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EASTERN
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U.S. DEPARTMENT
OF AGRICULTUREe

FOREST SERVICE

MISCELLANEOUS
PUBLICATION NO. 1175

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Historic, archived document

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scientific knowledge, policies, or practices.

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LIBRARY

DC BRANCH

WHITEFORD L. BAKER

PRINCIPAL ENTOMOLOGIST, RETIRED

FOREST INSECT RESEARCH BRANCH

FOREST INSECT AND DISEASE RESEARCH DIVISION
FOREST SERVICE

EASTERN oe

INSECTS

U.S. DEPARTMENT
OF AGRICULTUREe
FOREST SERVICE

MISCELLANEOUS
PUBLICATION NO. 1175

FEBRUARY 1972

THIS PUBLICATION SUPERSEDES ‘INSECT
ENEMIES OF THE EASTERN FORESTS,"’
MISCELLANEOUS PUBLICATION NO. 657

For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402—Price $5

ACKNOWLEDGMENTS

The author gratefully acknowledges the invaluable assistance
of many of his colleagues in the Forest Insect Branch of the
USDA Forest Service in the preparation of this publication. He
is also greatly indebted to the following individuals for their
helpful reviews of different sections of it: Dr. A. E. Brower,
Maine Forest Service; Dr. R. L. Giese, Purdue University; Dr. D.
M. Benjamin, University of Wisconsin; Dr. R. F. Anderson, Duke
University; Dr. Jack L. Krall, State University of New York
College of Forestry at Syracuse; Dr. R. T. Franklin, University
of Georgia; Dr. C. E. Atwood, University of Toronto, Canada;
Dr. J. A. Slater, University of Connecticut; Dr. John A. Davidson,
Dr. T. L. Bissell, and Dr. F. E. Wood, University of Maryland;
Dr. D. L. Collins and Dr J. A. Wilcox, New York State Museum
and Science Service, Albany, New York. Finally, he wishes to
make acknowledgement to the Connecticut Agricultural Experi-
ment Station, Illinois Natural History Survey, and Duke School
of Forestry for the loan of many photographs for illustrations.

LIBRARY OF CONGRESS CATALOG NO. 76—607316

| CONTENTS

HAA ETEO GLUNC EL O10 arene ear nee et UL ee ee ean tuicss erst
Imseetswand elated Oreanisms 2. ee. mee
@laccs#Hexanoda—=INSsectsy ee ee re
GlacscseGrustacea——WiOOdNCe ee ee !
Class Diplopoda——Miallipedes: 23.) ded ,
Class-Chilopoda=—Centipedes)) 24.72 isso Sine 3
Class Arachnida—Spiders, mites, ticks, scorpions, ete. .......
Order vwAraneida=—Splders)..- 2 eee &
Order-Acarina—-Mites-and! ticks 25. 3.2.2 ne
Order ScorpionideAa—ScorpioMs oie... ene nee:
Order Pseudoscorpoinidae—F alse scorpions .........................
Order Phalangida—Daddy-long-legs 0... ae
Phylum? Mollusea=_Ship worms =< 3.60. ee
Honest MinSeCi sin tise Mee. Rete We me ed Oe ern ee | ?
Borest insect causedlosses 2 ee py Cae
MOrest IMSeCClISUEVEYS. |. ee oe Ed ile |
Forest insect control... APN eget ECL ee aw Ne oa oy atetsely
INaturaleconerolin Scrat Fi tte Seed Bee as
AMC CMC OMER Ole Met anes eee rs Ns ee
Sivacultumalscontrol et ee uy
Physical and mechanical methods of control...
Reoulatory control 20s: ee |
Biolosicalecontrolo =e ee Cy, Se a ee
Chemicalecontroles 396 ee ope
Other approaches to control =e. )
teerated=controle as hee hs ae
Practical keys to the orders, families, and genera of forest
Imsectssoasedv on types of Injury 72 2
RAMA ya VISIONS Ole KOY. ie ee Se :
important orders of forest insects. 3...
Order Anoplura—sSuckineg lice 20d
Order Mallophaga—Chewing Vice 00... icc
Order Siphonoptera—Fleas ea,
Order rhysanura—bristietails
Order Collembola—Sprinetails ©... ecccccccecceenseueee ee
Ordercchysanoptera——Eehrips (0 Se ee
Order Psocoptera—Book lice and psocids ......................... aie
Ordersneuroptera i 6.2 re OO GS
Order Ephemeroptera—Mayflies 00... ccc. nae ace
Order Plecoptera—Stonefhies oi... ccsssesesssssceeeee Nes
Order Odonata—Dragonflies and damselflies ...................... 7
Order-lsoptera—— Termites 2 ee
Order Orthoptera—Cockroaches, grasshoppers and allies
Order Hemiptera—True bugs i ceccccccccccccecsessssseeneeee |
Order Homoptera—Aphids, scales, mealybugs, etc. ._.
Order Coleoptera—Beetles .cscccecccetecsessesecssscstssstsrtestieneace
Order Lepidoptera—Butterflies, moths, skippers...
OrdernoDiptera=—Mlies ota ee :
OndersryMeNOPCerar. oe ee cone eae nies
imsectseand. Tree: Diseases: 5 nc sea cee
HP OR A LUC sO AUCC her ah ed ie eee eeu NN Os ei Mie ye
indexatosiisects: by- Host: Plants... oe
mMsecteinGdexis sen. wer We Aonsierh Sips say i nae

OMAP PWNNH

INTRODUCTION

Interest in the protection of the Nation’s forests from insects
has grown considerably in recent years. This has come about
largely because of increased awareness of the destructive capaci-
ties of insects and the heavy toll they take of our dwindling sup-
plies of commercial timber. Interest has been further heightened
by the fact that more and more people have come to realize that
much of the damage caused by insects need not necessarily hap-
pen—that given adequate knowledge of the habits and behavior
of injurious insects, it should be possible to either prevent or
reduce most of it.

Increased interest in the protection of our forests from insect-
caused losses has resulted in increased research to develop
cheaper, safer, and more effective methods of control. Much prog-
ress has been made in recent years, especially since World War II,
in studies of the biology and ecology of many important species,
leading to new or improved control methods. Much new informa-
tion has also been obtained on many other species of potential
pests. The results of these investigations appear in many publica-
tions, some of limited circulation and more or less inaccessible to
the general public.

It is the purpose of this publication, which supersedes U.S.
Department of Agriculture Miscellaneous Publication 657 (167),!
to review the literature on eastern forest insects which has been
published since 1940, the cutoff date for nearly all of the informa-
tion contained in M.P. 657. Insects discussed are those which occur
either entirely in portions of the United States lying east of the
100th meridian or which occur in both the eastern and western
halves of the country.2 The major portion of the publication is
devoted to the identification, distribution, host relationships, and
life histories of insects occurring in eastern forests.

This publication includes not only new information obtained
since 1940, but also much of the still pertinent information ob-
tained earlier and reported in M.P. 657. The common names of
insects preceding the scientific names are approved (85); the
common names following the scientific names have not been
approved.

‘Italic numbers in parentheses refer to Literature Cited, p. 503.
2A companion volume covering western forest insects is also available.

if

INSECTS AND RELATED ORGANISMS

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, milli-
pedes, centipedes, spiders, and mites. Members of the phylum are
distinguished by having the body composed of a series of more or
less similar rings or segments joined together and in having some
of these segments bear jointed legs. In certain forms, the segmen-
tations of the body may be obscure and not evident from cursory
examination. In most instances this is due to a secondary modifi-
cation of form, a result of adaptation to special modes of life
(152).

Insects comprise the most abundant and important group of
arthropods, but many other members of the phylum are also com-
mon and often important. Many of the latter are harmful, either
to trees and other vegetation or to wildlife and man. Many are so
small they are seldom seen, although enormously abundant;
others, 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 Hexapoda (In-
sects) ; Crustacea (crawfish, shrimp, crabs) ; Diplopoda (milli-
pedes) ; Chilopoda (centipedes) ; and Arachnida (spiders, mites,
and scorpions). Mention is also made of shipworms which belong
to the phylum Mollusca.

Class HEXAPODA

Insects

An insect is an air-breathing arthropod with a distinct head,
thorax, and abdomen. 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 parts of their lives in water.

The majority of insects hatch into wormlike larvae which grow
by periodically shedding the outer skin, finally transforming into
an inactive pupal stage from which the adult emerges. This sort
of development is known as complete metamorphosis. The imma-
ture stages of 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.
This represents 80 to 90 percent of all the known kinds of animals.
The actual number of insect species, however, is believed to be
much larger and to run into the millions (628). 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 man.

2

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

Several general references to insects are available for consulta-
tion: Leonard (454), Brimley (101), Essig (232), Comstock
(152), Imms (391), Matheson (510), Brues, Melander and Car-
penter (114), Ross (624), and Borror and DeLong (90).

Class CRUSTACEA
Wood Lice

Crustaceans are aquatic organisms, mostly marine. They occur
throughout the world, and several species attack the wood of most
species of trees when they are placed in salt water. The surface
of heavily infested wood may be so completely honey-combed that
it resembles a sponge. When this damaged wood is 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 1 inch
of wood per year, Infested pilings often have an hour-glass 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 40 to 70 feet or more (9).

Limnoria ignorum (Rathke), commonly called the wood 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 salt water
only and spreads slowly, usually in infested driftwood.

Members of the genus Sphaeroma are beetle-like 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 dam-
age 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, however, even though the burrows are some-
what wider and penetrate to depths of 3 or 4 inches.

The genus Chelura contains the largest of the wood-boring
crustaceans. None of these initiate attacks on wood, but they do
invade and enlarge burrows made by shipworms. C. terebrans 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, worm-like organisms,
usually with 30 or more pairs of legs. They are usually 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 ring-like joints, each of which
bears two pairs of legs. Spirobolus marginatus Say, one of the
larger species, is dark brown, narrowly ringed with red, and
about 100 mm. long.

Millipedes sometimes become very abundant. At such times they
may invade camps, old buildings, and residences in large numbers.
Many species emit a fluid with a cyanide-like odor through open-
ings along the sides of the body. Removal of moisture and of
accumulations of vegetable matter from infested areas should be
helpful in their control. Williams and Hefner (768) discuss many
of the species occurring in eastern United States.

Class CHILOPODA

Centipedes

Centipedes are worm-like animals, with elongate, flattened seg-
mented 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 usu-
ally 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 man.

The largest centipedes belong to the family Scolopendridae,
some of which may be more than one foot long. The largest species
found in the United States, however, is only 125 mm. long. Mem-
bers of this family are also the most venomous of all centipedes.
The bite of the largest ones are not only quite painful but also are
occasionally fatal (90). The larger and more dangerous species
are southern in distribution; those occurring in the North are
usually too small to be harmful to man. Bailey (16) discussed the
centipedes of New York.

The removal of debris used as hiding places is helpful in centi-
pede control.

4

Class ARACHNIDA
Spiders, ticks, mites, scorpions, etc.

The class Arachnida is a large group of air-breathing arthro-
pods in which the body is usually comprised of two regions: (1)
the cephalothorax, and (2) the abdomen. They generally have six
pairs of appendages (the chelicera, pedialps, and four pairs of
legs) but are without antennae.

Order ARANEIDA
Spiders

Spiders are found almost everywhere, both indoors and out-
doors, and are so familiar as to need no description (153). Ob-
jects of fear and revulsion to many people, their presence in large
numbers in and around places of habitation or other areas fre-
quented by people, is often considered intolerable. Fortunately,
the majority of species pose no hazard to man, 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 man
and should be avoided (14).

The black widow spider, Latrodectus mactans (Fab.), is prob-
ably the most poisonous spider in the United States. 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 hour glass-like red mark on the underside of the 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.

Loxosceles reclusa Gertsch and Muliak, the brown recluse spider,
is also quite poisonous to man, 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 man. It has been
recorded from several Midwestern and Southern States—Okla-
homa, Arkansas, Kansas, Texas, Louisiana, Mississippi, Alabama,
and Tennessee. It may be found in almost any situation where
protection, food, and dryness are adequate, but it probably occurs
most commonly inside buildings. Its web is medium-sized and
irregular, with a maze of threads extending in all directions with-
out definite pattern or plan (354).

It is usually impractical or impossible to eliminate spiders com-
pletely. Populations can be reduced, however, by the removal of
favorite breeding places.

Order ACARINA
Mites and ticks

The order Acarina contains a large number of important pests
of plants and man and 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 Arachinids in having the mouth parts more or less distinctly
set off from the rest of the body on a false head and in never
having the body so divided that a distinct cephalothorax and
abdomen are clearly recognizable (18).

MITES.—Numerous species of mites feed on and weaken or
kill a wide variety of valuable plants, including trees. Many others
attack man and various other forms of animal life, often causing
extreme irritation, and sometimes illness and death. A consider-
able number are either parasitic or predaceous on various species
of destructive insects and harmful species of their own kind.
Many others feed on dead materials of all kinds on the forest floor.

The family Tetranychidae (480, 281) contains a large number
of species commonly known as spider mites, many of which are
important pests of trees and shrubs. These mites vary in color
from yellowish, greenish, orangish, and reddish to red, and all
are less than 1 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 dis-
colored, disfigured, or killed.

The spruce spider mite, Oligonychus ununguis (Jacobi), feeds
on a number of conifers, especially spruce, cedar, and some of the
pines and is widely distributed in the United States. Infested
trees may become brownish-gray and appear unhealthy, or they
may be completely defoliated. Outbreaks over large forested areas
have been recorded. Periods of drought appear to be most favor-
able for pepulation build-ups. Outbreaks have also occurred fol-
lowing widespread spraying with DDT (403). Young nursery
stock and recently planted trees are especially subject to serious
injury.

Oligonychus milleri (McG.) feeds on pines throughout the
South and north to Pennsylvania and the Lake States. Out-
breaks covering millions of acres have been recorded during
recent years from North Carolina to Florida, in northern Louisi-
ana and southern Arkansas, and in Mississippi and Texas. Young,
Spee sown, even-aged stands appear to be susceptible to severe
attack.

Other important species in the genus Oligonychus include: (1)
O. bicolor (Banks)—long recognized as a pest of oaks, especially
shade oaks and sometimes injures beech; (2) the southern red mite,
O. ilicis (McG.)—damages azalea and camellia; (8) O. aceris
(Shimer)—sometimes a serious pest of maples; (4) O. letch-
wortht Reeves—occasionally seriously injures hop hornbeam; (5)
O. newcomert McG.—often seriously infests shadbush; (6) O.
cunliffei P. & B.—feeds on longleaf pine in Florida; (7) O. bou-
dreauxt P. & B.—feeds on cypress in the South; and (8) O.
Bans P. & B.—feeds on oaks from Washington, D. C. to North

arolina.

6

The carmine spider mite, Tetranychus cinnabarinus (Bois.), is a
common and widely distributed species that feeds on a great
many species of plants, trees, and ornamentals. Several genera-
tions are produced during the summer months, often giving rise
to tremendous populations. Heavily infested plants may be en-
tirely defoliated, especially during hot, dry weather. T. ellipticus
Garm. is often abundant on honey locust in the Midwest. T.
homorus P. & B. feeds on hickory and ash in North Carolina;
T. magnoliae Boud. occurs on magnolia and yellow poplar in
Louisiana; the four-spotted spider mite, 7. canadensis (McG.),
feeds on elm, basswood, horse chestnut, osage orange, and poplar
throughout eastern United States and southern Canada; and the
Schoene spider mite, 7. schoenet McG., infests elm and black locust
throughout eastern United States.

The genus Hotetranychus also contains a number of common
and frequently important species. E. populi (Koch) and E. wel-
dont (Ewing) are found on poplars and willows; E. hicoriae
(McG.) occurs on pecan, hickory, horse chestnut, and various
oaks; and E.. matthyssei Reeves attacks elm in New York. Heavy
infestations may cause severe browning and cupping of the under-
sides of leaves. EF. querci Reeves has caused severe browning of
pin oaks in New York.

Platytetranychus multidigituli (Ewing) feeds on the leaves of
honey locust, causing them to turn yellow, and P. thujae (McG.)
attacks arborvitae, juniper, and cypress. 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 (414). Many produce open pouch-like or blister-like 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 bladder-gall mite, Vasates quadripedes (Shimer), is
a common species. It ranges in length from about 1/20 to 1/5 of a
mm. and feeds on the under-surface of silver maple leaves, caus-
ing the formation of pouch-like or bladder-like galls up to 3 mm.
in diameter (fig. 1). 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 pear leaf blister
mite, EZ. pyri (Pagen)—an introduced pest of pear and apple, has
also been recorded on mountain ash and shadbush. It is sometimes
abundant enough on mountain ash to cause noticeable injury. E.
parapopuli (K.)—stunts the growth of poplar by producing
woody galls around the buds. EH. caulis Keifer—causes large, de-
forming, hairy growths on the petioles of black walnut leaves. The

7

F-506694

FIGURE 1.—Galls of the maple

bladder-gall mite, Vasates quad-
ripedes, on leaf of maple.

pine bud mite, Phytoptus pini Nal.—causes the yellowing and
dropping of pine needles. Aceria phloceoptes (Nal.)—deforms
fruit spurs and produces woody galls on plum. Nalepella tsugi-
foliae Keifer—has been reported damaging hemlock in nurseries
in New York. Trisetaceus cupressi (K.)—attacks southern red
cedars, causing distortion and deformation of young trees.

Mites of the family Sarcoptidae are skin parasites of warm
blooded animals. The itch mite, Sarcoptes scabet (DeG.), is a well-
known species. It attacks man and causes severe itching as it bur-
rows into the skin.

The family Trombiculidae contains the notorious red bugs or
chiggers. The common chigger in eastern United States is the first
instar larva of the species, Eutrombicula alfreddugesi (Ouder-
mans). In its later stages, it feeds on other insects, snakes, birds,
lizards, and rodents. When it attaches itself to the skin of man
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 Sea-
board north to New Jersey.

Many other species of mites occur in association with bark
beetles,? and some are known to be parasitic. So far, very little
information is available on their effectiveness in natural con-
trol 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 com-
mon, 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.

* Unpublished manuscript by J. C. Moser and L. M. Roton. Mites asso-
ciated with southern pine bark beetles in Allen Parish, La.

8

The American dog tick, Dermacentor variabilis (Say), occurs
throughout eastern United States, but is most abundant in coastal
areas and in the Mississippi River Valley. The larvae and nymphs
feed largely on rodents; the adults feed on dogs, horses, hogs,
cattle, and many species of wild animals. Man also may be bitten,
but he is not a preferred host (477). 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 trans-
mits Rocky Mountain spotted fever and tularemia, both serious
diseases of man. It also transmits anaplamosis, a disease of cattle.

The black-legged tick, Ixodes scapularis Say, occurs along the
Atlantic Coast. The larvae and nymphs feed on rodents and rep-
tiles, 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 along game trails, paths, and roadways where the ticks
wait on vegetation for people or animals to come by. Adults are
most prevalent during fall and early winter. This species trans-
mits anaplamosis to cattle and piroplasmosis to dogs (726).

Order SCORPIONIDEA
Scorpions

Scorpions are fairly common in eastern United States, espe-
cially 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 14 to 714 inches in length. The abdomen is
divided into a large anterior portion of seven segments and a
long, narrow tail-like posterior portion of five segments. The
latter ends in a vesicle which bears a poisonous sting. When a
scorpion runs, it holds it large claw-like pedialps forward, and
the posterior end, bearing the sting, is usually curved upward.

Scorpions are capable of inflicting painful and sometimes 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 eastern United States is Centu-
roides vittatus (Say), the so-called striped scorpion. This is a
fairly small species, being only about 58 to 60 mm. long. Its sting
results in a sharp pain which usually lasts about 15 to 20 minutes.
Stings inflicted from late March to early May, however, may re-
main painful for several hours (15).

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

Order PSEUDOSCORPIONIDAE
False scorpions

False scorpions bear a striking resemblance to their larger rel-
atives, the true scorpions, but differ in having wider, shorter
abdomens and in not having terminal poison glands. They are
seldom more than 5 mm. long.

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, and in chicken houses
and beehives. Their food consists of mites, ants, and various other
insects. They have also been reported as biting man but without
inflicting appreciable injury. One species, Chelifer cancroides
(Linn.), is frequently found in association with man. Adults are
about 2.5 to 3 mm. long (860).

Order PHALANGIDA
Daddy-long-legs

Daddy-long-legs 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, usu-
ally appearing to totter about, and they occur commonly 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 are
harmful to man.

Phylum MOLLUSCA
Shipworms

Wood submerged in salt water is attacked by several species of
bivalve mollusks, commonly known as shipworms. Damage to
pilings is often severe, 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 be-
cause the damage they cause is similar to and often confused with
that caused by wood-boring insects.

The genera Teredo and Bankia contain the so-called shipworms.
Their bodies are long and soft. They are armed with small, chisel-
like 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 the shell, which is no longer needed for protection, 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.

10

Because a single piling may contain several thousand ship-
worms, it may be literally honeycombed (fig. 2). In such situa-
tions, tunnels may be no more than 14, of an inch in diameter and
only a few inches long. Under the most favorable conditions, how-
ever, the tunnels may reach a diameter of 1 inch and a length of
4 feet.

The genus Martesia also contains a number of destructive
species. They differ from the shipworms in being clam-like. 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 about 25 mm. in diameter and
60 mm. long. As they grow, they enlarge their cavities to accomo-
date 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 impreg-
nation of the wood with a suitable preservative. Coal tar 1
creosote is recommended for treating waterfront timbers of
Douglas-fir and southern pine. Copper naphthenate is usually
used for the protection of wood in boats. Coatings of paint and
metal are also effective as long as they remain intact.

F-519930
FIGURE 2.—Cross-section of a piece of timber riddled with tunnels made
by a molluscan shipworm of the genus Bankza.

11

FOREST INSECTS

Insects are by far the most numerous of all the forms of animal
life inhabiting the forest. They are remarkably well adapted to
their surroundings and occupy an incredibly wide variety of eco-
logical niches. They serve many functions in the economy of the
forest and are as essential a part of the complex association of
organisms comprising it as are the trees themselves (412). While
the majority of species are either beneficial or innocuous, many
species are exceedingly injurious.

Beneficial forest insects are represented by a great many
species, some of which are enormously abundant. Some feed on
forest debris and aid in its deterioration; others feed on organic
matter in the duff and soil and contribute to improvements in soil
fertility. Some contribute to stand improvement by attacking and
killing decadent, diseased, or overmature trees and making way
for younger growing stock. Many others are parasites or preda-
tors of destructive species of insects.

Harmful forest insects are those that are responsible for eco-
nomic loss. They include (1) species that damage or destroy the
flowers and seeds of trees and which are particularly important
pests in seed orchards and seed production areas; (2) species that
stunt, deform, or kill young trees by damaging or destroying the
terminals, laterals, or roots of reproduction and in plantations;
(3) species that cause loss of vitality, growth reduction, and often
the death of trees by eating the foliage; and (4) species that feed
under the bark or in the wood of living trees and girdle and kill
them or riddle them with tunnels are important pests in the forest.
These are not all: Numerous others that bore into and damage or
destroy green logs, storm-felled timber, green-sawn and seasoned
lumber, rustic construction, poles, posts, crossties, mine props,
and all manner of finished products from flooring to furniture are
highly destructive.

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 normally
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 in-
creases in numbers when favorable environmental conditions
prevail.

Eastern forests are also inhabited by many species of intro-
duced insects, a few of which are widely distributed and extremely
destructive. These include the gypsy moth, European pine shoot
moth, balsam woolly aphid, 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 composi-
tion, (2) in overmature rather than immature stands, and (3) in

12

ner

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plantations rather than natural stands. They may also develop
in stands weakened or decimated by hailstorms, flood, wind,
drought, tree diseases, fire, or defoliation or during logging opera-
tions. Any time a breakdown occurs 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,
outbreaks are likely to occur.

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 hundreds of
thousands of acres and last for many years before subsiding. From
1957 to 1966, a total of 154 different species were recorded in out-
break status in eastern forests.4

The following general discussions on forest insects are avail-
able for reference: Anderson (9); Doane, et al (196); Graham,
K. (302); Graham and Knight (309) ; Thatcher (710) ; Beal and
Massey (41); Beal, et al (40); Becker (50); Houser (381);
Kotinsky (437); MacAloney and Ewan (472); MacAloney and
Schmiege (473); and Shenefelt and Benjamin (644).

Forest Insect Caused Losses

Insects are among the most destructive agents affecting forest
and shade trees. During 1952 (considered an average year), they
killed an estimated 1 billion cubic feet of young, growing trees
and 5 billion board feet of older, more mature trees of sawtimber
size. In addition, they were responsible for a loss in growth of 1.8
billion cubic feet in the young, growing stock and 8.6 billion board
feet in surviving sawtimber trees (729). The.losses in sawtimber
alone were enough to build 114 million homes. Losses incurred
during epidemics are sometimes much heavier. Additional losses
of a considerable but unknown amount are caused by insects that
bore into the wood of living trees or dead and fallen timber.

Insects are not only responsible for the killing of valuable trees
and the loss of growth or recoverable volumes from surviving
damaged trees, for the weakening or destruction of wood products
of all kinds, but also for serious losses of other kinds, both
tangible and intangible. Management plans are often upset and
fire hazards increased; watersheds and wildlife habitats are im-
paired or destroyed; water in streams and lakes is polluted; the
incidence and severity of floods are increased; and the attractive-
ness of parks and other recreational areas is reduced or destroyed.
Heavy investments in tree improvement programs are also jeopar-
dized 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; and innumerable shade and ornamental trees around
homes, along roadsides, or along the streets of towns and cities
are killed or their attractiveness greatly reduced or destroyed.

*Forest Insect Conditions Annual Reports, 1957-1966 inclusive, USDA
Forest Service.

13

Forest Insect Surveys

It has long been recognized that successful control of forest
insect outbreaks depends not so much on the control method used
as upon a knowledge of the insect situation on immediate, sur-
rounding areas. When an outbreak is discovered, it needs to be
known whether it is increasing or decreasing. It is also impor-
tant to locate the center of greatest concentration and to deter-
mine the direction of spread (169). 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 sup-
pressed 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, non-outbreak
types of insects, to determine if, when, and where their numbers
or damage are reaching economic levels of concern. Where inten-
Sive management of the forest for timber or other objectives is
being practiced, it is also important to have up-to-date informa-
tion on all destructive pests to allow for their early control where
needed. Such knowledge as described above is sought or acquired
through surveys.

Forest insect surveys of one sort or another have been made in
the United States since the early days of this century (88), but
prior to the passage of the Forest Pest Control Act in 1947, they
were usually conducted on a more or less unorganized basis. Pas-
sage 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 Agri-
culture, either directly or in cooperation with other Departments
of the Federal Government, with states and other public agencies,
and with private organizations and individuals, to conduct surveys
to detect and appraise insect infestations before they develop to
outbreak proportions; to appraise their potential destructiveness ;
and to determine the needs for their control. Forest insect surveys
are also conducted in Canada. There, the aim is 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 (481).

Basically, forest insect surveys are of two kinds, detection and
evaluation, and they may be conducted separately or at the same
time. Detection surveys, as the term applies, are primarily for the
purpose of discovering threatening infestations. Evaluation sur-
veys are usually far more intensive and complex, being concerned
with such factors as the intensity and 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, 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 sur-
veys, the choice depending on many factors, such as the insect

14

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 (3, 339, 341) ; or by aerial photography (745, 2, 338,
340, 746, 142). Unfortunately, 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. 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 highly complex and require the services of highly
trained personnel. Ground surveys for evaluation purposes pre-
sent 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. In addition to 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: what likely will happen if no
action is taken to suppress the infestation, and what might be
expected to happen if suppressive measures are applied.

Much progress has been made in recent years in reducing the
time and cost of conducting surveys, particularly through the use
of sequential sampling techniques. These techniques provide for
flexibility in sample size in contrast to conventional techniques
which 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 dis-
tinguished by specified limits (743). Sampling plans have been
published for several important forest insect pests (158, 157, 393,
894, 542, 612, 692).

Forest Insect Control

The impact of naturally occurring adverse environmental fac-
tors 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 abundant at times to be
considered ‘“‘pests” are also subject to control by these factors,
though usually to a lesser degree and frequency. Depending on the
insect and the value of the forest resources affected, a given de-
gree of natural control may be economically effective.

When natural control factors fail to hold populations to eco-
nomically tolerable levels, it may be necessary to apply arti-
ficial controls. Depending on the insect and its biology and eco-
logical 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 repro-

15

duction, or to kill it; or they may be applied to the environment
to render it less favorable for the insect either through alteration
in food supply, in microclimatic factors, or in the abundance and
effectiveness of natural enemies.

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

Artificial control methods designed to suppress existing out-
breaks, usually have short-term effects only; those that are used
for the prevention of outbreaks may be long lasting. Control costs
vary considerably, depending on the insect and its habits, the
methods and materials used, and the size and accessibility of in-
fected stands. It may be limited to only a few dollars per acre
where infested stands are treated with aerial applications or
insecticides or it may cost several dollars per tree when treat-
ments are made to individual trees. Occasionally, control costs
may be retrieved through the salvage and sale of infested timber.
Some types of control may be used with complete safety; others,
in which hazardous materials are involved, must be used with
great care.

NATURAL CONTROL

Natural control results when naturally occurring adverse en-
vironmental 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 en-
tirely, but its impact may fluctuate greatly from place to place
and time to time (9, 302, 309).

PHYSICAL FACTORS.—Temperature is one of the most im-
portant 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 75° and 80° F. As temperatures depart
from this range, in either direction, activity gradually declines and
eventually ceases. When they climb to 120° or drop to around 0°
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 summer, 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.
Adults find relief from heat by moving into the interior of stands,
to the shaded portions of trunks, or to the interior of crowns. Dur-
ing 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 utilize areas in

16

which to hibernate either in the egg, larval, pupal, or adult stage
where the likelihood of lethal temperatures occurring are greatly
lessened. Because of physiological changes which occur in the dor-
mant state, many northern species are able to survive tempera-
tures well below 0°. Southern species are much less likely to
Survive exposure to such temperatures.

Outbreaks of leaf eating insects may be suppressed by late
spring frosts which 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 slow-down in the rate of larval development. Prolonged de-
velopmental 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 is related to photoperiod and the length of the season of
biologically effective temperatures. In the South, a species may
produce five or more generations per year; whereas in the North,
it may produce only one. 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 powder-post
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 intensi-
fied during or following periods of drought.

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
one species of tree. Some of the most serious bark beetle pests.
belong to this group. Many species of destructive wood borers also
confine their attacks to a single species or to closely related
species. In contrast, numerous other species feed on several to 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 manner of places; others deposit them on
host trees only. This often leads to heavy losses among newly-
hatched larvae, especially where acceptable food is not readily
available. Infestations of a single-host 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 which contain a low ratio
of preferred to non-preferred 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

17

often severe. Occasionally, as when a virus epizootic develops, the
results may be catastrophic. 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 col-
lected 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 unwise interference
by man; 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 pre-
daceous; many of these attack forest insects. The majority of
parasitic forms belong to only a few families in the orders Hy-
menoptera and Diptera. Important predators are found in several
families of the orders Coleoptera, Diptera, Neuroptera, Hemip-
tera, and Hymenoptera.

Parasitic and predaceous 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 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 (284, 535). Small mammals such as
shrews are often effective in controlling species that spend parts
of their lives on the ground.

The literature contains many references to the role of parasites
and predators in insect control. A good introduction to the subject
is found in publications by DeBach (187), Sweetman (701),
Clausen (144), Balch (25), Turnbull and Chant (722), Thompson
(714), and Burkner (118).

Many species of forest insects are also subject to a consider-
able degree of control by pathogenic micro-organisms. Some of
the latter, especially the viruses, frequently occur in the form of
epizootics and decimate or eliminate outbreaks over large areas.
Notable examples are (1) a polyhedrosis virus disease which has
played a leading role in the control of gypsy moth outbreaks in
the Northeast for many years, and (2) a disease of the same type
which caused the collapse of an outbreak of the European spruce
sawfly over a region of several thousand square miles in eastern
Canada and the Northeastern States during the late 1930’s.

Well over 1100 different kinds of micro-organisms, most of
which are pathogenic, have been found associated with insects.

18

This total includes 90 species and varieties of bacteria, 260 species
of viruses and rickettsia, 460 species of fungi, 255 species of proto-
zoa, and 100 species of nematodes (557). Generally speaking, they
gain entry into the insect either by being ingested with food,
through wounds or other damaged areas in the integument, or
through the integument or trachea. Most of them rely on wind,
rain, streams, healthy or contaminated insects, small mammals,
or birds for dispersal throughout an insect infestation. 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. Once inside the host
these viruses enter the cell nuclei, replicate, and become encapsu-
lated in many-sided, rodlike crystals, known as polyhedra. In-
vaded cells are soon destroyed, the polyhedra are then released in
the body cavity, and the host dies. Diseased larvae usually become
sluggish, 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 liquefied. Eventually
they completely disintegrate and dry up on the trees. Some of the
more important eastern forest insects, in addition to the gypsy
moth and European spruce sawfly, that are subject to polyhedrosis
virus diseases are eastern tent and forest tent caterpillars, fall
webworm, linden looper, white-marked tussock moth, winter
moth, European pine sawfly, red-headed pine sawfly, jack-pine
sawfly, Virginia pine sawfly, and Neodiprion excitans.

Forest insects are also affected by several other kinds of
viruses—cytoplasmic polyhedrosis, granulosis, polymorphic inclu-
sion, and non-inclusion. Generally speaking, these viruses appear
to be less effective and less specific in control than those of the
nuclear-polyhedrosis type. For further information on viruses the
reader is referred to Bergold (61), Bird (67), Hughes (389),
Martignoni and Langston (501), and Steinhaus (690).

Several species of bacteria produce so-called milky diseases in
the insect host, the best known being the milky disease of Japa-.
nese bettle larvae (212). 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 Hymen-
optera, Coleoptera, Diptera, and Orthoptera are susceptible to
diseases caused by various crystalliferous bacteria, the best
known of which are several varieties of Bacillus thuringrensis
(G3), 332).

Some species of fungi occasionally cause significant reductions
in insect populations. Aphids are particularly susceptible to at-
tack by species of Entomophora (=Empusa). The species, E.
megasperma, was credited with causing a high degree of control
of the forest tent caterpillar in Ontario from 1949 to 1952. An-
other species, Beawveria bassiana, reportedly killed over 90 per-
cent of the larvae of the smaller European elm bark beetle in a
number of infested elm trees in Connecticut (195).

Many species of pathogenic Protozoa are also associated with
forest insects. Generally speaking, they prolong the length of the
larval stage and reduce the fecundity of surviving adults.

tg

Numerous species of nematodes 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; and some cause injury but not death. Certain bark beetle-
infesting species 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 (508). Welch (753)
reviewed the literature on the subject.

APPLIED CONTROL

Applied control consists of the regulation by man of insect
activities, distribution, and abundance through the use of prac-
tices designed to enhance, supplement, or serve as substitutes for
natural control. The action taken may be either direct or indirect;
i.e., 1t may be directed either against the insect itself to suppress
currently destructive populations or toward the alteration of en-
vironmental conditions in such a way that destructive populations
cannot develop. Whatever the objective, the aim is to use the
cheapest, safest, and most economical methods available.

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 care-
fully designed cultural or management practices. Efforts directed
toward this end are commonly called silvicultural control.

Possibilities of silvicultural control occur during the establish-
ment and throughout the lives of stands, 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 thin-
nings 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-
Ae trees may be removed in sanitation-salvage cuttings (413,
631).

For further discussions of silvicultural control, see Keen (412),
Graham and Knight (309), and Prebble (605).

