Xylosandrus zimmermanni (Hopkins) has been recorded in southern Florida,
Mexico, and northern South America. Its hosts are listed as red maple and ardisia
species. X. germanus (Blandford) breeds in the branches, logs, and stumps of a
wide variety of hosts including maple, oak, hickory, beech, dogwood, elm, ash,
other hardwoods, and pine. It occurs from Connecticut and New Jersey west to
Illinois and Kentucky. Heavy infestations have been found in elms killed by the
Dutch elm disease. It is capable of transmitting Dutch elm disease fungus to healthy
trees (167). It has also been found associated with Fusarium cankers on yellow-
poplar and black walnut (22).

374

The black twig borer, X. compactus (Eichhoff), introduced from southeast
Asia, was first detected in Florida and Cuba in 1941. It has invaded Georgia,
Mississippi, and Louisiana, and will probably spread throughout much of the
Southeastern United States. Vigorous terminal twigs of more than 200 known host
species are attacked and killed. Hosts include maples, hickories, magnolias, oaks,
willows, and many others. This aggressive species is a serious pest in deciduous
forests and in horticultural areas. In Florida, smaller twigs (1 to 7 mm in diameter)
of dogwood were infested by single females, and larger twigs contained several
females (927).

Xylosandrus morigerus (Blandford), the red shothole borer, was introduced
from southeast Asia, and has spread from Mexico to Brazil. It has been intercepted
at several ports of entry in the United States but it has not yet become established. It
breeds in broken or cut branches about | to 3 cm in diameter (/356).

Xylosandrus crassiusculus (Motschulsky), also from southeast Asia, was found
near Charleston, S.C., in 1975 on cherry. Its galleries resemble those of Xyleborus
dispar (F.) (1356).

Family Platypodidae
Platypodid Beetles

Almost all members of this family are ambrosia beetles and they occur prin-
cipally in the tropics and subtropics. Only one genus has been recorded from the
United States (J355). The adults differ from those of other ambrosia beetles in
having longer and more slender bodies and wide heads flattened in front, not
covered by the pronotum. The first segment of the tarsus 1s as long as the other three
tarsal segments combined, and there are spinelike projections at the seam of the
elytra of the males (20/).

Members of the family are usually more destructive than other ambrosia beetles.
Their tunnels are more extensive, and they extend deeper (25 to 30 cm) into the
sapwood and heartwood. Dying, weakened, or recently felled trees are usually
preferred; however, vigorous, healthy trees are also attacked if dead areas of bark
are present. Eggs are laid in small loose clusters in the tunnels. Larvae and adults
are also found in these tunnels.

Platypus flavicornis (F.) occurs in the Eastern United States from New Jersey to
Florida and west to Texas and Mexico. It breeds commonly in various species of
pines and occasionally in several species of hardwoods. Dead and dying trees,
stumps, and logs cut or left in the woods during the summer are preferred. It is
commonly found in the lower | m of trees killed by southern pine beetle (245).
Adults are reddish brown and about 5 mm long. The front of the head is flat and
clothed with moderately long hairs; the pronotum is longer than broad and densely
but shallowly punctured; and the elytra are elongate and striate, with the third, fifth,
seventh, and ninth interspaces produced into blunt, toothlike processes on the
declivity of the male. The adult bores a horizontal gallery in the sapwood (fig. 174).
Here, it may branch extensively and extend into the heartwood. The lower portions
of infested trees are sometimes literally riddled. In the South, this species is so
abundant that very few dying pines, stumps, or logs escape attack. Large amounts
of white downy frass is evidence of attack (76).

Platypus quadridentatus (Olivier) occurs throughout the South from Florida to
Texas and north to West Virginia. Various species of hardwoods, especially the
oaks, are most commonly attacked. Adults are dark reddish brown and about 4.5
mm long. The front of the head is shallowly and densely punctured and sparsely
clothed with moderately long hairs. The pronotum is longer than broad, and in the

375

i
e
=

et
F-532822

Figure 174.—Egg cradles of the ambrosia beetle,
Platypus flavicornis.

female, it bears two large pits just behind the middle. The third, fifth, and seventh
interspaces of the elytra are produced into toothlike processes on the elytral
declivity of the male. Two large tuberosities also occur on the lower edge of the
declivity in the male, and two hooklike spines are on the fourth abdominal segment
(76).

Platypus compositus (Say) occurs throughout the Southern States northward to
southern New York and southern I[Ilinois. It breeds in a wide variety of deciduous
trees such as hickory, pecan, birch, poplar, oak, chestnut, basswood, elm, beech,
sweetgum, tupelo, magnolia, baldcypress, and persimmon (fig. 175). Recently
felled or girdled baldcypress is often seriously damaged. Adults are light reddish-
brown and about 4.5 mm long. The front of the head is densely punctured above,
and there are two centrally located pits just behind the middle of the pronotum. The
first. third, fifth, and seventh interspaces of the elytra of the male are produced into
small tubercles on the declivity. The declivity of the male also bears two large
tridentate teeth at the outer apical angle (76).

Courtesy Duke Univ. Sch. For.

Figure 175.—Tunnels and larval cradles
of the ambrosia beetle, Platypus
compositus, in the wood of persimmon.

376

Platypus parallelus (F.) apparently is the most destructive ambrosia beetle in the
world. Although it occurs in Florida, Texas, and southern California, the unfavora-
ble climate prevents this tropical species from becoming sufficiently abundant to
cause more than minor damage to hardwood trees in North America.

Ambrosia beetle control is largely a matter of preventing damage to recently cut
logs through quick removal from the woods, proper storage of logs at the mill, and
by proper handling of lumber and milled products (76).

Order Hymenoptera—Ants, Bees, Sawflies, Wasps,
and Allies

The order Hymenoptera is one of the largest orders of insects, with more than
20,000 species in America north of Mexico. Most are beneficial and many are very
important to forestry and agriculture either as parasites or predators of pests or as
pollinators of more than 100 commercially grown crops. The honey bee is one
important pollinator. Among the relatively few injurious Hymenoptera are the
sawflies, some of which are stem borers attacking crops or serious defoliators of
coniferous trees (697).

The winged members of the order characteristically have four membranous
wings, the front pair larger and more completely veined than the hind pair. The
hindwings have a row of tiny hooks along the anterior margin by which they are
attached to the forewings. Some forms, or groups, such as workers of the common
ant, are wingless. The ovipositor is usually well developed and in the higher forms
is modified into a sting.

The order is divided into two suborders—Apocrita (= Clistogastra) and Sym-
phyta (= Chalastogastra) (697). Each of these, in turn, is divided into a number of
superfamilies.

Suborder Symphyta (= Chalastogastra)

Members of the suborder Symphyta are distinguished by having the abdomen of
the adult broadly joined to the thorax—the second abdominal segment is not
constricted into a petiole as in the suborder Apocrita. The ovipositor of the female
is well developed and fitted for making incisions in the leaves or stems of plants. In
the majority of species it is sawlike. Because of this, these members of the suborder
are known as sawflies.

The larvae of all species, except for the family Orussidae, are phytophagous, the
majority feeding externally on the foliage. The remainder bore into stems, fruit,
wood, or leaves, and some form galls. Orussid larvae are parasitic on wood-boring
coleopterous larvae. While externally leaf-feeding larvae look like lepidopterous
larvae, they have only one simple eye on each side of the head and have six or more
pairs of prolegs on the abdomen, none of which bears hooks or “‘crochets.”

The suborder Symphyta is represented in the United States and Canada by more
than 1,000 species, many of which are highly destructive of forest and shade trees
and of young trees in nurseries and plantations.

Superfamily Megalodontoidea—Family Xyelidae
Xyelids

Members of the family Xyelidae are medium-size to small sawflies, mostly less
than 19 mm long. They differ from all other sawflies in having the third antennal
segment elongate with a slender terminal filament of 9 to 25 segments, and in
having three marginal cells in the forewing. Unlike all other sawflies except the
Pamphiliidae, they have the costal cell divided by a longitudinal vein.

SW

The genus Xyela contains several species that feed as larvae on the developing
pollen of the male strobili of various species of pines and one species forms shoot
galls on several species of pines. The adults are often found feeding on pollen of
catkins of willow, alder, and other trees in flower during the adult flight period, and
occasionally on the pollen of pines. Only 17 species are known in North America,
of which 7 are found in eastern pine forests. X. bakeri Konow, reared or collected
from longleaf, slash, jack, Virginia, and shortleaf pines, is known to occur from
Quebec to Florida, westward to British Columbia, and California. X. minor Nor-
ton, whose known hosts include Virginia, slash, loblolly, and longleaf pines,
ranges from Quebec to Florida, westward to British Columbia, and California. X.
obscura (Strobl), reared or collected from Virginia, -jack, slash, longleaf, and
loblolly pines, is known from Newfoundland to Florida westward to California,
Northwest Territories, British Columbia. X. dodgei Greenbaum, collected beneath
sand pine, is known only from Cedar Key, Fla. X. alpigena (Strobl), whose known
hosts are white pines, ranges from Quebec, Maine, to Maryland, westward to Utah,
New Mexico. X. styrax Burdick, collected from Virginia pine is known from New
York, Maryland, Virginia, and Georgia. X. gallicaulis Smith, the pine shoot gall
sawfly, is collected and reared from spindle-shaped galls on new shoots of loblolly,
shortleaf, and slash pines in Virginia and Georgia (fig. 176). Adults emerge in
December and oviposit presumably in the dormant buds of host trees. Egg hatch 1s
timed with bud elongation, and the larvae form a gall. Full-grown larvae drop to the
soil where prepupae form cocoons. Adults emerge 19 months later. The cur-
culionid, Conotrachelus caroliniensis Schoof, feeds externally on sawfly galls,
which damages weakened shoots and causes them to break in the wind. Xyelid
sawflies occurring on slash and longleaf pines in Florida are discussed in the
literature (355, 547).

Pleroneura brunneicornis Rohwer is recorded from balsam and white firs from
eastern Canada and from Maine and New York to Minnesota (//03). Mature larvae
are white, measuring 4 to 7 mm long. The thoracic legs are weakly developed and
prolegs are reduced to slight swellings. In New Brunswick, Canada, adults fly
during late May. Eggs are deposited within tightly packed needle clusters on
expanding buds, and larvae feed in tunnels excavated in the center of new shoots.
Mined shoots die and turn reddish brown, an effect similar to that caused by late
frost. Mature larvae drop to the ground and overwinter in thin, oval cocoons spun
within an earthen cell. There is one generation per year (/260).

Several species of Megaxyela have been observed feeding as larvae on hickory
and walnut, and M. langstoni Ross has been reared from pecan. Macroxyela
ferruginea (Say) has been recorded as feeding on elm.

Family Pamphiliidae
Webspinning Sawflies

The family Pamphiliidae is represented in North America by 4 genera and 72 or
more species, of which more than half occur in eastern America. The adults are
moderately large with long, many-segmented antennae. The abdomen is flattened
and has sharp lateral margins; the ovipositor of the female is short. Larvae have the
venter flattened and are about 15 to 25 mm long. The thoracic legs are well
developed, abdominal legs are absent, and the last abdominal segment bears a pair
of three-segmented subanal appendages. The larvae of certain species are gregari-
ous and feed together in nests that they prepare by webbing together leaves of their
hosts. Others roll the edges of leaves or spin silken tubes in which to live (845).

The pine false webworm, Acantholyda erythrocephala (L.), an introduced

378

species first recorded in Pennsylvania in 1925, now occurs in Connecticut, New
Jersey, Pennsylvania, and Ontario. Its preferred hosts are eastern white and red
pines, but it also attacks several other pines, including Scotch, Austrian, Swiss
mountain, and Japanese red. Female adults have orange-red heads and steel-blue
bodies; males are almost entirely steel-blue. Full-grown larvae are pale greenish-
gray and about 16 to 20 mm long. The head is clay yellow with dense, small dark-
brown spots, and there are longitudinal stripes of purplish red on the dorsum,
venter, and sides. The prepupa is bright apple-green.

Winter is spent in the prepupal stage, pupation occurs in early spring, and adults
appear from about mid-April to mid-May. Eggs, in short rows of 3 to 10 each, are
deposited over small slits cut into the previous year’s needles. The larvae spin loose
webbing about themselves and feed gregariously on the old needles. Young larvae
cut the needles off just above the bundle sheaths and pull them into the webbing,
where they are consumed. Older larvae feed singly from within individual silken
tubes spun around themselves along twigs. Considerable amounts of frass and bits
of needles usually adhere to the exterior of these tubes (/333) (fig. 177). Full-grown
larvae drop to and enter the ground for hibernation in an earthern cell. There is one
generation per year. Heavy infestations sometimes develop locally, causing severe
defoliation (505).

F-531259 Courtesy Conn. Agric. Exp. Stn.
Figure 176.—Pine shoot gall sawfly, Figure 177.—Damage and webbing by
Xyela gallicaulis, on new shoot of the pine false webworm,
loblolly pine in Georgia. Acantholyda erythrocephala.

379

Acantholyda zappei (Rohwer), the nesting-pine sawfly, occurs from Quebec to
New Jersey west to Michigan. Its hosts are various species of pines such as red,
jack, Austrian, pitch, and Japanese red. A full-grown larva is green, with a brown
head and a dark-green dorsal stripe, and is about 18 to 25 mm long. Eggs are laid
singly on young needles of the current year’s growth from late June to early July.
Young larvae spin webs about themselves and fasten the outer threads to the
needles. Needles are cut off near the base and drawn into the web where they are
consumed. Webs are increased in size as the larvae develop, and may reach a length
ot 13 cm by the time the larvae are full grown. Winter is spent as prepupae in cells
in the soil, and pupation occurs in the spring. There is one generation per year
(1370).

Acantholyda circumcincta (Klug) has been recorded from New Brunswick,
Quebec, Pennsylvania, Georgia, and Florida. During May and June 1968, it
severely defoliated approximately 40 hectares of sand pine in west Florida. Known
details of its life history are similar to those of A. erythrocephala (203).

Some additional eastern species of Acantholyda, along with known hosts and
known areas of distribution, include the following: A. apicalis Westwood—loblolly
pine in North Carolina, Georgia, Florida, Mississippi, Arkansas; A. pini Rohwer—
red pine in Quebec, Ontario, Michigan, south to Georgia; A. angulata (Mac-
Gillivray)—eastern white, pitch, jack, Austrian, and Japanese red pines in New
Brunswick, Quebec, south to North Carolina, west to Minnesota; A. luteomaculata
(Cresson)—white and jack pines, preferably eastern white, from New Brunswick
to Connecticut, west to Ohio, Minnesota; A. floridana Greenbaum— slash and sand
pines in Florida; A. maculiventris (Norton)—ftrs and white spruce from Labrador
to North Carolina, west to British Columbia, Wyoming, California.

Cephalcia fascipennis (Cresson) occurs in Quebec, Nova Scotia, New
Hampshire, west to British Columbia. Its hosts are listed as blue and white spruces.
Full-grown larvae have a black head and thorax, and a green body. They are about
25 mm long. Ornamental spruce and hedges are sometimes rendered unsightly by
the presence of larval nests (375). C. fulviceps (Rohwer) feeds on jack and red pines
in southern Canada, New Hampshire, Connecticut, and New Jersey. C. marginata
Middlekauff feeds on young red pines in Nova Scotia, Quebec, New York, Mas-
sachusetts, Connecticut, Pennsylvania, Virginia, and West Virginia. Its life history
is discussed (277).

Pamphilius phyllisae Middlekauff is widely distributed in Eastern United States
and Canada (376). Only one outbreak has been recorded for P. phyllisae, and it
occurred in Pennsylvania following a period of several droughty years. At this time
the host severely impeded parasitism by encapsulating nearly all of the parasites
(376). The life cycle may require | or more years, with about 6 weeks spent as
feeding larvae.

The plum webspinning sawfly, Neurotoma inconspicua (Norton), occurs in
Quebec, Massachusetts to Pennsylvania, west to British Columbia, Montana, and
Kansas. Its hosts are listed as hawthorn and various wild piums and wild cherries.
The adults are black with supraocular spots, their wings are hyaline with faint bands
beneath the stigma, and their legs are mostly reddish brown beyond the coxae. The
larvae are gregarious; they web the foliage together to make webs somewhat similar
to those of the fall webworm. Heavily infested trees no more than 1.8 m tall may
support up to 25 webs, some of which may enclose entire branches. Such trees may
be completely defoliated.

380

Neurotoma fasciata (Norton), the cherry webspinning sawfly, occurs from
Quebec to Florida, west to Wisconsin, Missouri, and Arkansas. Its hosts are listed
as black and pin cherries. The full-grown larva has a shiny black head and
prothoracic plates, and a deep green body; it is about 18 mm long. The larvae are
gregarious and construct dirty-brown nests on branches and shoots, especially on
young trees. Prepupae overwinter in cells in the soil and some of these pupate in the
spring. The remainder do not pupate until late fall or the following spring. The
related species, N. crataegi Middlekauff, feeds on hawthorn from Massachusetts
and New York, west to Manitoba, Wisconsin, and Illinois.

Superfamily Tenthredinoidea
Sawflies

Members of the superfamily Tenthredinoidea are commonly called sawflies
because of the sawlike ovipositor of the female. The ovipositor is composed of three
main pieces held within protecting sheaths. The upper piece is rigid and consists of
a pair of fused lances, with ventral grooves along which the other two pieces slide.
These last two are called the lancets, or saws. They consist of thin plates with their
flat inner surfaces together. Each plate is usually shaped like a long, acute triangle
with the dorsal edge along a groove of the lance. The ventral edge is usually
sawtoothed. The rod that attaches the blade to the abdomen and controls its
movements originates at the narrow base of the lance.

Sawfly larvae resemble lepidopterous larvae but are usually naked. A few,
however, are spined, hairy, or covered with a gummy or waxy secretion. The best
single feature for distinguishing sawfly larvae from lepidopterous larvae is the
presence of only one simple eye on each side of the head—lepidopterous larvae
have six on each side. When disturbed, the larvae of certain sawfly species curl up
and lie on their sides, whereas others hold their abdomens aloft over their heads or
raise the head and thorax. The majority are external feeders on foliage, either eating
entire leaves or skeletonizing them. Some feed from within; a few feed within
mines in the leaves, leaf petioles, and twigs; some produce galls on the leaves or
shoots; and a few feed on catkins, buds, or fruits. When they reach maturity they
either spin cocoons or construct cells in which to pupate. Cocoons may be spun on
leaves, twigs, or other parts of the host, or they may be spun in the litter or soil
beneath the trees. Cells are formed in pith, bark, and brashy wood, or in the
ground. Leafmining species sometimes form their cocoons within their mines.

Sawfly adults frequently resemble small bees or wasps, except for their antennae
and the broad connection between thorax and abdomen. The antennae may be
feathered, clubbed, threadlike, forked, or may bear spurs on some of the joints.

Sawflies constitute one of the most destructive groups of insect defoliators in
eastern forests. Outbreaks occur frequently and sometimes spread over large areas
and persist for several years before subsiding. Damage is often severe, especially in
coniferous plantations. Outbreaks of an increasing number of species infesting
pines have been reported during recent years, possibly as a result of the establish-
ment of numerous pine stands in planting programs. Lists of the species in the
superfamily occurring in America north of Mexico have been published (697).
Family Pergidae
Pergid Sawflies

The family Pergidae is represented in Canada and the United States by only one
genus, Acordulecera. Four species are now listed from the Eastern United States
west to Arizona (697). Full-grown larvae are usually greenish with light or blackish
heads and distinct lateral lobes, and measure less than 12 mm long. Each body

381

segment bears transverse rows of slight tubercles armed with short, stiff setae. The
larvae feed gregariously on the foliage of oak, butternut, hickory, hawthorn, and
pecan, and occasionally are abundant enough to attract attention. Adults appear in
May and June; larvae are present from June to August, and overwinter in cocoons in
the ground. There is usually one generation per year, but some species occasionally
have a partial second.

Family Argidae

Argid Sawflies

The family Argidae is represented by 8 genera and 59 species in the United
States and Canada, the majority of which occur in the Southwestern United States
(697). The adults are medium- to small-size, stout-bodied sawflies. They can be
recognized by their three-segmented antennae, the third segment of which is very
long and bifurcate in males of some groups. The more common species are usually
reddish brown or bluish black, with more or less dark-brown wings. Larvae are
yellowish green or red and are usually spotted. The body is often rather thick-set,
widest on abdominal segments one to three, tapering toward the rear end, and the
venter is often flattened.

The birch sawfly, Arge pectoralis (Leach), 1s widely distributed in eastern North
America. Its preferred hosts are various birches, especially gray and paper. It also
has been recorded feeding on willow and alder. Full-grown larvae are about 18 mm
long. The head is reddish yellow; the body yellowish, with six rows of black spots
on top and three on each side. Adults appear during June and July and deposit their
eggs in slits cut in the margins of leaves. Larvae are present from July to September,
and winter is spent as prepupae in cocoons spun in the litter on the ground. The
species is of no economic importance, although it occasionally becomes abundant
enough to cause noticeable defoliation over limited areas.

Additional species of argid sawflies likely to be encountered on trees and shrubs
in the Eastern United States are: A. clavicornis (F.)—on willow, birch, hawthorn,
plum, hornbeam, and azalea; A. scapularis (Klug)—on oak, alder, elm, and birch;
A. coccinea (F.)—on sumac; A. abdominalis (Leach)—on birch and azalea; A.
humeralis (Beauvois)—on poison-ivy and poison-sumac; and Sterictiphora spp.—
on plum and cherry.

Family Cimbicidae
Cimbicid Sawflies

The family Cimbicidae is represented in the United States and Canada by 3
genera and 12 species, some of which have been recorded feeding on trees in the
Eastern States (697). The adults are large with clavate antennae. The tibiae are
without preapical spurs; however, there are single apical spurs on the front tibiae.
Full-grown larvae have large heads, and their bodies taper toward the rear, and they
are sometimes covered with a waxy bloom.

The elm sawfly, Cimbex americana Leach, is the largest of the North American
sawflies. It is widely distributed, occurring from coast to coast in southern Canada
and throughout the United States. The larvae feed primarily on the foliage of elm
and willow, but are also observed occasionally on basswood, birch, maple, poplar,
and alder. The antennae of adults are tinged with orange; the head and thorax,
black; the wings, smoky brown; and they are about 25 mm long. The female’s
abdomen is usually steel blue, with three or four yellowish spots on each side and a
faint, whitish spot near the thorax. In the male, the whitish spot near the thorax is
distinct, but there are no yellowish spots on the sides. Full-grown larvae are
yellowish white or greenish white with a pebbly skin and a black dorsal stripe, and

382

about 43 mm long. While feeding, the larva usually coils its rear end around a stem
or twig; at rest, it usually lies coiled like a snail (fig. 178).

Courtesy Conn. Agric. Exp. Stn.

Figure 178.—-Larva of elm sawlfly,
Cimbex americana.

Adults appear from about mid-May to mid-August, and the eggs are deposited in
pockets cut into leaf tissues. Larvae occur from June until October, depending on
location. When they become full grown, they crawl to the ground and spin tough,
papery cocoons in the litter or just below the surface of the soil. Winter is spent in
the prepupal stage, and pupation occurs in the spring. There is one generation per
year.

The elm sawfly is of minor importance as a defoliator of forest trees, but
occasionally seriously defoliates shade tree elms. Willow also has been heavily
attacked in th. Northern Great Plains area. The larvae partially or entirely defoliate
the trees, while the adults chew the thin, tender bark of twigs, girdling and killing
them (//52).

Trichiosoma triangulum Kirby occurs quite commonly in the Eastern United
States. The larvae are solitary feeders on the foliage of ash, birch, poplar, willow,
and wild cherry. At maturity, they are 37 mm long. The head is creamy white, the
body greenish white, and the eye spots and spiracles blackish. The life cycle is
similar to that of the elm sawfly.

Family Diprionidae
Conifer Sawflies

The family Diprionidae includes many of the most serious defoliators of coni-
fers. The majority of species are native to the continent but several of foreign origin
are well established. The family is divided into two subfamilies, Monocteninae and
Diprioninae, and six genera (/099). Adults are described as follows: Antennae
composed of 13 or more segments, serrate in the female, and pectinate or bipecti-
nate in the male. Mesosterno-pleural sutures atrophied, mesoscutellum with ante-
rior margin V-shaped, posterior margin with an extremely narrow and cordlike
postergite (1/041). The larvae usually range in length from about 18 to 25 mm. The
body is usually whitish, yellowish, or grayish with brown or black stripes or rows of
more or less distinctly separated spots (1369).

Many species of diprionids are serious pests in both forest stands and plantations.
Outbreaks occur periodically, sometimes over extensive areas, resulting in loss of
growth and sometimes tree mortality (29, 3/7, 238).

383

The genus Monoctenus is represented in the United States and Canada by three
species, all of which occur in eastern North America on Juniperus spp., and Thuja
spp. None is considered a serious pest of forest or shade trees. M. suffusus
(Cresson) and M. fulvus (Norton) occur in southern Canada and in the Northeastern
States west to Kansas. Their hosts are redcedar, northern white-cedar, and juniper.
Full-grown larvae are dull green and about 18 mm long. The head is light brown:
the body is marked with three, dark longitudinal stripes: and the legs are black.
Adults appear in May, and larvae are active during June and July.

The genus Neodiprion contains more than 30 species in North America (697).
Many of the most destructive species of sawflies are in this genus. Outbreaks of
several species have occurred, resulting in serious damage to young pines on
thousands of hectares of pine plantations in the Eastern United States, and some-
times spreading over thousands of hectares of forested areas before subsiding.
Several authors have discussed the taxonomy of the genus (30, /042). A list of
world species and their distribution is available (/099). Life cycle information, in
addition to preserved specimens of reared adults and feeding larvae, is often needed
for positive identification of Neodiprion sawflies (697).

The redheaded pine sawfly, NV. /econtei (Fitch), occurs in southeastern Canada
and throughout the Eastern United States. Its preferred hosts are jack, red, short-
leaf. loblolly, slash, longleaf, pitch, and Swiss mountain pines. Eastern white pine,
larch, deodar cedar, and Norway spruce may also be defoliated, especially where
they are growing close to trees of preferred species. Full-grown larvae are about 20
to 30 mm long. The head is reddish and the body is yellowish white, with six rows
of black spots (fig. 179).

Pupation occurs in early spring and the adults appear in a few weeks. Eggs are
deposited in the tissues of current or previous year’s needles, a single female
depositing up to 150 eggs. The larvae feed gregariously on new and old needles and
also on the tender bark of young twigs. Sometimes they completely defoliate a tree,
progressing from the top downward, before they reach maturity. When this hap-
pens, larvae may abandon the tree and migrate for distances of several meters in

F-532021

Figure 179.—Larvae of the redheaded pine sawlfly,
Neodiprion lecontei, on loblolly pine.

384

search of new foliage. Full-grown larvae drop to the ground, enter the soil, and spin
tough, reddish-brown cocoons in which they become adults or spend the winter as
prepupae. In the South there may be three generations per year; in some Northern
States and Canada there is only one. Population development can be monitored by
use of a synthetic attractant (/296).

The redheaded pine sawfly 1s one of the most widespread and destructive of the
pine sawflies. It usually feeds on young trees, preferably those from 0.3 to 4.6 m
tall (769). Trees growing under stress on shallow soils, very wet or dry sites, or
subject to severe competition from hardwoods, bracken fern, or other vegetation are
especially susceptible to infestation, heavy defoliation, and damage (3/). Out-
breaks occur frequently throughout the range of this sawfly.

A nuclear polyhedrosis virus formulated for field use at the Canadian Forest Pest
Management Institute has proved to be effective in controlling the redheaded pine
sawfly (292, 1293). The native egg parasite, Closterocerus cinctipennis Ashmead,
and the larval parasites such as Spathimeigenia spp. are also effective in helping to
control this sawfly (92, 342).

A number of management practices have been suggested for preventing damage
in plantations by the redheaded pine sawfly: (1) Remove competing vegetation such
as hardwoods or dense bracken fern before planting sites to hard pines; (2) avoid
planting on high hazard sites covered with hardwoods or dense vegetation, in frost
pockets, or on soils that are excessively wet, dry, or very low in nutrients; and (3)
promote early closure of plantations by planting pines with spacing not greater than
1.8 by 1.8 m in the North Central States.

The Virginia pine sawfly, NV. pratti pratti (Dyar), has been recorded from New
Jersey and Maryland to North Carolina westward to Illinois. Its main hosts are
Virginia and shortleaf pines; other pines are sometimes infested. Newly hatched,
feeding larvae are pale green, with black head capsules, and are about 3 mm long.
Full-grown larvae (fig. 180) are spotted or marked with longitudinal black stripes
and are from 16 to 23 mm long.

F-519912

Figure 180.—Larvae of the Virginia pine
sawfly, Neodiprion pratti pratti.

385

Winter is spent in the egg stage, and hatching occurs in early spring. Newly
hatched larvae feed gregariously on the previous year’s needles, beginning about 10
to 15 mm below the tip. Feeding in groups of 2 to 15 larvae each, they consume the
outer portion of the needle, leaving the inner vascular tissue intact. After the first
two instars, the larvae consume entire needles except for the basal portion within
the sheath. They may also feed on portions of developing buds and on the tender
bark of twigs. By mid-May they become full grown and drop to the ground. Here
they spin cocoons in the surface litter or in the soil. Pupation occurs in September
and the adults appear in October and November. Eggs are deposited individually
within slits made by the female ovipositor along the flat sides of needles, usually
before the end of November. There is one generation per year (872, 1058).

Before the development of an outbreak in Maryland, Virginia, and North Car-
olina during the late 1950's, this species was not considered an important pest in the
Middle Atlantic States. Surprisingly, the outbreak occurring at that time swept over
more than 5.6 million hectares of pine and pine-hardwood type before it subsided.
Tree mortality was not serious although some scattered killing was reported, but
growth loss was severe.

White-footed mice and ants destroy large numbers of cocoons and prepupae in
heavy infestations. The ichneumonid, Exenterus nigrifrons Rohwer, is an effective
parasite of prepupae before they spin cocoons (/34). Dahlbominus fuscipennis
(Zetterstedt), an introduced parasite of sawfly cocoons, has been released and
established in infested stands in Virginia. A native polyhedrosis virus has destroyed
up to 70 percent of sawfly larvae when applied from an airplane (790).

Neodiprion pratti (Dyar) subsp., the sand pine sawfly, was detected in outbreaks
in west Florida for the first time during 1977. Both Ocala and Choctawhatchee
races of sand pine are hosts. Life stages are similar in appearance to those described
for the Virginia pine sawfly, but the supraspiracular markings consistently blend
into a black lateral stripe in mature, feeding larvae of the sand pine sawfly.

Winter is spent in the egg stage and hatching occurs during February and March,
after which larvae feed on previous year’s foliage, twig bark, buds, and strobill.
Cocoons are spun in soil or duff during March and April, and turn dark brown.
Adults emerge from cocoons during November and December and eggs are laid in
shoots fully exposed to sunlight. Females typically lay two to three yellow eggs
under the flat side and toward the apex of a needle. An average of about 30 needles
per shoot is infested. There is only one generation per year.

Plantations 10 years old and older have been most subject to defoliation,
especially along stand edges and within those stands of 750 or fewer trees per
hectare. Defoliation exceeding 90 percent per tree has resulted in subsequently

_reduced tree growth and some tree mortality has occurred during spring droughts in
association with infestations of cerambycid beetles. Known parasites include the
ichneumonids, Exenterus spp. and Endasys subclavatus (Say), and a bombyliid,
Villa sinuosa sinuosa (Wiedemann). Ice storms can cause heavy mortality in young
larvae.

The jack pine sawfly, NV. pratti banksianae Rohwer, occurs in Canada from New
Brunswick to Manitoba, and in the Lake States. Jack pine is the favored host, but
red and Scotch pines are occasionally attacked if they are growing with heavily
infested jack pine. Full-grown larvae are yellowish green and about 22 mm long.
The head is black and there are two longitudinal pale greenish-gray stripes running
down the back. There is also a single row of 11 black spots on each side.

386

Winter is spent in the egg stage and hatching occurs in May or June. The larvae
feed gregariously on previous year’s needles only. Feeding is completed by early
July. The larvae then drop to the ground and spin cocoons in the duff and top layers
of the soil. Adults appear in late August and early September and lay three to five
eggs in each needle, with the full complement of eggs on the needles of one twig.
There is one generation per year.

Older, open-grown trees are more subject to infestation than young trees or trees
in closed stands. Even-aged stands and plantations are especially attractive. Mor-
tality from a single defoliation seldom occurs, but it may result from several
successive defoliations. Generally, the most serious damage results from gradual
reduction in vigor and growth of the affected trees. Late spring frosts and a
polyhedrosis virus disease are often effective in the control of heavy infestations.
Direct and preventive methods of control are discussed (387).

Neodiprion pratti paradoxicus Ross has been recorded from Maryland to Nova
Scotia and Ontario. Its hosts are pitch, Scotch, shortleaf, and jack pines. Full-grown
larvae have black heads, and the body is marked with very pale lateral stripes.
Supraspiracular spots are usually partially or entirely lacking in the middle of the
body. Widespread outbreaks have occurred in pitch and shortleaf pine stands in
New Jersey.

Neodiprion taedae taedae Ross, the spotted loblolly pine sawfly, has been
recorded from eastern Virginia. Its favored host is loblolly pine, but it also
occasionally feeds on shortleaf pine. Full-grown larvae are greenish white and
about 22 mm long. The head is reddish brown, and there is a dull grayish-green
longitudinal stripe on each side of the dorsum. There is also a row of black spots
just above the spiracular line that extends from the second thoracic to the ninth
abdominal segment, and there are two black blotches on top of the tenth segment.

Winter is spent in the egg stage, and hatching occurs in late April or early May.
The larvae feed on old needles and reach maturity in about 30 days, fall to the
ground where they enter the soil a few centimeters, and spin cocoons. Adults appear
about mid-October and lay their eggs in rows on the sides of needles. There is one
generation per year.

Since the new foliage is not eaten, infested trees are not completely defoliated or
killed. Infestations occur more commonly in understocked pine stands where the
trees retain their lower branches. The life history of this species was discussed
under the name N. americanum (Leach) (563).

The loblolly pine sawfly, N. taedae linearis Ross, long recognized as a pest of
loblolly pine in Arkansas, is now known to occur also in Louisiana, southeast
Texas, Mississippi, South Carolina, Missouri, Ohio, and Illinois. As far as known,
loblolly and shortleaf pines are its only hosts; loblolly is preferred. Full-grown
larvae are dull green and about 25 mm long. There are heavy black stripes along
each side and often two lighter stripes below the heavier, black ones.

Winter is spent in the egg stage. Hatching occurs from early March to early May,
depending on location. Young larvae feed gregariously in groups, often encircling
the needles about half way from end to end, and partially girdling them. Infested
terminals soon take on a reddened appearance. Older larvae feed singly or in pairs
and consume the entire needle, leaving short stubs on the branch. They still retain
their gregarious habit, however, and move in a group from branch to branch (fig.
181). For the most part, only the older foliage is eaten, but on shortleaf pine the
terminal buds and tender bark on the new growth are also occasionally eaten (256).
Full-grown larvae drop to the ground and spin mahogany-colored cocoons in the

387

litter or topsoil. Pupation occurs in October or November and the adults emerge
soon thereafter. Eggs are laid in slits cut into the needles, usually 2 to 10 per needle.
Each female lays from 90 to 120 eggs, often all in the needles of one twig (/25/).

This sawfly is found chiefly on medium-size or large trees in forest stands.
Several outbreaks have been recorded. One, which lasted four seasons, spread over
an area of about |.2 million hectares in Arkansas before it subsided. Trees suffering
spring defoliation exceeding 75 percent per tree have shown an average net growth
loss of 51 percent | year following defoliation and 29 percent the second year (fig.
182). Important natural control factors are a polyhedrosis virus disease, cold, rainy
weather in the spring, and two larval parasites, a bombyliid, Villa sinuosa sinuosa
(Wiedemann), and the ichneumonid, Exenterus nigrifrons Rohwer (256, 1251).

Neodiprion warreni Ross is found in the Southeastern States from Arkansas into
north Florida. Known hosts include spruce and shortleaf pines. Mature feeding
larvae have a shiny black head and broad black subdorsal, lateral, and sublateral
longitudinal stripes. Spaces between stripes are reduced to narrow, pale lines.
Mature, feeding larvae have been collected during October in Arkansas and Flor-
ida.

The Swaine jack pine sawfly, NV. swainei Middleton, one of the most important
of the pine-infesting sawflies in eastern Canada, is known to have been present in
the Lake States since the early 1950’s. It is now widely distributed in the Upper
Peninsula of Michigan, and in north-central Minnesota and Wisconsin. Jack pine is
its favored host, but red, Scotch, and eastern white pines growing in close prox-
imity to heavily infested jack pines are also subject to defoliation by migrating
larvae (/333). Full-grown larvae differ in color in different portions of the infested
region. Those found in the Lake States have bright-orange heads and yellow bodies
and there usually are two longitudinal, pale stripes on each side. Bright-yellow
specimens without stripes are found occasionally.

F-486627
Figure 181.—Larvae of the loblolly pine sawfly,
Neodiprion taedae linearis.

388

F-486626

Figure 182.—Large pines heavily defoliated
by the loblolly pine sawfly, Neodiprion
taedae linearis.

In the Lake States, winter is spent as prepupae in cocoons in the litter or top-
soil, and pupation and adult emergence occur in early to late spring. Eggs are
laid in the current year’s needles, one to three eggs per needle, during June or
July. The larvae are gregarious and feed primarily on old foliage, usually on ex-
posed trees during July and August, until they become mature and drop to the
ground. There is one generation per year (80).

Numerous outbreaks have occurred at about 8-year intervals in jack pine stands
in Ontario and Quebec, with foci in poorer jack pine sites on outwash plains (S03).
Heavy tree mortality occurred in overmature stands and practically all surviving
trees over many hectares remained stag-headed for several years. A sawfly popula-
tion simulation model for such stands has been developed (804). Some of the most
severe infestations in the Lake States have occurred in jack pine plantations and
windbreaks. Aerial application of a nuclear polyhedrosis virus spray has provided
satisfactory control for at least | year in tests conducted in Quebec (/095).

The red pine sawfly, N. nanulus nanulus Schedl, occurs in the Northeastern
States west to Minnesota and in adjacent Canada. Its favored hosts are red and jack
pines, but it also feeds on eastern white, Japanese red, and Swiss mountain pines.
Full-grown larvae are dull grayish-green to black on top, greenish white beneath,
and about 18 mm long. The head is black and two olive-green stripes run down the
back to a darker area near the rear end. There are two longitudinal dark stripes on
each side, with the upper one a little darker than the lower one.

389

Winter is spent in the egg stage. Hatching occurs in early May and larvae are
present until July or August, depending on focation. They are gregarious and
usually consume all of the mature needles from one branch before migrating to
another. Prepupae drop to the ground and spin cocoons in the duff. Adults emerge in
early fall and deposit their eggs in slits in the round face of current season’s needles
near the tips of well-exposed branches. The female will usually oviposit her entire
complement of eggs on one twig. There is one generation per year (650).

Little or no mortality results from a single defoliation because only the older
needles are eaten, but where defoliation continues for several years a high percent-
age of the affected trees may be killed. Overmature trees are particularly susceptible
to damage.

The white pine sawfly, N. pinetum (Norton), occurs on eastern white pine
throughout the range of its host in eastern North America. Pitch, shortleaf, red, and
Swiss mountain pines are sometimes infested. Full-grown larvae are pale yellow
and about 25 mm long. The head is black and four rows of black spots extend from
the head to a black spot at the posterior end.

In the spring, the female deposits three to four eggs per needle, and the full
complement is placed in the needles of a single twig. The larvae feed gregariously
on both old and new needles, and when one branch is defoliated they migrate to
another. When they reach maturity they drop to the ground, spin cocoons in the soil,
and overwinter as a prepupa; pupation occurs in the spring. There is sometimes a
partial second generation per year, depending on location.

The white pine sawfly attacks trees of all sizes both in plantations and in forest
stands. Because the larvae feed on both old and new needles, they are capable of
completely defoliating a tree. Widespread outbreaks occur occasionally, whereas
local ones are reported almost every year. The egg parasite, Closterocerus
cinctipennis, 1S Sometimes an important natural control agent (/0/7/).

The balsam fir sawfly, N. abietis (Harris), occurs from coast to coast in
southern Canada and Northern United States. Its favored host is balsam fir but it
also attacks black and white spruces. Full-grown larvae are dark green and are from
18 to 25 mm long. The head is black and the body is marked with six longitudinal
dark stripes or bands on the back, the lowest of which may be broken up into small
dots or spots.

Winter is spent in the egg stage and hatching occurs in May or June. Newly
hatched larvae feed gregariously on old needles, eating only parts of the needle.
Feeding is completed in about 1 month. The mature larvae then spin tough, silvery
or light-brown cocoons on the needles or in the litter on the ground. Adults appear
from late July to early September and lay their eggs singly in slits cut in the edges of
the needles. There is one generation per year.

Balsam fir growing in open stands, in pastures, along lakeshores in Ontario, and
on islands off the coast of Maine is often heavily defoliated. Trees are seldom killed
by a single defoliation but some are killed by repeated defoliations. Spruce trees in
farm shelterbelts and ornamental plantings in the Prairie Provinces of Canada also
have been damaged. The introduced parasite, Dahlbominus fuscipennis (Zet-
terstedt), has destroyed up to 40 percent of cocoons in the duff in Maine. Cocoons
spun among the needles on the trees in the same stands escape parasitization.

Neodiprion abbotii (Leach) has been recorded in Ontario, Quebec, Wisconsin,
Virginia, North Carolina, South Carolina, Georgia, and Florida. Its hosts are listed
as loblolly, slash, longleaf, shortleaf, and red pines. The full-grown larva has a
brown to black head with a light spot on the frons, and the thorax and abdomen are

390

pale green. There are also four dark-green to black lateral stripes. N. abbotii larvae
feeding on slash and longleaf pines in south Florida have pale-amber head capsules
and pale, longitudinal stripes.

Winter is spent as a prepupa in a cocoon and pupation occurs in the spring.
Adults emerge as early as late March in the South. Eggs are laid in single rows of 15
to 20 each, mostly on previous year’s needles, with the eggs of a row almost
touching each other. Newly hatched larvae feed gregariously; older ones tend to
feed singly. Prepupae spin tough, silvery-brown cocoons on the needles of the tree.
There appear to be at least three generations per year in the South. First-generation
larvae are found in May, second-generation larvae during July and August, and
third-generation larvae from October to December (564).

The redheaded jack pine sawfly, N. rugifrons Middleton, occurs in southern
Canada and the Lake States where it feeds on jack pine. Mature feeding larvae are
15 to 20 mm long, have an orange-brown head, and two dark, subdorsal stripes run
the full length of the body to a black patch on the last abdominal segment. A row of
conspicuous black supraspiracular spots extends backward along each side from the
prothorax to the ninth abdominal segment. The underlying body color is white.

Winter is spent as prepupae in the soil. Adults emerge annually in May and June,
and oviposit in 1-year-old or older needles. Larvae of this early generation feed only
on older foliage and damage to the host tree is slight. In those years when two
generations develop, adults of the second generation emerge during July and
August, Oviposit principally in current-year needles, and larvae feed on foliage of
all ages. The chemical basis for this feeding behavior is the presence of relatively
high concentrations of two antifeedant organic acids in new foliage during the
growing season (624). When heavy infestations develop, jack pine may be almost
completely defoliated by September or October and planted pines from 0.9 to 6.1 m
tall may be killed. The tops of trees retaining 10 percent or less foliage often die the
following spring. Trees in isolated plantations and windbreaks are most subject to
severe damage (1295).

The brownheaded jack pine sawfly, NV. dubiosus Schedl, also occurs on jack
pine with N. rugifrons, and is somewhat similar in biology and appearance. N.
dubiosus differs principally in the underlying yellow color of larvae and in the first
annulus of the adult female saw, which has less than 10 teeth. A comparison of
these two species has been published (82).

Neodiprion virginianus Rohwer is found in Virginia pine from the Piedmont of
North Carolina north into southern Pennsylvania, west into eastern Kentucky, and
on sand pine in peninsular Florida. Eggs are laid in a row underlying the round side
of a needle, beginning near the point of needle divergence and ending near the
apex. Mature, feeding larvae are relatively short and stocky with a shiny black head
and a lateral-longitudinal row of distinct and evenly spaced subquadrate black
spots. There are at least three generations per year in Florida. A 62-hectare outbreak
occurred in the Tampa Bay area of Florida during the fall of 1978, resulting in
growth loss and some tree mortality.

The European pine sawfly, NV. sertifer (Geoffroy), an introduced species first
recorded in North America at Somerville, N.J., in 1925 (1060), is now widely
distributed. It has been recorded from New England to southwestern Ontario,
Michigan, Wisconsin, Ohio, Illinois, lowa, South Dakota, and south to Missouri. It
has many hosts, particularly Scotch, red, jack, Japanese red, Table Mountain, and
Swiss mountain pines. Eastern white, Austrian, ponderosa, shortleaf, and pitch
pines also are fed on to some extent when they occur in mixture with more favored
species.

391

Full-grown larvae are grayish green and from 18 to 25 mm long. The head,
thoracic legs, and upper part of the anal plate are black and there is a longitudinal
light stripe down the back. There are also two light-green stripes and one intense
green or black stripe on each side.

Winter is spent in the egg stage and hatching occurs from early April to mid-
May. The larvae feed gregariously on the previous year’s foliage and devour all of
these needles on one branch before moving to another one. New needles are never
eaten. Larvae may also feed on the bark of new shoots. This may result in shoot
deformation or death (/324). When the previous year’s foliage on a tree is com-
pletely eaten before the larvae reach maturity, the tree is vacated and the larvae
crawl to other trees to feed. Full-grown larvae either drop to the ground and spin
tough, light to dark golden-brown cocoons in the duff, or they spin them in
protected locations on the tree. Pupation occurs during late August or early
September and the adults appear from early September to late fall. Eggs are laid in
slits cut in the edges of current year’s needles in needle clusters near the end of a
branch. From 6 to 8 eggs are usually laid in a single needle, and about 10 to 12
needles in a single cluster are usually infested. There is one generation per year.

Trees defoliated by the European pine sawfly are seldom killed, since the new
foliage is never eaten and the tree is never entirely stripped of its foliage. Bark
feeding may cause some twig mortality but it 1s not serious. Losses in height and
diameter growth may be severe, especially following repeated defoliations. Damage
is most severe to Scotch pines grown in Christmas tree plantations.

During the late 1930's and early 1940's, colonies of the two European parasites,
Dahlbominus fuscipennis (Zetterstedt) and another believed to be Exenterus abrup-
torius (Thunberg), were received from Canada for release in eastern infestations of
this sawfly. D. fuscipennis became established in New Jersey and within a few years
became rather abundant locally (487). E. abruptorius also became established in
New Jersey. So far, neither species has given adequate control. The European
ichneumon, Lophyroplectus oblongopunctatus (= luteator) (Hartig) was released
in Canada in 1962 and transferred to Wisconsin in 1979. It is established in both
locations. A supply of a European polyhedrosis virus of the sawfly was also
received from Canada for use in its control (//2, //3). It has proved to be very
effective when applied as a spray either by aircraft or by knapsack or hydraulic
sprayers (fig. 183). The literature on the biology, ecology, and control of the
European pine sawfly has been reviewed (687, 76/).

as

F-519945

Figure 183.—Colony of larvae of the European pine
sawfly, Neodiprion sertifer. Larvae hanging head
downward were killed by a polyhedrosis virus.

392

The blackheaded pine sawfly, N. excitans Rohwer, occurs from Virginia to
Florida and west to Arkansas and Texas. It also occurs in Central America (1/297).
Loblolly and shortleaf pines are its preferred hosts in the United States. Slash,
longleaf, pond, spruce, sand, and Sonderegger pines are also attacked but to a much
lesser extent (//94). The full-grown larva is olive green and about 25 mm long (fig.
184). Its head is glossy black, there are two longitudinal black stripes on the
dorsum, a row of black spots on each side, and a large black spot on the last
abdominal segment.

F-514368

Figure 184.—Larva of the blackheaded
pine sawfly, Neodiprion excitans.

Winter is spent mostly in cocoons, but occasionally in the egg state or as older
larvae (1292). Oviposition begins in March when each female lays one egg per
needle in individual pockets sawed just above the fascicle sheaths of needies on a
given shoot. Newly hatched larvae are gregarious, with a circle of larvae feeding on
a single needle. Older larvae feed in steadily decreasing numbers per needle.
Previous year’s foliage is preferred during the growing season, but all foliage may
be consumed when needle growth ceases during the fall. When a branch or tree is
heavily defoliated, the larvae migrate in groups to other branches or trees. Full-
grown larvae spin golden-brown cocoons in ground litter or topsoil, but sometimes
remain on the tree and spin their cocoons on twigs, needles, or in bark crevices on
the lower trunk. There are three to four generations per year in the Gulf region (564,
SOR 94s):

Heavy infestations typically develop during the fall in moderate to dense stands
of sawtimber, especially when overmature trees are present. Overwintering trees
stripped of more than 90 percent of their foliage suffer growth loss and may be
attacked and killed by /ps engraver beetles, the black turpentine beetle, cerambycid

393

beetles, and ambrosia beetles. Several large outbreaks have occurred in Florida and
Texas; however, they subsided after one or two seasons. Starvation and reduced
reproductive capability of the females are among the factors that help bring
outbreaks to an end. The ichneumonid, Endasys subclavatus (Say), is an effective
parasite attacking cocoons spun in the litter and topsoil (342), while the pteromalid,
Dibrachys cavus (Walker), commonly attacks cocoons spun above ground.

The slash pine sawfly. NV. merkeli Ross, is known to occur in Georgia, Florida,
Mississippi, and on Grand Bahama Island. Both varieties of slash pine are hosts in
the United States, and Caribbean pine is infested in the Bahamas. Full-grown
female larvae are 25 to 30 mm long, with brown subdorsal and supraspiracular
stripes, and two widely separated, black semioval spots near the posterior end. The
head is 2.3 to 2.5 mm wide, reddish above the eyes and sooty black below. This
two-tone coloration is most evident in alcohol-preserved larvae. Eggs are laid in
growing, current year’s needles, with a row of 10 to 15 egg pockets typically
underlying the flat side of a needle, beginning at the fascicle sheath (/294).

Winter is spent as a prepupa in dark mahogany-brown cocoons usually spun in
the soil. Adult emergence occurs approximately during May, July, and September
in Florida. Larvae feed only on old foliage during the growing season, but may
completely defoliate trees during the fall when the largest populations typically
develop. Extensive fall outbreaks have occurred in 5- to 10-year-old open planta-
tions growing on periodically flooded flatwood sites in Florida and Mississippi,
subsequently resulting in greatly reduced tree growth when defoliation exceeds 90
percent per tree. Outbreaks in south Florida have been terminated within one
growing season by a nuclear polyhedrosis virus disease. Flies in the genus
Spathimeigenia are important parasites attacking mature, feeding larvae (342), and
Villa sinuosa sinuosa (Wiedemann) is an important cocoon parasite during
droughts. Chance flooding of cocoons immediately following adult sawfly
emergence resulted in complete mortality of parasitoids within cocoons, followed
by a large outbreak of this sawfly in south Florida.

A number of other species of Neodiprion also occur in eastern forests: N.
nigroscutum Middleton occurs on jack pine in the Lake States and Ontario. Mature,
feeding larvae have reddish-brown heads, unbroken subdorsal and supraspiracular
stripes, and arcuate markings above the legs. An epiproctal spot is not present (8/).
N. pinusrigidae (Norton) feeds on pitch pine from New Jersey to Maine. Full-
grown feeding larvae are dull green with a black head, solid black subdorsal stripe,
and broken black lateral stripe. Sometimes a double row of black dots occurs below
each lateral stripe. N. hetricki Ross has been observed feeding on loblolly pine in
Virginia and North Carolina and pond pine in South Carolina. Mature, feeding
larvae have black heads, black subdorsal stripes, and black lateral stripes that may
be either broken or solid. There is a single generation each year in Virginia, with
females depositing overwintering eggs during October (564). N. compar (Leach) is
reported from southeastern Canada, Virginia, Georgia, and Florida. Known hosts
include red and jack pines in Canada and longleaf pine in Florida. Mature feeding
larvae from Florida are pale green, with barely discernible longitudinal stripes. The
pale-amber head bears a yellow triangular spot within a large black pentagonal
marking. N. maurus Rohwer occurs on jack pine in southeastern Canada and the
Lake States.

The genus Diprion is represented by only one species in the United States and
Canada, and this species is of foreign origin.

394

The introduced pine sawfly, D. similis (Hartig), was first recorded in New
Haven, Conn., during 1914, and spread to western Wisconsin by 1944 (247). It was
first found in North Carolina during 1977 and by 1981 had been detected in two
distinct outbreaks covering 24,000 square kilometers in North Carolina, Tennessee,
and Virginia (339, 474). It is now known to occur from Maine to North Carolina, in
the Central and Lake States, and in southern Ontario and Quebec. Eastern white
pine is the favored host but Scotch, jack, red, and Swiss mountain pines are
commonly reported secondary hosts. Shortleaf and Virginia pines are also attacked
but are not considered to be threatened.

Full-grown feeding larvae (fig. 185) are about 20 to 25 mm long, with shiny
black heads. The body has a double black stripe bordered by yellow along the
dorsal midline, and the sides are dark with numerous rounded yellow and white
spots. Ventrally the larvae are pale yellow. Cocooned larvae have gray heads; the
body is pale gray-green and bears markings similar to those of feeding larvae, but
much lighter. The pupa is light green. Cocoons are cylindrical with rounded ends
and relatively thick, tough walls that vary from very light to dark brown in color.

Winter is spent as cocooned larvae. Pupation occurs in early spring and adults
emerge principally during May and early June. The females produce a potent sex
attractant (239). Eggs are laid in rows of about 10 per needle, each deposited in a
slit cut into the edge of the needle (fig. 186). Young larvae feed gregariously; older
larvae, singly. First-generation larvae prefer to feed on previous year’s needles,
while larvae of the second generation eat both old and new needles. Young larvae
eat only the outer, tender parts of needles; older larvae consume entire needles and
will feed on twig bark when most of a tree’s foliage has been consumed. Both first-
and second-generation cocoons are spun in crown foliage, at the base of small
branches, in bark crevices on the trunk, on host or nonhost trees, shrubs, grasses,
or other understory objects. Pupation occurs mainly during July, and second-
generation larvae feed through September. Small numbers of third-generation adults
(mostly males) are reported to emerge during some years in Wisconsin, but most

F-531260

Figure 186.—Introduced pine sawfly,
Diprion similis, ovipositing in
eastern white pine needles. Eggs
of this species are covered with a
pulpy mass.

Courtesy H. C. Coppel, Univ. Wis.

Figure 185.—Larvae of the
introduced pine sawfly, Diprion
similis.

395

second-generation sawflies overwinter as diapaused larvae within cocoons. Because
of its variable development, all stages of this sawfly may be present at the same time
during the summer (24/, 474, 846, 1325).

Defoliation is usually most severe in the upper half of tree crowns, but heavily
infested trees may be completely defoliated in one season. When this happens late
in the season after the winter buds have formed, many branches and occasionally
trees may be killed.

Some factors in natural control include very low or rapidly fluctuating tem-
peratures or heavy rainfall during the egg and early larval stages, bird predation of
cocooned larvae, and the establishment of three European parasitic wasps. The
ultimate-stage larval parasitoid, Exenterus amictorius (Panzer), and two cocoon
parasitoids, Dahlbominus fuscipennis (Zetterstedt) and Monodontomerus dentipes
(Dalman), are important control agents in Wisconsin (24/). Suppression of the
introduced pine sawfly to low population levels in North Carolina was achieved by
1982, primarily through mass-rearing and augmentative releases of M. dentipes
(474).

A comprehensive review of the European and North American literature on D.
similis is available (24/).

The genus Gilpinia is represented by two species in the United States and Canada
and both are of foreign origin.

Gilpinia frutetorum (F.) was discovered in North America in Massachusetts and
Rhode [sland in 1932, and now occurs in Quebec, from Maine south to New Jersey,
Pennsylvania, Wisconsin, and Minnesota. Its favored hosts are red and Scotch
pines, but it may also attack various other hard pines. Full-grown larvae are light
green with reddish-brown heads and are about 20 mm long. The body is marked
with six longitudinal dark-green stripes, two on the dorsum and two on each side
(fig. 187).

Winter is spent as a cocooned prepupa and pupation occurs in the spring. Adults
appear from late May to late July. Eggs are laid in slits cut in the needles. The larvae
feed singly, preferably on the older needles. Because of their greenish color they
blend into the background and are difficult to see. The first sign of infestation
usually is the presence of frass and green needle fragments on the ground beneath
the tree. Heavy infestations occur occasionally in pine plantations (/06/).

The European spruce sawfly, G. hercyniae (Hartig), was first recorded in North
America near Ottawa, Canada, in 1922. It has spread throughout the eastern spruce

F-519578
Figure 187.—Larvae of Gilpinia frutetorum.

396

forests from Newfoundland to Pennsylvania and west to Minnesota and Manitoba.
White spruce appears to be the favored host; however, red, black, and Norway
spruces also are attacked. Young larvae are yellowish green; older feeding larvae are
darker green, marked with five longitudinal white lines, and about 20 mm long
(fig. 188). The white lines are absent and the body is somewhat shortened in
cocooned larvae.

ee
—

nett.