PHYSICAL AND MECHANICAL METHODS OF CONTROL

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

20

Trap trees and trap logs are sometimes effective in the control
of bark beetles (552). 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. Bag-
worms on ornamental trees may be controlled by handpicking.
Subterranean termites can be kept out of structures by capping
foundations with a 4-inch 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.

The logging and milling of infested timber and destruction of
the slabs containing the broods is often effective in preventing
bark beetle outbreaks, but this 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 disposed of by the storage of infested logs in water ;
by peeling infested logs and burning the bark; by sawing infested
trees into short lengths and burning them; by peeling the bark of
infested trees and exposing the brood to desiccation and to natural
enemies such as ants, birds, and rodents; or by felling infested
trees in a north-south direction in the open where the bark is
exposed to heat from direct exposure in the sun.

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

Ips bark beetle populations which 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 cut-
tings throughout the summer months. In situations where it is
necessary to cut on a small seale or discontinuously during the
summer, control may be obtained by piling and burning the slash
before the beetles complete their development in it. The gathering
and burning of severed branches late in the fall or in the 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 powder-post beetle damage can
be reduced by the long storage of susceptible sapwood in water or
by steaming it at high temperatures. Bark-beetle damage to bark-
covered poles and slabs used in rustic construction can be reduced
or prevented 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 Lake States
can be reduced by removing the lower limbs. This forces the over-
wintering stages of the insect into buds on the upper branches
above the snow line where they are killed by lethal low tempera-
tures during the winter.

21

REGULATORY CONTROL

Regulatory control is aimed at the prevention of entry and es-
tablishment of foreign plant and animal pests, or at the suppres-
sion, 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 importa-
tion and interstate movement of articles that might spread insect
pests. It was reinforced in 1912 by passage of the Plant Quaran-
tine Act, which authorized the Secretary of Agriculture to enforce
necessary regulations to protect the agricultural economy of the
United States by preventing the introduction of insects and plant
diseases, and by passage of the McNary-McSweeney Act in 1928,
which established a federal policy regarding the use of legislative
means for combatting forest insects and diseases. Then, 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.

Prior to the passage and enforcement of quarantine laws, many
species of forest, shade tree, and ornamental insects, several of
which are highly destructive, gained entry into 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 Eastern United States.

Order and Species Country of origin Principal food plant

Orthoptera
Gryllotalpa gryllotalpa (L.) Europe Nursery stock
Hemiptera
Stephanitis rhododendri Harv. Europe Rhododendron
Isoptera
Coptotermes formosanus Shiraki Far East Wood products
Homoptera
Adelges abietis L. Europe Spruce
piceae (Ratz.) Europe True firs
strobilobius Kalt. Europe Larch
Asterolecanium variolosum (Ratz.) Europe Oak
Cryptococcus fagi (Baer) Europe Beech
Gossyparia spuria (Mod.) Europe Elm
Matsucoccus resinosae (B. & G.) Red Pine
Pealius azaleae (B. & M.) Asia Azalea

Phenacoccus acericola King Europe Sugar maple

Pineus strobi (Htg.) Europe Pine

Pseudococcus comstocki (Kuwani) Far East Deciduous trees

Quadraspidiotus perniciosus (Comst.) China Deciduous trees
Coleoptera

Amphimallon majalis (Raz.) Europe Various plants

Anobium punctatum (DeG.) Europe Wood products

Anomala orientalis Waterhouse
Chlorophorus annularis (F.)

Philippines
Japan, India

Nursery stock
Stored bamboo

Chrysomela interrupta F. Europe Willow, poplar
crotchi Brown Europe Willow, poplar
Cryptorhynchus lapathi (1L.) Europe Poplar
Crytepistomus castaneus (Roelofs) Japan Oak
Heterobostrychus aequalis (Waterh.) Far East Oak & mahog-
any boards
Hylotrupes bajulus (L.) Europe Seasoned conif-

22

erous wood

Order and Species

Maladera castanea (Arrow)
Minthea rugicollis (Walk.)
Nacerdes melanura (L.)
Phyllobius oblongus (L.)
Plagiodera versicolora (Laich.)
Polydrusus impressifrons Gyll.
Popillia japonica Newm.

Pseudocneorhinus bifasciatus (Roelofs)
Pyrrhalta luteola (Mueller)

Scolytus multistriatus Marsh.
Thylacites incanus (L.)

Xestobium rufovillosum

Zengophora scutellaris Suffr.

Lepidoptera

Aegeria apiformis (Clerck)
Anthophila pariana Clerck
Archips rosanus (L.)

Chrysoclista linneella (Clerck)
Cnidocampa flavescens (Wlk.)
Coleophora laricella (Hbn.)
fuscedinella (Zell.)
limosipennella (Dup.)
Dichomeris marginellus (D & S.)
Dioryctria abietella (D. & 8S.)
Epinotia nanana (Treit.)
Gracillaria syringella (F.)
Homadula anisocentra Merrick

Nepticula sericopeza Zell.
Nygmia phaeorrhoea (Don.)
Orgyia antiqua (L.)
Phalonia rutilana (Hbn.)
Porthetria dispar (L.)
Rhyacionia buoliana (Schiff. )
Samia cynthia (Drury)

Spilonota ocellana (D. & S.)
Stilpnotia salicis (L.)

Zeiraphera ratzeburgiana (Ratz.)
Zeuzera pyrina (L.)

Diptera

Monarthropalpus bux. (Lab.)
Oligotrophus betulae ( Winn.)
Rhabdophaga salicis (Schr.)

Hymenoptera

Acantholyda erythrocephala (L.)
Caliroa cerasi (L.)

Diprion frutetorum (F.)
hercyniae (Htg.)
similis (Htg.)
Fenusa dohrnii (Tisch.)
pusilla (Lep.)
ulmi (Sund.)
Hemichroa crocea (Fourcroy)
Heterarthrus nemoratus (Fallen)
Neodiprion sertifer (Geoff. )
Pristiphora erichsonu (Htg.)
geniculata (Htg.)
Sirex juvencus (L.)
Solenopsis saevissima richteri Forel
Trichiocampus viminalis (Fall.)

Country of origin

Orient
Antilles (7?)
Europe
Europe
Europe, Japan
Europe

Japan

Japan

Europe
Europe
Europe
Europe
Europe

Europe
Europe
Europe

Europe

Asia

Europe, Japan
9

Europe
Europe
Europe
Europe
Europe
9

Europe
Europe
Europe
Europe
Europe
Europe
Asia

Europe
Europe
Europe
Europe

Europe
Europe
Europe

Europe, Japan
Europe

Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Eurasia
Europe
Europe
South America
Europe

Principal food plant

Deciduous trees

Oak flooring

Wooden wharves

Deciduous trees

Willow

Poplar

Wide variety of
plants

Ornamentals

Elm

Elm

Pine

Wood products

Poplar

Poplar

Apple, thorn

Deciduous trees,
privet

Linden

Norway maple

Larch

Birch

Elm

Juniper

Pines

Spruce

Lilac

Mimosa,
honeylocust

Norway maple

Apple, pear

Deciduous trees

Juniper

Oak, etc.

Pines

Ailanthus,
cherry

Apple, oak

Poplar, willow

Spruce

Elm, maple

Boxwood
Birch
Willow

Pine

Cherry,
shadbush

Pine

Spruce

Pine

Alder

Birch

Elm

Alder

Birch

Pine

Larch

Mountain ash

Pine, fir, spruce

General feeder

Poplar

23

BIOLOGICAL CONTROL

Biological control is possible because of the existence in nature
of a continuing interplay between the abundance of insects and
their natural enemies, and because many of the latter are subject
to manipulation one way or another. Traditionally, it has con-
sisted of the use of parasites, predators, and disease pathogens to
hold populations in check, but within recent years the concept has
been broadened to include various other biotic methods for reduc-
ing and/or maintaining insect populations at tolerable levels. So
far, almost all biological control efforts against forest insects
have been confined to the use of parasites, predators, and
pathogens.

Generally speaking, biological control efforts against forest in-
sects 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; (8) the augmentation of
established parasite and predator populations with fie!d-collected
or laboratory-reared individuals; and (4) the use of microbial
sprays to control outbreaks.

The majority of importations of parasites and predators were
made prior to World War II, with emphasis on enemies of the
gypsy moth, brown-tail moth, satin moth (387, 124), European
pine shoot moth, European pine sawfly, European spruce sawfly,
balsam woolly aphid, and larch casebearer. Since World War II,
importations have been limited mostly to enemies of the balsam
woolly aphid and the smaller European elm bark beetle. Alto-
gether, a total of 156 species of parasites and predators had been
imported and liberated against 59 species of introduced and
native pests by 1960. Of these, a total of 44 were successfully
established (205). Since the early 1930’s, the Canadian govern-
ment has also imported large numbers of many species of para-
sites and predators against a number of pests, many of which also
occur in the United States (484). Some of these 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 popu-
lations and is almost unnoticeable in light infestations. Limited
field trials have indicated that direct spraying of the virus in
water formulations has considerable promise for direct control.

Applications of polyhedrosis virus sprays have been used to
suppress populations of the European pine sawfly (66), the Vir-
ginia pine sawfly (485), and the Swaine jack pine sawfly (654).
Virus epizootics have been initiated in populations of the forest
tent caterpillar by disseminating the virus during one generation
(685).

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.

Progress has been made in the control of a number of lepidop-
terous defoliators with Bacillus thuringiensis. Aerial applications

24

re

caused some mortality of the gypsy moth but the results were
inconclusive. Higher mortalities of two species of loopers (EHran-
mis tiliaria and Phigalia titea), which also were present in the
sprayed areas, were recorded.

CHEMICAL CONTROL

The use of chemicals to suppress forest insect populations is a
method of last resort. It is the policy 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. Depending
on the situation, chemicals may be applied to a single tree or to
forested areas covering thousands of acres or square miles. 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 (601). Aerial applications are made by
fixed-wing aircraft (fig. 3) or helicopters to large areas (29, 214,
57, and 392). Individual trees, small groups of trees, and seed
orchard trees (fig. 4) may be treated by ground equipment such
as mist blowers (602) or by knapsack sprayers. Large individual
trees may be treated by mist blowers and by hydraulic sprayers.
Logs are treated by power sprayers. Fogging machines are
sometimes used around resorts and campgrounds. Chemicals
may be introduced into the tissues of living trees for bark beetle
control (171, 52, and 762). Systemic insecticides may be applied
by trunk implantation or injection, by banding or spraying of the
circumference of the trunk, by spraying the foliage, by treating
the soil around the base of trees, or by dipping cuttings prior to
plantine= (570-587-199, 515, 36, and 447).

The effectiveness of chemicals in suppressing many forest insect
populations has been amply demonstrated during the past 25 years

er 3 Rah

F-482299
FicgurRE 3.—Airplane spraying for spruce budworm control

25

F-517833
FIGURE 4.—Mist blower applying insecticide to pine trees in a seed orchard.

—a period during which a large number of new insecticides and
methods of application have come into extensive use, and during
which outbreaks covering hundreds of thousands of acres of forest
have been suppressed. As effective as insecticidal control has
been, however, it has not proved to be an unmixed blessing. It has
given rise to many complex problems associated with known or
suspected adverse side effects caused by some of the more com-
monly used chemicals. This stems from the fact that these chemi-
cals, like most other chemicals used as insecticides, are non-specific
and can be expected to be harmful to at least some other exposed
animal species. The problem is compounded in that some chemical
insecticides are very persistent, that all of the material applied in
a given environment may not remain in that environment and
that free-ranging animals cannot be excluded from sprayed areas.
Public awareness of known and possible hazards associated with
the use of these chemicals in insect control projects, therefore,
has given rise to a considerable degree of concern over their con-
tinued widespread use.

Efforts are unceasing to discover and develop new and safer
insecticides and to determine the danger points of insecticidal
accumulations in the tissues of various forms of wildlife, as a
basis for preventing undesirable damage to the biota. Application
techniques and equipment are being refined in order to provide
better control of the placement of insecticides in the environment
and to further lessen the dangers of undesirable side effects.
Studies are being made to improve sampling and biological evalu-
ation techniques as a basis for improving the timing of application
and to insure that insecticides are applied only when and where
they are needed.

The rapid changes occurring in the development and use of
insecticides in forest insect control make it inadvisable to include

26

control recommendations in this publication. If you are going to
use pesticides, remember: Because some States have restrictions
on the use of certain pesticides, check your State and local regula-
tions. Also, because registrations of pesticides are under constant
review by the U. S. Department of Agriculture, consult your
county agricultural agent or State Extension specialist. For fur-
ther information on chemical control, in addition to U. S. Depart-
ment of Agriculture Handbook No. 331, many publications are
available. The following are of particular interest: Billings (65),
Brown (109), Brown (110), Gorham (301), Hickey (350), Hoff-
man, et al. (363), Hoffman and Merkel (362), Hoffman and
Linduska (361), National Academy of Science (555, 556), Whit-
ten (761), and Rudd and Genelly (625).

OTHER APPROACHES TO CONTROL

Many conventional methods of suppressing forest insect popu-
lations are very effective, but they are all found wanting in many
important respects. Insecticides are often entirely effective, but
they usually cannot be applied to large areas without endangering
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 out-
break before unacceptable damage has been incurred. They are
also incapable of preventing outbreaks, although they may reduce
their frequency of occurrence and their magnitude, intensity, and
duration. Because of these deficiencies, as well as those of other
current methods of control, intensive research is underway to
develop new or improved methods which may be used as comple-
ments, supplements, or substitutes for them.

Sexual sterilization is receiving much current attention be-
cause of its potential use as a new method of control. The effec-
tiveness of releasing males sterilized by gamma radiation was-
established in the mid-1950’s when they were used to eradicate the
screw-worm from the West Indian island of Curacao and from the
southeastern portions of its range in the United States. Knipling
discussed the principal of sexual sterilization in several papers
(425, 428, 426, 429, 427). Several authors reported the research
that led to the method and its use against the screw-worm (456,
457, 129, 180, 131). Some progress has also been reported in the
sterilization cf a few important forest insects by gamma radia-
tion (300, 401). Further research must be conducted before the
possibilities of using the sterile male release technique for forest
insect control can be determined.

Chemically produced sterility also offers promise in control.
The appropriate chemicals are called chemosterilants because of
their capacity to deprive insects of the ability to reproduce (89).
Most of the chemosterilants that offer promise for practical con-
trol are mutagenic and can be used only in ways that will avoid
all contact between them and non-target animals (557). Chemo-
sterilants may be administered with the insect’s food or by apply-
ing them to surfaces to which the insects are attracted. Those
mixed with food appear to produce maximum effectiveness and
minimum hazard. So far, none are available for use in forest
insect control.

27

Insect attractants are being given increased research atten-
tion to improve or reduce the cost of surveys and to open up
hew opportunities in integrated control of forest insects. Some of
these materials are powerful enough to lure insects over consider-
able 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.

tion (665, 666, 667).

INTEGRATED CONTROL

Integrated control is defined as a pest Management system that
in the context of the associated environment and the population
dynamics of the pest species, utilizes all suitable techniques and
methods in as compatible manner as possible and maintains the
pest populations at levels below that causing economic injury.
In forest insect control, this would seem to encompass many of
the methods discussed in preceding sections on applied control.

28

Most of the important pest species in eastern forests lend them-
selves to some form and degree of integrated control. The long
life of the forest crop and the fact that many species of commer-
cially valuable trees can withstand some degree of infestation
without serious damage provides the opportunity in the majority
of cases to utilize different methods of control.

Since the early days of this century, considerable emphasis has
been placed on the importation and augmentation of natural ene-
mies of many species of introduced pests. In many cases, the rate
of dispersal of these imported enemies has been speeded up by
the liberation of colonies in many parts of infested areas. The
build-up in numbers after successful establishment of several
species has also been speeded up by the liberation of large num-
bers of field-collected or laboratory-reared individuals.

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 ap-
pearance of a virus disease in heavily infested stands usually por-
tends 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, such as the appli-
cation of insecticide. Observation of abnormally high percentages
of natural control provided by insect parasites and predators may
lead to a similar decision. In other situations where insecticidal
control is deemed necessary, it is often possible to limit spraying
to designated portions of infested areas because of the abundance
of natural enemies in other portions (423).

Knowledge of the population dynamics of a pest species pro-
vides opportunities for various other means of preventing or
reducing losses caused by it, thereby obviating the need for di-
rect, suppressive measures to control outbreaks. Depending on
the situation, outbreaks of certain leaf-feeding insects may be
prevented or minimized by modifying the composition and den-
sity of susceptible stands. It is often possible to prevent or sup-
press outbreaks of some bark beetles by the removal of infested
slash or the salvage of infested trees before the emergence of beetle
broods from the bark. The utilization of mature trees growing on
poor sites before they begin to deteriorate and become attractive
to borers, such as the bronze birch borer, is a recommended con-
trol practice. The avoidance of unfavorable sites in planting pro-
grams is recommended for preventing damage by certain species
(318). Adherence to good construction methods in the building
of structures obviates the need for chemical control of termites
later on (630). The destruction of elm material suitable for breed-
ing by the smaller European elm bark beetle reduces the need for
the use of insecticides in Dutch elm disease control (763).

Some of the new approaches to biological control, many of
which are yet to be perfected, may be expected to provide addi-
tional opportunities for the integrated approach to control in
the next few years. For a discussion of the integrated control
concept, see Stern et al. (691). Comprehensive treatments of
insect-pest control management and control has also been pub-
lished (557, 416).

29

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

The following keys are designed for the use of those unfamiliar
with the orders, families, and genera of insects. The insects 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 to other
groups or subdivisions.

PRIMARY DIVISIONS OF KEY

Te Insects injurious to seeds, seedlings, young plantations, and
small reproduction.
A. To seeds, cones, and fruits.
B. To seedlings and small reproduction.
II. Insects injurious to large reproduction, forest trees, and
shade trees.
A. Acarina (red spiders).
B. Defoliators, leaf miners, etc.
C. Twig and tip damage, etc.
D. Borers in wood and bark.
E. Galls, swellings, etc.
F. Sucking Insects.
III. Insects injurious to forest products.
A. Defects in green timber.
B. Insects in round logs.
C. Insects in lumber.
D. Insects in material in ground.
E. Defects in wood in salt and brackish water—marine
borers.

DIVISION I

INSECTS INJURIOUS TO SEEDS, SEEDLINGS, YOUNG PLANTATIONS,
AND SMALL REPRODUCTION

This group includes the insects that attack the fruit and seeds
of forest trees, also young seedlings in the nursery, plantation, or
forest. Insects attacking seedlings more than 4 or 5 years old and
somewhat shrubby and woody are discussed under Division II.

A. Insects Attacking Seeds, Cones, and Fruits

1. Larvae without well-developed head capsule; maggot-like:
With a sclerotized structure like a breastbone near an-
terior end; in seeds of fir, cypress, 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 walnut husks
Diptera, Tephritidae
Larvae with distinct head capsule . 2

30

2. Larvae active; body extended; abdominal prolegs present
Larvae less active; no abdominal prolegs; body curved ....
3. In the shucks of pecans, hickories, and walnuts and in
ACOLMS +e Lepidoptera, Blastobasidae, Olethreutidae
TRC ONCS Bes oars 8 SON) a lat Tt Lepidoptera, Phycitidae
In fruits of wild cranberries and blueberries...
Lepidoptera, Phycitidae
4. In acorns, walnuts, chestnuts, hickory nuts, and filberts....
imnvlesuminouseseeds: 2. 228 Coleoptera, Bruchidae
COMES) ee ek ere aA) ee cole lien 1 wae en ree
5. First abdominal spiracle vestigal; body spindle-shaped
Hymenoptera, Cynipidae
First abdominal spiracle normal; body curved like a
closed ingen tee! Coleoptera, Curculionidae

6. Cones abnormal and dropping prematurely
Coleoptera, Scolytidae

Cones maturing; damage to seeds alone

Warvaevlegless: ....5.2..c hadi... Hymenoptera, Chalcididae
Larvae with true legs ............000... Coleoptera, Anobiidae

B. Insects Attacking Seedlings and Small Reproduction

a

Plants cut off near ground line or stems lacerated and
GINTIVClCO shee orn Fo, meet Named ree ee eke
Plants wilting or fading, easily plucked from ground be-
cause of severed stem or roots below ground... |
Gall-like swellings Onethe Stems. Oe es Oe eae
Bark gnawed in patches along the stem of conifers... |
Foliage off-color, yellowish or rusty, and often covered
with very fine cobweb-like threads or matting
Tetranychididae
Roots showing small cottony or globular objects at time of
CRANSMIAMCINO 210 Fee cae Homoptera, Aphididae
Leaves or cotyledons cut off and carried away; Southern
SENECA aRe oT Hymenoptera, Formicidae
Other types of damage. See Division II.

8. Small, tender plants:
Small, maggotlike larvae causing injury
Diptera, Muscidae
Large, smooth, dark-colored caterpillars present
Lepidoptera, Phalaenidae
Larger plants, hardwoods, with woody part tunneled:
Tunnels longitudinal ................. Coleoptera, Cerambycidae
Tunnels across the grain, wood stained
Coleoptera, Scolytidae
Tunnels irregular, chiefly underground .................. lsoptera
9. Curved, grublike larvae in the soil
Coleoptera, Scarabaeidae
Elongate, cylindrical, hard-shelled larvae
Coleoptera, Elateridae
Small, molelike tunnels usually near the surface of the
SOU ee .. Orthoptera, Gryllotalpinae

10
Jit

31

10. Bark swollen and gnarled at or just below ground line
Lepidoptera, Phycitidae
pouthern: States he... we Homoptera, Fulgoridae
Northern States .......... ncaa Hymenoptera, Formicidae
11. Northern, Southern, and Lake States

Coleoptera, Curculionidae

More southern and western States and Lake States
Orthoptera, Acrididae

DIVISION II
INSECTS INJURIOUS TO LARGE LIVING TREES AND TO
SMALL TREES MORE THAN 4 OR 5 FEET TALL

This group includes insects and related organisms which com-
monly attack living forest and shade trees other than 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 ITI.

Injury consisting of discolored, yellowish, rusty, or mottled

foliage covered with fine cobweblike threads or matting
Tetranychidae A

Injury consisting of defoliation, leaf rolling, leaf tying, leaf

and petiole mining, or bast or epidermis miners on green-
barked ‘Stems. oes a eo es B

Injury occurring on new growth, twigs, branches, or small
trees, consisting of mining, pruning, withering, or flagging OC

Injury caused by larvae or beetles boring in the bark, under

the bark, orci the wood" 2.9 fee ee eee D
Injury consisting of a gall or swelling on stem, branch, or

11-5) aaa aemnnrTOne Ss muUnnUTNS Son oh ERD Sinemmact GV NA Mer CCT NAT iy ec E
Injury caused by sucking insects feeding on leaves, twigs, or

bark surfaces, usually the softer tissues of the plant ............. F

B. Defoliation and Other Injury

1. injury esrused-by beetles’ sc.) nn tk hoe Ce eee 2
Injury*caused-by larvae. 8 ee Gea ee eee 4
Injury caused by other forms or insect not present ....... wrID

2. Adults and larvae associated on the leaves
Coleoptera, Chrysomelidae
Adults’ only’ present... ee ee eee 3
3. Beetles usually found feeding at night
Coleoptera, Scarabaeidae
Small, bright-colored, jumping beetles, Flea beetles,
Coleoptera, Chrysomelidae
Dull black, purplish, or gray, soft-bodied beetles, Blister
Weetles i set OEE oy A ee Coleoptera, Meloidae
Small snout beetles _..... Coleoptera, Curculionidae

32

4, Prolegs, usually two or five pairs os Lepidoptera
Prolegs, usually six or more pairs—or none
Hymenoptera, Tenthredinidae
Prolegs inconspicuous; leaf- or bast-mining forms 5
5. Circular holes cut in the leaves
Hymenoptera, Megachilidae
Leaves rolled into a small, compact bundle
Coleoptera, Curculionidae
Grasshoppers associated with injury
Orthoptera, Acrididae
Walkingsticks associated with injury
Orthoptera, Phasmatidae

C. Twig Pruning and Other Injury

6. Injured portion hollowed or mined; injury caused by
larvae or bark beetles, which are usually present ff
Injury caused by external feeding or ovipositing, which
removes a portion of the bark or causes a definite
mechanical injury or a resin-infiltrated scar ........ 13
Cottony-masses on tips of twigs concealing the insects ;
conifers: .).= _Homoptera, Coccidae, Phylloxeridae
7. Injury on two or more whorls of the terminal of conifers;
inactive, curved larvae under bark or in pupal cells in
WOOC ew ers | cen sree Coleoptera, Curculionidae
Twigs or brances of hardwoods or conifers containing
bark beetles or powder-post beetles or a cylindrical
shotlike hole, usually darkly stained, directly entering
injured portion
Coleoptera, BUR Bostrichidae

In ysOUuneRWISG 4 eee) es ae 8

8. Twigs not mined below fading portion .__. 9
Twigs mined far below fading portion, ‘tunnel often ex-
tending to the ground ..... Coleoptera, Cerambycidae

rea ONCE Seater ee ea ene ee LO

UAC OOGSS mite (tote) ss onal sooth Srey ial
10. Larvae with prolegs; often pitch masses at point of
INJURY. = Lepidoptera, Olethreutidae
Larvae without conspicuous pr olegs; usually a spine on
lastseement! 2125 26.23. Hymenoptera, Tenthredinidae

11. Larvae with well-developed prolegs; usually colored;
usually in more tender parts of twigs
Lepidoptera, Olethreutidae, Aegeridae, Cossidae
Larvae with under- peve oped prolegs in pees portions
OMFtWIGS eS LN ence geome ys
12. Elongate, flat larvae; ‘mines filled
Coleoptera, Buprestidae
Cylindrical larvae; mines open
Coleoptera, Cerambycidae

> There seems to be no simple and practical method of separating the leaf
miners 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 Curcu-
lionidae, Chrysomelidae, or Buprestidae of the Coleoptera; or they may
be Diptera of the families Agromyzidae or Cecidomyiidae.

33

13. Obvious scar and pitching of wood at base of injury or

14.

15.

16.

U7.

18.

Ig

34

along twigs; conifers
Scale bodies present on twig
Homoptera, Margarodidae

Scale bodies absent .................. Coleoptera, Cerambycidae
Numerous phloem scars on twigs; spittle masses may
be’ present #02". 0) anes ae Homoptera, Cercopidae

(Hail injury is similar except that the scars are al-
ways on top side of branch)
Twigs slit with a lacerated wound at base of injury or at
point of breaking
Homoptera, Cicadidae, Membracidae
Orthoptera, Gryllidae

D. Borers in Wood and Bark

Borers in the phloem and outer corky bark of living
trees rarely scarring the: wood 444 6 ee
Borers in callous tissue around wounds

On various hardwoods ........ Coleoptera, Curculionidae
On maplesa:treer oti sees Lepidoptera, Aegeriidae
Oni coniters7: es 25 Lepidoptera, Aegeridae, Phycitidae

Borers in the dead wood beneath fire scars, turpentined
faces, blazes, cavities, and similar wounds ....................
Borers under the bark or in the wood (other than be-
neath scars or catfaces) of living trees 0...
Bark beetles associated with their larvae under the bark
Coleoptera, Scolytidae
Root borers or mining at base of tree
White, unpigmented larvae
Coleoptera, Cerambycidae
Highly pigmented larvae
Hymenoptera, Tenthredinidae
Serpentine mines just under the epidermis of chestnut
and oak _.. Lepidoptera, Nepticulidae
Ambrosia beetles; 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 conifers | Coleoptera, Buprestidae
Larvae with heavy, chitinous armature on last segment;
chestnut, oak, maple ... Coleoptera
Larval mines extended under the bark and also deep into
the wood in later stages
Larval mines entirely under the bark or only in wood of
current annual ring
Pitch exuding from larval mines; larvae with prolegs
present; conifers Lepidoptera,
No pitch, but often water and frass exuding
Head of larvae globular, protuberant
Head of larvae somewhat flattened and embedded in
prothorax Coleoptera, Cerambycidae

15

16
ey

22

18
21

i
20

20.

21.

22.

23.

24,

25.

26.

ale

Prolegs absent; last segment often heavily armed
Coleoptera, Tenebrionidae
Prolegs absent; larvae curved, grublike; in willow, pop-
larvandwpalmettoy. 2.2... Coleoptera, Curculionidae

Prolegs present; last segment not heavily armed
Lepidoptera, Hepialidae, Aegeriidae

Larvae depressed, flat-headed or pestle-shaped

Coleoptera, Buprestidae
Larvae curved, grublike ............ Coleoptera, Curculionidae
Larvae slender; thoracic segments not noticeably en-

larged; cause pitch flecks in wood, birch, ete.
Diptera, Agromyzidae
Larvae with prolegs; poplar, willow, alder, ash, persim-

MOM Aare Neer os Lepidoptera, Hepialidae, Aegeriidae
Larvae without prolegs
Inghardwoodsies<5 2405.2. Coleoptera, Cerambycidae

In conifers; associated with pitch mass
Coleoptera, Curculionidae, Scolytidae

EB. Galls §

Galls of more or less open, exposed, simple structure, or,
when enclosed, the insects maintain permanent open-
ings, or the gall is dehiscent to permit the escape of

rarely, several insects to a cavity; occasionally a per-
manent opening is maintained by the feeding larva.
Mites present having 2 pairs of legs; galls of various
shapes but always provided with an opening to the
exterior and lined on the inside with hairy or fuzzy
POMC INSGe ee ee he Se EN. ci Brag ee A cnn ee hee Acarina
Galls otherwise ......... Satie Bir ane Siow ath de leanne es 7 |
Insects=not.fitted. for JUMPING ...8..60.8 ban
Insects with hind legs developed for jumping
Homoptera, Psyllidae
Leaf galls on hardwoods, chiefly elm, poplar, hickory,
ash, sumac, and witch-hazel
Homoptera, Aphididae
Conelike galls on tips of spruce twigs
Homoptera, Chermidae
Pitlike galls on twigs of hard pines
Homoptera, Margarodidae
Pitlike galls on twigs of white oak
Homoptera, Asterolecanidae
Galls inhabited by larvae with a well-developed head
capsule ..... NaC e ee hae eee ae eee a ae ne
Larvae without well-developed head capsule, maggot-
like; white to yellowish or reddish in color:

24

27

25
26

28

Salt seems impossible to devise a key that will separate all the varied
types of galls into family or order groups. On the other hand, 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.

30

28.
29.

30.

ol.

32.

33.

36

Larvae with a distinct structure like a breastbone near

Anterior End eee Diptera, Cecidomyiidae
Larvae without breastbone; mouth parts well de-
veloped? 11.2. 2 See eee cs mees Diptera, Berouiyzide

Larvae legless or with only minute legs 0.0...
Legs well developed, also prolegs present 0...
Woody galls containing plain evidences of mining ac-
tivity of the larvae; larvae with a well-developed head
capsule and mandibles
Coleoptera, Buprestidae, Cerambycidae, Curculionidae
Larval mines not obvious; white larvae, curved or grub-
like in form, legless, and with distinct head capsule,
each contained in a specialized cell
Hymenoptera, Cynipidae, Chalcididae
On, willow. 26-2 Hymenoptera, Tenthredinidae
On locust, poplar, maple... Lepidoptera, Olethreutidae

F. Sucking Insects

Injury or insects present on leaves .....
Injury primarily confined to twigs ... i
Injury primarily confined to branches and main stem __
Leaves off-color, yellowish or spotted from feeding punc-
tures of active, jumping insects:
Homoptera, Cicadellidae
Hemiptera, Tingidae
Leaves bearing galls or abnormal spots:
On Nackberty- 2) Sane Homoptera, Phyllidae
On elm, poplar, willow, witch-hazel, hickory, oak,
chestnut, etc.
Homoptera, Aphididae, Phylloxeridae
On: conifers2542 54 eae Homoptera, Phylloxeridae
Exposed insects on the leaves:
Sealelike, 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 4-winged flies
Homoptera, Aleyrodidae
Soft-bodied insects with long conspicuous antennae
Homoptera, Aphididae
Insects surrounded by a conspicuous frothy mass of
spittle; ends of branches and trees slowly dying in
severe infestations; pines ......... Homoptera, Cercopidae
Injury consisting of ragged slits in the twigs, often
breaking at incision; the tips of the branches hanging
with withered leaves; hardwoods
Homoptera, Cicadidae
Homoptera, Membracidae
Injury consisting of gall-like or gouty swellings on limbs

ANG CWilLeiSrOl tlie ee. ee Homoptera, Chermidae
Tips of hard pines flagged (needles yellowing’) ; scales
embedded in pits in bark Homoptera, Coccidae

29
30

32
33
34

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;
COMIMETS te ee. 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
WOO RMN Re ee Homoptera, Chermidae
Beech trees unhealthy and slowly dying, with dead areas
of bark on stems covered with whitish masses of soft-
bodiedinsects = Homoptera, Margarodidae
Trees infested with scalelike, gall-like, or soft grublike
insects, and covered with wax in the form of powder
ORL UES Heres Fear oo is 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, 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 bor-
ing 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 occuring in the wood of green logs or lumber, re-
vealed as the logs are sawed, usually as darkly stained,
pitch-infiltrated wood, or scar (callous) tissue

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 mill, or logs utilized for
LUStiG work, ete; 0s.