Courtesy Can. For. Serv., Gt. Lakes For. Res. Cent.

Figure 188.—Larva of European spruce sawfly, Gilpinia
hercyniae.

In the Northeastern States there may be one or two generations per year or a
partial third, depending on location. Farther north there is only one. In the
Northeastern States, adults of the first generation appear from early May to June.
Males are rare and reproduction is parthenogenetic. Eggs are laid in slits cut in old
needles and hatching occurs within a few days. Larvae of all ages feed singly.
Young larvae feed by chewing out small pieces of the needle or by eating all but the
vascular bundle. Older ones consume the entire needle. Old needles are usually
preferred, but full-grown new needles may be eaten also. Second-generation adults
appear in early July or later and lay their eggs in needles. Hatching occurs shortly
thereafter, and the larvae feed for 3 or 4 weeks. At maturity, they drop to the ground
and spin cocoons in the litter in which they spend the winter. In the northern portion
of the species’ range, some remain in diapause for several years before pupating
(999).

The European spruce sawfly found conditions favorable for a rapid increase in
numbers once it gained a foothold in the spruce forests of North America. In 1932,
serious infestations were found over an area of about 5,200 square kilometers of the
Gaspé. Two years later evidence of defoliation was also apparent in Quebec, New
Brunswick, and adjacent areas in the United States. By 1935, the Gaspé outbreak
covered about 15,500 square kilometers, and tree mortality in some areas had
reached serious proportions. Collapse of the outbreak by 1942 was caused by an
accidentally introduced nuclear polyhedrosis virus disease (49). Since then popula-
tions have fluctuated at very low levels, and the spruce sawfly is no longer
considered to be an economic pest (9/3).

During the 1930’s and 1940’s, the Canadian government imported and liberated
approximately 20 species of parasites in infested areas (788). Substantial numbers
of these parasites were also released in the United States. Two species of cocoon
parasites and five species of larval parasites became established (328).

397

Family Tenthredinidae
Sawflies

The family Tenthredinidae is represented by more than 730 species in the United
States and Canada. The larvae are all leaf feeders, leafminers, gall formers, or fruit
borers. Many species are important pests of forest and shade trees and forest
plantations.

The antennae of the adults most commonly have nine segments in the Eastern
United States and range in shape from setaceous and filiform to clavate. The
mesothorax is without sterno-pleural sutures, the anterior of the scutellum is V-
shaped, and the posterior margin usually has a distinct posttergite. The tibiae are
without preapical spurs, and the apical spurs of the front tibiae usually have the
longer spur cleft at the apex. The larvae range in length from 10 to 37 mm and are
usually largest in diameter at the thorax. The body is greenish or variously colored,
sometimes with distinct markings, and is either smooth, glabrous, setiferous,
tuberculate, or spinous (/04/, 1369).

Heterarthrus nemoratus (Fallén), the birch leafmining sawfly, an introduced
species first recorded from Nova Scotia during 1908, is now widely distributed in
the Northeastern United States and southeastern Canada (/097). Its hosts are
various species of birch, with gray, paper, yellow, and European white being
preferred. The full-grown larva is somewhat flattened and whitish, with the head
and joints of the thoracic legs brownish, and is about 10 mm long.

In Maine, winter is spent in the prepupal stage and pupation occurs in late spring.
Female adults (no males have been found) appear during June and early July and
deposit their eggs singly in slits cut in the edges of mature leaves, apparently at all
levels in the tree. The larvae feed in the tissues between the upper and lower
surfaces of the leaf, producing large blisterlike or blotch mines free of frass (fig.
189). Each full-grown larva constructs a cocoon within its mine. The leaf then falls

F-519525

Figure 189.—Mines and cocoons of
Heterarthrus nemoratus, the birch
leafmining sawfly, in leaf of paper birch.

398

to the ground and the prepupa remains in its cocoon throughout the winter. There is
one generation per year (977).

Heavy infestations of the birch leafmining sawfly occurred in Maine during the
1920’s and 1930’s, and severe defoliation of birch occurred in many areas. Very
little tree mortality occurred, but there was a considerable loss in annual growth.
During this period several species of parasites were imported against the sawfly
(327), two of which, Chrysocharis laricinellae (Ratzeburg) and Phanomeris phy-
llotomae Muesebeck, became established.

The birch leafminer, Fenusa pusilla (Lepeletier), an introduced species first
recorded from North America in Connecticut in 1923, now occurs from New-
foundland to Maryland, west to Ontario, Minnesota, and Iowa; also in Washington
and Oregon (/097). Full-grown larvae are somewhat flattened, yellowish white in
appearance, and about 6 mm long. Black spots occur on the venter of the thorax and
the first abdominal segment.

Pupation occurs in the spring and the adults begin to appear about mid-May.
Eggs are deposited singly in slits cut in the central areas of young leaves, usually
near the tips of branches. The larvae feed on the tissues between the leaf surfaces.
At first they feed singly, forming small kidney-shaped mines near the egg. As the
individual mines increase in size, they coalesce and form large, hollowed-out
brown areas in the leaf. These areas (fig. 190) wrinkle and turn brown. Full-grown
larvae chew their way out of the leaf and drop to and enter the ground where they
form small earthen cells, 2.5 to 5 cm below the soil surface, in which they
overwinter as prepupae in cocoons. There are three or four generations per year in
the southern part of the range of this leafminer (450).

Outbreaks occur frequently in the Northeastern States and result in the browning
of birch stands over wide areas. Some trees may be killed, but the greatest damage
is the weakening of affected trees, which leads to attack by other insects.

Courtesy Conn. Agric. Exp. Stn.

Figure 190.—Mines of the birch leafminer, Fenusa
pusilla, in leaves of gray birch.

399

The elm leafminer, F. u/mi Sundevall, an introduced species, occurs in south-
eastern Canada and the Northeastern United States west to the Lake States. Its
preferred hosts are English, Scotch, and Camperdown elms. American elm is also
attacked occasionally. Full-grown larvae are about 6 mm long, flattened, and
whitish with a greenish cast. The head is brown and the legs are encircled with
brown.

Winter is spent as prepupae in brown papery cocoons in the topsoil. Pupation
occurs in the spring and the adults appear in May, usually during the first half of the
month. Eggs are laid in the upper surfaces of leaves. The larvae mine the tissue
between the leaf surfaces, causing large blotch or blisterlike mines. Several attacks
may occur on a single leaf. When this happens, the various mines may coalesce and
the entire leaf be hollowed out (fig. 191). These leaves soon wither and fall. Where
only a small portion of a leaf is mined, the surfaces dry out and crack, leaving holes
in the leaf. The larvae usually become mature in late June. Then they vacate their
mines and drop to and enter the ground to spin their cocoons. There is one
generation per year (/097). This species appears to be most injurious to small trees
in nurseries and ornamental plantings.

: A < ‘
ce, é
Courtesy Conn. Agric. Exp. Stn.

Figure 191.—Mines of the elm leafminer, Fenusa
ulmi, in elm leaves.

The European alder leafminer, F. dohrnii (Tischbein), an introduced species,
occurs in southern Canada and the Northern United States. Its hosts are listed as
alders, especially the introduced European alders. Winter is spent in the prepupal
stage in cocoons in the ground. Pupation occurs in the spring, and the adults appear
and lay eggs during late May and early June. Larvae feed in the tissues of the leaf
for about 3 weeks, forming blotch mines (/097). Up to 12 larvae may feed on a
single leaf. Full-grown larvae drop to the ground to pupate, and a second generation
of adults appears from late July to early September. They also lay eggs and give rise
to a second generation of larvae. These become full grown by late fall and then
enter the ground for the winter.

400

Profenusa thomsoni (Konow), the ambermarked birch leafminer, possibly an
introduced species from Europe, 1s widely distributed in Quebec, Maine, Vermont,
Connecticut, Ontario, Wisconsin, Illinois, and Manitoba. Its hosts are gray, paper,
and yellow birches. This leafminer is one of the four sawflies that feeds in leaves of
Betula spp. (1097). In Ontario, female adults were observed in late July and early
August. Eggs are laid in the tissues of leaves. The larvae mine the tissues, forming
light-colored blotch mines. Up to 40 larvae inhabit a single mine. When all of the
tissues of a leaf are consumed, all of the larvae, regardless of age, vacate the mine
and drop to the ground. Of these, only those in the latter part of the fifth instar are
able to enter the soil and survive. In light infestations, sucker growth up to about
1.2 m tall in shaded locations is preferred. In contrast, trees up to 10.7 m tall in all
types of habitats are attacked in heavily infested areas. Damage is not considered
serious because defoliation occurs late in the season (S2/).

Profenusa alumna (MacGillivray) has been observed mining the leaves of oak,
primarily red oak, from Maine to Virginia west to Wisconsin and [llinois. Full-
grown larvae are about 6 mm long and have prognathous, octagonal heads more
than twice as wide as long. Winter is spent as prepupae in cells in the duff. Adults
appear in the spring and the female lays her eggs on the upper surfaces of the
leaves. The larvae bore into the leaf and mine the tissues, causing blotching and
severe browning. Heavily infested leaves may drop by September, leaving bare
branches in the top of the tree. Although males have not been collected in Maine,
they have been taken in other areas of its range (/097).

Profenusa canadensis (Marlatt), the hawthorn leafmining sawfly, has been re-
corded mining the leaves of hawthorn and cultivated cherry. It 1s widely distributed
in eastern North America (/097). Serious infestations have occurred in Mas-
sachusetts and New York. P. /ucifex (Ross) attacks white oak in Maine, New York,
and Illinois, and bur oak in Ontario (737).

Messa populifoliella (Townsend), the poplar leafmining sawfly, mines the leaves
of poplar from New Brunswick southwestward to New Mexico and west to Califor-
nia, also in South Dakota and Manitoba (/097). In New Brunswick, adults are
present the latter part of May. Females oviposit generally near the leaf tip. Larvae
feed for 2 to 3 weeks, making blotch mines. They then drop to the soil to overwinter
and pupate (/2/7).

The pear sawfly, Caliroa cerasi (L.), an introduced species, occurs from coast
to coast in southern Canada and the Northern United States. Although best known
as a pest of cherry and pear in the United States, it also feeds occasionally on
hawthorn, plum, quince, mountain-ash, black cherry, and serviceberry. Full-grown
larvae are tadpole-shaped, sluglike, and about 12 mm long. The body is covered
with a shiny, olive-green material secreted by the larva.

Winter is spent in earthen cells lined with a substance secreted by the larvae.
Pupation occurs in June. Eggs are deposited singly in small semicircular slits cut in
the leaf tissue. The larvae feed mostly on the upper surface of the leaf, eating the
parenchyma only. Heavily infested trees appear as if scorched, and their leaves drop
prematurely. Full-grown larvae drop to the ground and form cells in the soil in
which pupation occurs. Reproduction is parthenogenetic (/097). Females appear in
2 or 3 weeks and lay eggs. Larvae of this generation are present in August and
September. When they become mature, they also drop to the ground. There are two
generations per year throughout most of the range of this sawfly.

The scarlet oak sawfly, Caliroa quercuscoccineae (Dyar), has been recorded
from scarlet, black, pin, and white oaks from Maine to North Carolina west to

401

Minnesota, Illinois, Missouri, and south to Louisiana. Full-grown larvae are shiny
green and about 12 mm long. Adults appear to be most numerous in October. Eggs
are laid in slits cut in the lower surface of the leaf. They are placed singly in rows
along the sides of the midribs and larger veins, and all hatch within a few days after
being laid. The larvae feed on the epidermis of the leaf, leaving it almost colorless
and transparent. Evidence of feeding is apparent during late summer. There may be
two generations per year, and the winter may be spent in the larval stage (74).

Caliroa petiolata Smith occurs on pin oak in New Jersey. Three other species of
Caliroa are known on oaks, one on tupelo, one on chestnut, and one on Prunus spp.

The genus Periclista is represented in eastern forests by 10 species that feed on
the leaves of oaks and hickories. The larvae are usually light green or have the
dorsum grayish, and are armed with rows of small, single spines that may be simple
or forked.

The larch sawfly, Pristiphora erichsonii (Hartig), was first recorded in North
America in 1880. It now occurs in all Canadian Provinces, in Alaska, and all of the
northern tier of States plus Maryland, North Carolina, and West Virginia. Its hosts
are listed as tamarack and Western, subalpine, European, Japanese, Dahurian, and
Siberian larches. In eastern America, tamarack is most seriously infested, but
planted exotic larches are coming under increasingly serious attack. Full-grown
larvae are whitish beneath and gray-green along the dorsum, have jet-black heads,
and are about 16 mm long. Female adults are black and from 6 to 9 mm long. The
abdomen has a broad orange or red band, tapers sharply posteriorly, and is keeled
longitudinally along the midventral line.

In the Lake States, winter is spent in the prepupal stage in the ground. Pupation
occurs in the spring and the adults appear from mid-May to August, depending on
temperature and location. Eggs are laid in rows under the bark of currently
elongating shoots on the branches, and hatching occurs in about 8 days. The
emerged larvae move to and feed in groups upon tufts of needles, which are found
on short shoots on the older twigs. Feeding is completed in about 20 days and
mature larvae drop to the ground, enter the duff, and spin papery, brown cocoons.
Because of the long period of adult emergence, feeding larvae, cocooned larvae,
pupae, adults, and eggs all may be found at the same time in early summer (fig.
192). There is one generation per year, but occasionally small numbers of second-
generation adults are produced too late for any larvae to complete feeding. A small
number of larvae enter diapause and require 2 years to complete one generation
(332, 496).

Because larvae have the feeding behavior of rejecting single needles found on
elongating shoots in favor of needle tufts found on short shoots, 100 percent
defoliation seldom occurs. The chemical basis for this differential feeding appears
to be the presence of relatively high concentrations of five deterrent chemicals in
single needles from mid-July into August (945). Heavily defoliated trees commonly
refoliate after a few weeks; however, repeated defoliations can result in trees with
thinned foliage, reduced radial and terminal growth, reduced production of normal
shoots with a tendency toward adventitious growth, and branch mortality. Marked
loss of radial increment occurs after 4 to 6 years of outbreak, and tree mortality
occurs after 6 to 9 years of moderate to heavy defoliation. Many widespread
outbreaks have been recorded since 1880 and losses have been severe. Since the
middle of the 1950's, an estimated volume loss of 40 percent in valuable sawtimber
and pulp stands has occurred in managed forests in Minnesota.

402

A, F-485946; B, F-485947;
C, F-531261; D, F-531899

Figure 192.—The larch sawfly, Pristiphora erichsonii. A,
Ovipositing female; B, eggs in new shoot; C, mature
larvae; D, cocoons.

Overwintering cocooned sawfly larvae are destroyed in enormous numbers by
small mammals, especially shrews and voles. High surface water in tamarack bogs
also destroys large numbers of cocoons at times. A moderate number of species of
parasites occur but only two, the imported ichneumonid Mesoleius tenthredinis
Morley and the tachinid Bessa harveyi (Townsend), have been common during the
current outbreak in central Canada and Minnesota, which began about 1938.
Initially, M. tenthredinis was quite effective in control; however, the sawfly Gevel-
oped an immunity to it in central Canada, Minnesota, and Wisconsin (335).

In 1961, the ichneumonid Olesicampe benefactor Hinz was introduced from
Europe to Canada; this has added a very promising parasite to the biota for control
of the larch sawfly. This parasite was first established in the Prairie Provinces,
transferred eastward in Canada, and then to several locations in the Central and
Eastern United States (341, 704, 1216).

403

The mountain-ash sawfly, P. geniculata (Hartig), a probable introduction from
Europe, was first recorded in North America at Haines Falls, N.Y., in 1926 (J059).
It is now recorded from the Northeast to Minnesota, southeast to West Virginia, and
Newfoundland to Ontario. Its principal hosts are American and European moun-
tain-ashes. It also occurs occasionally on showy mountain-ash and the hybrid,
Sorbaronia. Full-grown larvae are pale greenish to yellow with yellow heads and
yellow thoracic legs. All body segments except the last are marked with black spots
of uneven size and shape (fig. 193). The spots occur in irregular rows, four along
each side of the body and two broken ones down the dorsum.

In eastern Canada, winter is spent in the prepupal stage. Pupation occurs in the
spring and the adults appear from late May to early July. Eggs are deposited in slits
cut around the edges of leaflets and hatching occurs in about 1 week. Newly
hatched larvae straddle the edge of the leaf and feed around the periphery. When
disturbed they raise their abdomens in the form of an S. Larvae in the first two
Stages are gregarious. Fourth and fifth instars often feed singly. They usually
consume all of a leaflet except the midrib. When they have consumed one leaflet
they move to another. Feeding is completed in about 2 or 3 weeks and the larvae
drop to the ground. Here they spin cocoons in the duff and topsoil. About 20
percent of these pupate and appear as second-generation adults in July (432).

The mountain-ash sawfly is primarily a pest of shade and ornamental trees. The
esthetic value of these trees is seriously reduced by defoliation, but they usually
survive even when completely defoliated.

Other species of Pristiphora occurring in eastern forests include P. chlorea
(Norton) on oak, P. siskivouensis Marlatt on birch, and P. sycophanta Walsh on
willow.

The yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer), is wide-
spread, occurring from Newfoundland to Alaska, Massachusetts, Michigan, Min-
nesota, Wyoming, Idaho and British Columbia. Its hosts are white, black, red,
blue, Norway, and Engelmann spruces (/320). Full-grown larvae (fig. 194) are
olive green above and lighter green below and are about 18 mm long. The head is
chestnut brown or reddish yellow and is sometimes mottled with various shades of
brown. Each side of the body bears a gray-green longitudinal stripe near the midline
of the dorsum, a broad one beneath this, and a darker one farther down. There is
also a dark line or spot just above the base of each leg.

Winter is spent as prepupal larvae in tough, dark-brown cocoons. Pupation
occurs in the spring, and the adults appear from late May to mid-June. Eggs are
deposited in slits cut in current season’s needles, usually at the base and usually
only one per needle. Sometimes they are also found in the tender bark of the stem
between needles. There are five feeding instars in males and five or six in females
(1228). Larvae prefer to feed on new needles, but older larvae will eat old needles
when new foliage is lacking. No significant differences in the susceptibility of
white spruce from different seed sources has been found (237). The larvae become
full grown in early to late July and drop to the ground. Here they spin cocoons in the
duff or top 2 cm of soil. There is one generation per year (906).

The yellowheaded spruce sawfly is a serious defoliator of young spruce planta-
tions and natural reproduction growing in open areas; ornamental, shelterbelt, and
nursery spruces are also infested. Young plantation trees are usually not attacked
until the 3rd to Sth year after planting; trees older than 5 years may be killed after 3
or 4 consecutive years of moderate to heavy defoliation. Small mammals and other
predators as well as numerous insect parasites are important causes of mortality in

404

Ss

Courtesy Can. For. Serv. Courtesy Can. For. Serv.,

Figure 193.—Larvae of the Gt. Lakes For. Res. Cent.
mountain-ash sawfly, Figure 194.—Larvae of the
Pristiphora geniculata. yellowheaded spruce sawfly,

Pikonema alaskensis.

late-stage feeding larvae and cocooned populations (607, /202). An attractant
involving female and synergistic host components has recently been identified (60,
61).

The greenheaded spruce sawfly, P. dimmockii (Cresson), is recorded from
Newfoundland to Northwest Territories, New Hampshire, Colorado, Wyoming,
and Idaho, and feeds on various species of spruce. Populations are usually low and
apparently cause little injury.

The willow sawfly , Nematus ventralis Say, occurs in southeastern Canada, south
to Georgia, and west to Utah, Colorado, and Montana. Its favored host is willow,
but it also feeds on poplar. Full-grown larvae are black or greenish-black, with large
light-yellow spots on the sides of the body, and are about 18 mm long.

Winter is spent in the prepupal stage in cocoons in the litter or topsoil beneath the
trees. Pupation and adult emergence occur in the spring. Eggs are deposited in
pockets cut in the tissues of the leaves. Young larvae feed in colonies, eating small
holes in the leaves. Later, entire leaves are eaten. There may be one to several
generations per year, depending on location. Some individuals may remain in
diapause up to 20 months before completing their development, especially in
northern areas.

The willow sawfly occasionally heavily defoliates willows in ornamental plant-
ings and along streams. Basket willows in the South have been damaged severely.

Nematus salicisodoratus Dyar occurs from Maine to Virginia, west to Wiscon-
sin, Illinois, and Arkansas. It also feeds on willow and poplar. Full-grown larvae are
light green and about 15 mm long. The head is black and there are three longitudi-
nal rows of closely spaced black spots on the dorsum. There are also two rows of
black spots on each side; those in the lower row are larger. Winter is spent in the
prepupal stage in cocoons on the ground. Pupation occurs in the spring, and adults

405

of the first generation appear in late May or June. Larvae of this generation are
found in June and July. Adults of a second generation appear from late July to early
September. This species is often abundant locally in the Northeastern States.

Approximately 40 additional species in the genus Nematus have been recorded
from eastern forests. Known hosts of a few species are as follows: black locust—N.
abbotii (Kirby); black locust and honey locust—N. tibialis Newman, paper birch—
N. viridescens Cameron and N. pinguidorsum Dyar; willow and possibly poplar—
N. fulvicrus Provancher; poplar—N. hudsoniimagnus Dyar; hophornbeam-——N.
ostryae (Marlatt); American hornbeam—N. carpini (Marlatt); alder—JN.
erythrogaster Norton; poplar and willow—N. limbatus Cresson; and willow—N.
oligospilus Foerster.

Trichiocampus viminalis (Fallén), the poplar sawfly, an introduced species,
occurs across southern Canada and the Northern United States. Its hosts are various
species of poplar and sometimes willow. Full-grown larvae are orange-yellow,
sparsely clothed with yellow hairs, and about 18 mm long. Each side of the body is
marked with two rows of more or less rounded black spots. Those of the lowest row
are small (fig. 195).

x

Courtesy Conn. Agric. Exp. Stn.

Figure 195.—Larvae of the poplar sawfly, 7richiocampus
viminalis, on leaves of poplar.

Winter is spent in the prepupal stage in cocoons in the duff beneath the trees.
Pupation occurs in the spring and the adults appear in May. Young larvae feed side
by side on the leaves.

Older larvae tend to scatter and eat all but the midribs and larger veins. They
become full grown in June or early July. Some pupate and give rise to a second
generation of adults in August; the remainder do not complete their development
until the following spring. Carolina and Lombardy poplars grown for ornamental
purposes may be seriously defoliated.

The dusky birch sawfly, Croesus latitarsus Norton, is the most commonly
encountered species in the genus, and occurs in eastern Canada and south through
the Eastern States to Florida and west to Alaska, British Columbia, and Utah
(1098). Its hosts are various species of birch, preferably gray birch. Full-grown
larvae are yellowish green with shades of black and are about 24 mm long. There is
a row of more or less distinct black blotches on each side and a series of black spots
in the subspiracular area.

406

Winter is spent in the prepupal stage in a cocoon in the topsoil. Adults appear
during May and June, or earlier in the Deep South. There may be several overlap-
ping generations per year and larvae may be found from spring until fall. The larvae
are gregarious and feed along the edges of the leaf (fig. 196). Colonies are often
found defoliating small saplings, but this usually does not occur over wide areas.

Other species of Croesus include C. curvarius Smith on hazel, C. castaneae
Rohwer on chestnut, and C. varus (Villaret) on alder, the latter sawfly of foreign
origin.

2
Courtesy Conn. Agric. Exp. Stn.

Figure 196.—Larvae of the dusky birch sawfly, Croesus
latitarsus.

The striped alder sawfly, Hemichroa crocea (Geoffroy), probably an introduced
species, occurs from coast to coast in southern Canada and the Northern United
States south to New Mexico. Its hosts include various species of alder and occasion-
ally birch, hornbeam, and willow (//0/). Full-grown larvae are yellowish and about
20 mm long. The head is shiny black, and there is a dark-brown subdorsal stripe on
each side running from the second thoracic to the tenth abdominal segment. Two
broken subspiracular stripes composed of blotches and dashes extend along each
side to the ninth segment.

Winter is spent in the prepupal stage in cocoons just beneath the surface of the
soil. The cocoon is very thin-walled and is formed within a cell made by cementing
together particles of soil. Adults appear during late May and deposit their eggs in
slits cut in the sides of the midrib on the undersurface of leaves. The larvae are
gregarious and usually eat all but the midrib and larger veins of the leaf. They
become full grown in July, and adults of a second generation appear during late July
and August. Larvae of this generation are found during August and September.
There are two generations per year. This species occasionally severely defoliates
alder in the Lake States, Northeastern States, and Canada.

The maple petiole borer, Caulocampus acericaulis (MacGillivray), is known to
occur in Ontario, Connecticut, New York, Maryland, Michigan, Illinois, and
Alabama. The larvae feed by boring into and tunneling the petioles of maple leaves.
Full-grown larvae are light yellow with light-brown heads and are about 8 mm long.

Winter is spent in the prepupal stage in a cell 5.0 to 7.5 cm below the surface of
the soil. Adults appear early in May and deposit their eggs near the bases of the

407

petioles of maple leaves. The larvae tunnel in the petioles, usually breaking them
near the leaf blade, and the leaves fall (fig. 197). After the petioles break, the larvae
continue to feed for about | week to 10 days in those portions of the petioles
remaining on the tree. When these also break and fall, the larvae vacate them and
enter the soil to pupate. There is one generation per year.

Courtesy Conn. Agric. Exp. Stn.

Figure 197.—Petiole of maple leaf severed by larvae of
the maple petiole borer, Caulocampus acericaulis.

Damage by the maple petiole borer is slight. Nevertheless, heavy infestations on
valuable shade trees may be undesirable because of the presence of large numbers of
dead leaves during the summer.

The brownheaded ash sawfly, Tomostethus multicinctus (Rohwer), occurs in
southern Canada and throughout the Eastern United States west to the Great Plains,
and also in Oregon and California. Its hosis are red and white ash trees. Full-grown
larvae are greenish or yellowish white and from 14 to 20 mm long.

Winter is spent in the prepupal stage in cocoonlike cells in the topsoil. Adults
appear as early as April in the southern portions of its range and lay their eggs in
slits cut along the outer margin of young leaflets, several eggs per leaflet. Young
larvae chew holes in the leaflets, and the older ones consume them entirely. Larvae
become full grown and move to and enter the ground by late May in the South.
Farther north, they become full grown at progressively later dates. Pupation occurs
in the spring and there is only one generation per year. The larvae are such
voracious feeders that a heavily infested tree may be completely defoliated in |
week. Shade trees are especially subject to serious defoliation (7/3).

The blackheaded ash sawfly, Tethida cordigera (Palisot de Beauvois), occurs
throughout much the same area as the brownheaded ash sawfly, except Oregon and
California. The habits, food plants, and life histories of the two species are also
very similar. Full-grown larvae are whitish with a yellowish tinge and are about 18
mm long. The head is shiny black, and the thoracic legs are eae brown. This
species 1s occasionally a pest of shade trees.

408

Eriocampa juglandis (Fitch), the butternut woollyworm, occurs from New
Brunswick to North Carolina west to Ontario, Minnesota, and Nebraska. Its hosts
are butternut, black walnut, and hickory. Full-grown larvae are green, with indis-
tinct black spots on the sides, and are about 18 mm long. The body is covered with
flocculent white tufts that rub off when touched, and the head is white with black
eyes. Larvae feed gregariously, often causing considerable defoliation locally.
When they become full grown they move to and enter the ground, where they form
cocoons made of soil cemented together. Winter 1s spent in the cocoons and there is
one generation per year.

The genus Hoplocampa is represented in eastern America by a number of
species, all of which presumably feed as larvae in the fruit of their hosts. Eggs are
laid in the calyx of flowers and the larvae bore into and hollow out the developing
fruit. Some of the species and their known hosts are: H. oskina Ross—hawthorn
from Maine, New York west to Michigan, lowa and Kansas; H. halcyon (Norton) —
serviceberry from Maine to North Carolina west to Alberta and Illinois; H. pallipes
MacGillivray—serviceberry from New Hampshire, New York to Michigan, Al-
berta, British Columbia, Oregon, and California; H. lacteipennis Rohwer—
~chokecherry in southeastern Canada and from Maine to New York, west to Man-
itoba, Alberta, Montana, and Colorado; and H. montanicola Rohwer—choke-
cherry in the Northern United States and southern Canada.

The genus Anoplonyx contains four species that occur in the Northern United
States and Canada. The larvae of all species feed on various species of larch. Full-
grown larvae of the two eastern species, A. canadensis Harrington, the onelined
larch sawfly, and A. luteipes (Cresson), the threelined larch sawfly, are green and
range in length from about 9 to 15 mm. The thorax is larger than the abdomen,
producing a humpbacked appearance and causing the body to taper posteriorly
(144).

The genus Euura is represented in the Eastern United States and Canada by 10
species, and all are gall-makers on various species of willow. Larvae are usually
yellowish or greenish white with black eyes, and the head 1s often tinted brown. All
species apparently have one generation per year. Winter is spent as prepupae either
in cocoons in the ground or in galls on the host. Adults appear in the spring and lay
their eggs in the shoots. Larvae feed on the tissues and become enveloped in galls
on stems, twigs, petioles, or buds.

Pontania spp. form leaf galls on willow, while Phyllocolpa spp. live in rolled
leaves or rolled leaf edges.

The genus Macremphytus is represented in eastern forests by three species, all of
which feed as larvae on dogwood. Full-grown larvae of some species are creamy
yellow on top with grayish-black crossbands or spots, and the legs and venter are
yellowish. The head is shiny black, and the body is covered with a white powdery
secretion.

Superfamily Siricoidea—Family Siricidae
Horntails

Members of the family Siricidae are commonly known as horntails because of the
presence of a hornlike projection on the last abdominal segment of the adult. This
process is short in the male; in the female it is much longer and often spear-shaped.
The ovipositor of the female is long and fitted for boring. In this respect it differs
from the ovipositor of sawflies that consists of sawlike plates. Horntails attack both
hardwoods and coniferous trees. A few species have been recorded infesting
vigorous trees, but they usually prefer trees or parts of trees that are dead or in a

409

badly weakened condition. Horntail larvae are parasitized by several members of
the ichneumonid genera Rhyssa and Megarhyssa. Members of the latter genus are
of special interest because of their striking appearance. Using their extremely long
Ovipositors, the females bore deep holes into the wood and deposit their eggs on or
near horntail larvae in their galleries. The ovipositor often becomes caught in the
wood and the female, unable to escape, dies. M. atrata (F.), M. macrurus macrurus
(L.), and M. greenei Viereck are parasites of Tremex columba (L.), but not of any
other horntails.

Horntail adults are medium to large in size and are usually metallic blue or black.
Some are varicolored with combinations of black, red, and yellow. The head,
thorax, and abdomen are of equal width; the wings are well developed; the antennae
are long and filiform, with about 15 segments; and the anterior tibia is armed with a
single apical spur, cleft at the apex.

Damage caused by members of the family can be prevented or reduced by
promptly using infested logs or submersing them in water, and by kiln-drying green
lumber sawed from infested logs.

The genus Sirex is represented by four eastern species. S. juvencus (L.) occurs in
eastern Canada and the Northeastern and Midwestern States. Its hosts are listed as
pine, fir, and spruce. Canadian studies indicate that S. juvencus larvae do not
develop in the absence of the fungus, Stereum chailletii (Pers.) Fr. (1159). In New
Brunswick, the males precede the earliest females by 2 to 4 weeks, before each of
the sexes continues emerging sporadically throughout the summer. Peak emergence
continues for a month from mid-August. Oviposition begins within a day of
emergence; a 2- to 10-mm puncture into the sapwood is completed in about 8 to 10
minutes. One to three eggs are deposited at intervals along the oviposition site.
Eggs or early-stage larvae overwinter. Pupation occurs after 5 to 11 molts, and the
pupal stage lasts 4 to 6 weeks. It takes 2 to 3 years for S. juvencus to complete its
life cycle (1/59). The blue horntail, S. cyaneus F., attacks spruce, larch, and pine
in southern Canada and the northern tier of Eastern States. S$. edwardsii Brulle has
been taken from pitch pine in the Atlantic Coast States and is recorded from Quebec
to Georgia west to Saskatchewan, Wisconsin, and Arkansas. S. nigricornis F. has
been recorded from shortleaf, eastern white, and Virginia pines and 1s distributed
from Quebec to Florida west to Saskatchewan, Wisconsin, Arkansas, and Loui-
siana.

Sirex areolatus (Cresson) has been introduced from western North America into
the Southeastern United States, but may not have become established.

The genus Urocerus Fourcroy is represented by four eastern species. U. albicor-
nis (F.), the whitehorned horntail, attacks many species of conifers and occurs
throughout boreal America. Adults are blue-black or black and about 25 to 30 mm
long. The middle of the antennae, cheeks, bases of the tibiae, and tarsi are white.
White spots also occur at times on the sides of the abdomen. This species has also
been observed attacking freshly sawed lumber. U. gigas flavicornis (F.), the yellow-
horned horntail, attacks spruce and other conifers in New England and Canada.
Adults are black and from 20 to 37 mm long. Females have the first, sixth, and part
of the seventh abdominal segments yellow; males have the second through fifth
segments orange-yellow. The fungus, Stereum chailletii, is also associated with this
siricid (1159). U. cressoni Norton, the black and red horntail, occurs from
eastern Canada to Florida, west to Wisconsin and Minnesota, and breeds in fir,
spruce, and pine. U. taxodii (Ashmead) infests baldcypress in Florida.

The pigeon tremex, 7remex columba (L.), 1s the most common of the horntails.

410

It occurs throughout eastern North America and breeds in a wide variety of dead or
weakened deciduous trees such as beech, maple, birch, elm, hickory, oak, and
sycamore. The adult female is 37 to 50 mm long. The head, antennae, and thorax
are reddish and black; the abdomen is black with ocher-yellow bands and spots
along the sides; and the wings are smoky brown with a wingspread of 50 mm or
more. Males are reddish, with some black, and are about 18 to 37 mm long. Full-
grown larvae are whitish, cylindrical, and about 50 mm long. The abdomen ends in
a short, strongly sclerotized and compressed process armed with two pairs of small
teeth.

The female bores through the bark into the wood to a depth of about 12 mm to
deposit her eggs. Although laid singly, several eggs may be found near each other
in a limited area. The fungus, Daedalea unicolor Bull. ex Fr., 1s essential for larval
development (//60). The larvae feed by excavating tunnels entirely in the wood.
This frequently weakens the tree and leads to windbreakage. Pupation occurs at the
end of the larval tunnel and the adults emerge through circular holes about 8 mm in
diameter. There appears to be one generation per year, although in New Brunswick,
it has a minimum of a 2-year life cycle (1/60).

Eriotremex formosanus (Matsumura), the Formosan horntail, a species pro-
bably introduced from Taiwan or Indochina, has been collected from dead water
oak and hardwood logs in Alabama and has been reported from Florida and
Georgia. Female adults are black with yellow hairs on the head and body, yellow
pronotum, yellow bands and spots on the abdomen, and yellowish to reddish-
brown wings with the area around the stigmata of the forewings and apical margin
of each wing infuscated. E. formosanus probably prefers dead, dying, or weaken-
ed hardwood tress and may be similar in habits to Tremex columba (1100).
Family Xiphydriidae
Xiphydriids

The family Xiphydriidae is represented in the United States and Canada by eight
species, all of which occur in the Eastern United States. One species occurs west to
the Northwest Territories, British Columbia, and Oregon. The adults are somewhat
similar to those of the family Siricidae, but are only about 12 to 18 mm long, and
the female ovipositor sheath is seldom longer than the last tergite. Adults are
reddish, black and yellowish, or entirely black. Full-grown larvae are about 18 mm
long and the abdomen ends with a brown concave prong ornamented with teeth on
the underside. As a rule, the larvae feed in moderately sound to partly decayed
wood of deciduous trees, usually in small branches. Adults sometimes emerge from
firewood brought inside, but they are no threat to people or property (//02). Few, if
any, are ever very abundant.

Eastern species and their known hosts and distribution are as follows: Xiphydria
abdominalis Say—basswood from southern Canada to North Carolina, Wisconsin,
and lowa; X. maculata Say—maple in Canada and most of the Eastern United
States except the Gulf States; X. tibialis Say—elm, birch, beech, American
hornbeam, oak, and hawthorn in southeastern Canada from Nova Scotia, Quebec,
south to Florida and west to Wisconsin, Illinois, and Kansas; X. hicoriae Rohwer—
hickory and elm in southeastern Canada and from Massachusetts and New Jersey to
Illinois; X. mellipes Harris—birch from Nova Scotia to North Carolina west to the

Northwest Territories, British Columbia, and Oregon.
Family Orussidae
Orussids

Five species of the family Orussidae have been recorded from the United States
and Canada, three of which occur in the Eastern United States. The adults are

411

somewhat similar to those of the Siricidae, but are much smaller, ranging from only
about 8 to 14 mm in length. As far as known, the larvae are all parasitic on
woodborers.

Superfamily Cephoidea—Family Cephidae

Stem Sawflies

The larvae of all members of this family and superfamily are borers in the stems
of plants such as grasses or berries, or in the tender shoots of trees and shrubs. Only
12 species are recorded from the United States and Canada. Adults are slender-
bodied and seldom more than 18 mm long. The body is black or dark colored,
occasionally marked with narrow yellow bands. The antennae are filiform, with 20
to 30 segments, and are either spindle-shaped or club-shaped. The front tibia has a
single apical spur, cleft at the apex.

The willow shoot sawfly, Janus abbreviatus (Say), occurs in southern Canada
west to Manitoba and South Dakota, and over much of Eastern United States
(1136). Its hosts are willow and poplar. Full-grown larvae are white, cylindrical,
and about 12 mm long. The thoracic legs are indistinctly jointed and fleshy. There
is a single pair of small prolegs on the last abdominal segment and a short, tubular
prong on the tip of the abdomen. Adults appear in late May and June and the
females deposit their eggs in punctures in the shoots of their hosts. Sometimes they
girdle and weaken the shoots above the oviposition site. The larvae feed by boring
down through the pith, which kills the shoot for varying distances. Winter is spent
within the shoot in cocoonlike structures. There is one generation in the North, and
three generations in Mississippi (//36).

Suborder Apocrita

Members of the suborder Apocrita have the base of the abdomen constricted into
a slender petiole or “‘waist.”’ The constricted portion is the first abdominal seg-
ment, which is fused to the thorax. Thus, what appears to be the first segment of the
abdomen is actually the second. The adult female is equipped with a piercing
ovipositor. In some species it is used as a tool for boring deep holes into the wood in
which eggs are deposited: other species use it for thrusting eggs into the bodies of
other insects; in still others, it is connected to poison glands and is used as a Sting.
The larvae are usually grublike or maggotlike. Some feed as parasites or predators
on other insects and some feed on plants. Adults feed chiefly on flowers, sap, or
other plant materials: some parasitic species feed occasionally on body fluids of the
host.

Considered as a whole, members of this suborder are far more beneficial than
harmful. Only a few species are harmful to trees or wood products. A number of
species are injurious to tree seed and cone crops. Some species of ants are
destructive to young trees in nurseries, plantations, and natural regeneration areas,
whereas others are destructive to finished wood products. At present, there are
approximately 15 times more species in this suborder than in the Symphyta.
Superfamily Ichneumonoidea

This superfamily constitutes one of the largest groups of parasitic insects (more
than 5,000 species) and from the point of view of effectiveness in holding in check
the numerous pests that infest plants, it probably ranks first (2/6). It comprises five
families (Stephanidae, Aphidiidae, Braconidae, Hybrizontidae, and Ich-
neumonidae), containing more than 30 subfamilies, hundreds of genera, and thou-
sands of species (697). Only a small portion of the important species parasitic on

412

forest insects is mentioned. A complete list of the species known to occur in the

United States and Canada ts available (697).
Family Stephanidae
Stephanids

This small family of rather rare insects, parasitic on coleopterous and hymenop-
terous wood borers, is represented in eastern forests by only one genus and two
species. Adults are usually collected on dead trees harboring their wood-boring
hosts. The adult is odd looking, having a crown of teeth on its spherical head which
is situated at the end of a long neck.

Family Aphidiidae
Aphidiids

Until recently this family was considered a subfamily of the Braconidae. It
consists of 3 subfamilies, some 16 genera, and 114 species. Most species are
solitary endoparasites of both larval and adult aphids. Pupation may take place
inside or outside the mummified host. Several species have been introduced from

abroad in biological control programs (697).
Family Hybrizontidae
Hybrizontids

This small family, consisting of one genus and two species, was originally
considered a subfamily of the Braconidae. Because of their rarity, little is known of

their host relationships. They may be parasites of ants or aphids (697).
Family Braconidae
Braconids

The family Braconidae constitutes one of the major groups of insect parasites.
The majority of species, nearly 2,000 for North America, are parasitic in the larvae
of Lepidoptera, but a large number are also parasitic in the larvae of several other
orders, especially the Coleoptera. Braconid adults are seldom more than 15 mm
long. They resemble those of the family Ichneumonidae in lacking a costal cell but
differ in not having more than one recurrent vein. Many species pupate in silken
cocoons on the outside of the body of the host, whereas others spin cocoons entirely
apart from their hosts. There are from one to many generations per year, depending
on the species. In some species the life cycle may be completed in less than 2 weeks
(216).

In addition to the large number of native species attacking forest insects, several
species have been imported from abroad to parasitize important introduced pests.
Following is a brief description of a few of these.

Agathis pumila (Ratzeburg) was imported into the United States and Canada
against the larch casebearer during the 1930's. It is now widely distributed through-
out eastern Canada and the Northeastern United States and is providing a high
degree of suppression in some areas. It has also been transferred from the Northeast
into Idaho where the larch casebearer was first discovered on western larch in 1957.
Genetic stock of A. pumila from Wisconsin was also released and established in
Oregon and Washington in 1973 and 1974 (/047). Winter is spent in the larval stage
within the host and there is one generation per year.

Meteorus versicolor (Wesmael), a parasite of the browntail moth and various
other species of Lepidoptera, was introduced from Europe into the United States
early in the century, and is known to have been established since 1909. It is now
widely distributed throughout the range of the browntail moth in New England, but
appears to be of little value in its suppression. Attempts to establish it on the satin
moth in the Northeast have failed. It is established on the satin moth in the Pacific
Northwest, however, where it is considered an important control factor (2/8, 328).

413

Dolichogenidea (= Apanteles) lacteicolor Viereck (825), a parasite of the
browntail moth in Europe, was introduced in 1907, established in the United States
by 1908, and is now generally distributed throughout the range of its host. Studies
made several! years after its establishment showed a rate of parasitization less than
10 percent of overwintering larvae over a wide area (2/8). Winter is spent within
young browntail moth larvae. The adults appear in the spring, and two generations
may develop utilizing alternate hosts (1/73, 894).

Cotesia (= Apanteles) melanoscelus (Ratzeburg) (825), a European parasite of
the gypsy moth, was introduced to New England in 1911 and 1912 and became
established quickly. Later, it was recolonized widely and is now generally estab-
lished over the infested area of New England. Parasitization is sometimes high in
localized areas in the Northeast. Its abundance is greatly reduced in the spring by
hyperparasites that attack overwintered cocoons. Under the name Apanteles soli-
tarius (Ratzeburg), this parasite was released in New England against the satin moth
in 1927. Parasitization as high as 67 percent has been recorded. The winter is spent
either as a first instar in a satin moth larva or as a prepupa in a satin moth cocoon.
Thus, two separate broods of adults arise from the overwintering generation, each
of which produces a second generation during the summer (957).

Phanomeris phyllotomae Muesebeck was imported from Austria to New England
and New York in the early 1930’s against the birch leaf-mining sawfly. It became
established but its effectiveness in suppression has not been determined. There is
one generation a year (327).

Orgilus obscurator (Nees) was imported by the United States and Canada in the
1930’s against the European pine shoot moth and is now widely distributed in most
shoot moth infested areas. Parasitization is variable, and in the United States, has
ranged from very low to as high as 50 percent. Significant levels of parasitization
have also been recorded in parts of Canada. Winter is spent as a first or second

instar within the hibernating host larva and there is one generation per year.
Family Ichneumonidae
Ichneumons

There are more than 3,000 described species of ichneumons in the United States
and Canada, and probably at least 5,000 undescribed ones (/208). The family has
been divided into 27 subfamilies and 502 genera (697). All members of the family
are parasites of the larvae and pupae of holometablous insects, or of spiders, spider
egg sacs, or pseudoscorpions. The majority of insect hosts belong to the order
Lepidoptera, but many species of Hymenoptera, especially the sawflies, and a
considerable number of Coleoptera are attacked.

Ichneumon adults vary greatly in size, form, and coloration. They resemble
slender stinging wasps, but differ in having the antennae longer and with more
segments, in having the trochanter two-segmented, in having the ovipositor perma-
nently extended, and in lacking a costal cell in the front wing. They also resemble
adult braconids but differ in having two recurrent veins in the forewing instead of
just one.

Forest and shade tree insects are parasitized by a great many species of ich-
neumons, far too many to mention here (697). The majority are native to this
continent and attack native hosts. In addition to these, a number of species have
been introduced from abroad against several important introduced hosts (328).

Pleolophus basizonus (Gravenhorst), a European parasite of sawfly cocoons, was
introduced originally to Canada against the European spruce sawfly during the
1930’s, and several colonies were made available for liberation in the United States

414

against both the European spruce sawfly and the European pine sawfly. It became
established on the European pine sawfly, which it parasitizes heavily at times. P.
basizonus also has been reared from cocoons of the introduced pine sawfly in North
Carolina and from Neodiprion pratti paradoxicus in New Jersey (328). It has not
been recovered from the European spruce sawfly in the United States.

Exenterus amictorius (Panzer), a parasite of the European pine sawfly in Europe,
was imported to Canada during the 1930’s against the European spruce sawfly and
European pine sawfly. Colonies were later made available for release in the United
States. It is an important parasite of the introduced pine sawfly in Wisconsin and
North Carolina, and has been recovered from several Neodiprion species including
N. sertifer, N. lecontei, N. nanulus nanulus, and N. swainei (697).

Mesoleius tenthredinis Morley was imported to Canada from England in 1910
and 1911 and liberated against the larch sawfly in Ontario, Canada. A small colony
was also released against the sawfly in Michigan. It became established and for
many years effected considerable suppression. Gradually though, the host devel-
oped an immunity to the parasite in many portions of its range, thereby greatly
reducing its effectiveness. A Bavarian strain of M. tenthredinis is showing consider-
able promise in Canada (1216). Olesicampe benefactor Hinz was introduced from
Europe to Canada to supplant M. tenthredinis in suppression attempts against the
larch sawfly. It is doing this very well in the Prairie Provinces and is established in
Minnesota (629, 704, 896), and a recent release seems promising in Pennsylvania
(341).

Coccygomimus turionellae turionellae (L.), a European parasite of the pupae of
many Lepidoptera, was imported against the European pine shoot moth during the
1930’s without success. Releases have been made against other lepidopterans, and
though it has not been recovered in the United States, it is apparently established in
southern Ontario (697).

Temelucha interruptor (Gravenhorst) was imported from England and Europe
against the European pine shoot moth during the 1930’s. It became established and
was recovered in Connecticut, New Jersey, and southern New York in 1937. Ten
years later, however, it had almost disappeared, being collected at only one point in
Connecticut. This species is cleptoparasitic on Orgilus obscurator and thus is
detrimental to the biological control program against the European pine shoot moth
(697).

Eastern forest insects also are attacked by numerous native species of ich-
neumonid parasites. /toplectis conquisitor (Say) parasitizes a tremendous number
of species, and is especially important. /. quadricingulata (Provancher) and /.
viduata (Gravenhorst) also attack a great many species. Megarhyssa macrurus (L.)

is a conspicuous parasite of the pigeon tremex.
Superfamily Chalcidoidea
Chalcids

This superfamily contains more species than any other superfamily in the order
Hymenoptera. It contains, among its families, probably a majority of all ento-
mophagous insects, with an extremely wide range in form, habits, host preferences,
and host relationships. The entomophagous species make up the majority and are
mostly beneficial, but there also are a number of phytophagous species, many of
which are economic pests. The species occurring in the United States and Canada
originally were divided into 21 families and more than 460 genera (974); however,
current researchers divide the superfamily into 11 families, 490 genera, and more
than 2,000 species (697).

415

Practically all of the more common orders of insects serve as hosts for the
parasitic and predacious members of the Chalcidoidea, with the Lepidoptera,
Diptera, Coleoptera, and Homoptera being preferred. The majority of the injurious
tree-infesting species in this country are those that destroy the seeds of their hosts.

Chalcids are mostly small to minute in size—some are less than 0.25 mm in
length. The antenna is elbowed, the pronotum does not extend back to the tegula,
the trochanter is two-jointed, the forewing is without either a stigma or closed cells,
the ovipositor issues some distance before the apex of the abdomen, and a prepectus

is present (697).
Family Mymaridae
Mymarids

This family, closely related to the Eulophidae, is represented by more than 100
species in the United States and Canada, the majority of which occur in the East.
All members of the family are internal parasites in the eggs of other insects,
particularly of Homoptera. Adults are mostly black or yellow and are extremely
minute, usually less than | mm long.

Polynema striaticorne Girault is an important parasite of several species of
membracids. It also attacks various aphids and other insects. Its bionomics in Ohio
and Illinois have been studied (52). Acmopolynema bifasciatipenne (Girault) para-
sitizes the eggs of several species of tree crickets. Ooctonus aphrophorae Milliron
attacks the Saratoga spittlebug (697).

Family Trichogrammatidae
Minute Egg Parasites

The family Trichogrammatidae, with some 43 species, consists of extremely
small insects all of which are internal parasites in the eggs of other insects (/5,
697). Hosts have been recorded from the orders Lepidoptera, Coleoptera,
Hymenoptera, Neuroptera, Diptera, and Hemiptera, but the Lepidoptera are pre-
ferred.

Trichogramma minutum Riley parasitizes the eggs of a great many species of
insects, including many important enemies of trees. The following is a partial list of
important eastern hosts: locust leafminer, gypsy moth, browntail moth, or
angestriped oakworm, saddled prominent, walnut caterpillar, satin moth, hickory
shuckworm, European pine shoot moth, Nantucket pine tip moth, forest tent
caterpillar, spruce budworm (as many as 75 percent of spruce budworm eggs may
be attacked), eastern blackheaded budworm, bagworm, elm sawfly, fall webworm,
yellowheaded spruce sawfly, and fall cankerworm (974).

The adult is less than 0.5 mm long, and females insert their eggs directly into
host eggs in arboreal habitats. During warm weather, the life cycle may be

completed within 9 to 16 days and there may be 12 or more generations per year.
Family Eulophidae
Eulophids

Adults of this family are very small, ranging in length from | to 3 mm. Well over
100 species are known to parasitize tree-infesting insects, several of which are
important pests (974). A number of foreign species have been imported to the
United States and Canada in efforts to suppress several species of introduced pests.
The family contains more than 500 North American species (697).

Chrysocharis laricinellae (Ratzeburg) (fig. 198), a parasite of the larch case-
bearer and the birch leafminer, was introduced from Europe to New England and
Canada in the late 1920’s and 1930’s. It is now widely established. Adults are bright
metallic-green with pale-yellow legs, and are about 2 to 3 mm long. There may be
three generations per year in the casebearer, but there is only one and a partial

416

second in the leafminer. C. laricinellae was reported ineffective against the case-
bearer because it acted as a secondary parasite, attacking the more effective A gathis
pumila (327). Recent studies indicate it can be effective, especiaily in concert with
A. pumila, as a successful multiple parasite. The species has been transferred to the
Northwest where it 1s established on the larch casebearer (/047).

F-519579

Figure 198.—Adult of the parasite Chrysocharis
laricinellae.

Dahlbominus fuscipennis (Zetterstedt), a parasite of several species of sawflies in
Europe, was introduced to Canada in 1934 for release against the European spruce
sawfly. The following year, shipments were received from Canada for release
against the same species in Maine. Since then, releases have been made against
several other sawflies in Canada and the United States. So far, it has been recovered
in this country from the European spruce sawfly in New England, the European
pine sawfly in New Jersey, the redheaded pine sawfly in Michigan, the Virginia
pine sawfly in Virginia, Neodiprion pratti paradoxicus Ross and N. pinusrigidae
(Norton) in New Jersey, the red pine sawfly and the introduced pine sawfly in
Wisconsin and North Carolina, the balsam fir sawfly in Maine, Gilpinia frutetorum
(F.) in Connecticut, and others (328, 697).

Adults are 2.3 to 2.8 mm long. The head, thorax, and abdomen are black; the
wings have a smoky tinge; the legs are white to light brown, except the femur which
is black; and the antennae are elbowed and black, except for a white scape in the
female. This species is an external parasite of sawfly larvae and of pupae within
their cocoons. There are from two to seven generations per year, depending on
location. It has never become a very effective parasite of the spruce sawfly. Yet,
almost 50 percent parasitization of the European pine sawfly has been recorded
locally in New Jersey.

The genus Jetrastichus contains a large number of parasites of important forest
insect pests. Descriptions of most North American species and information on their
distribution and hosts have been published (/5, /75).

Tetrastichus turionum (Hartig) was imported from Europe during the 1930’s and
released in New England, New York, and New Jersey against the European pine
shoot moth. Several years later it was also released in Ontario. It occurs in
Massachusetts, New Jersey, Connecticut, on Long Island, N.Y., and in southern
Ontario. The adult is a tiny, iridescent, blue-green insect. The antennae are brown
and the apices of the femora and the tarsi are light yellow or white. This is a pupal
parasite and it has one generation per year. So far, it has been of no consequence in
suppression of the shoot moth in this country.

417

Tetrastichus brevistigma Gahan 1s a pupal parasite of the elm leaf beetle. Adults,
are black with a slight metallic-green tinge and are only about 0.5 to 1.5 mm long.
An average of 12 parasite larvae develop in each pupa and there are three or four
generations per year. Parasitization as high as 50 to 80 percent occurs commonly in
the vicinity of Boston, Mass., where it was introduced from California. T. holbeini
Girault and T. rugglesi Rohwer attack several species of Chrysobothris and Agrilus,
respectively.

Dimmockia incongrua (Ashmead) parasitizes the gypsy moth and many other
species of Lepidoptera, although it 1s almost always a secondary parasite (697).
Elachertus cacoeciae Howard parasitizes various species of Lepidoptera. Members
of the genus Hyssopus parasitize several species of shoot and tip moths and seed-

and cone-infesting insects.
Family Encyrtidae
Encyrtids

This is a large family of parasites. Insect hosts are widely distributed among the
various orders, but the majority of species are parasitic on aphids, scales, and
whiteflies. Adults are | to 2 mm long.

Coccophagus insidiator (Dalman), a European parasite of the introduced Euro-
pean elm scale, was discovered at Ithaca, N.Y., in 1924, where it was parasitizing
the scale quite heavily. It had apparently been introduced to the area by accident.
The female is an endoparasite of the scale; the male an ectoparasite of the larval
stage of the female parasite. There may be three to five generations per year.

Ooencyrtus kuvanae (Howard), a parasite of gypsy moth eggs, was introduced to
the United States in 1908 and 1909 from Japan (/5, 267). The adult 1s black and
about | mm long. Winter is spent in the adult stage, during which mortality may be
severe.

Surviving adults appear during April and lay their eggs in the overwintered eggs
of the host. There may be one or two spring generations; for the entire year there
may be four or five generations. This species is an important parasite of the gypsy
moth in the southern portion of its range. Parasitization of 40 to 50 percent
frequently occurs in Massachusetts and Connecticut. O. ennomophagus Yoshimoto
(fig. 199), a parthenogenetic member of this family, was responsible for the
suppression of the elm spanworm in Connecticut in the early 1970’s (/5). It
develops only in unembryonated eggs of its hosts (653). O. trinidadensis Crawford
parasitizes eggs of the leaffooted pine seed bug and of the shieldbacked pine seed
bug, both pests of loblolly pine cones and seed.

~~

F-531262

Figure 199.—Ooencyrtus ennomophagus parasitizing
eggs of Eutrapela clemataria, the purplish-brown
looper.

418

Habrolepis dalmanni (Westwood) is sometimes fairly common as a parasite of
the golden oak and oystershell scales in this country. It has been introduced to New
Zealand from New England and is credited with having saved the oaks in that
country. Kermes pubescens Bogue, another oak scale, is attacked by at least five
other species of encyrtids. The European fruit lecanium is attacked by more than 20
different species. The San Jose scale and oystershell scale are also attacked by

several species.
Family Eupelmidae
Eupelmids

Members of the family Eupelmidae parasitize a wide range of insects including
Coleoptera, Orthoptera, Diptera, Lepidoptera, Hymenoptera, Hemiptera, and Ho-
moptera. Many other species are hyperparasitic and a few are phytophagous.