Injury to freshly sawed lumber, seasoned lumber, stored and
manufactured materials, or wood in buildings

Injury to materials in contact with the ground, such as cross
ties, posts, poles: foundation materials, piling above water,
Cl Cer ‘

A. Defects in Green Timber

1. In hardwoods
MS COMITECTS inet 3 |
2. Holes small, ‘‘pinholes,”’ 14 inch or less in cross section;
circular, open 1.€., never filled with boring dust
Holes larger, “grub holes,” up to “4 inch in diameter,
usually oval in cross section, usually open, not filled
with boring dust

37

38

Pith flecks in wood, birch, maple, etc.
Diptera, Agromyzidae
Pinholes, about 1, inch in size, of uniform diameter
throughout, wood stained in streaks, in oaks and yeilow
PODIAL: cn eNVRee eee ee ee Coleoptera, Scolytidae
Holes tapering, several sizes grouped together and origin-

ating in a wound:

Holes up to 14 inch in diameter, in chestnut and

chinaguapin = ene Coleoptera, Lymexylidae
Holes up to 144 inch in diameter, in oak and other
WO OGS in. retain eee ee ee Coleoptera, Brentidae

Variable-sized holes grouped and radiating from wounds
or cavities
Coleoptera, Cerambycidae, Tenebrionidae, Brentidae
Large grub holes, up to 1 inch in diameter, usually ap-
pearing singly and not associated with wounds
Invhickonyeie a Se ee Coleoptera, Cerambycidae,
Lepidoptera, Cossidae
In poplar and cottonwood
Coleoptera, Cerambycidae,
Lepidoptera, Cossidae, Aegeriidae

In maple
Coleoptera, Cerambycidae, Tenebrionidae,
Lepidoptera, Cossidae, Aegeriidae
Invashist s<wieceed. 40 eee iene Lepidoptera, Aegeriidae
In| Persimmon ss 4¢ -aa eee Lepidoptera, Aegeriidae
In lOCUSt eases oot. ok mene Lepidoptera, Cossidae,

Coleoptera, Cerambycidae
Pitch pockets in the wood
Lepidoptera, Aegeriidae, Phycitidae
Coleoptera, Scolytidae
Holes filled with boring dust, associated with turpentine
faces or fire scars :

In: the:South, pines 2242... Coleoptera, Buprestidae

In the North, pines or other conifers
Coleoptera, Cerambycidae, Buprestidae

B. Insects in Round Logs

Sawdust exuding from small, round “pinholes” (1/10 inch
or less in diameter) on the surface of the bark; wood
usually stained around the holes

Coleoptera, Scolytidae

Sawdust exuding from larger holes; larvae present under
the barksormin: woods. =. 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
Larvae flat-headed .....................Coleoptera, Buprestidae
Larvae curved, legless; only one larva to a burrow
Coleoptera, Curculionidae

Larvae curved, legless; several larvae in a burrow,

each usually separated by a pith or clay partition
across the gallery
Hymenoptera, Sphecidae, Vespidae, Apidae
Larvae and bark beetles associates
Coleoptera, Scolytidae

C. Insects in Lumber

7. Fine sawdust exuding from small ‘“‘pinholes’” (less than
1/10 inch in diameter) in green lumber; holes usually
darkdysstamed g..0o10 eo ake. Coleoptera, Scolytidae

Sawdust, if exuding, coming from larger holes in drier
lumber, cut a month or more ................. alle oer eT SS ba

8. Damage to lumber with bark present:

Larvae elongate 20.0... Coleoptera, Cerambycidae
Marnvae curved .....8% 2 ss Coleoptera, Bostrichidae
Damage not associated with presence of bark on material

9. Fine sawdust exuding from circular or oval holes:

Small, curved larvae ............. Coleoptera, Lyctidae
Klongate larvae Coleoptera, Cerambycidae
Large, black ants associated with damage; sawdust accu-
mulating in large piles from damp wood
Hymenoptera, Formicidae
Damage concealed and sawdust usually not falling from
JOON SIS) 2a GS sass ke i RS ed a amet ALAI Sash alee etal ee base eh cil at
10. Larval tunnels packed with sawdust:
Larvae elongate, cylindrical
Coleoptera, Cerambycidae
Larvae elongate, flatheaded
Coleoptera, Buprestidae
Tunnels open:
Irregular cavities following the grain of the wood
loosely filled with fine impressed pellets ......... lsoptera
Round holes 14 inch or less in diameter, often with
cross partitions or cells
Hymenoptera, Apidae, Vespidae, Sphecidae

D. Insects in Materials 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 wood, representing
spring wood; usually extending with the grain of the
wood; sides of cavities plastered with claylike excre-
IG Mien: eet evetetr «hue has or ue noes dines bear A erate Isoptera
Large, irregular cavities eaten into wood, usually cutting
across the grain, surfaces smooth, no excrement, large
piles of sawdust accumulating outside; large, black ants
associated with injury in moist or damp wood
Hymenoptera, Formicidae

10

39

Important orders of forest insects

The primary purpose of this publication is to list and briefly
discuss the more important insects affecting forest and shade
trees and wood products. Its secondary purpose is to discuss some
of the other insects that are also abundant in the forest, but are
not usually thought of as forest insects. Many of these insects
are also of economic importance, either as pests of man or of
other forms of animal life. Given the ever-increasing use of for-
ested areas for recreational purposes and the growing public
concern about our wild life resources, it seems no longer advisable
to omit these insects from a discussion of important forest insects.

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 flat-
tened; 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 segment with a single large claw. This
claw is opposed by a toothed projection on the tibia. The body
louse, Pediculus humanus L., and the crab louse, Phthirus pubis
(L.), attack man, and a number of other species attack various
kinds of livestock and other animals. For treatments of the order
see Ewing (236) and Ferris (243). Eddy et al. (220) and Cole
et al (145) discussed the control of species attacking man.

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 are known to
attack man. The family is divided into six families. Members of
the family Trichodestidae attack various species of domestic ani-
mals; the other five (Menoponidae, Laemobothridae, Philopteri-
dae, Gyroptidae, and Ricinidae) feed mostly on birds and poultry.

Order SIPHONAPTERA
Fleas

Fleas are small wingless, hard-bodied, jumping insects. The
body is strongly flattened laterally and is armed with numerous
backward-projecting spines or bristles. There may be no eyes; the
mouth parts are formed for sucking; and the legs are long. Adults
feed on the blood of birds, wild and domestic animals, and man. 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 Eastern United States are attacked by
more than 50 different species of fleas (261).

40

Some of the more important eastern species are as follows: (a)
the human flea, Pulex irritans Lin., (b) the cat flea Ctenocepa-
lides felis (Bouché), (c) the dog flea, C. canis (Curtis), and (d)
the oriental rat flea, Xenopsylla cheopsis (Rothschild). The latter
species is the principal vector of bubonic plague and may also
transmit endemic typhus to man.

Order THYSANURA
Bristletails

Bristletails occur abundantly in the forest in rotting wood and
debris, under stones, and among fallen leaves, but none are in-
jurious. They are distinguished by the style-like appendages on
some of the abdominal segments, by the two or three tail-like ap-
pendages 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 is
capable of propelling the insect several inches into the air (512).

Springtails are widely distributed and are among the most
abundant of insects. They are found in all sorts 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 of their
help in the reduction of litter and in the formation of humus.
Bellinger (54) published a key to species occurring in eastern
America.

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
sucking type; the antennae are usually short and 6- to 10-seg-

Al

mented; 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, and fungi and in debris. A number of
species cause considerable damage to cultivated plants, but only
a few have been reported injurious to trees (17).

Liothrips umbripennis Hood became abundant enough on chest-
nut oak in New Jersey in 1937 to cause the curling of leaves.
Gnophothrips fuscus (Morgan) has damaged pine seedlings in
nurseries in New York and Rhode Island. Jack pine growing in
mixed stands on rocky slopes in Ontario has also been slightly
damaged. Female strobili of slash pine growing in Florida have
been severely damaged (216). Damage to the strobili consisted of
punctures and abrasions on the scales and bracts of flowers and
conelets. Heavy infestations resulted in shriveled conelets and
the death of affected flowers. The flewer thrips, Frankliniella
tritici (Fitch), feeds on the flowers and flower buds of hawthorn.
This sometimes prevents the buds from opening.

Order PSOCOPTERA
Book lice and psocids

Book lice and psocids are small, soft-bodied, winged or wing-
less insects, usually less than 6 mm. long. The more typical pso-
cids have well-developed wings and bear a striking resemblance
to aphids of the order Homoptera. The wings are held rooflike and
almost vertically over the body while at rest. Book lice are either
EIS or possess only vestigial wings, and are about 1 mm.
ong.

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.

Book lice occur most commonly in damp, dark rooms, not gen-
erally 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 NEUROPTERA

This order contains a wide variety of terrestrial and aquatic
insects. The adults have two pairs of large, membranous, leaf-
like wings which they hold roof-like 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 carbiform and are usually armed with very large,
curved mandibles.

FAMILY CORYDALIDAE
DOBSONFLIES

The best-known species of this family is the dobsonfly, Cory-

dalus cornutus (L.). The adult, especially the male, occasionally
reaches a length of 100 mm., has two long curving pincers or

42

mandibles, and has a wing expanse of 100 mm. to 125 mm. Fe-
males are similar in appearance except that the mandibles are
smaller. The full-grown larvae, commonly known as a hellgram-
mite, is also large and formidable in appearance. Hellgrammites
are found under stones in stream beds, especially where the water
runs swiitest. After about 214 years, they leave the water and
construct cells in which to pupate under stones, logs, or other ob-
jects on or near the bank of the streain, usually during early
summer. The hellgrammites are highly prized as fish bait; other-
wise, members of the family are of little or no economic impor-
tance.

FAMILY CHRYSOPIDAE
GREEN LACEWINGS

Members of the family Chrysopidae are all predaceous in both
the adult and larval stages on the bodies of 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 cer-
tain species of scale insects are also attacked. The adults are
green or yellowish-green and have delicate, lacelike wings. Eggs
are usually laid at the ends of gelatinous stalks, 3 to 4 mm. long,
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 (661). The golden-eye lacewing, Chrysopa oculata Say,
is an important predator of spruce gall aphids in the Lake States.

FAMILY HEMEROBITDAE
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 predaceous on other insects, principally aphids, but
also chermids, mealybugs, white-flies, mites, and occasionally dia-
spine scales. The larvae are similar in general appearance to the
larvae of green lacewings, but they do not carry packets of trash
on the dorsum. Eggs are laid on the surface of leaves, and the
winter is spent in either the larval, pupal, or adult stages, depend-
ing on species. The number of generations per year varies from
one to many, depending on species and climate.

FAMILY MYRMELEONTIDAE
ANT LIONS

Ant lion larvae, or doodle-bugs, 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 1!% to 2 inches

43

wide and from 1 to 2 inches 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 EPHEMEROPTERA
Mayflies

Mayflies are frail, delicate insects, with medium-sized, 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 are large and triangular; the hind
pair, when present, are small and rounded. The antennae are
bristle-like and inconspicuous; the mouth parts vestigial. The
nymphs are elongate and cylindrical or flattened, and have leaf-
like 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 vegetation. The adults occa-
sionally appear in enormous numbers, but they seldom live longer
than a day or two. Occasionally, their dead bodies literally pile
up along the shore, on bridges, or in the streets of nearby towns
(560).

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

Order PLECOPTERA
Stoneflies

Stoneflies are small to medium-sized, drab-colored insects, with
soft, flattened bodies. The wings are membranous and have nu-
merous 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 bases 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 the rapids of streams. When they reach maturity, they leave
the water and climb up on nearby objects to transform to the
adult stage (558).

Stoneflies may be a nuisance at times, especially when they ap-
pear in swarms in recreational areas. The nymphs are an impor-
tant source of fish food.

44

Order ODONATA
Dragonflies and damselflies

Dragonflies and damselflies are relatively large and often beau-
tifully 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
bristle-like; 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 abdomen 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 be-
cause of their large size and 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 containing 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 ex-
tended. The nymphs breathe by means of gills. In the dragonflies
gills are in the rectum; in damselflies gills are three leaflike struc-
tures at the end of the abdomen. Mature nymphs craw] out of the
water and transform to the adult stage, usually on rocks or
vegetation.

Order ISOPTERA
Termites

The order Isoptera consists entirely of termites, one of the
oldest and most primitive groups of insects. Termites are social
insects and live in colonies. In all but a few species, there is a>
definite caste system with each caste performing an essential
function in the life of the colony. Termites feed on cellulose which
they obtain from wood and other plants. Normally, cellulose is
indigestible to animals, but termites have single-celled protozoan
organisms within their digestive tracts (421) which convert it
to simple digestible foods. Many species of fungi are associated
with termites (342). Their influence on the activity of the insects
is not completely understood, but certain species such as Lenzites
trabea are known to produce an attractant (231).

Only 40 species of termites are believed to be native to conti-
nental United States. Of these, only 15 are known to occur in
the eastern half of the country, exclusive of Texas (677).

In addition to these native species, two introduced species, Cryt-
otermes brevis (Walker) and Coptotermes formosanus Shiraki,
also occur in the East. C. brevis may have been introduced into
southern Florida from a nearby island. C. formosanus is believed
to have been introduced into the Gulf Coast during World War II.

Termites are small, flat, soft-bodied insects sometimes without
eyes. The antennae are moniliform, and in the winged adults, the
number of antennal segments varies from 12 to 25 or more. The

45

median ocellus is absent. In certain species it is replaced by a
more or less distinct opening of a gland known as the fontanel.
The wings are long and narrow and, when folded on the back,
extend well beyond the end of the abdomen. In all North Ameri-
can species, the fore and hind wings 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. For example,
there are no internal insect parasites, and predaceous enemies are
largely confined to ants and birds. The predators sometimes con-
sume large numbers of termites during flight periods. This may
have some adverse effect on the establishment of new colonies, but
it has no effect on the parent colony. Fungi, nematodes, and bac-
teria usually kill a small percentage of the individuals in a colony.
Moulds, occasionally, cause much heavier losses.

In many parts of eastern United States, particularly the South,
termites are almost universally present in stumps, dead logs and
other woody materials in contact with the ground in forested
areas. Termites are largely beneficial in forested areas because
of their assistance in the decomposition of dead wood and its re-
incorporation into the soil. On the other hand, when termites in-
fest wooden structures and other useful wood products, they are
often extremely destructive. The extent of the monetary losses
they cause is not known, but has been estimated at $400,000,000. to
$500,000,000. per year in the United States for damage, repairs,
and control.

For further information on termites, the reader is referred to
the following (31, 229, 435, 670, 669, 677, 674, 673, 676, 441).

Keys to Termites of Eastern United States Based on Soldiers

1. Head long, much longer than broad 2
Head short, truncated in front . Abie Ye 9
2. Mandibles with prominent mar ginal teeth - igh Beg 3
Mandibles without prominent marginal Pere 3
3. Hind femora swollen; third antennal segment not equal to
fourth and fifth ............... Kalotermes approximatus
Hind femora swollen; third antennal segment slightly
longer than fourth . _Neotermes jouteli
Hind femora swollen; third antennal segment enual to
fourth and fifth . AD <A Dice De he soca eae 4
4, Anterior margin of pr onotum dentate
Incisitermes synderi
Anterior margin of pronotum not dentate
Incisitermes schwarzt
5. Labrum rounded at tip Prorhinotermes simplex
Labrum pointed at tip ere. 6
6. Fontanelle large and tub: lar; head oval and yellow;
length of body 5.8 mm. ..... Coptotermes formosanus
Fontanelle not large and tubular, sometimes very small,
circular and distinct . 7

7. Gula less than twice as broad in front as in middle
Reticulitermes tibialis
Gula fully twice as broad in front as in middle 8

8. Head 114 times as long as broad; length 4.6 to 4.9 mm.
Reticulitermes arenicola
Head 124 times as long as broad; length 6 to 7 mm.
Reticulitermes flavipes
Head 134, times as long as broad
Reticulitermes hageni
Head twice as long as broad ......... Reticulitermes virginicus
9. Head cleft and bilobed in front; front of head blackish
Calcaritermes nearcticus
Head not cleft or bilobed in front; front of head blackish
tor dark castaneous: behind’ :22.20. 4 ee A)
10. Anterior of head tuberculate 00.0000... Crytotermes brevis
Anterior of head not tuberculate
Cryptotermes cavifrons

Subterranean Termites

Subterranean termites 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 is otherwise available. In
order to attack wood located away from the ground where a sup-
ply of water is not otherwise available, they construct covered
passageways, commonly called shelter tubes (fig. 5A).

Regardless of the extent of damage they cause to the interior
of wood (fig. 5B), these termites always leave a covering shell
intact. Because of this, there usually is no external evidence of
infestation, even though the interior of the wood may be de-
stroyed. The first indication of infestation may be the swarming

F-519932-33
FIGuRE 5.—Subterranean termite: A, shelter tube on foundation wall;
B, damage to a 2 x 4 inch pine stake.

AZ

of 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,
telegraph poles, fence posts, and other wood products, but also
to paper, fiberboard, and various types of fabrics derived from
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 feed on the soft spring
wood only. As a result their galleries run parallel with the grain.
These galleries are characteristically 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.

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

The eastern subterranean termite, Reticulitermes flavipes (Kol-
lar) (fig. 6), one of the most destructive subterranean termites
in eastern America, is widely distributed, occurring from the
Gulf of Mexico north to Ontario, and from the East Coast to the
Great Plains. Infestations normally occur in the forest where the
dead wood of practically all species of trees is attacked. Infesta-
tions also occur in the woodwork of buildings, in cellulose mate-
rials stored therein, in poles and posts, and occasionally in living
trees, shrubbery, flowers, and crop plants.

Each colony of the eastern subterranean termite comprises
three castes: reproductives or adults, workers, and soldiers. The
reproductives (fig. 6A) are soft-bodied and brown or black, have
two pairs of long, whitish opaque wings of equal size, and are
10 to 12 mm. long. Workers, (fig 6B) are soft-bodied, grayish-
white, and slightly less than 6 mm. long. Soldiers (fig. 6C) are
similar to the workers except for their much larger, longer heads,
their longer and more formidable mandibles, and their slightly
greater length. Winged reproductives are often mistakenly iden-
tified as flying ants. They differ from ants, however, in having
the abdomen broadly joined to the thorax. The abdomen of an
ant is reduced to threadlike proportions where it joins the thorax.
The wings of ants are also transparent, and the two pairs are of
unequal size.

Colonizing flights occur most frequently after the first warm
days of spring, often following a warm rain. They may also occur
during the remainder of the warm season and sometimes even

48

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

A few days after mating the female lays from six to twelve
eggs. Hatching occurs in about 30 to 90 days and the nymphs
reach maturity in about one year. Later, both soldiers and repro-
ductive nymphs appear in the colony, and reach maturity in 1
and 2 years, respectively. Mating continues at irregular inter-
vals, and the colony continues to increase in size. The original
pair of reproductives live together for life, sometimes for many
years. In well-established colonies, there may be hundreds of
thousands of individuals. Such colonies may be so widely spread
out that it is difficult or impossible to estimate their size or to
locate their main parts.

Reticulitermes tibialis Banks, the arid-land subterranean termite,
is probably the most widely distributed species of Reticulitermes
in the United States. It is largely confined to the western half of
the country, but also occurs eastward into the North Central
States as far as the Chicago area. In the latter area, it is found
among sand dunes where it infests small pieces of wood partly

F-107105
23 FIGURE 6.—Eastern subter-
ranean termite, Reticulit-
ermes flavipes: A, adult;
B, worker; C, soldier.

A9

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 it is fully capable of damaging build-
ings, it is not very often injurious to them because of the thinly
populated regions in which it occurs. Winged adults are almost
entirely shiny black and about 10 mm. long. The tibiae are also
blackened and the pronotum is broad.

Reticulitermes virginicus Banks occurs in the southeastern and
central western parts of the United States. It is often mistaken
for R. flavipes, but is smaller. Winged adults are about 8 mm.
long, and soldiers 4.5 to 5 mm. long. R. hageni Banks occupies
about the same territory as R. virginicus. Winged adults are yel-
lowish and about 8 mm. long. Soldiers are similar to those of R.
virginicus except for shorter heads and paler color. These two
species are reported to occur more commonly in dry, forested
areas than R. flavipes. R. arenicola Groellnes occurs in the Chi-
cago area and has been found in and under decaying pieces of
wood in sandy places near sand dunes. Its habits are similar to
those of Ff. tibialis which occurs in the same area. Winged adults
have grayish wings and are about 9 to 10 mm. long. Soldiers
range in length from 4.6 to 4.9 mm.

The Formosan subterranean termite, Coptotermes formosanus
Shiraki, is an introduced species. Active colonies were first re-
corded in North America in a shipyard warehouse in Houston,
Texas, in 1965. Since then additional colonies have been discov-
ered in Houston and Galveston, Texas; Charleston, South Caro-
lina; and New Orleans and Lake Charles, Louisiana. The species
was described from individuals collected in Formosa in 1909.
Since then it has been reported from Ceylon, mainland China,
Guam, Hawaii, Japan and South Africa. Incipient colonies gen-
erally 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 may
be found almost anywhere with favorable conditions. Colonies
have been observed on boats, ships, barges, dredges, water tanks,
piers, floating drydocks, in living and dead trees, and in wooden
buildings.

Workers are over-all grayish-white. 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 long and the legs are slender and hairy. Soldiers (fig. 7) have
oval heads and slender bodies. The pronotum is short, elliptical,
and notched at the middle of the frontal margin. A small, short
tube-like process extends from the frontal gland and exudes a
milky, acidulous secretion. The abdomen is slender, entirely pu-
bescent, and has a 3-segmented caudal appendage. Males have
large, hyaline wings, 10.6 mm. long, about three times as long as
the abdomen, and about three times as long as broad. The head is
hexagonal and brown; the frons is irregularly concave at the
center where a globular projection occurs; the pronotum is semi-
circular and as broad as the head; the legs are short, large, and
vellowish-brown; and the abdomen is short, eliiptical and yel-
lowish-brown. Females are similar to males but are slightly
larger. Their bodies are marked with minute spots, and their
wings are 12 to 14 mm. long.

50

F-520104
FIGURE 7.—Soldier castes of the Formosan subterranean termite
Coptotermes formosanus.

Formosan subterranean termite nests are constructed from a
friable material called carton. It consists of a mixture of masti-
cated wood, saliva, and excrement. A nest may be several cubic
feet in size. Tunnels radiate from nests constructed in the soil.
They can be found at depths of 10 feet in the earth, and they can
extend horizontally to 200 feet. Their walls are lined with essen-
tially the same materials used in constructing the nest and they
are nearly impervious to water. Primary queens can lay up to
1,000 eggs per day, and a single colony may contain hundreds of
thousands of individuals. Winged reproductives swarm during
May and June. They are poor fliers, and the majority drop to the
ground within 100 yards of the nest unless carried farther by
the wind. First evidence of a colony’s presence may be the ap-
pearance of these winged adults at swarming time.

Large colonies can cause severe damage in a short time (fig.
8A). In Hawaii, walls of new buildings have been hollowed out
in three months’ time. Living trees are also hollowed out (fig. 8B)
and weakened. Known susceptible trees in the South are the
Chinese elm, several species of oak, hackberry, and Arizona ash.
Dead trees are highly susceptible. Extensive damage all the way
to the top of 70-foot-tall cypress snags has been observed in
Louisiana (43).

51

F-519935

FIGURE 8.—Formosan subterranean termite colony and damage: left, carton

nest and damaged wood above a window inside a building; right, colony
of workers in a cypress log.

Nonsubterranean Termites

Several species of nonsubterranean termites occur in the East-
ern States. They are found throughout the state of Florida; also
in a narrow strip along the Atlantic Coast as far north as south-
eastern Virginia, and westward along the Gulf Coast to Mexico.
Infestations are found in structural timber and other woodwork
in buildings; in furniture; in utility poles; in wooden derricks and
piled lumber; in wood pulp or fiber insulation boards; in other
products containing cellulose; and in trees and other plants. Be-
cause of their ability to live in wood that is frequently moved,
nonsubterranean termites are often found in regions far removed
from their normal range, including Canada. None of these are
able to establish permanent infestations (675).

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 spring
wood and the harder summer wood. Their cavities contain com-
pressed pellets of partly digested wood. Some pellets are also
pushed to the exterior through small holes and are found in piles
on the floors of infested buildings (fig. 9).

Nonsubterranean termite colonies contain only two castes of
adults: reproductives and soldiers. Since there is no worker caste,
the work of the colony is performed by young nymphs. Winged
adults are distinguished by the presence of branches between the
upper rim of the wing and the first long vein. These are absent in
winged subterranean termites. Soldiers differ from the soldiers
of subterranean termites in having their mandibles lined with
teeth on the inner edge.

Winged adults usually swarm during the early evening hours
of late winter or spring. Unless carried by the wind, they fly for
only short distances in search of places to found a new colony.
Once they have succeeded, they shed their wings and bore directly

52

into the wood. Then they plug the opening, thereby sealing them-
selves in. Colonies grow very slowly, are never confined to a single
chamber and never contain more than a few thousand individu-
als. Piles of pellets on the floor may be the first evidence of in-
festations. Other tell-tale evidence consists of pitted and rough-
ened surfaces of infested floor surfaces, door frames, and other
wood.

Nonsubterranean termites usually are less injurious than sub-
terranean species in the United States. However, large public
and other buildings in several Florida cities; New Orleans, Lou-
isiana; Savannah, Georgia; Charleston, South Carolina; and less
commonly elsewhere, have been seriously damaged (672).

Incisitermes snyderi (Light) is the most widely distributed and
injurious of the eastern nonsubterranean termites. It occurs from
South Carolina to Florida and west to Brownsville, Texas, mainly
along the coast. Infestations occur in the woodwork of buildings;
in untreated utility poles, and in dead trees, logs, and branches.
Winged adults have yellowish heads and are 11 to 12 mm. long.
Soldiers are 7 to 8 mm. long and have the anterior margin of the
pronotum dentate.

Incisitermes schwarzi (Banks), the southern dry-wood termite,
occurs in Florida south of Pensacola, where it is the commonest
species. Infestations are found in the woodwork of buildings, in
the bases and tops of utility poles, and in dead trees, logs, and
stumps. Winged adults have yellowish heads and are 15 to 16 mm.
long. Soldiers resemble the soldiers of J. snyderi except that the
anterior margin of the pronotum is not dentate.

Kalotermes approximatus Snyder has been recorded from
northern Florida; New Orleans, Louisiana; and southern Vir-
ginia. Infestations occur in the woodwork of buildings, in living
and dead trees, and in logs and stumps. Winged adults are black-
ish in color (249).

Neotermes jouteli Banks occur in southern Florida, including
the Keys. It occasionally occurs in the moist foundation timbers
of buildings. In nature, it lives in dead trees and in logs and
branches lying on the ground. Adults are yellowish or light cas-
taneous and about 10 mm. long. .

eS
@¢;

ere 6
#n8 92%

% .

F-519928
FIGURE 9.—Excretal pellets of
nonsubterranean termites

Do

Neotermes castaneus (Burmeister) occurs in southern Florida,
including the Keys. It has not been found damaging the woodwork
of buildings. Infestations have been found in the dead wood of
trees, logs, stumps, and branches; also in living citrus trees.

Prorhinotermes simplex (Hagen) occurs in swampy areas of
southern Florida where it is found in moist logs. It has been
known to attack the woodwork of buildings, but then extremely
rarely. Adults are yellowish-brown and about 9 mm. long.

Cryptotermes brevis (Walker), the tropical rough-headed pow-
der-post termite, occurs commonly in southern Florida and New
Orleans, Louisiana. Isolated infestations have also been found
elsewhere, including Memphis, Tennessee. This species is found
only in the woodwork of buildings and in furniture. It not only
destroys these products, but the constant dropping of pellets from
infested wood is a source of annoyance to the householder. Winged
adults are 10 mm. long. Soldiers have the front of the head black-
ish to dark castaneous behind and its anterior margin tuberculate.

Cryptotermes cavifrons Banks, a powder-post termite, occurs
in southern Florida where it occurs in dead trees, logs, stumps,
and branches. Winged adults are about 8.5 mm. long. Soldiers
resemble the soldiers of C. brevis except that the anterior margin
of the head is not tuberculate.

Calcaritermes nearcticus (Snyder) is also a powder-post ter-
mite. It, too, is known only from Florida. Winged adults are small,
have smoky wings, and are coarsely punctate. Soldiers have the
anterior of the head bilobed. The head is also blackish in front
and light castaneous behind.

Order ORTHOPTERA

Cockroaches, grasshoppers and allies

The order Orthoptera contains some of our most familiar in-
sects, 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 gen-
erally long and narrow, many-veined, and leathery or parchment-
like. 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: (82,
LulfDp, OF BOO), Cid oI, DO, Bil UO, Biles, WOO)

FAMILY MANTIDAE
PRAYING MANTIDS
Praying mantids are large, elongate, rather slow-moving in-
sects. The eyes are very large; the head is wider than 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

D4

long and the front femora and tibiae are armed with strong spines
and fold together to form a pincer. Praying mantids are preda-
ceous and feed on a wide variety of other insects. To capture 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 lightning-like speed to grasp it.
Because of their habit of holding their front legs in an upright
position, these insects are commonly called “praying mantids.”’

Praying mantids lay their eggs in the fall in paper-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 (Johannson), is
the most common species in the Southern States. It occurs from
the Atlantic Coast to New Mexico and north to Nebraska, Penn-
sylvania, and Illinois. Adults are 75 to 100 mm. long. The male is
grayish-brown with smoky-brown outer wings and often with
greenish-yellow body and legs. Females are either colored like the
males or are greenish-yellow with bright green forewings. S. flori-
densis Davis, a somewhat more slender and longer species, occurs
in Florida.

The Chinese mantid, Tenodera aridifolia sinensis Sauss., an in-
troduced 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. 10). Females are green or greenish-
yellow; males are the same color or wholly brown, or brown with
margins of the forewings green. The narrow-winged mantid, 7.
angustipennis Sauss., also an introduced species, is similar to but
more slender and smaller than the Chinese mantid. It is widely
distributed in the Eastern States.

Two other eastern species are the European mantid, Mantis ve-
ligiosa L., an introduced species (medium sized and greenish-
yellow) and Litaneutria minor (Scudder). The latter is about 25
mm. long, and occurs on the Great Plains (316).

FAMILY PHASMATIDAE

WALKINGSTICKS

Eastern species of walkingsticks are long, slender, and subcyl-
indrical insects. The head is free and nearly horizontal; the an-
tennae are long, and the eyes small; the abdomen is elongate; the
legs very long and slender; and the wings are rudimentary, except
for one species found in Florida. Walkingsticks are slow-moving
insects, and all are plant feeders. The eggs are hard-shelled and
are often dropped promiscuously to the ground.

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

Adults are 62 to 87 mm. long and, while motionless, closely re-
semble the twigs of their hosts (fig. 11). The body color is vari-

D5

able. 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-
colored band on one edge. It is about 2 mm. long.

Winter is spent in the egg stage and hatching occurs in May or
early June. The nymphs feed at first on shrubs such as sweet fern,
blueberry, strawberry, and juneberry. 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 deposition.

Severe outbreaks occur frequently in the Lake States. They also
occur occasionally south of a line drawn from Nebraska to Dela-
ware. 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 (304). The

COURTESY CONN. AGR. EXPT. STA.

FIGURE 10.—Adult and egg
mass of the Chinese man-
tid, Tenodera aridifolia
sinensis.

F-504615

FIGURE 11.—Adults of the

walkingstick, Diaphero-

mera femorata, on leaves
of oak.

hymenopterous parasite, Mesitiopterus kahlit Ashmead; various
predators such as crows, robins, and other birds; and dry weather
during the period of egg hatch are important natural control
factors (775).

Several other species of walkingsticks also occur in eastern
United States. The two-striped walkingsticks, Anisomorpha bupres-
toides (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 eggs, each in small pits dug in the soil.
A. ferruginea (P de B.) feeds on various trees and shrubs from
southeastern Nebraska and Arkansas through the high country to -
Georgia and the Carolinas. Diapheromera velit Walsh and D.
blatchleyi (Caud.) feed on grasses and tall shrubs. D. vel occurs
on the Great Plains; D. blatchleyi, from the Great Plains to the
Atlantic Coast. Megaphasma denticrus (Stal.), the giant walking-
stick, sometimes attains a length of 150 mm. Its habits are similar
to those of D. femorata, but it is apparently never abundant
enough to be injurious.

FAMILY ACRIDIDAE
SHORT-HORNED GRASSHOPPERS

This family contains most of the well-known grasshoppers,
many of which are frequently very destructive to agricultural
crops. Ordinarily, they are not very injurious to trees, but they
may be seriously damaging during outbreaks. Young trees in nur-
series, shelterbelts, and plantations are particularly vulnerable,
especially on the Great Plains, in the upper Mississippi Valley,
and in the Lake States. The adults are distinguished by their
short filiform, or three-jointed, clubbed, antennae; short and in-

o7

conspicuous ovipositors; auditory organs on the sides of the first
abdominal segment; greatly enlarged hind legs; narrow front
wings; and broad, membranous, fanlike hindwings.

The majority of short-horned 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
distances. All species have much the same life history. Nearly all
lay their eggs in pods at depths of 1 to 3 inches 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 species may be injurious in nurseries, planta-
tions, and shelterbelts during epidemics: (1) The lesser migratory
grasshopper, Melanoplus sanguinipes—the adult is about 2 mm.
long and reddish-brown with a distinct patch of black on the neck
or collar; (2) The differential grasshopper, M. dijfferentialis
(Thos.)—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 hind legs. (8) The two-striped grasshopper, M. bivittatus
(Say)—the adult is about 30 mm long. It is slighty more robust
than the differential grasshopper and is greenish-yellow beneath,
with two yellow stripes running the full length of the dorsum.
(4) The red-legged grasshopper, M. femurrubrum (DeG.)—the
adult is about 18 mm. long. It is reddish-brown above, yellowish
beneath, has colorless hindwings, and has red-tinged hind wings.
(5) The clear-winged grasshopper, Camnula pellucida (Scudd.)—
the adult is about 12 mm. long. It is yellow to brown; the front
wings are blotched with large brown spots; and the hindwings
are clear or pellucid.

A few species of grasshoppers are more closely associated with
trees than others. One, Dendrotettix quercus Pack., the post-oak
locust, is widely distributed from east central Texas to the Lake
States, Long Island, and southeastern Canada. Oaks are its pre-
ferred hosts, but it has also been observed on pines in the Lake
States. Several outbreaks have been recorded, some of which
covered several square miles. 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: (1) D. australis
(Morse)—apparently feeds exclusively on Virginia pine in the
Southeastern States; (2) Melanoplus punctulatus (Scudd.)—has
been observed defoliating young white pines in plantations in
Connecticut; (8) M. bruneri Secudder—inhabits coniferous for-
ests; and (4) the eastern lubber grasshopper, Romalea microptera
(P de B.)—feeds on shrubs in Florida and Alabama.

FAMILY TETTIGONIDAE
LONG-HORNED GRASSHOPPERS AND KATYDIDS

Long-horned grasshoppers and katydids are mostly large with
hair-like 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 file-like 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 in color 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 com-
mon species, none of which are very injurious, are discussed
below.

The fork-tailed bush katydid, Scudderia furcata (Brun.), so-
called because of the forked appendages at the tip of the abdomen
of the male, is widely distributed. It occurs on but is not restricted
to trees. The related species, S. curvicauda (DeG.), lives commonly
on oak.

The broad-winged katydid, Microcentrum rhombifolium (Sauss.)
is widely distributed in the East. Adults are 25 to 35 mm. long
and leaf green. The angular-winged katydid, M. retinerve (Burm.),
a smaller species, is more southerly in its distribution. Both
species have long, narrow wings, and the vertex is narrowed
anteriorally.

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 col-
onies 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, rarely spotted
with brown, which occurs on pines in the Southeastern States, and
Conocephalus brevipennis (Scudder) 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 Hald. Although primarily western in distribu-
tion, it occasionally occurs in destructive numbers as far east-
ward as the Dakotas and Kansas. It feeds mostly on various trees
and shrubs. The ovipositor of the female is sword-shaped and is
up to 87 mm. long.

FAMILY GRYLLIDAE
CRICKETS

Crickets are medium-sized insects, usually with long filiform
antennae, three-jointed tarsi, spear-shaped ovipositors, and hind

59

legs fitted for jumping. The wings of certain species are fully-
developed; in others, they are either abbreviated or absent. They
lie flat on the back and bend down abruptly at the sides of the
body. The winged forms possess auditory tympanae on one or
both sides of the fore tibia. The males also possess sound-produc-
ing 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 (4). 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-jointed, 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, tree
hoppers, and scales; also on the leaves, flowers, and bark of trees.

The snowy tree cricket, Oecanthus fultonti T. J. Walker [O.
niveus], 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 antennial
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 appearance, 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 Beut. occurs
on pines and lays its eggs in regular rows on the bark. O. latt-
pennis Riley occurs commonly on shrubs and low trees, especially
on scrub oaks in dry open areas in the Eastern States.

Anurogryllus muticus (DeG.), the 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 (748).

FAMILY GRYLLOTALPIDAE
MOLE CRICKETS

Gryllotalpa gryllotalpa (L.), the European mole cricket, an in-
troduced 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 vel-
vety hairs and are up to 37 mm. long. The front legs terminate in
four dactyls which are used for digging. This species feeds at
night by tunneling in the upper inch or two 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 in-
jurious in nurseries.

The changa, Scapteriscus vicinus Seudder, 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 Scudd. occurs
in Georgia and Florida. Adults are brownish-fuscous and blotched

60

with yellow. The southern mole cricket, S. acletus R. & H., occurs
from Georgia to Texas. It is pinkish-buff in color and somewhat
more slender than S. abbreviatus. All members of this genus feed
on earthworms, roots, and other 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, are of no economic impor-
tance 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 ovipositors in the
females.

Order HEMIPTERA

True bugs

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

The majority of the Hemiptera have the basal portion of the
front wing 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 membranous and
usually slightly shorter than the front wings. Both pairs of wings
lie flat over the abdomen with the membranous distal portion of
the front ones overlapping. 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 odors when disturbed. Man usually finds
these odors offensive.

Blatchley (83) and Britton (103) have discussed the Hemiptera
of eastern North America.

61

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-seg-
mented 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 diffi-
cult to see when they rest on the bark of limbs and trunks of
trees. Most species are plant feeders, but some feed on other in-
sects, and some feed on both.

Tetrya bipunctata (H.-S.) has been recorded feeding on slash
and longleaf pine cones in the South. Adults are dark reddish-
brown and about 15 mm. long. Because of their coloring, they are
not readily noticeable when motionless on a cone. Females lay
eggs on needles near cone clusters. The nymphs are gregarious.
When they reach the third instar they begin to feed on the seeds
within green closed cones. This type of feeding is continued in
the adult stage (188).

Other plant feeding species and their hosts include the follow-
ing: (a) Brochymena quadripustulata (Fab.)—feeds through the
bark of limbs and trunks of elm, oak, and willow. Brownish, hair-
like lines running across the grain in the cambial region are evi-
dence of its attack. Adults frequently overwinter in houses. (b)
B. carolinensis (Fabr.) —feeds through the bark of slash and long-
leaf pines. (c) Elasmuche lateralis (Say)—feeds on the leaves and
catkins of yellow birch (also reported feeding on the larvae of
Tetralopha asperatella in the Lake States). (d) Pitedia uhleri
(Stal.)—feeds on the larger branches and trunks of white pines
in the Lake States. |

Predacious species include: (a) the spined soldier bug, Podisus
maculiventris (Say). It 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 punc-
tures and is about 12 mm. long. (b) Stiretus anchorago (F.) feeds
on the larvae of many species, including the gypsy moth and tent
ei area Adults are shiny, dark metallic, and 8 to 11 mm.
ong.

FAMILY TINGIDAE
THE LACE BUGS

Lace bugs, so-called because of their broad gauzelike or lace-
like wing covers, are small and usually whitish. They are also
flat, oval, or rectangular with reticulated surfaces and are about
5 or 6 mm. long. The head is often hidden beneath a large hood
on the front of a greatly modified pronotum, which projects be-
yond the sides of the body. The abdomen is completely hidden by
the wing covers, which are frequently transparent. The nymphs
are black and often covered with long spines.

Many species of lace bugs live and feed on the under-surfaces
of leaves. A given species usually occurs either on a single host

62

or on closely related ones. The upper surfaces of infested
leaves may be either whitened, or brownish, or dead in appear-
ance. The undersurfaces are speckled with eggs, excrement, and
cast skins of the developing nymphs. The leaves of heavily in-
fested 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 te the undersurfaces of leaves or
embedded in leaf tissues). Most species have two generations a
year.

The sycamore lace bug, Corythucha ciliata (Say), occurs
throughout the Eastern United States (725) and in southern Can-
ada. Its preferred host is sycamore, but it also feeds on ash,
hickory, and mulberry occasionally. The adult (fig. 12) 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 stip-
pling of foliage. Heavily infested leaves of sycamore turn white and
drop prematurely. During dry weather this may result in severe
injury.

COURTESY CONN. AGR. EXPT. STA.
FIGURE 12.—Adults of the syca-
more lace bug, Corythucha ciliata.

The oak lace bug, Corythucha arcuata (Say), feeds on white,
bur, and chestnut oaks from Alabama and the Carolinas to
southern Canada. Connell and Beacher (156) discussed its life
history and control. 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.

The elm lace bug, Corythucha ulmi O. & D. feeds on American
elm in many Eastern States and southern Canada, and on Si-
berian elm on the Northern Great Plains. It is capable of defoli-
ating its host.

Heavy infestations of Corythucha mollicula O. & D. (=salicis
O. & D.) (=canadensis Parsh.) may seriously injure willow, its
only known host, throughout the Eastern States. Heavy infesta-
tions of the hackberry lace bug, C. celtidis O. & D., often occur on
hackberry in the Midwest. The species has also been reported
from Florida. C. pallipes Parsh. (=betulae Drake=cyrta Parsh.)

63

is often abundant on young yellow birch. Other hosts include
white birch, beech, eastern hophornbeam, willow, mountain-ash,
and maple. C. pergandei Heidemann feeds principally on alder;
also occasionally on hazel, elm, and birch. C. pruni O. & D.
(—pyriformis Parsh.) feeds on wild cherry; C. juglandis (Fitch)
on black walnut, butternut, and linden; C. elegans Drake on wil-
low, balsam poplar, quaking aspen, and bigtooth aspen; C. aescult
O. & D. on buckeye; C. associata O. & D. on wild cherry; C. bellula
Gibson on hawthorn; and the hawthorn lace bug, C. cydoniae
Fitch, on hawthorn and pyracantha.

The rhododendron lace bug, Stephanitis rhododendri Harv., an
introduced species, occurs from New England to North Carolina
and Ohio, and infests rhododendron, mountain-laurel, and azalea.
The adult is about 4 mm. long, and has a triangular hood and
long, slender antennae. Its wings are rounded at the apex and are
nearly twice as long as the abdomen. Winter is spent in the egg
stage and there are at least two generations per year. Rhododen-
dron plants infested continuously for several ae turn yellow
and lose their attractiveness.

The azalea lace bug, Stephanitis pyrioides (Scott), resembles S.
rhododendri except for its duskier wings. It feeds on azalea, pre-
ferably the evergreen varieties. The leaves of heavily infested
plants become almost white and many of them dry out completely
and drop. There may be three generations per year.

The basswood lace bug, Gargaphia tiliae (Walsh), often occurs
in large numbers 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.
Others are blood-sucking and frequently bite man. 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 well over 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 which usually curves downward in the form of a
semi-loop, 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.

The wheel bug, Avilus cristatus (L.), is an important predator of
various forest insects. As a young nymph it feeds on aphids;
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: (a) Sinea spi-
nipes (H.-S.) which feeds on the fall webworm in the South; (b)

64

Acholla multispinosa (DeG.), an enemy of the pine webworm;
(c) Zelus exsanguis Stal., an enemy of the gypsy moth; and
(d) Melanolestes picipes (H.-S.), an enemy of May beetles and
their larvae. Readio (611) discussed the biologies of members of
the family.

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, members of this family of small bugs
are predaceous on other insects. The adults may be found on
flowers, under loose bark, in leaf litter, or in decaying fungi.
Anthocoris musculus (Say) (=borealis Dallas) 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 the pine twig gall scale in Mas-
sachusetts. The majority of flower bugs are black with white
markings and are only about 3 to 5 mm. long. Orius insidiosus
(Say) is a common predator of insect eggs.

FAMILY MIRIDAE

THE PLANT BUGS

This is the largest family in the order Hemiptera, with about
1600 species occurring in the United States and Canada alone
(424). The majority of species appear to be phytophagous. Many
others are predaceous and feed on a wide variety of young and/or
soft-bodied insects. The adults are 2 to 9 mm. long, and are usu-
ally fragile. 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. Tropidosteptes amoenus Reuter [= Neoborus amoenus
Reuter] has caused noticeable injury to ash seedlings in nurseries
in the Lake States; the tarnished plant bug, Lygus lineolaris (P. de
B.), is often injurious to ornamentals and to forest nursery trees.
Young, succulent growth of elm is frequently damaged by the
feeding of Neolygus invitus (Say).

65

FAMILY COREIDAE
COREID BUGS

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

Leptoglossus corculus (Say) occurs commonly in longleaf and
slash pine seed orchards in north Florida. First and second instar
nymphs feed on the needles and first year conelets; older nymphs
and adults feed on the seeds. Adults are reddish-brown to black,
have long legs, and average 18 mm. in length. A white zig-
zag jine crosses the wings, and the hind tibiae are fiattened and
leafiike. Infested cones usually show no external damage symp-
toms (188). The related species, L. occidentalis Heidemann, a
serious pest of Dougias-fir seed in California (424) has been ob-
served feeding on the needles and green cones of Austrian pine in
Missouri. ;

FAMILY RHOPALIDAE

The boxeider bug, Leptocoris trivittatus (Say), often becomes
a pest wherever boxeider is grown as a shade tree in the United
States and Canada. In heavily infested areas, it will feed on ash
and mapie. Adults (fig. 13) are somewhat fiattenea, brownish-black

F-508517
IGURE 18.—Adult of the boxelder
bug, Leptocoris trivittatus, on
seed of boxelder.

on top, and about 12 mm. long. There are also three red longi-
tudinal stripes on the thorax; the margins of the basal half of the
wings are red; and the abdomen is bright red. Nymphs are wing-
less 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, they be-
come active and come out of hiding, only to retreat again when it
turns cold. During the spring, they emerge and fly to 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 in leaves, fruits, or soft seeds by
inserting their beaks into the tissues. Feeding continues through-
out 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 or spring burning of leaf litter also discour-
ages the congregating of the insects (791).

FAMILY LYGAEIDAE
LYGAEID BUGS

This is a rather large family, most members of which feed on
mature seed. The adults are smal! 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 her-
baceous vegetation in the forest. One species, Kleidocerys resedae
geminatus (Say), feeds on the catkins of yellow and gray birch,
rhododendron, etc.

FAMILY THAUMASTOCORIDAE

Xylastodoris luteolus Barber, the reyal palm bug, is the only
member of this family known to occur in the United States. It
feeds on the royal palm, Roystonea regia, in southern Florida.
Adults are pale yellow, flattened, and about 2 to 2.5 mm. long.
Eggs are deposited on the undersurfaces of leaflet midribs, usu-
ally on older trees. Feeding occurs on unfolded leafiets of newly-
emerging fronds and to some extent the spike or projecting part
of the terminal bud. Fronds exposed to heavy feeding may turn
brown (32).

Order HOMOPTERA

Aphids, scales, mealybugs, etc.

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.

67

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 man-
dibes and maxillae). Many species are wingless, at least in the
female sex under certain conditions. Where wings are present,
they are usually four in number with the front pair longer and
the hind pair often wider. The wings do not overlap much at the
tip, and their bases are never abruptly thicker than their tips.
They usually stand sloping 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.

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 alteration 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. Members of
different genera differ greatly in body and form. Those occurring
in this country are usually less than 12 mm. in length, and many
have the head greatly modified, with the part in front of the eyes
greatly enlarged and snoutlike. A useful characteristic for recog-
nizing them is the position and form of the antennae. The two
basal segments are stout, whereas the remainder consist of a
nearly pear-shaped basal segment and a segmented, bristle-like
terminal part. They also have a few large spines on the hind tibiae.
Two fairly common species are Anormenis septentrionalis (Spin.)
and Metcalfa pruinosa (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 dis-
turbed. Osborn (574) and Van Duzee (731) discuss the group.

FAMILY CICADELLIDAE
THE 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 hav-

68

ing one or more rows of small spines extending the length of the
hind tibiae. Because of the nature of their feeding, which con-
sists of piercing plant tissues with their mouth parts and
sucking the juices, the damage they cause is usually not recog-
nized or is attributed to other factors. About the only visible
effects of leafhopper feeding are the white stippling of foliage or
the browning, withering, and curling of leaves. Because of the
difhculty in observing or evaluating 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 con-
tribute significantly to the general unproductiveness of many
stands of valuable hardwoods in Eastern United States. Many
species are also vectors of destructive plant diseases, especially
diseases of virus origin.

Some species of leafhoppers spend the winter in the egg stage;
others as adults. Overwintering eggs usually hatch in May or
June, and the young nymphs feed on new, tender leaves. Over-
wintering adults emerge during the first warm days in spring and
lay eggs as soon as the leaves of their hosts are fully developed.
These eggs 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
Hrythroneura, Empoasca, and Typhlocyba (189). Several species
of Idiocerus feed on willow and poplars. Various species of Mac-
ropsts feed on poplars, willow, and honeylocust. Certain species of
Scaphoideus, Gyponana, and Ponana oecur on American elm; oaks
are infested by species of Alebra, EHutettix, 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 white-banded elm leafhopper, Scaphoideus
luteolus Van D., the vector of elm phloem necrosis, a virus disease
of American elm (21). Adults are difficult to separate from those
of closely related species, but not so the nymphs (fig. 14). 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
Eastern United States, from New York west to Kansas, Nebraska,
and Iowa, and south to Georgia and Alabama.

The white-banded elm leafhopper lays its eggs in the cork par-
enchyma of elm bark. When the eggs hatch the young nymphs
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 mid-
ribs 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 (22).

Elm phloem necrosis virus is transmitted by the adults which
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 virus. All of the details of the life cycle of the species

69

F-519937-8
FIGURE 14.—Adult and nymph of the white-banded elm leafhopper,
Scaphoideus luteolus.

have not 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), H. insolita (Wlk.), Oncometopia
orbona (F.) [= O. undata], Cuerna costalis (F.), and Grapho-
cephala versuta (Say) transmit the virus causing phony peach, a
destructive disease of peach trees in the South.

FAMILY MEMBRACIDAE
TREEHOPPERS

Treehoppers are characterized by the prolongation of the pro-
notum 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 tip of the abdomen and completely covers the
wings. The hind legs are long and adapted for jumping, and the
female’s ovipositor is 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. Osborn
(575) discussed the species occurring in Ohio, and Funkhouser
(276) listed the species occurring in Connecticut.

The buffalo tree hopper, Stictocephala bubalus (F.), (=Ceresa),
is occasionally injurious to young trees such as ash and elm. Injury
results from two opposing slits cut in the bark by the female dur-
ing the act of oviposition. Eggs are placed in the slit and the
portion of the stem beyond it often aies. Adults are light green to
yellowish and about 9 mm. long. The pronotum is sharply ele-
vated, is widest at the tip, bears two sharply pointed horns which
extend at right angles to the body, and ends beyond the tip of the
abdomen in a narrowed acute process.

70

The three-cornered alfafa hopper, Spissistilus festinus (Say) oc-
casionally damages black locust seedlings in nurseries in the
South. Injury results from the feeding of the nymphs on the
stems, usually 1 to 2 inches above the base. Gall-like swellings or
calluses which develop just above the feeding punctures kill many
of the seedlings. Other seedlings are lost when they break at these
points. Stictocephala militaris (Gibson and 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 rec-
livata Fitch— common to abundant on basswood; T. decorata Ball
—fairly common on aspen; Vanduzea arquata (Say)—abundant
on black locust; Thelia bimaculata (F.)—abundant on black lo-
cust; Cyrtolobus disciodalis (Emmons)—common on red oak;
Carynota stupida Walker—sometimes abundant on _ yellow
birch; Playcotis vittata (F.)—abundant on oak; and the
two-marked treehopper, Hnchenopa binotata (Say)—common on
butternut.

FAMILY CERCOPIDAE
SPITTLEBUGS

Spittlebugs are represented in our fauna by 25 species, most of
which belong to the subfamily Aphrophorinae (686). 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 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 Ala-
bama. Its favorite host appears to be Scotch pine, but it is also
known to attack pitch, white, Virginia, jack, slash, loblolly, and
Japanese pines; Norway, white, and red spruces; and balsam fir
and hemlock. Trees of all ages and sizes are attacked. During out-
breaks, infested Scotch pines may be severely stunted or killed
(681). 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. 15).

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

The fungus, Entomophora aphrophora, and high temperatures
during the nymphal period occasionally cause heavy mortality of

71

COURTESY CONN. AGR. EXPT. STA.
FIGURE 15.—Adult and nymphal spittle masses of the
pine spittlebug, Aphrophora parallela.

the pine spittle bug. The opening up of closed stands and pruning
of dead and dying limbs to reduce the amount of material suitable
for oviposition are recommended as control practices.

The Saratoga spittlebug, Aphrophora saratogensis (Fitch), oc-
curs in southeastern Canada and eastern United States from
Maine to Minnesota, and south through the Appalachians to
Florida. In the nymphal stage it feeds on a wide variety of herbs
and shrubs. Sweet fern, brambles, prairie willow, and aspen
sprouts are preferred by fourth and fifth instar nymphs. The
adults feed on red, jack, white, Virginia, pitch, and loblolly pines;
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, hight brown to tan in color, with
a light, irregular stripe on the head and pronotum and with
oblique, wavy markings on the wing covers. The abdomen of the
fourth instar nymph is scarlet, edged with black. Fifth instar
nymphs are dark brown.

Adults are active from late June to late September. In the Lake
States, the majority appear in late July. They feed by inserting
their mouth parts into the cortex of new and old shoots of the
host, mostly on two-year-old internodes, and extracting sap. This
causes the formation of necrotic resin-filled pockets in the phloem
and xylem tissues. Damaged pines are characterized by the pres-
ence of reddish-brown “flags” of dead foliage, by numerous punc-
tures on the twigs covered with small drops of resin, and by light

72

i

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 (235).

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 dead twigs. The
winter is spent in the egg stage and the eggs hatch in the spring
just about the time red and jack pine buds begin to elongate.
Young nymphs craw] 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. 16.). To find these masses it
is frequently necessary to brush the litter away from the base of
the stem. The nymphs reach maturity in about 40 to 70 days, de-
pending on the weather. Then they climb to the upper parts of
their alternate hosts and transform to adults. 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 com-
plete by mid-July.

F-489365

FIGURE 16.—Spittle mass of

the Saratoga _ spittlebug,

Aphrophora saratogensis,
at base of sweet fern.

Nymphal populations are often greatly reduced by late spring
frosts in the Lake States. Hot, dry weather also kills many young
nymphs, especially in open plantations. Insect parasites and pre-
dators provide a certain amount of control, but are unable to
prevent outbreaks. Recommended preventive control measures in-
clude selection of sites for red or jack pine plantations that are
comparatively free of hosts favored by the nymphs, dense plant-
ing so as to secure early crown closure, and planting on good sites.
Methods of direct control are discussed by Wilson and Millers
(784) and Wilson and Kennedy (782).

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, wild
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 ob-
served feeding on the young twigs of Australian pine in Florida.
In heavily infested areas, the adults may be attracted to lights in
such large numbers they become a nuisance. The alder spittlebug,

73

C. obtusa (Say), a common species, feeds on various shrubs and
trees, including hickory, birch, and alder. The pecan spittlebug,
C. achatina Germ., 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 nigricollis
Fitch, has been recorded on pine. Omalophora salicis (DeG.) fre-
quently 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 of all of the order Homoptera occurring
in the United States. The adults are stocky, heavy-bodied insects
with large compound eyes and membranous 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
cicada, 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, red-
dish legs, and reddish wing veins. The life cycle is 18 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. 17 A),
is widely distributed in Eastern United States and it lays its eggs
in more than 70 species of trees and other plants. The most sus-
ceptible of the trees appear to be the oaks, hickory, honeylocust,
dogwood, apple, and peach; however, many others such as sweet-
gum, 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.

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 3 inches long (fig. 17 B).
When the eggs hatch, the nymphs fall to and enter the ground
where they feed on suitable roots. Usually they are found at
depths of 18 to 24 inches beneath the surface. When they become

74

: j te COURTESY TLL. NAT. HIST. SURV.
FIGURE 17.—The periodical cicada, Magicicada septendecim: A, Adults
on the bark of a tree; B, oviposition scars in branch of tree.

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, lit-
erally deafening in wooden 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, es-
pecially 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
in U.S. Department of Agriculture Leaflet No. 540 issued in 1966
2A):

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

75

FAMILY PSYLLIDAE
JUMPING PLANT LICE

Psyllids are very small, about 2 to 5 mm. long, and look very
much like miniature cicadas. There is also some resemblance to
winged aphids, but psyllids differ in having stouter legs, with the
hind pair fitted for 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.

Hackberry is attractive to many gall-forming species, all of
which belong to the genus Pachypsylla. Although hackberry
(Celtis) is cosmopolitan, Pachypsylla is known only in North
America. Of the five species of hackberry in the United States (C.
laevigata Willd., C. occidentalis L., C. pallida Torr., C. reticulata
Torr. and C. tennifolia Nutt.) psyllid galls are recorded for all
except C. pallida, a spiny bush occurring in the Texas, Arizona,
and New Mexico deserts.

Psyllids fall into two species groups, a) the leaf gall makers,
and b) the woody gall makers. Adults of the leaf feeding forms
emerge from galls in the fall; the wood feeding forms, in the
spring. Leaf gall makers are further subdivided into two groups,
blister galls and nipple galls.’

Pachypsylla celtidisvesicula Riley, the blistergall psyllid, forms
a small, monothalamous (one psyllid per gall) blister gall. It is
believed to occur throughout the range of Celtis occidentalis in
the United States and is the most abundant of the hackberry
psyllids. Fifty or more may infest a single leaf. Adults often be-
come a household nuisance when thousands accumulate on screens
in the fall prior to hibernating. Two undescribed species of
blistergall psyllids occur on C. Laevigata and C. reticulata.

Pachypsylla celtidisastericus Riley, the hackberry stargall, pro-
duces a gall which closely resembles the blistergall formed by the
blistergall psyllid, except it has star-like growth on the lower side
of the leaf. It can occur on all hackberry tree species except Celtis
occidentalis.

The hackerry-nipple-gall maker, Pachypsylla celtidismamma (Ri-
ley), is perhaps the best known hackberry psyllid. It produces light
green, nipple-shaped galls on the underside of Celtis occidentalis
leaves. It is monothalamous, although at times it may appear to be
polythalamous due to as many as seven galls of P. celtidisvesicula,
which may be incorporated into a single nipple-gall (537). Like
the blister-gall psyllid, adults cluster on screens during the fall.
The nipple-gall psyllid may disfigure leaves when infestations are
(ae (662). Methods of control are discussed by Thompson

713) \

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 in the
fifth nymphal instar in the gali (739).

“Riemann, J. G. 1961. A study of the hackberry gallmaker genus Pachyp-
sylla. (Unpublished thesis. Univ. Texas, 201 p., illus.)

76

Pachypsylla venusta (O.8.), the petiolegall psyllid, is the largest
of the hackberry psyllids, and it forms the largest galls. The galls
are polythalamous and usually contain six cells. Sometimes there
are as many as 30. These galls may persist on the trees for several
years and seriously damage the appearance of shade trees. Dis-
tribution of the species is mostly limited to below the 40th parallel.

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

Several species in the genus Psylla also feed on various trees,
shrubs, and ornamentals. Large quantities of honeydew are pro-
duced which drip down and cover their hosts. A black mold grow-
ing in this material often makes infested plants look unhealthier
than they are.

The boxwoed psyllid, Psylla buat (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 waxy ma-
terial. 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. Weiss and St.
George (752) discuss methods of control. Other species of Psylla
and their hosts include: P. annulata Fitch, paper birch and maple;
P. carpinicola Crawf., birch; P. floccosa Patch and P. galeaformis
Patch, alder; and P. trvimaculata Crawf., wild 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 which frequently coalesce,
causing the needles to die and drop prematurely. The persimmon
psylla, T. diospyrt (Ashm.), and T. magnoliae (Ashm.) feed on
persimmon and magnolia, respectively, in Florida.

Psyllids are attacked by a large number of hymenopterous and
dipterous parasites. Jensen (402) listed more than 150 species
from the United States alone.

FAMILY APHIDIDAE
APHIDS, PLANT LICE

The aphids or plant lice 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, have
fairly long antennae, and 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 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,

Aa

especially to shade and ornamental trees and young trees in plan-
tations. Honeydew dropping from heavily infested shade and
ornamental trees is often a nuisance as 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 proper functioning of their leaves. Numerous publica-
tions have been issued on the identity, distribution, and hosts of
aphids (296, 380, 583, 584, 585, 586).

10.

aL

78

Field Key to the Aphids, Based Principally on Host Plants

Causing galls or gall-like formations .__ 2
Not causing galls or gall-like formations _. Ace de Se 19
On contters::2ye4ernekd ee ee oe Apia dog ds oh NT ed aT, Oy ted
Onvhardwoodsi. = = ee ene ue Feet AliO

On balsam fir twigs, branches, and ees causing
“gouty” swellings at tips of twigs and stems, often
causing leaves to turn red and trees to die. Insects 14 to
114 mm. long; color dark brown to black, covered with
white flocculence o.oo... «cb bide Se i Adelges piceae

ON, SPEUCCs Sei td Belt sien Ba a eel etiah lence anes ee a

Causing only terminal galls c or gall-like formations

Causing galls other than terminal galls .

Terminal galls, compact, small .

Terminal galls, larger, composed of entire new growth
with needles thickened at base, or loose and on current
DRO Wil Menges ecw eectaleal eg ee tse eee eee we dxirke a part E Neste NOR iam

Terminal galls conelike, compact; on new growth. On
black, Engelmann, red, or Sitka spruce

Pineus pinifoliae

Terminal galls small, not conelike; on current year’s

growth. On spruce, mostly red and black
Adelges strobilobius

Terminal galls comprising entire new growth and causing

thickening of needles at base. On black and red spruce
Pineus floccus

Terminal gall a loose growth. On Engelmann, Norway,
white, black, and red spruce .... oa... Pineus similis

Causing galls on entire new growth of spruce host. On
blue, Engelmann, oriental, and Sitka spruce

Adelges cooleyt

Causing pineapple-shaped galls only at base of new

So CO O1}-

growth of spruce 7 | | fae 9
Galls with very short needles. On white and Norway

spruce | Adelges lariciatus
Galls with longer needles. ‘On white, red, and Norway

spruce .... Adelges abietis

On hickory or pecan, causing green, hollow, bullet-like
galls on twigs or leaf stems in June
Phylloxera spp.

On other hardwoods lel:
On elm fue
On other hosts 15

a:

12.

13.

14,

155

16.
gfe

18.

cD.
20.

21.

22.

23.

Producing spindle-shaped galls up to 1 inch long. On
slippery elm leaves. Insect 1 to 114 mm. long, reddish
Gobaishia ulmifusa
Galls not spindle-shaped. On various elms ..........................
Causing leaf curl or root swellings... Eriosoma
Causing cockscomb or pouchlike galls on upper surface
Olpleave ste Meta Aik) Secs Eh iitaale usty lak pence ee, «
Pale yellow, translucent, or head greenish-black and
abdomen and thorax dark green, covered with white
waxy powder. On leaves of slippery and wych elm,
producing cockscomb or pouchlike galls on upper
SUPT ACCR Rie etree tec et By SE OTN Colopha ulmisacculi
Olive green to dark olive brown or reddish-brown; 14 to
114 mm. long. On leaves, causing only cockscomb galls
ON: WOPCLRSUT LACEY ia an = eek aS Colopha ulmicola
On apple or serviceberry, causing leaf curl and leaf
swellings. About 2 mm. long; yellowish or brown to
reddish, usually with white or bluish flocculence
Eriosoma
On other hosts .... Wp eC ee Se ark. agile ie 5 ee ac
One pop laine =: ne ean eh Se Sees Ole ee eee ey |
Omngnwieclnave le tome te. eet he ee ee ae!
Causing convolute galls at tips of branches. Insects 1 to
4 mm. long; head and thorax black, abdomen pale;
oviparous female with head, thorax, and abdomen
goldens vellowrr oes | ois. ee ee Mordwilkoja
Occurring on leaves and petioles, causing galls at base of
leaf, juncture of leaf and petiole, or on petiole. Insects
1 to 3 mm. long; greenish or yellowish-green to
greenish-white, head and thorax usually dark; body
OL LEN DIUNIMOSCLL ent is eee le rise: Snell, Pemphigus
Causing conical galls on upper surface of leaves. Insects
1 to 2 mm. long, pale yellowish to greenish
Hormaphis
Causing spiny galls from buds on stems. Insects 1 to
2 mm. long, pale yellowish to greenish
Hamamelistes

QnecOmiueLrs os ee a eee
On=hardwoods: <2. 28... ft

Onepinews. he ee eS USE VRC L RAS FRc es ee ates
Grr ouhereconibens: «visi foe ea Ne ee ee
Small, dark insects covered mith white flocculence. Oe
curring on trunks and branches of Austrian, Scotch,
IGM bep DINCs eee Soret ace h ye ie canons Pineus strobi
Insects not covered with white flocculence, at most
pruinose. Occurring on twigs, branches or leaves _

Shining black with white powdery spots on sides and
white median dorsal line; legs black; body hairy, hairs
long and stiff. On branches and twigs of white pine

Cinara strobt

Noteassabove. 2.

Olive brown, body nents « sparse at cuvinabike. On leaves
of Scotch, ponderosa, shortleaf, and white pine

EFulachnus rileyt

13

14

29
21

23

Pale green with white pruinosity. On leaves and twigs
Mindarus abietinus
24. On larch 2.2204 ere ee ee ee ee
On other conifers) se 1 Se ee ee ee
25. Appearing as white woolly masses on the needles, as
dark individuals on the underside of twigs, or as
clusters of dark individuals at the base of the leaves
Adelges strobilobius
26. Nymphs dark green. On Douglas-fir, often causing ab-
normal toliace dropre..2- ee ee Adelges cooley
Noti as above:...... 8.20 ie ae ee a, ee
27. Cinnamon brown with 4 rows of black spots on abdo-
men; length about 3 mm.; many specimens covered
with fluffy white wax above ; body hairs long and fine.
On black spruce branches and twigs
Cinara abietis
Not as'above 202 ee ee etre ee ee eco
28. Brownish-black with a slight pruinosity, legs brown to
black, antennae pale; length about 5 mm. On branches,
twigs, stems, and roots of fir and Atlas cedar
Cinara curvipes
Pale green with pruinosity, or with white flocculent
covering when on balsam fir; length about 2 mm. On
fir and spruce twigs and leaves, causing curling and
roushenine Of-twits 202. Mindarus abietinus
29. Large species, about 6 mm. long; ash gray with triangu-
lar spots on pronotum; body covered with bluish-white
bloom:
On various hosts; common on hickory and sycamore
— Longistigma
Smaller species, usually 5 mm. or less in length:
Alder: Causing mottling or dropping of leaves.
Insects 114 to 2 mm. long; green or greenish-
yellow, abdomen uniformly colored or dusky-
spotted. 4282 ).8U- 38%. EGE eee ree Myzocallis
On underside of leaves or on stems. Insects 2 to 4
mm. long; brownish, covered with white flocculence
Prociphilus
Beech: On underside of leaves. Insects 2 to 3 mm.
long; bluish-white to greenish-yellow, covered with
white. 1LocculenCe =e ea ee Phyllaphis
On underside of branches or on trunk. Insects 2 to
4 mm. long; brownish, covered with white floccu-
VS acy Mi ot Site lt de ore ce apd ek eet gt .... Prociphilus
Birch: On ieins and leaves; 3 to 4 mm. long; whitish
to yellow, with or without black stripes on head
and thorax and dark bands on dorsum of abdomen
Calaphis
On twigs and leaves; 2 to 3 mm. long; light green,
with or without dark patch on abdomen; hind
tibiae pale with distal part and tarsi black or with
dark brown stripe ..... ...... Huceraphis
Boxelder: On leaves and twigs; 2 to 2% mm. long;
yellowish-green marked with brown on thorax and

80

25
26

27

28

abdomen or with head, antennae, thoracic lobes,
sternal plates, and cornicles black or yellowish to
apple green; antennae rather hairy
Periphyllus
Butternut: On leaves; 114 to 2 mm. long; pale yellow,
costal vein yellow or brown, or a brown band ex-
tending beyond stigma; antennae ringed with
DEOWiine teeth Ain eee Wer area Monellia
Elm: Causing mottling or dropping of leaves; 114 to
2 mm. long; green or greenish-yellow, abdomen
uniformly colored or spotted dusky
Myzocallis
Hickory:
Large species, about 6 mm. long; mostly on
branches and twigs; ash gray with triangular
spots on pronotum covered with bluish-white

GUN OSEUV getter. race Oey eee tt © ee certine Longistigma
Smaller species, 1144 to 2 mm. long; occurring
on leaves:

Causing mottling or dropping of leaves;
green or greenish-yellow, abdomen uni-
formly colored or spotted dusky
Myzocallis
Pale yellow, costal vein yellow or brown or
a brown band extending beyond stigma;
antennae ringed with brown ......Monellia
Linden: On underside of leaves; 2 to 3 mm. long;
yellow; antennae uniformly yellow or ringed with
EOWA LN RE A, ee Myzocallis
Maple:
Brownish, often covered with white flocculence:
Without white flocculence; egg-laying fe-
males with 7th and 8th abdominal seg-
ments prolonged into ovipositor
Drepanaphis
Usually with white flocculence; 4th anten-
nal segment about one-half or less as long
as third segment; length 2 to 4 mm.; al-
ways with white flocculence on underside
of leaves or on trunks .............. Prociphilus
Either yellowish, reddish, or shading to green:
On soft maple leaves and twigs; 3 to 4 mm.
long; dusky reddish to dark yellowish-
GRECH EY Oe ch, SOLE eS Se Pterocomma
On Norway and sugar maple, on underside
of leaves; 2 to 214 mm. long; yellowish-
green marked with brown on thorax and
abdomen, or with head, antennae, thor-
acic lobes, sternal plates, and cornicles
black or yellowish to apple green; anten-
mae xather jhainy $26. ee Periphyllus
On Norway, sycamore, or English maple,
on underside of leaves; 2144 to 3 mm.
long; yellow or reddish to green, with

81

er me ram RET

black markings or bands; legs and cor-
nicles yellow to dark orange
Drepanosiphum
On underside of leaves; 214 mm. long;
olive green or pale sordid green
Drepanaphis
Oak:

Causing mottling or dropping of leaves; 114 to
2 mm. long; green or greenish-yellow, abdo-
men uniformly colored or spotted dusky

Myzocallis

On leaves; 114 to 2 mm. long; pale yellow; costal
vein yellow or brown or brown band extend-
ing beyond stigma, antennae ringed with
brown i hee eee eee ee eee Monellia

On underside of leaves; 2 to 3 mm. long; yellow,
antennae uniformly yellow or ringed with
Drown ee eee ee ee Mayzocalits

Poplar:

On sprouts or tender growth; 2 to 214 mm. long;
yellowish or greenish, with brown mottling;
head: brownie. a eee Chaitophorus

On leaves and twigs; 3 to 4 mm. long; dusky
reddish to dark yellowish-green

Pterocomma

On leaves and twigs; about 5 mm. long; ash
gray due to white pruinosity over base color
of brown, abdomen marked with black spots;
large black tubercle on 4th abdominal segment

Lachnus
Sycamore:
Length about 6 mm.; body covered with bluish-
white\ PEUINOSIty | 27 ee Longistigma
Length 2 to 4 mm.; covered with white floccu-
ETI CEs ieee ae es coe opie eres enn Prociphilus

Walnut: On leaves; 114 to 2 mm. long; pale yellow
or brown, or brown band extending beyond
stigma; antennae ringed with brown

Monellia .
Willow:
Length 3 to 4 mm.; dusky reddish to dark yel-
lowish-green ............ _Pterocomma

Length about 5 mm. - ash. gray due to pruinosity
over base color of brown, abdomen marked
withdarkespotse. . 70. ee ee Lachnus

The giant bark aphid, Longistigma caryae (Harris), the largest
of all North American aphids, occurs throughout much of Eastern
United States and feeds on a wide variety of deciduous trees such
as elm, pecan, sycamore, oak, maple, basswood, birch, beech, wal-
nut, and willow. Adults are about 6 mm. long, and are covered
with a bluish-white bloom. Males are winged; egg-laying females
wingless. Eggs are laid on twigs, a single female laying many.
The aphids usually occur in clusters on the undersides of twigs

82

~*~” —"" _ Evie

and small limbs. Depending on locality, there may be several gen-
erations per year. Heavily infested twigs may be seriously injured
or killed.

The genus Cinara contains a number of conifer-infesting spe-
cies, only a few of which are mentioned here.