Anastatus disparis Ruschka, an egg parasite of the gypsy moth in Europe and
Japan, was introduced to New England in 1906, and quickly became established
(15, 267). It is now generally distributed throughout the infested parts of New
England. Adult females are marked with green and brown, have two broad fuscous
bands on the wings, and are 2 to 3 mm long. Males are greenish black with hyaline
wings, and are only about 2 mm long. The winter is spent as a mature larva within
the gypsy moth egg. Adults appear in June and July and lay their eggs in egg masses
of the host. Hatching occurs quickly and larval development is rapid, as the
hibernating stage is reached within about 2 weeks. Yet, there is only one generation
per year. This species is usually scarce in areas where the egg parasite, O. kuvanae,
also occurs. Interspecific competition may be the cause. A. reduvii (Howard) is an
egg parasite of the leaffooted pine seed bug and the shieldbacked pine seed bug.

A few other species of eupelmid parasites and some of their hosts are: Eupelmella
vesicularis (Retzius)—gypsy moth, satin moth, forest tent caterpillar, European
pine shoot moth, and several species of pine sawflies; Eupelmus cyaniceps Ash-
mead—webbing coneworm, Nantucket pine tip moth, bagworm, smaller European
elm bark beetle, European pine shoot moth, and bagworm; E. pini Taylor—white
pine weevil; EF. allynii (French)—various species of tree crickets; and Metapelma

spectabile Westwood—buprestid and cerambycid borers.
Family Eucharitidae
Eucharitids

The family Eucharitidae comprises only 6 genera and 28 species. As far as
known, all species are parasitic on the pupae of ants. Adults are distinguished by the
configuration of the scutellum, which is frequently produced backward in the form
of powerful spines (697).

Family Torymidae
Torymids

Most members of this fairly large family (27 genera and 175 North American
species) are parasites of gall-forming insects (697). A number of others are parasitic
on various lepidopterous larvae; many feed in the seeds of various plants; and some
act as hyperparasites.

The genus Jorymus contains a large number of species that parasitize the
immature stages of gall-forming cynipids and gall midges. T. rugg/esi Milliron has
been collected from the seeds of American holly in Delaware.

Members of the genus Megastigmus are all phytophagous, developing in the
seeds of plants. Eastern species include M. amelanchieris Cushman—on ser-
viceberry; M. laricis Marcovitch—on larch; and M. specularis Walley—on balsam
fir. The latter has destroyed up to 40 percent of balsam fir seed during certain years
in eastern Canada.

419

The genus Monodontomerus contains several important parasites of various
species of Lepidoptera and sawflies. M. dentipes (Dalman), a European species
which probably entered this country with its European host, the introduced pine
sawfly, is now widely distributed in southeastern Canada, in the Northern States
from Maine to the Lake States, and in North Carolina. It is not only an effective
parasite of the introduced pine sawfly, but also an important parasite of several
other sawflies attacking conifers (fig. 200). It spends the winter as a prepupa inside
the host cocoon. Adults appear over a fairly long period in the spring and lay their
eggs through the host cocoon, depositing several upon the prepupa. There are
probably two generations per year in the North.

F-531263
Figure 200.—Monodontomerus dentipes ovipositing in a
cocoon of the introduced pine sawfly.

Monodontomerus aereus Walker was introduced to New England from 1906 to
1910 against the gypsy and browntail moths. It was released originally as a primary
parasite but is much more common as a hyperparasite, attacking both hymenop-
terous cocoons and tachinid puparia. It is also parasitic on the whitemarked tussock
moth and the eastern tent caterpillar.

Other species of Monodontomerus recorded as parasites of important forest
insects in the Eastern States include M. indiscretus Gahan—on birch leafminer; M.
montivagus Ashmead—on spruce budworm; and M. minor (Ratzeburg)—on east-

ern tent caterpillar, cecropia moth, and spruce budworm.
Family Pteromalidae
Pteromalids

The family Pteromalidae is the largest in the superfamily (129 genera and 395
North American species), and its members act as parasites or hyperparasites of
almost all orders of insects (697). The adults are minute, black or metallic-green or
bronze insects. Many have a more or less triangular abdomen.

Perilampus hyalinus Say, a common species throughout the United States, may
act as a secondary parasite of a large number of insects in which it develops at the
expense of many species of primary tachinid and ichneumonid parasites. The adult
is a bright metallic bluish green and is from 2 to 4 mm long. The thorax is large and
the abdomen triangular. More than 20 species of Orthoptera, 16 species of Lepidop-
tera, 12 species of sawflies, and many species of ichneumonid, sarcophagid, and
tachinid parasites are listed as hosts (974); studies of its biology as a secondary
parasite of the fall webworm and as a primary parasite of the Swaine jack pine
sawfly have been reported (//07, 12/1).

420

Schizonotus latus (Walker), a primary parasite of the imported willow leaf beetle
and several allied species of Chrysomelidae, is widely distributed in the Eastern
United States. The larva feeds externally on the pupa of its host. High percentages
of parasitization were recorded over a 3-year period in the vicinity of Boston,
Mass., but host populations were not materially reduced (326).

Dibrachys cavus (Walker) 1s a hyperparasite of many primary parasites, but also
acts as a primary parasite. It is extremely destructive to many beneficial parasites.
In common with the adults of many other parasites, the adults often feed at puncture
holes made by their ovipositors. This results in the death of many larvae that are not
parasitized.

Family Eurytomidae
Eurytomids

This family contains a number of both phytophagous and entomophagous spe-
cies. Many form galls in the stems of grasses and other plants; some are parasites of
gall-forming Diptera and Hymenoptera; a few are egg parasites of Orthoptera; and
others are parasites of various tree-infesting Coleoptera. Adults are usually black
and the abdomen is rounded or oval and somewhat compressed.

Eurytoma pissodis Girault is one of the most important parasites of the white pine
weevil. The adult is dull black on the thorax, glossy black on its shining abdomen,
has conspicuous red eyes, and is from 3 to 6 mm long. Eggs are laid on mature
weevil larvae and the winter is spent as a prepupa within the pupal cell of the host.
Parasitization of at least 50 percent has been recorded in some white pine weevil
infestations.

Species parasitic on other important forest insects include: E. pini Bugbee—on
European pine shoot moth, Nantucket pine tip moth, pitch pine tip moth, and
possibly the introduced pine sawfly; E. tylodermatis Ashmead—on hickory shuck-
worm, European pine shoot moth, and Nantucket pine tip moth; and E. magdalidis

Ashmead—on southern pine beetle and Pityophthorus liquidambarus Blackman.
Family Chalcididae
Chalcids

The family Chalcididae contains many primary and secondary parasites of
Lepidoptera, Diptera, Coleoptera, and Orthoptera. The adults of certain species are
fairly large and conspicuous; all are solitary in habit and practically all develop
inside their hosts.

A few of the more important species and their hosts are: Haltichella rhyacionia
Gahan—Nantucket pine tip moth, pitch pine tip moth, and a western pine tip moth;
H. xanticles (Walker)—European pine shoot moth and oak skeletonizer; Phas-
gonophora sulcata Westwood—bronze birch and flatheaded appletree borers; Trig-
onura elegans (Provancher)—several species of woodborers including flatheaded
appletree borer, red elm bark and black elm bark weevils; Spilochalcis albifrons
(Walsh)—numerous hosts including locust and arborvitae leafminers, and larch
casebearer; S. flavopicta (Cresson)—Nantucket pine tip moth; S. mariae (Riley) —
bagworm and several species of saturniid moths; Brachymeria compsilurae
(Crawford)—tachinid flies, especially Compsilura concinnata (Meigen) and
Blepharipa pratensis (Meigen) (325); and B. ovata (Say)—a wide variety of
lepidopterous hosts including whitemarked tussock moth, hemlock looper, and
bagworm.

Superfamily Cynipoidea

The superfamily Cynipoidea consists of very small, dark-colored wasps. Many
species are gall makers or gall inquilines. The remainder are parasitic on other
insects. This superfamily is divided into six families—Ibaliidae, Liopteridae,

421

Figitidae, Eucoilidae, Alloxystidae, and Cynipidae (697). The liopterids are mostly

exotic, and their habits are unknown.
Family Ibaliidae
Ibaliids

Members of this family are all parasitic on horntails of the family Siricidae. At
least four species are recorded from the Eastern United States—J/balia anceps Say,
I. leucospoides ensiger Norton, 1. maculipennis Haldeman, and I. scalpellator
Westwood (697).

Ibalia scalpellator, a common species in the Eastern United States and south-
eastern Canada, is a parasite of pigeon tremex. The adult is marked with a yellow
and dark-brown pattern, has two conspicuous dark-brown to black bands on the
forewings, and is about 12 mm long.

Family Figitidae
Figitids

This family of 60 North American species is represented in eastern America by
about 30 species. The majority are parasitic in dipterous puparia; a few are parasitic
in the cocoons of chrysopids.

Family Cynipidae
Cynipids or Cynipid Gall Wasps

This family of several hundred species consists mostly of species that induce
galls on their plant hosts in which they live and feed during the larval stage. Many
others live as inquilines in galls produced by other insects. The adults are small to
extremely small and usually black. They are distinguished by the abdomen that is
oval, shining, somewhat compressed, and almost covered by the first tergite. Many
gall-forming species produce two quite different generations per year. One genera-
tion develops during the summer in one type of gall. Adults appear in the fall and
consist entirely of parthenogenetic females. Eggs laid by these females give rise to
larvae that induce an entirely different kind of gall. Adults of this generation consist
of both males and females, and may be quite different in appearance from those of
the first generation.

It is estimated that 86 percent of the known gall-forming species induce galls on
oaks and are confined to them (668). There are 717 species listed as occurring in
the United States and Canada, 76 percent of which induce galls on oaks.

The females deposit their eggs in the tissues of all parts of the host, from the
roots to the flowers. Gall production is believed to result from the reaction of the
cambium and other meristematic tissues to stimuli produced by the larvae. The
great majority of species are of little or no economic importance. However, certain
species that induce large irregular galls on the smaller branches are capable of
causing injury (fig. 201). Infested branches may be disfigured or even killed;
occasionally entire trees are killed. On the other hand, galls induced by certain
other species are economically valuable—some have long been used in the man-
ufacture of ink and in dyeing and tanning, others serve as a source of winter food
for bees. One, a deciduous oak gall, is occasionally abundant enough on black oak
in Missouri to be used as food for hogs, cattle, sheep, turkeys, and chickens (667,
669). Some of the more common and important gall-inducing species are discussed
below.

Callirhytis floridana (Ashmead) occurs from Virginia to Florida and Missouri
and Arkansas. It induces slender, elongate swellings from 12 to 75 mm long on
branches close to the ground of Chapman, post, and sand post oaks. Large areas of
sand post oaks were nearly all killed during an outbreak in eastern North Carolina
(669).

422

N

is .
F-520107

Figure 201.—Heavy infestation of galls on twigs of
Nuttall oak in the Mississippi River Delta region.

Callirhytis quercuspunctata (Bassett), the gouty oak gall, occurs from southern
Canada to North Carolina and west to Illinois. It induces galls (fig. 202) on the
twigs and smaller limbs of scarlet, pin, and black oaks. These galls are about 12 to
38 mm long and they frequently occur so close together that they form practically
continuous masses. C. quercuspunctata has alternate generations. The first induces
small blisterlike galls on the leaves near the veins in the spring. The second induces
gouty galls during the summer. In heavy infestations, twigs, fairly large branches,
and even entire trees may be killed. Shade trees are especially subject to damage.

Callirhytis cornigera (Osten Sacken), the horned oak gall, occurs from southern
Canada to Georgia and west to Iowa. It induces galls on the twigs of pin, scrub,
black, blackjack, and water oaks. Injurious infestations have been observed on
roadside and woodland oaks in the Niagara area of Canada.

Callirhytis quercusgemmaria (Ashmead), the ribbed bud gall, occurs from
Massachusetts to Florida, and west to Illinois. It induces somewhat conical,
strongly ribbed galls about 5 mm long on the twigs of black oaks. These galls
sometimes occur in such large numbers as to cause infested twigs to split and die.
Entire young trees are sometimes killed.

423

Callirhytis quercusoperator (Osten Sacken) occurs from New England to North
Carolina and west to Iowa. It causes the formation of woolly galls on the staminate
flowers of various oaks. When the adults emerge, they oviposit in immature acorns.
This results in the formation of so-called acorn pip galls within the acorn cups. In
heavy infestations, acorn crops are reduced. Callirhytis quercusfutilis (Osten
Sacken) induces globose, grayish galls up to 11.4 cm long on the main roots of
young white oaks just below the ground line.

Neuroterus quercusbatatus (Fitch), the oak potato gall (fig. 203). occurs from
Ontario and Rhode Island to Florida and west to Illinois on white oaks. There are
two generations per year. Adults of the first generation emerge in May from galls
produced on the preceding year’s growth. Second-generation adults emerge from
green galls. Females of this generation lay their eggs in the same galls from which
they emerge.

Neuroterus floccosus (Bassett), the oak flake gall, induces small. hemispherical
galls from 1.5 to 3.5 mm in diameter on the lower surface of terminal leaves of
white oaks. These galls are covered with white hairs and often occur in large
numbers on a single leaf. Heavily infested leaves curl and are unsightly on shade
trees.

Neuroterus noxiosus (Bassett), the noxious oak gall, occurs on swamp white
oaks from New England to Virginia and in the Central States. Heavily infested trees
may be severely disfigured.

/ ff
Courtesy Ill. Nat. Hist. Surv. ~ Courtesy Am. Mus. Nat. Hist.
Figure 202.—Gouty oak galls caused by Figure 203.—Oak potato gall on white
Callirhytis quercuspunctata. oak caused by Neuroterus
guercusbatatus.

424

Amphibolips confluenta (Harris), the large oak-apple gall (fig. 204), occurs
from southern Canada to Virginia. It produces galls on the leaves or leaf petioles of
various Oaks, principally red, black, and scarlet. These galls are large, from 12 to
50 mm in diameter, and greenish to brownish, depending on age. The related
species, A. quercusfuliginosa (Ashmead), produces globose galls on the sides of the
acorn cups of willow and laurel oaks in Florida.

Dryocosmus quercuspalustris (Osten Sacken), the succulent oak gall, occurs
from southern Canada and New England to Florida and west to lowa and Louisiana.
It induces somewhat circular, succulent galls from 9 to 12 mm in diameter on the
leaves or axils of staminate flowers of red oaks. The galls are fleshy walled and
hollow, except for a free-rolling cell about 2.5 mm in diameter. D. kuriphilus
Yasumatsu, native to Japan and Korea, is a pest of Chinese and Japanese chestnuts
in Georgia where it was found in 1974 (967).

Xanthoteras quercusforticorne (Walsh), the oak fig gall, induces galls on the
leaves, twigs, and stems of white oaks in the Eastern States north of Virginia and
west to Minnesota. Early in the season they are reddish and bladderlike. Sometimes
they occur around a twig in dense clusters up to 8 cm long, and the irregular masses

look like pressed figs.
Superfamily Evanioidea—Family Evaniidae
Ensign Wasps

Members of this small family of spiderlike wasps are parasitic in the egg capsules
of cockroaches. Adults are about 10 to 15 mm long and are distinguished by very

2 Courtesy Can. For. Serv., Can. Dep. Environ.,
; Sault Ste. Marie, Ont.

Figure 204.—Large oak-apple gall caused by
Amphibolips confluenta.

425

small, oval abdomens attached by petioles to the propodeum considerably above the
base of the hind coxae. The majority of described species occur in the Eastern

United States.
Family Aulacidae
Aulacid Wasps

The family Aulacidae consists of slender ichneumonlike wasps. The female has
an ovipositor about as long as the body. A number of species are parasitic on wood-
boring insects; the remainder are parasitic in the nests of bees and wasps in twigs or
wood. Pristaulacus rufitarsis (Cresson) is a parasite of the hemlock borer and
poplar borer in eastern America; P. bilobatus (Provancher) is.also parasitic on the
hemlock borer. Aulacus burquei (Provancher), A. digitalis Townes, A. lovei (Ash-

mead), and A. pallipes Cresson are parasitic on various species of Xiphydria.
Superfamily Pelecinoidea—Family Pelecinidae
Pelecinid Wasps

This family appears to be represented in North America by only one species,
Pelecinus polyturator (Drury), a parasite of June beetle larvae. The female is a large
shiny black insect, often up to 62 mm long. The abdomen is slender, about five
times as long as the head and thorax combined, and is without a sting. Males are
much smaller and have the posterior of the abdomen swollen.

Superfamily Ceraphronoidea—Family Megaspilidae
Megaspilid Wasps

A number of species in this family have been taken from colonies of ants. Most
are known to be hyperparasitic primarily on various hymenopterous and dipterous
parasites. Conostigmus virginicus (Ashmead) is a hyperparasite of Blepharipa
pratensis (Meigen), an introduced parasite of the gypsy moth.

Superfamily Proctotrupoidea

As far as known, all members of this superfamily are parasitic on the immature
stages of other insects. The group as a whole seems to be little known, and most of
the North American species are still undescribed. The adults of the majority of
species are black, often shiny, and small to extremely small in size. The smaller
ones resemble chalcids, but differ in having the pronotum extend laterally to the
tegulae and the ovipositor issue from the end of the abdomen. In many of the
smaller species, the wings are almost veinless; in others, the wings are entirely
absent.

Family Scelionidae
Scelionid Wasps

As far as known, all members of this large family of small insects are parasitic in
the eggs of other insects and spiders (/5). Species attacking some of the more
important forest and shade tree insects are: Telenomus dalmani (Ratzeburg)—
whitemarked tussock moth; 7. californicus Ashmead—satin moth and Douglas-fir
tussock moth, Orgyia pseudotsugata (McDunnough); 7. alsophilae Viereck—fall
cankerworm (fig. 205); 7. bifidus Riley—tfall webworm; T. catalpae Muesebeck—
catalpa sphinx; 7. droozi Muesebeck—elm spanworm; T. coelodasidis Ashmead—
saddled prominent; Te/enomus sp.—cherry scallop shell moth; and 7. clisiocampae
Riley—forest tent and eastern tent caterpillars. Gryon pennsylvanicus (Ashmead)
parasitizes eggs of the leaffooted pine seed bug.

Family Platygastridae
Platygastrid Wasps

The majority of species in this family are parasites of Diptera, especially of the
families Cecidomytidae and Tipulidae. A number of others are also important as
enemies of mealybugs and whiteflies. The introduced species, Allotropa burrelli

426

F-532019

Figure 205.—The parasite Jelenomus alsophilae
ovipositing in eggs of fall cankerworm.

Muesebeck, an important parasite of the Comstock mealybug, has been widely
released in the Eastern United States. Two native species, A. ashmeadi Muesebeck
and A. convexifrons Muesebeck, also are important parasites of the Comstock

mealybug.
Superfamily Bethyloidea—Family Chrysididae
Cuckoo Wasps

Members of this family are brilliantly metallic blue and green wasps and are
popularly known as gold or cuckoo wasps. The majority of species are external
parasites of various wasps and bees. A number of species deposit their eggs in the
cells of their hosts, and the larvae feed on the original occupants of the cells or on
the food prepared for them.

Chrysis shanghaiensis Smith, the only species known to be parasitic upon
lepidopterous larvae, was introduced to Massachusetts from Japan in 1917 and 1918
against the oriental moth, also an introduced species. Recoveries were made the

year following its release (956), but it has not been recorded since that time.
Subfamily Cleptinae

This subfamily is represented in North America by one genus, Cleptes, which is
represented by seven species. One of these, C. semiauratus (L.), occurs in the East.

All parasitize sawflies in their cocoons (697).
Subfamily Amiseginae

Mesitiopterus kahlii Ashmead, M. floridensis Krombein, Microsega bella
Krombein, and Ade/phe anisomorphae Krombein are egg parasites of walk-
ingsticks.

Family Bethylidae
Bethylid Wasps

Members of the family Bethylidae are parasitic almost exclusively on the larvae
of Lepidoptera and Coleoptera. More than 100 species have been recorded from the
United States, about two-thirds of which occur in the eastern portion of the country.
The adults are small to medium in size. Females are often wingless and antlike and
differ so much in appearance from the males that the two sexes are not easily
correlated.

A number of species of the genus Sclerodermus are often parasitic on larvae of
the family Cerambycidae. Additional species of importance as parasites of forest
insects and their hosts are Pseudisobrachium prolongatum (Provancher)—black
carpenter ant; Parasierola punctaticeps Kieffer—hickory shuckworm; Goniozus

427

electus Fouts—Nantucket pine tip moth: and G. longinervis Fouts—a western pine
tip moth.

Family Dryinidae

Dryinid Wasps

This is a small family of rarely collected insects. As far as known, all species are
parasitic on the nymphs of Homoptera, especially the membracids, cicadellids,
flatids, and cercopids. The females of some genera are wingless and antlike. Also.
in most species they differ so much in appearance from the males that the two sexes
can be associated only by rearing them.

During the larval stages, these insects are intermediate between ectoparasites and
endoparasites in the abdomens of their hosts. Usually one or more external gall-like
cysts develop on the integument of the host. These cysts, which may be as large as
the abdomen of the host, contain the parasite larvae. The larva absorbs food from
the host via the cyst. Aphelopus theliae Gahan, a parasite of the membracid, Thelia
bimaculata (F.), which feeds on young black locust, lays a single egg in a nymph of
its host. During oviposition the females secrete a substance that stops host meta-
morphosis. Polyembryonic development takes place and from 50 to 75 parasites are
produced in the nymph. When they reach maturity, they bore through the body wall
and drop to the ground to pupate (46/).

Superfamily Scolioidea—Family Tiphiidae
Tiphiid Wasps

The family Tiphiidae contains a number of important species parasitic on
scarabaeid larvae in soil. As far as known, all species are external parasites, usually
feeding on the final instar of the host. The adults of most species are moderate in
size, hairy, and black. The remainder are mostly black and yellow. A number of
species have been imported against several introduced pests.

Tiphia inornata Say 1s one of the most common and important parasites of white
grubs in the United States. Infestations are heaviest during the years when full-
grown white grubs are present. At other times, populations are usually very low
(1348).

Tiphia vernalis Rohwer was introduced against the Japanese beetle from Japan
and Korea during the period 1924 to 1933. It became established and now plays an
important role in control of the beetle in certain parts of the East, especially in
Pennsylvania (2/7). T. popilliavora Rohwer, also introduced against the Japanese
beetle during the 1920’s and 1930’s, is now well established from New Hampshire
to Virginia and Ohio. T. asericae Allen & Jaynes was introduced against the Asiatic
garden beetle and Serica peregrina Chapin during the 1920’s, and is established in

Pennsylvania, New York, and New Jersey (697).
Family Mutillidae
Velvet Ants

Members of the family Mutillidae are commonly known as velvet ants. The
females are wingless, have heavy coatings of fine hairs, and are usually observed
running back and forth over the ground. In certain regions they are also known as
‘“cow-killers.”” Adults are usually brightly colored with red, orange, or yellow
markings, and some of the females are 25 mm or more in length. The majority of
species are parasitic on ground-nesting bees and wasps (544). A common species in
the South, where members of the family are most prevalent, is Dasymutilla

occidentalis occidentalis (L.). Velvet ants are capable of inflicting painful stings.
Family Scoliidae
Scoliids

The family Scoliidae is represented by about 12 species in the Eastern United

428

States and, as far as known, all are external parasites of scarabaeid larvae in the
soil. The adults are large, hairy, and usually black except for spots or bands of
yellow or red, and the wings are often dark brown with purple or green iridescence.
Campsomeriellia annulata (F.), a native of China and Japan, was introduced against
the Japanese beetle in the 1920’s but did not become established.
Family Sapygidae
Sapygids

As far as known, all members of this family are parasitic on the larvae of
leafcutting bees and possibly of the large carpenter bees. Less than a dozen species
are recorded from the Eastern United States. Adults are moderate in size, short-

legged, and usually spotted or banded with yellow.
Family Formicidae
Ants

Ants are among the most abundant and widespread of all insects and are found in
practically all terrestrial habitats. The majority of species nest in soil. Others nest in
wood, timbers, in or under the bark of decaying trees, or in hollow stems of plants.
Some of the more primitive species feed on insects or other small animals which
they are able to kill. Many others feed on sweet fluids such as sap exuding from
wounds, on nectar, or on honeydew produced by other insects. Certain leafcutting
species cultivate fungi on which they feed.

Ants differ from their near relatives in having the abdomen divided into two
distinct regions, the pedicel and the gaster. Also, the antennae are elbowed, with the
first segment greatly elongated in females and workers. They also differ from
termites, with which they are often confused, by having a strong constriction or
“waist”? between the thorax and abdomen, and by having two pairs of wings of
unequal size.

Ants are social insects and live in nests or colonies, each containing from a few
to several thousand individuals. A colony consists of three castes—reproductive
females, males, and workers. Females generally are winged, the wings being
discarded after they mate. Males are usually smaller than females and generally
retain their wings until death. Workers are wingless and usually smaller than
reproductive females or males.

With the exception of the carpenter ants and certain leafcutting species, ants are
of minor importance as enemies of trees or wood products. Many species, however,
are nuisance pests in forested areas, especially in picnicking or other recreational
areas. Several publications on the biology and control of carpenter ants are available
(458, 1052, 1083, 1110).

The black carpenter ant, Camponotus pennsylvanicus (De Geer), nests in live
and dead standing trees, rotting logs and stumps, telephone and telegraph poles,
and wood of houses and other buildings. It is widely distributed in the Eastern
United States and Canada, occurring from North Dakota to Quebec and Ontario and
south to Texas and Florida. These large ants (fig. 206) include workers ranging in
length from 6 to 13 mm. The body is typically black, but in some individuals the
pleuron, petiole, and legs are reddish. The gaster is covered with dense, long,
appressed, pale yellowish and ashy pubescence. Body hairs are suberect or erect,
yellowish, and moderately abundant.

The black carpenter ant does not eat wood. It simply removes it in order to
produce galleries which serve as its nest. Its natural food consists largely of dead
and live insects, honeydew, sap, juices of well-ripened fruits, and refuse. It also
feeds on various household foods such as different kinds of sweets, raw and cooked
meats, and fruits.

429

Courtesy Conn. Agric. Exp. Stn.

Figure 206.—Black carpenter ant, Camponotus
pennsylvanicus: A, adult winged female; B, adult
winged male.

Live trees are infested occasionally but usually only when the ants are able to
enter through cracks, scars, knotholes, and decayed or other faulty places. Once
inside, they remove faulty wood and often extend their galleries into adjacent sound
wood. A wide variety of trees such as poplar, cherry, eastern white and pitch pines,
balsam fir, elm, willow, and red, white, scarlet, black, and post oaks have been
infested. Infestations frequently are located near the base but may occur very high
in a tree. Infested trees are often subject to serious injury. They are frequently
weakened to the point that they are subject to windbreakage. The wood also may be
rendered worthless for lumber or pulpwood (fig. 207).

Often houses are invaded by carpenter ants coming from nearby nests. This
happens frequently where houses are located in the vicinity of trees, logs, or
stumps. Entry is frequently gained through openings around the foundation of the
house or from tree branches in contact with the house. The woodwork may be
attacked in any number of places, but the most commonly damaged parts are
supporting timbers, porch pillars, sills, girders, joists, studs, window casings, and
external trim. The galleries are similar to those constructed by termites except that
they run across the grain, are sandpaper smooth, and free of frass. Evidences of
infestation are the presence of large black ants running about the house, swarms of
large black, winged ants about the house in spring, piles of sawdustlike borings,
slitlike holes in woodwork such as window and door casings, and faint rustling
sounds in walls, floors, woodwork, and flush-panel doors. Where infestations are
of long standing, damage to structural timbers may be severe and require extensive
repairs. Telephone and telegraph poles also are subject to serious damage (fig. 208)
(451).

430

F-504087

Figure 207.—Cross section of an oak log showing
galleries made by carpenter ants.

Courtesy Conn. Agric. Exp. Stn.

Figure 208.—Colony of carpenter ants in a 0.6-m
section of a telephone pole in winter. Arrow
indicates ground level.

431

Overwintering males and females in colonies over 3 years old engage in nuptial
flights from May to late July. Fertilized females then establish nests in cavities,
usually under the bark of a tree, log, or stump, and seal themselves inside. Here
they rear their first broods of workers to maturity on salivary secretions. These
workers, being inadequately fed, are smaller than normal. Subsequent broods are
fed by the workers, and individuals are larger. Long-established colonies contain
workers of various sizes, some of which are extraordinarily large. Such colonies
may consist of a reproductive female, scores of winged males and virgin females,
and several thousand workers.

Tightly constructed houses with concrete foundations, good clearance, and full
basements are fairly safe from invasion by black carpenter ants. Removing all wood
from near or under a house site before construction, making certain that infested
wood is not brought into the house, and cutting back branches in contact with the
house are also helpful in preventing infestations in buildings.

The red carpenter ant, C. ferrugineus (F.), occurs over much of the same
portion of the United States as the black carpenter ant, but is apparently less
common. It appears to prefer wooded areas where it nests in or beneath well-rotted
logs and stumps. Under these conditions, it often extends its galleries for consider-
able distances in the soil. Nests are also found in dead, standing trees but rarely in
houses. The workers are about the same size as those of the black carpenter ant.
Most of the thorax, petiole, base of the gaster, and much of the legs are yellowish
ferruginous. The remainder is black. Hairs and pubescence are more golden yellow
than those of the black carpenter ant, especially on the gaster. This species has
caused considerable damage to standing northern white-cedar in Minnesota (494).

The Florida carpenter ant, C. abdominalis floridanus (Buckley), occurs from
North Carolina to Florida and Alabama. It is one of the most important house-
infesting species in Florida. Workers are about 5.5 to 10 mm long. The head is
reddish, the thorax and petiole yellowish or yellowish red, the scape and gaster
blackish or black, and the body is covered with many long yellowish hairs. This
species builds its nest in various places such as in the ground beneath objects, in
dead branches in trees, in and beneath rotting logs and stumps, and sometimes in
the woodwork of porches, roofs, kitchen sinks, and paneling. Outdoors, the ants
feed largely on living and dead small insects, and on honeydew which they secure
by tending aphids, mealybugs, and scales. Indoors, they feed on such items as
molasses, honey, and raw and cooked meats.

Camponotus nearcticus Emery usually nests in small colonies in dead twigs and
branches of trees, in or beneath the bark of dead and living trees, in insect galls and
pine cones, in the hollow stems of plants, and in wooden posts. Nests have also
been found in the woodwork of houses, especially in roofing. It occurs from New
York to Ontario, North Dakota and Colorado, and south to Mississippi and Florida.
Workers are about 4.5 to 7.5 mm long, and their bodies are usually black and rather
shiny.

Camponotus sayi Emery occurs from North Carolina to Florida and west to
Colorado and California, but is apparently most common in the Gulf Coast States.
It nests in small colonies in tunnels made by borers in the twigs and branches of
various hardwoods, in insect galls, in cavities in the stalks of plants, under the bark
of trees, and in logs, stumps, wooden posts, and houses. Galleries in branches may
be anywhere from 2 cm to over 1.5 m long. Workers are 4 to 9 mm long. The head,
thorax, and petiole are usually yellowish red or reddish, and the gaster is blackish or
black. C. caryae discolor (Buckley), a species similar in appearance and habits to

432

C. sayi, has been recorded from Ohio to Kansas and Iowa and south to Texas and
Florida. It is most common in the lower Mississippi Valley. C. castaneus (Latreille)
occurs throughout the South but is most common in the Southeast. It nests in rotten
Jogs and stumps or in the soil. Workers are yellowish to yellowish red and from 7 to
10 mm long. C. tortuganus Emery occurs in the southern half of Florida and
apparently nests in small colonies in rotting wood or in the soil beneath stones. It
also occurs in houses where it may be a pest. C. mississippiensis M. R. Smith has
been found nesting in the new growth of white ash branches in Mississippi.
Galleries are apparently limited to | year’s growth. C. pylartes fraxinicola M. R.
Smith nests in the dead branches of various hardwoods. It has been recorded from
Mississippi.

The Texas leafcutting ant, Affa texana (Buckley), is a serious pest of pine
seedlings in eastern Texas and west-central Louisiana. It also defoliates and
damages a wide variety of other plants, including orchard trees. Injury to pines is
especially severe during the winter when there is a dearth of other green foliage.

Texas leafcutting ants are rusty red. The head is strongly bilobed, the antennae
are 11-segmented and without a well-defined club, the thorax bears three pairs of
prominent spines on top with the anterior ones the largest, and the legs are
extraordinarily long. The queen is about 18 mm long; workers, from 1.5 to 12 mm
long.

Nests of the Texas leafcutting ant are constructed in the ground, usually in well-
drained sand or loamy soiis and commonly on slopes facing the south or west (95).
The interior of the nest may reach a depth of 6 m. It may contain a thousand or more
entrance holes. Nest areas are usually marked by many crescent-shaped mounds up
to 12 to 35 cm high and 30 cm in diameter. Each mound surrounds an entrance hole
(fig. 209). The nest consists of many cavities connected by narrow tunnels. There
are vertical tunnels that extend to mound openings and lateral tunnels that lead
outward, sometimes for 90 m or more. Above ground, sharply defined foraging
trails sometimes extend hundreds of meters to the plants under attack. Ants move in
procession along these trails, each carrying a fragment of leaf or other material to
the nest. These fragments, which may be several times the size of the ants carrying
them, are borne upright over the head like a parasol.

7

F-482849

Figure 209.—Mounds of the Texas leafcutting ant, Atta
texana.

433

The ants do not eat the foliage that they remove from plants. Instead, they cut it
up into small fragments, shape it into small pellets, and carry it into their under-
ground chambers. Here it is placed upon so-called gardens, where it serves as a
medium for the growth of a fungus. It is this fungus that serves as the food of the
colony. Through the summer, most of the foliage is brought into the nest during the
night; in the fall, winter, and spring, most of it is brought in during the day, unless it
is too cold or wet (369).

Winged males and females appear during May and June and fly from the colony
and mate. Mated females lose their wings and dig into the soil where they establish
nests. Here they become the queens of new colonies.

The Allegheny mound ant, Formica exsectoides Forel, 1s a serious pest of young
eastern white, red, and Scotch pines, eastern redcedar, and spruce in the Eastern
and Northeastern States. It nests in the ground and constructs mounds that may be
up to 1.2 m in height and 1.8 m across. In forested areas, these mounds are most
often found in openings or along the edges of stands. All vegetation, except large
trees, may be destroyed in an area 12 to 15 m in diameter around a mound. Trees
from 2 to 15 years old are especially susceptible to attack. Damage may be severe in
forest plantations. The adult ant is about 3 to 6 mm long. The head 1s reddish brown
and about as wide as it is long, the thorax is reddish brown and feathered, the anal
region is reddish and surrounded by a fringe of hairs, and the legs are sometimes
brownish or dark red.

The Allegheny mound ant does not feed on trees or other vegetation. Its food
consists of living and dead insects and honeydew excreted by various species of
sucking insects. It appears that the only reason it attacks vegetation is to kill it to
Keep it from shadowing the mounds. Trees are killed by the injection of formic acid
into their tissues. Apparently this results in the coagulation of cell contents and the
prevention of the downward movement of foods in the inner bark (976).

The life history of the Allegheny mound ant is not too well understood, but this
ant is known to forage for food from April to September and to spend the winter in
its nest. There are several generations per year and both queens and workers are
known to live for several years.

Crematogaster cerasi (Fitch) occurs from southern Canada to Georgia. Its nests
are found in the ground, in rotting stumps, logs, or branches, and in empty nuts on
the ground. Nests also may be found in various parts of houses such as the roof,
siding, ceiling, and porch, but most often in and around door and window frames.
C. clara Mayr occurs from Indiana to New Jersey and south to Texas and Florida,
but is most common in the lower Mississippi Valley. Its nests are found in cane
stems, branches, trees, rotten stumps, and sometimes in the woodwork of houses.
C. lineolata (Say) occurs in southern Canada and throughout the Eastern United
States. It constructs fairly large nests, usually in the soil, but also in logs, stumps,
dead trees, or in the woodwork of houses. When alarmed, the workers bite fiercely
and give off a repulsive odor. C. laeviuscula Mayr and C. ashmeadi Mayr have
been recorded nesting in various dead hardwoods in Mississippi.

The Argentine ant, /ridomyrmex humilis (Mayr), an introduced species first
recorded in this country at New Orleans in 1891, now occurs in many localities in
the Southern States and California. Local infestations have also been found in St.
Louis, Baltimore, and Chicago (///0). Indoors, it feeds on almost every kind of
food, especially sweets, meats, pastries, fruit, eggs, dairy products, animal fats,
and vegetable oils; outdoors, it feeds partly on honeydew produced by aphids,
mealybugs, and scale insects. By fostering and protecting the latter insects from

434

many of their enemies, they encourage the development of heavy infestations that
are capable of causing serious damage to affected plants. They also remove seeds
from seed beds and feed on the sap or fruit juices from various trees and plants.

Argentine ants are more or less uniformly light brown or brown, the antennae are
12-segmented and without a club, and the petiole scale is well developed and
inclined. Workers are from 2.2 to 2.6 mm long; when freshly crushed, they emit a
stale, greasy, musty odor. Nests occur in many kinds of places, such as in the soil,
in rotten wood, in cavities in trees, in refuse piles, and in bird nests and beehives.
The number of ants present in well-established infested areas is beyond comprehen-
sion. Fortunately, they are usually of minor importance as pests in forested areas.

Iridomyrmex pruinosus (Roger) occurs throughout the Eastern and Southern
States north to New York and Wisconsin. It nests in the ground and under the bark
of logs and stumps, and is sometimes a nuisance in houses. The odor of freshly
crushed workers resembles that of rotten coconut.

The genus Solenopsis is represented by eight species in the United States, seven
of which occur in the Southeastern States. They are commonly called fire ants.
Nests are usually constructed in the soil, but also sometimes in rotten wood and in
houses. The workers are aggressive and practically omnivorous.

The primary diet of fire ants is insects, spiders, myriapods, earthworms, and
other small invertebrates. They also may feed on plants, particularly seedlings or
germinating seeds. Other economically important problems created by the imported
fire ants are stinging of livestock, damage to farm machinery that strike mounds,
loss of hay and grazing area, refusal of workers to enter heavily infested fields to
cultivate or harvest crops, and hazards to human health from stings that may cause
systemic reactions or complications from secondary infections (749).

Both the red imported fire ant, Solenopsis invicta Buren, and the black
imported fire ant, S. richteri Forel, were introduced from South America into the
United States at Mobile, Ala.—the former, about 1940 and the latter, about 1918.
S. richteri is more or less stable in a relatively small area in northeastern Mis-
sissippi and northwestern Alabama. S. invicta is more widespread and is now found
in nine States from the Carolinas to Texas. It is still spreading in all directions
although the northward spread is slow (749). The two species are often very
difficult to distinguish on the basis of color alone. Detailed examination of mor-
phological characters such as the shapes of the head, thorax, and postpetiole is
needed to separate these species (/72).

Imported fire ant mounds are found in almost all kinds of soil, but most
commonly in open areas. Few if any are ever found in heavily timbered areas,
especially in hardwood stands. An average mound is about 30 cm high and 60 cm
across; some are up to 90 cm tall. The larger ones are usually dome-shaped or
conical, and may contain more than 100,000 ants. In heavily infested areas there
may be more than 30 nests per hectare. Nests are also found in rotting logs, around
tree trunks, and occasionally under buildings. The biologies of these species have
been discussed (50/). Comprehensive treatments of ants may be found in the
literature (265, 1109, 1282).

Superfamily Vespoidea

. This superfamily comprises two families, the Vespidae and the Pompilidae.
Many species are social in habit and live together in colonies; others are solitary
and live alone. The adults generally feed on nectar, sap, honeydew, or similar
materials, whereas the larvae feed on other insects or spiders provided by the adult
female. The larvae of some species feed on pollen and nectar. A few species are of

435

some importance as pests of forest or shade trees; quite a number of other species
may create problems when they occur in abundance in picnicking areas, camp-
grounds, or other recreational areas.

Family Vespidae

Hornets, Yellowjackets, and Potter Wasps

The family Vespidae contains the well-known stinging wasps, hornets, and
yellowjackets. More than 360 species have been recorded in America north of
Mexico, about one-third of which occur in the Eastern United States. Considered as
a whole, these insects are not very important as forest pests since very few of them
are capable of inflicting serious injury to trees. Adults of the family differ consider-
ably in size and appearance, but all are distinguished by the very long discoidal cell
in the forewings, and by their common habit of folding their wings longitudinally
while at rest. Some of the more common species live in nests or colonies containing
a few to several hundreds or thousands of individuals. A synoptic catalog of species
occurring in America north of Mexico is available (697).

Social species construct nests out of papery material consisting of wood or
foliage chewed up and elaborated by the insect. Hornet nests contain several to
many tiers of hexagonal cells enclosed in a papery envelope and are usually
attached to the limbs of bushes or trees or to the eaves of buildings. They are
roughly spherical and often quite large, measuring 75 mm or more in diameter.
Yellowjackets also construct nests of papery material, but these are usually placed
out of sight in the ground, in stumps, or under objects. An exit hole leads to the
outside. Wasp nests usually consist of a single horizontal comb or layer of cells
attached to a support by a slender stalk. They may be found in many places, such as
under the eaves of roofs, under porches, in open sheds, and on bushes, shrubs, and
trees. The adults of all members of the family feed commonly on nectar, ripe fruits,
sap, and honeydew. The larvae feed on other insects provided by the adults.

The European hornet, Vespa crabro germana Christ, an introduced species, has
been recorded from Quebec and Massachusetts to Alabama and North Dakota.
According to reports, hollow trees, hollow posts, sheds, barns, porches, and even
attics are preferred as nesting sites. These nests occasionally become very large,
reaching a length of 90 cm and a diameter of 50 cm. There are reports of damage to
trees and shrubs by the adults (/077). They may girdle small twigs and gnaw holes
in the bark of larger branches, possibly in search of nest-building materials and sap.
Injuries have been reported to lilac, birch, ash, and buckeye. The adults possess a
long stinger and a large poison sac. This species is capable of inflicting painful
stings, but does not appear to be as likely to attack when disturbed as is the
baldfaced hornet.

The baldfaced hornet, Vespula maculata (L.), a well-known member of the
family, is widely distributed in the United States and Canada. Adults are 12 to 19
mm long, largely black with white markings on the face and thorax, and with the
posterior third of the abdomen white. It commonly attaches its nests to the limbs of
trees or bushes, also occasionally to the walls and windows of houses. These nests
may reach a diameter of 38 cm and contain up to several hundred hornets each.
Baldfaced hornets attack at once when their nests are disturbed, and are capable of
inflicting extremely painful stings.

Vespula arenaria (F.), the yellowjacket wasp, and V. maculifrons (Buysson) are
two of the more common species of yellowjackets in the Eastern United States. The
former also occurs throughout the remainder of the United States, in Canada, and
Alaska north to the Arctic Circle. V. arenaria belongs to the group of aerial nesters

436

(subgenus Dolichovespula) and is not a scavenger. Nests (fig. 210) are typically
built in trees, low brush, or just above ground in vegetation, not beneath the
ground. Thus they would not be picnic pests unless their nest was built on or very
close to the picnic table.

Courtesy Conn. Agric. Exp. Stn.

Figure 210.—Nest of the yellowjacket wasp, Vespula
arenaria.

The eastern yellowjacket, V. maculifrons (subgenus Vespula), along with V.
squamosa (Drury), may be the two most troublesome species in the Eastern United
States (778). V. maculifrons also occurs in southeastern Canada. Adults of V.
maculifrons are black and yellow with black predominating. The legs are yellow
with the femora partly black. Nests are usually built in the ground, frequently in
yards, and may attain very high densities in recreational areas. This species also
inflicts severe stings and it often attacks in force when its nest is endangered. It is a
common species around picnic tables where it is often a nuisance. The adventive
species, V. germanica (L.), nests above ground and commonly builds its nest in
attics and walls of homes.

Vespula squamosa is the most common yellowjacket from central Georgia into
Florida and probably along the Gulf Coast. This species was recently found to be a
social parasite of V. maculifrons (779) and the extent of its scavenging habits is still
in question. A discussion of various methods of control of yellowjackets and their
effectiveness has been published (778).

The genus Polistes contains the familiar wasps whose nests are so frequently
encountered hanging under the eaves of buildings and in a variety of other places.
They differ in appearance from the hornets and yellowjackets in having long and
spindle-shaped abdomens. The nest consists of a single comb suspended by a
peduncle, and it is not enclosed in an envelope. It is usually rather small, although

437

some may be 75 mm or more in diameter. The adults of a number of species collect
lepidopterous larvae as food for their young, and where abundant, are apparently
capable of exercising a considerable degree of control of their hosts. Like hornets
and yellowjackets, these insects are vicious stingers, and their nests should be
approached with caution.
Family Pompilidae
Spider Wasps

Members of this family are predacious or parasitic on various species of spiders.
The adults are often seen visiting flowers and are noted for their extreme activity
and ability to run. Their nests are usually found in the ground, but those consisting
of mud cells are constructed under logs or stones and in other protected places such
as holes in wood.
Superfamily Sphecoidea

All members of this superfamily are predacious on other insects. The majority of
Species nest in burrows in the soil; the remainder build nests of clay, mud, or sand,
or in the stems of plants or various kinds of cavities in which they store paralyzed
prey for their progeny. Host preferences are varied and include spiders and most of
the more common orders of insects. Adults generally can be recognized by the
structure of the pronotum that does not extend back to the tegulae, by their
unjointed trochanters, by the absence of dilation in the hind tarsus, and by the
simple pubescence of the head and thorax.
Family Ampulicidae
Ampulicids

This family is represented in North America by only two genera and three
species. As far as known, they nest in twigs, under bark, and under litter on the

ground. Their prey consists of immature cockroaches.
Family Sphecidae
Cicada Killers, Mud Daubers, and Sand Wasps

This family is represented by 111 genera and more than 1,100 species in the
United States and Canada. The majority are solitary nest-building wasps that
provision their nests with other insects or spiders. Many species nest in the ground;
some construct nests or mud and attach them to the ceilings or walls of buildings, or
to the lower surfaces of other objects; others construct their nests in the stems of
plants. Adults are distinguished by the collarlike pronotum that has a straight hind
margin, the cylindrical rather than flattened hind basitarsus, and absence of
branched body hair. In a few species, such as the familiar daubers, the abdomen has
a long, cylindrical petiole.

The cicada killer, Sphecius speciosus (Drury), which provisions its nest with
adult cicadas, is one of the more conspicuous members of the family. The adult
female is a large, black or yellow wasp up to 37 mm long. There are prominent
black and yellow bands or spots on the abdomen. Adults are usually present from
midsummer to early fall and they burrow into the ground for nesting purposes. The
female paralyzes a cicada by stinging it and then flies it to her nest, stores it in a
cell, and deposits an egg between its legs. When the larva hatches it feeds on the
cicada. There is one generation per year.

Cerceris fumipennis Say provisions its nest with a wide variety of wood-boring
buprestids. Species of the genus Psen prey on various membracids, cercopids, and
leafhoppers; Crossocerus annulipes (Lepeletier & Brulle) preys on various species
of tree-infesting cicadellids (28/): Stictia carolina (F.) is frequently seen hovering
over livestock in search of horse flies.

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Superfamily Apoidea
Bees

It has been estimated that more than 3,500 species of bees in 7 families occur in
North America. Some Apidae such as the honey bee and bumble bee, are so
common as to require no description. Fortunately, none of these insects is particu-
larly injurious to forest or shade trees. On the contrary, as a group they are highly
beneficial because of the prominent role so many of them play in the pollination of
flowers. While the majority of species provision their nests with pollen and honey, a
few lay their eggs in the cells of other bees where their young live as inquilines or
parasites and feed on food stored by their hosts. Members of the superfamily are
distinguished by the following characteristics: the pronotum does not extend back to
the tegulae, the trochanter is single-jointed, the hind basitarsi are dilated or
thickened, and the head and thorax are covered with branched hairs.
Family Megachilidae
Leafcutting and Mason Bees

Leafcutting and mason bees are small to medium size and black, blue, brown,
gray, metallic, or purplish. Some are marked with yellow. They build their nests in
rotten wood, in holes in solid wood, and in the hollow stems of plants, and they line
the walls of their cavities with mud, resin, or circular or oval-shaped pieces of
leaves cut from various species of plants. A few species have been recorded
damaging shade trees and ornamentals in various parts of the country; otherwise,
the group is noninjurious.
Family Anthophoridae
Digger, Carpenter, and Cuckoo Bees

The family Anthophoridae is represented by more than a thousand species in the
United States and Canada. It has been divided into three subfamilies—An-
thophorinae, Xylocopinae, and Nomadinae.

The subfamily Anthophorinae contains the so-called mining or digger bees and
the cuckoo bees. A number of species collect pollen and nest in the ground. Certain
others, such as the cuckoo bees or Nomadinae, are parasitic in the nests of other
bees. The adults are usually wasplike in appearance. Members of the subfamily
Xylocopinae construct their nests in wood or plant stems and are commonly known
as carpenter bees. The genus Xylocopa contains the large carpenter bees. The adults
look like bumble bees but differ in having the dorsum of the abdomen largely bare.
They nest in solid wood.

Considered as a whole, members of the family are far more beneficial than
harmful. Many species are highly efficient plant pollinators, and the only destruc-
tive members of the family are the carpenter bees, several species of which attack
and damage wood in use.

The carpenter bee, X. virginica (L.), is an important pest because of its habit of
tunneling into the solid wood of beams, rafters, telephone poles, or structural
timbers. This activity may lead to structural damage, especially when the same
piece of timber is attacked for several years. Dead but sound wood of baldcypress,
thuja, eastern white and hard pines, and redwood seems to be preferred. The adult,
which is about 25 mm long, bores a hole about 9 mm in diameter straight into the
wood for a short distance, then makes a right-angle turn and follows the grain of the
wood for a distance of 15 to 20 cm. Sometimes two bees use a common entrance
hole. When this happens the tunnel is extended in opposite directions from the
entrance hole. Eggs are deposited singly in separate chambers in the tunnel, each of
which is largely provisioned with pollen. Each larva lives and feeds in its chamber
until mature. Adults feed on pollen during daylight hours. Females spend the night

439

in their burrows, and males, under boards or in other protected places. Winter is
spent as young adults in their tunnels. There is one generation per year.

Living bees in tunnels can be killed by running a stiff wire all the way to the end
of the tunnel.

One other species of carpenter bee, X. micans Lepeletier, also occurs in the
Eastern United States from North Carolina to Florida and westward along the Gulf
Coast.

Order Diptera—Flies

The Diptera constitutes one of the largest orders of insects. The majority of
species differ from other insects to which the term “fly” is applied, such as the
sawflies, stoneflies, and the dragonflies, by the fact that they posses only one pair
of wings, the forewings. Their hindwings are reduced to small knobbed structures
called halteres. The majority of species are soft-bodied and small to minute in size.
The larvae, commonly known as maggots, are legless, and vary in form from
slender and elongate to stout and cylindrical. The pupae may be free, loosely
enclosed, or held immobile in the last larval skin. In the latter case it is known as a
puparium.

Many species of Diptera are destructive pests and are of great economic impor
tance. Bloodsucking forms such as the mosquitoes, black flies, punkies, and horse
flies are serious pests of animals, including humans. Some of these as well as some
of the scavenger species, such as the house fly, are important vectors of the
causative organisms of such serious diseases as malaria, yellow fever, filariasis,
dengue, sleeping sickness, and dysentery. A number of others are important pests
of agricultural crops, a few are pests of trees and ornamental plants, and many are
important parasites or predators of injurious species of insects.

Several native and introduced species are highly effective parasites of some of
our most destructive tree-defoliating insects, especially of the order Lepidoptera. A
few aquatic species, some of which are nuisance pests in the adult stage, are
economically important as fish food in the larval stage.

Several comprehensive treatments of the order Diptera have been published (/65,

273, 11975, 1192 413513).
Families Oestridae and Cuterebridae
Bot and Warble Flies

Bot and warble flies are endoparasites of animals, and several are serious
economic pests. They are best known as enemies of domestic animals, but many
species also attack various wild animals. The adults are medium- to large-size flies
resembling bees. Members of the genus Cephenemyia intest the nasopharyngeal
region of deer, moose, elk, caribou, and reindeer. The northern cattle grub,
Hypoderma bovis (L.), and the common cattle grub, H. /ineatum (Villers), are
subcutaneous parasites of cattle, causing serious damage to the tissue and hides.
The genus Cuterebra contains a large number of species that are subcutaneous

parasites of various rodents and lagomorphs.
Family Culicidae
Mosquitoes

Mosquitoes are important pests of people and other animals. Not only are their
bites extremely annoying, but they transmit many of the most serious diseases of
humans and other animals, such as malaria, dengue, yellow fever, and encephalitis
(431). As nuisances, they often seriously interfere with public enjoyment of parks,
vacation sites, and other recreational areas. Woods workers, fishers, hunters,

440

vacationers, and hikers are also pestered by them. The health of domestic animals
and wildlife suffers with serious losses of weight and disease.

Twelve genera and 148 species of true mosquitoes occur in the United States and
Canada (//73). The larvae and pupae live in standing or slowly moving bodies of
water that range in size from small accumulations held by plants to vast salt
marshes. Many of these are included in a review of mosquitoes in the Southeast
(666).

Family Ceratopogonidae
Biting Midges

This family contains a number of species of very tiny flies, usually | to 4 mm in
length, that feed on the blood of humans and other animals. They are often
abundant in the vicinity of fresh water inlets along the seashore or near fresh water
streams, ponds, and pools. Woods workers, hikers, hunters, fishers, picnickers,
and others frequenting these areas often find their presence almost intolerable
because of their very burning and painful bites. Populations are heaviest during late
summer. At this time, these flies bite chiefly in the evening and very early in the
morning. Some 27 genera and 348 species occur in America north of Mexico
(1341).

Family Chironomidae
Midges

Midges are small, slender flies, rarely over 10 mm in length (//77). They
resemble mosquitoes, but do not have fringes of scalelike hairs on the wings. They
differ also by the discontinuation of the costal vein at the end of the third vein.
Midges are frequently seen in great swarms, dancing in the air, usually in the
evening. The larvae of most species are found in the water of streams, usually
attached to the surface of stones, sticks, and other objects. Many are red and are
commonly known as bloodworms. Midge larvae are fed upon by many freshwater
fishes and other aquatic animals.

Family Simuliidae
Black Flies

Black flies occur in nearly all parts of the United States and Canada (//74). In
different parts of their range they have various common names such as buffalo gnats
and turkey gnats, as well as black flies. The females feed on the blood of humans
and other animals and their bites cause swelling, itching, and sometimes bleeding.
Their habit of hovering about the face and getting into the eyes, ears, and nostrils
makes them a nuisance. When they appear in large numbers, birds and animals may
be literally smothered by flies drawn into their air passages. In the woods and
mountains of the Northern United States and Canada, they are often so abundant in
the spring that they are almost unbearable. Black flies have also been incriminated
in the transmission of several diseases of wild and domestic birds.

Black flies usually lay their eggs on grass and other materials just below the
surface of water in swiftly flowing streams. The larval stage is spent in the water,
usually attached to sticks, stones, or living vegetation. Adults are not strong fliers
and usually are encountered in large numbers not too far from streams. Sometimes,
however, the wind blows them a considerable distance.

Family Bibionidae
March Flies

March flies are slender-bodied, stout-legged, rather hairy, and have short, many-
jointed antennae. They are usually dark, but may be marked with red or yellow.
Full-grown larvae are distinguished by a fully developed false segment, armed with
spines behind the head. The common name, March flies, has been applied to the

441

group because of the frequent appearance of the species, Bibio albipennis Say, in
large numbers in March. Because of its huge numbers, Plecia nearctica Hardy, the
lovebug, is a nuisance to drivers in the Gulf States. The larvae usually feed on

decaying vegetable matter, but a few feed on the roots of grass and other plants.
Family Sciaridae
Darkwinged Fungus Gnats

Darkwinged fungus gnats are moderately small, slender, delicate, mosquitolike
insects. The antennae have 12 to 17 segments, the wings are large, the coxae are
prominent and elongated, and the tibiae are armed with spurs. The adults are often
common in dark, humid habitats in wooded areas. Many species breed on mush-
rooms or on fungi growing on trees and logs. A number of others are predacious and
are found under bark or in the galleries of wood-boring insects.

Family Cecidomyiidae
Gall Midges

There are more than 1,000 described species of Nearctic cecidomyiids, of which
several hundred are known to attack various trees and shrubs (430). The oaks are
especially favored as hosts but several other species of trees are also infested, such
as willow, elm, maple, walnut, hickory, and the pines. The larvae of about two-
thirds of the species cause the formation of galls or pronounced swellings by their
feeding (400). Others feed in such places as patches of pitch exuding from injured
limbs, on fungi, in the excrement of insects, birds, and mammals, and in galls
produced by other insects. A few are predators on small insects such as aphids,
mites, psyllids, and scales.

Gall midges are small, mosquitolike flies with relatively long antennae and legs.
Young larvae are slender, somewhat flattened, and taper toward each end. Full-
grown larvae are distinguished by the presence of a sclerotized structure, commonly
known as a spatula, on the underside of the front end.

The balsam gall midge, Paradiplosis tumifex Gagné, produces swollen, oval
galls about 3 mm in diameter near the bases of needles of balsam fir (95/7). It most
likely also occurs on Fraser fir and wherever these hosts are found in North
America. Damage in Christmas tree plantations may be severe because galled
needles drop from the twigs in October.