The white-pine aphid, Cinara strobi (Fitch), occurs from New
England to the Lake States and Carolinas and feeds on white
pine. Winged forms are almost 6 mm. long. The body is 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 females lay up to 5 or 6 eggs each, in
straight lines on white pine needles. Hatching occurs in the spring
and wingless females produce living young which live in colonies
up to 8 or 4 inches long clustered around a branch or the leader.
Several generations later, winged females are produced which
migrate and also produce living young. Toward fall, winged males
and females mate, and a new crop of overwintering eggs are laid.
Young trees or individual branches of large trees may be killed
by heavy infestations or their growth may be seriously reduced.

Cinara sabinae (Gill. & Palm.), a small, reddish-brown species
about 3 mm. long and covered with a white powdery secretion,
feeds on red cedar in 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 honey-
dew on the foliage. The related species, C. canadensis Hottes &
Bradley and C. juniperivora (Wilson), also feed on red cedar.

Cinara pint (Linn.) is a large reddish-brown species with
numerous dark specks and a pair of large spots behind the cor-
nicles. It has been recorded feeding on young Scotch, shortleaf,
and Virginia pines and seriously injuring them. Additional pine-
infesting species of Cinara include: C. carolina Tissot—loblolly,
shortleaf, slash, longleaf, pond, and Virginia pines in the South;
C. taedae Tissot—loblolly and Virginia pines; C. watsoni Tissot—
loblolly, shortleaf, slash, spruce, and pond pines in Florida and
red, pitch, Scotch, and Virginia pines in Pennsylvania; and C.
atlantica (Wilson) —various pines in the Southeast.

Other pine-infesting aphids include the following: Hulachnus
rileyt Williams—Scotch, red, pitch, Virginia, shortleaf, and white
pines; FE. -agilis (Kaltenbach)—red and Scotch pines; Essigella
pint Wilson—loblolly, slash, and pond pines in Florida (has also
been recorded from Maryland and Pennsylvania) ; Schizolachnus
piniradiatae (Davidson)—jack and red pines in the Eastern
States; S. lanosus Hottes—loblolly pine in Maryland; and Proci-
philus bumeliae (Schrank) —roots of white pines.

Several other aphids have also been recorded from other coni-
fers. For example, Cinara abietis (Fitch) —various firs; C. twja-
filina (Del G.)—arborvitae, cypress, and other species (heavy in-
festations have been reported in nurseries in Florida and Dela-
ware); and the balsam twig aphid, Mindarus abietinus Koch—
wide variety of hosts including balsam fir, Siberian fir, Alpine fir,
white spruce, juniper, and Fraser fir (damaged needles curl, and
the bark of heavily infested twigs becomes roughened).

The boxelder aphid, Periphyllus negundinis (Thomas), feeds on
the leaves and twigs of boxelder wherever it grows in the United

83

States and Canada. Its body is usually yellowish-green with
brownish marks on the thorax and abdomen. Walks, benches, or
cars parked beneath infested trees are often badly soiled. The
related species, the Norway-maple aphid, P. lyropictus (Kessler),
often completely defoliates Norway and sugar maple trees. Honey-
dew dripping from infested leaves is a nuisance. P. americanus
(Baker), Drepanaphis acerifoliae (Thomas), D. carolinensis
Smith, D. nigricans Smith, D. sabrinae Miller, and Drepanosi-
phum platanoides (Schrank) are also found on maples.

Numerous species of Myzocallis such as M. bella (Walsh), M.
alhambra Davidson, M. discolor (Fitch), and M. melanocera
Boudreaux and Tissot feed on various oaks. Other common mem-
bers of the genus and their hosts include M. alnifoliae (Fitch) —
alder; M. tiliae (L.)—native and introduced lindens; the crape-
myrile aphid, M. kahawaluokalani Kirkaldy—crapemyrtle; the
black pecan aphid, M. caryaefoliae (Davis)—hickory and pecan;
and the elm leaf aphid, M. ulmifolii (Mon.)—elm.

Many other species of free-living aphids also occur commonly
on various deciduous trees. These include Monellia caryella
(Fitch), M. costalis (Fitch), M. microsetosa Richards and Movnel-
liopsis nigropunctata (Granovsky)—hickories; Chaitophorus ste-
vensis Sanborn, C. populicola (Thomas) and Pterocomma populi-
foliae (Fitch)—poplars; P. smithae (Monell)—poplars and
willows; P. salicis (L.)—willow; Macrosiphum lirtodendri (Mo-
nell) —yellow-poplar; Aphis craccivora Koch—black locust seed-
lings; Calaphis betulaecolens (Fitech)—birch (often in large
numbers); C. betulella (Walsh)—bireh and beech; Huceraphis
betulae (Koch)—various birches, especially yellow birch; LF.
lineata Baker—gray birch; and E. mucidas (Fitch)—river birch.

Three species of woolly aphids are commonly found on elms in
Eastern United States. The woolly apple aphid, Hriosoma lani-
gerum (Hausm.), feeds on new terminal leaves, causing them
to curl or appear in the form of rosettes (It also attacks
apple, pear, hawthorn, and mountain-ash. Damage to elm is not
especially severe.). The presence of large numbers of distorted
leaves on shade trees is unsightly. The woolly elm aphid, FE’. ameri-
canum (Riley) has elm as a primary host and shadbush 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. Then, members of the
second generation fly to shadbush and attack its roots. The
Summer is spent here. During the fall, a winged generation de-
velops which flies back to elm where it lays a crop of over-
wintering eggs. #. rileyi Thomas, the woolly elm bark aphid, at-
tacks American and slippery elm. 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, EF.
crataegi (Oest.). It occurs in dense colonies on 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, Prociphilus imbricator (Fitch)
feeds on the trunk and undersides of leaves and branches of beech
from New England to Georgia and Illinois. Its body is covered

84

with a white, cottony substance that strings out in fairly long
threads and forms a tuft at the rear end. Heavy infestations oc-
casionally develop on forest-grown trees. The woolly alder aphid,
P. tessellatus (Fitch) is often abundant on alder and silver maple.
P. fraxinifolu Riley feeds on ash and P. corrugatans (Sirrine) on
shadbush.

Certain species of.aphids produce galls by their feeding. The
elm coxcomb-gall 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. It feeds on leaves, causing the for-
mation 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
is spent in the egg stage on elm, and 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 be-
come unattractive. The related species, C. ulmisaccult (Patch)
produces pedunculated, bladder-like galls up to 25 mm. long on the
upper surfaces of elm 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.
18) up to 5 inches in diameter. These galls may occur singly or in
clusters of three to five galls each. The winter is usually spent in
the egg stage in old galls or in nearby bark crevices (390). Other
gall producing aphids and their hosts include Hormaphis hama-
melidis (Fitch)—conical galls on the upper surfaces of witch-
hazel leaves; Hamamelistes spinosus (Shiner)—causes galls to
form on the stem buds of witch-hazel; and Gobaishia ulmifusa
(Walsh and Riley)—spindle-shaped, saclike galls on the leaves of
slippery elm.

F-506747
FIGURE 18.—Galls of the poplar vagabond aphid, Mordwilkoja vagabunda,
on poplar.

85

FAMILY PHYLLOXERIDAE

Members of this family resemble those of the family Aphididae
but differ in that eggs are laid by both sexually perfect and im-
perfect females. The majority of important eastern species feeds
on the needles, twigs, limbs, and trunks or in galls on coniferous
trees.

The eastern spruce gall aphid, Adelges abietis (L.), an introduced
species, occurs in southeastern Canada and the Northeastern and
Lake States. Its preferred hosts appear to be Norway and white
spruce, but it is also found occasionally on red and Colorado blue
spruce. The winter is spent as small nymphs under coverings of
waxy threads at the bases of buds on the undersides of twigs
(765). The nymphs molt in the spring and become stem mothers
which lay eggs on the needles. Hatching occurs in about 2 weeks
and the young nymphs craw! to the bases of expanding buds. Here
they feed, causing the formation of pineapple-shaped galls (597)
in which they live and continue their development (fig. 19). Dur-
ing late August to October the galls open and the full- -grown
nymphs or “pupae” emerge. These transform to winged adults in
a couple of days and, although they can fly, many-remain on the
tree. The females insert their mouth parts through the bark,
deposit upwards to 60 eggs each and then die. These eggs hatch
within 16 days and the young nymphs immediately crawl to over-
wintering sites. There is one generation per year.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 19.—Galls of the east-

ern spruce gall aphid, Adel-
ges abietis, on spruce.

86

The eastern spruce gall aphid is a serious pest in nurseries and
Christmas tree plantations and on park and other ornamental trees.
Its damage is limited primarily to a reduction in the esthetic value
of the trees.

The Cooley spruce gall aphid, Adelges cooleyi (Gill.), occurs
from coast to coast in northern United States and throughout the
range of white spruce in Canada. Its primary hosts are recorded
as white, Colorado blue, Sitka, Engelmann, and big-cone spruce.
It also has an alternate host, Douglas-fir.

The Cooley spruce gall aphid winters as immature stem-mothers
under bark scales near the terminals of twigs of spruce. The
female becomes mature in early spring and deposits a large num-
ber of eggs under a mass of white, cottony wax. The eggs hatch
in about a week and the nymphs settle down to feed at the bases
of young needles. Galls (fig. 20) begin to form immediately and
develop rapidly, enclosing the nymphs. Young galls are green or
purple in color; older ones are 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, allow-
ing the nymphs to escape and crawl to the needles. Here they

F 519584

FIGURE 20.—Galls of Cooley

spruce gall aphid, Adelges
cooleyi, on blue spruce.

transform to winged adults and fly to Douglas-fir, if present.
Eventually, a winged generation is produced on this host, and it
returns to spruce (179). Where spruce and Douglas-fir do not
occur close enough together for the aphid to move back and forth
from one to the other, generations may be produced continuously
on either species.

The Cooley spruce gall aphid is not usually considered an im-
portant 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 plantations are
often seriously damaged by excessive numbers of unsightly galls.

The balsam woolly aphid, Adelges piceae (Ratz.), an introduced
species first recorded in North America from Brunswick, Maine,
in 1908, now occurs throughout the Maritime Provinces, Canada,
and the Northeastern States (except the northern parts of Maine
and New Brunswick Province). Infestations also occur over much
of Newfoundland, in the Gaspé Peninsula, in the southern Appala-
chians, and in the Pacific Northwest. Its hosts are balsam and
Fraser firs in eastern America. Full-grown aphids are roughly
spherical in shape, less than a millimeter long, and almost in-
visible to the naked eye. Because of a covering of white wax
threads, however, they appear as dots of white “wool” (fig. 21).

Eggs are deposited in late spring and early summer, each egg
being attached to the bark behind the female’s body by a silken
thread. Newly hatched larvae crawl rapidly over the bark until
they find suitable feeding places. When these are found, they insert
their feeding 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 short period of rest they develop into
the second generation. Adults of this generation deposit eggs
during midsummer. Hatching soon occurs and all stages are found
until late fall. The winter is spent as a first-instar larva. 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 the Northeast.
There are two generations per year in the Northeast; in the

SAFER he
t fae > ae ee ‘
a+ = fue"

CF &s

othe
“a “0
we

‘

bail

Ye

Tey?
a”

-

ou

F 519571

FIGURE 21.—Infestation of the

balsam woolly aphid, Adel-

ges piceae, on trunk of
Fraser fir.

southern Appalachians two and occasionally three generations are
produced (24, 5).

The balsam woolly aphid 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 which 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. Branchlets
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 “compres-
sion’ wood, is produced in greatest amount on moderately infested
fast-growing trees. Trees suffering from heavy stem attack may
be killed in 3 or 4 years (fig. 22).

The balsam wooly aphid 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 aphid have
been found in North America. Quite a large number of predators
have been recorded, however, but they have proved incapable of
preventing outbreaks. A few species of introduced predators, such
as the derodontid beetle, Laricobius erichsonti, offer promise of
greater benefits in control in the future (7, 27).

Outlying infestations of the balsam wooly aphid may be
controlled by clear cutting infested stands, especially if this is fol-
lowed by burning. The spread of infestations may also 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 re-
ducing infestations in New Brunswick (28).

Adelges strobilobius (Kalt.), an introduced species, occurs in
southern Canada and south and west in the Eastern States to
Washington, D. C. and the Lake States. Its hosts are recorded as
larch and spruce, principally red and black. Infestations on larch
appear as white wooly masses on the needles and as clusters of
aphids at the bases of needles. Infestations on spruce may be
recognized by the presence of small galls at the tips of new
growth. A related species, A. lariciatus (Patch), occurs on larch
and Norway spruce. It produces galls similar to those produced
by the eastern spruce gall aphid. Cumming (180) discussed its
life history. 4

The pine bark aphid Pineus strobi (Htg.), an introduced species,
occurs over most of the United States wherever white, Scotch,
and Austrian pines grow. It is small, dark, and covered with floc-
culent 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. 23) or at the bases of needles on
twigs, or on buds. The trunks of heavily infested trees often ap-
pear as if whitewashed.

89

F-519567
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ooly aphid, Adelges
EXPT
of

pine bark aphid, Pineus strobi,

Infestation
on the trunk of pine.

the balsam w
COURTESY CONN. AGR.
Don

piceae.

FIGURE 22.—Fr

FIGURE

poopy s-nsamataens ips sels gin eae

Sy Knee +
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ee Pa re / OCR AT NH if é Va ae ES Deh 4
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90

a

Eggs are laid in the spring by overwintering females and the
eggs hatch into both winged and wingless females. The wingless
forms remain on the pine host and reproduce repeatedly. Some of
the winged ones may fly to spruce where they settle on the needles,
lay eggs, and die. Nymphs hatching from these eggs also soon die.
Five generations per year have been recorded as far north as the
Lake States (619).

Trees in parks and recreational areas, ornamentals and small
nursery stock may be seriously damaged by the pine bark aphid.
The needles turn yellow, and small trees may be stunted or killed.
Large forest trees are usually not seriously injured.

The pine leaf chermid, Pineuws pinifoliae (Fitch), occurs in both
Eastern and Western United States. Its range in the East coin-
cides with that of its primary hosts, red and black spruce, where
they grow close to its alternate host, white pine. The adult is
scalelike, about 1.5 mm. in diameter, and bears a fringe of white
hairs.

The life cycle takes two 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.®

Infestations on spruce result in ‘the production of terminal com-
pact galls which have the appearance of true cones and contain a
single aphid in each chamber. These galls are of minor impor-
tance except on ornamentals where they may be undesirable.
Heavily infested white pines, especially young pines in planta-
tions, may be severely injured. Needles turn yellow; growth is
reduced; deformities are produced; and occasionally the trees are
killed. Several outbreaks have been recorded since 1900, some of
which spread over large areas before subsiding.

Pineus floccus (Patch) feeds on spruce and pine, spending 1
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 chermid. Heavy infestations on spruce may kill the
tips of branches, or they may cause an over production of laterals
which lead to bushy, deformed trees. Damage to pine is usually
not serious.

Pineus similis Gillette produces terminal cone-shaped galls on
Norway, white, red, black, Colorado blue, and Engelmann spruces.
The galls are shorter and thicker than those produced by the
Cooley spruce gall aphid and the chambers are intercommunicat-
ing. 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.

The genus Phylloxera contains several species which 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, globe-
like structures up to 18 mm. in diameter.

® Lowe, J. H. 1965. Biology and dispersal of Pineus pinifoliae (F.). (Un-
pub. PhD. thesis, Yale Univ., 104 p. illus.)

91

Phylloxera caryaecaulis (Fitch), the hickory gall aphid, 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 aphids vacate them, they
turn brown or black. The pecan phylloxera, P. devastatrix Per-
gande, and the pecan leaf phylloxera, P. notabilis Pergande, pro-
duce galls on pecan. P. rileyi Riley is found on white and post
oaks, and P. nyssae Pergande infests blackgum.

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 white powdery wax, are whitish
in appearance. The larvae are very small and scalelike and 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, Tetraleurodes mori (Quaint.) feeds on
mulberry, dogwood, azalea, hackberry, holly, mountain-laurel
basswood, maple, and sycamore. The larvae are less than 1 mm.
long, jet black, and ringed around with a white fringe. Adults are
active from June to September. The rhododendron whitefly, Dia-
leurodes chittendeni Laing, an introduced species, feeds on
rhododendron. The adult is pale yellow; the larvae and pupae
greenish-yellow. Infested leaves have a yellow, mottled appear-
ance, and their margins curl. The azalea whitefly, Pealiws azaleae
(Baker & Moles), feeds on azalea as far north as Rhode Island
and Ohio, and sometimes causes severe defoliation. Alewrochiton
forbesi (Ashm.), feeds on maple, and the citrus whitefly, Dialewr-
odes citri (Ashm.), feeds on chinaberry and on crapemyrtle in
Florida.

SUPERFAMILY COCCOIDEA
(SCALE INSECTS)

Seale insects are among the most destructive agents of shade
trees and ornamentals. At times they may also cause serious dam-
age to forest growth. Injury results either from the withdrawal
of sap or the production of galls while feeding.

Male scale insects are usually winged but the females are en-
tirely wingless. Adult females also either have no appendages, or
they are atrophied. As a result, the body is either scalelike or gall-
like. It is also covered with wax, either in the form of powder,
tufts, or plates or as a thin layer covering the insect. Females usu-
ally appear formless because of the difficulty of separating their
bodies into head, thorax, and abdomen.

Some scale insects are highly specific in their selection of hosts;
others feed on a wide variety of hosts. Because of their habit of
feeding on many different parts of plants, many species have been

92

widely distributed by shipping nursery stock or cuttings (500,
ID)

Field Key to the Genera of Scale Insects Based Principally
on Host Plants

1. Restricted feeders on specific host plants
General feeders found on a variety of plant hosts
2. On conifers ... ad Se cngte
On hardwoods (see appropriate host).
Beech branches and trunks; mature female oblong,
1 mm. or less, yellow, body covered with white,
WO Oly aRWia Xie. a0 we | ‘Cryptococcus
Celestrus or euonymus leaves and stems, female ob-
long, broadened posteriorly, dirty gray to nearly
black, small white male scales usually abundant
and conspicuous in clusters on leaves
Unaspis euonymer
Elm bark, branches and stems; mature female oval,
about 2 mm., red-brown body with white waxy
fringe about margin _ . _..Gossyparia
Horse chestnut twigs and branches, mature female
oblong, 1 to 2 mm., dark gray to black with lighter
border... Aspitiotus, in part
Magnolia twigs and branches, mature female round
or slightly oblong, about 1 cm. in diameter, rich
dark brown, very convex .. 3 Neolecanium
Maple:
On leaves. Mature female 4 mm. or more:
Dark purplish, large white egg sac with 2
longitudinal grooves and several transverse
KId@eS = | Pulvinaria
Yellow, body covered with large white cot-
tony mass of wax thread. On sugar maple
Phenacoccus
On stems and branches (Southern States).
Mature female round to oval, about 2 mm.
gray, dark nipple concentrically placed
Chrysomphalus

Wwe bo

Oak:

Mature female gall or berrylike in shape, 3 to 4
mm. in diameter, brown with mottling. On
twigs and sometimes on leaves Kermes

Mature females nearly round, 1 to 1!4 mm. in
diameter, reddish-brown, body covered with
translucent greenish wax “‘test,’’ causes small
craterlike or pitlike galls on twigs, branches,
and stems. Various species of white oak group

Asterolecanium

Pecan branches, stems and fruit in Southern States,
not producing red coloration under infested bark;
mature female round to oval, about 2 mm., gray,
nipple concentrically placed Chrysomphalus
Yellow tulip twigs and branches, sometimes on mag-
nolia and poplar; mature female hemispherical, 5

93

to 8 mm. in diameter, dark brown
Toumeyella, in part
3. On pines:
Hard pines at base of needles on new growth during
summer; mature female oval, woolly, wax covering
1 cm. or more, white woolly covering
Pseudophilippia
Scotch, jack, lodgepole, and other hard pines on twigs
and branches; mature females oval, 5 to 7 mm.,
reddish-brown, dorsal surface very convex and
pitted: or irregular) = Toumeyella, in part
Hard pines on new shoots and under bark scales, ma-
ture female long oval, 2 to 4 mm., dark brown to
black, produces long, flat white egg sac under bark
scales of branches and trunk; immature instars
causes small pit galls in epidermis of new growth,
new growth dying in July, August, and September
Matsucoccus
Pine leaves, various species; mature female pear-
shaped, 114 to 3 mm., dirty white to dirty gray
Phenacaspis
Pine, various species, and Douglas-fir leaves; mature
female oval, 1 to 2 mm., dark gray, concentric nip-
ple orange... SNe abet SA ears ae Aspidiotus, in part
On other conifers:
Arborvitae twigs; mature female round to oval,
3 to 4 mm., reddish or reddish-brown, some-
times mottled; dorsal surface very convex
Lecanium, in part
Hemlock leaves and twigs; mature female oblong,
1 to 2 mm., dark gray to black with lighter
border ......... _..... Aspidiotus, in part
Juniper leaves and twigs; mature female nearly
circular, about 2 mm. in diameter, light gray
to white, central or concentric nipple yellow
Diaspis
Spruce, various species at base of buds on new
growth; mature female round, 3 to 4 mm. in
diameter, light brown, dorsal surface very

CONVEX. - eee late ue! SP lysOnenmes
4, Body of adult female covered by a “scale” made up of
secreted matter, plus the cast eee of the two larval

stages = iach ety oe mea
Body of adult female otherwise: naked, covered with
waxy or woolly secretion, enclosed in a secreted shell,
never covered by a scale as described above
5. Scale of adult female elongated, narrowed anteriorly with
the larval cast-skins terminal, broadened posteriorly
Seale of adult female circular to oval with exuviae cen-
tral or subcentral . Aspidiotus, in part
6. Scale of adult female white or dirty white
Chionaspsis, in part
Seale of adult female dark, usually dark brown
Lepidosaphes

94

7. Body small, not exceeding 2 to 3 mm. in diameter, en-
closed in a thin transparent shell with a fringe of se-

cretion around margin ee ae Asterolecanium, in part
Body not enclosed in a shell, mostly considerably larger,
ASTRNTTAR OV SIN OVC geo cetera) oe Teen oe aioe 1S ere ete ee Smee 8

8. Adult female at maturity with body remaining soft,
fringed at hind end with a large ovisac showing con-
SICUOUS ILC Ss eee eee eee are Icerya

Body at maturity becoming hard and eventually brittle ;
ovisac, if present, loosely formed, not fluted . 7 9
9. Witha short, stout posterior ovisac . _.... Pulvinaria
Body naked ‘at maturity, eggs deposited beneath it
Lecanium

FAMILY MARGARODIDAE
MARGARODID SCALES

Margarodid scales occur throughout most of the earth and vary
greatly in their habits. Some species feed in exposed positions on
their hosts; others beneath the bark or in gall-like pits on the
leaves, branches, trunks, or roots. Some species have only one or
a few hosts; others have many. Eggs may be deposited in an in-
ternal pouch or marsupium; in a posterior ovisac; within a gall-
like pit or heavy-walled tent in which the female develops; or in
loose cottony materials beneath the female’s body. Morrison (544)
published a classification of the family.

The red-pine scale, Matsucoccus resinosae Bean and Godwin,
probably an introduced species, was first recorded in North
America at Easton, Connecticut, in 1946. Since then it has spread
slowly and now also occurs on Long Island, in southeastern New
York, and in northern New Jersey. Red pine is its only known
native host. Other recorded hosts are three species of imported
ornamental pines: Chinese, Japanese red, and Japanese black
(322).

Eggs of the summer generation are laid in May in bark crevices
or under bark scales and hatch by the first of June. First-stage
larvae move around for a brief period and then settle down under
bark scales until they become full-grown (fig. 24). Adults emerge
in early August and lay eggs in late August or early September.
Hatching occurs soon thereafter. The winter is apparently spent
as first-instar larvae. Growth is resumed in the spring, and adults
emerge by May (46).

The red-pine scale is one of the most destructive insect pests of
red pine in the Northeast. Thousands of trees ranging in size
from nursery stock to mature trees have been killed, and many
more have been severely injured (44). The foliage of infested
trees turns olive green, then yellow, and, finally, brick red when
the trees die. The bark on the branches and trunks of heavily in-
fested trees appears swollen and cracked and there is a patch of
dead tissue beneath each feeding scale.

Several species of predators attack the scale but none have been
effective in control. Low winter temperatures of —10° F. are
almost entirely fatal to the species. This may prevent its spread

95

COURTESY CONN. AGR. EXPT. STA.

FIGURE 24.—Masses of male cocoons

of the red-pine scale, Matsucoccus

eee on lower side of branch
axil.

much farther northward than its present range. The cutting and
sae of infested trees is effective in reducing the rate of local
spread.

Matsucoccus gallicola Morrison, the pine twig gall scale, is also
an important tree-infesting species. Its hosts include pitch, short-
leaf, table mountain, Virginia, ponderosa, loblolly, and spruce
pines. It has been recorded from New England to Florida, and
west to Ohio and Missouri. Trees of all sizes, from three-year-old
seedlings to mature specimens, are attacked.

Mature females are generally much flattened and about 2 to 5
mm. long. They deposit their eggs under bark scales on the larger
branches and trunks. Heavy infestations may be quite injurious,
especially to young trees. Damage appears to have been most
severe to pitch pine in the Northeast (579).

Matsucoccus alabamae Morrison has been observed feeding in
cracks and crevices of heavy bark on pines in Alabama; and M.
matsumurae (Kuw.) infests Virginia and pitch pines on Long
Island.

Xylococculus betulae (Perg.) occurs in eastern Canada and
from New England to Virginia. Its hosts are paper and yellow
birches and beech. Adults are orange-red, about 4 mm. long, and
covered with white wax. Females live and deposit eggs in cells in
the bark. Young larvae crawl from these cells into lenticels and
roughened spots on the bark to feed. Here, they lose their legs
and produce a mass of wax around themselves, forming pearllike
cells (645). Honeydew is excreted through hairlike tubes up to 2
inches long. Large necrotic areas develop in infested areas on

96

birch. Damage to beech consists of roughened or swollen spots of
bark up to 2 inches in diameter. These spots dry out as they age,
and additional cracks form around them. Damaged spots may be
found over the entire trunk, but they usually occur in narrow
longitudinal strips, starting at old branch stubs.

FAMILY KERMOCOCCIDAE

Kermes pubescens Bogue is often a serious pest of oaks in the
eastern half of the United States, especially in the Midwest (476).
Adult females are light brown, almost spherical, and about 2.5
mm. in diameter. Eggs are laid on new growth from late June to
late July. Newly-hatched larvae migrate to the trunk and larger
limbs where they hide, in cracks or beneath loose bark. They
remain here without much growth until the following spring, at
which time they craw! to the new growth and resume feeding.
Heavy infestations on petioles and midveins may cause the distor-
tion or death of leaves and twigs. On heavily infested trees, the
flagging of terminals may be so severe by late July that the
beauty of shade and ornamental trees is greatly reduced. K. galli-
formis Riley, K. kingw Ckll., K. pettiti Ehrh., and K. trinotatus
Bogue are also fairly common on oaks in the Eastern States but
they are seldom injurious (fig. 25).

FAMILY ERIOCOCCIDAE

The beech scale, Cryptococcus fagi (Baer.), an introduced
species, was first recorded in North America in Nova Scotia
around 1890 and in the United States at Boston, Massachusetts,
in 1929. It now occurs throughout the beech-growing areas of
New England, eastern New York, Pennsylvania and New Jersey,
and the Maritime Provinces of Canada. Its hosts are American
and European beech and their varieties. Female scales (no males
have been found) are circular-shaped, 0.5 to 1 mm. long, and.
covered with a woollike wax.

Young larvae overwinter and their development is completed in
the spring. Egg laying begins in early June or later, depending on
location, and continues into July. Hatching begins by August 1 in
some areas. The young larvae crawl over the bark until a suitable
feeding site is found. There they insert their stylets, become
quiescent, excrete a white, wooly wax and eventually spend the
winter. There is one generation per year.

2

z 5 8
7 at
mt! : La
SEL 4
“tea a
aay i i
i a
+o x
<

Bae

F-519574

FIGURE 25.—Female scales of

Kermes sp. on white oak
twigs and leaves.

The first sign of infestation is the appearance of isolated,
minute, white wooly 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. 26) and lower sides of branches may be
completely whitened.

F-502240

FIGURE 26.—Infestation of the beech scale
Cryptococcus fagi, on the trunk of beech.

Light infestations are not particularly injurious but when they
increase to an intensity of about 100 scales per square inch the
outer layer of bark is killed and turns brown. Depressions or pits
form in bark tissues around these wounds and are frequently
numerous on young trees. The death and shrinkage of groups of
cells within feeding areas also causes ruptures in the bark. How-
ever, this usually does not happen until 2 to 5 years of infestation.
A fungus, probably Nectria coccinea var. faginata, also probably
introduced, gains entry through these ruptures (223). It pene-
trates to the cambium and sapwood, killing tissues and interfering
with the conduction and storage processes of the tree. Red fruit-
ing bodies produced by the fungus become so abundant on
severely infected trees that large areas on the trunk turn red.
This leads to death of irregular-shaped areas of bark. Individual
fungal lesions also coalesce and girdle the trunk, leading to crown
deterioration and finally, the death of the tree. The fungus is
entirely dependent on the scale for its incidence and spread.

Enormous quantities of beech have been killed and the only
known way to prevent such losses is through the cutting and re-
moval of infested trees (646). This is true despite the fact that
some degree of natural control is provided by the twice-stabbed
lady beetle and that winter temperatures of —35° F. are highly
effective in the control of infestations exposed above the snow
line.

The European elm scale, Gossyparia spuria (Mod.), an intro-
duced species, occurs over most of southern Canada and the
United States, wherever its hosts, native or European elms, grow.
The adult female is oval and olive green to reddish-brown, has a
white, waxy fringe along the body margin, and is about 114 to
114 mm. long. The males, some of which are winged, are much
smaller. Trees of all ages are infested.

The winter is spent in bark crevices on the trunk and large
branches. Eggs are laid from spring to midsummer, and newly-

98

hatched larvae are quite active. Some crawl to the undersurface
of leaves where they settle down and feed; others settle down on
twigs and branches. During late fall those on the leaves migrate
back to the bark and settle in crevices where they spend the
winter. There is only one generation per year.

The European elm scale is often very injurious, especially to
young or transplanted elms. Heavy losses have been reported from
many communities in the Midwest. Sooty mold developing in
honeydew secreted by the scale also reduces the beauty of surviv-
ing trees. Thompson (712) discussed methods of control.

FAMILY PSEUDOCOCCIDAE
MEALYBUGS

Members of this family are known as mealybugs because of the
mealy or waxy secretions that cover their bodies. Females are
elongate-oval and have segmented bodies and well developed legs.
Some species lay eggs; others give birth to living young. Mc-
Kenzie (490) discussed the taxonomy and biology of the group.

Phenacoccus acericola King, the maple phenacoccus, an intro-
duced species, occurs throughout the Northeastern, Central, and
Lake States. Its preferred host appears to be sugar maple, but it
has also been found on hornbeam, basswood, and horse chestnut.
Adults are about 5 mm. long and are covered with masses of
cottony wax. Evidence of infestation is the presence of cottony
masses on the undersides of leaves during the summer. The winter
is spent as a partly grown nymph in a hibernaculum beneath a
flake of bark. There are three generations per year. This species
is usually not very important, although it may damage shade
trees. Other tree-infesting species in the genus include Peliococcus
serratus Ferris which feeds on birch in the Northeastern States
and P. dearnessi King which occurs on hawthorn in the Lake
States.

Dysmicoccus cuspidatae (Rau.) attacks all species of Taxus in
the Northeast and Midwest. Infestations have also been recorded
on basswood, cedar, maple, and rhododendron. Females are 9 to
10 mm. long and 5 mm. wide and are covered with white wax
through which reddish body fluids show in longitudinal lines. The
winter is spent in the first nymphal instar in bark crevices or
under waxy secretions left by previous year adults. There appears
to be only one generation per year (in New York State). The
branches and trunks of heavily infested trees are often completely
covered with these insects and their white, waxy secretions. Con-
tinuous heavy infestations stunt the growth or kill Tazus in nur-
series and ornamental plantings.

An infestation of Dysmicoccus obesus (Lob.) was reported in
a 15-year-old ornamental planting of loblolly pine in Delaware.
Infested trees were spindly, short, and the foliage was sparse and
off-color. Conspicuous white powdery deposits occurred in cracks
and crevices of the bark on the lower four feet of the trunks of
infested trees (534).

Spilococcus juniperi (Ehrh.) [=Pseudococcus juniperi Ehrh.]
is a serious pest of native red cedars in ornamental plantings in

99

Kansas, Indiana, and Oklahoma. The lower and inner branches are
usually attacked first. The foliage of heavily infested trees turns
brown and drops. Entire trees are sometimes defoliated. The long-
tailed mealybug, Pseudococcus adonidum (L.), a widely distributed
species, feeds on many ornamental plants. Females are oval 2.4
to 8 mm. long, grayish to light yellow, covered with a fine, pow-
dery, waxlike whitish secretion, and bear two unusually long
fringe-like filaments at the tail end. Oracella acuta (Lob.) feeds
on the tips of loblolly pine terminals in Florida. Infestations have
also been observed on shortleaf pine in Pennsylvania.

The grape mealybug, Pseudococcus maritimus (Ehrh.), a
widely distributed species, feeds on all species of Taxus in Ohio;
also on grape, euonymus, and sycamore. The first winter is spent
as a first-instar nymph mostly near the main stem in the interior
of the plant. The nymph may be found in a mass of white waxy
threads produced by the female or beneath webbing, bark scales,
and other debris webbed together. During early spring the young
nymphs migrate to the twigs to feed. Adults emerge by mid-June
and the female deposits her eggs in masses of white fibrous mate-
rial. When the eggs hatch, the young nymphs migrate to the
foliage to feed. There are two generations a year. Damage to
Taxus in nurseries may be serious enough to warrant control
(562).

The Comstock mealybug, Pseudococcus comstocki (Kuwana), an
introduced species, occurs throughout most of the Eastern States
and feeds on a wide variety of trees such as apple, peach, catalpa,
boxwood, holly, horse chestnut, maple, mulberry, osage orange,
Taxus, and poplar. Full-grown females are 2 to 5 mm. long, and
covered with a white waxy material. Short, soft, spines pro-
ject from the edges of the body forming a sort of fringe. The two
posterior spines are three or four times longer than the others.
The winter is spent in the egg stage under white cottony masses
in bark crevices, under bark scales, and in the axils of twigs and
leaves. There are several generations per year. Adults feed at
pruning scars and in splits in the bark, causing the formation of
knotlike galls. Important parasites include the hymenopterons,
Pseudaphycus malinus Gahan, an introduced species from Japan,
and Allotropa convexifrons Muesebck. The latter is often very
effective in the Northeast.

Heavy infestations of the striped mealybug, Merrisia virgata
(Ckll.), have been observed on azalea in Maryland (351). It has
also been recorded on azalea in Pennsylvania and Virginia and
on magnolia in Alabama. The adult is covered with flossy threads,
much longer than the body. Young nymphs feed on tender growth,
such as flower buds; later, on all parts of the plant except the
main stem and roots.

FAMILY ASTEROLECANIIDAE
PIT SCALES
Pit scale insects are small and elongate-oval in shape. Their
legs are either vestigial or lacking, their antennae are short and

4- to 6-segmented, and their bodies are either covered by a tough
waxy film or embedded in a waxy mass. The genus Asterolecanium

100

contains a number of species that are often destructive of forest,
shade, and ornamental trees. Females are smooth and shiny, light
yellowish to brownish, and about 1 to 3 mm. wide. Males are un-
known in most of the species occurring in this country. Morrison
and Morrison (545) published on the subfamily. Asterolecaniinae.
Russell (617) published a classification of the genus
Asterolecanium.