Dasineura balsamicola (Lintner) originally was believed to be the cause of the
gall formed by P. tumifex (482). However, the role played by D. balsamicola is that
of inquiline, and this midge kills the gall-former P. tumifex where they occur
together. Closer study of other species of Dasineura may reveal similar rela-
tionships. D. gleditchiae (Osten Sacken), the honeylocust pod gall midge, pro-
duces oblong, podlike galls on the new leaflets of honeylocust seedlings in nurser-
ies in the Midwest, New England, Oregon, California, and Pennsylvania. D.
pseudacaciae (Fitch) attacks the young leaves of black locust and causes them to
fold, and D. communis Felt, the gouty vein midge, produces greenish or reddish
pouch galls on the veins of the leaves of red and sugar maples.

The boxwood leafminer, Monarthropalpus buxi (Laboulbene), an introduced
species, occurs from Rhode Island south to Delaware and Maryland. Its hosts are
different varieties of boxwood, the tree boxwood and the glossy leafed boxwood, in
particular. Adults are orange-yellow and about 2.5 mm long. The larvae feed within
the tissues of the leaf, causing blisterlike blotches up to 2.5 mm long by fall.
Heavily infested leaves turn gray or yellowish brown and often drop prematurely.

Contarinia juniperina Felt, the juniper midge, is a pest of redcedar and other
junipers in the Midwest and California. Adults are very small, only about 1.5 mm
long, and have bright-red abdomens. Eggs are laid on the needles of new growth

442

near the tips of twigs. Larvae bore into the twigs at the bases of the needles and kill
the portion beyond the entrance hole (533). Heavily infested trees turn brown in the
fall and most of the infested twigs break off during the winter. The hymenopterous
parasite, Platygaster pini Fouts, is reported to have exerted a high degree of control
of an outbreak in Missouri during the late thirties.

Contarinia negundifolia Felt, the boxelder gall midge, produces fleshy galls on
the leaves of boxelder. It has been recorded from Virginia, the Lake States, and the
Prairie Provinces of Canada (/326). Other fairly commen species of Contarinia
include: C. virginianiae (Felt), the chokecherry midge, which feeds on the fruit of
common chokecherry, causing it to become swollen and deformed; C. ce-
rasiserotinae (Osten Sacken), which produces bright-red or yellow irregular bud or
terminal galls on black cherry; C. canadensis Felt, which produces galls on the
leaves of ash; C. catalpae (Comstock), whose larvae feed on the young leaves and
seed pods of catalpa; and C. verrucicola Osten Sacken, the linden wart gall, which
produces wartlike galls on the leaves of basswood.

Asphonaylia ilicicola Foote larvae feed in the berries of American holly, and the
infested berries remain green all winter. It has been recorded from New Jersey,
Maryland, Virginia, and West Virginia (569). The related species, A. azaleae Felt,
produces the so-called pinkster bud gall on rhododendron.

Thecodiplosis piniresinosae Kearby, the red pine needle midge, has been
recorded causing the formation of gall-like structures in the bases of needle fascicles
of red pine in Wisconsin (657). Damaged needles turn brown in the fall and drop
during the winter. The laterals and terminals of heavily infested trees may be killed.
Heavy infestations appear to be confined to slow-growing trees.

Cecidomyia resinicola (Osten Sacken), the gouty pitch midge, causes swellings
and malformations on the twigs of Virginia and pitch pines in eastern North
America. It has a broad host range among hard pines and is widely distributed.
Dead needles, dead or dying twigs, and distorted, twisted terminals are evidence of
its attack (355). The generally gregarious larvae of C. resinicola (fig. 211) are
found in small patches of fluid resin exuding from wounds caused by their feeding.
C. piniinopis Osten Sacken is another widespread resin midge with many host
species among the hard pines (460). Usually an individual larva occupies a drop of
pitch at the base of a needle fascicle. When full grown the larva moves away from
this site and spins a white cocoon in which it pupates. There may be several
generations annually. C. poculum Osten Sacken produces clusters of pale or red
saucerlike galls that are attached by slender stalks to the undersurfaces of oak
leaves. C. pellex Osten Sacken produces reddish-brown bullet galls on ash. C.
ocellaris Osten Sacken, the ocellate gall midge, causes the yellow galls margined
with red that are often seen on the upper surface of red maple leaves.

Macrodiplosis foliora Russell & Hooker produces marginal fold galls on the
leaves of black, red, and pin oaks.

Sequoiomyia cupressi (Schweinitz) feeds in the seeds and causes a leaf gall on
baldcypress. Taxodiomyia cupressiananassa (Osten Sacken) also causes leaf galls
on baldcypress.

Apagodiplosis papyriferae Gagné larvae feed in the buds of paper birch in the
Lake States, causing the formation of galls (fig. 212). The galls surround the basal
portions or all of one or two leaf petioles and either a bud or a part of the stem near
the bases of the petioles (/329). Damaged leaves fall prematurely and buds and
branches are killed. Trees from 0.7 to 10 m tall are attacked. The younger ones are
often badly deformed.

443

USDA, ARS, SEL F-519915
Figure 211.—Resin masses caused by Figure 212.—Damage to paper birch by
feeding activity of larvae of the cecidomyid, Apagodiplosis
Cecidomyia resinicola. papyriferae.

The willow beaked-gall midge, Mayetiola rigidae (Osten Sacken), produces
apical, beaked galls on the lower branches of many species of willow (fig. 213). In
Michigan, eggs are laid singly on or near the buds of the host. Newly hatched larvae
penetrate the bud and a gall begins to develop by the end of the first instar and
continues to enlarge until fall. Winter is spent as a larva inside the gall, and
pupation occurs in the spring. The gall deforms the stem and occasionally a galled
branch dies or breaks off (/328).

F-519914
Figure 213.—Gall produced on willow by
the willow beaked-gall midge,
Mayetiola rigidae.

elt

Many other species of gall midges have been recorded attacking various species
of eastern trees. A few of the more common ones are: Parallelodiplosis florida
(Felt)—produces elongate, pocketlike swellings on the veins and midribs of scrub
and pin oak leaves; Obolodiplosis robiniae (Haldeman)—causes the margins of
black locust leaves to fold; Caryomyia holotricha (Osten Sacken), C. sanguinolenta
(Osten Sacken), and C. tubicola (Osten Sacken)—produce galls on hickory; Poly-
stepha pilulae (Beutenmuller)—produces subglobose or globose, irregular,
wrinkled, reddish galls on red oak leaves; Janetiella coloradensis Felt—causes
swellings at the bases of Virginia pine needles; Prodiplosis morrisi Gagné—feeds
on the young leaves of cottonwood in the South, causing them to turn black, unfold
improperly, and frequently drop off; terminal growth is sometimes severely stunted.
The dogwood clubgall midge, Resseliella clavula (Beutenmiller), produces club-
like swellings on small twigs of dogwood, which sometimes kills up to 8 cm of the
damaged twigs. Semudobia spp. feed on and destroy the seeds of birch.

Aphidoletes thompsoni Mohn, a European predator of the balsam woolly adelgid,
has been released in aphid-infested stands of balsam and Fraser firs in the Northeast
and the Southern Appalachians, but establishment is doubtful in the Southern
Appalachians.

Family Xylophagidae
Ichneumonlike Flies

These flies, as their common names suggests, often resemble members of the
hymenopterous family, Ichneumonidae. The adults are usually observed feeding on
sap, the nectar of flowers, or other liquid matter in forested or wooded areas. The
larvae appear to be either predators or scavengers in rich soil, in decaying vegetable
matter, under the bark of trees, or in decaying logs. Xylophagus lugens Loew larvae
may occur in large numbers under the bark of elm in association with the elm borer.

The larvae of X. abdominalis Loew feed on beetle larvae under the bark of pine.
Family Stratiomyidae
Soldier Flies

Soldier flies are brightly colored, moderately large, nearly bare, and thinly
pilose. Many species are wasplike in appearance but the majority are broad and
greatly flattened, and their wings lie parallel upon each other while at rest. These
flies occur chiefly in wooded or forested areas or in meadows near water. Many are
attracted to flowers. The larvae are usually terrestrial but sometimes aquatic and act
mostly as scavengers. Zabrachia polita Coquillett has been reared from decaying

pine logs and from beetle-infested elm logs.
Family Tabanidae
Deer and Horse Flies

Many members of this family are important bloodsucking pests of livestock and
wild animals. Some species also attack people and can be extremely annoying.
Their bites are sharp and painful, and many are capable of removing considerable
quantities of blood from their hosts, especially when they attack in force. They may
also transmit such animal diseases as anthrax, anaplasmosis, surra, swamp fever,
and tularemia.

Most of the large horse flies usually seen belong to the genus TZabanus. These
flies breed mostly in marshes, swamps, bogs, and ponds. The largest and best
known species is the black horse fly, 7. atratus F. The adult is up to 25 mm long
and its bite can result in the loss of a considerable quantity of blood. Other
important species are: the striped horse fly, 7. Jineola (F.), T. abactor Phillip, T.
quinquevittatus Wiedemann, T. sulcifrons Macquart, and 7. nigrovittatus Mac-
quart.

445

Deer flies belong to the genus Chrysops and are smaller than horse flies. They
are active during the hottest weather and are often quite abundant during rainy
spells. More than 60 species occur in the Eastern United States. Diachlorus
ferrugatus (F.) is acommon and notorious pest in swampy areas from New Jersey to
Florida and Louisiana, especially in the Carolinas.

Family Rhagionidae
Snipe Flies

Snipe flies are commonly found in the woods, especially in moist areas. The
majority are brownish or gray; others are black with spots or stripes of white,
yellow, or green. Both the adults and larvae feed on other insects. Adults of the
genus Symphoromyia fly persistently about the head and are very annoying because
their bites are painful.

Family Asilidae
Robber Flies

Robber flies are mostly large to very large (fig. 214). Some have long, tapering
abdomens; others are stout-bodied and resemble bumble bees. All are rather hairy.
They are predacious on other insects and occasionally on their own kind. The larvae
of Jolmerus notatus (Wiedemann), Leptogaster flavipes Loew, Laphria index
McAtee, L. flavicollis Say, and L. thoracica F. have been collected from decaying
wood infested with cerambycid and other coleopterous larvae. Larvae of certain
other species feed on grasshopper eggs and white grubs.

Courtesy Conn. Agric. Exp. Stn.
Figure 214.—Adult of a robber fly, Asi/us sericeus.

Family Bombyliidae
Bee Flies

These flies are moderately large, densely hairy, and look like bees. The adults
are nectar and pollen feeders. The larvae of certain species are parasitic on the
larvae of other insects, especially of the orders Lepidoptera, Hymenoptera, and

Coleoptera. Others feed on the egg pods of grasshoppers.
Family Empididae
Dance Flies
Members of this family are commonly known as dance flies because of their habit

of flying in swarms—up and down, up and down—1in the woods, along streams, or

446

on the shores of ponds and lakes. The flies range in length from | to about 15 mm
and probably are all predatory, mostly on small Diptera. The larvae live in damp
earth, in decaying wood or other vegetation, under the bark of trees, or in the water.
Tachydromia sp. has been reared from eastern white pine leaders infested with the
white pine weevil.

Family Dolichopodidae
Longlegged Flies

Members of this family are very small, longlegged flies, rarely more than 10 mm
long. Adults are predacious on other insects and are found on the foliage or trunks
of trees, or on damp earth, usually in swamps or along lightly shaded streams. Most
of the larvae appear to be aquatic, but those of a few species occur under the bark of
trees where they feed on other insects. Dolichopus vittatus Loew has been reared
from beetle-infested hickory, and Medetera sp. from beetle-infested larch and pine.

Family Pipunculidae
Bigheaded Flies

Pipunculids are small, dark flies with large heads composed mostly of large,
approximated eyes. The larvae are small, elliptical, thick, depressed, naked, and
narrowed at each end. They are parasitic on various families of Homoptera,
especially the Cicadellidae.

Family Syrphidae
Flower Flies

This is one of the largest families in the order Diptera, and it is almost entirely
beneficial (840, 1342). The adults are strongly attracted to flowers and play
important roles in the cross-pollination of plants and trees. The larvae of many
species feed on and aid in control of many destructive insects such as aphids,
scales, psyllids, spittlebugs, mealybugs, and lepidopterous larvae. A few species
are plant feeders but they are seldom injurious.

Adults are usually brightly colored and are frequently striped, spotted, or banded
with yellow. Some resemble wasps; others look like small bumble bees. The maies
have the peculiar habit of hovering almost completely motionless in the air and then
darting swiftly to one side when disturbed. The maggots of insect-feeding species
are Sluglike. The body tapers toward the front end and the body contents are visible
through the integument.

Family Conopidae
Thickheaded Flies

Conopid flies are thinly pilose or nearly bare, elongate, and of moderate size.
The head is broader than the thorax, and the abdomen is elongated and constricted.
The antennae are three-segmented, the third segment bearing a dorsal arista. Adults
fly slowly and are usually seen around flowers. The larvae are solitary internal
parasites, mainly of Hymenoptera. One species parasitizes grasshoppers and
crickets.

Family Tephritidae
Fruit Flies

Fruit flies are fairly small and usually have spotted or banded wings. The larvae
are usually pale yellowish and taper slightly toward the front. Eggs of most species
are deposited in healthy, living tissue and larvae feed in various parts of plants.
Certain species produce root and stem galls and a few are leafminers. Others
develop in fleshy fruits or in the seeds or ovaries of flowers. Many species are
highly destructive of fleshy fruits and vegetables throughout the world. A few of the
species occurring in the woodlands or forests of the Eastern United States are
discussed below.

447

The apple maggot, Rhagoletis pomonella (Walsh), an important pest of apple,
also breeds in the fruit of hawthorn, common chokecherry, plum, and dogwood.
The adult is dark colored and a little smaller than the house fly. Each wing is
crossed by four dark bars that merge together. Three or four white bands run across
the dorsum of the abdomen. The cherry fruit fly, R. cingulata (Loew), breeds in
the fruit of cherry. Adults are black except for yellow margins on the thorax, two
white crossbands on the abdomen, and a dark band on each wing. They are smaller
than house flies. The black cherry fruit fly, R. fausta (Osten Sacken), breeds in
cherry throughout the same area as does the cherry fruit fly. Adults resemble those
of the cherry fruit fly except for the abdomen, which is entirely black. The walnut
husk fly, R. completa Cresson, breeds in the husks of black walnut in the Central
States. Feeding by the larvae produces a slimy condition that causes the husks to
turn black, stick to and stain the shell. Nuts with damaged shells cannot be sold
even though the contents are sound. The wings of the adult are transparent with
dark crossbars. Toxotrypana curvicauda Gerstaker breeds in papaya in Florida and
Texas.

Family Chamaemyiidae
Aphid Flies

The larvae of a number of species in several genera of these small, yrayish flies
are predators of aphids, scale insects, and mealybugs. One species, Leucopis
obscura Haliday, a native of Europe, was introduced into Canada against the
balsam woolly adelgid in the early thirties. It quickly became established and,
following additional colonizations, spread over most of the infested areas in eastern
Canada and into northern New England. Since 1954, colonies have been released in
adelgid-infested stands in Vermont, New Hampshire, New York, North Carolina,
and the Pacific Northwest. It is usually found on heavily infested trees only, where
it feeds mostly on adults that have laid many eggs.

Cremifania nigrocellulata Czerny, another European species, also imported into
eastern Canada and the Pacific Northwest against the balsam woolly adelgid, is now
established. A fairly large number have developed on stem-infested trees, but their
spread has been very slow.

Family Lonchaeidae
Lonchaeids

Adults of this family are shiny black and about 5 to 6 mm long. The larvae are
very small and are covered by minute spines. These flies act mostly as scavengers
or as predators on other insects. Lonchaea polita Say has been reared from bark-
beetle infested wood. L. corticis Taylor has been recorded as an important parasite
of the white pine weevil (762).

Family Drosophilidae
Vinegar Flies

These are the flies that are so often seen around spoiled fruit, slime fluxes, and
fungi. Usually yellowish except for black markings on the abdomen, adults seldom
exceed 5 mm in length. Few if any species are of importance as enemies of forest,
shade, or ornamental trees. There is a possibility, however, that certain species are
involved in the transmission of the oak wilt fungus, Ceratocystis fagacearum
(Bretz) Hunt (507). Members of the genus Drosophila are very important in the

field of genetics research.
Family Chloropidae
Chloropid Flies

These flies are small to very small with numerous clothing hairs and some
bristles. Some species are brightly colored with yellow and black. The larvae of

448

certain species parasitize or prey on other insects; others have been found feeding in
the seeds and cones of conifers. Adults of the genus Hippelates are attracted to the
eyes of humans and of other animals, and are very annoying. Certain species are
also reported to transmit yaws and pinkeye. Gaurax apicalis Malloch, Oscinella
coxendix (Fitch), and Hippelates sp. have been reared from the leaders of eastern
white pine infested by the white pine weevil (762).

Family Agromyzidae

Leafminer Flies

These flies are very small and light or dark colored (449). The body covering
ranges from sparse bristles to dense hairs. The larvae are plant feeders; some mine
the cambium, but the majority mine the leaves.

Larvae of the genus Phytobia feed in the cambium of living trees, making long,
thin, gradually widening mines (fig. 215). These mines sometimes originate in the
top of the tree and extend all the way to the base and into the roots. They cause
defects known as pitch-ray flecks. Heavily infested logs may be rendered unfit for
some uses.

F-506078, 506069

Figure 215.—Cambium miner work: left, streaklike,
longitudinal mines on trunk and roots of infested trees;
right, pitch-ray flecks in the wood.

Phytobia setosa (Loew), the red maple cambium borer, attacks red and sugar
maples (523). Adults are small and dark colored. The larvae are opaque white and
about 16 mm long. P. pruinosa (Coquillett) infests cherry, maple, and sweet and
river birches. Adults are about 3 to 4 mm long; the larvae are up to 30 mm long.
This species apparently lays its eggs in the forks of branches near the tops of trees
and the larvae tunnel all the way down into the roots. P. amelanchieris (Greene), the
amelanchier twig borer, attacks serviceberry, and P. pruni (Grossenbacher) at-
tacks cherry.

The native holly leafminer, Phyfomyza ilicicola Loew, 1s a serious pest of
American holly in the Eastern United States. The adult is a small, grayish-black fly
about 2.5 mm long. The female punctures leaves with her ovipostor and feeds on
the juices exuding from the wounds. She also deposits eggs in the undersurfaces of
leaves in punctures made near the midrib. The larvae mine the tissues between the
leaf surfaces. The mine is hairlike at first but gradually widens as the larva
continues to grow. Eventually it becomes blotchlike (fig. 216). Heavily infested
leaves become unsightly and usually drop prematurely. Leaves damaged by feeding
punctures become roughened, twisted, and stunted. The winter is spent as second

449

or third instars or as pupae in the mine (705). Pupation occurs from early March to
early April and the adults emerge from mid-May to late June. There is one
generation per year.

Courtesy Conn. Agric. Exp. Stn.

Figure 216.—Injury by the native holly leafminer,
Phytomyza ilicicola: left, undamaged leaves; right,
mined leaves.

Several other members of the family also mine the leaves or produce galls on
their hosts. Japanagromyza viridula (Coquillett) produces blotch mines in the
leaves of red oak; Hexomyza schineri (Giraud) causes the formation of slight
swellings on the smaller twigs of poplar; H. tiliae (Couden) produces swellings
about | cm long on the twigs of basswood; Nemorimyza posticata (Meigen)
produces blotch mines in the leaves of sweetgum; and Trilobomyza_ pleuralis

(Malloch) produces blotch mines on catalpa.
Family Anthomyiidae
Anthomyiids

Members of this family are quite similar in appearance to those of the family
Muscidae. The maggots vary in habits; some feed on the roots of plants, some are
scavengers, and others are parasites of other insects. The seedcorn maggot,
Hylemya platura (Meigen), damaged redcedar seedlings in a forest nursery in
Tennessee (/29/). In this case, the larvae chewed through the bark of the main stem
just below the ground line and fed on the roots. Large numbers of seedlings were

killed.
Family Muscidae
House Flies, Stable Flies, and Allies

Flies of this family vary in length from 2 to 12 mm. Some are yellowish or black,
but the majority are gray or brown. There are several economically important
species, such as the well-known house fly, Musca domestica L., and the stable fly,
Stomoxys calcitrans (L.). Tsetse flies, which transmit the organisms responsible for

450

sleeping sickness and other diseases, are native to Africa and do not occur in this
country. A number of species in the genus Muscina are parasites of various species

of Coleoptera.
Family Hippoboscidae
Louse Flies

Louse flies may be winged or wingless. Many look like lice. The body is flat and
leathery-looking, and the legs are short, strong, and broadly separated by the
sternum. The tarsi are short and armed with strong claws, and the abdomen is
saclike. All species feed on birds and mammals. The best known member of the
family is the sheep ked, Melophagus ovinus (L.), an important parasite of sheep.
Other important species are /costa americana (Leach)—parasitizes grouse, hawks,
and owls; Olfersia fumipennis (Sahlberg)—attacks the bald eagle; species of Or-
nithoica and Ornithomyia—teed on various small birds; and species of Lipop-

tena—parasitize deer.
Family Calliphoridae
Blow Flies

Blow flies have metallic blue, green, or yellow bodies and are usually about the
size of house flies. The arista of the antenna is plumose at the tip and the hindmost
posthumeral bristle is almost always longer than the presutural bristle. In the
maggots, the posterior spiracles are flush. A few species are economic pests, but
the majority serve a useful purpose in helping to rid the landscape of such
undesirable materials as dead animal bodies and animal excrement.

The screwworm, Cochliomyia hominivorax (Coquerel), a long-time pest of
livestock in the Southwest, was first recorded from the Southeastern States in 1933.
Since then, or until its eradication from the region through the mass release of
sexually sterile males flies (735), it caused tremendous losses to livestock. Big
game, such as deer, and smaller animals, such as raccoons, rabbits, and opossums,
have also been attacked and injured. The female deposits up to 300 eggs around
wounds; the maggots feed on the tissues, and produce foul-smelling wounds. This
attracts additional flies and compounds the infestation. Heavily infested animals
may die within a few weeks unless the maggots are killed and the wounds treated.

Other blow flies occasionally attacking living animals include: Phaenicia se-
ricata (Meigen), the greenbottle fly; the black blow fly, Phormia regina (Meigen);
and the secondary screwworm, C. macellaria (F.). They also oviposit on wounds
or sores. Larvae of the genus Protocalliphora are bloodsucking parasites of nes-
tlings.

Family Sarcophagidae
Flesh Flies

Larvae of this family feed on a wide variety of foods. A number of species are
scavengers, feeding on dead insects, dung, and other decaying materials. Some
species are parasitic in various insects. Adults are 6 to 12 mm long, the sides of the
face are hairy, the aristae of the antennae are feathery for about half their length,
there are three black stripes on the thorax, and the abdomen is checkered. The
larvae have their posterior spiracles located in a pit.

Sarcophaga aldrichi Parker, an important parasite of the forest tent caterpillar,
occurs in southern Canada and from New England to the Lake States and Southern
Appalachians. It significantly suppresses forest tent caterpillar outbreaks in the
Lake States. Living young are normally deposited on cocoons and the larvae feed as
scavengers on the prepupae or pupae inside the cocoons (576). Full-grown larvae
are about 13 mm long. The adults are strong, active fliers; during tent caterpillar
outbreaks, they may occur in enormous numbers, swarming over everything,

451

livestock and people included. During intervals between outbreaks, the species is
hard to find. In the Southern States, the related species, S$. houghi Aldrich,

parasitizes the forest tent caterpillar and elm spanworm.
Family Tachinidae
Tachina Flies

The family Tachinidae contains many of the most important species of insect
parasites. All species appear to be internal parasites of many kinds of insects,
especially Lepidoptera, but also various species of Coleoptera, Hemiptera, Orthop-
tera, and Hymenoptera (/049).

Tachinid flies look very much like house flies but many differ in having an
entirely bare arista; however, the arista on some species may be pubescent or as
plumose as that of house flies. The maggots are clothed with minute spinules, and
the posterior spiracles are flush with or raised from the adjacent area.

The majority of species are oviparous, but a few give birth to living young. Eggs
are deposited on the skin of the host, on leaves or other parts of plants on which
their hosts feed, or on the soil. Living young are deposited on or under the skin of
the hast. on leaves and other parts of plants frequented by their hosts, or on the
ground. The number of generations per year varies from | to 10, depending on
species and climate.

A large number of native species and a few introduced species parasitize various
native and introduced species of eastern forest insects. More than 125 species have
been recorded attacking lepidopterous larvae in this country (/062, 1063). A few of
the more important introduced species are discussed here.

Compsilura concinnata (Meigen) (fig. 217) was imported against the gypsy and
browntail moths early in this century. It is now widely distributed throughout most
of the Northeast and in southeastern Canada where it attacks at least 200 species of
Lepidoptera. Parasitization of the gypsy moth averages from 10 to 50 percent in

F-519564

Figure 217.—Adult and puparium of Compsilura
concinnata, a parasite of the gypsy moth and other
defoliators.

452

much of the infested area. Even higher percentages are recorded in browntail and
satin moth infestations (1/73, 269, 328, 1262).

Blepharipa pratensis (Meigen) (fig. 218), a parasite of the gypsy moth, was
imported and became established by 1911. In most of the areas where it occurs, it
parasitizes from 5 to 25 percent of the gypsy moth population. As far as known, it
confines its attack to the gypsy moth in this country, whereas in Europe it has
several lepidopterous hosts. Its life history and habits are discussed (/73).

Courtesy P. Godwin, Hamden, Conn.

Figure 218.—Adult of Blepharipa pratensis, a parasite of
the gypsy moth and other defoliators.

Parasetigena silvestris (Robineau-Desvoidy) and Exorista larvarum (L.) were
imported against the gypsy moth from 1924 to 1933. P. silvestris, a single-brooded
species, 1S probably the most important parasite of the nun moth in Europe. E.
larvarum, a multibrooded species, attacks about 45 different hosts in Europe. Both
species are now important parasites of the gypsy moth through the generally
infested area in the United States. EF. /arvarum is also a parasite of the satin moth
and browntail moth in this country.

Two species, Zownsendiellomyia nidicola (Townsend) and Carcelia laxifrons
Villeneuve, were imported against the browntail moth early in this century. 7.
nidicola quickly spread throughout the infested area. It has frequently parasitized
an average of 17 percent of overwintering browntail moth larvae. C. laxifrons also
occurs throughout the infested area. Unfortunately, it parasitizes a very low per-
centage of the population.

Chaetexorista javana Brauer & Bergenstamm, a native of Japan, was imported
against the oriental moth in 1929 and 1930. As early as 1933, parasitism in the
infested area around Boston, Mass., averaged 60 percent. Erynniopsis antennata
Rondani, a native of Europe, has been successfully introduced into California
against the European elm leaf beetle. Colonies have also been liberated against the
beetle along the Atlantic Seaboard but without success. Palexorista bohemica
(Mesnil), a native of Europe, was successfully introduced into Canada against the
European spruce sawfly during the thirties and forties. This is probably the same
species that was introduced into the United States around 1906 as P. inconspicua

453

(Meigen) against the gypsy moth, browntail moth, and redheaded pine sawfly
(328). It was recovered at gypsy moth colonization sites for several years thereafter,
but it apparently did not become permanently established. It probably occurs now
in northern Maine as a result of spread from New Brunswick and Quebec, where it
appears to be one of the most effective parasites introduced against the European
spruce sawfly.

454

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zappei. Conn. Agric. Exp. Stn.
Bull. 226, p. 179-182.

Common and Scientific Names of Host Plants

acacia
sweet, See: huisache
ailanthus
alder
andromeda
apple
apricot
arborvitae
See also: white-cedar, northern
aralia
ardisia
ash
black
European
green
pumpkin
red, See: ash, pumpkin
velvet
white
aspen, See: poplar
Australian-pine
See: casuarina, horsetail
azalea, See: rhododendron

baldcypress
bamboo
banak
banyan, See: fig, shortleaf
basswood
American
bayberry
beech
American
European
oriental
birch
European white
gray
paper
‘river
sweet
white, See: birch, paper
yellow

Acacia

Allanthus altissima
Alnus

Andromeda

Malus spp.

Prunus armeniaca
Thuja

Aralia

Ardisia

Fraxinus

Fraxinus nigra
Fraxinus excelsior
Fraxinus pennsylvanica
Fraxinus profunda

Fraxinus velutina
Fraxinus americana

Taxodium distichum
Bambusa spp.
Virola_ spp.

Tilia

Tilia americana
Myrica

Fagus

Fagus grandifolia
Fagus sylvatica
Fagus orientalis
Betula

Betula pendula
Betula populifolia
Betula papyrifera
Betula nigra
Betula lenta

Betula alleghaniensis

529

bittersweet
blackgum, See: tupelo

blue-beech, See: hornbeam, American

blueberry

boxelder

boxwood, European
Brazil nut tree
buckeye

bumelia

butternut

camellia
camphor-tree
cancer-root
caragana
casuarina, horsetail
catalpa
cedar

Atlas

deodar

of Lebanon

See also: juniper; arborvitae;

white-cedar, northern

cherry |

black

chokecherry

Japanese flowering

pin

sour

wild, See: cherry, black
chestnut

American

Asiatic, See: chestnut, Chinese &

Japanese

Chinese

Japanese

See also: chinquapin
chinaberry
chinquapin
chokeberry
coffeetree, Kentucky
cotoneaster
cottonwood

See also: poplar
crapemyrtle
creeper, Virginia
cryptomeria
cucumbertree
cypress

530

Celastrus

Vaccinium

Acer negundo
Buxus sempervirens
Bertholletia excelsa
Aesculus

Bumelia spp.
Juglans cinerea

Camellia japonica
Cinnamomum camphora
Conopholis americana
Caragana arborescens
Casuarina equisetifolia
Catalpa spp.

Cedrus

Cedrus atlantica
Cedrus deodara

Cedrus libani

Prunus

Prunus serotina
Prunus virginiana
Prunus serrulata
Prunus pensylvanica
Prunus cerasus

Castanea
Castanea dentata

Castanea mollissima
Castanea crenata

Melia azedarach
Castanopsis spp.

Pyrus arbutifolia
Gymnocladus dioicus
Cotoneaster pyracantha
Populus spp.

Lagerstroemia indica
Parthenocissus quinquefolia
Cryptomeria Japonica
Magnolia acuminata
Cupressus spp.

dogwood Cornus spp.

Douglas-fir Pseudotsuga menziesii
elder Sambucus
American Sambucus canadensis
elm Ulmus
American Ulmus americana
Camperdown Ulmus glabra camperdownt
cedar Ulmus crassifolia
English Ulmus campestris
European Uimus effusa
Japanese Ulmus japonica
rock Ulmus thomasii
Scotch Ulmus glabra
September Ulmus serotina
Siberian Ulmus pumila
slippery Ulmus rubra
euonymus Euonymus spp.
fig Ficus
shortleaf Ficus citrifolia
fir Abies
balsam Abies balsamea
Fraser Abies fraseri
Siberian Abies sibirica
subalpine Abies lasiocarpa
white Abies concolor
forsythia Forsythia
fremontia Fremontodendron
fringetree Chionanthus virginicus
gardenia Gardenia spp.
ginkgo Ginkgo biloba
gooseberry-tree Phyllanthus acidus
grape Vitis spp.
greenbrier Smilax laurifolia
hackberry Celtis occidentalis
netleaf Celtis reticulata
hawthorn Crataegus spp.
haze! Corylus
hemlock Tsuga
eastern Tsuga canadensis
hickory Carya
- pignut Carya glabra
holly Ilex
American (typical) Ilex opaca var. opaca
Chinese Ilex cornuta
Japanese Ilex crenata

531

honeylocust
hophornbeam
eastern
hops
hoptree
hornbeam, American
See also: hophornbeam

horsechestnut, See: buckeye

huckleberry
huisache

incense-cedar

Ironwood, See: hornbeam, American

Japan cedar, See: cryptomeria

juniper
Ashe
California
common
Irish
oneseed
western

larch

Dahurian

eastern

European

Japanese

Siberian

subalpine

western
laurel, See: mountain-laurel
lilac
linden. See: basswood
locust

black
luan/meranti

magnolia
Star
See also: cucumbertree
mahogany
mahonia
mangrove
maple
Japanese
mountain
Norway
planetree

532

Gleditsia triacanthos
Ostrya

Ostrva virginiana
Humulus americanus
Prelea trifoliata
Carpinus caroliniana

Gaylussacia spp.
Acacia farnesiana

Libocedrus decurrens

Juniperus

Juniperus ashei
Juniperus californica
Juniperus communis

Juniperus communis var. stricta

Juniperus monosperma
Juniperus occidentalis

Larix

Larix gmelini
Larix laricina
Larix decidua
Larix leptolepis
Larix sibirica
Larix lyallii
Larix occidentalis

Svringa vulgaris

Robinia spp.
Robinia pseudoacacia

Shorea spp.

Magnolia spp.
Magnolia stellata

Swietenia mahagoni
Mahonia spp.
Rhizophora mangle
Acer

Acer palmatum
Acer spicatum

Acer platanoides
Acer pseudoplatanus

red

silver

sugar

See also: boxelder
mesquite
mimosa
moss, Spanish
mountain-ash

American

European

hybrid

showy
mountain-laurel
mulberry

nettle

oak
black
blackjack
bur
Chapman
cherrybark
chestnut
laurel
live
northern pin
northern red
Nuttall
overcup

pin
post

runner, See: oak, sand post & sand

live
sand live
sand post
scarlet
scrub
southern red
swamp white
turkey
water
white
obeche
oleander
olive
orange, sour
Osage-orange

Acer rubrum
Acer saccharinum
Acer saccharum

Prosopis

Albizia julibrissin
Tillandsia usneoides
Sorbus

Sorbus americana
Sorbus aucuparia
Sorbaronia

Sorbus decora
Kalmia latifolia
Morus

Urtica
Quercus

Quercus velutina
Quercus marilandica

Quercus
Quercus

macrocarpa
chapmanii

Quercus falcata var. pagodifolia
Quercus prinus

Quercus
Quercus
Quercus
Quercus
Quercus
Quercus
Quercus
Quercus

Quercus
Quercus
Quercus
Quercus
Quercus
Quercus
Quercus
Quercus
Quercus

laurifolia
virginiana
ellipsoidalis
rubra
nuttallii
lyrata
palustris
stellata

virginiana var. geminata

stellata var. margaretta

coccinea

spp.

falcata

bicolor
laevis
nigra
alba

Triplochiton scleroxylon
Nerium oleander

Olea europaea

Citrus aurantium
Maclura pomifera

533

pachysandra
palm
Canary Island date

Pachysandra spp.
Palmae family
Phoenix canariensis

palmetto Sabal
cabbage Sabal palmetto
papaya Carica papaya
pawpaw Asimina tribola
peach Prunus persica
pear Pyrus communis
pecan Carya illinoensis
persea Persea spp.
persimmon Diospyros virginiana
phylox Phylox spp.
pieris Pieris spp.
pine Pinus
Austrian Pinus nigra
Caribbean Pinus caribaea
Chinese Pinus tabulaeformis
Corsican Pinus nigra poiretiana
eastern white Pinus strobus
European black, See: pine, Austrian
jack Pinus banksiana
Japanese white ; Pinus parviflora
Japanese black Pinus thunbergiana
Japanese red Pinus densiflora
loblolly Pinus taeda
lodgepole Pinus contorta var. latifolia
longleaf Pinus palustris
mugho, See: pine, Swiss mountain
pitch Pinus rigida
pond Pinus serotina
ponderosa Pinus ponderosa
red Pinus resinosa
sand Pinus clausa
Choctawhatchee Pinus clausa var. immuginata
Ocala Pinus clausa var. clausa
Scotch Pinus sylvestris
shortleaf Pinus echinata
Slash Pinus elliottii var. elliottii
Sonderegger Pinus sondereggeri
South Florida slash Pinus elliottii var. densa
spruce Pinus glabra
Swiss mountain Pinus mugo
Table Mountain Pinus pungens
Virginia Pinus virginiana

white, See: pine, eastern white
plane, London
planetree, See: sycamore
pondcypress

Platanus acerifolia

Taxodium distichum var. nutans

534

plum
American
poison-ivy
poison-sumac
poplar
balsam
bigtooth
Lombardy
quaking

See also: yellow-poplar

prickly ash
privet
pyracantha
See also: cotoneaster

quince

rattan-vine
redbud
redcedar, eastern

See also: juniper
redgum, See: sweetgum
rhododendron
royalpalm, Florida

sassafras
serviceberry

Siberian peashrub, See: caragana

silktree, See: mimosa
smilax, See: greenbrier
sourgum, See: tupelo
sourwood
spicebush
spirea
spruce

black

blue

Colorado blue, See: spruce, blue

Engelmann
Norway
red
Sitka
white
strawberry
sugarberry
sumac
staghorn
sweetfern
sweetgum

Prunus spp.

Prunus americana
Toxicodendron radicans
Toxicodendron vernix
Populus

Populus balsamifera
Populus grandidentata
Populus nigra var. italica
Populus tremuloides

Zanthoxylum
Ligustrum
Pyracantha

Cydonia oblonga

Berchemia scandens
Cercis canadensis
Juniperus virginiana

Rhododendron spp.
Roystonea elata

Sassafras albidum
Amelanchier spp.

Oxydendrum arboreum
Lindera benzoin
Spiraea

Picea

Picea mariana

Picea pungens

Picea engelmannii
Picea abies

Picea rubens
Picea sitchensis
Picea glauca
Fragaria

Celtis laevigata
Rhus spp.

Rhus typhina

Myrica (Comptonia) aspleniifolia

Liquidambar styraciflua

Index to Insects by Host Plants

This index should enable foresters and
others not trained in entomology to identify
many of the insects and related forms causing
damage to particular species of trees or
shrubs. Only those insects having definite
host plants are included. The common names
of insects, where known, are used; other-
wise, the scientific names are given. Where
applicable, the part of the host injured or
destroyed is listed in the following catego-
ries: foliage; bark, wood, or twigs; buds,
shoots, or roots; flowers, seeds, or fruits.

fig tree borer, 308

Oberea pallida, 300
poplar-and-willow borer, 334
Sinodendron rugosum, 267
Sthenopis argenteomaculatus, 124

sucking insects—

alder spittlebug, 74
Chionaspis lintneri, 111
Corythucha pergandei, 65
Psylla floccosa, 77
galeaformis, 77
Pterocallis alnifoliae, 79
woolly alder aphid, 80

The readily identifiable sucking insects are Andromeda:

listed separately from the feeding categories cottony maple leaf scale, 98
because of their primarily oligophagous Apple:

feeding habits. In addition, wood products is foliage —

listed as a host. *)

A

Acacia:
Cryptothelea gloverii, 127
mesquite borer, 287
Oncideres pustulatus, 301
Thysanoes fimbricornis, 367
Acacia, sweet:
Oncideres pustulatus, 301
Ailanthus:
ailanthus webworm, 140
Asiatic garden beetle, 270
cynthia moth, 206
Alder:
foliage—
Acronicta distans, 236
alder flea beetle, 263
alder tubemaker, 181
Arge scapularis, 382
birch sawfly, 382
birch skeletonizer, 128
Bruce spanworm, 189
Caloptilia pulcheila, 132
Chrysomela interrupta, 260
Clastoptera obtusa, 74
Corythucha pergandei, 65
cottonwood leaf beetle, 260
Croesus varus, 407
Dichelonyx elongata, 270
elm sawfly, 382
European alder leafminer, 400
European snout beetle, 320
Evora hemidesma, 149
Himatolabus pubescens, 319
Tridopsis larvaria, 192
large aspen tortrix, 168
Nematus erythrogaster, 406
smeared dagger moth, 236
spear-marked black moth, 190
stout looper, 195
striped alder sawfly, 407
threelined leafroller, 161
western tent caterpillar, 204
bark, wood, twigs—
alder borer, 298
birch and beech girdler, 310
buds, shoots, roots—
alder borer, 298
birch and beech girdler, 310
Dicerca lurida, 284
European snout beetle, 320

Acronicta distans, 236
interrupta, 236

apple bucculatrix, 128

apple flea weevil, 334

apple-and-thorn skeletonizer, 145

basswood leafminer, 263

browntail moth, 235

cecropia moth, 206

crinkled flannel moth, 176

Datana major, 218

definite-marked tussock moth, 228

eastern tent caterpillar, 202

fall cankerworm, 188

filament bearer, 195

flatheaded appletree borer, 281

gypsy moth, 229

Haploa clymene, 226

io moth, 208

linden looper, 193

locust leafminer, 264

oriental moth, 177

palmerworm, 137

pearleaf blister mite, 31

pepper-and-salt moth, 200

Phyllonorycter crataegella, 130

Polydrusus impressifrons, 321

red spotted purple, 175

redbanded leafroller, 169

resplendent shield bearer, 126

roundheaded appletree borer, 297

Schizura ipomoeae, 223
leptinoides, 223

smeared dagger moth, 236

Sparganothis sulfureana, 161

Sphinx drupiferarum, 214

spring cankerworm, 193

twinspot sphinx, 215

unicorn caterpillar, 223

unspotted leafminer, 131

variable oakleaf caterpillar, 220

velleda lappet moth, 206

western tent caterpillar, 204

white admiral, 175

whitemarked tussock moth, 227

willow flea weevil, 334

winter moth, 189

bark, wood, twigs—

American plum borer, 186
apple bark borer, 142
apple twig borer, 257
banded hickory borer, 289

537

flatheaded appletree borer, 281
larger shothole borer, 354
leopard moth, 145
Lichenophanes bicornis, 257
Lymantor decipiens, 361
Pseudolucanus capreolus, 267
roundheaded appletree borer, 297
Saperda cretata, 298
buds, shoots, roots—
Polydrusus impressifrons, 321
flowers, seeds, fruits—
apple fruit moth, 140
apple maggot, 448
birch casebearer, 134
cigar casebearer, 134
plum curculio, 336
quince curculio, 336
sucking insects—
apple mealybug, 91
cottony maple scale, 97
European fruit scale, 120
Forbes scale, 119
oystershell scale, 114
periodical cicada, 75
scurfy scale, 109
striped mealybug, 91
woolly apple aphid, 79
Apricot:
European fruit lecanium, 96
Aralia:
twobanded Japanese weevil, 322
Arborvitae: See: white-cedar, thuja
Ardisia:
Xylosandrus zimmermanni, 374
Ash:
foliage—
Agonopterix nigrinotella, 132
American dagger moth, 236
Cecidomyia pellex, 443
cecropia moth, 206
Contarinia canadensis, 443
definite-marked tussock moth, 228
elm spanworm, 196
fall cankerworm, 188
fruittree leafroller, 163
giant walkingstick, 51
great ash sphinx, 214
green fruitworm, 237
hickory horned devil, 211
lilac leafminer, 131
Machimia tentoriferella, 132
Melanolophia canadaria, 191
Olcerlostera angelica, 201
Pachybrachis othonus, 266
Papaipema furcata, 237
Plagodis kuetzingi, 195
polyphemus moth, 207
promethea moth, 207
regal moth, 211
stout looper, 195
Tetranychus homorus, 31
tiger swallowtail, 173
Trichiosoma triangulum, 383
velleda lappet moth, 206
walkingstick, 50
waved sphinx, 213
bark, wood, twigs—
American dagger moth, 236

538

ash and privet borer, 308
banded ash borer, 294
Chrysobothris sexsignata, 283
eastern ash bark beetle, 341
eastern Hercules beetle, 272
Hemicoelus carinatus , 253
Hylesinus pruinosus, 341
ivory-marked beetle, 302
leopard moth, 145
lilac borer, 143
painted hickory borer, 287
powderpost beetle, 255, 256
redheaded ash borer, 294
Xvleborus ferrugineus, 373
Xyvlosandrus germanus, 374
Xvloterinus politus, 371
flowers, seeds, fruits—
Eriophyes fraxiniflora, 31
sucking insects—
Asterolecanium arabidis, 106
boxelder bug, 67
buffalo treehopper, 71
Chionaspis kosztarabi, \\1
common falsepit scale, 104
cottony maple scale, 97
Dysmicoccus difficilis, 94
Japanese maple scale, 115
Melanaspis nigropunctata, 116
oystershell scale, 114, 115
periodical cicada, 75
Prociphilus fraxinifolii, 80
sycamore lace bug, 64
Tropidosteptes amoenus, 66
walnut scale, 119
Ash, black:
lilac leafminer, 131
oak-bark scaler, 309
Ash, green:
banded ash clearwing, 144
carpenterworm, 146
Ash, red:
blackheaded ash sawfly, 408
brownheaded ash sawfly, 408
Ash, velvet:
Formosan subterranean termite, 60
Ash, white:
follage—
blackheaded ash sawfly, 408
brownheaded ash sawfly, 408
laurel sphinx, 214
mountain-ash sawfly, 404
notch-wing geometer, 196
Sparganothis diluticostana, 161
reticulatana, 161
bark, wood, twigs—
banded ash clearwing, 144
Camponotus mississippiensis, 433
Hemicoelus carinatus, 253
Aspen: See: poplar
Azalea: See: rhododendron

B

Baldcypress:
foliage—
Anacamptodes pergracilis, 191
bagworm, 126
Coleotechnites apicitripunctella, 135
variella, 136
imperial moth, 212

Oiketicus abbotii, 126
Oligonychus boudreauxi, 31
pine colaspis, 263
Platytetranychus thujae, 31
Systena marginalis, 265
Taxodiomyia cupressiananassa, 443
bark, wood, twigs—
Buprestis striata, 281
Chrysobothris sexsignata, 283
cypress bark borer, 306
flatheaded baldcypress sapwood borer,
Dag
ivory-marked beetle, 302
Micracisella opacicollis, 367
Oeme rigida rigida, 309
Platypus compositus, 376
southern cypress beetle, 35i
Trachykele lecontei, 285
Urocerus taxodii, 410
Xvleborinus saxeseni, 374
Xvleborus affinis, 374
ferrugineus, 373

flowers, seeds, fruits—

baldcypress coneworm, 185
Sequoiomyia cupressi, 443

sucking insects—

Cinara tujafilina, 78
Platytetranychus thujae, 31
Quadraspidiotus taxodii, 121

drakei, 268

implicita, 269

prunina, 269
Plagiometriona clavata, 266
polyphemus moth, 207
purplish-brown looper, 199
question-mark butterfly, 173
red spotted purple, 175
redhumped oakworm, 219
Sparganothis pettitana, \61
stout looper, 195
threelined leafroller, 161
tiger swallowtail, 173
variable oakleaf caterpillar,
velleda lappet moth, 206
walkingstick, 50
white admiral, 175
whitemarked tussock moth, 227
winter moth, 189
yellownecked caterpillar, 216

i
i)
oS

bark, wood, twigs—

American plum borer, 186

Chrysobothris azurea, 283

Columbian timber beetle, 369

Dicerca lurida, 284

flatheaded sycamore-heartwood borer,
283

Hexomyza tiliae, 450

linden bark borer, 135

Bamboo: linden borer, 297
Asterolecanium bambusae, 106 Platypus compositus, 376
miliaris miliaris, 106 Pseudothysanoes rigidus, 356
miliaris robustum, 106 Saperda imitans, 298
bamboo powderpost beetle, 257 lateralis, 298
Chlorophorus annularis, 317 sapwood timberworm, 248
Banyan: Trigonarthris proxima, 306
Lymire edwardsii, 226 twig girdler, 301
Basswood: Xiphyvdria abdominalis, 411
foliage— buds, shoots, roots—

American dagger moth, 236
Archips purpuranus, 164
bagworm, 126

basswood blotchminer, 130
basswood leafminer, 263
basswood leafroller, 178
cecropia moth, 206
Coleophora tiliaefoliella, 133
Datana drexelii, 217

elm dagger moth, 236

elm sawfly, 382

elm sphinx, 213

fall cankerworm, 188
fourspotted spider mite, 31
fruittree leafroller, 163
giant walkingstick, 51
gypsy moth, 229

hemlock looper, 198
hickory tussock moth, 223
Japanese beetle, 271

linden borer, 297

linden looper, 193

linden wart gall, 443
Melanolophia canadaria, 191
notched-wing geometer, 196
palmerworm, 137
Pandemis lamprosana, 161

broadnecked root borer, 309
Japanese beetle, 271
Phyllophaga crenulata, 269

drakei, 268

implicita, 269

prunina, 269

flowers, seeds, fruits —

fruittree leafroller, 163
Japanese beetle, 271

sucking insects—

apple mealybug, 91
basswood aphid, 79
basswood lace bug, 65
common falsepit scale, 104
cottony maple scale, 97
elm armored scale, 111
elm scurfy scale, 108
giant bark aphid, 77
Japanese maple scale, 115
maple mealybug, 92
mulberry whitefly, 87
Parrott scale, 105

Putnam scale, 112
Telamona reclivata, 71
tuliptree scale, 99

walnut lace bug, 65
walnut scale, 119

Basswood, American:
Hemicoelus carinatus, 253

Phigalia titea, 193
Phyllophaga crenulata, 269

539

Ptilinus ruficornis, 254
Bayberry:

Oiketicus abbotii, 126
Beech:

ftoliage—

Pseudopityophthorus pruinosus, 363
Scolytus fagi, 356
Xiphydria tibialis, 411
Xyleborus ferrugineus, 373
validus, 374

Acleris chalybeana, 171
Asiatic oak weevil, 322
Bruce spanworm, 189
Bucculatrix packardella, 129
Choristoneura fractivittana, 169
Coleophora alniella, 133
Colocasia propinquilinea, 237
dark tussock moth, 228
Datana angusi, 217
fall cankerworm, 188
flatheaded appletree borer, 281
green oak caterpillar. 219
Heterocampa biundata, 222
imperial moth, 212
io moth, 208
Lambdina fervidaria athasaria, 199
locust leafminer, 264
luna moth, 207
Macrarocampa marthesia, 222
maple leafcutter. 125
maple trumpet skeletonizer. 157
maple webworm, 180
Norape ovina, 176
notched-wing geometer. 196
Oligocentria lignicolor, 222
Oligonychus bicolor, 30
Pandemis lamprosana, 161
Phyllophaga drakei, 268
forsteri, 269
implicita, 269
luctuosa, 268
prunina, 269
Plagodis serinaria, 195
polyphemus moth. 207
Prolimacodes badia, 178
redhumped oakworm. 219
saddled prominent. 220
Schizura ipomoeae, 223
leptinoides, 223
spotted tussock moth, 224
Tetralopha asperatella, 180
unicom caterpillar. 223
variable oakleaf caterpillar. 220
walnut sphinx, 215
Xanthonia decemnotata, 266
bark, wood, twigs—
Actenodes acornis. 285
beech borer, 293
birch and beech girdler, 310
birch bark beetle, 362
Buprestis rufipes, 281
Chrysobothris sexsignata, 283
Columbian timber beetle. 369
Eucrada humeralis, 255
flatheaded appletree borer. 281
flatheaded sycamore-heartwood borer.
283
leopard moth. 145
Lichenophanes bicornis, 257
New York weevil, 331
oak timberworm, 318
pigeon tremex. 410
Platypus compositus, 376

Xylosandrus germanus, 374

Xyloterinus politus, 371

buds. shoots, roots—

Asiatic oak weevil, 322

Phyllophaga drakei, 268
forsteri, 269
implicita, 269
luctuosa, 268
prunina, 269

Tlowers, seeds, fruits—

filbertworm, 159

sucking insects—

beech blight aphid, 79

birch lace bug. 65

birch margarodid,. 90

Calaphis betulella, 79

giant bark aphid, 77

large hickory lecanium, 95

oystershell scale. 114

Peliococcus serratus, 94

Phyllaphis fagi, 79

Beech, American:
beech scale, 101
Hemicoelus carinatus, 253
Psilocorsis cryptolechiella, 132
Prilinus ruficornis, 254
Xyleborinus saxesensi, 374
Beech, European:
beech scale, 101
Beech, Oriental:
beech scale. 101
Birch:
foliage—

Acronicta distans, 236
innotata, 236
interrupta, 236

ambermarked birch leafminer. 401

apple-and-thorn skeletonizer. 145

Ancylis spp.. 158

Arge abdominalis, 382
clavicornis, 382
scapularis, 382

basswood leafminer, 263

birch skeletonizer, 128

bronze birch borer, 128, 277

cecropia moth. 206

Charadra deridens, 237

Colocasia propinquilinea, 237

crinkled flannel moth, 176

elm prominent. 222

elm sawfly. 382

elm sphinx, 213

European snout beetle. 320

fruittree leafroller, 163

gypsy moth, 229

Haploa lecontei, 226

hickory tussock moth, 223

lappet moth, 206

linden looper. 193

locust leafminer, 264

Lomgarapha vestaliata. 190

luna moth, 207

Machimia tentoriferella, 132

maple leafcutter, 125
Melanolophia canadaria, 191
Pachybrachis peccans, 266
paddle caterpillar, 236
Phyllophaga crenulata, 269

drakei, 268

forsteri, 269

luctuosa, 268
pinkstriped oakworm, 209
Polydrusus impressifrons, 321
polyphemus moth, 207
poplar dagger moth, 236
Pristiphora siskiyouensis, 404
promethea moth, 207

Protoboarmia porcelaria indicataria,

200
Psilocorsis cryptolechiella, 132
red spotted purple, 175
rose chafer, 270
saddleback looper, 192
spotted tussock moth, 224
striped alder sawfly, 407
Syneta ferruginea, 266
tiger swallowtail, 173
Trichiosoma triangulum, 383
twinspot sphinx, 215
western tent caterpillar, 204
white admiral, 175

bark, wood, twigs—

Actenodes acornis, 285
alder borer, 298
birch and beech girdler, 310
birch bark beetle, 362
Brachyleptura vagans, 306
Chrysobothris azurea, 283
Conotrachelus anaglyptius, 337
pigeon tremex, 410
Platypus compositus, 376
poplar-and-willow borer, 334
Pselaphorhynchites cyanellus, 319
sapwood timberworm, 248
Tropideres fasciatus, 318
Trypodendron betulae, 371
Xiphydria mellipes, 411
tibialis, 411
Xyleborinus saxesensi, 374
Xyleborus obliquus, 374
Xyloterinus politus, 371

buds, shoots, roots—

European snout beetle, 320
Phyllophaga crenulata, 269
drakei, 268
forsteri, 269
luctuosa, 269

flowers, seeds, fruits—

Semudobia spp., 445
sucking insects—

alder spittlebug, 74

birch margarodid, 90
Calaphis betulella, 79
Chionaspis lintneri, 111
common birch aphid, 79
Corythucha pergandei, 65
giant bark aphid, 77

large hickory lecanium, 95
oystershell scale, 114
Psylla carpinicola, 77

Birch, European white:

birch casebearer, 134
birch leafmining sawfly, 398

Birch, gray:

foliage—
ambermarked birch leafminer, 401
Archiearis infans, 188
birch casebearer, 134
birch leafminer, 399
birch leafmining sawfly, 398
birch sawfly, 382
chainspotted geometer, 199
cigar casebearer, 134
Compton tortoiseshell, 174
Datana angusi, 217
dusky birch sawfly, 406
Epinotia stroemiana, 160
filament bearer, 195
gypsy moth, 229
Japanese beetle, 271
Tortricidia flexuosa, 178
willow flea weevil, 334
bark, wood, twigs—

Pseudopitvophthorus asperulus, 363

flowers, seeds, fruits—
Japanese beetle, 271

sucking insects—
Euceraphis lineata, 79
Kleidocerys resedae geminatus, 68

Birch, paper:

foliage—
Acleris logiana, 172
tripunctana, 172
ambermarked birch leafminer, 401
American dagger moth, 236
Archiearis infans, 188
Archips purpuranus, 164
Bibarrambla allenella, 132
birch casebearer, 134
birch leafmining sawfly, 398
birch sawfly, 382
birch skeletonizer, 128
birch tubemaker, 181
bronze birch borer, 277
Bruce spanworm, 189
chainspotted geometer, 199
Choristoneura fractivittana, 169
Compton tortoiseshell, 174
definite-marked tussock moth, 228
Epinotia solandriana, 158
giant walkingstick, 51
graybanded leafroller, 171
green oak caterpillar, 219
gypsy moth, 229
hemlock looper, 198
Heterocampa biundata, 222
io moth, 208
Tridopsis larvaria, 192
large aspen tortrix, 168
large maple spanworm, 199
Lophodonta ferruginea, 219
Nematus pinguidorsum, 406
viridescens, 406
Nites betulella, 132
notched-wing geometer, 196
oak beauty, 195
obliquebanded leafroller, 168
Pandemis lamprosana, 161

54]

Plagodis serinaria, 195
purplish-brown looper, 199
redhumped caterpillar, 222
redhumped oakworm, 219
Schizura ipomoeae, 223
leptinoides, 223
Sparganothis diluticostana, 161
reticulatana, 161
spear-marked black moth, 190
stout looper, 195
threelined leafroller, 161
unicorn caterpillar, 223
variable oakleaf caterpillar, 220
walkingstick, 50
whitemarked tussock moth, 227
willow flea weevil, 334
yellownecked caterpillar, 216
bark, wood, twigs—
bronze birch borer, 277
Hemicoelus carinatus, 253
Prilinus ruficornis, 254
buds, shoots, roots—
Apagodiplosis papyriferae, 443
sucking insects—
birch lace bug, 65
Psylla annulata, 77
Birch, river:
Acleris logiana, 172
Acrobasis betulivorella, 181
Agrilus betulae, 280
Euceraphis mucida, 79
larger elm leaf beetle, 262
Phytobia pruinosa, 449
Xyleborus affinis, 374
Birch, sweet:
Bucculatrix coronatella, 129
Dichelonyx elongata, 270
oriental moth, 177
Phytobia pruinosa, 449
Birch, white: See: paper
Birch, yellow:
foliage—
Acleris chalybeana, 171
ambermarked birch leafminer, 401
American dagger moth, 236
birch leafmining sawfly, 398
bronze birch borer, 277
Caloptilia pulchella, 132
hemlock looper, 198
Plagodis serinaria, 195
saddled prominent, 220
Sciaphillus asperatus, 320
whitemarked tussock moth, 227
yellownecked caterpillar, 216
buds, shoots, roots—
Sciaphillus asperatus, 320
bark, wood, twigs—
bronze birch borer, 277
Chrysobothris sexsignata, 283
Columbian timber beetle, 369
Hemicoelus carinatus, 253
Prilinus ruficornis, 254
sucking insects—
birch lace bug, 65
Carynota stupida, 71
Elasmuche lateralis, 64
European birch aphid, 79
Kleidocerys resedae geminatus, 68

Bittersweet:
euonymus scale, 121
Petalium seriatum, 254
Blackgum: See tupelo, black
Blue-beech: See hornbeam, American
Blueberry:
azalea bark scale, 103
chainspotted geometer, 199
common falsepit scale, 104
giant walkingstick, 51
walkingstick, 50
Boxelder:
follage—
American dagger moth, 236
Archips negundanus, 164
Asiatic garden beetle, 270
boxelder gall midge, 443
boxelder leafroller, 132
fall cankerworm, 188
flatheaded appletree borer, 281
green fruitworm, 237
greenstriped mapleworm, 210
smeared dagger moth, 236
spotted tussock moth, 224
threelined leafroller, 161
variable oakleaf caterpillar, 220
bark, wood, twigs—
boxelder twig borer, 156
Columbian timber beetle, 369
flatheaded appletree borer, 281
hardwood stump borer, 315
buds, shoots, roots—
Archodontes melanopus melanopus,
315
flowers, seeds, fruits—
green fruitworm, 237
sucking insects—
boxelder aphid, 78
boxelder bug, 67
Boxwood:
folhage—
boxwood leafminer, 442
sucking insects—
boxwood psyllid, 76
common falsepit scale, 104
oystershell scale, 114
striped mealybug, 91
Boxwood, European:
Eurytetranychus buxi, 31
Brazil nut tree:
Hypothenemus obscurus, 368
Buckeye:
foliage—
Asiatic garden beetle, 270
Derocrepis aesculi, 266
Eotetranychus hicoriae, 31
filament bearer, 195
fourspotted spider mite, 31
fruittree leafroller, 163
Japanese beetle, 271
Phyllophaga crenulata, 269
bark, wood, twigs—
Petalium bistriatum, 254
sapwood timberworm, 248
buds, shoots, roots—
Proteoteras aesculana, 156
flowers, seeds, fruits—
Japanese beetle, 271

sucking insects—
elm armored scale, 111
common falsepit scale, 104
Comstock mealybug, 93
Corythucha aesculi, 65
maple mealybug, 92
Parrott scale, 105

Bumelia:

dentate scale, 123

holly pit scale, 106

Parrott scale, 105

Butternut:

foliage—
Acordulecera spp., 381
butternut woollyworm, 409
Datana angusi, 217
Eugnamptus collaris, 319
hickory horned devil, 211
hickory tussock moth, 223
luna moth, 207
pecan leaf casebearer, 181
polyphemus moth, 207
regal moth, 211
walnut caterpillar, 217
walnut sphinx, 215
yellownecked caterpillar, 216

bark, wood, twigs—
Agrilus juglandis, 280
Brachyleptura vagans, 306
hickory bark beetle, 355
painted hickory borer, 287
Saperda discoidea, 298

buds, shoots, roots—
pecan leaf casebearer, 181

flowers, seeds, fruits—
Conotrachelus retentus, 337

sucking insects—
twomarked treehopper, 71
walnut lace bug, 65

C

Camellia:
camellia scale, 115
camphor scale, 118
Cerococcus kalmiae, 105
common falsepit scale, 104
cottony camellia scale, 98
cottony taxus scale, 98
Indian wax scale, 94
Japanese wax scale, 94
peony scale, 117
southern red mite, 31
tea scale, 113
Camphor-tree:
camphor scale, 118
Caragana:
ashgray blister beetle, 243
caragana blister beetle, 243
Nuttall blister beetle, 243
Casuarina, horsetail:
Australianpine borer, 282
Clastoptera undulata, 74
Catalpa:
catalpa sphinx, 213
Contarinia catalpae, 443
Trilobomyza pleuralis, 450
sucking insects—
common falsepit scale, 104
Comstock mealybug, 93

dentate scale, 123
elm armored scale, 111
striped mealybug, 91
Cedar spp.:
deodar weevil, 330
Forbes scale, 119
Nuculaspis pseudomeyeri, 117
shortneedle evergreen scale, 117
Cedar, Atlas:
deodar weevil, 330
Cedar, deodar:
deodar weevil, 330
Pityophthorus pulicarius, 363
redheaded pine sawfly, 384
Cedar of Lebanon:
deodar weevil, 330
Cherry:
folage—
Acronicta distans, 236
interrupta, 236
Anacampsis innocuella, 137
basswood leafminer, 263
cecropia moth, 206
chainspotted geometer, 199
cherry leaf beetle, 262
crinkled flannel moth, 176
cynthia moth, 206
eastern tent caterpillar, 202
European snout beetle, 320
fall cankerworm, 188
giant walkingstick, 51
Haploa lecontei, 226

hawthorn leafmining sawfly, 401

hemlock looper, 198

i0 moth, 208

lappet moth, 206

locust leafminer, 264
Lomgarapha vestaliata, 190
oriental moth, 177
palmerworm, 137
pepper-and-salt moth, 200

Phyllonorycter crataegella, 130

promethea moth, 207
resplendent shield bearer, 126
smalleyed sphinx, 215
Sparganothis sulfureana, 161
Sphinx drupiferarum, 214
Sterictiphora spp., 382
Tetracis cachexiata, 199
threelined leafroller, 161
tiger swallowtail, 173
velleda lappet moth, 206
walkingstick, 50
western tent caterpillar, 204
willow flea weevil, 334

bark, wood, twigs—
birch bark beetle, 362
black carpenter ant, 430
Chrysobothris viridiceps, 283
larger shothole borer, 354
Olethreutes quadrifidum, 160
peach bark beetle, 350
Phytobia pruinosa, 449

pruni, 449

powderpost beetle, 255
Pseudolucanus capreolus, 267
Saperda lateralis, 298
Xylosandrus crassiusculus, 375

543

buds, shoots, roots—

European snout beetle, 320

flowers, seeds, fruits—

apple fruit moth, 140
birch casebearer, 134
black cherry fruit fly, 448
cherry fruit fly, 448

cigar casebearer, 134
plum curculio, 336

sucking insects—

Forbes scale, 119
Prosapia bicincta, 74
Psylla trimaculata, 77

Cherry, black:
foliage—

Apatelodes torrefacta, 200
apple bucculatrix, 128
Archips purpuranus, 164
Cerura borealis, 223

cherry scallop shell moth, 190
cherry webspinning sawfly, 381
Heterocampa biundata, 222
pear sawfly, 401

Prolimacodes badia, 178

red spotted purple, 175
spotted tussock moth, 224
uglynest caterpillar, 161

white admiral, 175

bark, wood, twigs—

Hypothenemus eruditus, 368
Oberea ulmicola, 300
Trigonarthris minnesotana, 306
Xyloterinus politus, 371

buds, shoots, roots—

Contarinia cerasiserotinae, 443

Cherry, choke:
foliage—

Bruce spanworm, 189

graybanded leafroller, 171

Hoplocampa lacteipennis, 409
montanicola, 409

prairie tent caterpillar, 203

uglynest caterpillar, 161

western tent caterpillar, 204

flowers, seeds, fruits—

apple maggot, 448
chokecherry midge, 443

Cherry, Japanese flowering:
white prunicola scale, 118
Cherry, pin:
foliage—

544

Acronicta oblinita, 236

Bruce spanworm, 189

cherry leaf beetle, 262

cherry webspinning sawfly, 381

definite-marked tussock moth, 228

filament bearer, 195
Machimia tentoriferella, 132
Packardia geminata, 178
Pero honestaria, 197
Prolimacodes scapha, 178
purplish-brown looper, 199
Schizura ipomoeae, 223
leptinoides, 223
smeared dagger moth, 236
stout looper, 195
Tortricidia flexuosa, 178
unicorn caterpillar, 223

unspotted leafminer, 131
wildcherry looper, 190
bark, wood, twigs—
American plum borer, 186
sucking insects—
Corythucha associata, 65
pruni, 65
Cherry, sour:

apple-and-thorn skeletonizer, 145

unspotted leafminer, 131
Chestnut:
foliage—
Caliroa sp., 401
Croesus castaneae, 407

Dyseriocrania auricyanea, 124

oak skeletonizer, 128
pinkstriped oakworm, 209
bark, wood, twigs—
chestnut bark borer, 304
chestnut timberworm, 248
Enaphalodes atomarius, 292
ivory-marked beetle, 302
leopard moth, 145
Magdalis salicis, 333
oak sapling borer, 293
oak-stem borer, 303
Platypus compositus, 376

Pseudopityophthorus asperulus, 363

pubescens, 363

Pseudothysanoes lecontei, 356

Scobicia bidentata, 257
twolined chestnut borer, 277
Xyleborus obliquus, 374
xylographus, 374
buds, shoots, roots—
broadnecked root borer, 309
tilehorned prionus, 309
flowers, seeds, fruits—
acorn moth, 133
Curculio spp., 333
filbertworm, 159
Chestnut, American:
large chestnut weevil, 333
small chestnut weevil, 333
Synanthedon castaneae, 143
Chestnut, Chinese:
Asiatic oak weevil, 322
Dryocosmus kuriphilus, 425
large chestnut weevil, 333
small chestnut weevil, 333
Chestnut, Japanese:
Asiatic oak weevil, 322
Dryocosmus kuriphilus, 425
large chestnut weevil, 333
small chestnut weevil, 333

Chinaberry:

citrus whitefly, 87

Chinquapin:

Dyseriocrania auricyanea, 124

Coffeetree, Kentucky:

Epicauta torsa, 243

Conifers:

foliage—
Amorbia humerosana, 161
Dasychira plagiata, 229
rusty tussock moth, 227
small conifer looper, 190
spruce-fir looper, 191

Tortricinae, 160
bark, wood, twigs—
brown prionid, 315
Callidium antennatum antennatum,
304
Cossonus impressus, 337
Crypturgus borealis, 362
Gnathotrichus materiarius, 372
pole borer, 307
striped ambrosia beetle, 371
Tetropium cinnamopterum, 308
whitehorned horntail, 410
Xyleborus spp., 372
Xylotrechus sagittatus sagittatus, 311
yellowhorned horntail, 410
buds, shoots, roots—
Couper’s collar weevil, 326
pales weevil, 323
Polyphylla variolosa, 269
Warren’s collar weevil, 326
flowers, seeds, fruits—
Cimberis spp., 318
Conophthorus spp., 364
Diodyrhynchus spp., 318
Dioryctria spp., 182
Cotoneaster:
apple mealybug, 91
twocirculi mealybug, 92
Cottonwood: See also: poplar
foliage—
poplar vagabond aphid, 80
Prodiplosis morrisi, 445
Zengophora scutellaris, 265
bark, wood, twigs—
aspen carpenterworm, 146
carpenterworm, 146
cottonwood borer, 289
Paranthrene dollii, 144
tabaniformis, 145
poplar borer, 295
buds, shoots, roots—
cottonwood borer, 289
Crapemyrtle:
citrus whitefly, 87
crapemyrtle aphid, 79
Creeper, Virginia:
grape leaffolder, 178
Cryptomeria:
cryptomeria scale, 107
juniper scale, 108
minute cypress scale, 108
Cucumbertree:
Archips magnolianus, 164
Cypress:
cryptomeria scale, 107
juniper scale, 108
minute cypress scale, 108
Nuculaspis pseudomeyeri, 117
Phloeosinus scopulorum neomexicanus,

392
D

Dogwood:
foliage—
Asiatic oak weevil, 322
dark tussock moth, 228
Epinotia lindana, 160
giant walkingstick, 51
io moth, 208

Macremphytus spp., 409
Phyllophaga drakei, 268
implicita, 269
redhumped caterpillar, 222
twobanded Japanese weevil, 322
walkingstick, 50
bark, wood, twigs—
Agrilus cephalicus, 280
Chrysobothris azurea, 283
dogwood borer, 141
dogwood clubgall midge, 445
dogwood twig borer, 299
Hypothenemus eruditus, 368
Oberea ulmicola, 300
Petalium bistriatum, 254
pitted ambrosia beetle, 370
Synanthedon geliformis, 142
twig girdler, 301
Xyleborinus saxeseni, 374
Xvleborus rubricollis, 374
buds, shoots, roots—
Asiatic oak weevil, 322
Phyllophaga drakei, 268
implicita, 269
flowers, seeds, fruits—
apple maggot, 448
sucking insects—
calico scale, 94
common falsepit scale, 104
cottony maple leaf scale, 98
cottony maple scale, 97
dogwood scale, 108
Forbes scale, 119
Melanaspis nigropunctata, 116
mulberry whitefly, 87
periodical cicada, 74
striped mealybug, 91
taxus mealybug, 91
Dogwood, flowering: See: dogwood
Douglas-fir:
bark, wood, twigs—
firtree borer, 307
old house borer, 316
pales weevil, 323
red turpentine beetle, 347
Stephanopachys substriatus, 257
Xylotrechus undulatus, 312
buds, shoots, roots—
eastern pine shoot borer, 154
European chafer, 270
pales weevil, 323
sucking insects—
black pineleaf scale, 116
Cooley spruce gall adelgid, 82
cryptomeria scale, 107
elongate hemlock scale, 113
fiorinia hemlock scale, 113
Leptoglossus occidentalis, 67
pine needle scale, 109

E
Elder:
elder borer, 303
Elder, American:
elder leaftier, 178
Elm:
foliage—
Acronicta interrupta, 236
American dagger moth, 236

apple flea weevil, 334 whitemarked tussock moth, 227

Arge scapularis, 382 willow flea weevil, 334
bagworm, 126 Xanthonia decemnotata, 266
Brachys aeruginosus, 285 yellownecked caterpillar, 216
browntail moth, 233 bark, wood, twigs—

Canarsia ulmiarrosorella, 187 banded ash borer, 294

cecropia moth, 206 beech. borer -293

Charadra deridens, 237 black carpenter ant, 429

Choristoneura fractivittana, 169 black elm bark weevil, 331

claycolored leaf beetle, 266 Buprestis rufipes, 281

comma butterfly, 173 carpenterworm, 146

Dasychira cinnamomea, 229 Columbian timber beetle, 369

definite-marked tussock moth, 228 dogwood twig borer, 299

elm calligrapha, 262 elm bark borer, 307

elm casebearer, 133 flatheaded appletree borer, 281

elm dagger moth, 236 ivory-marked beetle, 302

elm flea beetle, 263 larger shothole borer, 354

elm leaf beetle, 260 leopard moth, 145

elm prominent, 222 Lichenophanes bicornis, 257

elm sawfly, 382 Magdalis barbicornis, 333

elm sphinx, 213 pandura, 333

Eotetranychus matthyssei, 31 midges, 442

European snout beetle, 320 native elm bark beetle, 20

fall cankerworm, 188 Neoclytus scutellaris, 295

fourspotted spider mite, 31 Oberea ulmicola, 300

fruittree leafroller, 163 pigeon tremex, 410

giant walkingstick, 51 Platypus compositus, 376

graybanded leafroller, 171 powderpost beetle, 255

hemlock looper, 198 red elm bark weevil, 331

hickory tussock moth, 223 Saperda lateralis, 298

imperial moth, 212 Scobicia bidentata, 257

io moth, 208 shothole borer, 356

linden looper, 193 smaller European elm bark beetle, 352

locust leafminer, 264 Tetraneura ulmi, 80

Machimia tentoriferella, 132 Thysanoes berchemiae, 367

Macroxyela ferruginea, 378 Trigonarthis minnesotana, 306

maple leafcutter, 125 twig girdler, 301

maple webworm, 180 twig pruner, 302

mourningcloak butterfly, 174 Xiphydria hicoriae, 411

Nerice bidentata, 219 tibialis, 411

notched-wing geometer, 196 Xylosandrus germanus, 374

Oiketicus abbotii, 126 buds, shoots, roots—

Pachybrachis othonus, 266 Phyllophaga forsteri, 269

paddle caterpillar, 236 implicita, 269

Pandemis lamprosana, 161 prunina, 269

pepper-and-salt moth, 200 tristis, 268

Phigalia titea, 193 sucking insects—

Phyllophaga drakei, 268 buffalo treehopper, 71
forsteri, 269 camphor scale, 118
implicita, 269 common falsepit scale, 104
prunina, 269 Corythucha pergandei, 65
tristis, 268 cottony maple scale, 97

polyphemus moth, 207 elm armored scale, 111

puss caterpillar, 175 elm leaf aphid, 79

question-mark butterfly, 173 elm scurfy scale, 108

redhumped caterpillar, 222 European elm scale, 103

redhumped oakworm, 219 fourhumped stink bug, 64

Schizura ipomoeae, 223 giant bark aphid, 77
leptinoides, 223 large hickory lecanium, 95

Schoene spider mite, 31 Neolygus invitus, 66

spiny-elm caterpillar, 174 oystershell scale, 114

spring cankerworm, 193 Parrott scale, 105

stout looper, 195 periodical cicada, 74

threelined leafroller, 161 Putnam scale, 112

twinspot sphinx, 215 tarnished plant bug, 66

unicorn caterpillar, 223 walnut scale, 119

velleda lappet moth, 206 whitebanded elm leafhopper, 70

walkingstick, 50 woolly apple aphid, 79

woolly elm aphid, 79
546

Elm, American:
elm borer, 298
elm calligrapha, 262
elm cockscombgall aphid, 80
elm lace bug, 65
elm leaf beetle, 260
elm leafminer, 400
European elm scale, 103
Gyponana spp., 70
Japanese beetle, 271
Ponana spp., 70
Scaphoideus spp., 70
variable oakleaf caterpillar, 220
whitebanded elm leafhopper, 70
winter moth, 189
woolly elm bark aphid, 79
Elm, Camperdown:
elm leafminer, 400
Elm, English:
elm casebearer, 133
elm leafminer, 400
Japanese beetle, 271
Elm, European:
elm leaf beetle, 260
Elm, Japanese:
larger elm leaf beetle, 262
Elm, rock:
elm cockscombgall aphid, 80
Elm, Scotch:
elm casebearer, 133
elm leafminer, 400
Elm, Siberian:
elm lace bug, 65
elm leaf beetle, 260
Japanese beetle, 271
Elm, slippery:
dark tussock moth, 228
elm borer, 298
elm cockscombgall aphid, 80
Aylocurus langstoni, 367
Kaltenbachiella ulmifusa, 80
larger elm leaf beetle, 262
woolly elm bark aphid, 79
Euonymus:
common falsepit scale, 104
euonymus scale, 121
Indian wax scale, 94
Japanese wax scale, 94
Lepidosaphes yanagicola, 115
lilac leafminer, 131

F
Fig tree:
fig tree borer, 308
Lymire edwardsii, 226
Fir:
foliage—

Acantholyda maculiventris, 380

false hemlock looper, 198

filament bearer, 195

Pero morrisonaria, 197
bark, wood, twigs—

black and red horntail, 410

fir coneworm, 183

firtree borer, 307

flatheaded fir borer, 284

Hylastes tenuis, 340

Melanophila acuminata, 284

old house borer, 316

pales weevil, 323
Phymatodes dimidiatus, 306
red turpentine beetle, 347
Sirex Juvencus, 410
Stephanopachys substriatus, 257
flowers, seeds, fruits—
fir coneworm, 183
spruce bud moth, 157
spruce coneworm, 184
sucking insects—
Cinara confinis, 78
pilicornis, 78
elongate hemlock scale, 113
fiorinia hemlock scale, 113
hemlock scale, 106
Nepytia pellucidaria, 198
pine needle scale, 109
round conifer scale, 107
shortneedle evergreen scale, 117
Fir, balsam:
foliage—
Argyrotaenia occultana, 17\
balsam fir sawfly, 390
balsam gall midge, 442
chainspotted geometer, 199
chameleon caterpillar, 237
early brown looper, 190
eastern blackheaded budworm, 171
Eufidonia notataria, 191
fir needle inchworm, 190
fringed looper, 200
Griselda radicana, 160
hemlock looper, 198
Lexis bicolor, 226
pine tussock moth, 228
Pleroneura brunneicornis, 378

Protoboarmia porcelaria indicataria,

200
redbanded leafroller, 169
small conifer looper, 190
spruce budworm, 164
spruce harlequin, 237
spruce-fir looper, 191
whitemarked tussock moth, 227
whitespotted sawyer, 313
bark, wood, twigs—
balsam bark weevil, 331
balsam fir bark beetle, 357
balsam fir sawyer, 314
black carpenter ant, 429
Chrysobothris neopusilla, 283
scabripennis, 283
Cryphalus fraseri, 363
Crypturgus pusillus, 362
four-eyed spruce bark beetle, 352
Gnathotrichus materiarius, 372
hemlock borer, 284
Marmara spp., 131
Neacanthocinus obsoletus, 304
pusillus, 304
northeastern sawyer, 314
old house borer, 316
Orthotomicus caelatus, 358
Pityophthorus balsameus, 364
cariniceps, 364
opaculus, 363
pulchellus, 364
spruce scolytus, 356

547

Stictoleptura canadensis, 304
whitespotted sawyer, 313
flowers, seeds, fruits—
white pine cone borer, 156
sucking insects—
balsam twig aphid, 78
balsam woolly adelgid, 83
cryptomeria scale, 107
pine spittlebug, 72
Prociphilus bumelia, 78
Saratoga spittlebug, 73
Fir, Fraser:
balsam gall midge, 442
balsam twig aphid, 78
balsam woolly adelgid, 83
Cryphalus fraseri, 363
Dryocoetes autographus, 361
Fir, Siberian: See: balsam fir
Fir, subalpine:
balsam twig aphid, 78
Fir, white:
Pleroneura brunneicornis, 378
Forsythia:
twobanded Japanese weevil, 322
Fremontia:
azalea bark scale, 103
Fringetree:
laurel sphinx, 214
Fruit trees, various:
foliage—
carpenterworm, 146
eastern tent caterpillar, 202
eyespotted bud moth, 154
gypsy moth, 229
Papilio cresphontes, 173
redhumped caterpillar, 222
Texas leafcutting ant, 433
yellownecked caterpillar, 216
bark, wood, twigs—
apple twig borer, 257
carpenterworm, 146
dogwood twig borer, 299
New York weevil, 331
shothole borer, 356
twig girdler, 301
Xyleborus dispar, 373
flowers, seeds, fruits—
fruittree leafroller, 163
plum curculio, 336
sucking insects—
barnacle scale, 94
Florida red scale, 111
Forbes scale, 119
oystershell scale, 114
San Jose scale, 120
scurfy scale, 109
terrapin scale, 95

G
Gardenia:

barnacle scale, 94
Ginkgo:

American plum borer, 186
Gooseberry:

azalea bark scale, 103

prairie tent caterpillar, 203
Grape:

black vine weevil, 321

grape leaffolder, 178

548

grape mealybug, 93

H

Hackberry:

foliage—
flatheaded appletree borer, 281
hackberry butterfly, 175
hackberry nipplegall maker, 76
hackberry stargall, 76
mourningcloak butterfly, 174
Oiketicus abbotii, 126
oriental moth, 177
puss caterpillar, 175
question-mark butterfly, 173
Sphinx drupiferarum, 214
spiny-elm caterpillar, 174
tawny emperor, 175
whitetringed beetle, 322

bark, wood, twigs—
Agrilus celti, 280

lecontei, 280
Chramesus chapuisi, 352
subopacus, 352

Enaphalodes atomarius, 292

flatheaded appletree borer, 281

Formosan subterranean termite, 59

hackberry engraver, 356
Hylocurus langstoni, 367
rudis, 367

Lichenophanes bicornis, 257
Pachypsylla celtidisinteneris, 76
painted hickory borer, 287
petiolgall psyllid, 76
Phloeotribus dentifrons, 350
Pseudothysanoes lecontei, 356
redheaded ash borer, 294
Scobicia bidentata, 257
Scolytus fagi, 356
Thysanoes fimbricornis, 367
twig girdler, 301
twig pruner, 302
Xvleborus affinis, 374

buds, shoots, roots—
budgall psyllid, 76

Archodontes melanopus melanopus,

315
sucking insects—
azalea bark scale, 103
common falsepit scale, 104
cottony maple scale, 97
dentate scale, 123
elm scurfy scale, 108
hackberry lace bug, 65
mulberry whitefly, 87
Neosteingelia texana, 90
Parrott scale, 105
sour-gum scale, 110
Hackberry, netleaf:
blistergall psyllid, 76
Hardwoods:
foliage—
Amorbia humerosana, 161
hag moth, 177
Phyllophaga forsteri, 269
prunina, 269
rugosa, 268
rusty tussock moth, 227
Serica spp., 269
Svmmerista albifrons, 219

Tortricinae, 160 sucking insects—

bark, wood, twigs— apple mealybug, 91
brown prionid, 315 Corythucha bellula, 65
Camponotus pylartes fraxinicola, 433 cottony maple scale, 97
sayi, 432 Eriosoma crataegi, 79
Cossonus concinnus, 337 hawthorn lace bug, 65
impressus, 337 Parrott scale, 105
platalea, 337 scurfy scale, 109
Crematogaster ashmeadi, 434 sour-gum scale, 110
laeviuscula, 434 striped mealybug, 91
divergent beech beetle, 284 twocirculi mealybug, 92
Goes spp., 292 woolly apple aphid, 79
hardwood stump borer, 315 Hazel:
horntails, 409 foliage—
Phloeotribus spp., 350 apple flea weevil, 334
Platypus parallelus, 377 Asiatic oak weevil, 322
pole borer, 307 Cameraria corylisella, 130
red shothole borer, 375 Croesus curvarius, 407
rustic borer, 312 Dichelonyx subvittata, 270
spined bark borer, 303 Himatolabus pubescens, 319
Stenoscelis brevis, 337 larger elm leaf beetle, 262
Xvleborus spp., 372 palmerworm, 137
Xvlosandrus crassiusculus, 375 pinkstriped oakworm, 209
germanus, 374 Sciaphillus asperatus, 320
buds, shoots, roots— spiny oakworm, 208
Phyllophaga rugosa, 268 buds, shoots, roots—
flowers, seeds, fruits— Sciaphillus asperatus, 320
Orthosia hibisci, 237 flowers, seeds, fruits —
Hawthorn: filbertworm, 159
foliage— hazelnut weevil, 333
Acordulecera spp., 381 sucking insects—
apple bucculatrix, 128 Corythucha pergandei, 65
apple flea weevil, 334 Hemlock:
apple-and-thorn skeletonizer, 145 foliage—
Arge clavicornis, 382 Anacamptodes ephyraria, 191
Argyrotaenia quadrifasciana, 171 Asiatic garden beetle, 270
cecropia moth, 206 black vine weevil, 321
Cryptothelea gloverti, 127 eastern blackheaded budworm, 171
eyespotted bud moth, 154 false hemlock looper, 198
hawthorn leafmining sawfly, 401 filament bearer, 195
Hedya chionosema, 160 fringed looper, 200
Hoplocampa oskina, 409 hairy leaf beetle, 266
larger elm leaf beetle, 262 hemlock looper, 198
leaf crumpler, 181 joker, 237
locust leafminer, 264 Lambdina fervidaria athasaria, 199
maple trumpet skeletonizer, 157 Nalepella tsugifoliae, 32
Neurotoma crataegi, 381 purplish-brown looper, 199
pear sawfly, 401 spruce-fir looper, 191
Phyllonorycter crataegella, 130 strawberry root weevil, 321
plum webspinning sawfly, 380 twobanded Japanese weevil, 322
polyphemus moth, 207 bark, wood, twigs
unspotted leafminer, 131 Buprestis maculipennis, 281
whitefringed beetle, 322 striata, 281
bark, wood, twigs— Chrysobothris pusilla, 282, 283
apple bark borer, 142 scabripennis, 283
roundheaded appletree borer, 297 sexsignata, 283
Saperda cretata, 298 Dryocoetes autographus, 361
shothole borer, 356 flatheaded fir borer, 284
thorn-limb borer, 298 hemlock borer, 284
Xiphydria tibialis, 411 Ips latidens, 361
flowers, seeds. fruits— old house borer, 316
apple fruit moth, 140 pales weevil, 323
apple maggot, 448 Stephanopachys substriatus, 257
birch casebearer, 134 Stictoleptura canadensis, 304
cigar casebearer, 134 tanbark borer, 306
flower thrips, 47 Tropideres dorsalis, 318
quince curculio, 336 Trypodendron scabricollis, 371

Xyleborinus saxeseni, 374

549

Xyleborus dispar, 373
Xyloterinus politus, 371
buds, shoots, roots—
black vine weevil, 321
joker, 237
strawberry root weevil, 321
flowers, seeds, fruits—
Nepytia pellucidaria, 198
sucking insects—
elongate hemlock scale, 113
fiorinia hemlock scale, 113
Florida wax scale, 94
hemlock scale, 106
Japanese fiorinia scale, 114
Nuculaspis pseudomeyeri, 117
pine needle scale, 109
round conifer scale, 107
shortneedle evergreen scale, 117
Hemlock, eastern:
Coleotechnites apicitripunctella, 135
cryptomeria scale, 107
Eufidonia notataria, 191
Indian wax scale, 94
Japanese wax scale, 94
pine spittlebug, 72
spruce budworm, 164
white pine cone borer, 156
Hickory:
foliage—
Acordulecera spp.. 381
Acrobasis caryivorella, 181
elyi, 181
exsulella, 181
Acronicta innotata, 236
lithospila, 236
American dagger moth, 236
Anthonomus suturalis, 337
Archips griseus, 164
infumatanus, 164
rileyanus, 164
Asiatic oak weevil, 322
Attelabus bipustulatus, 319
Bassareus literatus, 266
butternut woollyworm, 409
Carvomyia holotricha, 445
sanguinolenta, 445
tubicola, 445
Catocala spp., 236
Colocasia flavicornis, 237
Conotrachelus aratus, 337
Cryptothelea gloverii, 127
dark tussock moth, 228
Datana angusi, 217
elm spanworm, 196
Eotetranychus hicoriae, 31
Eugnamptus collaris, 319
striatus, 319
fall cankerworm. 188
flatheaded appletree borer, 281
fruittree leafroller, 163
gall midges, 442
Glyptoscelis barbata, 266
Haploa clymene, 226
hickory horned devil, 211
hickory leafroller, 170
hickory spiral borer, 279
hickory tussock moth, 223
Homoeolabus analis, 319

550

imperial moth, 212
i0 moth, 208
lappet moth, 206
linden looper, 193
luna moth, 207
Machimia tentoriferella, 132
Megaxyela spp., 378
oriental moth, 177
Pachybrachis peccans, 266
paddle caterpillar, 236
pecan cigar casebearer, 133
Periclista spp., 402
Phigalia titea, 193
Phyllophaga crenulata, 269
tristis, 268
polyphemus moth, 207
raspberry leafroller, 160
redhumped caterpillar, 222
regal moth, 211
Schizura ipomoeae, 223
leptinoides, 223
Tetranychus homorus, 31
unicorn caterpillar, 223
walkingstick, 50
walnut caterpillar, 217
walnut sphinx, 215
white oak borer, 292
bark, wood, twigs—
Actenodes acornis, 285
Agrilus otiosus, 280
banded ash borer, 294
banded hickory borer, 289
black twig borer, 375
Brachyleptura vagans, 306
Buprestis rufipes, 281
Chramesus hicoriae, 352
Chrysobothris adelpha, 283
sexsignata, 283
Conotrachelus anaglyptius, 337
Dicerca lurida, 284
obscura, 284
Drvophthorus americanus, 337
Enaphalodes atomarius, 292
flat powderpost beetle, 299
flatheaded appletree borer, 281
hickory bark beetle, 355
hickory borer, 293
hickory gall phylloxera, 86
hickory spiral borer, 279
Hylocurus bicornus, 367
biorbis, 367
harnedi, 367
rudis, 367
spadix, 367
Hypothenemus dissimilis, 367
eruditus, 368
interstitialis, 368
quercus, 368
rotundicollis, 368
ivory-marked beetle, 302
Lichenophanes bicornis, 257
Lymantor decipiens, 361
Magdalis olyvra, 333
Neoclytus mucronatus mucronatus,
295
scutellaris, 295
New York weevil, 331
Oberea ulmicola, 300

Orchesia castanea, 247
painted hickory borer, 287
pecan carpenterworm, 146
pecan leaf phylloxera, 86
pecan phylloxera, 86
Phymatodes varius, 306
pigeon tremex, 410
Platypus compositus, 376
powderpost beetles, 255
Pseudopitvophthorus pruinosus, 363
Pseudothysanoes dislocatus, 357
redheaded ash borer, 294
redshouldered shothole borer, 257
Saperda discoidea, 297
imitans, 298
lateralis, 298
Scobicia bidentata, 257
Thysanoes fimbricornis, 367
Trigonarthris minnesotana, 306
proxima, 306
twig girdler, 301
twig pruner, 302
white oak borer, 292
Xiphydria hicoriae, 411
Xyleborinus saxeseni, 374
Xyleborus affinis, 374
celsus, 373
devexulus, 374
ferrugineus, 373
lecontei, 374
obliquus, 374
rubricollis, 374
Xyloterinus politus, 371
buds, shoots, roots—
Asiatic oak weevil, 322
Phyllophaga crenulata, 269
tristis, 268
walnut shoot moth, 181
flowers, seeds, fruits—
acorn moth, 133
Conotrachelus affinis, 337
elegans, 337
hicoriae, 337
Curculio spp., 333
hickory shuckworm, 158
pecan weevil, 333
sucking insects—
alder spittlebug, 74
black-margined aphid, 79
black pecan aphid, 79
Chionaspis caryae, 111
Dysmicoccus morrisoni, 94
Forbes scale, 119
hickory scale, 113
large hickory lecanium, 95
Monellia caryella, 79
microsetosa, 719
nigropunctata, 79
Neosteingelia texana, 90
periodical cicada, 74
sycamore lace bug, 64
walnut scale, 119
Hickory, pignut:
mulberry bark borer, 303
oak-bark scaler, 309
Holly:
foliage—
velleda lappet moth, 206

bark, wood, twigs—
Xyvleborinus saxeseni, 374
sucking insects—
camellia scale, 115
common falsepit scale, 104
Comstock mealybug, 93
cottony maple leaf scale, 98
cottony taxus scale, 98
euonymus scale, 121
fem scales 17
Florida red scale, 111
holly pit scale, 106
Indian wax scale, 94
Japanese wax scale, 94
mulberry whitefly, 87
peony scale, 117
Prosapia bicincta, 74
striped mealybug, 91
tea scale, 113
walnut scale, 119
Holly, American:
Asphondylia ilicicola, 443
native holly leafminer, 449
Holly, Chinese:
cottony camellia scale, 98
cottony taxus scale, 98
Indian wax scale, 94
Japanese wax scale, 94
Holly, Japanese:
Indian wax scale, 94
Japanese wax scale, 94
Honeylocust:
foliage—
claycolored leaf beetle, 266
Epicauta torsa, 243
honeylocust pod gall midge, 442
imperial moth, 212
Machimia tentoriferella, 132
mimosa webworm, 138
Nematus tibialis, 406
oriental moth, 177
Platytetranychus multidigituli, 31
Semiothisa ocellinata, 191
Sparganothis sulfureana, 161
yellownecked caterpillar, 216
bark, wood, twigs—
Agrilus difficilis, 280
Hylocurus langstoni, 367
Hypothenemus dissimilis, 367
rotundicollis, 368
ivory-marked beetle, 302
painted hickory borer, 287
twig girdler, 301
Xvleborinus saxeseni, 374
flowers, seeds, fruits
Amblycerus robiniae, 259
sucking insects—
honeylocust plant bug, 66
Lopidea incurva, 66
Lygocoris tinctus, 66
Macropsis spp., 70
Micrutalis calva, 71
periodical cicada, 74
Pilophorus walshii, 66
Plagiognathus delicatus, 66
Taedia gleditsiae, 67

Hophornbeam:

foliage—
Arge clavicornis, 382
basswood leafminer, 263
luna moth, 207
maple leafcutter, 125
Nematus ostryae, 406
Nites betulella, 132
Oligonychus letchworthi, 31
Sciaphillus asperatus, 320
winter moth, 189
yellownecked caterpillar, 216

bark, wood, twigs—
Hypothenemus dissimilis, 367

Pseudopityophthorus pruinosus, 363

Pseudothysanoes lecontei, 356

Thysanoes fimbricornis, 367
buds, shoots, roots—

Sciaphillus asperatus, 320
sucking insects—

birch lace bug, 65

maple mealybug, 92

Hophornbeam, eastern:

Acleris chalybeana, 171

banded hickory borer, 289

birch lace bug, 65

Cinvra gracilipes, 285

luna moth, 207

Parrott scale, 105

pitted ambrosia beetle, 370

Tymnes tricolor, 266

winter moth, 189
Hops:

comma butterfly, 173
Hoptree:

Agonopterix pteleae, 132
Hornbeam: See: hophornbeam
Hornbeam, American:

Acrobasis carpinivorella, 181

birch and beech girdler, 310

Chionaspis kosztarabi, 111

Dicerca lurida, 284

Nematus carpini, 406

pitted ambrosia beetle, 370

Pseudopityophthorus pruinosus, 363

pubescens, 363

striped alder sawfly, 407

Xiphydria tibialis, 411
Horsechestnut: See: buckeye
Huckleberry:

azalea bark scale, 103

chainspotted geometer, 199

I

Incense-cedar:
juniper scale, 108
minute cypress scale, 108
Ironwood:
Pandemis lamprosana, 161
Stilbosis ostryaeella, 135

J
Juniper:
foliage—

Aethes rutilana, 172
Coleotechnites juniperella, 136
Cudonigera houstonana, 169
juniper midge, 442
juniper webworm, 138

‘Nn
Nn
N

Monoctenus fulvus, 384
suffusus, 384
Platytetranychus thujae, 31
bark, wood, twigs—
blackhorned juniper borer, 304
cedartree borer, 307
Oeme rigida rigida, 309
pales weevil, 323
Phloeosinus scopulorum
neomexicanus, 352
small cedar-bark borer, 308
buds, shoots, roots—
strawberry root weevil, 321
sucking insects—
balsam twig aphid, 78
Fletcher scale, 96
Gillette eriococcin, 104
juniper mealybug, 93
juniper scale, 108
Maskell scale. 115
minute cypress scale, 108
Nuculaspis pseudomeyert, 117
round conifer scale, 107
shortneedle evergreen scale, 117
Juniper, California:
Gillette eriococcin, 104
Juniper, Irish:
juniper webworm, 138
Juniper, oneseed:
juniper mealybug, 93
Juniper, western:
Gillette eriococcin, 104

L
Larch:
foliage—

Anomala lucicola, 271

Argyrotaenia occultana, 171

chainspotted geometer, 199

false hemlock looper, 198

fir needle inchworm, 190

green larch looper, 191

hemlock looper, 198

joker, 237

larch casebearer, 134

larch sawfly, 402

Melanolophia canadaria, 191

Nepytia pellucidaria, 198

onelined larch sawfly, 409

Pero honestaria, 197
morrisonaria, 197

Protoboarmia porcelaria indicataria,
200

redbanded leafroller, 169

redheaded pine sawfly, 384

spruce budworm, 164

threelined larch sawfly, 409

Tolype laricis, 206

whitemarked tussock moth, 227

whitespotted sawyer, 313

bark, wood, twigs—

balsam fir bark beetle, 357

blue horntail, 410

Callidium violaceum, 304

Chrysobothris blanchardi, 283
dentipes, 283

Crypturgus pusillus, 362

Dryocoetes affaber, 361

eastern larch beetle, 349

firtree borer, 307
flatheaded fir borer, 284

four-eyed spruce bark beetle, 352
Gnathotrichus materiarius, 372

hemlock borer, 284
Hylobius congener, 326
Aylurgops pinifex, 340

Neoclytus muricatulus muricatulus,

295
Orthotomicus caelatus, 358
pales weevil, 323
Phymatodes dimidiatus , 306
pine spittlebug, 72
red turpentine beetle, 347
spruce scolytus, 356
whitespotted sawyer, 313
buds, shoots, roots—
eyespotted bud moth, 154
larch shoot borer, 140
spruce budworm, 164
flowers, seeds, fruits—
Nepytia pellucidaria, 198
Zeiraphera improbana, 157
sucking insects—
Adelges laricis, 84
lariciatus, 84
Larch, Dahurian:
larch sawfly, 402
Larch, eastern: See: tamarack
Larch, European:
Adelges laricis, 84
eastern larch beetle, 349
larch sawfly, 402
Larch, Japanese:
larch sawfly, 402
Larch, Siberian:
larch sawfly, 402
Larch, subalpine:
larch sawfly, 402
Larch, western:
larch sawfly, 402
Laurel:
dogwood twig borer, 299
eyespotted bud moth, 154
rhododendron borer, 142
Lilac:
great ash sphinx, 214
laurel sphinx, 214
lilac borer, 143
lilac leafminer, 131
Olcerlostera angelica, 201
oystershell scale, 114
promethea moth, 207
waved sphinx, 213
white prunicola scale, 118
Linden: See: basswood
Locust:
cottony maple scale, 97
locust leafroller, 186
twig pruner, 302
Locust, black:
foliage—
Agonopterix robiniella, 132
bagworm, 126

Chrysaster ostensachenella, 130

claycolored leaf beetle, 266
crinkled flannel moth, 176
Dasineura pseudacaciae, 442

giant walkingstick, 51
hickory tussock moth, 223
io moth, 208
locust leafminer, 264
locust leafroller, 186
Nematus abbotii, 406
tibialis, 406
Nephopteryx virgatella, 186
Parectopa robiniella, 131
pepper-and-salt moth, 200
Pero honestaria, 197

Phyllonorycter robiniella, 130

redhumped caterpillar, 222
Schoene spider mite, 31
Semiothisa ocellinata, 191
silverspotted skipper, 172
spotted tussock moth, 224
walkingstick, 50
bark, wood, twigs—
Agrilus egenus, 280
carpenterworm, 146
claycolored leaf beetle, 266
lesser cornstalk borer, 186
Lichenophanes bicornis, 257
locust borer, 286
locust twig borer, 160
painted hickory borer, 287
Xyleborus affinis, 374
buds, shoots, roots—
Obolodiplosis robiniae, 445
sucking insects—
cowpea aphid, 79
Thelia bimaculata, 71

threecornered alfalfa hopper, 71

Vanduzeea arquata, 71

M

Magnolia:
foliage—
Endopiza liriodendrana, 160
Odontopus calceatus, 334
Phyllocnistis magnoliella, \31
Phyllophaga forsteri, 269
Tetranychus magnoliae, 31
bark, wood, twigs—
black twig borer, 375
Euzophera magnolialis, 187
ostricolorella, 187
Platypus compositus, 376
Xyvloterinus politus, 371
buds, shoots, roots—
Euzophera magnolialis, 187
sucking insects
camellia scale, 115
common falsepit scale, 104
dentate scale, 123
magnolia scale, 96
oleander scale, 107
redbay scale, 107
striped mealybug, 91
Trioza magnoliae, 77
tuliptree aphid, 79
tuliptree scale, 99
Magnolia, star:
calico scale, 94
Mahogany:
oriental wood borer, 258
Mahonia:
Parrott scale, 105

553

Mangrove: Aylocurus rudis, 367

Australianpine borer, 282 ivory-marked beetle, 302
Maple: leopard moth, 145
foliage— Lymantor decipiens. 361
Acronicta retardata, 236 Micracisella opacicollis, 367
Apatelodes torrefacta, 200 oak-bark scaler, 309
Asiatic garden beetle. 270 Orchesia castanea, 247
bagworm, 126 Phytobia pruinosa, 449
basswood leafminer, 263 pigeon tremex. 410
browntail moth, 233 Pseudopityophthorus pruinosus, 363
Caloptilia bimaculatella, 132 Synanthedon acerrubri, 143
cecropia moth, 206 Thysanoes fimbricornis, 367
Charadra deridens, 237 lobdelli, 367
Colocasia propinquilinea, 237 Trigonarthris proxima, 306
elm sawfly. 382 twig pruner, 302
fall cankerworm, 188 Xiphydria maculata, 411
filament bearer. 195 Xvleborinus saxesensi, 374
fruittree leafroller, 163 Xvloterinus politus, 371
gall midges, 442 buds, shoots, roots—
greenstriped mapleworm, 210 Asiatic garden beetle. 270
Haploa clymene, 226 hard maple budminer, 125
lecontei, 226 Proteoteras aesculana, 156
hemlock looper, 198 sucking insects—
Heterocampa biundata, 222 Aleurochiton forbesii, 87
umbrata, 222 azalea bark scale, 103
imperial moth, 212 birch lace bug, 65
io moth, 208 birch margarodid, 90
Lambdina fervidaria athasaria, 199 boxelder bug, 67
lappet moth, 206 calico scale, 94
large maple spanworm, 199 camphor scale, 118
linden looper, 193 Chionaspis acericola, 111
Machimia tentoriferella, 132 Comstock mealybug. 93
Macrarocampa marthesia, 222 cottony maple leaf scale, 98
maple bladdergall mite. 31 dentate scale, 123
maple petiole borer. 407 Drepanaphis carolinensis, 79
notched-wing geometer, 196 nigricans, 79
Oligonychus aceris, 31 sabrinae, 79
Phyllophaga drakei, 268 elm armored scale, 111
forsteri, 269 Florida wax scale. 94
implicita, 269 giant bark aphid, 77
tristis, 268 gloomy scale, 116
pinkstriped oakworm, 209 Japanese maple scale, 115
polyphemus moth, 207 maple mealybug, 91
Prolimacodes badia, 178 mulberry whitefly, 87
promethea moth, 207 oystershell scale, 114
Psilocorsis cryptolechiella, 132 painted maple aphid, 79
purplish-brown looper, 199 Parrott scaly 105
puss caterpillar, 175 Periphyllus americanus, 79
saddleback looper, 192 Psylla annulata, 77
Sparganothis pettitana, 161 Putnam scale, 112
reticulatana, 161 sycamore maple aphid, 79
spotted tussock moth. 224 striped mealybug, 91
stout looper, 195 taxus mealybug. 91
twobanded Japanese weevil, 322 terrapin scale, 95
velleda lappet moth. 206 walnut scale, 119
yellownecked caterpillar, 216 Maple, Japanese:
bark, wood, twigs— Japanese beetle, 271
Agrilus masculinus, 280 Maple, mountain:
birch and beech girdler, 310 Acleris chalybeana, 171
black twig borer, 375 maple webworm, 180
boxelder twig borer, 156 Sciaphillus asperatus, 320
Buprestis rufipes, 281 Maple, Norway:
carpenterworm, 146 folage—
Choragus zimmermanni, 318 Caloptilia packardella, 132
Chrysobothris azurea, 283 Isa textula, 178
flatheaded sycamore-heartwood borer. Japanese beetle, 271
283 Obrussa sericopeza, 125
gallmaking maple borer, 310 oriental moth, 177

554

Phyllonorvcter trinotella, 130
whitemarked tussock moth, 227
flowers, seeds, fruits—
Japanese beetle, 271
Obrussa sericopeza, 125
sucking insects—
Norway maple aphid, 79
Maple, red:
foliage—
Acleris chalybeana, \7\
American dagger moth, 236
Choristoneura fractivittana, 169

definite-marked tussock moth, 228

elm spanworm, 196
gouty vein midge, 442
green oak caterpillar, 219
Itame pustularia, 191
Lithophane laticinerea, 237
maple leafcutter, 125
maple leafminer, 130
maple trumpet skeletonizer, 157
maple webworm, 180
ocellate gall midge, 443
Pandemis lamprosana, 161
Phigalia titea, 193
Phyllonorycter trinotella, 130
Plagodis serinaria, 195
Sciaphillus asperatus, 320
Sparganothis acerivorana, 161
willow flea weevil, 334
winter moth, 189
bark, wood, twigs—
Actenodes acornis, 285
Chrysobothris sexsignata, 283
viridiceps, 283
Columbian timber beetle, 369
gallmaking maple borer, 310
maple callus borer, 142
red maple cambium borer, 449
Xylosandrus zimmermanni, 374
buds, shoots, roots—
Sciaphillus asperatus, 320
sucking insects—
Cryptococcus williamsi, 102
Maple, silver:
foliage —
flatheaded appletree borer, 281
maple bladdergall mite, 31
Vasates aceris-crummena, 31
whitemarked tussock moth, 227
bark, wood, twigs—
Columbian timber beetle, 369
flatheaded appletree borer, 281
Hemicoelus carinatus, 253
Ptilinus ruficornis, 254
sucking insects—
cottony maple scale, 97
woolly alder aphid, 80
Maple, sugar:
foliage—
Acleris chalybeana, 17\
American dagger moth, 236
Bruce spanworm, 189
Caloptilia packardella, 132
Choristoneura fractivittana, 169
European snout beetle, 320
forest tent caterpillar, 204
gouty vein midge, 442

green oak caterpillar, 219
maple leafcutter, 125
maple leafminer, 130
maple trumpet skeletonizer, 157
maple webworm, 180
orangehumped mapleworm, 220
Pandemis lamprosana, 161
Phigalia titea, 193
redhumped oakworm, 219
saddled prominent, 220
Sciaphillus asperatus, 320
Sparganothis acerivorana, 161
Vasates aceris-crummena, 31
bark. wood, twigs—
Columbian timber beetle, 369
Hemicoelus carinatus, 253
maple callus borer, 142
pitted ambrosia beetle, 370
Prilinus ruficornis, 254
red maple cambium borer, 449
sugar maple borer, 290
buds, shoots, roots—
European snout beetle, 320
Proteoteras moffatiana, 156
Sciaphillus asperatus, 320
sucking insects—
Cryptococcus williamsi, 102
maple mealybug, 92
Neosteingelia texana, 90
Norway maple aphid, 79
Mesquite:
Aneflus protensus, 308
banded ash borer, 294
mesquite borer, 287
Oncideres pustulatus, 301
Mimosa:
Epicauta torsa, 243
Moss, Spanish:
Quadraspidiotus tillandsiae, \21
Spanish moss orthezia, 90
Mountain-ash:
foliage—
apple-and-thorn skeletonizer, 145
Hedya chionosema, 160
Japanese beetle, 271
Lomgarapha vestaliata, 190
Machimia tentoriferella, 132
mountain-ash sawfly, 404
Paria quadrinotata, 266
pear sawfly, 401
pearleaf blister mite, 31
yellownecked caterpillar, 216
bark, wood, twigs—
American plum borer, 186
apple bark borer, 142
roundheaded appletree borer, 297
shothole borer, 356
flowers, seeds, fruits—
apple fruit moth, 140
Japanese beetle, 271
sucking insects—
birch lace bug, 65
Cerococcus kalmiae, 105
scurfy scale, 109
woolly apple aphid, 79
Mountain-ash, American:
mountain-ash sawfly, 404

555

Mountain-ash, European:
mountain-ash sawfly, 404
Mountain-ash, showy:
mountain- ash sawily, 404
Mountain-laurel:
Cerococcus kalmiae, 105
common falsepit scale, 104
laurel sphinx, 214
mulberry whitefly, 87
rhododendron stem borer, 300
Mulberry:
foliage—
io moth, 208
large maple spanworm, 199
Phyllophaga luctuosa, 268
bark, wood, twigs—
American plum borer, 186
Dorcaschema alternatum, 303
Aylocurus langstoni, 367
mulberry bark borer, 303
mulberry borer, 303
painted hickory borer, 287
Phloeotribus frontalis, 350
sucking insects—
common falsepit scale, 104
Comstock mealybug, 93
large hickory lecanium, 95
mulberry whitefly, 87
striped mealybug, 91
white peach scale, 118
sycamore lace bug, 64

N
Nettle:
comma butterfly, 173

O
Oak:
foliage—
Acordulecera spp., 381
Acronicta lithospila, 236
modica, 236
American dagger moth, 236
Anisota discolor, 210
peigleri, 209
pellucida, 210
Archips georgianus, 164
griseus, 164
Arge scapularis, 382
Argyrotaenia alisellana, 171
Asiatic oak weevil, 322
Attelabus bipustulatus, 319
bagworm, 126
Basicladus celibatus, 126
basswood leafminer, 263
Bibarrambla allenella, 132
Brachys ovatus, 285
browntail moth, 233
Bucculatrix packardella, 129
quinquenotella, 129
buck moth, 208
Caliroa spp., 401
Caloptilia quercinigrella, 132
Cameraria bethunella, 130
Catocala spp., 236
Cecidomyia poculum, 443
chainspotted geometer, 199
Charadra deridens, 237
crinkled flannel moth, 176
Cryptothelea gloverii, 127

556

Dasychira dorsipennata, 229
meridionalis, 229
tephra, 229
Datana angusi, 217
contracta, 217
Dichelonyx subvittata, 270
Dyseriocrania auricyanea, 124
elm spanworm, 196
Eotetranychus hicoriae, 31
fall cankerworm, 188
filament bearer, 195
forest tent caterpillar, 204
fruittree leafroller, 163
goldsmith beetle, 272
graybanded leafroller, 171
green oak caterpillar, 219
greenstriped mapleworm, 210
gregarious oak leafminer, 129
gypsy moth, 229
Haploa lecontei, 226
Heterocampa umbrata, 222
Homoeolabus analis, 319
Hyparpax aurora, 223
perophoroides, 223
imperial moth, 212
io moth, 208
Isa textula, 178
Lacosoma chiridota, 200
Lambdina fervidaria athasaria, 199
lappet moth, 206
large maple spanworm, 199
linden looper, 193
locust leafminer, 264
Lophodonta angulosa, 219
luna moth, 207
Machimia tentoriferella, 132
Macrarocampa marthesia, 222
maple webworm, 180
oak beauty, 195
oak skeletonizer, 128
Oligocentria lignicolor, 222
Oligonychus bicolor, 30
orangestriped oakworm, 209
oriental moth, 177
Pachybrachis carbonarius, 266
palmerworm, 137
Pandemis lamprosana, 161
Periclista spp., 402
Phyllophaga tristis, 268
pinkstriped oakworm, 209
Plagiometriona clavata, 266
polyphemus moth, 207
postoak locust, 51
Pristiphora chlorea, 404
Prolimacodes badia, 178
Psilocorsis quercicella, 132
reflexella, 132
saddleback looper, 192
Scudderia curvicauda, 52
Seirarctia echo, 226
solitary oak leafminer, 129
Sonoran tent caterpillar, 203
Sparganothis diluticostana, 161
spiny oakworm, 208
spotted tussock moth, 224
Syneta ferruginea, 266
Systena marginalis, 265
threelined leafroller, 161

Tortricidia flexuosa, 178 Pseudolucanus capreolus, 267

twobanded Japanese weevil, 322 Pseudopityophthorus asperulus, 363
twostriped walkingstick, 51 pruinosus, 363
Tymnes tricolor, 266 pubescens, 363
velleda lappet moth, 206 Pseudothysanoes lecontei, 356
walkingstick, 50 red oak borer, 291
white oak borer, 292 redheaded ash borer, 294
Xanthonia decemnotata, 266 sap beetles, 243
yellownecked caterpillar, 216 Saperda lateralis, 298
bark, wood, twigs— Scobicia bidentata, 257
Actenodes acornis, 285 Svnanthedon geliformis, 142
banded hickory borer, 289 sapygaeformis, 143
beech borer, 293 Thysanoes berchemiae, 367
black twig borer, 375 lobdelli, 367
Buprestis rufipes, 281 twig girdler, 301
Callirhytis quercusoperator, 424 twig pruner, 302
carpenterworm, 146 white oak borer, 292
chestnut bark borer, 304 Xiphydria tibialis, 411
Choragus zimmermanni, 318 Xyleborinus saxesensi, 374
Cinyra gracilipes, 285 Xvleborus affinis, 374
Columbian timber beetle, 369 ferrugineus, 374
Conotrachelus anaglyptius, 337 rubricollis, 374
Cupes concolor, 241 xylographus , 374
Dicerca obscura, 284 Xvloterinus politus, 371
dogwood borer, 141 buds, shoots, roots—
Enaphalodes atomarius, 292 Asiatic oak weevil, 322
Eucrada humeralis, 255 broadnecked root borer, 309
flat powderpost beetle, 299 Charadra deridens, 237
flatheaded sycamore-heartwood borer, eyespotted bud moth, 154
283 oak bark beetle, 362
Hadrobregmus notatus, 255 tilehorned prionus, 309
hardwood stump borer, 315 flowers, seeds, fruits—
hickory spiral borer, 279 Conotrachelus carinifer, 336
Hypothenemus dissimilis, 367 naso, 336
interstitialis, 368 posticatus, 336
quercus, 368 Curculio iowensis, 333
rotundicollis, 368 nasicus, 333
ivory-marked beetle, 302 proboscideus, 334
leopard moth, 145 sulcatulus, 334
Lichenophanes arminger, 257 oak bark beetle, 362
bicornis, 257 sucking insects—
little carpenterworm, 148 Alebra spp., 70
Marmara spp., 131 Asterolecanium minus, 105
Micracisella nanula, 367 quercicola, 105
opacicollis, 367 camphor scale, 118
Neoclytus fulguratus, 295 cottony maple scale, 97
jouteli jouteli, 295 Eutittix spp., 70
scutellaris, 295 false oak scale, 121
New York weevil, 331 fourhumped stink bug, 64
oak bark beetle, 362 giant bark aphid, 77
oak branch borer, 294 golden oak scale, 105
oak timberworm, 318 greater striped red oak aphid, 79
oak-bark scarrer, 292 island oak scale, 121
oak-stem borer, 303 Kermes galliformis, 101
Oberea ulmicola, 300 pubescens, 101
Orchesia castanea, 247 large hickory lecanium, 95
oriental wood borer, 258 Myzocallis discolor, 79
painted hickory borer, 287 melanocera, 79
Paranthrene simulans, 144 punctatus, 79
pecan carpenterworm, 146 oak lecanium, 96
Petalium bistriatum, 254 oak eriococcin, 104
seriatum, 254 Osborn scale, 112
Phymatodes varius, 306 Penthimia spp., 70
pigeon tremex, 410 periodical cicada, 74
Platypus compositus, 376 Platycotis vittata, 71
quadridentatus, 375 Quadraspidiotus socialis, 121
Pselaphorhynchites aeratus, 319 Quernaspis quercicola, 121
cyanellus, 319 sour-gum scale, 110

western oak scale, 121

eit)