The golden oak scale, Asterolecanium variolosum (Ratz.), is
one of our most important species of pit scale insects. It feeds
almost exclusively in small pits on the twigs, branches, and trunks
of white oaks, and probably occurs wherever its hosts grow in this
country. English oak appears to be favored above all other
species. In its absence, chestnut oak is preferred (580). Females
give birth to living young which move rapidly over the surface of
twigs and branches. Once they settle down and begin to feed,
however, they never move again. Small pits develop as they feed
and the scale appears to be embedded in the bark. Thse pits re-
main after the scale dies. As a result, the twigs and small
branches of heavily infested trees become quite rough. Adults
overwinter, and there is one generation per year in the North-
eastern States. Heavy infestations may kill young trees and the
branches of larger trees (578). Heavily infested mature trees are
killed during periods of dry weather.

Asterolecanium quercicola (Bouché) feeds in pits on the twigs
branches, and trunks of white, swamp white, red, chestnut, and
English oaks from New York to Ohio and North Carolina. Heavy
infestations kill twigs, branches, and occasionally entire trees. A.
luteolum Russell, the yellow oak scale, and A. minus Lindinger
attack several species of oaks in Pennsylvania and Ohio. The
holly scale, A. puteanuwm Russell, attacks two species of holly (Ilex
vomitoria Ait. and I. opaca Ait.) and Bumelia sp. from Delaware
and Pennsylvania to Alabama and Florida. A. arabidis (Sign.),
the English ivy scale, occurs on green ash from Massachusetts and
New Jersey to Ohio. The oleander scale, A. pustulans (Ckll.), feeds
on Acacia spp., the marmalade tree, magnolia, mulberry, and
many other species. It is sometimes quite injurious. A. bambusae
(Bdv.), the bamboo scale, is a common and injurious pes: of
bamboo in the Southern States. A. miliaris miliaris (Bdv.) and
A. miliaris robustum Green also feed on bamboo.

FAMILY COCCIDAE
SOFT SCALES

Soft scale females are either bare or enclosed in waxy or cot-
tony secretions; hence the common name, soft scales. They may
be recognized by an anal cleft running anteriorally from the pos-
terior apex of the abdomen. The anal opening lies at the top of
this cleft and is covered by two small triangular plates. Forest,
shade, and ornamental trees are attacked by many species, several
of which are serious pests.

The cottony maple scale, Pulvinaria innumerabilis (Rathvon),
an important pest of shade tree maples, occurs throughout much
of the United States and Canada. Its favored host is silver maple,
but a large number of other deciduous trees are also attacked,

101

such as other species of maple, basswood, white ash, boxelder,
dogwood, honey and black locusts, hackberry, sycamore, beech,
elm, willow, and poplar. The female is pale to dark brown, bare,
convex to oval in shape, has a more or less distinct median ridge,
and is about 4 to 6 mm. long. They are most conspicuous during
the summer when large, white, cottony egg sacs extend from the
rear. Males are flat, winged, and much smaller than the females.

Eggs are laid from April to June in cottonlike masses under
the female. These masses are several times the size of the scale.
As a result, the rear of the scale is often elevated at an angle to
the twig (fig. 27). Hatching occurs in June and July in the more
northern States. Young larvae crawl to the undersurfaces of
leaves and feed on the principal veins. They become mature in
August and September and mate. The fertilized immature females
crawl back to the twigs and small branches where they over-
winter. Development is resumed in early spring and the females
become mature by April.

F-504123

FIGURE 27.—The cottony maple

é scale, Pulvinaria innumera-

vb bilis, on twigs and leaves of
soft maple.

o

: %

eae ny

Damage to heavily infested maples may be severe. Branches are
killed, the foliage turns a sickly yellow, and the vigor of the tree
is reduced. Such trees are then frequently attacked by other
species of insects and further damaged. Honeydew dripping from
infested shade trees on sidewalks, automobiles, or people is often
a nuisance. Sooty mold developing in the honeydew also detracts
greatly from the attractiveness of the trees. Fortunately, trees
are seldom attacked heavily for two successive years. Insect pred-
ators, including the dipteron, Lewcopomyia pulvinarie, larvae of
the lepidopteron, Laetilia coccidivora, and the coccinellids, Chilo-
corus bivulnerus and Adalia bipunctata sometimes play a primary
role in the suppression of heavy populations.

Pulvinaria acericola (Walsh and Riley), the maple leaf scale,
feeds on soft and sugar maples throughout most of the Eastern
States and in southern Canada. Female scales are dark purple,
have a brownish-yellow mid-dorsal stripe and three rows of small
waxy spots on each side, and are about 5 mm. long. The body
usually bears tiny tufts of white, cottonly filaments and a long
cottony egg sac at the posterior end.

Partly grown nymphs spend the winter on the branches of the
host. Growth is completed in the spring. Males appear first and

102

fertilize immature females. During May the females migrate to
the undersides of leaves and complete their development. A mass
Eggs deposited in the sac hatch in June. The nymphs feed on the
of white cottony threads is secreted which forms the egg sac.
leaves until late fall and then migrate to the branches where they
spend the winter. There is one generation per year (12).

Pulvinaria floccifera (W.) feeds on Taxus in the Northeast and
Midwest. Adult females are pale yellow above and reddish-brown
or brightly ochreous below. The undersides of branches of heavily
infested plants become covered with irregular, white, cottony
masses. Damage is occasionally severe in the Northeast.

The magnolia scale, Neolecaniuwm cornuparvum (Thro), occurs
throughout Eastern United States and feeds almost exclusively on
various species of magnolia. This is one of the largest and most
conspicuous scale insects found in the United States. Adult fe-
males are about 12 mm. in diameter; elliptical; dark, shining
brown; convex; smooth; and covered with a white waxy bloom
(fig. 28). The nymphs are small, flat, and blackish with a waxy
bloom. The winter is spent as a first-instar nymph on the newer
wood. Females become mature by late July and early August as
far north as Ohio and give birth to living young. The latter move
to the current year’s twig growth where they settle down and
feed. Sometimes they are abundant enough to cover many
branches of the tree. There is one generation a year. Heavily in-
fested trees may be seriously weakened, and, if infested continu-
ously for several years, are killed. Heavily infested trees are also
rendered unsightly by sooty mold developing in honeydew secreted
by the scale.

The tuliptree scale, Towmeyella liriodendri (Gmelin), occurs
from New York and Connecticut to Florida and westward to the
Mississippi River Valley. It has also been recorded in California.
Yellow poplar is the preferred host (fig. 29), but the scale also
attacks basswood, native and cultivated magnolias, persimmon,
loblolly-bay, buttonbush, catalpa, and walnut. Mature females are
dark brown, about 8 mm. in diameter and approximately hemi-
spherical when not crowded. Males are small, inconspicuous, dark
brown, and two-winged. Young scales are very small and range
from gray to black.

Winter is spent as a second-instar larva in regions as far north
as Pennsylvania and Ohio, and in all developmental stages in the
Deep South. Male scales emerge first in the spring and mate with
partially grown females. The female gives birth to living young
in the fall. Newly-hatched larvae are very active and crawl over
most of the host. During the winter they are usually found on the
less exposed areas of branches and on the trunks of saplings,
usually on wood less than 4 years old.

The lower branches of mature trees and entire trees up to 5
inches in diameter may be killed by heavy infestations. The ter-
minal shoots of surviving saplings are often killed, leading to
crooks which makes the trees worthless for future use as veneer
or lumber. Black sooty mold developing in honeydew excreted by
the scale detracts from the attractiveness of shade trees (200).

The pine tortoise scale, Towmeyella numismatica (P. & McD.),
occurs in southeastern Canada and from New York and New

103

COURTESY CONN. AGR. EXPT. STA.

FIGURE 29.—The tuliptree scale,

Toumeyella liriodendri, on yellow-
poplar.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 28.—The magnolia scale,

Neolecanium cornuparvum, on
magnolia.

Jersey to Nebraska and the Dakotas. Its favored hosts appear to
be Scotch and jack pines, but it also feeds on several other pines.
Austrian pine is fairly commonly attacked. Infestations have also
been found on red pines growing among or adjacent to heavily
infested jack pines. Mature females are reddish-brown, oval, very
convex, and about 6 mm. long. Males are tiny, fragile, and winged
(469).

Partly-grown females spend their first winter on twigs. Growth
is completed by June or earlier of the following year, and each
female deposits about 500 eggs under her body. Hatching occurs
in June or July, and the young larvae begin feeding on the twigs
immediately. A white, powdery substance develops on the margins
of the larvae. Male adults emerge in about 3 weeks and fly in
search of immature females with which they mate. A few days
later, the males die. The female continues to develop slowly until
late fall, then goes into hibernation. There is one generation per
year.

Pole sized stands of pine are occasionally severely infested, but
seedlings and young saplings usually suffer the greatest damage.
Heavy feeding may result in considerable branch mortality or the
death of entire trees. Damage in Christmas tree plantations in
the Midwest is often severe. In heavy infestations, a large per-

104

centage of the trees may die following one or two seasons of
attack.

Heavy infestations of the pine tortoise scale are frequently
wiped out by various species of coccinellid beetles. Hyperaspis
congressis is especially effective. Several other species including
Hyperaspis signata, Chilocorus bivulnerus Muls., Scymnus lacus-
tris, are also important in different parts of the scale’s range. The
larva of the moth, Laetilia coccidivora, is also effective at times.

Toumeyella sp. near numismaticum feeds on Virginia pine in
Pennsylvania, West Virginia, and Maryland; T. pini (King) has
been recorded feeding on mugho and Scotch pines in Pennsyl-
vania, Connecticut, Michigan, and Florida; and T. parvicornis
(Ckll.) has been observed feeding on pines in Alabama.

The terrapin scale, Lecanium nigrofasciatum Perg. occurs in
southern Canada and over most of the eastern part of the United
States. Its preferred hosts appear to be the maples, sycamore,
boxelder, and hawthorn; however, it also attacks many other
forest and shade trees such as ash, mulberry, birch, cottonwood,
redbud, basswood, and live oak. Mature females are convex, oval,
and about 2 mm. in diameter. The body is dark reddish-brown,
smooth and shining, and has 10 or 12 dark radiating bands ex-
tending from the high center of the dorsum to the crimped or
fue edges of the body. Males are two-winged and about 1 mm.
ong.

Partly grown females overwinter attached to branches. Growth
is completed in the spring and the female gives birth to living
young from early June to mid-July. The young scales migrate to
the leaves and settle on the midrib or larger veins on the under-
surfaces. They remain here for about 6 weeks and then move back
to the branches. Male adults emerge and mate with immature fe-
males. The latter continue their growth until cold weather and
hibernate. There is one generation per year.

The terrapin scale is most important as a pest of shade trees
because it frequently appears in great numbers and seriously re-
duces the vitality of affected trees. The twice-stabbed lady beetle
exerts a considerable degree of control.

The European fruit lecanium, Lecaniwm corni (Bouché), also
commonly known as the brown elm scale, is widely distributed in
the United States and southern Canada. It feeds on a wide variety
of forest, shade, and fruit trees. Important species include elm,
maple, yellow-poplar, white and black oaks, sycamore, beech, bass-
wood, hickory, mulberry, walnut, black and honey locusts, hack-
berry, poplar, ash, magnolia, aborvitae, and juniper. Mature fe-
males are dark brown to reddish, circular to oval, strongly convex,
3.5 to 6 mm. in diameter, and often covered with whitish powder.
Living specimens are soft and plastic; at death the body becomes
a hard, brown shell loosely attached to the bark.

Eggs are laid in early summer. Newly-hatched larvae crawl to
the undersurfaces of leaves and congregate near the veins. They
remain here until late summer and then they migrate to the bark
of small branches where they spend the winter. Two generations
per year have been recorded as far north as Pennsylvania. Dam-

105

age to shade trees, especially elm, is often severe. Small branches
are sometimes killed and weakened trees become subject to attack
by various species of boring insects or bark beetles.

The Fletcher scale, Lecanium fletcheri Ckll., is widely dis-
tributed in eastern and midwestern United States and Canada.
Heavy infestations have been observed on Taxus, juniper, and
arborvitae in nurseries in Ohio, Indiana, and Connecticut, and on
arborvitae in North Dakota. Because of its prevalence on and in-
jury to Taxus, it is sometimes called the taxus lecanium. Eggs are
laid in late May and hatch by early July. Adult females are light
yellowish-brown, almost globular, and about 3 to 5 mm. in di-
ameter. Foliage on the inner portions of heavily infested plants
is often blackened by the sooty mold growing on the honeydew
excreted by the scales. The oak lecanium, L. quercifex Fitch, at-
tacks many species of oaks throughout the Eastern States and
southern Canada. L. caryae Fitch occurs commonly on beech,
birch, hickory, elm, oak, willow, and walnut. This is one of the
largest members of the genus—mature specimens are up to 12
mm. long. L. quercitronis Fitch is occasionally abundant on oaks
in the South.

The spruce bud scale, Physokermes piceae (Schr.), feeds on
Norway spruce in the Northeastern States and southern Canada.
Mature scales are about 3 mm. in diameter and usually occur in
clusters of 3 to 5 at the bases of branchlets. The body is reddish-
brown with irregular flecks of yellow, and is covered with a deli-
cate coating of powdery wax. The winter is spent as immature in-
dividuals clustered about terminal buds and there is one generation
per year. Heavily infested trees are weakened and rendered un-
sightly by black mold developing in the honeydew which the scale
excretes.

Pseudophilippia quaintancit Ckll., the woolly pine scale, feeds
on young trees of several species of pines from New England to
Florida and Louisiana. It forms white, wooly masses at the base
of needles of new growth (240).

FAMILY DIASPIDIDAE
ARMORED SCALES

The family Diaspididae contains more than one-half of all
species of scale insects. Members of the family live under scales
composed of cast skins of earlier stages held together by waxy
excretions of the insects. The terminal segments of the body are
fused and strongly chitinized. Adult females are small, flattened,
disclike, and legless. The scale coverings vary from long and
threadlike, to oyster shape, to circular. The females of some
species lay eggs; others give birth to living young. Adult males
and their scale covers are usually elongated. Various trees and
shrubs are the principal hosts. Kosztarab (436) discussed the
species occurring in Ohio.

The San Jose scale, Quadraspidiotus (= Aspidiotus) perniciosus
(Comstock), a probable introduction from China, was first re-
corded in North America at San Jose, California in 1880. In 1893
it was also found in Virginia. It now occurs throughout the United
States and southern Canada. Its hosts include a wide variety of

106

forest, shade, ornamental, and fruit trees. Some of its most im-
portant forest and shade tree hosts are: elm, ash, poplar, linden,
willow, redbud, magnolia, and mountain-ash. Female scales are
flat, circular, 1 to 2 mm. in diameter, and have a slightly raised,
dark or yellow central area or nipple (fig. 30). Immature scales
are small, nearly black, and have a black nipple surrounded by a
ee ring. Male scales are oblong-oval and nearly twice as long
as wide.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 30.—San Jose scale, Quad-

raspidiotus perniciosus. Note

large number of immature
scales.

The winter is spent as a partly grown larva. Mature males ap-
pear in May and mature females in June. Fertilized females give
birth to 200 to 600 living young each. Young larvae or “crawlers”

swarm over the tree before settling down to feed. The bark of

heavily infested trees may be literally coated with a grayish
crust.Male scales reach maturity in 3 to 4 weeks; the females in
about 7 weeks. There are one to several generations per year,
depending on locality.

For many years after it spread over the United States, the San
Jose scale was considered one of the most serious pests of orchard,
shade, and ornamental trees in the country. In recent years it has
generally been less troublesome. Two species of parasites, Pros-
paltella perniciosa Tower and Aphelinus proelia (Walker), have
been quite effective in its control in the Northeast.

The walnut scale, Quadraspidiotus (=Aspidiotus) juglansregiae
(Comstock) has a wide variety of hosts including walnut, ash,
maple, boxelder, horsechestnut, cottonwood, buckeye, and linden.
Female scales are flat, circular, pale grayish-brown, and about 3
mm. in diameter. The exuvium is subcentral and reddish-brown.
Young scales usually settle down around the female in a regular
formation. Infestations heavy enough to kill twigs and branches
have been observed in the South. There are two or more genera-
tions per year. The related species, Q. forbesi (Johns), has been
observed feeding on cherry and Q. gigas (Thiem & Gerneck) on
poplar and willow.

107

—

a

The Putnam scale, Diaspidiotus (=Aspidiotus) ancylus (Put-
nam), occurs in southern Canada and throughout most of the
United States. Silver maple and linden seem to be its favored
hosts, but it also feeds on several other important forest and shade )
trees such as elm, ash, hackberry, poplar, dogwood, black locust,
and beech. Female scales are circular and about 1 to 1.5 mm. in
diameter. They are dark gray to nearly black and have an off-
center, brick-red nipple. Otherwise, they closely resemble females
of the San Jose scale.

Winter is spent as an immature scale, and growth is resumed
and completed in the spring. Eggs are laid over a period of sev-
eral weeks in early summer, and there is one generation per year.
Ordinarily, this scale is not very injurious; although, it may kill
small twigs and branches on heavily infested trees.

Diaspidiotus liquidambaris (Kotinsky) feeds on the leaves,
twigs, and buds of sweetgum in the South and Midwest. Infesta-
tions have also been observed on magnolia and red maple. During
the summer it occurs primarily in pockets on the lower surfaces
of leaves. Small, discolored galls occur on the upper surface close
to the pockets. Damage has been serious in nurseries in Missouri.
D. (=Aspidiotus) aesculi (Johns) has been observed on horse-
chestnut.

The hemlock scale, Abgrallaspis (=Aspidiotus) ithacae (Ferris)
feeds on hemlock, fir, spruce, and pine in Eastern United States.
It is small, oblong, dark gray or nearly black, and is usually found
attached to the undersides of needles. The needles of heavily in-
fested twigs and branches may drop prematurely. The related
species, A. townsendi (Ckll.), has been observed feeding on
yellow-poplar.

The gloomy scale, Melanaspis (=Chrysomphalus) tenebricosa
(Comstock), occurs from Maryland to Ohio, Florida, and Texas.
It is often quite destructive of soft maples in the South. Other
hosts include sugar maples, elm, boxelder, and hackberry. Female
scales are gray and almost the same color as the bark. They re-
semble San Jose scales except for their more concentrically placed
nipples (517).

The obscure scale, Melanaspis obscura (Comstock) (=Chrys-
omphalus obscurus), occurs from Massachusetts to Florida and
Arkansas and feeds on a wide variety of forest and shade trees,
including oak, maple, elm, hickory, willow, dogwood, hackberry,
and pecan. Female scales are dark gray to blackish, almost circu-
lar, and about 3 mm. in diameter. The first exuvium is subcentral,
nipplelike, and shining black with a white ring. Damage to pecan
is often serious in the South. Chestnut and white oaks are injured,
and, in Maryland, willow and pin oaks are severely damaged.

The elm scurfy scale, Chionaspis americana Johnson feeds on
elm and hackberry throughout most of Eastern United States.
Female scales are 2 to 3.2 mm. long, oystershell-shaped, and al-
most pure white. Twigs, branches, and small trees may be killed
in heavy infestations, and large trees, seriously weakened. There
are two generations per year in the North, but in the South there
are probably more.

The scurfy scale, Chionaspis furfura (Fitch), feeds on the
leaves, branches, and trunks of a wide variety of trees in southern
Canada and the United States. Important forest and shade tree

108

hosts include elm, ash, maple, hickory, walnut, aspen, willow,
mountain-ash, and horse chestnut. Female scales vary from pure
white to dirty gray, are somewhat pear-shaped, and are from 2 to
3 mm. long. C. lintnert Comstock feeds on willow, birch, dogwood,
and other trees and shrubs in southeastern Canada and the North-
eastern States. C. corni Cooley often damages ornamental dog-
woods in the Midwest.

Chionaspis salicisnigrae (Walsh), the willow scurfy scale, occurs
across the Northern States and in southern Canada. Willow ap-
pears to be its favored host, but it also feeds on several other
trees such as dogwood, poplar, yellow-poplar, and shadbush.
Female scales are white, elongate-oval, convex, and about 8 to 4
mm. long. Infestations are often heavy enough to cover completely
the twigs and branches of its hosts. These may seriously injure
young trees.

The oystershell scale, Lepidosaphes ulmi (L.), occurs in south-
ern Canada and throughout the United States. Its hosts include
more than 100 species of forest, shade, fruit, and ornamental
trees. Important species of forest and shade trees include white,
European, and black ash, beech, silver maple, elm, willow, poplar,
birch, and lilac. As its common name indicates, this scale re-
sembles a miniature oystershell (fig. 31). The female is chestnut
brown or darker and about 2.5 to 3 mm. long.

Eggs overwinter under the protective covering of female scales.
Hatching occurs in early June, or late May, or even earlier, de-
pending on location. Newly-hatched nymphs wander over the bark
for a short time, then settle down to feed. Eggs are laid during
early summer in the South, and during late summer and early fall
in the North. There are two generations per year in the South but
only one in the North.

The oystershell scale often occurs in such large numbers as to
form a crust on the trunk or branches of its host. Damage to ash
has been severe during past years. According to some reports,
entire forest stands have been killed. Lilac and silver maple are
also subject to serious injury. A related species, Lepidosaphes
yanagicola Kuwana, first found and identified in Ohio in 1950,
feeds on Kuonymus.

The pine needle scale, Phenacaspis pinifoliae (Fitch), occurs in
southern Canada and throughout the United States. Its hosts in-
clude various species of spruces and the following species of
eastern pines: white, red, Scotch, Austrian, Virginia, slash, lob-
lolly, and mugho. Mature females are white, have a yellow pellicle
attached to the anterior end, and are about 3 mm. long. They are
usually widest toward the posterior end. The shape tends to vary,
however, depending on the shape and width of the needles on
which they live. Males are white and slender and not more than
half as long as the females.

Eggs spend the winter under the dead scale covering of the
female. Hatching occurs about the time the new needles appear
and the newly-hatched larvae crawl to green needles to feed. They
become mature in early July and the new females produce new
batches of eggs. Offspring of this generation reach maturity by
fall and lay eggs which overwinter. There are two generations a
year as far north as Ohio.

109

F-504093

FIGURE 31.—Oystershell seale,

Lepidosaphes ulmi, on bark of
poplar.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 32.—Pine needle _ scale,

Phenacaspis pinifoliae, on nee-
dles of red pine.

Pine needle scale infestations (fig. 32) are occasionally so heavy
that infested trees present a gray, unhealthy appearance. Heavy
infestations cause the needles to turn yellow and may kill parts of
branches or entire young trees. Damage is particularly noticeable
and troublesome on ornamental plantings in or near towns and
cities. The twice-stabbed lady beetle is an important enemy of

the species.

110

The black pine-leaf scale, Nuculaspis californica (Coleman), oc-
curs over most of North America and feeds on many different
species of conifers. Mature female scales are yellowish-brown to
black and almost circular in shape. They are about 1.5 mm. in
diameter, and there is a small elevated nipple in the center. Their
first winter is spent as partly-grown females attached to the
needles. “Crawlers” appear in late spring and move to the old
needles to feed. A second generation appears in midsummer and
feeds on needles of the current season. Damage has been serious
in nurseries and plantations and to natural reproduction in the
Lake States. Heavily infested needles become desiccated and die.
ane twice-stabbed lady beetle is an important enemy in the Lake

tates.

The juniper scale, Carulaspis (=Diaspis) caruelt (Targ.), oc-
curs in southern Canada and throughout the United States. It is
primarily a pest of juniper but also feeds on arborvitae, incense-
cedar, and cypress. The female scale is white with a yellow center,
circular and slightly convex, and about 1.5 mm. in diameter. Male
scales also are white, but are long, narrow, and smaller than the
females. Only fertilized females overwinter. Egg laying begins in
May and hatching occurs during June. Newly-hatched larvae or
“crawlers” migrate to feeding sites. Once they begin to feed they
never move again. There is one generation per year.

The juniper scale is also a serious pest of ornamental ever-
greens. It feeds on many different parts of its hosts including
the needles, twigs, and cones. Heavily infested plants make poor
growth and become grayish-brown in appearance.

The evonymus scale, Unaspis euonymi (Comstock), a common
and serious pest of deciduous and evergreen Euonymus, probably
occurs wherever its hosts grow in the United States. Mature fe-
male scales are grayish-brown, roughly pear-shaped, and from
1.6 to 2 mm. long. Males are snow white, somewhat narrower
than the females, and have three longitudinal ridges. Mature
females overwinter. First brood larvae appear from early to late
spring; a second brood appears in July. There are two to three
generations per year, depending on locality. Plants are often
heavily infested, the stems and leaves being practically covered
by the scales (fig. 33).

The camphor scale, Pseudaonidia duplex (Ckll.), a probable in-
troduction into Louisiana from Japan around 1920, is now known
to occur also in Texas, Mississippi, and Alabama. Its hosts include
more than 200 species of plants, but the camphor tree is the most
seriously injured. Uncrowded female scales are nearly circular,
convex, have a subcentral nipple, and are about 1.5 mm. in
diameter. There is no well-defined hibernation period, and egg
laying probably begins as early as January. There are three gen-
erations per year.

Fiorinia externa Ferris has been recorded from New York,
Massachusetts, Connecticut, Rhode Island, New Jersey, Mary-
land, Virginia, Pennsylvania, and Ohio. Its hosts are listed as
Canadian, Carolina, and Japanese hemlocks, also fir, spruce, and
yew (703). All developmental stages of both sexes are present
throughout the year. Two generations per year have been re-
ported from Maryland and Connecticut. Infested hemlock takes

dled

F-519580
FIGURE 33.—The euonymus scale, Unaspis euonymi, on Euonymus.

on a mealy, whitewashed appearance in the spring and fall.
Needles drop prematurely and twigs and branches are killed on
heavily infested trees. The related species, fF’. theae (Green), a
serious pest of camellia in the South, occurs on holly also.

Lecanodiaspis tessalatus (Ckll.), the persimmon scale, a large
hemispherical species, feeds on persimmon in the South. The sur-
face of heavily infested limbs and trunks become rough, lumpy,
or wartlike. The white peach scale, Pseudaulacaspis pentagona
(Targ.), feeds on ash, dogwood, chinaberry, and lilac; Chrys-
omphalus dictyospermi (Morgan) feeds on holly (in Virginia) ;
and A. hederae feeds on chinaberry.

112

ee a ee |, ee

Order COLEOPTERA

Beetles

The beetles comprise the largest order of insects. Over a quarter
of a million species have been described, and over 26,000 of this
number occur in the United States. They can be distinguished
from all other insects, except the earwigs, by the structure of the
forewings, these being horny “wing covers” or elytra which meet
in a straight line along the middle of the back (152). They cover
the membranous hindwings when the latter are present. Beetle
larvae vary considerably in form in different families. The ma-
jority are campodeiform or scarabaeiform, but some are platy-
form, some are elateriform, and a few are vermiform. All beetles
have complete metamorphosis, and all have chewing mouth parts.
The life cycle varies greatly in length, from 10 or more genera-
tions per year to one generation in many years.

Beetle larvae and adults present a wide range of habits.
Some species are aquatic, but the majority are terrestrial. Some
feed on vegetable matter; others on animal matter. Many are
phytophagous, many are predaceous, some are scavengers, some
feed on mold or fungi, and a few are parasitic.

Many species of the phytophagous beetles are external feeders
on the foliage, and many others feed under the bark. Some feed
in the wood or fruit, on parts of blossoms, as miners in leaves, or
on the roots of their hosts. Some of the predaceous species feed
both as larvae and adults on free-feeding lepidopterous larvae and
various sucking insects; others feed on various insects in the leaf
litter and under logs, or on the immature stages of bark beetles
and wood borers in their burrows in wood. A few of the parasitic
species feed on the egg masses of grasshoppers or in the nests of
bees.

The order Coleoptera contains many of the most destructive
forest insects. The adults of many leaf-eating species cause exten-
sive defoliation and damage. Many others kill valuable timber or
shade trees by introducing blue-staining fungi or other lethal
pathogens as they bore into their hosts. The larvae are generally
most destructive, however. For example, enormous losses of stand-
ing timber are caused each year by phloem-feeding larvae; severe
losses in timber degrade and to forest products are caused by the
larvae of various woodboring species. Young trees in nurseries
and plantations as well as natural reproduction are severely dam-
aged or destroyed by the larvae of other species.

For more extended treatment of the Coleoptera, the reader is
referred to the following works: Blatchley (81), Boving and
Craighead (91), Bradley (94), Leng (461), and Leng and
Mutchler (452, 453). Dillon & Dillon published a manual of
species occurring in eastern North America (193).

Key to the Families of More Important Beetle Larvae

1. Legs apparently 6-jointed, the 5th bearing 1 or 2 distinct
PORTO e HGlA WSs ie Ghee Tg Tas tee Shee

Legs either 5- jointed with. tarsus and claw fused into

a single claw ail terminal oe or less than 5-

jointed or absent | Boxe hee 6

bo

113

10.

dele:

12.

13.
14.
ily

16.
ie

114

Hypopharynx and ligula strongly chitanized; soft-bodied,
wood-boringe forms, 422. ee ee
Hypopharynx soft; ligula soft or vestigial; mandibles
falcitorm:, fitteds fom eas in cae ee eee
Ninth abdominal segment armed with 1 or 2 chitinous proc-
esses; mandible with large grinding molar structure ....
Ninth abdominal segment unarmed; mandibles of grasp-

ING ty Pe. ee ee eee eee eee wee Rhysodidae
Ninth abdominal segment dorsally produced into a
straight: pROCeSS .22 ee eee Cupesidae
Ninth abdominal segment bearing a dorsal and ventral
process, curved toward each other .......... Micromalthidae
Body fusiform, usually developed for active motion, no
abdominalhooks) <2). 3. eae ent en ere Carabidae

Body fleshy digitate; dorsal surface of 5th abdominal
segment bearing 2 or 3 pairs of hooks; larvae living in
earthen burrowsoin the-soili eee eee Cicindelidae

Ninth abdominal segment bearing 1- or 2-jointed, mov-
able appendages or cerci- 2-2) ee

Cerci-rigid: or absent. 0) ae ee eee

Mandibles of the grasping type without a grinding or
molar structure at base; gula reduced or represented
by a line, the sides of head meeting ventrally ................

Mandibles of the biting type; gula present ........ Silphidae

Spiracles bifore; cardo and stipites fused .......... Histeridae

Spiracles annular; stipites articulate on cardo

Staphylinidae

Curved, grublike larvae with strong well-developed legs;
mandible with a large grinding structure; lacinia and
galea usually distinctly divided; spiracles usually crib-
rif orm: NGO! CORCL) Oh re ae ee een nee

Form variable and not having the above combination of
CHAP ACTER Sis ee Ora ee Se ence en he een a eter

Gula present separating submentum from prothoracic

skins; laciniaeZ-joOmmtedip se eee ees Byrrhidae
Gula absent; submentum and prothorax continues in
a soft skinlike connection 0... Lamellicornia

Gular region or a median gular suture present ............

Gular region fleshy, i.e., submentum and presternum
of prothorax continuous in a skinlike connection .......

Parasitic; soft-bodied, swollen larvae with head and
body white; mandible without molar structure |

Not parasitic; head and body normally chitinized __.
Mouth parts peouracheGa ee Cucujidae
Mouth pantsenetracted te ee ..... Colydiidae

Labrum absent or fused with clypeus into a nasale

Labrum distinct. eee

Head structures reduced and specialized; body usually
much depressed; wood borers, the larvae cutting gal-

leries across the grain of the wood oer cee 7 |
Head structures not so modified ...............................Klateridae
Legs short, 5-jointed. .................... dais pope NFOSCICae

Legsrabpsent. or vestigial... eee Eucemidae
Maxillary articulating area indistinct ......... ,

10
Ht

12
40

13
14

15
seg

16

18

nS:

Ih.

20.

21.

22.

23.
24,

25.

26.

21.

28.

29.

30.
Sie

Maxillary articulating area large and distinct ..................
Ninth abdominal segment operculate, terminal; body
cylindrical, heavily chitinized 000.000.000.000... Rhipiceridae
OdWAOEherwiSeles eee a ES 00 ee eae ei eas hae ee
Prothorax enlarged and flattened, marked with a dis-
tinct circular or oval area above and below; legs ves-
Eiodalmore wanting) 0s... ee eae Ee ee Buprestidae
Prothorax not specialized; legs well developed ....0.............
Body fleshy; curved, inactive larvae feeding in fungi
Cisidae
Body digitate or active; predaceous forms or scav-
CTU CNS epee reel trea Para cuit ah es 9 Oey <n CN es ane ET ee Te
Ventral mouth parts retracted considerably behind a line
between the ventral mandibular articulations .................
Ventral mouth parts not retracted; predaceous forms

Cleridae
Lacinia armed with spurs; body hairy ........ Dermestidae
WACIMARWALHOULESDULS ie etme, reel ae Ser let ce
Epicranial suture nearly absent 0... Ostomatidae

Ventral mouth parts retracted considerably behind a
line between the ventral mandibular articulations;
molar or grinding structure of the mandible usually
WOGESCMiGe meer eerie em erie rs Gu Ouy § sexe heme ree. ee eee OF

Ventral mouth parts not retracted; larvae soft skinned,
fleshy; wood borers; molar structure absent

Cerambycidae

Spiracles on tubular process; sap feeders

Nitidulidae

Spiracles not noticeably projecting 0.

Free living predaceous forms crawling about on bark
or foliage; 3 ocelli present on each side of head

Coccinellidae

Concealed forms under bark or in wood or fungi...

Form very depressed, about twice as wide as thick;
adapted to living between bark and wood; clypeus and
EOMEROMLOMEA USEC. 8.0, kee @ emis abe sete cee) sealant,

RormeMoresrobvust 0200 Soe. AG ee eee Be.

Cardo not composed of 2 pieces; mala falciform; hypo-
pharynx without a chitinization 0. Cucujidae

Cardo of 2 pieces; mala obtuse; hypopharynx chiti-
TN CG erent hel ec iuerrey tue alk eerie enh sy ean Eh eee ls a

Fighth and ninth abdominal segments nearly equal; a
single pit present between terminal spines; asperities
on venter of 9th abdominal segment arranged in a
brokenmanchiwe: ss lar 2aet ah) oxi bens Salpingidae

Eighth abdominal segment at least twice as long as 9th
(terminal spines excluded) ; a pair of pits present be-
EWE ers Dlleciamet tke putty re. aii dade Cas aaa See Be

Ninth abdominal segment with asperities on venter ar-
raneed in aginele arch 0.0) (ee) ee Pyrochroidae

Mandibles with a reduced, smooth molar structure,
somewhat resembling the mandibles of the Ceramby-
CICACH ee Ae tay ma eare en cia ri ote tee! Way Melandryidae

Mandibles with normal molar structure 00...