Oak, black:
foliage—
Argyrotaenia quercifoliana, 170
flatheaded appletree borer, 281
giant walkingstick. 51
gouty oak gall. 423
large oak-apple gall, 425
Macrodiplosis foliora, 443
walkingstick, 50
bark. wood, twigs—
black carpenter ant, 429
Callirhytis cornigera. 141
flatheaded appletree borer. 281
horned oak gall. 423
red oak borer. 291
ribbed bud gall. 423
scarlet oak sawfly. 401
Thysanoes fimbricornis. 367
twolined chestnut borer. 277
Xvleborus validus. 374
sucking insects—
Kermes galliformis, 101
kingii, 101
Prociphilus longianus, 80
Oak, blackjack:
horned oak gall, 423
whitefringed beetle, 322
Oak, bur:
Bucculatrix recognita, 129
Kermes pubescens, 101
Melsheimer’s sackbearer, 200
oak lace bug. 64
oak webworm, 162
Profenusa lucifex, 401
redhumped oakworm, 219
Oak, Chapman:
Callirhytis floridana, 422
Oak, cherrybark:
Paranthrene simulans, 144
Oak. chestnut:
Liothrips umbripennis, 46
mulberry bark borer. 303
oak lace bug. 64
oak sapling borer. 293
Oak. laurel:
Amphibolips quercusfuliginosa, 425
Oak, live:
Archodontes melanopus melanopus, 315
claycolored leaf beetle. 266
Curculio fulvus, 333
Oiketicus abbotii, 126
Oak, northern pin:
Archips semiferanus, 163
Oak. northern red:
Archips semiferanus, 163
giant walkingstick, 51
Hemicoelus carinatus, 253
Phigalia titea, 193
walkingstick. 50
winter moth, 189
Oak, Nuttall:
Paranthrene simulans, 144
Oak, overcup:
spotworm borer. 279
white cutworm,. 237
white oak borer, 292
Oak, pin:
Argyrotaenia quercifoliana, 170

558

Caliroa petiolata, 402
Callirhytis cornigera, 141
Ectodemia heinrichi, 125
Eotetranychus querci, 31
gouty oak gall. 423
horned oak gall, 423
Macrodiplosis jfoliora, 443
obscure scale, 116
Parallelodiplosis florida. 445
scarlet oak sawfly. 401
Oak, post:
black carpenter ant, 429
Callirhytis floridana, 422
Phylloxera rilevi. 86
Oak, red:
foliage—
Argyrotaenia quercifoliana, 170
Bruce spanworm, 189
Choristoneura fractivittana. 169
definite-marked tussock moth, 228
giant walkingstick. 51
Hedya chionosema, 160
Japanagromyza viridula, 450
large oak-apple gall, 425
Macrodiplosis foliora, 443
oak leaftier, 171
oak skeletonizer, 128
Phigalia titea, 193
Polystepha pilulae. 445
Profenusa alumna, 401
Psilocorsis cryptolechiella, 132
succulent oak gall, 425
walkingstick, 50
willow flea weevil, 334
bark, wood, twigs—
black carpenter ant. 429
carpenterworm, 146
Chrysobothris viridiceps, 283
Paranthrene simulans, 144
red oak borer, 291
Thysanoes fimbricornis, 367
flowers, seeds, fruits—
Curculio sulcatulus, 334
sucking insects—
Cyrtolobus discoidalis. 71
Kermes andrei, 101
galliformis, 101
kingti. 101
Oak, runner: See: sand post
Oak, sand post:
Callirhytis floridana, 422
Seirarctia echo, 226
Oak, scarlet:
Archips semiferanus, 163
black carpenter ant, 429
Caliroa quercuscoccineae, 401
gouty oak gall, 423
large oak-apple gall, 425
red oak borer, 291
Oak, scrub:
Argyrotaenia quercifoliana, 170
Brachys tesselatus, 285
horned oak gall. 423
Oecanthus latipennis, 53
oak webworm, 162
Oak, swamp chestnut:
Coleophora querciella, 133

Oak, swamp white:
noxious oak gall, 424
Oak, turkey:
puss caterpillar, 175
Oak, water:

Archodontes melanopus melanopus, 315

Formosan horntail, 411
horned oak gall, 423
Oak, white:
toliage—
Argyrotaenia quercifoliana, 170
Bucculatrix luteella, 129
Coleophora querciella, 133
dark tussock moth, 228
flatheaded appletree borer, 281
giant walkingstick, 51
oak fig gall, 425
oak flake gall, 424
oak potato gall, 424
Phylloxera rilevi, 86
Profenusa lucifex, 401
redhumped oakworm, 219
scarlet oak sawfly, 401
solitary oak leafminer, 129
variable oakleaf caterpillar, 220
walkingstick, 50
bark, wood, twigs—
Agrilaxia flavimana, 285
banded ash borer, 294
black carpenter ant, 429
chestnut timberworm, 248
Chrysobothris azurea, 283
sexsignata, 283
Cinyra gracilipes, 285
flatheaded appletree borer, 281
Hemicoelus carinatus, 253
oak fig gall, 425
oak sapling borer, 293
oak-bark scaler, 309
oak-bark scarrer, 292
Paranthrene simulans, 144
Phylloxera rileyi, 86
Physocnemum violaceipenne, 307
powderpost beetle, 255
white oak borer, 292
buds, shoots, roots—
Callirhytis quercusfutilis, 424
sucking insects—
golden oak scale, 105
Kermes andrei, 101
pubescens, 101
oak lace bug, 64
Oleander:
oleander scale, 107
Olive:
oleander scale, 107
Orange, sour:
Cryptothelea gloverii, 127
Osage-orange:
fourspotted spider mite, 31
mulberry borer, 303

P

Pachysandra:
euonymus scale, 121
Palm:
elongate hemlock scale, 113
fern scale, 117
fiorinia hemlock scale, 113

Florida red scale, 111
palm leaf skeletonizer, 135
palmetto billbug, 337
royalpalm bug, 68
Xvleborus lecontei, 374
Palm, Canary Island date:
palm leaf skeletonizer, 135
Palmetto:
Carvobruchus gleditsiae, 259
Palmetto, cabbage:
palm leaf skeletonizer, 135
palmetto billbug, 337
Seirarctia echo, 226

Papaya:

Toxotrypana curvicauda, 448
Pawpaw:

zebra swallowtail, 173
Peach:

cherry leaf beetle, 262

lesser peachtree borer, !43

plum curculio, 336

quince curculio, 336

Sphinx drupiferarum, 214

sucking insects—
cottony maple scale, 97
Cuerna costalis, 71
European fruit lecanium, 96
Forbes scale, 119
Graphocephala versuta, 71
Homalodisca coagulata, 71
Oncometopia orbona, 7\
periodical cicada, 74
white peach scale, 118

Pear:

foliage—
apple-and-thorn skeletonizer, 145
Argyrotaenia quadrifasciana, 171
browntail moth, 233
cigar casebearer, 134
flatheaded appletree borer, 281
graybanded leafroller, 171
oriental moth, 177
palmerworm, 137
pear sawfly, 401
pearleaf blister mite, 31
unspotted leafminer, 131

bark, wood, twigs—
birch bark beetle, 362
flatheaded appletree borer, 281
leopard moth, 145

buds, shoots, roots—
Polydrusus impressifrons, 321
tilehorned prionus, 309

flowers, seeds, fruits—
cigar casebearer, 134
quince curculio, 336

sucking insects—
cottony maple scale, 97
oystershell scale, 114
San Jose scale, 120
woolly apple aphid, 79

Pecan:

foliage—
Acordulecera spp., 381
Acrobasis carvivorella, 181
Anthonomus suturalis, 337
Archips infumatanus, 164
Eotetranychus hicoriae, 31

599

hickory spiral borer, 279 bark, wood, twigs—

larger elm leaf beetle, 262 Agrilus fuscipennis, 280
Megaxyela langstoni, 378 American plum borer, 186
pecan cigar casebearer, 133 Dicerca obscura, 284
pecan leaf casebearer, 181 persimmon borer, 143
pecan leaf phylloxera, 86 Platypus compositus, 376
walnut caterpillar, 217 powderpost beetle, 255
walnut sphinx, 215 redheaded ash borer, 294
whitefringed beetle, 322 redshouldered shothole borer, 257
bark, wood, twigs— twig girdler, 301
Chrysobothris adelpha, 283 Xyleborinus saxeseni, 374
dogwood borer, 141 Xyleborus affinis, 379
hickory bark beetle, 355 buds, shoots, roots—
hickory borer, 293 Archodontes melanopus melanopus,
hickory spiral borer, 279 by be
Lichenophanes bicornis, 257 persimmon borer, 143
pecan carpenterworm, 146 sucking insects—
pecan phylloxera, 86 camphor scale, 118
Platypus compositus, 376 Cerococcus kalmiae, 105
powderpost beetle. 255 common falsepit scaie, 104
redshouldered shothole borer, 257 cottony maple leaf scale, 98
Synanthedon geliformis, 142 persimmon psylla, 77
twig girdler, 301 Parrott scale, 105
twig pruner, 302 white peach scale, 118
Xyleborinus saxeseni, 374 Phlox:
buds, shoots, roots— Asterolecanium arabidis, 106
Acrobasis exsulella, 181 Pieris:
broadnecked root borer, 309 azalea bark scale, 103
pecan carpenterworm, 146 Pine:
pecan cigar casebearer, 133 foliage—
pecan leaf casebearer, 181 Asiatic garden beetle, 270
pecan nut casebearer, 181 bagworm, 126
Prionus spp., 309 Basicladus celibatus, 126
tilehorned prionus, 309 Brachyderes incanus, 321
walnut shoot moth, 181 Citheronia sepulcralis, 211
flowers, seeds, fruits— Dichelonyx subvittata, 270
Curculio spp., 333 Diplotaxis liberata, 270
hickory shuckworm, 158 gall midge, 442
pecan nut casebearer, 181 Gilpinia frutetorum, 396
pecan weevil, 333 gypsy moth, 229
sucking insects— Hubbellia marginifera, 52
black pecan aphid, 79 imperial moth, 212
camphor scale, 118 Lapara coniferarum, 215
Dysmicoccus morrisoni, 94 nesting-pine sawfly, 380
giant bark aphid, 77 Oecanthus pini, 53
obscure scale. 116 Oligonychus milleri, 30
Parrott scale, 105 pales weevil, 323
pecan spittlebug. 74 Pero morrisonaria, 197
Persea: Phyllophaga forsteri, 269
elongate hemlock scale, 113 micans, 269
fiorinia hemlock scale, 113 _prunina, 269
striped mealybug, 91 pine bud mite, 32
redbay scale, 107 pine chafer, 270
Persimmon: pine colaspis, 263
foliage— pine false webworm, 378
bagworm. 126 postoak locust, 51
hickory horned devil, 211 spruce budworm, 164
imperial moth, 212 Texas leafcutting ant, 433
luna moth. 207 tufted white pine caterpillar, 237
Phyllophaga crenulata, 269 zebra caterpillar, 237
luctuosa, 268 bark, wood, twigs—
prununculina, 268 balsam fir bark beetle; 357
tristis. 268 black and red horntail, 410
redhumped caterpillar, 222 blue horntail, 410
regal moth. 211 Brachyleptura vagans, 306
Seirarctia echo. 226 Buprestis maculipennis, 281
variable oakleaf caterpillar, 220 striata, 281

560

Callidium antennatum antennatum,
304
violaceum, 304
carpenter bee, 439
Carphoborus bifurcus, 352
Cecidomyia piniinopis, 443
Chalcophora georgiana, 283
liberta, 283
Chrysobothris floricola, 283
scabripennis, 283
Cossonus corticola, 337
Crypturgus alutaceus, 362
pusillus, 362
Cupes concolor, 241
deodar weevil, 330
Dicerca punctulata, 284
dogwood borer, 141
Dryocoetes affaber, 361
autographus, 361
four-eyed spruce bark beetle, 352
gall midges, 442
gloomy borer, 284
gouty pitch midge, 443
hairy pine borer, 307
Hylastes exilis, 340
porculus, 340
tenuis, 340
Hylurgops pinifex, 340
Ips spp., 358
large flatheaded pine heartwood borer,
283
Magdalis perforatus, 332
Melanophila acuminata, 284
aeneola, 284
notata, 284
Monarthrum fasicatum, 370
Monochamus carolinensis, 315
Neacanthocinus obsoletus, 304
Neoclytus muricatulus muricatulus,
295
old house borer, 316
Orthotomicus caelatus, 358
pales weevil, 323
Phloeosinus pini, 352
pine engraver, 359
pitcheating weevil, 326
Pityogenes hopkinsi, 357
meridianus, 357
plagiatus, 357
Pityophthorus annectens, 364
balsameus, 364
biovalis, 364
cariniceps, 364
pulchellus, 364
pulicarius, 363
Pygoleptura nigrella, 306
red turpentine beetle, 347
Sirex juvencus, 410
six-spined engraver, 358
small southern pine engraver, 359
southern pine beetle, 343
southern pine engraver, 359
. southern pine sawyer, 313
spotted pine sawyer, 315
Stephanopachys cribratus, 257
densus, 257
hispidulus, 257
substriatus, 257

Stictoleptura canadensis, 304
Trvpodendron rufitarsis, 371
Tylocerina nodosa, 304
white pine weevil, 328
Xyleborus dispar, 373
ferrugineus, 373
intrusus, 374
pubescens, 374
Xyloterinus politus, 371
Xvlotrechus sagittatus sagittatus, 311

buds, shoots, roots—

Brachyderes incanus, 321
Hylastes exilis, 340

porculus, 340

tenuis, 340
pine chafer, 270
pitcheating weevil, 326
Polyphylla occidentalis, 269
southern pine root weevil, 326
Xyleborus xylographus, 374

flowers, seeds, fruits—

bark beetles, 339

eastern pine seedworm, 159
longleaf pine seedworm, 159
Moodna ostrinella, 187
Nepytia pellucidaria, 198
Phyllophaga micans, 269

red pine cone borer, 155
southern pine coneworm, 183

sucking insects—

Alabama pine scale, 89
black pineleaf scale, 116
cryptomeria scale, 107
dogwood spittlebug, 74
Dysmicoccus obesus, 94
Florida wax scale, 94
McComb scale, 113
pine needle scale, 109
pine twig gall scale, 89
slash pine flower thrips, 46
Trioza tripunctata, 77

Pine, Austrian:
foliage—

Acantholyda angulata, 380
European pine sawfly, 391
nesting-pine sawfly, 380
pine bud moth, 136

pine false webworm, 378

bark, wood, twigs—

black turpentine beetle, 346
northern pine weevil, 329
pine root collar weevil, 324
pitch mass borer, 142
Zimmerman pine moth, 182

buds, shoots, roots—

Battaristis vittella, 138

eastern pine shoot borer, 154
European pine shoot moth, 149
pine root collar weevil, 324

sucking insects—

Eulachnus agilis, 78
Leptoglossus occidentalis, 67
pine tortoise scale, 99

Pine, Caribbean:
slash pine sawfly, 394
subtropical pine tip moth, 153
Pine, Chinese:
pine tortoise scale, 99
red pine scale, 88

561

Pine, Choctawhatchee sand:
sand pine sawily. 386
Pine, Corsican:
pitch pine tip moth, 151
Pine. eastern white:
foliage—
Acantholyda angulata, 380
luteomaculata, 380
chainspotted geometer. 199
Eacles imperialis pini, 212
Eufidonia notataria, 191
European pine sawily. 391
hairy leaf beetle. 266
introduced pine sawfl
Magdalis austera, 33
hispoides, 333
Melanoplus punctulatus, 52
pine bud moth, 136
pine false webworm, 378
pine tree sphinx, 214
pine tube moth, 169
pine tussock moth, 228
pine webworm, 179
red pine sawfly, 389
redheaded pine sawfly, 384
Semiothisa bisignata, 191
small pine looper, 190
spruce-fir looper, 191
Swaine jack pine sawily, 388
white pine sawfly, 390
Xenotemna pallorana, 172
bark, wood. twigs—
Allegheny mound ant, 434
black carpenter ant, 429
carpenter bee, 439
Chrysobothris blanchardi, 283
dentipes, 283
harrisi, 283

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fir coneworm, 183
hemlock borer. 284
Aylobius congener, 326
Ips latidens, 361
lodgepole pine beetle, 350
Marmara fasciella, 130
northeastern sawyer, 314
northern pine weevil, 329
pine root collar weevil, 324
Pissodes affinis, 331
pitch mass borer, 142
Pitvogenes hopkinsi, 357
Pityophthorus opaculus, 363
ramiperda, 364
Sirex nigricornis, 410
southern pine beetle, 343
white pine weevil, 328
whitespotted sawyer. 313
buds, shoots. roots—
eastern pine shoot borer, 154
European pine shoot moth. 149
fir coneworm, 183
pine root collar weevil, 324
pine root tip weevil, 326
flowers, seeds, fruits—
fir coneworm, 183
white pine cone beetle, 365
white pine cone borer, 156

sucking insects—
Canadian pine scale. 89
Eulachnus agilis, 78
leaffooted pine seed bug. 67
pine bark adelgid. 84
pine spittlebug. 72
Pitedia uhleri, 64
powdery pine needle aphid. 78
Prociphilus bumelia, 78
red spruce adelgid, 86
Saratoga spittlebug, 73
shieldbacked pine seed bug. 6
white pine aphid. 78
Pine, jack:
Toliage—
Acantholyda angulata, 380
luteomaculata, 380

ee)

brownheaded jack pine sawily, 391

Cephalcia fulviceps, 380
chameleon caterpillar, 237
Dichelonyx albicollis, 270
Diplotaxis sordida, 270
Eacles imperialis pini, 212
European pine sawily, 391
hemlock looper. 198
introduced pine sawfly, 395
jack pine budworm, 167
jack pine sawfly, 386
Neodiprion compar, 394
maurus, 394
nigroscutum, 394
pratti paradoxicus, 387
nesting-pine sawfly, 380
pine bud moth, 136
pine needle sheathminer, 140
pine needleminer, 136
pine tree sphinx, 214
pine tussock moth, 228
pine webworm, 179

Protoboarmia porcelaria indicataria,

200
red pine sawfly, 389
redheaded jack pine sawfly, 391
redheaded pine sawfly. 384
small pine looper, 190
Sparganothis tristriata, 161
Swaine jack pine sawfly, 388
whitespotted sawyer, 313
Xenotemna pallorana, 172
Xyela bakeri, 378

obscura, 378

bark, wood. twigs—

Chrysobothris orono, 282
fir coneworm, 183
Ips latidens, 361

perroti, 361
lodgepole pine beetle. 350
Neacanthocinus pusillus, 304
northern pine weevil, 329
pine root collar weevil, 324
Pissodes affinis, 331
Pityogenes hopkinsi, 357
white pine weevil, 328
whitespotted sawyer. 313

buds. shoots, roots—

eastern pine shoot borer. 154

European pine shoot moth. 149
jack pine tip beetle, 366

northern pitch twig moth, 154
Petrova pallipennis, 154
pine root tip weevil, 326
Rhyacionia sonia, 153
western pine tip moth, 152
white pine weevil, 328
flowers, seeds, fruits—
Cimberis elongatus, 319
fir coneworm, 183
red pine cone beetle, 365
sucking insects—
Cinara pergandei, 78
pinivora, 78
watsoni, 78
pine spittlebug, 72
pine tortoise scale, 99
Saratoga spittlebug, 73
shieldbacked pine seed bug, 63
slash pine flower thrips, 46
woolly pine needle aphid, 78
Pine, Japanese black:
European pine shoot moth, 149
red pine scale, 88
Pine, Japanese red:
Acantholyda angulata, 380
European pine sawfly, 391
European pine shoot moth, 149
nesting-pine sawfly, 380
pine false webworm, 378
red pine sawfly, 389
red pine scale, 88
Pine, Japanese white:
pine spittlebug, 72
Pine, loblolly:
follage—
Acantholyda apicalis, 380

Nantucket pine tip moth, 151
pales weevil, 323

Petrova taedana, 153

pitch pine tip moth, 151
pitcheating weevil, 326
Rhyacionia aktita, 153
Sparganothis sulfureana, 161
subtropical pine tip moth, 153

flowers, seeds, fruits—

Atlantic pine coneworm, 186
blister coneworm, 184
loblolly pine coneworm, 186
longleaf pine seedworm, 159
pine conelet looper, 198

slash pine seedworm, 159
south coastal coneworm, 185
southern pine coneworm, 183
webbing coneworm, 184
white pine cone beetle, 365

sucking insects—

Cinara atlantica, 78

pergandei, 78

pinivora, 78

taedae, 78

watsoni, 78
leaffooted pine seed bug, 67
Oracella acuta, 94
pine spittlebug, 72
pine tortoise scale, 99
pine twig gall scale, 89
powdery pine needle aphid, 78
Saratoga spittlebug, 73
shieldbacked pine seed bug, 63
speckled pine needle aphid, 78
Virginia pine scale, 100
woolly pine scale, 98

blackheaded pine sawfly, 393
eastern pine looper, 198
Lapara coniferarum, 215
loblolly pine sawfly, 387
Neodiprion abbotii, 390
hetricki, 394
Phyllophaga luctuosa, 268
prununculina, 268
pine chafer, 270
pine needleminer, 136
pine shoot gall sawfly, 378
pine webworm, 179
redheaded pine sawfly, 384

spotted loblolly pine sawfly, 387

turpentine borer, 280

bark, wood, twigs—

black turpentine beetle, 346
Buprestis lineata, 281
deodar weevil, 330
HAylastes salebrosus, 340
Orthotomicus caelatus, 358
pitcheating weevil, 326
Pityoborus comatus, 364
southern pine beetle, 343
Trypodendron scabricollis, 371
turpentine borer, 280
Xvleborinus saxeseni, 374

buds, shoots, roots—

Battaristis vittella, 138
deodar weevil, 330
Hylastes salebrosus, 340
lesser cornstalk borer, 186

Pine, lodgepole:

lodgepole pine beetle, 350
northern pitch twig moth, 154
striped pine scale, 100
Xvlotrechus undulatus, 312

Pine, longleaf:

follage—

blackheaded pine sawfly, 393

Coleotechnites chillcotti, 136
Lapara coniferarum, 215
Neodiprion abbotii, 390
compar, 394
Oligonvchus cunliffei, 31
Phyllophaga luctuosa, 268
prununculina, 268
pine needleminer, 136
pine webworm, 179
redheaded pine sawfly, 384
bark, wood, twigs—
black turpentine beetle, 346
Buprestis lineata, 281
Chrysobothris dentipes, 283
deodar weevil, 330
Hylastes salebrosus, 340
Pityoborus comatus, 364
southern pine beetle, 343
turpentine borer, 280
buds, shoots, roots—
blister coneworm, 184
deodar weevil, 330

European pine shoot moth, 149

Hylastes salebrosus, 340

563

loblolly pine coneworm, 186
subtropical pine tip moth, 153

flowers, seeds, fruits—

Battaristis vittella, 138
blister coneworm, 184
Holcocera lepidophaga, 133
loblolly pine coneworm, 186
longleaf pine seedworm, 159
slash pine seedworm, 159
south coastal coneworm, 185
southern pine coneworm, 183
webbing coneworm, 184
Xyela bakeri, 378

minor, 378

obscura, 378

sucking insects—

Brochymena carolinensis, 64
Cinara atlantica, 78
leaffooted pine seed bug, 67 |
shieldbacked pine seed bug, 63
Virginia pine scale, 100
woolly pine scale, 98

Pine, Ocala sand:

sand pine sawfly, 386
Pine, pitch:

foliage—

Acantholyda angulata, 380

Choristoneura pinus maritima, 168

eastern pine looper, 198
European pine sawfly, 391
Neodiprion pinusrigidae, 394
pratti paradoxicus, 387
nesting-pine sawfly, 380
pine needleminer, 136
pine tree sphinx, 214
pine webworm, 179
redheaded pine sawfly, 384
Semiothisa granitata, 191
turpentine borer, 280
white pine sawfly, 390

bark, wood, twigs—

black carpenter ant, 429
black turpentine beetle, 346
Buprestis lineata, 281
salisburyensis, 281
Chrysobothris blanchardi, 283
harrisi, 283
pusilla, 282, 283
sexsignata, 283
gouty pitch midge, 443
northern pine weevil, 329
pine gall weevil, 331
pine root collar weevil, 324
Sirex edwardsii, 410
six-spined engraver, 358
southern pine beetle, 343
southern pine engraver, 359
turpentine borer, 280

buds, shoots, roots—

564

European pine shoot moth, 149
Nantucket pine tip moth, 151
pales weevil, 323

pine gall weevil, 331

pine root collar weevil, 324
pitch pine tip moth, 151

pitch twig moth, 153
Rhyacionia aktita, 153

sucking insects—

Cinara atlantica, 78

pergandei, 78

pinivora, 78

taedae, 78

watsoni, 78
leaffooted pine seed bug, 67
pine spittlebug, 72
pine twig gall scale, 89
Pineus coloradensis, 86
powdery pine needle aphid, 78
Saratoga spittlebug, 73
striped pine scale, 100
woolly pine scale, 98

Pine, pond:
blackheaded pine sawfly, 393
Cinara atlantica, 78

pinivora, 78
taedae, 78
watsoni, 78

eastern pine looper, 198

Lytta polita, 243

Neodiprion hetricki, 394

speckled pine needle aphid, 78
Pine, ponderosa:

European pine sawfly, 391

European pine shoot moth, 149

pine twig gall scale, 89

western pine tip moth, 152
Pine, red:

foliage—

Acantholyda pini, 380
Cephalcia fulviceps, 380
marginata, 380
chainspotted geometer, 199
Diplotaxis sordida, 270
eastern pine looper, 198
European pine sawfly, 391
European pine shoot moth, 149
Gilpinia frutetorum, 396
introduced pine sawfly, 395
jack pine budworm, 167
jack pine sawfly, 386
Neodiprion abbotii, 390
compar, 394
nesting-pine sawfly, 380
pine bud moth, 136
pine candle moth, 137
pine false webworm, 378
pine needleminer, 136
pine tree sphinx, 214
pine tussock moth, 228
pine webworm, 179
red pine needle midge, 443
red pine sawfly, 389
redheaded pine sawfly, 384
Sparganothis acerivorana, 161
strawberry root weevil, 321
Swaine jack pine sawfly, 388
western pine tip moth. 152
white pine sawfly, 390
Xenotemna pallorana, 172

bark, wood, twigs—

Allegheny mound ant, 434
Chrysobothris orono, 282
fir coneworm, 183
Hylobius congener, 326
Ips latidens, 361

perroti, 361
lodgepole pine beetle, 350
Neacanthocinus pusillus, 304
northern pine weevil, 329
pine gall weevil, 331
pine root collar weevil, 324
Pissodes affinis, 331
Pityogenes hopkinsi, 357
Pityophthorus ramiperda, 364
southern pine beetle, 343
white pine weevil, 328
whitespotted sawyer, 313
buds, shoots, roots—
Dioryctria resinosella, 186
eastern pine shoot borer, 154
European pine shoot moth, 149
Lambdina pellucidaria, 198
pine bud moth, 136
pine root collar weevil, 324
pine root tip weevil, 326
pine tree sphinx, 214
pitch pine tip moth, 151
red pine cone beetle, 365
Sparganothis sulfureana, 161
Zimmermann pine moth, 182
flowers, seeds, fruits—
Dioryctria resinosella, 186
fir coneworm, 183
webbing coneworm, 184
sucking insects—
Cinara pinea, 78
pinivora, 78
watsoni, 78
Eulachnus agilis, 78
pine needle scale, 109
pine tortoise scale, 99
pine twig gall scale, 89
Pineus coloradensis, 86
powdery pine needle aphid, 78
red pine scale, 88
Saratoga spittlebug, 73
shieldbacked pine seed bug, 63
striped pine scale, 100
woolly pine needle aphid, 78
Pine, sand:
Acantholyda circumcincta, 380
floridana, 380
blackheaded pine sawfly, 393
Cinara atlantica, 78
pergandei, 78
pinivora, 78
taedae, 78
watsoni, 78
Neodiprion virginianus, 391
pine conelet looper, 198
sand pine sawfly, 386
shieldbacked pine seed bug, 63
Xvela dodgei, 378
Pine, Scotch:
foliage—
European pine sawfly, 391
Gilpinia frutetorum, 396
introduced pine sawfly, 395
jack pine sawfly, 386
Magdalis austera substriga, 333
Neodiprion pratti paradoxicus, 387
pine candle moth, 137
pine false webworm, 378

pine needleminer, 136

pine webworm, 179
Sparganothis acerivorana, 161
strawberry root weevil, 321
Swaine jack pine sawfly, 388
Xenotemna pallorana, 172

bark, wood, twigs—

Allegheny mound ant, 434
Hylobius congener, 326

Ips latidens, 361

northern pine weevil, 329
pine root collar weevil, 324
pine spittlebug, 72

Pissodes affinis, 331

pitch mass borer, 142
Zimmermann pine moth, 182

buds, shoots, roots—

Couper’s collar weevil, 326
eastern pine shoot borer, 154
European pine shoot moth, 149
Nantucket pine tip moth, 151
northern pitch twig moth, 154
pine bud moth, 136

pine candle moth, 137

pine root collar weevil, 324
pine root tip weevil, 326
pitch pine tip moth, 151
Warren’s collar weevil, 326
western pine tip moth, 152

flowers, seeds, fruits—

Battaristis vittella, 138
webbing coneworm, 184

sucking insects—

Cinara atlantica, 78
pinea, 78
watsoni, 78
Eulachnus agilis, 78
meadow spittlebug, 74
pine spittlebug, 72
pine tortoise scale, 99
powdery pine needle aphid, 78
striped pine scale, 100

Pine, shortleaf:
foliage—

blackheaded pine sawfly, 393
eastern pine looper, 198
European pine sawfly, 391
hairy leaf beetle, 266
introduced pine sawfly, 395
loblolly pine sawfly, 387
Nantucket pine tip moth, 151
Neodiprion abbotii, 390
pratti paradoxicus, 387
pine needleminer, 136
pine webworm, 179
redheaded pine sawfly, 384
spotted loblolly pine sawfly, 387
turpentine borer, 280
white pine sawfly, 390

bark, wood, twigs—

Chrysobothris dentipes, 283
pusilla, 282, 283

deodar weevil, 330

Gnathotrichus materiarius, 372

Hylastes salebrosus, 340

Ips avulsus, 359

northern pine weevil, 329

pitcheating weevil, 326

Pityoborus comatus, 364
Platypus flavicornis, 375
Sirex nigricornis, 410
southern pine beetle, 343
turpentine borer, 280
Xyleborinus saxeseni, 374

buds, shoots, roots—

blister coneworm, 184
deodar weevil, 330
Nantucket pine tip moth, 151
pales weevil, 323

Petrova houseri, 154

pine shoot gall sawfly, 378
pitcheating weevil, 326

flowers, seeds, fruits—

blister coneworm, 184
Lytta polita, 243
Nantucket pine tip moth, 151
pine conelet looper, 198
shortleaf pine cone borer, 155
webbing coneworm, 184
Xvela bakeri, 378
sucking insects—
Cinara atlantica, 78
pergandei, 78
pinivora, 78
watsoni, 78
leaffooted pine seed bug, 67
pine tortoise scale, 99
pine twig gall scale, 89
powdery pine needle aphid, 78
shieldbacked pine seed bud, 63
striped pine scale, 100

Pine, slash:
foliage—

Acantholyda floridana, 380
Apantesis radians, 226
arboreal short-tailed cricket, 53
blackheaded pine sawfly, 393
Nantucket pine tip moth, 151
Neodiprion abbotii, 390

pine webworm, 179

redheaded pine sawfly, 384
slash pine sawfly, 394
turpentine borer, 280

bark, wood, twigs—

black turpentine beetle, 346
pitcheating weevil, 326
Pityoborus comatus, 364
southern pine beetle, 343
turpentine borer, 280

buds, shoots, roots—

blister coneworm, 184
Nantucket pine tip moth, 151
pine shoot gall sawfly, 378
pitch pine tip moth, 151
pitcheating weevil, 326
Rhyacionia aktita, 153

slash pine flower thrips, 46
subtropical pine tip moth, 153

flowers, seeds, fruits—

blister coneworm, 184
Holcocera lepidophaga, 133
loblolly pine coneworm, 186
longleaf pine seedworm, 159
pine conelet looper, 198
slash pine flower thrips, 46
slash pine seedworm, 159

south coastal coneworm, 185
southern pine coneworm, 183

Xyela bakeri, 378
minor, 378
obscura, 378

sucking insects—

Brochymena carolinensis, 64

Cinara atlantica, 78
pinivora, 78
taedae, 78
leaffooted pine seed bug, 6
pine spittlebug, 72

powdery pine needle aphid,

shieldbacked pine seed bug

speckled pine needle aphid,

Pine, Sonderegger:
blackheaded pine sawfly, 393
Pine, South Florida slash:
slash pine seedworm, 159
Pine, spruce:
blackheaded pine sawfly, 393
pine twig gall scale, 89
southern pine beetle, 343
Pine, Swiss mountain:
follage—
European pine sawfly, 391
introduced pine sawfly, 395
pine bud moth, 136
pine candle moth, 137
pine false webworm, 378
red pine sawfly, 389
redheaded pine sawfly, 384
Strawberry root weevil, 321
white pine sawfly, 390
bark, wood, twigs—
pine root collar weevil, 324
buds, shoots, roots—
Battaristis vittella, 138

7

78
Oo
78

eastern pine shoot borer, 154

sucking insects—
pine tortoise scale, 99
striped pine scale, 100
woolly pine scale, 98
Pine, Table Mountain:
Cinara atlantica, 78
pinivora, 78
taedae, 78
watsoni, 78
European pine sawfly, 391
leaffooted pine seed bug, 67
mountain pine coneworm, 186
northern pine weevil, 329
pine twig gall scale, 89
woolly pine scale, 98
Pine, Virginia:
foliage—

Choristoneura pinus maritima, 168

Dendrotettix australis, 52
hairy leaf beetle, 266
introduced pine sawfly, 395
Janetiella coloradensis, 445
Neodiprion virginianus, 391
pine needleminer, 136
pine webworm, 179
Semiothisa granitata, 191
Virginia pine sawfly, 385
bark, wood, twigs—
Buprestis lineata, 281

Chrysobothris blanchardi, 283
dentipes, 283
harrisi, 283
gouty pitch midge, 443
northern pine weevil, 329
pine gall weevil, 331
Sirex nigricornis, 410
southern pine beetle, 343
buds, shoots, roots—
European pine shoot moth, 149
Nantucket pine tip moth, 151
pine gall weevil, 331
pitch pine tip moth, 151
red pine cone beetle, 365
flowers, seeds, fruits—
Cimberis pilosus, 319
leaffooted pine seed bug, 67
red pine cone beetle, 365
shieldbacked pine seed bug, 63
webbing coneworm, 184
Xvela bakeri, 378
minor, 378
obscura, 378
stvrax, 378
sucking insects—
Cinara atlantica, 78
pergandei, 78
pinea, 78
pinivora, 78
taedae, 78
watsoni, 78
pine spittlebug, 72
pine tortoise scale, 99
pine twig gall scale, 89
powdery pine needle aphid, 78
Saratoga spittlebug, 73
striped pine scale, 100
Virginia pine scale, 100
Pine, white:
Acantholyda luteomaculata, 380
Chrysobothris trinervia, 283
Lepturopsis biforis, 306
pales weevil, 323
pine bark adelgid, 84
pine leaf adelgid, 86
pine tortoise scale, 99
strawberry root weevil, 321
Xvela alpigena, 378
Planetree: See: sycamore
Plum:
follage—
apple-and-thorn skeletonizer, 145
Arge clavicornis, 382
Argyrotaenia quadrifasciana, 171
browntail moth, 233
cherry leaf beetle, 262
cynthia moth, 206
hickory leafroller, 170
oriental moth, 177
palmerworm, 137
pear sawfly, 401
Phyllonorycter crataegella, 130
Polydrusus impressifrons, 321
Sphinx drupiferarum, 214
Sterictiphora spp., 382
twinspot sphinx, 215
unspotted leafminer, 131
western tent caterpillar, 204

bark, wood, twigs—
American plum borer, 186
banded hickory borer, 289
leopard moth, 145
buds, shoots, roots—
Polydrusus impressifrons, 321
flowers, seeds, fruits—
Acalitus phloeococotes, 32
apple fruit moth, 140
apple maggot, 448
cigar casebearer, 134
plum curculio, 336
sucking insects—
cottony maple scale, 97
European fruit lecanium, 96
Plum, American:
lesser peachtree borer, 143
plum webspinning sawfly, 380
prairie tent caterpillar, 203
Plum, wild: See: plum, American
Poison-ivy:
Arge humeralis, 382
Episimus argutanus, 148
Pondcypress:
baldcypress coneworm, 185
south coastal coneworm, 185
Poplar:
follage—
Acronicta distans, 236
Anacampsis innocuella, 137
Archips purpuranus, 164
aspen blotchminer, 130
aspen leaf beetle, 260
aspen leafminer, 131
bagworm, 126
blinded sphinx, 215
Bruce spanworm, 189
Cabera variolaria, 190
Caloptilia stigmatella, 132
cecropia moth, 206
Cerura borealis, 223
cinerea, 223
multiscripta canadensis, 223
occidentalis, 223
chainspotted geometer, 199
Chrysomela knabi, 260
Clostera albosigma, 216
apicalis, 216
brucei, 216
strigosa, 216
Compton tortoiseshell, 174
cottonwood dagger moth, 236
cottonwood leaf beetle, 260
Dasychira obliquata, 229
vagans, 229
elm sawfly, 382
Eotetranychus populi, 31
weldoni, 31
Epinotia solandriana, 158
Evora hemidesma, 149
false-sphinx, 218
forest tent caterpillar, 204
fourspotted spider mite, 31
fruittree leafroller, 163
giant walkingstick, 51
Gluphisia septentrionalis, 223
goldsmith beetle, 272
gypsy moth, 229

hickory tussock moth, 223 black carpenter ant, 429

Hyperaeschra stragula, 218 bronze poplar borer, 279

imported willow leaf beetle, 264 cottonwood borer, 289

io moth, 208 cottonwood twig borer, 156

Tridopsis larvaria, 192 Dicerca tenebrica, 284

lappet moth, 206 Hexomyza schineri, 450

large aspen tortrix, 168 hornet moth, 141

large maple spanworm, 199 leopard moth, 145

Macrarocampa marthesia, 222 linden borer, 297

maple webworm, 180 Micracis swainei, 366

mourningcloak butterfly, 174 oak timberworm, 318

Nematus fulvicrus, 406 Oberea schaumii, 300
hudsoniimagnus, 406 Platypus compositus, 376
limbatus, 406 Poecilonota cyanipes, 285
salicisodoratus, 405 poplar borer, 295

Nevada buck moth, 208 poplar carpenterworm, 146

notched-wing geometer, 196 poplar clearwing moth, 141

Notodonta simplaria, 223 poplar vagabond aphid, 80

obliquebanded leafroller, 168 poplar-and-willow borer, 334

oriental moth, 177 poplar-butt borer, 311

pepper-and-salt moth, 200 poplar-gall saperda, 298

Pero morrisonaria, 197 Saperda moesta, 298

Phyllonorycter salicifoliella, 130 sapwood timberworm, 248

Plagodis serinaria, 195 tarnished plant bug, 66

Polydrusus impressifrons, 321 Trypodendron retusum, 371

poplar dagger moth, 236 twig girdler, 301

poplar leafmining sawfly, 401 Xylotrechus annosus annosus, 312

poplar petiole gall aphid, 80 buds, shoots, roots—

poplar petiole gall moth, 125 Agrilus horni, 279

poplar sawfly, 406 broadnecked root borer, 309

poplar tentmaker, 215 cottonwood borer, 289

Protoboarmia porcelaria indicataria, Eriophyes parapopuli, 31
200 hornet moth, 141

Pseudexentera oregonana, 160 poplar clearwing moth, 141

Pseudosciaphila duplex, 160 poplar-butt borer, 311

Psilocorsis reflexella, 132 sucking insects—

purplish-brown looper, 199 azalea bark scale, 103

red spotted purple, 175 black willow aphid, 79

redhumped caterpillar, 222 black willow scale, 111

saddleback looper, 192 Chaitophorus populicola, 79

satin moth, 233 stevensis, 79

Schizura ipomoeae, 223 Comstock mealybug, 93

leptinoides, 223 Corythucha elegans, 65

smeared dagger moth, 236 cottony maple scale, 97

Sparganothis reticulatana, 161 dentate scale, 123

Sphinx luscitiosa, 214 Idiocerus spp., 70

spiny-elm caterpillar, 174 Macropsis spp., 70

spotted tussock moth, 224 oystershell scale, 114

stout looper, 195 Pterocomma populifoliae, 79

tiger swallowtail, 173 Telamona decorata, 71

Trichiosoma triangulum, 383 Tuberolachnus salignus, 79

twinspot sphinx, 215 willow scale, 119

twobanded Japanese weevil, 322 willow scurfy scale, 111

unicorn caterpillar, 223 Poplar, balsam:

velleda lappet moth. 206 Corythucha elegans, 65

viceroy, 175 large aspen tortrix, 168

western tent caterpillar, 204 Poplar, Lombardy:

white admiral. 175 Japanese beetle, 271

willow flea weevil, 334 Prickly-ash:

willow sawfly. 405 Agonopterix nigrinotella, 132

willow shoot sawfly, 412 Papilio cresphontes, 173

winter moth, 189 Privet:

yellow-marked caterpillar, 237 Archips rosanus, 161

Zengophora scutellaris, 265 ash and privet borer, 308

bark, wood, twigs— Asterolecanium arabidis, 106

Agrilus horni, 279 lilac borer, 143

American plum borer, 186 lilac leafminer, 131

aspen carpenterworm, 146 white prunicola scale, 118

568

Prunus spp.:
Caliroa spp., 401
elm scurfy scale, 108
Forbes scale, 119
oystershell scale, 114
San Jose scale, 120
scurfy scale, 109
taxus mealybug, 91
twocirculi mealybug, 92
white prunicola scale, 118
Pyracantha:
Corythucha cydoniae, 65
San Jose scale, 120

Q
Quince:
apple flea weevil, 334
cigar casebearer, 134
pear sawfly, 401
Phyllonorycter crataegella, 130
quince curculio, 336
twocirculi mealybug, 92
unspotted leafminer, 131

R

Rattan-vine:
Thysanoes berchemiae, 367
Redbud:
foliage—
Norape ovina, 176
twobanded Japanese weevil, 322
bark, wood, twigs—
Hypothenemus chapuisi, 368
dissimilis, 367
Micracis suturalis, 367
swainei, 366
Micracisella nanula, 367
opacicollis, 367
Pityophthorus lautus, 363
Ptosima gibbicollis, 285
buds, shoots, roots—
redbud leaffolder, 138
flowers, seeds, fruits—
Gibbobruchus mimus, 259
sucking insects—
periodical cicada, 74
Prosapia bicincta, 74

Redcedar:
foliage—
Argyresthia freyella, 140
bagworm, 126

Coleotechnites juniperella, 136
imperial moth, 212
juniper midge, 442
Monoctenus fulvus, 384
suffusus, 384
Paria sexnotata, 266
Trisetacus cupressi, 32
bark, wood, twigs—
eastern juniper bark beetle, 351
Chrysobothris texana, 283
buds, shoots, roots—
seedcorn maggot, 450
sucking insects—
Cinara canadensis, 78
Juniperivora, 78
sabinae, 78
juniper mealybug, 93
Redcedar, eastern:
arborvitae weevil, 320

Allegheny mound ant, 434
Callidium schotti, 304
Cudonigera houstonana, 169
Gillette eriococcin, 104
juniper webworm, 138
Phloeosinus canadensis, 352

Redgum: See: sweetgum
Rhododendron:

foliage —
Arge abdominalis, 382
clavicornis, 382
Asiatic oak weevil, 322
azalea leafminer, 132
black vine weevil, 321
Datana major, 218
laurel sphinx, 214
southern red mite, 31
twobanded Japanese weevil, 322
bark, wood, twigs—
dogwood twig borer, 299
pitted ambrosia beetle, 370
rhododendron borer, 142
rhododendron stem borer, 300
buds, shoots, roots—
Asiatic garden beetle, 270
Asiatic oak weevil, 322
Asphondylia azaleae, 443
black vine weevil, 321
rhododendron stem borer, 300
sucking insects—
azalea bark scale, 103
azalea whitefly, 87
Cerococcus kalmiae, 105
common falsepit scale, 104
Kleidocerys resedae geminatus, 68
mulberry whitefly, 87
peony scale, 117
rhododendron whitefly, 87
striped mealybug, 91
taxus mealybug, 91

Royalpalm, Florida:

royalpalm bug, 68

Sassafras:

foliage—
cecropia moth, 206
Datana drexelii, 217
Epimecis hortaria, 192
Odontopus calceatus, 334
Pandemis lamprosana, 161
polyphemus moth, 207
promethea moth, 207
sassafras leafminer, 132
spicebush swallowtail, 173
whitefringed beetle, 322
bark, wood, twigs—
Aypothenemus chapuisi, 368
Oberea ruficollis, 300
pitted ambrosia beetle, 370
Scobicia bidentata, 257
Tropideres fasciatus, 318
buds, shoots, roots—
Obera ruficollis, 300
sucking insects—
cottony maple leaf scale, 98
Parrott scale, 105

Seedlings:

foliage—
Acrobasis caryivorella, 181

569

Apantesis radians, 226
Aphelia alleniana, 172
arboreal short-tailed cricket, 53
arborvitae leafminer, 140
ashgray blister beetle, 243
Asiatic oak weevil, 322
Byrrhus spp., 241
honeylocust pod gall midge, 442
margined blister beetle, 243
obliquebanded leafroller, 168
Papaipema furcata, 237
Phyllophaga implicita, 269
Polydrusus impressifrons, 321
Say blister beetle, 243
Solenopsis spp., 435
Texas leafcutting ant, 433
threestriped blister beetle, 243
zebra caterpillar, 237
bark, wood, twigs—
Cryphalus spp., 363
Euzophera magnolialis, 187
hickory spiral borer, 279
lesser cornstalk borer, 186
longhorned beetles, 285
Lymantor decipiens, 361
northern pine weevil, 329
oak-stem borer, 303
Oberea schaumii, 300
Oncideres spp., 300
pales weevil, 323
pitcheating weevil, 326
Platypus quadridentatus, 375
Rhyacionia adana, 151
roundheaded borers, 285
seedcorn maggot, 450
threecornered alfalfa hopper, 71
twig girdler, 301
buds, shoots, roots—
European chafer, 270
green June beetle, 272
Phyllophaga spp., 267
Polyphylla occidentalis, 269
rose chafer, 270
strawberry root weevil, 321
flowers, seeds, fruits—
slash pine flower thrips, 46
sucking insects—
Aphis craccivora, 79
Tropidosteptes amoenus, 66
Serviceberry:
foliage—
Anisomorpha ferruginea, 51
apple bucculatrix, 128
Argyrotaenia quadrifasciana, 171
Bruce spanworm, 189
Diapheromera blatchleyi, 51
velit, 51
giant walkingstick, 51
Hoplocampa halcyon, 409
pallipes, 409
Oligonychus newcomer, 31
pear sawfly, 401
pearleaf blister mite, 31
walkingstick, 50
western tent caterpillar, 204
willow flea weevil, 334
bark, wood, twigs—
amelanchier twig borer, 449

570

apple bark borer, 142
apple fruit moth, 140
oak sapling borer, 293 |
roundheaded appletree borer, 297 |
sucking insects—
Chionaspis lintneri, 111
Prociphilus corrugatans, 80
twocirculi mealybug, 92
woolly elm aphid, 79
Silktree:

claycolored leaf beetle, 266

mimosa webworm, 138

Norape ovina, 176

Xvleborus affinis, 374
Smilax:

Melanaspis smilacis, 116
Sorbaronia: See: mountain-ash
Sourgum: See: tupelo
Sourwood:

dogwood twig borer. 299

hickory horned devil, 211

regal moth, 211

twig girdler, 301
Spicebush:

promethea moth, 207

spicebush swallowtail, 173
Spirea:

Hemileuca lucina, 208
Spruce:

foliage—

Argyrotaenia occultana, 171

bagworm, 126

black vine weevil, 321

Brachyderes incanus, 321

Eufidonia notataria, 191

European chafer, 270

greenheaded spruce sawfly, 405

hairy leaf beetle, 266

joker, 237

Lexis bicolor, 226

pales weevil, 323

Pero morrisonaria, 197

pine colaspis, 263

pine tussock moth, 228

Serica tristis, 269

spruce epizeuxis, 237

spruce spider mite, 30

spruce-fir looper, 191

tufted spruce caterpillar, 237

yellowheaded spruce sawfly, 404

bark, wood, twigs—

balsam fir bark beetle, 357

black and red horntail, 410

blue horntail, 410

Callidium violaceum, 304

Chrysobothris scabripennis, 283
trinervia, 283

Crypturgus pusillus, 362

Dryocoetes affaber, 361
autographus, 361
granicollis, 362

fir coneworm, 183

firtree borer, 307

flatheaded fir borer, 284

four-eyed spruce bark beetle, 352

gloomy borer, 284

Gnathotrichus materiarius, 372

Aylastes salebrosus, 340

tenuis, 340
Hylobius congener, 326
Aylurgops pinifex, 340
[ps spp., 358
Melanophila acuminata, 284
Neacanthocinus pusillus, 304
Neoclytus muricatulus muricatulus
295
old house borer, 316
Orthotomicus caelatus, 358
pales weevil, 323
Phloeosinus pini, 352
Phymatodes dimidiatus , 306
pine engraver, 359
pitch mass borer, 142
Pityophthorus balsameus, 364
biovalis, 364
cariniceps, 364
opaculus, 363
Pygoleptura nigrella, 306
red turpentine beetle, 347
Sirex juvencus, 410
spruce beetle, 349
Stictoleptura canadensis, 304
Trypodendron rufitarsis, 371
white pine weevil, 328
Xvlotrechus undulatus, 312
yellowhorned horntail, 410
buds, shoots, roots—
black vine weevil, 321
Aylastes tenuis, 340
joker, 237
pales weevil, 323
spruce bud moth, 157
strawberry root weevil, 321
Zeiraphera improbana, 157
flowers, seeds, fruits—
fir coneworm, 183
Nepytia pellucidaria, 198
white pine cone borer, 156
sucking insects—
Adelges lariciatus, 84
Cinara atlantica, 78
pergandei, 78
pinivora, 78
taedae, 78
watsoni, 78
cryptomeria scale, 107
elongate hemlock scale, 113
fiorinia hemlock scale, 113
hemlock scale, 106
leaffooted pine seed bug, 67
pine needle scale, 109
round conifer scale, 107
shortneedle evergreen scale, 117
Spruce, black:
foliage—
balsam fir sawfly, 390
Coleotechnites piceaella, 136
eastern blackheaded budworm, 171
European spruce sawfly, 396
false hemlock looper, 198
hemlock looper, 198
redbanded leafroller, 169
small conifer looper, 190
spruce budworm, 164
spruce needleminer, 149
spruce seed moth, 158
yellowheaded spruce sawfly, 404

bark, wood, twigs—
Crypturgus alutaceus, 362
hemlock borer, 284
northern pine weevil, 329
whitespotted sawyer, 313
flowers, seeds, fruits—
spruce seed moth, 158
sucking insects—
Adelges laricis, 84
pine leaf adelgid, 86
red spruce adelgid, 86
Spruce, blue:
foliage—
Cephalcia fascipennis, 380
Coleotechnites piceaella, 136
Epinotia nanana, 158
spruce needleminer, 149
spruce seed moth, 158
yellowheaded spruce sawfly, 404
buds, shoots, roots—
European chafer, 270
strawberry root weevil, 321
sucking insects—
Cooley spruce gall adelgid, 82
eastern spruce gall adelgid, 81
Spruce, Colorado blue:
See: spruce, blue
Spruce, Engelmann:
foliage—
Coleotechnites piceaella, 136
spruce needleminer, 149
spruce seed moth, 158
yellowheaded spruce sawfly, 404
bark, wood, twigs—
Dryocoetes caryi, 362
spruce scolytus, 356
sucking insects—
Cooley spruce gall adelgid, 82
Spruce, Norway:
foliage—
Coleotechnites piceaella, 136
Epinotia nanana, 158
European spruce sawfly, 396
redheaded pine sawfly, 384
spruce needleminer, 149
strawberry root weevil, 321
yellowheaded spruce sawfly, 404
bark, wood, twigs—
Crypturgus alutaceus, 362
southern pine beetle, 343
white pine weevil, 328
sucking insects—
Cinara pilicornis, 78
eastern spruce gall adelgid, 81
pine spittlebug, 72
spruce bud scale, 98
Spruce, red:
follage—
Coleotechnites piceaella, 136

eastern blackheaded budworm, 171

European spruce sawfly, 396

hemlock looper, 198

redbanded leafroller, 169

spruce budworm, 164

spruce seed moth, 158

yellowheaded spruce sawfly, 404
bark, wood, twigs—

Allegheny spruce beetle, 350

balsam bark weevil, 331
black turpentine beetle, 346
Cryphalus rubentis, 363
ruficollis, 363
Dryocoetes caryi, 362
hemlock borer, 284
northeastern sawyer, 314
northern pine weevil, 329
Pityogenes hopkinsi, 357
Pityophthorus dentifrons, 364
intextus, 364
pulchellus, 364
small spruce weevil, 331
southern pine beetle, 343
Strangalepta vittata, 306
Trachysida mutabilis, 306
Trigonarthris minnesotana, 306
whitespotted sawyer, 313
Xyloterinus politus, 371
sucking insects—
Adelges laricis, 84
eastern spruce gall adelgid, 81
pine leaf adelgid, 86
pine spittlebug, 72
red spruce adelgid, 86

Zeiraphera unfortunana, 157
flowers, seeds, fruits—
spruce coneworm, 184
spruce seed moth, 158
sucking insects—
balsam twig aphid, 78
Cooley spruce gall adelgid, 82
eastern spruce gall adelgid, 81
pine spittlebug, 72
Saratoga spittlebug, 73
spruce gall adelgid, 86
Tibicen canicularis, 75
Strawberry:
walkingstick, 50
Sugarberry:
blistergall psyllid, 76
Sumac:
foliage—
Anacampsis rhoifructella, 137
Arge coccinea, 382
Attelabus nigripes, 319
Episimus argutanus, 148
Nephopteryx subfuscella, 186
sumac datana, 217
bark, wood, twigs—

Dicerca obscura, 284
Pityophthorus crinalis, 363
lautus, 363
scriptor, 363
buds, shoots. roots—

Spruce, Sitka:
Cooley spruce gall adelgid, 82
spruce seed moth, 158
Spruce, white:
folage—

Acantholyda maculiventris, 380
balsam fir sawfly, 390
Cephalcia fascipennis, 380
chainspotted geometer, 199
Coleotechnites piceaella, 136
early brown looper, 190
eastern blackheaded budworm, 171
Epinotia nanana, 158
European spruce sawfly, 396
false hemlock looper, 198

fir needle inchworm, 190
Griselda radicana, 160
hemlock looper, 198

Protoboarmia porcelaria indicataria,

200
redbanded leafroller, 169
small conifer looper, 190
spruce budworm, 164
spruce harlequin, 237
spruce needleminer, 149
spruce seed moth, 158
strawberry root weevil, 321
whitespotted sawyer, 313
yellowheaded spruce sawfly, 404
bark, wood, twigs—
Allegheny spruce beetle, 350
Chrysobothris pusilla, 282, 283
Cryphalus ruficollis, 363
Dryocoetes caryi, 362
hemlock borer, 284
Ips latidens, 361
northern engraver, 361
northern pine weevil, 329
northern spruce engraver, 361
Pityophthorus dentifrons, 364
intextus, 364
spruce scolytus, 356
whitespotted sawyer, 313

sumac stem borer, 300
flowers, seeds, fruits—
Moodna ostrinella, 187
Sumac, poison:
Arge humeralis, 382
Pityophthorus crinalis, 363
Sumac, staghorn:
Pitvophthorus scriptor, 363
Sweetfern:
Saratoga spittlebug, 73
walkingstick, 50
Sweetgum:
foliage—
Coleotechnites dorsivittella, 136
forest tent caterpillar, 204
hickory horned devil, 211
imperial moth, 212
luna moth, 207
Nemorimyza posticata, 450
Oiketicus abbotii, 126

Phyllocnistis liquidambarisella, 131

purplish-brown looper, 199
redhumped caterpillar, 222
regal moth, 211
Tetralopha melanogrammos, 180
bark, wood, twigs—
American plum borer, 186
birch bark beetle, 362
Pityophthorus liquidambarus, 363
Platypus compositus, 376
twig pruner, 302
Xvleborinus saxeseni, 374
Xyleborus affinis, 374
ferrugineus, 373
sucking insects—
common falsepit scale, 104
Neosteingelia texana, 90
Parrott scale, 105

periodical cicada, 74
Stictocephala militaris, 71
sweetgum scale, 112
walnut scale, 119
Sweetleaf:
sour-gum scale, 110
Sycamore:

foliage—
Adoxophyes furcatana, 160
American dagger moth, 236
Ancylis platanana, 158
bagworm, 126
Datana contracta, 217
flatheaded appletree borer, 281
hickory horned devil, 211
io moth, 208
Japanese beetle, 271
Macrarocampa marthesia,
Misogada unicolor, 223
Neochlamisus platani, 266
Oiketicus abbotii, 126
Phyllophaga implicita, 269
Plagiometriona clavata, 266
polyphemus moth, 207
puss caterpillar, 175
regal moth, 211
sycamore plant bug, 67
sycamore tussock moth, 225
Tetralopha militella, 180
whitemarked tussock moth, 227

bark, wood, twigs—
American dagger moth, 236
beech borer, 293
Columbian timber beetle, 369
flatheaded appletree borer, 281

i)
N
i)

flatheaded sycamore-heartwood borer,

283
hardwood stump borer, 315
Lichenophanes bicornis, 257
Orchesia castanea, 247
pigeon tremex, 410
powderpost beetle, 255
Tropideres fasciatus, 318
buds, shoots, roots—
Japanese beetle, 271
sucking insects—
Chionaspis parki, 111
cottony maple scale, 97
elm scurfy scale, 108
giant bark aphid, 77
mulberry whitefly, 87
periodical cicada, 74
sycamore lace bug, 64
terrapin scale, 95

(t

Tamarack:
Adelges laricis, 84
eastern larch beetle, 349
larch sawfly, 402
Pitvophthorus opaculus, 363
Saratoga spittlebug, 73
Thuja:
carpenter bee, 439
cedartree borer, 307
Monoctenus spp., 384
small cedar-bark borer, 308
spruce spider mite, 30
strawberry root weevil, 321

Tupelo:
foliage—
Actrix nyssaecolella, 187
Caliroa spp., 401
Phyllophaga forsteri, 269
implicita, 269
luctuosa, 268
Prolimacodes scapha, 178
tupelo leafminer, 125
bark, wood, twigs—
Buprestis rufipes, 281
oak-bark scaler, 309
Platypus compositus, 376
Synanthedon rubrofascia, 143
sucking insects—
sour-gum scale, 110
Tupelo, black:
cottony maple leaf scale, 97
Dysmicoccus difficilis, 94
Phylloxera nyssae, 86
Prolimacodes scapha, 178
whitefringed beetles, 322
Tupelo, water:
forest tent caterpillar, 204

Vv

Viburnum:
Anacampsis rhoifructella, 137
Chionaspis lintneri, 111
dogwood twig borer, 299
hickory leafroller, 170
Synanthedon viburni, 143
twobanded Japanese weevil, 322

WwW
Walnut:
folhlage—

Acrobasis caryivorella, 181
Archips rileyanus, 164
Attelabus bipustulatus, 319
cecropia moth, 206
Colocasia propinquilinea, 237
Datana drexelii, 217
fruittree leafroller, 163
hickory horned devil, 211
hickory tussock moth, 223
Homoeolabus analis, 319
Japanese beetle, 271
Megaxyela spp., 378
Paria quadrinotata, 266
pecan cigar casebearer, 133
pecan leaf casebearer, 181
Phyllophaga forsteri, 269

implicita, 269

luctuosa, 268

prunina, 269
polyphemus moth, 207
regal moth, 211
saddleback looper, 192
Tymnes tricolor, 266
variable oakleaf caterpillar, 220
walnut caterpillar, 217
white oak borer, 292
yellownecked caterpillar, 216

bark, wood, twigs—

American plum borer, 186
banded hickory borer, 289
Chrysobothris sexsignata, 283
Enaphalodes atomarius, 292

373

Eugnamptus striatus, 319
gall midges, 442
Aylocurus rudis, 367
Hypothenemus interstitialis, 368
leopard moth, 145
Micracis suturalis, 367
painted hickory borer, 287
Petalium bistriatum, 254
powderpost beetle, 255
Pseudothysanoes lecontei. 356
twig pruner, 302
white oak borer, 292
Xyleborinus saxeseni, 374
Xyleborus ferrugineus, 373
rubricollis, 374
buds, shoots, roots—
Japanese beetle, 271
pecan leaf casebearer, 181
Phyllophaga forsteri, 269
implicita, 269
luctuosa, 268
prunina, 269
walnut shoot moth, 181
sucking insects—
Chionaspis caryae, 111
lintneri, 111
common falsepit scale, 104
giant bark aphid, 77
large hickory lecanium, 95
Orthezia pseudinsignis, 90
periodical cicada, 74
tuliptree scale, 99
twomarked treehopper, 7!
Walnut, black:
foliage—
butternut woollyworm, 409
Eriophyes caulis, 31
flatheaded appletree borer, 281
Gretchena bolliana, 160
concitatricana, 160
luna moth, 207
walnut sphinx, 215
bark, wood, twigs—
flatheaded appletree borer, 281
sapwood timberworm, 248
walnut sphinx, 215
Xylosandrus zimmermanni, 374
buds, shoots, roots—
Acrobasis demotella, 181
pecan cigar casebearer, 133
flowers, seeds, fruits—
Conotrachelus retentus, 337
walnut husk fly, 448
sucking insects—
walnut lace bug, 65
Walnut, English:
walnut scale, 119
Weigela:
Asterolecanium arabidis, 106
White-cedar:
cryptomeria scale, 107
false hemlock looper, 198
strawberry root weevil, 321
White-cedar, Atlantic:
bagworm, 126
White-cedar, northern:
foliage—
arborvitae leafminer, 140

574

Argvresthia aureoargentella, 140
freyella, 140
bagworm, 126
Coleotechnites thujaella, 135
Monoctenus fulvus, 384
suffusus, 384
pales weevil, 323
Platytetranychus thujae, 3
red carpenter ant, 432
bark, wood, twigs—
Melanophila acuminata, 284
pales weevil, 323
Phloeosinus canadensis, 352
buds, shoots, roots—
arborvitae weevil, 320
black vine weevil, 321
pales weevil, 323
sucking insects—
Cinara tujafilina, 78
Fletcher scale, 96
juniper scale, 108
minute cypress scale, 108
Nuculaspis pseudomeyeri, 117
Willow:
foliage—
Acronicta disians, 236
Agonopterix argillacea, 132
Altica subplicata, 263
American dagger moth, 236
Anacampsis innocuella, 137
apple-and-thorn skeletonizer, 145
Archips purpuranus, 164
Arge clavicornis, 382
Asiatic garden beetle, 270
bagworm, 126
basswood leafminer, 263
birch sawfly, 382
blinded sphinx, 215
browntail moth, 233
Bruce spanworm, 189
Cabera erythemaria, 190
variolaria, 190
Calligrapha multipunctata bigsbyana,
262

—

Caloptilia stigmatella, 132
cecropia moth, 206
Cerura borealis, 223
cinerea, 223
multiscripta canadensis, 223
occidentalis, 223
chainspotted geometer, 199
Chrysomela knabi, 260
Clostera apicalis, 216
brucei, 216
strigosa, 216
Compton tortoiseshell, 174
cottonwood dagger moth, 236
cottonwood leaf beetle, 260
Dasychira obliquata, 229
vagans, 229
definite-marked tussock moth, 228
elm sawfly, 382
Eotetranychus populi, 31
weldoni, 31
European snout beetle, 320
Euura spp., 409
Evora hemidesma, 149
false-sphinx, 218

flatheaded appletree borer, 281
goldsmith beetle, 272
gray willow leaf beetle, 262
graybanded leafroller, 171
green oak caterpillar, 219
gypsy moth, 229
Hyperaeschra stragula, 218
imported willow leaf beetle, 264
10 moth, 208
large aspen tortrix, 168
large maple spanworm, 199
luna moth, 207
maple webworm, 180
Marmara spp., 131
Melanolophia canadaria, 191
mourningcloak butterfly, 174
Nematus fulvicrus, 406

limbatus , 406

oligospilus, 406

salicisodoratus, 405
Nevada buck moth, 208
notched-wing geometer, 196
Notodonta simplaria, 223
obliquebanded leafroller, 168
Oiketicus abbotii, 126
oriental moth, 177
Pachybrachis tridens, 266
pepper-and-salt moth, 200
Pero morrisonaria, 197
Phyllocolpa spp., 409
Phyllophaga crenulata, 269

drakei, 268

forsteri, 269

implicita, 269

luctuosa, 268

prunina, 269

tristis, 268
Polydrusus impressifrons, 321
polyphemus moth, 207
Pontania spp., 409
poplar dagger moth, 236
poplar sawfly, 406
poplar tentmaker, 215
prairie tent caterpillar, 203
Pristiphora sycophanta, 404

Protoboarmia porcelaria indicataria,

200
Pyrrhalta tuberculata, 262
red spotted purple, 175
redhumped caterpillar, 222
satin moth, 233
Schizura ipomoeae, 223
leptinoides, 223
shortwinged meadow katydid, 52
smeared dagger moth, 236
Sparganothis sulfureana, 161
spear-marked black moth, 190
Sphinx luscitiosa, 214
spiny-elm caterpillar, 174
spotted tussock moth, 224
stout looper, 195
. Striped alder sawfly, 407
threelined leafroller, 161
Trichiosoma triangulum, 383
twinspot sphinx, 215
unicorn caterpillar, 223
viceroy, 175
western tent caterpillar, 204

white admiral, 175

willow flea weevil, 334
willow sawfly, 405

willow shoot sawfly, 412
winter moth, 189
yellow-marked caterpillar, 237

bark, wood, twigs—

American dagger moth, 236
black carpenter ant, 429
black twig borer, 375
blinded sphinx, 215
carpenterworm, 146
Chrysobothris azurea, 283
cottonwood borer, 289
cottonwood leaf beetle, 260
Dicerca lurida, 284
Euura spp., 409
flatheaded appletree borer, 281
hardwood stump borer, 315
hornet moth, 141
leopard moth, 145
Magdalis salicis, 333
Marmara spp., 131
Micracis suturalis, 367
swainei, 366
Oberea ferruginea, 300
Poecilonota thureura, 285
poplar borer, 295
poplar-and-willow borer, 334
poplar-gall saperda, 298
Pselaphorhynchites aeratus, 319
cyanellus, 319
Saperda imitans, 298
moesta, 298
mutica, 298
Synanthedon bolteri, 143
proxima, 143
sigmoidea, 143
Xylotrechus annosus annosus, 312

buds, shoots, roots—

Euura spp., 409
hornet moth, 141
Phyllophaga crenulata, 269

drakei, 268

forsteri, 269

implicita, 269

luctuosa, 268

prunina, 269

tristis, 268
Polydrusus impressifrons, 321
poplar clearwing moth, 141
Sthenopis thule, 124
Synathedon proxima, 143
willow beaked-gall midge, 444
willow flea weevil, 334

flowers, seeds, fruits—

Amphibolips quercusfuliginosa, 425

sucking insects—

azalea bark scale, 103

birch lace bug, 65

birch margarodid, 90

black willow scale, 111

Clastoptera salicis, 74

Corythucha elegans, 65
mollicula, 65

cottony maple scale, 97

dentate scale, 123

Dysmicoccus difficilis, 94

S75

fourhumped stink bug, 64
giant bark aphid, 77
Idiocerus spp., 70
Macropsis spp., 70
oystershell scale, 114
Pterocomma populifoliae, 79
salicis, 79
willow scurfy scale, 111
willow scale, 119
Willow, black:
Paranthrene dollii, 144
Wisteria:
Chramesus wisteriae, 352
locust leafroller, 186
silverspotted skipper, 172
Witch-hazel:
Cameraria hamameliella, 130
Datana drexelii, 217
Episimus argutanus, 148
Hormaphis hamamelidis, 80
witch-hazel gall aphid, 80

Wood products: includes insects infesting
lumber, piling, poles, woodwork, and

manufactured products.
ambrosia beetles, 368
anobiid beetles, 249
Anobium punctatum, 254
baldfaced hornet, 436
black carpenter ant, 429
bostrichid beetles, 256
brown prionid, 315
Calcaritermes nearcticus, 62

Canadian powderpost beetle, 254

carpenter bee, 439
centipedes, 28
cigarette beetle, 252
Crematogaster cerasi, 434

clara, 434

lineolata, 434
Cryptotermes brevis, 54

cavifrons, 62
deathwatch beetle, 254
dermestid beetles, 275
drugstore beetle, 252

marine borers—
Bankia spp., 25
Chelura terebrans, 27
Limnoria lignorum, 27
Martesia spp., 25
shipworms, 25
Sphaeroma spp., 27
Teredo spp., 25
wood louse, 27
millipedes, 27
Minthea rugicollis, 256
Monarthrum mali, 370
Narceus americanus, 27
Neotermes castaneus, 62
Jouteli, 62
Nicobium hirtum, 254
nonsubterranean termite, 60
old house borer, 316
Oligomerus alternans, 254
obtusus, 254
pole borer, 307
powderpost beetle, 255
Priobium sericeum, 254
Prorhinotermes—61
simplex, 63
Pselactus spadix, 337
Ptilinus pectinicornis, 254
red oak borer, 291
Reticulitermes arenincola, 58
hageni, 58
tibialis, 58
virginicus, 58
sap beetle, 243
southern lyctus beetle, 256
subterranean termite, 55
termites, 54
Tomolips quercicola, 337
Trogoxylon paralielopipedum, 256
Vespula germanica, 437
wharf borer, 247
white oak borer, 292
whitehorned horntail, 410
Xvletinus harristi, 254
peltatus, 252

eastern subterranean termite, 56 Y
Ernobius spp., 252 Yellow-poplar:

European hornet, 436

foliage—

576

flat powderpost beetle, 299
flatheaded borers, 276
Florida carpenter ant, 432
Formosan subterranean termite, 59
furniture beetle, 254
hairy pine borer, 307
Hexarthrum ulkei, 338
hide beetle, 276
Hylocurus langstoni, 367
Incisitermes milleri, 62
minor, 62
schwarzi, 62
snyderi, 62
Iridomyrmex pruinosus, 435
Kalotermes approximatus, 62
larder beetle, 276
lyctids, 255
Lyctus africanus, 256
brunneus, 256
planicollis, 256

Endopiza liriodendrana, 160
Epimecis hortaria, 192
Odontopus calceatus, 334
Phyllocnistis liriodendronella, 131
polyphemus moth, 207

promethea moth, 207

Tetranychus magnoliae, 31
whitefringed beetle, 322

bark, wood, twigs—

Buprestis rufipes, 281

Columbian timber beetle, 369
Dryocoetes autographus, 362
Euzophera ostricolorella, 187
flatheaded appletree borer, 281

flatheaded sycamore-heartwood borer,

283
Hololepta spp., 241
oak-bark scaler, 309
Orchesia castanea, 247
Plegaderus spp., 241
sapwood timberworm, 248

Xvleborinus saxeseni, 374
sucking insects—
common falsepit scale, 104
periodical cicada, 74
tuliptree aphid, 79
tuliptree scale, 99
walnut scale, 119
Yew:
Asiatic garden beetle, 270
black vine weevil, 321
sucking insects—
Comstock mealybug, 93
cottony camellia scale, 98
cottony taxus scale, 98
dentate scale, 123
elongate hemlock scale, 113
fiorinia hemlock scale, 113
Fletcher scale, 96
grape mealybug, 93
shortneedle evergreen scale, 117
taxus mealybug, 91

Sud

Insect Index

This is an alphabetical listing of the insect
species by both scientific and common
names. Where there are multiple entries, the
primary discussion is indicated by a
boldface page number.

Abgrallaspis ithacae, 106
Acalitus phloeococotes, 32
Acantholyda—
angulata, 380
apicalis, 380
circumcincta, 380
erythrocephala, 13, 378, 379, 380
floridana, 380
luteomaculata, 380
maculiventris, 380
pini, 380
zappei, 380
Acari, 30, 35, 36, 39
Acholla multispinosa, 66
Acleris—
chalybeana, 171
gloverana, 171
logiana, 172
tripunctana, 172
variana, 171
Acmaeodera pulchella, 277
Acmopolynemla bifasciatipenne, 416
Acordulecera, 381
Acossus—
centerensis, 146
populi, 146
Acrididae, 35, 37, 51
Acrobasis—181, 182
betulella, 181
betulivorella, 181
carpinivorella, 181
caryivorella, 181
demotella, 181
elyi, 181
exsulella, 181
indigenella, 181
Juglandis, 181
nuxvorella, 181
rubrifasciella, 181
Acronicta—
americana, 236
distans, 236
funeralis, 236
innotata, 236
interrupta, 236
leporina, 236
lepusculina, 236
lithospila, 236
modica, 236
morula, 236
oblinita, 236
retardata, 236
Actenodes acornis, 285
Actias luna, 207
Actrix nyssaecolella, 187
Acutaspis—
morrisonorum, 107
perseae, 107
Adalia bipunctata, 245
Adel ges—
abietis, 11, 81

cooleyi, 82

lariciatus, 84

laricis, 11, 84

piceae, 11, 83, 85
Adelgidae, 81, 86
adel gids—

balsam woolly, 1, 13, 83, 84, 85, 246,

331, 445, 448
Cooley spruce gall, 82, 83
eastern spruce gall, 81
pine bark, 84, 85
pine leaf, 86
red spruce, 86
spruce gall, 47, 86
Adelphe anisomorphae, 427
Adoxophyes furcatana, 160
Aethes rutilana, 12, 172
Agathidium oniscoides, 241
Agathis—

pini, 153, 154

pumila, 134, 413, 417
A gonopterix—

argillacea, 132

nigrinotella, 132

pteleae, 132

robiniella, 132
Agonoxenidae, 135
agonoxenids, 135
Agrilaxia flavimana, 285
Agrilus—242, 280, 418

acutipennis, 279, 280

anxius, 277

arcuatus torquatus, 279

betulae, 280

bilineatus, 277, 278

celti, 280

cephalicus, 280

difficilis, 280

egenus, 280

fuscipennis, 280

horni, 279

Juglandis, 280

lecontei, 280

liragus, 279

masculinus, 280

otiosus, 280
Agromyzidae, 38, 39, 41, 449
Airora cylindrica, 243
Alaus—

myops, 273

oculatus, 273

Alebra, 70

Aleochara, 242

Aleurochiton forbesii, 87

Aleyrodidae, 40, 87

Allotropa—

ashmeadi, 427

burrelli, 426

convexifrons, 93, 427

utilis, 92
Alloxystidae, 422
Alsophila—190, 199

pometaria, 188
Altica—

ambiens alni, 263

carinata, 263

subplicata, 263
Amblycerus robiniae, 259

579

Amiseginae, 427
Amorbia humerosana, 16]
Amphibolips—
confluenta, 425
quercusfuliginosa, 425
Amphicerus bicaudatus, 257
Amphimallon, see Rhizotrogus
Ampulicidae, 438
ampulicids, 438
Anabrus simplex, 52
Anacampsis—
innocuella, 137
rhoifructella, 137
Anacamptodes—
ephyraria, 191
pergracilis, 191, 192
Anastatus—
disparis, 231, 419
reduvii, 419
Anatis mali, 245
Ancylis—
discigerana, 158
logiana, 158
platanana, 158
Aneflomorpha subpubescens, 303, 305
Aneflus protensus, 308
Anisomorpha buprestoides, 51
ferruginea, 51
Anisota—
discolor, 210
peigleri, 209, 210
pellucida, 210
senatoria, 209
stigma, 208
virginiensis, 209, 210

Anobiidae, 35, 238, 239, 248, 249, 250,

iy |
anobiids, 250, 252
Anobium punctatum, 11, 254
Anomala—
lucicola, 270
oblivia, 270
orientalis, 11
Anomoea laticlavia, 266
Anomogyna elimata, 237
Anoplonyx—
canadensis, 409
luteipes, 409
Anoplura, 46
Anormenis septentrionalis, 69
Antheraea polyphemus, 207
Anthocoridae, 66
Anthocoris musculus, 66
Anthomyiidae, 450
anthomyiids, 450
Anthonomus suturalis, 337
Anthophila pariana, 12
Anthophoridae, 439
Anthophorinae, 439
Anthribidae, 317
Antispila nysaefoliella, 125
antlions, 47, 48
ants—241, 377, 412, 419, 426, 429
Allegheny mound, 434
Argentine, 434, 435
carpenter—430, 431
black, 427, 429, 430, 432
Florida, 432

580

red, 432
fire—435
black imported, 435
red imported, 435
Texas leafcutting, 433
velvet, 428
Anurogryllus arboreus, 53
Apagodiplosis papyrifera, 443, 444
Apanteles—
lacteicolor: See: Dolichogenidea
melanoscelus: See: Cortesia
solitarius, 414
Apantesis radians, 226
Apatelodes torrefacta, 200
Apatelodidae, 200
apatelodids, 200
Aphelia alleniana, 172
Aphelopus theliae, 428
Aphidecta obliterata, 246
Aphididae, 35, 39, 40, 77, 81
Aphidiidae, 412, 413
aphidiids, 413
Aphidoletes thompsoni 445
aphids—8, 47, 69, 76, 77, 78, 79, 245,
418, 432, 434, 442, 447, 448
balsam twig, 78
basswood, 79
beech blight, 79
black pecan, 79
black willow, 79
black-margined, 79
boxelder, 78
common birch, 79
cowpea, 79
crapemyrtle, 79
elm cockscombgall, 80
elm leaf, 79
European birch, 79
giant bark, 77
greater striped red oak, 79
Norway maple, 79
painted maple, 79
poplar petiole gall, 80
poplar vagabond, 80
powdery pine needle, 78
speckled pine needle, 78
sycamore maple, 79
tuliptree, 79
white pine, 78
witch-hazel gall, 80
woolly alder, 80
woolly apple, 79
woolly elm, 79
woolly elm bark, 79
woolly pine needle, 78
Aphis craccivora, 79
Aphrophora—
parallela, 72
saratogensis, 73
Apidae, 42, 439
Apocrita, 377, 412
Apoidea, 439
Arachnida, 26, 28
Araneae, 29
Archiearis infans, 188
Archips—
argyrospilus, 163
cerasivoranus, 161, 162

fervidanus, 162, 163 Atheta, 242

georgianus, 164 Atimia confusa confusa, 308
griseus, 164 Atrecus, 242
infumatanus, 164 Atta texana, 433
magnolianus, 164 Attelabidae, 317, 319
negundanus, 164 Attelabus—
purpuranus, 164 bipustulatus, 319
rileyanus, 164 nigripes, 319
rosanus, 12, 161 Atteva punctella, 140
semiferanus, 163 Aulacidae, 426
Archodontes melanopus melanopus, 315 Aulacus—
Arctids, 226 burquei, 426
Arctiidae, 223 digitalis, 426
Arge— lovei, 426
abdominalis, 382 pallipes, 426
clavicornis, 382 Automeris io, 208
coccinea, 382 B
humeralis, 382 bagworm, 9, 126, 127, 416, 419, 421
pectoralis, 382 Baliosus ruber, 263
scapularis, 382 Bankia, 25, 26
Argidae, 382 Basicladus celibatus, 126
Argvpon flaveolatum, 190 Basilarchia—
Argyresthia— archippus, 175
aureoargentella, 140 arthemis, 175
conjugella, 140 astyanax, 175
freyella, 140 basketworm, orange, 118
laricella, 140 Bassareus literatus, 266
thuiella, 140 Battaristis vittella, 138
Argyresthiidae, 140 bees—243, 377, 422, 426, 427, 428, 439
argyresthiids, 140 bumble, 439
Argyrotaenia— carpenter, 429, 439, 440
alisellana, 171 cuckoo, 439
juglandana, 170 digger, 439
mariana, 171 honey, 377, 439
occultana, 171 leafcutting, 429, 439
pinatubana, 169, 170 mason, 439
quadrifasciana, 171 beetles—19, 54, 237, 238, 239, 248, 249
quercifoliana, \70 alder flea, 263
velutinana, 169 Allegheny spruce, 350
Arhopalus rusticus obsoletus, 307 ambrosia, 6, 10, 24, 243, 250, 338,
Arilus cristatus, 66 346,,368, 369, 371, 372; 373,375,
Arrhenodes minutus, 318 376, 377, 394
Arthropoda, 26 pitted, 370
Asemum striatum, 307 stout, 374
Asilidae, 446 striped, 371
Asilus sericeus, 446 anobiid, 250, 252, 253, 254, 255
Asphondylia— Asiatic garden, 270, 428
azaleae, 443 bark—6, 10, 18, 23, 24, 243, 275, 317,
ilicicola, 443 338, 339, 340, 358
Aspidiotiphagus citrinus, 117 ash, 341
Aspidiotus— balsam fir, 357
cryptomeriae, 107 birch, 362
neril, 107 eastern ash, 341
Astenus, 242 eastern juniper, 351
Asterocampa— elm, 18,20, 354
celtis, 175 flat, 246
clyton, 175 four-eyed spruce, 352
Asterolecaniidae, 39, 105 hickory, 355, 356, 373
Asterolecanium— native elm, 20, 342, 343
arabidis, 106 oak, 22, 362
‘bambusae, 106 peach, 350
miliaris miliaris, 106 smaller European elm, 1, 8, 14, 19,
miliaris robustum, 106 202 T4340) 343. 352353 354:
minus, 105 355, 419
pustulans, 106 blister—243
puteanum, 106 ashgray, 243
quercicola, 105 caragana, 243
variolosum, 11, 105 margined, 243

581

flat, 299
red pine cone, 365, 366
rhinoceros, 273

Nuttall. 243
Say, 243
threestriped, 243

brentid, 318 rove, 242
carrion, 241 sap, 243, 244
cerambycid, 286, 393 scarab, 267

checkered, 274
cigarette, 252

scolytid, 338
seed, 259. 364

click —273 snout, 317

eyed, 273 European, 320
Columbian timber, 369, 370 southern cypress, 351
cone, 364 southern lyctus, 256

cucujid, 246 southern pine, 24, 339, 343, 344, 345,

darkling, 247 348, 375, 421
deathwatch, 249, 254 spider—259
dermestid, 275 brown, 259

whitemarked, 259
spruce, 349, 350

divergent beech, 284
drugstore, 249, 252

eastern Hercules, 272 stag, 267
eastern larch, 349, 350 giant, 267
elm flea, 263 tiger, 240
engraver, 24, 358-361, 393 timber, 247

furniture, 254

goldsmith, 272

green June, 272

ground, 239

hide, 276

hister, 241

ivory-marked, 302

jack pine tip, 366

Japanese, 8, 271, 272, 428, 429

June, 267, 426

lady—91, 92, 93, 94, 95, 96, 97, 98,
99. 100, 101. 103, 107, 108, 109,
110, Tile 113114, 145, 116. 317,

tooth-neck fungus, 246
trogositid, 243
tumbling flower, 248
turpentine—
black, 346, 347, 393
red, 347, 348, 349, 357
western pine. 18
whitefringed, 322
white pine cone, 365
wood-boring. 366
Bessa harveyi. 403
Bethylidae, 427
Bethyloidea, 427

118, 119, 120, 121, 122, 245
convergent, 245
fifteen-spotted, 245
nine-spotted, 245
transverse, 245
twicestabbed, 245

Bibarrambla allenella., 132
Bibio albipennis, 442
Bibionidae. 441

Biston betularia cognataria, 200
Bitoma carinata, 247
Blastobasidae. 35. 133

twospotted, 245 Blattidae, 53

larder, 276 Blepharipa—

leaf—259 ovata, 421
aspen, 260 pratensis, 421, 426, 453
cherry, 262 blotchminers—
claycolored. 266 aspen. 130

basswood. 130
Bombyliidae. 446
bombyliids, 386, 388

cottonwood, 260
elm, 260. 261. 418
European elm, 453

larger, 262 booklice. 46
gray willow, 262 borers—9, 19, 275. 276
hairy, 266 alder, 298

imported willow, 264, 265, 421
lodgepole pine, 350
longhorned, 285
lyctid, 255, 256
May, 66, 267, 269
Mexican bean, 245
nitidulid, 244
oedemerid, 247
passalid, 242
platypodid, 375 black twig, 375
powderpost, 6, 10, 249, 253, 255, 275. blackhorned juniper. 304
367 boxelder twig, 156, 157
bamboo, 257 broadnecked root, 309
Canadian, 254 bronze birch, 128, 277, 278, 421
false, 256, 257, 258 bronze poplar, 279

amelanchier twig. 449
American plum, 186. 187
apple bark. 142

apple twig, 257

ash and privet. 308
Australianpine, 282
banded ash, 294, 295
banded hickory, 289
beech, 293

582

cedartree, 307 red, 375

chestnut bark, 304 redshouldered, 257
cottonwood, 289, 290 small cedar-bark, 308
cottonwood twig, 156 spined bark, 303
cypress bark, 306 spotworm, 279
dogwood, 141 sugar maple, 290
dogwood twig, 299 sumac stem, 300
eastern pine shoot, 154, 155 tanbark, 306
elder, 303 thorn-limb, 298
elm, 298, 299 turpentine, 280, 281
elm bark, 307 twolined chestnut, 277, 278
fig tree, 308 western pine shoot, 18
firtree, 307 whart, 247
flatheaded, 276, 277 white oak, 292, 293
flatheaded apple tree, 281, 282, 421 white pine cone, 156
flatheaded baldcypress sapwood, 277 Bostrichidae, 37, 42, 239, 248, 251, 256
flatheaded fir, 284 bostrichids, 23, 250, 251. 255,256,257
flatheaded sycamore-heartwood, 283 Brachyderes incanus, 11, 321
gallmaking maple, 310 Brachyleptura vagans, 306
gloomy, 284 Brachymeria—
hairy pine, 307 compsilurae, 421
hardwood stump, 315 intermedia, 231
hemlock, 284, 285, 426 Brachys—284
hickory, 293 aeruginosus, 285
hickory spiral, 279 ovatus, 285
larch shoot, 140 tesselatus, 285
large flatheaded pine heartwood, 283 Braconidae, 246, 254, 412, 413
leadcable, 257 braconids, 413, 414
lesser cornstalk, 186 Brentidae, 38, 41, 317, 318
lesser peachtree, 143 bristletails, 44
lilac, 143, 144 Brochymena—
linden, 297 carolinensis, 64
linden bark, 135 quadripustulata, 64
locust, 6, 286, 287, 288 brown prionid, 315
locust twig, 160 Bruchidae, 35, 238, 259
longhorn, 249 bucculatrix, apple, 128
mangrove, 282 Bucculatrix—128, 129
maple callus, 142 ainsliella, 128, 129
maple petiole, 407, 408 canadensisella, 128
mesquite, 287 coronatella, 129
mulberry, 303, 305 luteella, 129
mulberry bark, 303 packardella, 129
oak branch, 294 pomifoliella, 128
oak sapling, 293 quinquenotella, 129
oak-stem, 303 recognita, 129
old house, 9, 249, 251, 316, 317 budminer, hard maple, 125
oriental wood, 258 budworms—
painted hickory, 287 eastern blackheaded, 171, 416
persimmon, 143 jack pine, 167, 168
pine bark, 304, 306 spruce, 14, 15, 18, 124, 164, 165-167,
pitch mass, 142 416, 420
pole, 307 western blackheaded, 171
poplar, 295, 296, 297, 426 bugs—
poplar-and-willow, 334, 335, 336 assassin, 65
poplar-butt, 311 boxelder, 67, 68
red maple cambium, 449 coreid, 67
red oak, 291 damsel, 66
red pine cone, 155 flower, 66
redheaded ash, 294 lace—64, 66
ribbed pine, 315 basswood, 65

- rhododendron, 142 birch, 65
rhododendron stem, 300 elm, 65
roundheaded, 285, 286 hackberry, 64
roundheaded appletree, 297 hawthorn, 65
roundheaded wood, 286 oak, 64
rustic, 312 sycamore, 64, 65
shortleaf pine cone, 155, 156 walnut, 65
shothole—356 lygaeid, 68

larger, 354

583

mealy: See: mealybugs
plant, 66
honeylocust, 66
sycamore, 67
tarnished, 66
rhopalid, 67
royalpalm, 68
seed bug—

leaffooted pine, 67, 418, 419, 426

shieldbacked pine, 63, 418, 419
spined soldier, 64
spittle: See: spittlebugs
stink bugs—63

fourhumped, 64
thaumastocorid, 68
true, 63
wheel, 66

Buprestidae, 37, 38, 41, 42, 238, 239,

248, 276
buprestids, 251, 276, 419, 438
Buprestis—280, 281
apricans, 280
lineata, 281
maculipennis, 281
rufipes, 281
salisburyensis, 281
striata, 281
butterflies—121, 173
brushfooted—173
comma, 173
Compton tortoiseshell, 174
hackberry, 175
monarch, 175
mourningcloak, 174, 175
question-mark, 173, 174
red spotted purple, 175
swallowtail—173
spicebush, 173
tiger, 173
zebra, 173
tawny emperor, 175
viceroy, 175
white admiral, 175
Byrrhidae, 241
byrrhids, 241
Byrrhus, 241
C

Cabera—
erythemaria, 190
variolaria, 190
Caenocara, 255
Calaphis—
betulaecolens, 79
betulella, 79
Calcaritermes nearcticus, 62
Caliroa—
cerasi, 13, 401
petiolata, 402
quercuscoccineae, 401, 402
Callidium—
antennatum antennatum, 304, 305
schotti, 304
texanum, 304
violaceum, 304
Calligrapha—
multipunctata bigsbyana, 262
scalaris, 262
Calliphoridae, 451

584

Callirhopalus bifasciatus, 11, 322
Callirhytis—
cornigera, 141, 423
floridana, 422
quercusfutilis, 424
quercusgemmaria, 423
quercusoperator, 424
quercuspunctata, 423, 424
Callosamia promethea, 207
Caloptilia—
azaleella, 132
bimaculatella, 132
negundella, 132
packardella, 132
pulchella, 132
quercinigrella, 132
sassafrasella, 132
stigmatella, 132
Calosoma—239
calidum, 239, 240
frigidum, 239, 240
scrutator, 196, 239
sycophanta, 231, 239, 240
willcoxi, 239
Cameraria—
aceriella, 130
bethunella, 130
cincinnatiella, 129
corylisella, 130
hamadryadella, 129, 130
hamameliella, 130
Camnula pellucida, 51
Campaea perlata, 200
Camponotus—
abdominalis floridanus, 432
caryae discolor, 432
castaneus, 433
ferrugineus, 432
mississippiensis, 433
nearcticus, 432
pennsylvanicus, 429, 430
pvlartes fraxinicola, 433
sayi, 432, 433
tortuganus, 433
Campoplex frustranae, 152
Campsomeriellia annulata, 429
Canarsia ulmiarrosorella, 187
cankerworms— 14

fall, 9, 188, 189, 193, 416, 426, 427

spring, 193, 194
Carabidae, 238, 239
carabids, 239, 240
Carabus auratus, 231
Carcelia—

lagoae, 176

laxifrons, 453
carpenterworm— 146, 147, 148

aspen, 146

little, 148

pecan, 146

poplar, 146
Carphoborus bifurcus, 352
Carulaspis—

Juniperi, 108

minima, 108
Carynota stupida, 71
Caryobruchus gleditsiae, 259

Carvomyia—
holotricha, 445
sanguinolenta, 445
tubicola, 445
casebearer—133, 181
birch, 134
cigar, 134
elm 7133
larch, 13, 134, 416, 417, 421
pecan cigar, 133
pecan leaf, 181
pecan nut, 181
caterpillar—
chameleon, 237
green oak, 219
paddle, 236
puss, 175
redhumped, 222
saddleback, 176
spiny-elm, 174
tent, 14, 64, 201
eastern, 14, 202, 203, 204, 420, 426
forest, 8, 14, 121, 204, 205, 416,
419, 426, 451, 452
prairie, 203
Sonoran, 203
western, 204
tufted spruce, 237
tufted white pine, 237
uglynest, 161, 162
unicorn, 223
variable oakleaf, 220, 221
walnut, 217, 218, 416
yellow-marked, 237
yellownecked, 216, 217
Zebras 257
Catocala, 236
Catogenus rufus, 246
Caulocampus acericaulis, 407, 408
Cecidomyia—
ocellaris, 443
pellex, 443
piniinopis, 443
poculum, 443
resinicola, 443, 444
Cecidomylidae, 34, 39, 426, 442, 444
centipedes, 25, 26, 28, 54
Centruroides vittatus, 28, 29
Cephalcia—
fascipennis, 380
fulviceps, 380
marginata, 380
Cephenemyia, 440
Cephidae, 412
Cephoidea, 412
Cerambycidae, 35, 37, 38, 41, 42, 238,
239, 246, 248, 249, 251, 285, 427
cerambycids, 308, 419
Ceraphronoidae, 426
Ceratomia—
amyntor, 213
catalpae, 213
undulosa, 213
Ceratopogonidae, 441
Cerceris fumipennis, 438
Cercopidae, 37, 40, 72
cercopids, 428, 438
Cerococcidae, 105

cerococcids, 105
Cerococcus—

kalmiae, 105

parrotti, 105
Ceroplastes—

ceriferus, 94

curipediformis, 94

floridensis, 94
Ceruchus piceus, 267
Cerura—

borealis, 223

cinerea, 223

multiscripta canadensis, 223

occidentalis ,. 223
Chaetexorista javana, 178, 453
chafer—

European, 270

lamellicorn leaf, 267

pine, 270, 271

rose, 270, 271
Chaitophorus—

populicola, 79

stevensis, 79
Chalastogastra, 377
Chalcididae, 35, 39, 421
Chalcidoidea, 415, 416
chalcids, 94, 415, 416, 421, 426
Chalcophora—

georgiana, 283

liberta, 283

virginiensis, 283, 284
Chalcophorella campestris, 283
Chamaemyiidae, 448
changa, 53
Charadra deridens, 237
Charhyphus, 242
Chariessa pilosa, 274
Chelifer cancroides, 29
Chelura terebrans, 27
chiggers, 32
Chilocorus stigma, 102, 245
Chilopoda, 26, 28
Chionaspis—

acericola, 111

americana, 108

caryae, 111

corni, 108

furfura, 109

heterophyllae, 109

kosztarabi, 111

lintneri, 111

nyssae, 110

parki, 111

pinifoliae, 109, 110

salicisnigrae, 111
Chironomidae, 441
Chlorophorus annularis, 11, 317
Chloropidae, 448
Choragus zimmermanni, 318
Choreutidae, 145
choreutids, 145
Choristoneura—

conflictana, 168, 169

fractivittana, 169

fumiferana, 164, 165, 166

pinus maritima, 168

pinus pinus, 167

rosaceana, 168, 169

585

Chramesus—
chapuisi, 352
hicoriae, 352
subopacus, 352
wisteriae, 352
Chrysaster ostensachenella, 130
Chrysididae, 427
Chrysis shanghaiensis, 427
Chrysobothris—281, 282, 418
adelpha, 283
azurea, 283
blanchardi, 283
dentipes, 283
femorata, 281
floricola, 283
harrisi, 283
neopusilla, 283
orono, 282
pusilla, 282, 283
scabripennis, 283
sexsignata, 283
texana, 283
tranquebarica, 282
trinervia, 283
viridiceps, 283
Chrysocharis laricinellae, 134, 399, 416,
417
Chrysoclista linneella, 12, 135
Chrysomelidae, 36, 259, 266, 421
Chrvsomela—
crotchi, 11, 260
interrupta, 11, 260
knabi, 260
scripta, 260
Chrysomphalus—
aonidum, \11
bifasciculatus, 111
dictvospermi, 111
Chrysopa oculata, 47
Chrysopidae, 47
chrysopids, 422
Chrvysops, 446
cicada killer, 438
cicadas—69, 74, 75, 438
dog-day, 74
periodical, 74, 75
Cicadellidae, 39, 69, 447
cicadellids, 428, 438
Cicadidae, 37, 40, 74
Cicindelidae, 238, 240
Cicindela spp., 240
Cicinnus melsheimeri, 200
Cimberis—
elongatus, 318
pilosus, 318
Cimbex americana, 382, 383
Cimbicidae, 382
Cinara—77
atlantica, 78
canadensis, 78
confinis, 78
Juniperivora, 78
pergandei, 78
pilicornis, 78
pinea, 78
pinivora, 78
sabinae, 78
strobi, 78

586

taedae, 78

tujafilina, 78

watsoni, 78
Cingilia catenaria, 199
Cinyra gracilipes, 285
Citheronia—

regalis, 211, 212

sepulcralis, 212
Clastoptera—

achatina, 74

obtusa, 74

proteus, 74

salicis, 74

undulata, 74
Clavaspis ulmi, 111
clearwing, banded ash, 144
Cleis picta, 246
Cleptes semiauratus, 427
Cleptinae, 427
Cleridae, 238, 274, 275
clerid, blackbellied, 275
Clistogastra, 377
Clostera—

albosigma, 216

apicalis, 216

brucei, 216

inclusa, 215, 216

strigosa, 216
Closterocerus—

cinctipennis, 385, 390

tricinctus, 264
Cnidocampa flavescens, 12, 177
Coccidae, 37, 40, 94
Coccinella—

novemnotata, 245

transversoguttata richardsoni, 245
Coccinellidae, 245
coccinelids, 246, 262
Coccodidae, 87
Coccophagus—

immaculatus, 103

insidiator, 418
Coccyvgomimus turionellae turionellae, 415
Cochliomyia—

hominivorax, 17, 451

macellaria, 451
Cochylidae, 172
cochylids, 172
cockroaches, 53, 54, 425, 438
Coeloides scolytivorus, 341
Colaspis pini, 263
Coleomegilla maculata, 262
Coleophora—

alniella, 133

atromarginata, 133

laricella, 12, 134,

laticornella, 133

querciella, 133

serratella, 12, 134

tiliaefoliella, 133

ulmifoliella, 12, 133
Coleophoridae, 133
Coleoptera, 6; 8, 11, 17, 35,36, 37; 38,

41, 42, 237, 238, 259, 317, 413, 414,

416, 419, 421, 427, 446, 451, 452
Coleotechnites—

apicitripunctella, 135

chillcotti, 136

dorsivittella, 136
Juniperella, 136
piceaella, 136
thujaella, 135
variella, 136
Collembola, 44
Colocasia—
flavicornis, 237
propinquilinea, 237
Colopha ulmicola, 80
Colydiidae, 247
colydiids, 247
Colydium lineola, 247
comma, 173
Compsilura concinnata, 217, 231, 421,
452
Compton tortoiseshell, 174
coneworm—
Atlantic pine, 186
baldcypress, 185
blister, 184
fir, 183
loblolly pine, 186
mountain pine, 186
south coastal, 185
southern pine, 183
spruce, 184
webbing, 184, 419
coniopterygid, 108
Conocephalus brevipennis, 52
Conophthorus—365
banksianae, 366
coniperda, 365
resinosae, 365, 366
Conopidae, 447
Conostigmus virginicus, 426
Conotrachelus—366, 337
affinis, 337
anaglyptius, 337
aratus, 337
carinifer, 336
caroliniensis, 378
crataegi, 336
elegans, 337
hicoriae, 337
Juglandis, 337
naso, 336, 337
nenuphar, 336
posticatus, 336
retentus, 337
Contarinia—
canadensis, 443
catalpae, 443
cerasiserotinae, 443
Juniperina, 442
negundifolia, 443
verrucicola, 443
virginianiae, 443
Coptodisca splendoriferella, 126
Coptotermes formosanus, 11, 54, 59, 60
Coreidae, 67
Corthylus—
columbianus, 369
punctatissimum, 370
Corticotomus cylindricus, 243
Corydalidae, 47
Corydalus cornutus, 47

Corythucha—
aesculi, 65
arcuata, 64
associata, 65
bellula, 65
celtidis, 65
ciliata, 64
cydoniae, 65
elegans, 65
Juglandis, 65
mollicula, 65
pallipes, 65
pergandei, 05
pruni, 65
ulmi, 65
Cosmopterigidae, 135
cosmopterigids, 135
Cossidae, 37, 41, 145
Cossoninae, 248, 337
Cossonus—
concinnus, 337
corticola, 337
impressus, 337
platalea, 337
Cossula magnifica, 146
Cotalpa lanigera, 272
Cotesia—
congregata, 214
melanoscelus, 231, 414
Cotinis nitida, 272
Cratichneuman sublatus, 222
Cregya oculata, 275
Crematogaster—
ashmeadi, 434
cerasi, 434
clara, 434
laeviuscula, 434
lineolata, 434
Cremifania nigrocellulata, 448
Cressonia juglandis, 215
crickets—48, 52, 53, 54, 447
arboreal short-tailed, 53
changa, 53
mole—53
European, 53
southern, 53
Mormon, 52
tree—416, 419
snowy, 53
Croesia semipurpurana, 171
Croesus—
castaneae, 407
curvarius, 407
latitarsus, 406, 407
varus, 407
Crossocerus annulipes, 438
Crustacea, 26, 27, 43
Crvyphalus—
fraseri, 363
rubentis, 363
ruficollis, 363
Cryptococcidae, 101
Cryptococcus—11, 101
fagisuga, 11, 101, 102
williamsi, 102
Cryptorhynchus lapathi, 11, 334, 335
Crvptotermes—
brevis, 54, 61, 62

cavifrons, 62
Cryptothelea gloverii, 127
Crypturgus—

alutaceus, 362

borealis, 362

pusillus, 362
Ctenicera—

nitidula, 274

triundulata, 274
Ctenocephalides—

canis, 48

felis, 48
Ctenuchidae, 226
ctenuchids, 226
Cucujidae, 246
Cucujus clavipes, 246
Cudonigera houstonana, 169
Cuerna costalis, 71
Culicidae, 440
Cupedidae, 241
cupedids, 241
Cupes concolor, 241
curculio—

plum, 336

quince, 336
Curculio—333, 336

caryae, 333

caryatrypes, 333

fulvus, 333

lowensis, 333

nasicus, 333

neocorylus, 333

proboscideus, 334

Sayl, 333

sulcatulus, 334

Curculionidae, 238, 248, 251, 317, 319,

320, 338, 378
Curculionoidea, 35, 36, 37, 38, 42, 317
Cuterebra, 440
Cuterebridae, 440
cutworms—237

white, 237
Cydia—

anaranjada, 158

caryana, 158

ingens, 158

strobilella, 158

toreuta, 158
Cymatodera bicolor, 275
Cynipidae, 35, 39, 422
cynipids, 141, 419, 422
Cynipoidea, 421
Cyrtepistomus castaneus, 11, 322
Cyrtolobus discoidalis, 71
Cyzenis albicans, 190

D
daddylonglegs, 29

Dahlbominus fuscipennis, 386, 390, 392,

396, 417
damselflies, 45
Danaus plexippus, 175
Dasineura—

balsamicola, 442

communis, 442

gleditchiae, 412

pseudacaciae, 442
Dasychira—

basiflava, 228

588

cinnamomea, 229

dorsipennata, 229

meridionalis, 229

obliquata, 229

pinicola, 228, 229

plagiata, 229

tephra, 229

vagans, 229
Dasymutilla occidentalis occidentalis, 428
Datana—

angusi, 217

contracta, 217

drexelii, 217

integerrima, 217, 218

major, 218 |

ministra, 216, 217

perspicua, 217
Dendrocranulus, 339
Dendroctonus—17, 18, 339, 343, 358

brevicomis, 18

frontalis, 343, 344, 345, 359

murrayanae, 350

punctatus, 350

rufipennis, 349, 350

simplex, 349, 350

terebrans, 346, 347

valens, 347, 348, 349
Dendrotettix—

australis, 52

quercus, 51
Dermacentor variabilis, 32
Dermestes—

lardarius, 276

maculatus, 276
Dermestidae, 248, 275
Derocrepis aesculi, 266
Derodontidae, 246
derodontids, 246
Desmia funeralis, 178
Desmocerus palliatus, 303
Diachlorus ferrugatus, 446
Dialeurodes—

chittendeni, 87

citri, 87
Diapheromera—

blatchleyi, 51

femorata, 50, 51

velii, 51
Diaphnocoris chlorionis, 66, 67
Diaspididae, 106
Diaspidiotus—

ancylus, 112

caryae, 113

liquidambaris, 112

mecombi, 113

osborni, 112
Dibrachys cavus, 394, 421

Dicerca—
divaricata, 284
lurida, 284

obscura, 284
punctulata, 284
tenebrica, 284
tenebrosa, 284
Dichelonyx—269
albicollia, 270
elongata, 270
subvittata, 270

Dichomeris—
ligulella, 137
marginella, 12, 138, 139
Dimmockia incongru, 418
Dinoderus minutus, 257, 258
Diodyrhynchus, 318
Dioryctria— 182, 183, 185
abietivorella, 183, 185
amatella, 183
clarioralis, 184
disclusa, 184, 185
ebeli, 185
merkeli, 186
pygmaeella, 185
reniculelloides, 184
resinosella, 186
taedae, 186
yatesi, 186
zimmermanni, 182, 186
Diplopoda, 26, 27
Diplotaxis—
liberata, 270
sordida, 270
Diprion, 394
similis, 13, 395, 396
Diprionidae, 383
Diprioninae, 383

Diptera, 6, 8, 12, 34, 35, 38, 39, 41, 416,

419, 421, 426, 440, 447
Dircaea quadrimaculata, 247
dobsonflies, 47

Dolichogenidea lacteicolor, 414

Dolichopodidae, 447
Dolichopus vittatus, 447
Dolichovespula, 437
doodlebugs, 48
Dorcaschema—
alternatum, 303
wildii, 303, 305
Dorcatoma, 255
Dorcus parallelus, 267
dragonflies, 45, 54, 440
Drepanaphis—
acerifoliae, 79
carolinensis, 79
nigricans, 79
sabrinae, 79
Drepanosiphum platanoidis, 79
Drosophila, 448
Drosophilidae, 448
Dryinidae, 428

Dryocampa rubicunda, 210, 211

Dryocoetes—

affaber, 361

autographus, 361

betulae, 362

caryl, 362

granicollis, 362
Dryocosmus—

kuriphilus, 425

quercuspalustris, 425
Dryophthorus americanus, 337
Dynastes.tityus, 272, 273
Dyseriocrania auricyanea, 124
Dysmicoccus—

difficilis, 94

morrisoni, 94

obesus, 94

wistariae, 91

E

Eacles—

imperialis, 212

imperialis pini, 212
Eburia quadrigeminata, 302
Ecdytolopha insiticiana, 160
Ectodemia—

heinrichi, 125

populella, 125
Ectropis crepuscularia, 192
Elachertus cacoeciae, 418
Elaphidion mucronatum, 303
Elaphidionoides—

incertus, 303

villosus, 302
Elasmopalpus lignosellus, 186
Elasmuche lateralis, 64
Elateridae, 35, 239, 273
Elatophilus inimica, 66
elm calligrapha, 262
elm prominent, 222
Empididae, 446
Empoasca, 70
Enaphalodes—

atomarius, 292

cortiphagus, 292

rufulus, 291
Enchenopa binotata, 71
Encvclops caerulea, 309
Encyrtidae, 418
encyrtids, 418, 419
Endasys subclavatus, 386, 394
Endopiza—

liriodendrana, 160

palliolana, 160
Endothenia albolineana, 149

engraver—
hackberry, 356
northern, 361

northern spruce, 361
pine, 357, 359
six-spined, 358

small southern pine, 359, 36i

southern pine, 359, 360
Ennomos—

magnarius, 196

subsignarius, 196, 197
Enoclerus—

lecontei, 275

nigripes, 275, 341
Eotetranychus—

hicoriae, 31

matthyssei, 31

populi, 31

querci, 31

weldoni, 31
Epargyreus clarus, 172
Ephemeroptera, 44
Epicauta—

fabricii, 243

lemniscata, 243

pestifera, 243

subglabra, 243

torsa, 243
Epilachna varivestis, 245
Epilachninae, 245
Epimecis hortaria, 192

589

Epinotia—

aceriella, 157

lindana, 160

nanana, 12, 158

solandriana, 158

stroemiana, 160
Epipaschiinae, 178, 179
Episimus argutanus, 148
Erannis—190

tiliaria, 193, 194, 195
Ericampa juglandis, 409
Eriococcidae. 101
eriococcids. 101
erlococcin—

oak, 104

Gillette, 104
Eriococcus—

azaleae. 103

gillettei, 104

quercus, 104
Eriocraniidae. 124
eriocraniids, 124
Eriophyes—

caulis, 31

fraxiniflora, 31

parapopulli, 31

pyri, 31
Eriophyidae, 31. 39
Eriosoma—

americanum, 79

crataegi, 79

lanigerum, 79

rileyi, 79
Eriotremex formosanus, 13, 411
Ernobius—252, 254

filicornis, 254

granulatus, 254

mollis, 254

tenuicornis, 254
Erynniopsis antenna, 453
Erythroneura, 70
Essigella pini, 78
Etainia sericopeza, 12
Eucallipterus tiliae, 79
Euceraphis—

lineata, 79

mucida, 79

punctipennis, 79
Eucharitidae. 419
eucharitids, 419
Eucoilidae, 422
Eucosma—

cocana, 155

gloriola, 154, 155

monitorana, 155

sonomana, 18

tocullionana, 156
Eucrada humeralis, 255
Eufidonia notataria, 191
Eugnamptus—

collaris, 319

striatus, 319
Eulachnus—

agilis, 78

rilevi, 78
Eulecanium—

caryae, 95

cerasorum, 94

590

Eulophidae, 416
eulophids, 416
Eupelimidae, 419
eupelimids, 419
Eupelmella vesicularis, 419
Eupelmus—
allynii, 419
cyaniceps, 419
pini, 419
Eupithecia—
filmata, 190
luteata, 190
palpata, 190
transcanadata, 190

Euproctis chrysorrhoea, 12, 233, 234, 235

Eurytetranychus buxi, 31
Eurytoma—
magdalidis, 421
pint, 421
pissodis, 421
tylodermatis, 421
Eurytomidae, 421
eurytomids, 421
Eustrophus tomentosus, 247
Eutittix, 70
Eutrapela clemataria, 199, 418
Eutrombicula alfreddugesi, 32
Eutromula pariana, 145
Euura, 409
Euxoa scandens, 237
Euzophera—
magnolialis, 187
ostricolorella, 187
semifuneralis, 186
Evaniidae, 425
Evanioidea, 425
Evora hemidesma, 149
Exenterus—
abruptorius, 392
amictorius, 396, 415
nigrifrons, 386, 388
Exorista larvarum, 231, 453
Exoteleia—
dodecella, 136
nepheos, 137
pinifoliella, 136