19

20

21
22

23

25

26

27

28

31

29

30

a2. Cardo of a:smngle pieces es ee 33

Cardo of: 2. pieces: 210 2.4 See ed eee on

33. Ninth abdominal segment bearing spines, latter with pit
between. Mareins |... ee ee ee ee 34

Ninth abdominal segment with or without spines, when
present pit absent between MAaLgins oe 35

64. Terminal spines biturcate- =.= ee Othniidae

Terminal spines ‘srmpley 2s. eee ee Colydiidae

35. Antenna inserted close to mouth frame; apices of pro-
thoraciczcoxae usually conticueus-. | 36

Antenna inserted some distance from mouth frame; apices

of prothoracic coxae separated 0... Lagriidae

36. Back of mandibles opposite cutting edge, sharp; body
cylindrical; terminal segment without spines and usu-

ally bluntly soundedts2) 2 ee ee Alleculidae
Back of mandibles rounded; body rarely cylindrical;
terminal segment often armed .......... Tenebrionidae
37. Second to 5th abdominal segments with venter bearing
asperatesampullae..= eo ee ee Oedemeridae
Abdominal segments normal, without ampullae . 38 |
38. Ninth abdominal venter bearing a pair of conical points )
Melandryidae |
Ninth abdominal venter simple, without such points 39
39. Submentum and gula fused and heavily chitinized; ter-
minal segment with soft, white spines ....Cephaloidae
40. Hypopharynx chitinized; mandibles with a molar or
erinding ‘structure (225-200 See eee Al
Hypopharynx fleshy; mandibles without a grinding
structure: 428.40 A eee ee eo ae eee 42
41. Legs well developed, 5-jointed; 9th abdominal segment
armed;-body ‘elongate 2.25522 fe Lymexylonidae
Legs weak, 9th abdminal segment unarmed; body
curved; larvae in bark or wood beneath the sporo-
phores, of :woody fungi (= eee _ Anthribidae
42. Ninth abdominal segment ending in a single truncate
spine:or in paired spines 2... | == Mordellidae
Ninth abdominal segment unarmed, except some first-
instar ‘larvae’ ee ee. eee 43 ;

43. Tenth abdominal segment provided with a pair of cush- .
ioned lobes (not anal lobes) separated by a median .
suture; fleshy, curved hairy forms, usually with slen-
der 4- or 5-jointed legs; wood-boring forms

Bostrichidae, Ptinidae

Tenth abdominal segment normal] ...... 44
44, Elongate buprestid-like larvae in the roots of hickory
CFEES> Var, i RO. Nie Peers? ton (Cera eer
Form otherwise 45

45. Legs vestigial, 2- jointed, cylindrical, fleshy, wood-boring
form, with chitinous asperities on prothor ax and ab-
dominal terga Brentidae

Legs 4- or 5-jointed except occasionally absent in some
leaf-mining forms; maxillary mala often divided; hy-
popharyngeal bracon absent 46

Legs wanting; mala always simple; bracon present 47

116

46. Mentum bearing a shieldlike plate; labial palpi absent
or rudimentary; feeding in the seeds of plants

Bruchidae
Mentum without such plate; labial palpi rarely lack-
TPG) eee Rh ee operant a IE OR Fa ca Hee ee OGL Chrysomelidae

47. Adults usually associated with the larvae and boring
characteristic galleries in the wood or under the bark
of shrubs, trees, or lumber...... Platypodidae, Scolytidae
larvae and adults not so associated (220°... 3 ake
48. Abdominal segments with 2‘ transverse folds; larvae
often found in a compacted roll cut from a leaf
Curculionidae
Abdominal segments with 3 or 4 transverse folds ......—s-—- 49)
49, Larvae boring in the moist sapwood of dead trees or
occasionally in beams of buildings, powder-posting the

FVCRCC Tel el ieaeeees ee het ae ee Bw Curculionidae
Habits of larvae variable; not working like powder-post
WeCCilCSmerr oe ee ee Curculionidae

FAMILY CARABIDAE
GROUND BEETLES

This is a large family, containing hundreds of species, nearly
all predaceous on other insects. There is considerable variation
among species in size, shape, and color, but most of them are
dark, shiny, and somewhat flattened. They are commonly found
under stones, logs, leaves, debris, or bark, or running about on
the ground. The larvae of bark-inhabiting species are white and
thin-textured, and the legs are five-jointed with a movable claw or
pair of claws on the fifth joint.

A number of species in the genus Calosoma Weber are impor-
tant predators of tree-defoliating lepidopterous larvae. The adults
are brilliantly colored, are about 25 mm. long, and emit a disa-
greeable odor when handled or disturbed.

Calosoma sycophanta (L.), a large bluish-black species with
golden-green elytra, and from 24 to 30 mm. long, (fig. 34) was
introduced from Europe against the gypsy moth in 1906. It be-
came established rather quickly and has since spread over most
of the gypsy moth infested area and beyond. Both the adults and
larvae climb trees and feed voraciously on the larvae and pupae of
their hosts. Large gypsy moth larvae and pupae are especially
attractive, but they will also feed on many other species of insects.
Adult beetles spend the winter in the ground and emerge about
June 1. Eggs are laid in the ground, and when the larvae hatch,
they climb trees in search of food. When they become full-grown,
they return to the ground to pupate. Adults live up to 4 years.
Populations of this species tend to increase rapidly during gypsy
moth outbreaks (122).

Several native species of Calosoma also feed on lepidopterous
larvae and pupae, sawflies, and other insects in the Eastern States.
C. frigidum Kirby, C. willcoxi LeC., C. scrutator (Fab.), and
C. calidum (Fab.) (fig. 35) are widely distributed in the region.

Many other members of the family are also associated with
insects attacking trees. Chlaenius erythropus Germ., Dicaelus
purpuratus Bon., Agonum spp., and Tachys spp. are frequently

Ly

COURTESY CONN. AGR. EXPT. STA.

SF Eg cp FIGURE 34.—Adult of
fh bps oo Calosoma sycophanta.

COURTESY CONN. AGR. EXPT. STA.
FIGURE 35.—Larva of Calosoma
calidum feeding on a gypsy moth larva

)

found in association with bark beetles under the bark. Galerita
janus F. and G. bicolor Drury are often found under loose bark
of logs or under stones in the forest. The adults are about 20 mm.
long and quite colorful. The thorax is red, the wingcovers bluish-
black, and the long legs red. Geopinus incrassatus (Dej.), Har-
palus spp., and Scarites subterraneus F. frequently feed on and
damage young Seedlings.

FAMILY CICINDELIDAE
TIGER BEETLES

Tiger beetles are of no particular importance as forest insects,
but their appearance and habits are such as to arouse the interest
and curiosity of almost anyone when they are encountered. The
genus Cicindela L. contains the majority of species. The adults
are mostly metallic green or bronze in color, with bands or spots
of yellow. They are strong flyers and fleet of foot, and are most
often seen along sandy beaches of streams or along woodland
trails on bright, sunny, summer days. Tiger beetles and larvae
are predaceous on other insects. The larvae live in vertical bur-
rows in the soil.

FAMILY CUPESIDAE

Cupesid beetles have flattened bodies and are densely scaly.
Cupes concolor Westwood is a common eastern species. Adults are
grayish brown and from 7 to 11 mm. long. Full-grown larvae are
about 25 mm. long; each leg is armed with a movable claw; and
the ninth abdominal segment ends in a short spine. The adults
feed on decaying and often moist wood, and are usually found in

118

basement timbers of pine, oak, and chestnut in old houses. They
are of no economic importance because the infested wood is al-
ready more or less destroyed by decay organisms.

FAMILY SILPHIDAE
CARRION BEETLES

Carrion beetles are mentioned here because they are so often
seen on animal excrement or around the bodies of dead animals
in the forest. Many of the more common species are brightly
colored, and their bodies are usually soft and somewhat flattened.
The larger and more conspicuous species belong to the genera
Silpha and Nicrophorus. Several small species are also found
under the bark of logs and dead trees where they probably act as
scavengers. Agathidium oniscoides Palisot commonly occurs in
the galleries of bark beetles in hardwoods.

FAMILY HISTERIDAE
HISTER BEETLES

Hister beetles are rather small, sluggish, and hard-shelled. The
body is often very flat and the wing covers are usually cut off
square at the tip, exposing the last one or two segments of the
abdomen. Many are shining black; others may be bronzed,
greenish metallic or marked with red, yellow, or orange. The
larvae are soft-bodied and sluggish and may be found in decaying
organic matter, in fungi, or under the bark of trees. Those of
Plegaderus spp. feed on the eggs and larvae of bark beetles.

FAMILY BYRRHIDAE

Members of this family are oval, convex beetles about 1 to 10
mm. long. The head bends downward and is concealed from above,
and the hind femora are wide and extended to the elytra. A few
species live under the bark of trees, but the majority occur in
sandy areas where they may be found under debris or at the roots
of grass and trees. Several species of the genus byrrhus occasion-
ally injure young trees in forest nurseries and plantations. Adults
of this genus are about 5 to 10 mm. long, black and grayish in
color, and densely covered with hairs.

FAMILY PASSALIDAE
PASSALID BEETLES

Passalid beetles are better known by some people by other
common names, such as betsey bugs, betsey beetles, bess bugs, and
patent-leather beetles. Three species are known to occur in the
United States, only one of which, the horned passalus, Popilius
disjunctus (Ill.), is found in the eastern part.

Horned passalus adults are large, shining black beetles, from
30 to 40 mm. long. The plates of the antennal club are rigid and
incapable of being opened or closed; there is a short, forward

119

bending horn on the top of the head; and the elytral striae are
deep and finely punctured. There also are roughened areas on the
undersides of the wings and on top of the abdomen. When dis-
turbed, the adult rubs these areas together, producing squeaking
sounds or the so called “bess”? notes. The larvae are bluish and
from 30 to 40 mm. long. The anal lobes consist of two large
lateral pads, and there are stridulatory organs on the third pair
of legs. The species lives in colonies in large, rough galleries ex-
cavated in damp rotten logs or stumps, each colony consisting of
a male and female and their progeny. The larvae feed on the wood
but not until after it has been reduced to pulp and treated with
digestive secretions by the parents (311).

FAMILY OSTOMATIDAE
OSTOMATID BEETLES

This family contains several important enemies of destructive
forest insects. The adults are rather flattened, elongate or oval,
blue, green, or black beetles. The antennae are 11-jointed, the last
three joints forming a loose club; the thorax is narrowest behind;
and the fourth joint of the five-jointed tarsus is very long. The
larvae are similar to clerid larvae except for the ventral mouth
parts. The latter are retracted below the point of attachment of
the mandibles in ostomatid larvae.

Temnochila virescens (F.) adults are bright, irridescent, bluish-
green; there is a pair of recurved hooks on the ninth abdominal]
segment; and each is about 10 to 18 mm. long. They feed on wood
borers in their tunnels, on bark beetles crawling about on the bark
surface, and on bark beetle eggs and larvae in the bark. This is
one of the most important insect enemies of the southern pine
beetle.

Other predaceous ostomatids occurring in eastern forests and
their insect hosts include the following: Tenebroides corticalis
(Melsh.) and T. dubiws (Melsh.)—feed on bark beetles; T. bima-
culatus (Melsh.)—feeds on Agrilus larvae in oaks; Corticotomus
(Nemosoma) cylindricus (LeC.) and Airora cylindrica (Serv.)—
feed on ambrosia beetles and cossonids.

FAMILY MELOIDAE
BLISTER BEETLES

Blister beetles are most commonly seen on forest vegetation ; they
are fairly large, soft-bodied, sprawling, and loose-jointed. The
head narrows abruptly behind to the neck; the elytra are soft; the
abdomen is often large and swollen; the tarsal claws are cleft or
appendaged; and the color is either green, gray, brown, or black.
The adults are plant feeders and often defoliate their hosts. The
larvae are parasitic of grasshopper eggs or in the nests of bees.

Several species, including the striped blister beetle, HE picauta vit-
tata (F.), and E. pestifera Werner, Macrobasis fabricu (LeC.),
M. unicolor (Kby.), Pomphopoea aenea (Say), and P. sayi LeC.
defoliate seedlings in nurseries and in ornamental plantings in
the Midwest. The ash-gray blister beetle, Epicauta fabriciit (LeC.)

120

E.. subglabra (Fall), and Lytta muttalli Say defoliate Caranga in
the Great Plains; E'picauta torsa (LeC.) defoliates ho'ly in Florida
and mimosa in Florida and Oklahoma; E. cinerea (Forst.) defoli-
ates black and honey locusts and hornbeam in Arkansas; and
Pomphopoea polita (Say) feeds on the pollen of shortleaf pine in
the South. Horsfall (379) discussed the biology and control of
common species.

FAMILY NITIDULIDAE

SAP BEETLES

Sap beetles are elongate or oval, usually less than 25 mm. long,
and often of contrasting black, red, and yellow colors. The an-
tennae are 11-segmented and club-shaped; the sides of the thorax
and elytra are often margined; and the elytra are shorter than
the body, leaving the end of the abdomen exposed. The larvae are
nearly white, flattened, and moderately elongate. The head is
broad and projecting with deeply retracted mouth parts. Paired
and forked horny armatures are often found on the ninth abdomi-
nal segment. Connell (155) published keys to the species occurring
in Delaware.

Both the adults and larvae feed on decaying fruit, on ferment-
ing juices under bark, and on sap exuding from freshly cut logs
and stumps or wounds on living trees. Patches of cambium are
often killed by bark infesting species. When these dead areas dry
up and heal over, they remain in the wood as “bark pocket” de-
fects and lower its value. A number of species also appear capable
of penetrating the bark of oak in the absence of injuries. When
this happens, the larvae feed in the cambium region and destroy
patches up to 2 inches across (fig. 36). Several members of the
family are also likely candidates as vectors of the oak wilt fungus,
Ceratocystis fagacearum. In studies conducted in West Virginia,
they occurred abundantly in and on moist wounds on wild-infested
trees and on mycelial mats of the fungus beneath the bark (201).

FAMILY COCCINELLIDAE
LADY BEETLES

More than 500 species of lady beetles occur in the United
States, by far the greater number of which feed on aphids and
scales, or on the eggs and larvae of larger insects. AS a group
they are among the most beneficial of all the insects. Being quite
common, they are also among the most widely recognized of all
species. The adults are small, usually less than 5 mm. long. The
body is round or hemispherical and extremely convex. The surface
is shining and often brilliantly marked with black, white, red, or
yellow spots. The larvae are alligator-shaped, black or gray in
color, or spotted or checkered with bright colors, and covered
with warts or spines. Eggs are light yellow and are laid in
clusters on the leaves or bark of a plant or tree. The winter is
spent in the adult stage, sometimes in enormous congregations of
beetles. A few of the more important species feeding on forest

121

F-519925-6
FIGURE 36.—Evidence of nitidulid attacks on oak: A, scars of overgrown
attacks; B, pockets in the wood beneath scars shown on A.

insects in eastern United States are discussed. Stehr (689) pub-
lished keys to the identification of species occurring in Minnesota.

The two-spotted lady beetle, Adalia bipunctata (L.), an intro-
duced species, is an important predator of the pine needle, jun-
iper, beech, and terrapin scales in the Northern States. Adults are

122

|
|
|

about 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 with black central spots.

The twice-stabbed lady beetle, Chilocorus stigma (Say), occurs
in the Northern States and feeds commonly on the pine needle,
beech, terrapin, and juniper scales. The adult is shining black and
about 4 to 5 mm. long. The ventral segments are red, and there is
a round red spot on the disk of each elytron.

The convergent lady beetie, Hippodamia convergens Guer. feeds
on aphids and scales throughout most of the United States. Adults
are 6 to 8 mm. long. The head is black except for a pale transverse
center spot; the thorax 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 of beetles.

Hyperaspis congressis Watson is widely distributed in eastern
United States and is an important enemy of the pine tortoise
scale. Adults are black, strongly punctate, shining, and about 2.5
mm. long. There is a rather large, yellowish spot on the middle of
the elytra, and often a fainter one on their tips.

Anatis 15-punctata (Oliv.) feeds on a wide variety of forest
insects, including the larvae of the gypsy moth and cankerworms,
in eastern United States. Adults are yellowish and about 8 to 10
mm. long. There are two pale spots enclosed in a black disk on
the thorax, a single spot on the scutellum, and seven black spots
on each elytron.

Coccinella novemnotata Hbst., the nine-spotted ladybird, 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 above and black beneath. Each elytron
bears four black spots, and there is one black spot on the elytral
suture. C. transversoguttata Fald. feeds on scale insects infesting
pines in the more northern States. Adults are reddish, nearly
hemispherical, and about 6 to 7.5 mm. long. The wing covers may
be spotless, 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 wooly aphid in recent years. The European species, Aphi-
decta obliterata (L.), is now firmly established in infested Fraser
fir stands in North Carolina (6). Eggs are laid on the needles or
trunks of infested trees, and the larvae and adults feed on all
stages of the aphid except the first larval instar. Attempts to
establish it in New Brunswick were unsuccessful (112). Scymnus
(Pullus) impexus (Muls.), one of the most important enemies of
the aphid in Central Europe, has been liberated in infested stands
of balsam fir in New England. The adults and larvae feed on
second and third instar larvae of the aphid.

Other important species of native coccinellids include Scymnus
lacustris LeC., an enemy of the pine tortoise scale in the Lake
States; Microwesia misella (LeC.), a minute species less than 1
mm. long that feeds on various species of scale insects; and Cleis
picta (Rand.), which often occurs in large numbers on pines in-
fested with the red pine scale in the Northeast.

123

FAMILY DERODONTIDAE

Derodontid adults are small brown or dullish-brown beetles
about 2 to 6 mm. long. Only about one-half dozen species have
been recorded from the United States.

Laricobius erichsonw Rosenh., 2 predator of the balsam woolly
aphid, has been imported from Europe during recent years and
released in aphid intested stands of fr in Canada, New England,
North Carolina, and the Pacific Northwest. The adult is an elong-
ate oval beetle about 2.2 to 2.4 mm. long. The body is covered with
yellowish to brownish to black hairs; the head is usually hidden
from above; and the central portions of the elytra, the antennae,
and the legs are reddish brown. The larvae are covered with flakes
of the aphid’s wool and small bits of bark.

Eggs are deposited deep within aphid egg clusters or under
lichens on the bark. The larvae feed on the host eggs until full-
grown at which time they drop to the ground and pupate in
cocoons of compacted soil particles. Adults are active on warm,
sunny days (113).

FAMILY CUCUJIDAE

FLAT BARK BEETLES

Members of this family are rather small and extremely fiat,
yellowish, brownish, reddish, or black beetles. The larvae, also
often very flat, are usually found under the bark of unhealthy
trees or newly-felled trees and logs. The majority are scavengers,
but a few are parasitic or predacious on mites, small insects,
wood boring larvae, or Hymenoptera.

Catogenus rufus (Fab.) is a common eastern parasite of borers
in sugar maple. Adults are elongate, reddish brown and up to 12
mm. long. The larvae are soft-bodied and white, with recurved
hooks on the ninth abdominal segment. They are usually found
beneath the bark in the pupal cells of their host. Cucujus clavipes
F. is another conspicuous eastern species. The adult is bright red,
exceedingly flat, and about 10 to 14 mm. long. The larvae are
usually found under the bark of recently dead trees, especially
ash and poplar.

FAMILY COLYDIIDAE

This family consists of small, elongate, slender beetles rarely
over 5 mm. long. They are reddish brown to nearly black and
beautifully sculptured. The larvae of certain species, some of
which are found under bark in association with bark beetles, are
scavengers. Others, such as those of Nematidium filiforme LeC.,
Bitoma carinata (LeC.), and Colydium lineola Say, feed on the
larvae of ambrosia beetles, cossonids, and certain other borers in
their tunnels.

FAMILY MELANDRYIDAE

Adults of this family are small to large, usually elongate to
oval, loosely jointed beetles. The thorax is margined at the sides;
the antennae are filiform; the front coxal cavities are open be-

124

_

hind; and the hind tarsi are four-jointed. Adults are seldom seen
but the larvae occur commonly under the bark or in the wood of
dead logs, usually with wood-decaying fungi.

Some of the more common species are: Melandrya striata Say—
occurs in many different hardwoods in association with black
lines of decay; Orchesia castanea Melsh.—occurs in oak, maple,
tulip poplar, sycamore, and hickory; Hypulus concolor (LeC.)—
occurs in rotting sapwood of pines; Serropalpus barbatus
(Schall.) and Eustrophus tomentosus Say—occur in various coni-
fers; and Dircacea quadrimaculata (Say) and Holostrophus bi-
fasciatus (Say)—occur in various hardwoods.

FAMILY OEKDEMERIDAE

Adult members of this family are usually slender, flattish, and
soft-bodied. Their color varies from pale to black, with red, yel-
low, or orange markings, and they range in length from about 5
to 25 mm. The elytra are rather soft and are sometimes covered
with fine silken hair; the next-to-last segment of each leg is di-
lated and bears a dense, hairy pad beneath; the middle coxae are
very large; and the hind tarsi are four-jointed. Adults are usually
seen on flowers or on old logs or stumps; the larvae feed in dead
and rotting wood.

The wharf borer, Nacerdes melanura (L.), the only economi-
cally important member of the family, is widely distributed, oc-
curring along both the Atlantic and Pacific Coasts and around
the Great Lakes. Adults are reddish-yellow and vary from 7 to 12
mm. length in the males to 10 to 15 mm. in the females. There is
a dark patch on the head and two dark patches on the thorax. The
tips of the elytra are also black. The larvae feed almost entirely
in very moist wood and occasionally are very destructive of pilings
under wharves and under buildings near the water. Boardwalks
along the shore and wood floors in damp basements are seriously
damaged at times. Adults occur outdoors from April to July.
Inside buildings they may be found during the winter.

FAMILY TENEBRIONIDAE
DARKLING BEETLES

This is a large family of beetles, and many species occur in the
forest. The adults are hard-shelled, usually dark brown or black
in color, and many are clumsy and slow-moving. The larvae are
long and slender, and are covered with tough, horny skins. The
majority of forest species feed as scavengers on dead vegetable
matter. They are commonly found under the bark of dead and
dying or rotten trees or 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

Timber beetles are slender and elongate; the head is deflexed
and narrows behind the eyes to form a neck; the eyes are large;

125

and the legs are slender. Larvae are also elongate and slender; |
the head is globular; the ninth abdominal segment is conspicu- |
ously armed; and the tenth segment is ventral.

The chestnut timberworm, Melittomma sericeum (Harr.), once a
destructive pest of chestnut, is now found chiefly in white oak. |
Adults are brown, clothed with fine, silky hair, and about to 11
to 15 mm. long. Eggs are laid in cracks on the surface of the
wood. The larvae bore deep into the wood, enlarging their tunnels
as needed. Pupation occurs in cells constructed near the surface.

The sapwood timberworm, Hylecoetus lugubris (Say), tunnels
under the bark and across the sapwood of various hardwoods such
as poplar, birch, tulip-poplar, basswood, buckeye, and black wal-
nut. Eggs are laid from April to July in the crevices of bark on
dying trees or on green logs left lying in the woods. Damage is
caused by pin hole defects. Each larva has a slender, barbed spine
on the ninth abdominal segment.

FAMILY MORDELLIDAE

TUMBLING FLOWER BEETLES
Members of this family are commonly called tumbling flower
beetles because they are usually found on flowers and because of
their speed in running, Jumping, or tumbling. They are usually
dark-colored, wedge-shaped, long legged, and from 3 to 14 mm.
long. The larvae of certain species are predaceous on other insects.

Powder-post Beetles

A number of species of Coleoptera belonging principally to the
families Lyctidae, Anobiidae, Ptinidae, and Bostrichidae are com-
monly known as powder-post beetles. The term powder-post, in
its broadest sense, is used to designate the type of damage in
which the interior of wood is reduced to a flour-like powder, some
of which is easily jarred from the wood in handling (40). It may
occur in the sapwood or heartwood of either hardwood or soft-
woods. Damage results from tunneling by the adults of certain
species and by the larvae of all species. Surface holes made by
emerging beetles also mar the surface of the wood. These holes
range in diameter from less than 1 mm. to several millimeters,
depending on the species involved.

A few species of cerambycids, buprestids, and curculionids also
have the powder-posting habit. They are discussed in the sections
devoted to these families. Members of families Lyctidae, Ano-
biidae, Ptinidae, and Bostrichidae, which are discussed below,
comprise the majority of destructive species.

Key to Genera of More Important Powder-Post Larvae

The four families and their more important genera, into which
powder-post beetle larvae are usually divided, are character-
ized and separated in the following key. The genera and species
are not described, because they have not been thoroughly worked
up and are difficult to characterize on the basis of present
knowledge.

126

1. Head almost entirely protruding from the prothorax;
mandible usually dentate; terga usually hairy or as-
OCT ALC Reet wre NF Fin sub erin rs ys we mryieels | Beate re te 3
Head deeply embedded in prothorax; cutting edge of man-
dible gougelike, rarely toothed; terga never asperate... 2
2. Last abdominal spiracle much larger than all others;
mandible with a fleshy process on dorsal inner edge
Lyctus
Last abdominal spiracle not abnormally large; mandible
with or without fleshy lobe oo... Bostrichidae 12
3. First spiracle pushed forward into anterior margin of
prothorax; terga without asperities; usually found in
stored plant material, seed, ete. oie Ptinus
First spiracle between prothorax and mesothorax; terga
usually asperate
4. Chitinization between labium and mentum narrow and
Ueshapedror lacking; mala divided 0... 5
Chitinization arrow-shaped and pointing posteriorly ; mala
simple, no inner lobe; spiracles large, annular without

spoutlike projection from COTO (teas am ce a a ce Ptilinus
5. Inner lobe of mala consisting of a strong, curved spine
Xestobium
Inner lobe of mala fleshy, usually with several smaller
GINS Ole SCCACE 2S. Moma mna ere ten edit Willey Ue 6
6. Inner lobe of mala smaller than outer. Spiracles with a
short or Jone. “spoutlike: process: 2. ee ql
Inner lobe of mala about as large as outer; spiracles with
a Short spouthike process]. Sac se a ee 9
7. Claw short and curved; a fleshy lobe at base; feeding in
stored plant products and Seeds ......................... Lasioderma
Claw usually slender, no lobe; wood borers ................................ 8
8. Spiracles with long spoutlike process; mandible with 3
lateral teeth on dorsal edge 20. Anobium

Spoutlike process on spiracles only indicated; each side
of prothorax curved rodlike impression. Mandible with

Selavetalecectnc Tee ane Hemicoelus

Mandible with 2 lateral teeth ...................... Microbregma

9. Labrum about twice as wide as long .. Ernobius
abtum about, as: wide as, lON@ oct eed 10

10. Tergal asperities in about 2 transverse rows... Xyletinus
Tergal asperities in about 4 transverse rowS |... 11

11. Posterior tergal fold with long, soft hairs .......... Nicobium

Posterior tergal fold densely beset with short, spine-

NIRS ag Settee wey DON Walaa ius bade iid, rio oa de ht le ne Trichodesma

12. Mandible with a large, grinding lobe and a soft, fleshy
lobe on upper inner margin; no impressed line on side

of prothorax; tarsi all bearing chitinized claws... 18
Mandible without chitinous or fleshy lobe; a curved im-
pressed line on side of prothorax, usually only first

Fansalenclaw: 2c Mitinl ZC Gea eee Ahan: Wied waren each ho 16

18. Prothorax bearing a chitinized tubercle in front of spir-
acle; no ocelli; cutting edge of mandible entire 14

No tubercle : ocelli present; cutting edge of mandible
Gdentatetinvi st: a seen Siva SU te 135" TA Teme ne 15
AY

j

14. Inner edge of molar lobe of mandible smooth

Polycaon

15. Six ocelli; breeds in conifers ..........00ccc0.0. Stephanopachys

Breedsiine- bamboo tere ee eee Dinoderus

16. Maxillary mala without projecting style or spine on
inner edveslabrunitcilobede = eee eee ILS

Maxillary mala with a free projecting style or spine on
inner edges labrum entire. eee 18

17. Chitinization of labium broken in middle; prothoracic
spiracle larger than antennal ring

Dendrobiella; Xylobiops

Chitinization of labium continuous across middle; pro-

thoracic=spinacle’smialilerea..) .. ee ee _ Scobicia

18. Spiracle with a spoutlike process; style shorter than

LACTNITA: Sse) ee Re eee We can oem. are Lichenophanes
Spiracles without process; style longer than lacinia

Amphicerus

FAMILY LYCTIDAE

POWDER-POST BEETLES

This family contains the most injurious of the powder-post
beetles occurring in North America. According to some authori-
ties their destructiveness to wood and wood products is exceeded
only by that of termites. The family has been revised by Gerberg.
Keys to all 12 genera in the world and to all 35 species in the new
world were also published (285).

Lyctid beetles are small, slender, elongate, reddish-brown to
black, and about 5 to 6 mm. long. The head is prominent, slightly
deflected, and constricted behind the eyes. Mature larvae vary in
size but are usually less than 5 mm. long. The body is small,
curved, and enlarged at the thorax.

The larvae of North American species bore in the sapwood of
seasoned hardwoods, making small tunnels up to 0.6 mm. in
diameter. Second growth ash, hickory, and oak which has been
seasoned and stored in the same place for 2 or more years are
most seriously damaged. Many other hardwoods such as elm,
maple, poplar, cherry, persimmon, sycamore, and walnut are also
infested. Because starch is an important component of the larval
diet, dry seasoned wood of high starch content is damaged most
seriously. The same wood may be infested repeatedly, and the in-
terior may be nearly reduced to powder, and rendered useless.
Damage is common in stored lumber, the interior woodwork of
buildings, furniture, tool handles, pallets, tent stakes, mallets,
mauls, and many other stored hardwood products. This is par-
ticularly true of wood that is left unfinished until it is placed on
the market or in use.

Lyctus spp. are attacked by several species of insect parasites
and predators; however, they cannot be relied upon to keep these
pests under control (671). The clerid, Tarsostenus univittatus
(Rossi) is a common predator of both adults and larvae. Various
species of predaceous mites also destroy large numbers at times.

128

The genus Lyctus Fab. is represented in the United States by
10 species, all of which feed in the sapwood portions of hard-
woods. The wood of ring-porous species is more susceptible to
attack than that of diffuse-porous species. Oak, hickory, and ash
are highly susceptible. Other species often attacked include wal-
nut, butternut, pecan, elm, sycamore, black locust, poplar, sweet
gum, wild cherry, and osage orange. The extent of infestation
and subsequent damage is proportional to the starch content of
the wood. Since heartwood is practically free of starch it is
immune. Wood infested for the first time may show very little
external evidence of attack, even though the interior is seriously
damaged; whereas the surface of older infested wood is per-
forated with small, round pin holes (fig 37.). Any slight jarring
of the wood causes a fine flour-like powder to sift through these
holes. The presence of small piles of this powder beneath infested
timbers is evidence of infestation.

F-519936, F-519929

FIGURE 37.—Lyctus powder-
post beetle damage: A,
frass packed tunnels in ply-
wood; B, adult exit holes in
arm of the frame of an up-
holstered piece of furniture.

The southern lyctus beetle, Lyctus planicollis LeC., is one of the
most common of the destructive powder-post beetles in this coun-
try. It is widely distributed, but is most injurious in the Southern
States. Adults are usually black or almost black, have a wider-
than-long prothorax with distinctly separated punctations on the
disk, and are from 4 to 6 mm. long (fig. 38). The elytra are
almost three times as long as wide, bear straight lines of fine
long hairs, and are marked with double rows of elongate punc-
tures. Seasoned or partially seasoned wood of oak, ash, and
hickory are especially subject to attack. Freshly cut wood is not
attacked.

Eggs are laid in the open ends of pores of longitudinal vessels
in the wood, at depths of 4 to 7.5 mm. Hatching occurs within a
week or two. The larvae burrow through the wood; their tunnels
run parallel with. the grain and are filled with fine powdery dust.

129

a

F-519911
FIGURE 38.—Adult of the southern lyctus beetle, Lyctus planicollis.

Full-grown larvae are yellowish-white and about 3 to 5 mm. long.
Usually there is only one generation per year, but under certain
conditions, a generation may be completed in 3 to 4 months or less.
Heaviest emergence and egg laying usually occur during late
ae or early spring. Winter is spent in the larval stage (139,
VG)

Lyctus linearis Goeze, probably an introduced species, occurs
throughout the United States. In the eastern half of the country
it is commonly encountered in commercial hardwood products,
especially in seasoned hickory, oak, ash, and walnut. It is often
quite destructive. The adult is brown or reddish-brown and about
2.5 to 5 mm. long. The pronotum is almost square; the elytra are
wider than the pronotum and are almost three times as long as
wide. The interspace is composed of a single row of large, ovoid
punctations, separated by single rows of fine hairs on the carinu-
lae. There is a fringe of fine hairs on the fifth sternite in both
sexes.

Other species of Lyctus frequently encountered in eastern
United States include L. cavicollis LeC.; the western lyctus beetle,
L. opaculus LeC.; and L. brunneus (Stephens), the Old World
lyctus beetle. The following distinguishing characteristics should
be helpful in identifying them: (1) L. cavicollis—elytral striae
composed of two rows of punctations. A triangular patch of hairs
on each side of median line of apical portion of fifth sternite of
female. (2) Western lyctus beetle—two or more rows of puncta-
tions separated by carinulae bearing fine hairs in the elytral inter-
space. The fifth sternite of the male rounded, with a broad fringe
of short hairs on the apical margin. (8) Old World lyctus beetle—
all carinulae of the elytra bear single rows of fine sparse hair
with longitudinal rows of hairs between interspaces. The fifth
abdominal sternite of the female bears a “pencil” of hairs.

The genus Trogoxylon LeC. is represented in eastern United
States by three species. The adults are similar to those of the
genus Lyctus. They differ in having the femur of the metathoracic
leg enlarged and the pubescense of the elytra confused (285).

130

Trogoxylon parallelopipedum (Melsheimer), a common and
| destructive species, occurs throughout the United States. The
; adult resembles the adult of the southern lyctus beetle, except
| that it is somewhat smaller in average size and usually reddish in
color. It is distinguished by the prominent, acute, anterior angles
of the pronotum; a rounded, depressed postclypeus; and an
abruptly expanded antennal club. It breeds in the same type and
species of wood as the southern lyctus beetle.

The winter is spent in the larval stage. Heaviest adult emer-
gence and egg laying occurs in the spring. Adults of a second
generation emerge in late summer or fall and lay the overwinter-
ing supply of eggs. The adults crawl much more rapidly than do
those of the southern lyctus beetle. Large numbers of adults once
were observed on lumber sawed from green logs only two weeks
earlier. Others were observed emerging from lumber that had been
sawed from green logs only four or five months earlier (139). The
related species, Trogoxylon aequale (Wallaston) and T. caseyi
Lesne, have been recorded from Texas.

Minthea rugicolis (Walker), a cosmopolitan species largely
confined to the tropics, has been recovered at many points in the
United States from wood products imported from abroad.

FAMILY ANOBIIDAE
DEATHWATCH AND DRUGSTORE BEETLES

Members of the family Anobiidae breed in old dry sapwood and
heartwood of both hardwoods and coniferous trees. In the forest,
many species breed in twigs, while others attack large limbs and
trunks. Many species also attack finished timber products, such
as the girders, beams, and supports of old houses. Some also in-
fest wooden furniture and other wood products in homes. Some-
times timbers in structures that have escaped attack for more
than 20 years become heavily infested and practically destroyed.
Other materials in buildings are also attacked, including drugs and -
groceries. Certain species are known as deathwatch beetles be-
cause of the ticking sounds they make as they bore through the
wood.

Anobiid beetles are usually reddish brown to black, covered
with fine yellow hairs, and from 1.5 to 8 mm. long. The head is
hidden by the pronotum; the antennae are inserted on the sides
of the head in front of the eyes, and are 11-segmented with the
last three segments enlarged; the coxae of the front legs are
small; and the tarsi are five-segmented, with the first segment
longer than the second. Full-grown larvae are grub-shaped,
whitish and about 8 mm. long. The tops of some or all of the
segments are roughened with small points or hooks; the thoracic
segments are enlarged; and the body is covered with long hairs.
White (760) discussed the taxonomy of the species occurring in
Ohio. Simeone (651) discussed the distribution of several species.