F

Fagiphagus imbricator, 79
false-sphinx, 218
Fascista cercerisella, 138
Fenusa—
dohrnii, 13, 400
pusilla, 13, 399
ulmi, 13, 400
Feralia jocosa, 237
Ferrisia virgata, 91
fiery hunter, 239, 240
Figitidae, 422
figitids, 422
filament bearer, 195, 196
filbertworm, 159
Fiorinia—113, 115
externa, 113
fioriniae, 114
Japonica, 114
theae, 113
Flatidae, 69
flatids, 428

fleas—48
cat, 48
dog, 48
human, 48
oriental rat, 48
flies—54, 93, 440
aphid, 448
bee, 446
bigheaded, 447
black, 440, 441
black blow, 451
black cherry fruit, 448
black horse, 445
blow, 451
bot, 440
cherry fruit, 448
chloropid, 448
conopid, 447

dance, 446

deer, 445, 446
flesh, 451

flower, 91, 99, 447
fruit, 188, 447
greenbottle, 451
harvest, 74

horse, 438, 440, 445
house, 448, 450
ichneumonlike, 445
leafminer, 449
longlegged, 447
louse, 451
March, 441
robber, 446
snipe, 446
soldier, 445
stable, 450
striped horse, 445
tachina, 452
tachinid, 421, 452
thickheaded, 447
tsetse, 450
vinegar, 448
walnut husk, 448
warble, 440
Formica exsectoides, 434
Formicidae, 35, 42, 429
Frankliniella tritici, 47
fruitworm, green, 237
Fulgoridae, 35
G
gall midges—91, 112, 419, 442, 443-445
balsam, 442
boxelder, 443
honeylocust pod, 442
ocellate, 443
willow beaked-, 444
galls—76, 423
blister gall makers, 76
~ gouty oak, 423, 424
hackberry nipplegall maker, 76
hackberry stargall, 76
horned oak, 423
large oak-apple, 425
leaf gall makers, 76
linden wart, 443
noxious oak, 424
oak fig, 425
oak flake, 424

oak potato, 424

pinkster bud, 443

ribbed bud, 423

succulent oak, 425
Gargaphia tiliae, 65
Gastropoda, 43
Gaurax apicalis, 449
Gelechiidae, 135
geometer—

chainspotted, 199

notched-wing, 196
Geometridae, 187
Geopinus incrassatus, 240
Gibbium psylloides, 259
Gibbobruchus mimus, 259
Gilpinia—

frutetorum, 13, 396, 417

hercyniae, 13, 396, 397
girdler—

birch and beech, 310

pecan twig, 301

twig, 301
Gluphisia septentrionalis, 223
Glycobius speciosus, 290
Glyptoscelis—

barbata, 266

pubescens, 266
Gnathotrichus materiarius, 372, 373
gnats—

buffalo, 441

darkwinged fungus, 442

turkey, 441
Gnophothrips fuscus, 46
Goes—

debilis, 294

pulcher, 293

pulverulentus, 293

tesselatus, 293

tigrinus, 292, 293, 294
Goniozus—

electus, 427, 428

longinervis, 428
Gossyparia spuria, 11, 103
Gracillaria syringella, 12, 131
Gracillariidae, 129
Graphium marcellus, 173
Graphocephala versuta, 71
Graphognathus, 322
grasshoppers—48, 51, 446, 447

clearwinged, 51

differential, 51

eastern lubber, 52

longhorn, 52

migratory, 51

postoak locust, 51

redlegged, 51

twostriped, 51
Gretchena—

bolliana, 160

concitatricana, 160
Griselda radicana, 160
grubs—

common cattle, 440

northern cattle, 440

white, 268, 428, 446
Gryllidae, 37, 52
Grvllotalpa gryllotalpa, 11, 53
Gryllotalpidae, 53

591

Gryllus, 53

Gryon pennsylvanicus, 426
Gyponana, 70

Gypsonoma haimbachiana, 156
Gyropdidae, 46

H

Habrolepis dalmanni, 419
Hadrobregmus notatus, 255
Haltichella—

rhyacionia, 421

xanticles, 421
Halysidota—

harristi, 225

tessellaris, 224, 225
Hamaelistes spinosus, 80
Haploa—

clymene, 226

lecontei, 226
Harpalus, 240
Hedya chionosema, 160
Heliozelidae, 125
hellgrammites, 47
Hemerobiidae, 47
Hemiberlesia—

lataniae, 114

rapax, 114
Hemichroa crocea, 13, 407
Hemicoelus carinatus, 253, 254
Hemicrepidus bilobatus, 274
Hemileuca—

lucina, 208

maia, 208

nevadensis, 208
Hemiptera, 6, 11, 39, 63, 66, 69, 416,

419, 452
Hepialidae, 38, 124
Hesperiidae, 172
Heterarthrus nemoratus, 13, 398
Heterobostrychus aequalis, 11, 258
Heterocampa—

bilineata, 222

biundata, 222

guttivitta, 220, 221, 222

manteo, 220, 221

umbrata, 222
Heterospilus flavicollis, 254
Hexarthrum ulkei, 337
Hexomyza—

schineri, 450

tiliae, 450
hickory horned devil, 211, 212
hickory shuckworm, 158, 416, 421, 427
Himatolabus pubescens, 319
Hippelates, 449
Hippoboscidae, 451
Hippodamia convergens, 245
Histeridae, 238, 241
Histeromerus canadensis, 254
Holcocera lepidophaga, 133
Hololepta, 241
Holostrophus bifasciatus, 247
Homadaula anisocentra, 12, 138, 139
Homaeotarsus , 242
Homaledra sabalella, 135
Homalodisca coagulata, 71
Homoeolabus analis, 319

Homoptera, 11, 35, 37, 39, 40, 69, 416,

419, 428, 447

592

Hoplocampa—

halcyon, 409

lacteipennis, 409

montanicola, 409

oskina, 409

pallipes, 409
Hormaphis hamamelidis, 80
horned passalus, 242
hornets—436, 437, 438

baldfaced, 436

European, 436
horntails—409, 410, 422

black and red, 410

blue, 410

Formosan, 411

whitehorned, 410

yellowhorned, 410
Hubbellia marginifera, 52
Hyalophora cecropia, 206
Hybrizontidae, 412, 413
hybrizontids, 413
Hydria prunivorata, 190
Hylastes—

exilis, 340

porculus, 340

salebrosus, 340

tenuis, 340
Hylecoetus lugubris, 248
Hylemya platura, 450
Hylesininae, 338, 340, 366
Hylesinus—

aculeatus, 341

pruinosus, 341
Hylobius—322, 323

aliradicis, 326

congener, 326

pales, 323, 324

pinicola, 326

radicis, 324, 325

rhizophagus, 326

warreni, 326
Hylocurus—366, 367

bicornus, 367

biorbis, 367

harnedi, 367

langstoni, 367

rudis, 367

spadix, 367
Hylotrupes bajulus, 11, 249, 316
Hylurgopinus rufipes, 342
Hylurgops pinifex, 340
Hymenoptera, 6, 8, 13, 17, 35, 36, 37,

38, 39, 42, 254, 377, 414, 415, 416,

419, 421, 446, 447, 452
Hyparpax—

aurora, 223

perophoroides, 223
Hyperaeschra stragula, 218
Hyperaspis congressis, 245
Hyphantria—

cunea, 225, 226

textor, 225
Hypoderma—

bovis, 440

lineatum, 440
Hypothenemus—366, 367

chapuisi, 368

dissimilis, 367

eruditus, 368
interstitialis, 368
obscurus, 364, 368
quercus, 368
rotundicollis, 368
Hypulus concolor, 247
Hyssopus, 418
I
Ibalia—
anceps, 422
leucospoides ensiger, 422
maculipennis, 422
scalpellator, 422
Ibaliidae, 421, 422
ibaliids, 422
Ichneumon navus, 192
Ichneumonidae, 413, 414

ichneumonids, 127, 386, 392, 394, 403,

410, 415, 420
Ichneumonoidea, 412, 445
ichneumons—392, 414
Icosta americana, 451
Idia aemula, 237
Idiocerus, 70
Illinoia liriodendri, 79
inchworm, fir needle, 190
Incisitermes—

milleri, 62

minor, 62

schwarzi, 62

snyderi, 62
Incurvariidae, 125
incurvariids, 125
Insecta, 26, 33
Ips—10, 18, 339, 358, 359, 372, 393

avulsus, 359, 361

borealis, 361

calligraphus, 24, 358, 359, 360

grandicollis, 24, 359, 360

latidens, 361

perroti, 361

perturbatus, 361

pini, 24, 359

typographus, 340
Tridomyrmex—

humilis, 434

pruinosus, 435
Tridopsis larvaria, 192
Isa textula, 178
Isoptera, 11, 42, 54
[tame pustularia, 191
Ithycerus noveboracensis, 331
Itoplectis—

conquisitor, 127, 415

quadricingulata, 415

viduata, 415
Ixodes scapularis, 33

Janetiella coloradensis, 445
Janus abbreviatus, 412
Japanagromyza viridula, 450
joker, 237

K

Kalotermes approximatus, 62
Kalotermitidae, 60, 61, 62
Kaltenbachiella ulmifusa, 80

katydids—
angularwinged, 52
broadwinged, 52
forktailed bush, 52
shortwinged meadow, 52
Kermes—100, 101, 133
andrei, 101
galliformis, 101
kingti, 101
pubescens, 101, 419
Kermesidae, 100
Kleidocerys resedae geminatus, 68
Knulliana cincta, 289
L
lacewings—47, 94
brown, 47, 93, 95
goldeneye, 47

green, 47593,,95, 98, 103; LE,

11925120122

Lacon—

avita, 274

discoidea, 274
Lacosoma chiridota, 200
Laetilia coccidivora, 187
Lagoa crispata, 176
Lambdina—

fervidaria athasaria, 199

fiscellaria, 198

pellucidaria, 198, 199
Lapara—

bombycoides, 214

coniferarum, 215
Laphria—

flavicollis, 446

index, 446

thoracica, 446
Laricobius erichsonii, 246
Lasiocampidae, 201
Lasioderma serricorne, 252
Lathrobium, 242
Latrodectus mactans, 29
leaf crumpler, 181
leafblotch miners, 129
leafcutter, maple, 125
leaffolder—

grape, 178

redbud, 138
leafhoppers—69, 70, 438
whitebanded elm, 21, 70, 71
leafminers— 148

ambermarked birch, 401

arborvitae, 140, 421

aspen, 131

azalea. 132

basswood, 263

birch, 399, 416, 420

boxwood, 442

elm, 400

European alder, 400

gregarious oak, 129

lilac, 131

locust, 264, 416, 421

maple, 130

native holly, 449, 450

sassafras, 132

solitary oak, 129, 130

tupelo, 125

unspotted, 131

114,

593

leafrollers—148, 149, 160
basswood, 178, 179
boxelder, 132
fruittree, 163
graybanded, 171
hickory, 170
locust, 186
obliquebanded, 168
raspberry, 160
redbanded, 169
threelined, 161
leaftier—
elder, 178
oak, 171
lecanium—
European fruit, 96, 97, 419
large hickory, 95
oak, 96
Lecanium—
caryae, See: Eulecanium
cerasorum, See: Eulecanium
corni, See: Parthenolecanium
fletcheri, See: Parthenolecanium
nigrofasciatum, See: Mesolecanium
quercifex, See: Parthenolecanium
Lecanodiaspididae, 104
Lecanodiaspis prosopidis, 104

Lepidoptera, 3. 12.17, 35; 30; 27, 38, 39,

41, 123, 124, 129, 236, 413, 414, 415,

416, 418, 419, 420, 421, 427, 440, 446,

452
Lepidosaphes—

camelliae, 115

pallida, \15

ulmi, 114, 115

yanagicola, 115
Leptacinus, 242
Leptocoris trivittatus, 67, 68
Leptogaster flavipes, 446
Leptoglossus—

corculus, 67

occidentalis, 67
Lepturopsis biforis, 306
Leschenaultia fulvipes, 208
Lespesia—

sabroskyi, 208

samiae, 207
Leucoma salicis, 12, 233, 234
Leucopis—108

obscura, 448
Lexis bicolor, 226
lice—

chewing, 46

sucking, 46
Lichenophanes—

arminger, 257

bicornis, 257
Limacodidae, 176
Limnoria lignorum, 27
Limnoriidae, 43
Liopteridae, 421
Liothrips umbripennis, 46
Lipoptena, 451
Litaneutria minor, 49
Lithophane—

antennata, 237

laticinerea, 237
locust, postoak, 51

594

Lomgarapha—

semiclarata, 190

vestaliata, 190
Lonchaea—

corticis, 448

polita, 448
Lonchaeidae, 448
lonchaeids, 448
Longistigma caryae, 77
looper—

early brown, 190

eastern pine, 198

false hemlock, 198

fringed, 200

green larch, 191

hemlock, 198, 421

linden, 14, 193, 195

pine conelet, 198

purplish-brown, 199, 418

saddleback, 192

small conifer, 190

small pine, 190

spruce-fir, 19]

stout, 195

wildcherry, 190
Lophocampa—

caryae, 223, 224

maculata, 224
Lophodonta—

angulosa, 219

ferruginea, 219
Lopholeucaspis Japonica, 115
Lophyroplectus oblongopunctatus , 392
Lopidea incurva, 66
louse—

body, 46

crab, 46

wood, 27
lovebug, 442
Loxosceles reclusa, 30
Lucanidae, 239, 267
Lucanus elaphus, 267
Lycia ursaria, 195
Lyctidae, 42, 238, 239, 248, 249, 250,

231 ¢ 2595 250
Lyctus—10, 249, 256

africanus, 256

brunneus, 256

planicollis, 256
Lygaeidae, 68
Lygocoris tinctus, 66
Lygus lineolaris, 66
Lymantor decipiens, 361
Lymantria dispar, 12, 229-233
Lymantriidae, 227
Lymexylonidae, 38, 41, 247
Lymire edwardsii, 226
Lyonetiidae, 128
lyonetiids, 128
Lypha dubia, 150
Lytta—

aenea, 243

nuttalli, 243

polita, 243

sayi, 243
M
Machimia tentoriferella, 132
Macrarocampa marthesia, 220

Macremphytus, 409
Macrodactylus subspinosus, 270, 271
Macrodiplosis foliora, 443
Macropsis, 70
Macroxyela ferruginea, 378
Magdalis—

armicollis, 331, 332

austera, 333

austera substriga, 333

barbicornis, 333

barbita, 331

hispoides, 333

olyra, 333

pandura, 333

perforatus, 332

salicis, 333
maggot—451, 452

apple, 448

seedcorn, 450
Magicicada—

cassini, 75

septendecim, 74, 75
Malacosoma—201

americanum, 202

disstria, 204, 205

californicum, 204

californicum lutescens, 203

tigris, 203
Maladera castanea, 11, 270
Mallophaga, 46
Mantidae, 48
mantids—48, 49, 54

Carolina, 49

Chinese, 49

European, 49

narrowwinged, 49
Mantis religiosa, 49
maple phenacoccus, 92
mapleworm—

greenstriped, 210

orangehumped, 220
margarodids—88

birch, 90
Margarodidae, 37, 39, 40, 88
Marmara fasciella, 130
Martesia, 25
Matsucoccus—

alabamae, 89

gallicolus, 66, 89

macrocicatrices, 89

resinosae, 11, 88, 89
Mayetiola rigidae, 444
mayflies, 44, 45

mealybugs—91, 94, 426, 432, 434, 447,

448

apple, 91, 92

Comstock, 93, 427

grape, 93

juniper, 93

maple, 92

striped, 91

taxus, 91

twocirculi, 92
Medetera, 447
Medon, 242
Megachilidae, 36, 439
Megacyllene

antennatus, 287

caryae, 287
robiniae, 286, 287

Megalodontoidea, 377
Megalopyge opercularis, 175
Megalopygidae, 175
Megaphasma denticrus, 51
Megarhyssa—

atrata, 410

greenei, 410

macrurus, 415

macrurus macrurus, 410
Megaspilidae, 426
Megastigmus—

amelanchieris, 419

laricis, 419

specularis, 419
Megaxyela langstoni, 378
Melanaspis—

nigropunctata, 116

obscura, 116

smilacis, 116

tenebricosa, 116
Melanchra picta, 237
Melandrya striata, 247
Melandryidae, 247
melandryids, 247
Melanocallis fumipennellus, 79
Melanolestes picipes, 66
Melanolophia canadaria, 191
Melanophila—

acuminata, 284

aeneola, 284

drummondi, 284

fulvoguttata, 284, 285

notata, 284
Melanoplus—

bivittatus, 51

bruneri, 52

differentialis, 51

femurrubrum, 51

punctulatus, 52

sanguinipes, 51
Melissopus latiferreanus, 159
Melittomma sericeum, 248
Meloidae, 36, 238, 243
Melophagus ovinus, 451
Melsheimer’s sackbearer, 200
Membracidae, 37, 40, 71
membracids, 416, 428, 438
Menoponidae, 46, 88
Mesitiopterus—

floridensis, 427

kahlii, 50, 427

Mesolecanium nigrofasciatum, 95

Mesoleius tenthredinis, 403, 415
Messa populifoliella, 401
Metapelma spectabile, 419
Metaphycus eriococci, 104
Metcalfa pruinosa, 69
Meteorus versicolor, 413
Micracis—366, 367

suturalis, 367

swainel, 366
Micracisella—366, 367

nanula, 367

opacicollis, 367
Microcentrum—

retinerve, 52

rhombifolium, 52
Microsega bella, 427
Microwesia misella, 246
Micrutalis calva, 71
midges—441

biting, 441

chokecherry, 443

dogwood clubgall, 445

gall, 442, 443, 444, 445

gouty pitch, 443

gouty vein, 442

juniper, 442

red pine needle, 443
millipedes, 25, 26, 27
Mimailonidae, 200
Mindarus abietinus, 78
miners, leafblotch, 129
Minthea rugicollis, 11, 256
minute egg parasite, 416
Mioptachys, 240
Miridae, 66
Misogada unicolor, 223

mites—25, 26, 28, 30, 47, 109, 111,

115; 416, 119, 122,442
itch, 32
maple bladdergall, 31, 32
pearleaf blister, 31
pine bud, 32
southern red, 31
spider—30, 245
carmine, 31
fourspotted, 31
Schoene, 31
spruce, 30
Mollusca, 25
Monarthropalpus buxi, 12, 442
Monarthrum—
fasciatum, 370
mali, 370
Monellia—
caryella, 79
costalis, 79
microsetosa, 79
nigropunctata, 79
Monocesta coryli, 262
Monochamus—312, 362
carolinensis, 315
maculosus, 315
marmorator, 314
notatus, 314
scutellatus, 313, 314
titillator, 313
Monocteninae, 383
Monoctenus—
fulvus, 384
suffusus, 384
Monodontomerus—420
aereus, 231, 420
dentipes, 396, 420
indiscretus, 420
minor, 420
montivagus, 420
Monophylla terminata, 275
Moodna ostrinella, 187
Mordellidae, 248
Mordwilkoja vagabunda, 80
mosquitoes, 440, 441

596

moths—123, 124

acorn, 133

American dagger, 236

apple fruit, 140

bagworm, 126, 127

blastobasid, 133

browntail, 13, 233, 235, 413, 414, 416,
420, 452, 453, 454

buck, 208

carpenterworm, 145, 148

casebearer, 133

cecropia, 206, 420

cherry scallop shell, 190, 426

clearwing, 141
poplar, 141

cottonwood dagger, 236

cynthia, 206

elm dagger, 236

ermine, 140

European pine shoot, 1, 10, 13, 18,
149, 150, 414, 415, 416, 417, 419,
421

eyespotted bud, 154

false-sphnix, 218

flannel, 175
crinkled, 176

gelechiid, 135

geometrid, 187

giant silkworm, 206

gypsy, 1, 7, 13, 14, 17, 42, 64, 66,
124, 229-233, 239, 240, 414, 416,
418, 419, 426, 452, 453, 454

hag, 177

hawk, 212

hepialid, 124

hornet, 141

hummingbird, 212

imperial, 212

io, 208

lappet, 206
velleda, 206

leafroller, 148

leopard, 145, 146

luna, 207

mimallonid, 200

Nantucket pine tip, 17, 151, 152, 416,
419, 421, 428

Nevada buck, 208

northern pitch twig, 154

notodontid, 215

nun, 453

olethreutid, 148, 160

oriental, 177, 427, 453

owlet, 236

pecan bud, 160

pecan carpenterworm, 146

pepper-and-salt, 200

pine bud, 136

pine candle, 137

pine tube, 169, 170

pitch pine tip, 151, 421

pitch twig, 153

polyphemus, 207

poplar dagger, 236

poplar petiole gall, 125

poplar tentmaker, 215, 216

promethea, 207

pyralid, 94, 95, 98, 99, 100

pyraustine, 178
regal, 211
royal, 208

satin, 13, 233, 234, 413, 414, 416, 419,

426, 453
saturniid, 421
shield bearer, 125
slug caterpillar, 176
smeared dagger, 236
southwestern pine tip, 153
spear-marked black, 190
sphinx, 212-215
spruce bud, 157
spruce seed, 158
subtropical pine tip, 153
tent caterpillar, 201
tiger, 223
tussock, 227
dark, 228
definite-marked, 228
Douglas-fir, 14, 17, 426
hickory, 223, 224
pale, 224, 225
pine, 228
rusty, 227
spotted, 224
sycamore, 225

whitemarked, 14, 227, 228, 420, 421,

426
underwing, 236
walnut shoot, 181
western pine tip, 152, 421, 428
winter, 14, 189
Zimmermann pine, 182, 183
mud daubers, 438
Musca domestica, 450
Muscidae, 450
Muscina, 451
Mutillidae, 428
Mymaridae, 416
mymarids, 416
myriapods, 435
Myrmeleontidae, 48
Myzocallis—
bellus, 79
discolor, 79
melanocera, 79
punctatus, 79

N
Nabidae, 66
Nabis sordidus, 66
Nacerdes melanura, 11, 247
Nacophora quernaria, 195
Nadata gibbosa, 219
Nalepella tsugifoliae, 32
Narceus americanus, 27
Neacanthocinus—

obsoletus, 304

pusillus, 304
needleminer—

pine, 136

spruce, 149
Neichnea laticornis, 275
Nematidium filiforme, 247
Nematocampa limbata, 195
Nematus—

abbotii, 406

carpini, 406

erythrogaster, 406
fulvicrus, 406
hudsoniimagnus, 406
limbatus, 406
oligospilus, 406
ostryae, 406
pinguidorsum, 406
salicisodoratus, 405
tibialis, 406
ventralis, 405
viridescens, 406
Nemonychidae, 317, 318
Nemorimyza posticata, 450
Neochlamisus platani, 266
Neoclytus—
acuminatus, 294
caprea, 294, 295
fulguratus, 295
Jouteli jouteli, 295
mucronatus mucronatus, 295, 296
mucicatulus muricatulus, 295
scutellaris, 295
Neodiprion—384, 394, 415
abbotii, 390, 391
abietis, 390
americanum, 387
compar, 394
dubiosus, 391
excitans, 393
hetricki, 394
lecontei, 384, 415
maurus, 394
merkeli, 394
nanulus nanulus, 389, 415
nigroscutum, 394
pinetum, 390
pinusrigidae, 394, 417
pratti, 386
pratti banksianae, 386
pratti paradoxicus, 387, 415, 417
pratti pratti, 385
rugifrons, 391
sertifer, 13, 391, 392, 415
swainei, 388, 415
taedae linearis, 387, 388, 389
taedae taedae, 387
virginianus, 391
warrenl, 388
Neolecanium cornuparvum, 96
Neolygus invitus, 66
Neoptychodes trilineatus, 308
Neosteingelia texana, 90
Neotermes—
castaneus, 62
jouteli, 62
Nephopteryx—
subcaesiella, 186
subfuscella, 186
virgatella, 186
Nepticula, 124
Nepticulidae, 38, 124
nepticulids, 124
Nepytia—
canosaria, 198
pellucidaria, 198
semiclusaria, 198
Nerice bidentata, 219
Neuroptera, 6, 47, 416

597,

Neuroterus—
floccosus, 424
noxiosus, 424
quercusbatatus, 424

Neurotoma—
crataegi, 381
fasciata, 381
inconspicua, 380

Nicobium hirtum, 254

Nicrophorus, 241

Nites betulella, 132

Nitidulidae, 243

nitidulids, 22, 108, 121, 224

Noctuidae, 35, 236

noctuids, 237

Nomadinae, 439

Norape ovina, 176

Notodonta simplaria, 223

Notodontidae, 215

Nuculaspis—
californica, 116
pseudomeyeri, 117
tsugae, 117

Nudobius, 242

Nymphalidae, 173

Nvymphalis—
antiopa, 174
vau-album, 174

O
oak-bark scaler, 309
oak-bark scarrer, 292
oak beauty, 195
oakworm—
orangestriped, 209, 210, 416
pinkstriped, 209, 210
redhumped, 219, 220
spiny, 208
Oberea—299, 300
ferruginea, 300
myops, 300
ocellata, 300
pallida, 300
ruficollis, 300
schaumii, 300
tripunctata, 299
ulmicola, 300
Obolodiplosis robiniae, 445
Obrussa—
ochrefasciella, 125
sericopeza, 125
Odonata, 45
Odontopus calceatus, 334, 335
Odontota dorsalis, 264
Odontotaenius disjunctus, 242
Oecanthus—
exclamationis, 53
fultoni, 53
latipennis, 53
pini, 53
Oecophoridae, 132
oecophorids, 132
Oedemeridae, 42, 247, 248
Oeme rigida rigida, 309
Oestridae, 440
Oiketicus abbotii, 126
Olcerlostera angelica, 201

Olesicampe benefactor, 403, 415

olethreutid, 148

598

Olethreutinae, 37, 148
Olethreutes—

permundana, 160

quadrifidum, 160
Olfersia fumipennis, 451
Oligocentria lignicolor, 222
Oligomerus—

alternans, 254

obtusus, 254
Oligonychus—

aceris, 31

bicolor, 30

boudreauxi, 31

cunliffei, 31

ics, 31

letchworthi, 31

milleri, 30

newcomeri, 31

propetes, 31

ununguis, 30
Olisthaerus, 242
Oncideres—300

cingulata, 301

pustulatus, 301

texana, 301
Oncometopia orbona, 71
Oniscus asellus, 27
Ooctonus aphrophorae, 416
Ooencyrtus—

ennomophagus, 196, 418

kuvanae, 231, 418, 419

trinidadensis, 418
Operophtera—189, 190

bruceata, 189

brumata, 189
Opiliones, 29
Oracella acuta, 94
Orchesia castanea, 247

Orgilus obscurator, 150, 414, 415

Orgyia—
antiqua, 12, 227
definita, 228
leucostigma, 227, 228
pseudotsugata, 426
Orius insidiosus, 66
Ornithoica, 451
Ornithomyia, 451
Orthezia—
pseudinsignis, 90
tillandsiae, 90
Ortheziidae, 90
Orthopleura damicornis, 275

Orthoptera—8, 11, 35, 37, 48, 419, 420,

421, 452
Orthosia hibisci, 237
Orthosoma brunneum, 315
Orthotomicus caelatus, 358
Orussidae, 377, 411
orussids, 377, 411
Oscinella coxendix, 449
Osmoderma—

eremicola, 273

scabra, 273
Ostomidae, 243
Otiorhynchus—

ovatus, 11, 321

sulcatus, 11, 321

P
Pachybrachis—
carbonarius, 266
othonus, 266
peccans, 266
tridens, 266
Pachylobius picivorus, 322, 326, 327
Pachypsylla—
celtidisastericus, 76
celtidisgemma, 76
celtidisinteneris, 76
celtidismamma, 76
celtidisvesicula, 76
venusta, 76
Packardia geminata, 178
Paleacrita vernata, 193, 194
Palexorista—
bohemica, 453
inconspicua, 453
palmerworm, 137
palmetto billbug, 337
Palthis angulalis, 237
Pamphiliidae, 378
Pamphilius phyllisae, 380
Pandemis—
lamprosana, 161
limitata, 161
Panthea—
acronyctoides, 237
furcilla, 237
Pantographa limata, 178, 179
Paonias—
excaecatus, 215
myops, 215
Papaipema furcata, 237
Papilio—
cresphontes, 173
glaucus, 173
troilus, 173
Papilionidae, 124, 173
Paraclemensia acerifoliella, 125
Paradiplosis tumifex, 442
Parallelodiplosis florida, 445
Parandra brunnea brunnea, 307
Paranthrene—
dollii, 144
simulans, 144, 145
tabaniformis, 145
Parasetigena silvestris, 231, 453
Parasierola punctaticeps, 427
Parastasia brevipes, 273
Parectopa vobiniella, 131
Paria—
quadrinotata, 266
sexnotata, 266
Parornix geminatella, 131
Parthenolecanium—
corni, 96
fletcheri, 96
quercifex, 96
Passalidae, 242
passalus, horned, 242
Pealius azaleae, \\, 87
Pediculoides ventricosus, 136
Pediculus humanus humanus, 46
Pelecinidae, 426
Pelecinoidea, 426
Pelecinus polyturator, 426

Pelectoma flavipes, 254
Pelidnota punctata, 273
Peliococcus serratus, 94
Pemphigus populitransversus, 80
Pentatomidae, 63
Penthimia, 70
Pergidae, 381
Periclista, 402
Perilampus hyalinus, 420
Periphyllus—
americanus, 79
lvropictus, 79
negundinis, 78
Pero—
honestaria, 197
morrisonaria, 197
persimmon psylla, 77
Petalium—
bistriatum, 254
seriatum, 254
Petrova—
albicapitana, 154
comstockiana, 153
houseri, 154
pallipennis, 154
taedana, 153
Phaenicia sericata, 451
Phanerotoma, 159
Phanomeris phyllotomae, 399, 414
Phasgonophora sulcata, 277, 421
Phasmatidae, 37, 49
Phenacoccus—
acericola, 11, 92
aceris, 91
dearnessi, 92
Pheosia rimosa, 218
Phigalia titea, 193
Philaenus spumarius, 74
Philonthus—242
cyanipennis, 242
Philopteridae, 46
Phloeonomus, 242
Phloeopora, 242
Phloeosinus—351, 352
canadensis, 352
dentatus, 351
pini, 352
scopulorum neomexicanus, 352
taxodii, 351
Phloeotribus—
dentifrons, 350
frontalis, 350, 351
liminaris, 350
Phlogistosternus dislocatus, 275
Phlyctaenia coronata, \78
Phobetron pithecium, \77
Phobocampe disparis, 231
Phormia regina, 451
Phryxe vulgaris, 194
Phycitinae, 178, 180
Phyllaphis fagi, 79
Phyllobius—
intrusus, 12, 320
oblongus, 12, 320
Phyllocnistis—
liquidambarisella, 131
liriodendronella, 131
magnoliella, 131
populiella, 131

599

Phyllocolpa, 409
Phyllodesma americana, 206
Phyllonorycter—

crataegella, 130

lucetiella, 130

robiniella, 130

salicifoliella, 130

tremuloidiella, 130

trinotella, 130
Phyllophaga—267, 268, 269, 270

crenulata, 269

drakei, 268

forsteri, 269

implicita, 269

luctuosa, 268

micans, 269

prunina, 269

prununculina, 268

rugosa, 268

tristis, 268
Phylloxera—86, 337

caryaecaulis, 86

devastatrix, 86

notabilis, 86

nyssae, 86

rileyi, 86
phylloxeras—

hickory gall, 86

pecan leaf, 86

pecan, 86
Phylloxeridae, 37, 39, 40, 86
Phymatodes—

dimidiatus, 306

testaceus, 306

varius, 306
Physocnemum—

andreae, 306

brevilineum, 307

violaceipenne, 307
Physokermes—

hemicryphus, 98

piceae, 98
Phytobia—

amelanchieris, 449

pruinosa, 449

pruni, 449

setosa, 449
Phytomyza ilicicola, 449, 450
pigeon tremex, 410, 415, 422
Pikonema—

alaskensis, 404, 405

dimmockii, 405
Pilophorus walshii, 66
pine colaspis, 263
pine needle sheathminer, 140
Pineus—

coloradensis, 86

floccus, 86

pinifoliae, 86

similis, 86

strobi, 11, 84, 85
Pinnaspis—

aspidistrae, 117

strachani, 117
Pipunculidae, 447
Pissodes—322, 327, 331

affinis, 331

approximatus, 329, 330

600

dubius, 331
fiskei, 331
nemorensis, 330
rotundatus, 331
strobi, 328, 330
Pitedia uhleri, 64
Pityoborus comatus, 364
Pityogenes—
hopkinsi, 357
meridianus, 357
plagiatus, 357
Pityokteines sparsus, 357 |
Pityophthorus—362, 363, 366 |
annectens, 364 |
balsameus, 364
biovalis, 364
cariniceps, 364
crinalis, 363
dentifrons, 364
intextus, 364
lautus, 363
liquidambarus, 363, 421
opaculus, 363
puberulus, 364
pulchellus, 364
pulicarius, 363
ramiperda, 364
scriptor, 363
Placusa, 242
Plagiodera versicolora, 12, 264, 265
Plagiognathus
albatus, 67
delicatus, 66
Plagiometriona clavata, 266
Plagodis—
kuetzingi, 195
serinaria, 195
planthoppers, flatid, 69
plantlice—77
jumping, 76
Platoeceticus gloveri, 118
Platybregmus canadensis, 254
Platycerus quercus, 267
Platycotis vittata, 71
Platygaster pini, 443
Platygastridae, 426
Platypodidae, 38, 248, 368, 375
Platypus—
compositus, 376
flavicornis, 375, 376
parallelus, 377
quadridentatus, 375
Platytetranychus—
multidigituli, 31
thujae, 31
Plecia nearctica, 442
Plecoptera, 45
Plectrodera scalator, 289, 290
Plegaderus, 241
Pleolophus basizonus, 414, 415
Pleroneura brunneicornis, 378
Plutellidae, 138
plutellids, 138
Podapion gallicola, 331, 332
Podisus maculiventris, 64
Podosesia syringae, 143
aureocincta, 143, 144

Poecilonota—

cyanipes, 285

thureura, 285
Polistes, 437
Polycaon stouti, 259
Polydrusus impressifrons, 12, 321
Polygonia—

comma, 173

interrogationis, 173, 174
Polygraphus rufipennis, 352
Polynema striaticorne, 416
Polyphylla—

hammondi, 269

occidentalis, 269

variolosa, 269
Polystepha pilulae, 445
Pompilidae, 435, 438
Ponana, 70
Pontania, 409
Popillia japonica, 12, 271
poplar tentmaker, 215, 216
poplar-gall saperda, 298
Priobium sericeum, 254
Priocera castenae, 275
prionid, brown, 315
Prionoxystus—

macmurtrei, 148

robiniae, 146, 147
Prionus—309

imbricornis, 309, 310

laticollis, 309

pocularis, 309
Pristaulacus—

bilobatus, 426

rufitaris, 426
Pristiphora—

chlorea, 404

erichsonii, 13, 402, 403

geniculata, 13, 404, 405

siskiyouensis, 404

sycophanta, 404
Probole—

alienaria, 195

amicaria, 195
Prochoerodes transversata, 199
Prociphilus—

bumelia, 78

corrugatans, 80

fraxinifolii, 80

longianus, 80

tessellatus, 80
Proctotrupoidea, 426
Prodiplosis morrisi, 445
Profenusa—

alumna, 401

canadensis, 401

lucifex, 401

thomsoni, 13, 401
Prolimacodes—

badia, 178

scapha, 178
prominent—

elm, 222

saddled, 220, 221, 416, 426
Prorhinotermes—61

simplex, 63
Prosapia bicincta, 74

Proteoteras—
aesculana, 156
moffatiana, 156
willingana, 156, 157

Protoboarmia porcelaria indicataria, 200

Protocalliphora, 451
Protozoa, 26
Pselactus spadix, 337
Pselaphorhynchites
aeratus, 319
cyanellus, 319
Psen, 438
Pseudaonidia—
duplex, 118
paeoniae, 117
Pseudaphycus malinus, 93
Pseudaulacaspis—
cockerelli, 119
pentagona, 118
prunicola, 118
Pseudexentera oregonana, 160

Pseudisobrachium prolongatum, 427

Pseudococcidae, 91
Pseudococcus—

comstocki, 11, 93

maritimus, 93

obscurus, 93
Pseudolucanus capreolus, 267
Pseudophilippia quaintancii, 98
Pseudopityophthorus—22, 362

asperulus, 363

minutissimus, 362

pruinosus, 363

pubescens, 363
Pseudosciaphila duplex, 160
Pseudoscorpiones, 28
pseudoscorpions, 28, 29, 414
Pseudotachinomyia slossonae, 194
Pseudothysanoes—

dislocatus, 357

lecontei, 356

rigidus, 356
Psilocorsis

cryptolechiella, 132

quercicella, 132

reflexella, 132
psocids, 46
Psocoptera, 46
Psyche casta, 126
Psychidae, 126
Psylla—76

annulata, 77

buxi, 76

carpinicola, 77

floccosa, 77

galeaformis, 77

trimaculata, 77
Psyllidae, 39, 76
psyllids—76, 442, 447

blistergall, 76

boxwood, 76

budgall, 76

petiolegall, 76
Pterocallis alnifoliae, 79
Pterocomma—

populifoliae, 79

salicis, 79

smithiae, 79

601

Pteromalidae, 420
pteromalids, 420
Pterophylla camellifolia, 52
Pthirus pubis, 46
Ptilinus—

pectinicornis, 254

ruficornis, 254
Ptinidae, 248, 259
Ptinus—

clavipes, 259

jur, 259
Ptosima gibbicollis, 285
Pulex irritans, 84
Pulvinaria—

acericola, 98

floccifera, 98

innumerabilis, 97
punkies, 440
Pygoleptura nigrella, 306
Pyralidae, 35, 38, 41, 178
Pyraustinae, 178
Pyrrhalta—

cavicollis, 262

decora decora, 262

luteola, 12, 260, 261

tuberculata, 262

Quadraspidiotus—
forbesi, 119
gigas, 119
Juglansregiae, 119
ostreaeformis, 120
perniciosus, 11, 120
socialis, 121
taxodii, 121
tillandsiae, 121
Quedius, 242
Quernaspis—
insularis, 121
quercicola, 121
quercus, 121
question-mark, 173, 174
quince curculio, 336

R
Raphia frater, 237
redbugs, 32
red spotted purple, 175
Reduviidae, 65
Resseliella clavula, 445
Reticulitermes—S58, 59, 61

arenincola, 58, 59

flavipes, 56, 57, 58

hageni, 58

tibialis, 58

virginicus, 58
Rhabdophaga salicis, 12
Rhagionidae, 446
Rhagium inquisitor, 315
Rhagoletis—

cingulata, 448

completa, 448

fausta, 448

pomonella, 448
Rheumaptera hastata, 190
Rhinotermitidae, 55, 60, 63
Rhipiphoridae, 238

Rhizotrogus majalis, 12, 270

Rhopalidae, 67

602

Rhyacionia—150, 153

adana, 151

aktita, 153

buoliana, 12, 149, 150

bushnelli, 152, 153

frustrana, 151, 152, 153

neomexicana, 153

rigidana, 151

sonia, 153

subtropica, 153
Rhynchaenus—

pallicornis, 334

rufipes, 334
Rhynchitidae, 317, 319
rhynchitids, 319
Rhynchophoridae, 317, 337
Rhynchophorinae, 337

Rhynchophorus cruentatus, 337

Rhyncophora, 317
Rhyssa, 410

Romalea microptera, 52
Roystonea elata, 68

S
saddled prominent, 220, 221, 416, 426

Samia cynthia, 12, 206
Sannina uroceriformis, 143
Saperda— 295
calcarata, 279, 295, 297
candida, 297
cretata, 298
discoidea, 298
favi, 298
imitans, 298
inornata, 298
lateralis, 298
moesta, 298
mutica, 298
obliqua, 298
tridentata, 298, 299
vestita, 297
Sapvgidae, 429
sapygids, 429
Sarcophaga—
aldrichi, 205, 451
houghi, 452
sarcophagid, 420
Sarcophagidae, 451
Sarcoptes scabiei, 32
Sarcoptidae, 32
Saturniidae, 206

sawflies—239, 273, 377, 381, 395, 398,

405, 414, 417, 420, 440
argid, 322

balsam fir, 390, 417
birch, 382

birch leafmining, 398, 399, 414

blackheaded ash, 408

blackheaded pine, 14, 393

brownheaded ash, 408

brownheaded jack pine, 391

cherry webspinning, 381
cimbicid, 382

conifer, 383

dusky birch, 406, 407
elm, 382, 383, 416

European pine, 1, 13, 14, 391, 392,

415, 417

European spruce, 13, 14, 274, 396, 397,
414, 415, 417, 453, 454

greenheaded spruce, 405

hawthorn leafmining, 401

introduced pine, 17, 18, 266, 395, 415,
417, 420, 421

jack pine, 14, 386

larch, 274, 402, 403, 415

loblolly pine, 387, 388, 389

mountain-ash, 404, 405

nesting-pine, 380

onelined larch, 409

pear, 401

pergid, 381

pine, 419

pine shoot gall, 378, 379

plum webspinning, 380

poplar, 406

poplar leafmining, 401

red pine, 389, 417

redheaded jack pine, 391

redheaded pine, 14, 384, 385, 417, 454

sand pine, 386

scarlet oak, 401

slash pine, 394

spotted loblolly pine, 387

stem, 412

striped alder, 407

Swaine jack pine, 14, 388, 420

threelined larch, 409

Virginia pine, 14, 385, 417

webspinning, 378

white pine, 390

willow, 405

willow shoot, 412

yellowheaded spruce, 404, 405, 416

sawyers—

balsam fir, 314
northeastern, 314
southern pine, 313, 315
spotted pine, 315
whitespotted, 313, 314

scales—22, 87, 106, 127, 245, 418, 432,

434, 442, 447, 448
Alabama pine, 89
armored, 106

azalea bark, 103
barnacle, 94

beech, 21, 101, 102, 126
bifasciculate, 111

black pineleaf, 116, 117
black willow, 111

calico, 94, 95

camellia, 115

camphor, 118

Canadian pine, 89
common falsepit, 104, 105
cottony camellia, 98
cottony maple, 97
cottony maple leaf, 98
cottony taxus, 98
cryptomeria, 107
dentate, 123
dictyospermum, 111
dogwood, 108, 109

elm armored, 111, 112
elm scurfy, 108

elongate hemlock, 113, 117

ensign, 90
euonymus, 121, 122
European elm, 103, 418
European fruit, 120
false oak, 121
falsepit, 104, 105
fern, 117

fiorinia, 114

fiorinia hemlock, 113
Fletcher, 96

Florida red, 111
Florida wax, 94
Forbes, 119

gall-like, 100
gloomy, 116

golden oak, 105, 419
greedy, 114
hemlock, 106, 107
hickory, 113

holly pit, 106

Indian wax, 94

island oak scale, 121
Japanese fiorinia, 114
Japanese maple, 115
Japanese wax, 94
juniper, 108

latania, 114

lesser snow, 117
magnolia, 96
margarodid, 88, 90
Maskell, 115
McComb, 113
minute cypress, 108
obscure, 116
oleander, 107
oleander pit, 106
Osborn, 112
oystershell, 114, 115, 419
Parrott, 105

peony, 117, 118
pine, 109

pine needle, 109, 110
pine tortoise, 99, 100
pine twig gall, 89
pit, 105

Putnam, 112

redbay, 107

red pine, 66, 88, 89, 246
round conifer, 107
San Jose, 120, 121, 419
scurfy, 109
shortneedle evergreen, 117
soft, 94

sour-gum, 110
spruce bud, 98, 99
striped pine, 100
sweetgum, 112

tea, 13

terrapin, 95

tuliptree, 99

Virginia pine, 100
walnut, 119, 120
wax, 94, 95

western oak, 121
white peach, 118
white prunicola, 118
willow, 119

willow scurfy, 111
woolly pine, 98

603

Scaphoideus luteolus, 70
Scapteriscus—

abbreviatus, 53

acletus, 53

vicinus, 53
Scarabaeidae, 35, 36, 239, 267
scarabaeids, 428, 429
scarabs, 267
Scarites subterraneus, 240
Scelionidae, 426
Schizolachnus piniradiatae, 78
Schizonotus latus, 265, 421
Schizura—

concinna, 222

ipomoeae, 223

leptinoides, 223

unicornis, 223
Sciaphillus asperatus, 12, 320
Sciaridae, 442
Sclerodermus, 427

Scobicia—
bidentata, 257
declivis, 257

Scoliidae, 428
scolds, 428
Scolioidea, 428
Scolopendridae, 28

Scolytidae, 35, 37, 38, 41, 42, 238, 248,
317, 337, 338, 339, 363, 364, 366

Scolytinae, 338, 352, 366
Scolytus—339, 352, 355

fagi, 356

mali, 354

multistriatus, 12, 352, 353

muticus, 356

piceae, 356

quadrispinosus, 355

rugulosus, 356
Scorpiones, 28
scorpions— 25, 26, 28, 54

striped, 28, 29
screwworm—17, 451

secondary, 451
Scudderia—

curvicauda, 52

furcata, 52
Scymnus—

impexus, 246

lacustris, 246
seedworm—

eastern pine, 159

longleaf pine, 159

slash pine, 159
Seirarctia echo, 226
Semanotus—

ligneus, 307

litigiosus, 307
Semiothisa—

bisignata, 191

granitata, 191

ocellinata, 191

sexmaculata, 191

signaria dispuncta, 191
Semudobia, 445
Sequoiomyia cupressi, 443
Serica—269

intermixta, 269

peregrina, 428

604

sericae, 269
tristis, 269
vespertina, 269
Serropalpus substriatus, 247
Sesia—
apiformis, 12, 141
tibialis, 141
Sesiidae, 37, 38, 41, 141
sheep ked, 451
shield bearers, 125
resplendent, 126
shipworms, 25, 26
Siagonium, 242
Sibine stimulea, 176
Silpha, 241
Silphidae, 241
Simuliidae, 441
Sinea spinipes, 66
Sinodendron rugosum, 267
Sipalia, 242
Siphonaptera, 48
Sirex—
areolatus, 410
cyaneus, 410
edwardsii, 410
Juvencus, 13, 410
nigricornis, 410

Siricidae, 37, 409, 411, 412, 422

Siricoidea, 409
skeletonizer—

apple-and-thorn, 145

birch, 128

maple trumpet, 157

oak, 128, 129, 421

palm leaf, 135
skippers—123, 124, 172

silverspotted, 172
Smerinthus jamaicensis, 215
Smodicum cucujiforme, 299
Solenopsis—435

invicta, 13, 435

richteri, 435
solpugids, 54
sowbug, common terrestrial, 27
Spanish moss orthezia, 90
spanworm—

Bruce, 189

cleft-headed, 200

elm, 196, 197, 232, 418, 426, 452

large maple, 199
Sparganothis—
acerivorana, 161, 171
diluticostana, 161
pettitana, 161
reticulatana, 161
sulfureana, 161
tristriata, 161
Spathimeigenia spp., 385, 394
Sphaeroma, 27
Sphecidae, 42, 438
Sphecius speciosus, 438
Sphecoidea, 438
Sphingidae, 212
sphinx—212, 213
blinded, 215
catalpa, 213, 214, 426
elm, 213
false-, 218

great ash, 214
laurel, 214
pine tree, 214
smalleyed, 215
twinspot, 215
walnut, 215
waved, 213
Sphinx—
chersis, 214
drupiferarum, 214
kalmiae, 214
luscitiosa, 214
spider mite destroyer, 246

spiders—25, 26, 28, 29, 54, 148, 414,

435

black widow, 29

brown recluse, 30
Spilochalcis—

albifrons, 421

flavopicta, 421

mariae, 421
Spilococcus juniperi, 93
Spilonota ocellana, 12, 154
Spissistilus festinus, 71
spittlebugs—69, 72, 447

alder, 74

dogwood, 74

meadow, 74

pecan, 74

pine, 72

Saratoga, 73, 416
springtails, 44
spruce epizeuxis, 237
spruce harlequin, 237
spruce scolytus, 356
Stagmomantis

carolina, 49

floridensis, 49
Staphylinidae, 238, 242
Staphylinus, 242
Stegobium paniceum, 252
Stenodontes dasytomus dasytomus, 315
Stenoscelis brevis, 337
Stephanidae, 413
stephanids, 413
Stephanitis rhododendri, \1
Stephanoderes, 367
Stephanopachys—

cribratus, 257

densus, 257

hispidulus, 257

rugosus, 257

substriatus, 257
Sterictiphora, 382
Stethorus punctum, 246
Sthenopis—

argenteomaculatus, 124

thule, 124
‘Stictia carolina, 438
Stictocephala—

bisonia, 71

militaris, 71
Stictoleptura canadensis, 304
Stilbosis ostryaeela, 135
Stiretus anchorago, 64
Stomoxys calcitrans, 450
stoneflies, 45, 440
Strangalepta vittata, 306

Stratiomyidae, 445
Strongylium, 247
Strophiona nitens, 304
sumac datana, 217
swallowtail—
spicebush, 173
tiger, 173
zebra, 173
Symmerista—
albifrons, 219
canicosta, 219, 220
leucitys, 220
Symphyta, 377, 412
Symphoromyia, 446
Synanthedon—
acerni, 142
acerrubri, 143
bolteri, 143
castaneae, 143
geliformis, 142
pictipes, 143
pini, 142
proxima, 143
pyri, 142
rhododendri, 142
rubrofascia, 143
sapygaeformis, 143
scitula, 141
sigmoidea, 143
virburni, 143
Syneta ferruginea, 266
Syrphidae, 447
Systena marginalis, 265

T
Tabanidae, 445
Tabanus—
abactor, 445
atratus, 445
lineola, 445
nigrovittatus, 445
quinquevittatus, 445
sulcifrons, 445
Tachinidae, 452
tachinids, 420, 452
Tachinus, 242
Tachydromia, 447
Tachyporus, 242
Tachyta, 240
Taedia gleditsiae, 67
Tarsostenus univittatus, 275
tawny emperor, 175
Taxodiomyia cupressiananassa, 443
Telamona—
decorata, 71
reclivata, 71
Telenomus—190, 426
alsophilae, 189, 426, 427
bifidus, 426
californicus, 426
catalpae, 426
clisiocampae, 426
coelodasidis, 222, 426
dalmani, 426
droozi, 196, 426
Temelucha interruptor, 415
Temnochila virescens, 243
Tenebrionidae, 41, 247, 248

605

Tenebroides—
bimaculatus, 243
corticalis, 243
Tenodera—
aridifolia sinensis, 49
augustipennis, 49
Tenthredinidae, 36, 38, 39, 398
Tentredinoidea, 381
Tephritidae, 34, 447
Teredinidae, 43
Teredo, 25
termites—19, 54, 55, 241, 429
eastern subterranean, 56, 57
Formosan subterranean, 59, 60
nonsubterranean, 60, 61
subterranean, 9, 55, 56, 252
Tethida cordigera, 408
‘Tetracis cachexiata, 199
Tetraleurodes mori, 87
Tetralopha—
asperatella, 180
melanogrammos, 180
militella, 180
robustella, 179
Tetraneura ulmi, 80
Tetranychidae, 30, 35, 36
Tetranychus—
canadensis, 31
cinnabarinus, 31
homorus, 31
magnoliae, 31
schoenei, 31
Tetrastichus—
brevistigma, 262, 418
holbeini, 418
rugglesi, 418
turionum, 417
Tetropium cinnamopterum, 308
Tettigoniidae, 52
Tetyra bipunctata, 63
Thanasimus dubius, 275
Thaumastocoridae, 68
Thecodiplosis piniresinosae, 443
Thelia bimaculata, 71, 428
thrips—46, 111, 119
flower, 47
slash pine flower, 46
Thyridopteryx ephemeraeformis, 126, 127
Thysanoes—356, 366, 367
berchemiae, 367
fimbricornis, 367
lobdelli, 367
Thysanoptera, 46
Thysanura, 44
Tibicen canicularis, 75
ticks—25, 28, 32
American dog, 32
blacklegged, 33
tilehorned prionus, 309, 310
timberworm—248
chestnut, 248
oak, 318
sapwood, 248
Tingidae, 64
Tinocallis—
kahawaluokalani, 79
ulmifolii, 79

606

Tiphia—
asericae, 428
inornata, 428
popilliavora, 272, 428
vernalis, 272, 428
Tiphiidae, 428
Tipulidae, 426
Tolmerus notatus, 446
Tolype—
laricis, 206
velleda, 206

. Tomolips quercicola, 337
Tomostethus multicinctus, 408

Tortricidae, 35, 39, 148
Tortricidia flexuosa, 178
Tortricinae, 160

tortrix, large aspen, 168, 169

Torymidae, 419
torymids, 419
Torymus rugglesi, 419
Toumeyella—
liriodendri, 99
parvicornis, 99, 100
pini, 100
virginiana, 100

Townsendiellomyia nidicola, 453
Toxotrypana curvicauda, 448

Trachykele lecontei, 285
Trachysida mutabilis, 306
Tragosoma depsarius, 307
treehoppers, 71

buffalo, 71

threecornered alfalfa hopper, 71

twomarked, 71
Tremex columba, 410, 411

Trichiocampus viminalis, 13, 406
Trichiosoma triangulum, 383

Trichiotinus, 273
Trichodestidae, 46

Trichogramma minutum, 165, 169, 416

Trichogrammatidae, 416
Tricorynus, 255
Trigonarthris—
minnesotana, 306
proxima, 306
Trigonura elegans, 421
Trilobomvyza pleuralis, 450
Trioza—
diospyri, 77
magnoliae, 77
tripunctata, 77
Trisetacus—
cupressi, 32
pini, 32
Trogositidae, 243

Trogoxylon parallelopipedum, 256

Trombiculidae, 32

Tropideres—
dorsalis, 318
fasciatus, 318

Tropidosteptes amoenus, 66

Trypodendron—339, 371
betulae, 371
lineatum, 371
retusum, 371
rufitarsis, 371
scabricollis, 371, 372

tubemaker— maple, 180

alder, 181 mimosa, 138, 139

birch, 181 oak, 162, 163
Tuberolachnus salignus, 79 pine, 179
Tylocerina nodosa, 304, 305 pine false, 378, 379
Tylonotus bimaculatus, 308 weevils—251, 275, 317, 319
Tymnes tricolor, 266 alfalfa, 320
Typhlocyba, 70 apple flea, 334
twig pruner, 302 arborvitae, 320

Asiatic oak, 322
] ; balsam bark, 331
D) 35
Ree ARAE O56 Bete black elm bark, 331, 421
DRG Rae ; black vine, 321

; cossonid, 243
eS jon cotton boll, 320

5 Couper’s collar, 326
eee deodar, 24, 330

fungus, 317
My grain, 320
Valentinia glandulella, 133 hazelnut, 333
Vanduzeea arquata, 7\ large chestnut, 333
Vasates— leaf-rolling, 319
aceris-crummena, 31 New York, 331
quadripedes, 31, 32 northern pine, 329
Velataspis dentata, 123 pales, 17, 323, 324
Vespa crabro germana, 436 pecan, 333
Vespidae, 42, 435, 436 pine-flower, 318
Vespoidea, 435 pine gall, 331, 332
Vespula— pine root collar, 324, 325
arenaria, 436, 437 pine root tip, 326
germanica, 437 pitcheating, 326, 327
maculata, 436 red elm bark, 331, 332, 421
maculifrons, 436, 437 small chestnut, 333
_Squamosa, 437 small spruce, 331
VICETOY, i southern pine root, 326
Villa sinuosa sinuosa, 386, 388, 394 strawberry root, 321
W twobanded Japanese, 322
walkingsticks—49, 50, 427 Warren’s collar, 326
giant, 51 white pine, 275, 320, 328, 329, 419,
twostriped, 51 421, 447, 448, 449
wasps—54, 377, 421, 426, 428, 436 willow flea, 334
aulacid, 426 white admiral, 175
bethylid, 427 whiteflies—87, 418, 426
chalcidoid, 91, 92, 93, 94, 95, 97, 98, azalea, 87
99, 100, 103, 104, 106, 107, 108, citrus, 87
109. TO tls 112,113, 116; 120, mulberry, 87
127 rhododendron, 87
cuckoo, 427 Winthemia—
cynipid gall, 422 cecropia, 207
dryinid, 428 datanae, 217
ensign, 425 wireworms, 273
gold, 427 woollyworm, butternut, 409
megaspilid, 426 x
parasitic, 101, 115, 117, 118, 119, 120, Xanthonia decemnotata, 266
12122 Xanthoteras quercusforticorne, 425
pelecinid, 426 Xenopsylla cheopis, 48
platygastrid, 93, 426 Xenotemna pallorana, 172
potter, 436 Xestobium rufovillosum, 12, 254
sand, 438 Xiphydria—4l11, 426
scelionid, 426 abdominalis, 411
spider, 438 hicoriae, 411
tiphiid, 428 maculata, 411
yellowjacket, 436, 437 mellipes, 411
webworms— 14 tibialis, 411
ailanthus, 140 Xiphydriidae, 411
fall, 14, 66, 225, 226, 380, 416, 420, xiphydriids, 411
_ 426 X vela—
juniper, 138, 139 alpigena, 378

607

bakeri, 378
dodgei, 378
gallicaulis, 378, 379
minor, 378
obscura, 378
styrax, 378
Xyelidae, 377
xyelids, 377
Xvlastodoris luteolus, 68
Xyleborinus saxeseni, 368, 374
X vleborus—372, 374
affinis, 374
celsus, 373
devexulus, 374
dispar, 373, 375
ferrugineus, 373, 374
intrusus, 374
lecontei, 374
obesus, 374
obliquus, 374
opimus, 374
planicollis, 374
pubescens, 374
rubricollis, 374
sayl, 374
tachygraphus, 374
validus, 374
viduus, 374
volvulus, 374
xylographus, 374
X vletinus—
harrisii, 254
peltatus, 252
Xvlobiops basilaris, 23, 257, 258
Xvlococculus betulae, 90
X vlocopa—
micans, 440
virginica, 439
Xylocopinae, 439
Xylophagidae, 445
X vlophagus—
abdominalis, 445
lugens, 445
Xyloryctes jamaicensis, 273
X ylosandrus—
compactus, 375
crassiusculus, 375
germanus, 374
morigerus, 375
zimmermanni, 374
Xyloterinus politus, 368, 371
X ylotrechus—
aceris, 310
annosus annosus, 312
colonus, 312
obliteratus, 311
quadrimaculatus, 310
Sagittatus sagittatus, 311
undulatus, 312

yellowjackets—436, 437, 438
eastern, 437
Yponomeutidae, 138, 140
Z
Zabrachia polita, 445
Zeiraphera—
canadensis, 157

608

improbana, 157
unfortunana, 157
Zelleria haimbachi, 140
Zelus exsanguis, 66
Zengophora scutellaris, 12, 265
Zenodochium coccivorella, 133
Zeuzera pyrina, 12, 145, 146

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