As the larvae feed, they excrete small pellets of partly digested
wood. These pellets differ from those excreted by drywood ter-
mites in being smaller and in tapering toward each end. Pupation
occurs in cells in the wood, and the adults emerge through small,
usually oval holes about 1.8 to 3 mm. in diameter. Pellets usu-

131

SOR ES one el a SS ee”

ally drop out through these holes when the wood is struck. The
life cycle may require anywhere from 1 to 10 years for comple-
tion, depending on the moisture and fungus content of the
wood (460).

The furniture beetle, Anobium punctatum (DeG.), an introduced
species, occurs in southeastern Canada and in the Northeastern
States near the Coast. It has also been found in scattered loca-
tions south to the Gulf Coast of Texas and along the west coast
of California. Adults are dark brown and about 2.7 to 4.5 mm. long.
The metasternum is abruptly excavated in front and the elytra
bear linear rows of punctures. Infestations occur in furniture and
in the frames and flooring of buildings. Eggs are usually laid in
slits, cracks, or crevices in the wood surface, or in old emergence
holes. Damage appears to be most serious in old, partly decayed
wood, or in old wood such as in antique furniture. The life cycle
requires from 1 to 8 years.

Xestobium rufovillosum (DeG.), commonly known as the death-
watch beetle, is also an introduced species. It is now widely dis-
tributed in the Northeastern States where considerable damage
has been reported in many old buildings. Adults are robust and
dark brown, mottled with patches of yellowish pubescence, and
are about 6 to 7.5 mm. long. Both the sapwood and heartwood
of hardwoods and softwoods are attacked. Poorly ventilated oak
timbers in old buildings are especially subject to attack. The life
cycle requires from one to several years.

Hemicoelus carinatus (Say), sometimes known as the eastern
death watch beetle, is widely distributed in the Northeastern and
North Central States, but appears to be most abundant in New
England and New York. It also occurs in southeastern Canada.
The adult is about 3.4 to 6.8 mm. long. Infestations have been
recorded in ash and basswood flooring in Ohio and Michigan, elm
floor joists in Minnesota, maple and birch in Indiana, and in sills
and flooring in Connecticut. Damage resembles that caused by
Lyctus spp., but differs in that the emergence holes are larger
and the frass coarser. H. gibbicollis, the softwood powder-post
beetle, infests flooring and beams in buildings, and H. wmbrosus
Fall. is sometimes a pest of furniture.

Xyletinus peltatus (Harris) occurs from northern New York
to Michigan, southeastward to Florida and southwestward to
Arkansas. It attacks both the sapwood and heartwood of seasoned
hardwoods and conifers. Adults are reddish-brown to brown and
from 3.4 to 6.3 mm. long. Eggs are deposited in cracks or de-
pressions in the wood surface or under small splinters or pieces
of debris. Damage is similar to that caused by Lyctus species
except that it occurs in both the sapwood and heartwood. It is
usually confined to the springwood of each annual ring. Damage
to joists and flooring in damp buildings is occasionally serious. In
North Carolina it is the commonest species found infesting pine
floor joists in buildings. In closed, unoccupied buildings, it may
be so severe it causes the floors to collapse. X. lugubris LeC. has
been observed in dead oak twigs in Ohio.

®°Moore, H. B. 1964. Observations on the biology of Xyletinus peltatus
(Harris) (Coleop,: Anobiidae) with notes on morphology. Unpub. PhD
Thesis, Entomol. Dep. .. North Carolina State Univ.

132

Ernobius mollis L., an introduced species, has been recorded
from southeastern Canada and southward to Florida and Texas.
It breeds in coniferous trees. The larvae usually feed under the
bark, but may bore into the wood where the bark is thin. Bark-
beetle weakened trees are particularly susceptible to attack.
Adults are reddish brown and about 3.5 to 5.5 mm. long. EL.
tenuicornis LeC. has been reported from many Eastern States
and has been reared from loblolly pine twigs in North Carolina
(40). Adults are uniformly yellowish brown and are about 2.5 to
3.5 mm. long. #. granulatus LeC. occurs in several Eastern States
and has been reared from pine twigs and cones. Adults are reddish
brown and from 2.5 to 4.5 mm. long.

Petalhum bistriatum bistriatum (Say) occurs from Massa-
chusetts 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 (40). Adults are about 1.5 to 2.3 mm. long;
the head, pronotum, and undersurface 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 about 1.5 to 2.5 mm. long.

Ptilinus ruficornis Say is a rather common and injurious pest
of woodwork in houses and stored products. Adults are about 2.8
to 4.8 mm. long. Males are either completely black, or black with
brown elytra, and have reddish-brown to reddish-yellow ap-
pendages. Females have the pronotum bright reddish-brown, the
elytra reddish-brown to reddish-black, and the appendages red-
dish-brown to reddish-yellow. P. pruinosus Casey has been re-
corded on cottonwood in Ontario, Canada; Indiana; and Ohio.
Adults are similar in size and appearance to those of P. ruficornis.
P. pectinicornis (L.), an introduced species, was first recorded
from North America in New York in 1950 (650). Infestations
were found in joists of beech and in sugar maple timber. Female
adults are about 3.9 to 4.7 mm. long. The head, thorax, and ab-
domen are dark brown. |

Trichodesma gibbosa (Say) occurs from New Hampshire to
western Ontario in the North and southward to Florida and
Texas. Infestations have been found in sweet gum joists and in
studding in old historic buildings in Tidewater Virginia. Adults
are about 4.5 to 6.8 mm. long and covered with grayish-white to
nearly white pubescence.

Microbregma emarginatum emarginatum (Duft.) is generally
present throughout the Northeastern States where it breeds in
the outer bark of pines and hemlock. Adults are brown to dark
brown and about 4 to 5 mm. long. Trypopitys sericeus (Say) oc-
curs from Nova Scotia and southward to Florida, Texas, and Ari-
zona. It breeds in the dead branches of oak, cherry, and hickory,
and occasionally in flooring, sills, and furniture in buildings.
Adults are reddish-brown and about 5 to 6 mm. long.

Coelostethus notatus (Say) occurs in southeastern Canada and
southward to North Carolina and Mississippi. It has been re-
corded feeding in dead oak branches in Ohio. Adults are dark
reddish-brown and about 3.2 to 4.3 mm. long. Eucrada humeralis
(Melsh.) larvae feed beneath the bark of dead oak and beech

133

trees. It occurs from Quebec to Michigan in the North, and south-
westward and southeastward to Iowa and South Carolina. Nico-
bium castaenum (Oliv.) a European species, is known to occur
in Virginia, South Carolina, and Louisiana. Certain species of
Tricorynus (—Catorania) have been found attacking stored pine
cones in the South; others have been reared from twigs of pine
and sweet gum. Platybregmus canadensis Fisher attacks maple
and basswood flooring and elm timbers in buildings. It is known
to occur in New Jersey and Ohio.

FAMILY BOSTRICHIDAE
FALSE POWDER-POST BEETLES

The family Bostrichidae contains many species, the larvae of
which bore in wood and cause typical powder-post damage. The
majority of species breed in the sapwood of hardwoods, but a few
attack conifers. Some attack freshly cut and partially seasoned
woods with the bark on; whereas others infest wood that has
been cut for sometime. Damage consists of circular holes 3 to 9
mm. in diameter and irregular longitudinal tunnels filled with
frass or coarse dust.

Bostrichid beetles are reddish-brown to black and about 3 to
6 mm. long. They resemble adults of the family Scolytidae but
differ in having a tuberculate and rasplike pronotum, five-seg-
mented tarsi, and straight rather than elbowed antennae. The
antennal club is three- or four-segmented; the thorax is usually
hood-like and covers the head; and the posterior portion of the
elytra is frequently tuberculate. The femora and tibia are broad
and the latter are frequently toothed on one margin (254).

Xylobiops basilaris (Say) is a common species in the Eastern
and Southern States. The adult attacks recently felled or dying
trees, logs, or limbs with the bark on. Hickory, persimmon, and
pecan are most frequently infested, but several other hardwoods
also are attacked. Healthy trees growing in close proximity to
heavily infested trees are occasionally attacked, but almost al-
ways without success. The adults (fig. 39A) are basically black
and about 3 to 5 mm. long. The basal part of the elytra is dull
reddish or yellow and the posterior end is concave, with three
spines or tubercles on each side.

The adult bores through the bark and into the sapwood of its
host. Then it constructs a tunnel across the grain just under the
wood surface. These tunnels may completely girdle limbs and
trunks of small diameter. Eggs are deposited at intervals along
the sides of the tunnel. The larvae feed mostly in the sapwood,
but also to some extent in the heartwood. Their mines run parallel
with the grain and are packed with fine, white, powder-like dust
(fig. 839B). Feeding may continue until the wood is quite dry. As
a result, adults often emerge after it has been processed. The
winter is spent mostly as mature larvae. Hickory and persimmon
wood used in the manufacture of small wood products such as
shuttle blocks, mallets, and mauls, is occasionally seriously
damaged.

134

nn

F-480474, 480481

FIGURE 39.—Xylobiops basilaris, a false powder-post beetle: A, adult; B,
larval damage (note openings to frass-frilled larval tunnels in cross-
section of a persimmon log).

The bamboo powder-post beetle, Dinoderus minutus (F.), a very
destructive pest of bamboo in the West Indies, occurs in Florida
and Louisiana. Adults are reddish-brown to brownish-black and
about 2.5 to 3.5 mm. long. The wing covers are often redder than
the rest of the upper surface. The female bores into bamboo culms
at breaks in the rind, usually at cut ends and trimmed nodes, and
then bores a tunnel across the grain. Eggs are deposited in the
exposed ends of tubular vessels of the fibrovascular bundles. The
interior of infested material is often reduced to masses of
powdered wood and fibers. This species is also intercepted fre-
quently in infested bamboo at ports of entry in this country.

135

In 1967, a heavy infestation of the Oriental bostrichid, Hetero-
bostrychus aequalis (Waterhouse), was discovered in oak and
mahogany boards in a mill yard in Florida (728). This is the
most common of the larger bostrichids known to attack packing
cases, boxes, plywood, and sapwood in furniture in India. It is
widely distributed in southeast Asia and has been recorded at-
tacking the wood of more than 85 species of trees. The adult is
reddish-brown to black, shiny, and from 6 to 13 mm. long. The
elytra are densely, deeply, and coarsely marked with rows of
punctures. Eggs are deposited on the rough surfaces of lumber or
logs, in cracks or holes, and in short tunnels made by the female.
The larvae, some of which may attain a length of 11 mm., exca-
vate winding tunnels up to 6 mm. in diameter and one foot in
length in both hardwood and softwood timber. The life cycle
requires from 1 to 6 years.

Stephanopachys rugosus (Oliv.) occurs in eastern United
States, mainly east of the Mississippi River, and attacks freshly
cut or recently milled pine. Adults are reddish-brown, have
roughened and dull, rusty-appearing bodies, and are about 3 mm.
long. The larvae normally reach maturity in one year, but may
require up to 5 years in seasoned wood. Infestations are occasion-
ally found in floor joists with the bark on in recently constructed
buildings. S. substriatus (Payk.) occurs in most of the Northern
States. It attacks Douglas-fir, hemlock, and all species of pines
and firs. Infestations have been reported in furniture and in
lumber used in buildings. Oak and hemlock tanbark is also subject
to attack and damage. S. densus (LeC.), S. ertbratus (LeC.), and
S. hispidulus (Casey) breed in pine.

The apple twig borer, Amphicerus bicaudatus (Say), breeds in
dying wood such as large prunings, exposed roots of maple, and
diseased and dying branches of most shade and fruit trees. It
occurs throughout most of the eastern half of the United States.

A number of other species of Bostrichids are also found in
eastern United States. Lichenophanes armiger (LeC.) breeds in
various hardwods, chiefly dead and dying oak. L. bicornis (Web.)
has been recorded breeding under dead bark and in the dead wood
of sycamore, hackberry, oak, pecan, hickory, apple, beech, and
elm. Lumber and other stored stock of oak, hickory, sycamore,
and black locust are also infested occasionally. Scobicia bidentata
(Horn) commonly infests freshly cut wood and lumber of hickory,
elm, oak, chestnut, hackberry, and sassafras in the Midwest.

Polycaon stoutw (LeC.), a destructive western species, is fre-
quently shipped to the Eastern States in infested wood. It is
especially injurious to veneer plywood which is infested while in
storage. Adults are coal black, broad-headed, and from 12 to 25
mm. long. Dendrobiella aspera (LeC.), D. sericans (LeC.), and
Xyloblaptus quadrispinosus (LeC.) are frequently encountered
in hardwoods shipped to the Eastern States from the southwest.

FAMILY PTINIDAE
SPIDER BEETLES

The beetles in the family Ptinidae are small, spider-like, and
from 2 to 4mm. long. The head and pronotum are much narrower

136

than the elytra; the legs are long; the antennae are long, filiform,
and inserted on the front of the head; and the elytra are usually
very convex and shiny. The majority of our native species breed
in old grass roots or in the bark of dead twigs of trees. Certain
other species are destructive in warehouses and museums where
they attack collections, to seasoned wood or logs left lying in the
woods too long, to wood in old buildings, and to stored products.

The white-marked spider beetle, Ptinus fur (Linnaeus), a cos-
mopolitan species, is frequently found damaging dried vegetable
and animal materials in warehouses and museums. Infestations
also occur in pine and oak woodwork in these structures.

The brown spider beetle, Ptinus clavipes Panzer, attacks pine
boards in old buildings.

FAMILY BRUCHIDAE

SEED BEETLES

Members of the family Bruchidae are distinguished by their
compact and usually oval bodies, their small heads which are pro-
longed into beaks, and their short wing covers which leave the tip
of the abdomen exposed. The larvae are quite small and feed al-
most entirely in the seeds of plants.

Amblycerus robiniae (F.) is occasionally a pest in eastern
United States. The female deposits her eggs on the pods of honey
locust 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 red-
bud and Caryobruchus gleditsiae (L.), in palmetto.

FAMILY CHRYSOMELIDAE
LEAF BEETLES

The leaf beetles comprise one of the larger families of Coleop-
tera, with more than 1800 species occurring in the United States
alone. The adults usually are medium-sized or small, short-bodied,
and more or less oval. The legs are generally short; but in some
species the femora of the hird 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 scale-like hairs.

Leaf beetle larvae are usually soft-bodied and frequently have
highly pigmented or well-chitinized sclerites on the integument.
They vary greatly in shape, depending on their feeding habits.
The head usually protrudes, except in the leaf miners, 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 leaf miner, 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

137

gregariously, whereas leaf mining larvae usually feed singly, each
within its own mine. Adult feeding is characterized by the pres-
ence of holes eaten all the way through the leaf, or by skeletoni-
zation usually of the lower surface of the leaf. Free-living larvae
elther fasten themselves to the surface of a leaf to pupate, or move
to the ground to do so. Leaf mining larvae pupate either within
the mine, or they vacate it and pupate in the ground. This family
has been treated by Leng (451), Leng & Mutchler (452, 453), and
Wilcox (264). A few of the more common eastern species are
discussed.

The cottonwoed 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 yel-
low 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. Eggs are deposited in groups of 15 to 75 eggs each on the
undersides of leaves. The young larvae are gregarious and, feed-
ing side by side, skeletonize the leaves. Later, they feed separately
and consume the entire leaf, excepting the larger veins. At ma-
turity they attach themselves to leaf surfaces, the bark, or to
weeds and grass beneath the trees to pupate. There are several
generations per year. Severe infestations occur occasionally and
cause considerable damage.

Chrysomela interrupta F. and C. crotcht Brown occur commonly
throughout the Eastern States. The adults and larvae of inter-
rupta feed on the leaves and at times on the tender bark of alder;
adults and larvae of crotchi feed on poplar. 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 (Mueller) (fig. 40), an
introduced species first recorded in North America at Baltimore,
Md. well over a century ago, now occurs throughout most of the
United States. Its hosts are all species of elm. Irn the Northeastern
States, American elm is often severely attacked and seriously
damaged. Farther west, Chinese 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 the
pupae are about 5 mm. long.

138

Wiidamagesttmtene—aa—y
~B, Elm leaf showing
é Woeimiesdina sonedia
mn skeletonization, and ee
tion; a.,egg-mass; Bye a f
ltoarv@e;c, pupa; and
Ck edelgehn leaf

COURTESY AGR. RES. SERV.

FIGURE 40.—Elm leaf bettle, Pynhalta luteola.

139

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 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 un-
folding leaves. Egg laying begins late in May or early June, each
female laying from 400 to 800 eggs (725). Hatching begins in
about one week and the larvae feed for 2 or 3 weeks on the under-
surfaces 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 and a partial third generations. Usually the first gener-
ation is the most injurious. Beetles maturing in the summer begin
entering hibernation quarters on or near the tree on which they
fed as early as August. :

Shade tree elms are often heavily defoliated, whereas those
growing in the forest are usually not seriously infested. The
native parasite, Tetrastichus brevistigma Gahan occasionally ex-
erts a considerable degree of control in the Northeastern States
(62). During damp weather the fungus, Sporotrichum globuli-
ferum, is also occasionally effective in control. Some degree of
direct control of beetles hibernating in attics can be obtained
locally by trapping them as they are leaving their hibernation
quarters in the spring. Most of the openings that admit light
should be covered with black paper. The remaining openings
should be covered with tightly stretched cellophane attached to
wooden frames. The inner surface of the cellophane should be
covered with a sticky substance such as Tanglefoot.

Pyrrhalta cavicollis (LeC.) occurs in southern Canada and in
the Eastern States southward through the Alleghenies and Appa-
lachians 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 larva return to the ground and pupate in the leaf mold or
soil. There is one generation per year. P. decora decora Say and
P. tuberculata (Say) feed on willows.

The elm calligrapha, Calligrapha scalaris (LeC.), has been re-
ported from eastern Canada and from several widely distributed
points in 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-colored with a black line down the middle of the dorsum.

140

The winter is spent in the adult stage in bark crevices, in
sheltered places around the base of the tree, or in the top 1 or 2
inches 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 crawl
down the trunk in search of places to pupate and spend the
winter. In heavy infestations, they frequently occur by the thou-
sands on the trunk or under the larger limbs. American elm is
sometimes heavily defoliated and damaged in the Midwest (186).

Calligrapha multipunctata bigsbyana (Kby.) sometimes com-
pletely 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 pro-
notum and most of the elytra are pale yellow.

The larger elm leaf beetle, Monocesta corylt (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 (8). Its hosts are recorded as
native and Japanese elms, river birch, pecan, hawthorn, and
hazelnut. 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 20 mm. long.

Winter is spent as a full-grown larva in a cell a few inches
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 genera-
tion per year. Outbreaks occurred in river bottoms in Piedmont
sections of the Carolinas and Alabama in 1964.

The basswood leaf miner, Baliosus ruber (Weber) (=B. ner-
vosus (Panzer), occurs in Canada and throughout most of the
eastern United States. Basswood is the favored host, but oak,
maple, willow, birch, hop hornbeam, 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 (357). 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.
41). 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 of miners per year.

The alder flea beetle, Altica ambiens subspecies alni Harr., oc-
curs 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

141

F-506746
FIGURE 41.—Leaf of basswood
mined by the basswood leaf
miner, Baliosus ruber.

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,
over-wintering 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 (794). In the North
there is one generation per year; in the South there may be two.
The related species, A. carinata Germ. feeds on elm and A. sub-
plicata (LeC.), on willow.

The pine colaspis, Colaspis pint Barber, occurs from Maryland
to central Florida and westward to east Texas. It feeds mostly on
southern pines but also occasionally on cypress 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.
Full-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. Pupa-
tion 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 in 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. Nichols!® discussed the biology of the species in
Louisiana.

10 Nichols, H. W. 1962. The biology, distribution, and insecticidal control
of pine colaspis (Colaspis pini Barber) in Louisiana. (Master’s thesis,
Louisiana State Univ.)

142

The locust leaf miner, Odontota (=Xenochalepus) dorsalis
(Thunb.), occurs in southern Canada and throughout most of
eastern United States. Its favored host is black locust, but several
other tree species such as apple, birch, beech, wild 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 posteriorally. The elytra are also deeply pitted, and each
elytron bears three longitudinal ridges. A full-grown larva is
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 eggs each. 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 single larva may mine
several leaves. Pupation occurs in the mine, and there is one
generation per year.

Outbreaks of the locust leaf miner occur practically every year
somewhere within its range and black locust trees on tens of
thousands of acres are often defoliated. 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 (381). The eulophid parasite, Clostocerus tricinctus Ash-
mead, is reported to have destroyed over 50 percent of the pupae
in West Virginia infestations (747). Direct control is seldom at-
tempted in the forest, but is sometimes desirable in parks and
other recreational areas.

The imported willow leaf beetle, Plagiodera versicolora
(Laich.) (fig. 42), an introduced species, was first reported in
this country from Staten Island, New York in 1915 (366). 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 in color, and sometimes tinged with red or bronze. Full-
grown larvae are almost jet black and about 5 mm. long. Rows of
protuberances run both transversely and longitudinally across and
along the body.

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
skeltonizing 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 and a partial fourth generations per year have been re-
corded in Massachusetts. Additional generations probably occur

143

COURTESY CONN. AGR. EXPT. STA.

FIGURE 42.—Adults, larvae, and pupae of the imported willow leaf beetle,

Plagiodera versicolora. Note skeletonization of leaves by larvae and holes
eaten in leaves by adults.

farther southward. 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
sieboldi (Ratzeburg) (203). Extremely cold winters are also
fatal to adults not well protected. .

Systena marginalis (Ill.) feeds on oak in the midwest and on
cypress in northern Florida and southern Georgia. The adult is
dull pale yellow in color 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 cypress leaflet to turn red and die. The beetles occur in
large swarms which tend to move about, spending only one to
three days in any one place. A single swarm may encompass more
than a dozen trees.

Zengophora scutellaris Suffr., 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 re-
mainder 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 large
black blotch mines. When they become full-grown they vacate
their mines and drop to and enter the ground. Here they construct
cells several inches below the surface in which to pupate. Trees
heavily fed on by both larvae and adults may be completely
defoliated.

144

Glyptoscelis pubescens (Fab.) occurs primarily east of the Mis-
Sissippi River from Canada to Georgia and feeds on various
species of pines and, reportedly, spruce and hemlock. The adult is
oblong-oval, robust, broadly rounded posteriorally, dark brown
with a brassy or golden sheen, and is from 7 to 10 mm. long. It is
also sparsely clothed with an intermixture of white and brownish
hairs. Adults feed on the edges of pine needles, causing them to
turn brown. Damage has been reported in seed orchards of Vir-
ginia, shortleaf, and 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 ex-
cept that it is smaller and its upper surface is shining brown.
Krause (440) reported on the genus Glyptoscelis in the United
States and Canada.

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: Pachybrachys peccans Suffr.—hickory and birch, P.
tridens (Melsh.)—willow, P. othonus (Say)—ash and elm, and
P. carbonarius Hald.—oak; Anomoea laticlavia (Forstr.)—
honey and black locusts, elm, live oak, and mimosa; Tymnes
tricolor (Fab.)—oak, walnut, and ironwood; Bassareus litur-
atus (F.)—hickory; Derocrepis aescult (Drury)—buckeye; Pla-
giometriona clavata (Fab.)—sycamore, basswood, and oak;
Chlamisus platant Brown—sycamore; Xanthonia decimnotata
(Say)—oak, beech, and elm; Paria sexnotata (Say)—red cedar;
P. quadrinotata (Say)—walnut and mountain ash; and Syneta
ferruginea (Germ.)—birch and oak.

FAMILY LUCANIDAE
STAG BEETLES

Stag beetles are distinguished by their very large mandibles
which, in the males of certain species, are branched like the ant-
lers 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 Fab., the most familiar
species, infests dead stumps in the South. Adults are large, fear-
some insects, up to 60 mm. long. Male mandibles are branched
and are more than half as long as the body. Pseudolucanus capre-
olus (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 Web., Ceruchus piceus
(Web.), Dorcus parallelus Say, and Sinodendron rugosum Mann.
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 1400 species, the majority occurring 1n. the eastern
half of the country. Depending on their feeding habits, members

145

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, decom-
posing 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 3 to 6 or 7 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 posteriorally, bent in the
shape of a crescent, and they have well-developed legs.

The genus Phyllophaga Harris is represented in the United
States and Canada by more than 100 species, the majority of
which occur in the eastern part of the continent (462). 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. 48 A). The wing covers are smooth and shiny or are
covered with short hairs. The antennae are lamellate and end in
3-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. 48 B). 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
(616). 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 3 to 7
inches, 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 tem-
peratures and frost levels. Pupation takes place in the soil at
depths of a few inches to a foot 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
(684).

Phyllophaga larvae, or white grubs, have caused heavy losses
in forest nurseries and plantations in the South, East, and Central
and Lake States during the past several years. 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 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.

146

F-494655, F-494654
FIGURE 43.—Phyllophaga species: A. adult; B, white grubs.

Phyllophaga luctuosa (Horn) occurs primarily along the At-
lantic 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,

147

birch, and loblolly and longleaf pines. The larvae are often de-
structive in nurseries and, probably, plantations.

Phyllophaga tristis (Fab.) occurs throughout 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 (Burm.) occurs in the South Atlan-
tic and Gulf Coast States. It is especially common in the Sand
Hills of South Carolina. Adults are reddish-brown to black, with
the surface shining and slightly pruinose, or dull smoky and are
12 to 18 mm. long. They feed on pines, preferably loblolly and
longleaf, and sometimes oaks and persimmon. The larvae have
caused serious losses in pine nurseries and plantations in South
Carolina.

Phyllophaga rugosa (Melsh.) 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 nur-
series in the Lake States.

Phyllophaga crenulata (Froel.) occurs throughout eastern
United States. Adults are brown, with a covering of short, recum-
bent hairs, and are about 17 to 20 mm. long. They feed on a wide
variety of hardwoods, preferably persimmon, hickory, basswood,
willow, birch, and horse chestnut. The larvae are often serious
pests in coniferous nurseries in the Lake States.

Phyllophaga forstert (Burm.) occurs generally throughout eas-
tern 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 (LeC.) 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 implicta Horn occurs mostly in the Mississippi
and Ohio River Valleys. Adults are orange-brown to brown (with
the head and thorax darker), shining, 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.

The genus Polyphylla Harris is represented by a number of
species in 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

148

seedlings, but they prefer the roots of grasses. They would appear
to be potentially harmful in nurseries and plantations. P. ham-
mendi LeC. occurs in the western part of the Central States and
has the interesting habit of depositing its eggs in rotten wood.

The genus Serica MacL. 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
(1ll.), which is often abundant; S. tristis LeC., which has been
known to defoliate spruce in plantations in southern Canada; and
S. vespertina (Gyll.) and S. wntermixta Blotch.

The genus Dichelonyx Harris is represented in eastern 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 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 re-
quires 2 to 8 years. Saylor (633) revised the genus.

Dichelonyx albicollis (Burm.) is a well-known species. It has
been recorded from New Jersey, Michigan, and Ontario and feeds
both during the day and night on the needles of pine, preferably
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 black birch and alder. The adult is
smaller and somewhat darker than the adult of D. albicollis. A
third species, D. subvittata LeC. has been recorded feeding on
oak, hazel, and pine from New England to the Lake States and in
southern Canada.

The genus Diplotaxis Kirby contains a number of species that
feed mostly on conifers, preferably 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 planta-
tions 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
(Germ.) 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 in-
troduced 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, horse chestnut, willow, boxelder, and Ailanthus. Young
pines, hemlocks, and yews in nurseries are defoliated occa-
sionally and seriously injured. The roots of rhododendron and

149

azalea are damaged occasionally by the larvae.

The rose chafer, Macrodactylus subspinosus (F.), is widely dis-
tributed in eastern United States. Adults are tan to reddish-
brown, densely covered with dull yellow scales or hairs, and have
long, reddish-brown legs. They feed on a wide variety of hosts,
including many species of forest and shade trees (fig. 44). 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
where they are abundant.

The European chafer, Amphimallon majalis (Raz.), an intro-
duced species first recorded in New York in 1940, now occurs in
several Eastern States and southern Ontario. The adult is oval-
shaped, light brown or tan, and about 14 mm. long. A distinguish-
ing 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.

The pine chafer, Pachystethus 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 in color and

COURTESY CONN. AGR. EXPT. STA.
FIGURE 44.—Birch leaf skele-
tonized by the rose chafer,
Macrodactylus subspinosus.

150

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 was recorded in several thousand
acres of young loblolly pines in southeastern North Carolina in
1959 and 1960. 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 several inches 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 South during recent years. The majority were
short-lived and limited in size, but several were quite extensive
and lasted for several years. Pachystethus 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 Newm., an introduced
species first recorded in North America near Riverton, New Jer-
sey in 1916, now occurs in all or parts of at least 14 States from
New Hampshire and Vermont to North Carolina and Ohio. 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 iast abdominal segment arranged in the
shape of a V.

The Japanese beetle feeds on the foliage, flowers, and fruits of
a wide variety of plants. Many species of forest and shade trees
are subject to defoliation, especially Japanese and Norway
maples, horse chestnut, sycamore, gray birch, walnut, Lombardy
poplar, basswood, mountain-ash, and American, English, and Chi-
nese elms. Feeding is usually confined to young, tender leaves.
Damaged leaves may be skeletonized or they may also have large,
irregular 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 1 to 4 inches 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 two years. bi

Disease pathogens, especially the bacterium, Bacillus popilliae
Dutky, often destroy large numbers of the larvae. Several intro-

151

duced parasites, Tiphia vernalis Rohwer and T. popilliavora
oe in particular, also exert a considerable degree of control
LAS) )

Cotalpa lanigera L., the goldsmith beetle, occurs throughout
eastern United States and feeds on the foliage of various hard-
woods, 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 elytra being lemon yellow, and the head and thorax burnished
golden. The venter is greenish to copper colored and covered with
whitish wool. The legs are reddish-yellow. Full-grown larvae are
about 48 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 in-
cluding young conifers. Adults have been observed fairly com-
monly 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 eastern United States, most commonly in the Atlantic
and Gulf Coastal 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 under-surface 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 fe-
males deposit their eggs in soil rich with organic matter. The
larvae feed on the organic matter during the remainder of the
season and then enter hibernation. 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 sometimes injured.

The genus Dynastes Kirby, which contains the largest known
beetles, is represented in eastern United States by only one
species, the eastern hercules beetle, D. tityus (L.). It has been re-
corded from New York, Indiana, and Arkansas south to the Gulf
of Mexico and breeds in decayed hardwood stumps and logs. It is
also found occasionally in cavities in the bases of living trees. The
adult is very large, from 40 to 60 mm. long and is usually green-
ish-gray or tan, except for mottlings or blotch-like areas of black.
The male is armed with a large pronotal horn which projects
forward and almost meets another horn which 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 Xyloryctes Hope is represented in eastern forests by
3 or 4 species, one of which, the rhinoceros beetle, X, jamaicensis
(Drury), is common in hardwood stands in the South. The adults
are robust, shiny beetles, about 28 mm. long. They are dark chest-
nut 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.

152

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 MacLeay occur in
eastern forests, one of which, P. punctata (L.), is 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 prono’:m. 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 Polymoechus LeC. is represented in eastern forests
by several species. The most common and widely distributed one
is P. brevipes LeC. The larvae are found most commonly in de-
cayed 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 Serville is represented in eastern forests
by at least four species, two of which, O. eremicola Knoch and O.
scabra (Beauv.), 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. eremi-
cola is dark chestnut brown, smooth and shiny, whereas O. scabra
is bronzy-purple black and rough in texture. They are approxi-
mately 25 mm. long. When handled, they emit a strong leather-
like odor.

The genus Trichiotinus Casey contains several species, the
adults of which are frequently seen around flowering trees and
shrubs, wild 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 dead logs and stumps.

FAMILY ELATERIDAE
CLICK BEETLES

Click beetles are so-called because of the presence of a spine on
the prothorax which 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 predomin-
antly phytophagous, but a few are predaceous on other insects,
such as wood borers. Vegetable feeding species usually confine
their attacks to dead and often well-decayed wood and are of little
economic importance. Dietrich (192) reported on the species
occurring in New York State. ;

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

153

ee and are only about 20 to 40 mm. long. They also have eye
spots.

Other predaceous species include Lacon discoidea (Web.) and
L. avita Say which are found in pines, and Hemicrepidus bilobatus
(Say) which occurs in hickory. Ctenicera triundulata (Rand.)
and C. nitidula (LeC.) have been observed feeding on cocoons of
the European spruce sawfly in Canada (541).

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 11-jointed and generally ser-
rate 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 18 mm. long.

Most species spend the winter in the larval stage. Others over-
winter as pupae or as adults in pupal cells in the bark. The larvae
travel down the tunnels of their hosts eating one larva after an-
other. Some are capable of consuming several times their own
weight of these larvae (91). A few of the more important preda-
tors of forest insects are discussed below.

Chariessa pilosa (Forst.) 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. 45) is a wedge-shaped, flattened beetle about 6 to 138
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.

Thanasimus dubius (Fab.) is one of the most important pred-
ators of destructive bark beetles in eastern United States. 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 cross bands of whitish hairs. The larva is
elongate, fusiform, and purplish with brown sclerotized areas.

Winter is spent in either 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
(BiG 27a).

154

F-519949

FIGURE 45.—Adult of Chari-

essa pilosa, a predator of
wood-boring larvae.

E’noclerus nigripes (Say) larvae feed on bark beetles in coni-
fers, and on wood borers in hardwoods. Adults are brightly col-
ored 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 cross bars on the wing covers. The
larvae are similar to those of Thanasimus dubius. The black-
bellied clerid EH. lecontet Wolc., 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
is 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) is an important enemy of round-
headed and flat-headed borers in hardwoods in 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 an-
tennae which are reddish-yellow mixed with black. The larvae
are purplish.

Tarsostenus univittatus (Rossi) is an important predator of
powder-post 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

155

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, slen-
der, and biack except for a spot on the head and the sides of the
thorax, which are golden yellow.

Many other clerids are also predaceous on various forest in-
sects in the Eastern States. Priocera castenae Newm. feeds on
bark beetles in conifers; Phlogistosternus 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. Knull (432) discussed the clerids occurring in Ohio.

FAMILY DERMISTIDAE
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.

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 beetie,
Dermestes maculatus DeGeer, 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 sur-
face 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 for-
est and shade trees. More than 150 species and varieties have
been recorded east of the Mississippi River, many of which are
discussed by Franklin and Lund (264). The larvae of all species
are borers, and they feed in all 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 sawlogs, often seriously
reducing or destroying their uesfulness as lumber.

Buprestid beetles are usually somewhat flattened or oval-
shaped, and are beautifully marked or metallic-colored. The head
is strongly deflexed and is inserted into the prothorax to the eyes.
The antennae are serrate, 1l-jointed, and inserted on the front;
the prosternum is prolonged behind and fits into the meso-
sternum; the elytra usually cover the abdomen; and the first two
of the abdominal sternites are fused.

156

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 sec-
ond and third thoracic segments (fig. 46). Leaf-mining 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.

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 46.—Typical larva of a 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 winding and usually oval in cross section.
Eventually, they terminate in elongated pupal cells which are
connected to the surface by short, oval exit holes. A characteristic
of the mines is that they usually are packed tightly with layers.
of sawdust-like 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 1 or 2 years, but in certain
Species it takes many years.

Key to the Adults of the Eastern Genera of Buprestidae

1. Hind coxal plates scarcely widened internally |... 2
Hind coxal plates distinctly widened internally, front
manreine straight, hind margin oblique «2..0.0200000022. 10
Zea laeOUNOKa XatLUNCAtetAt DASC ie) Lill ee a EE 3
Prounorax lobedvatibase ns. a ee ee 5
3. Mesosternum scarcely visible; breeds in small branches of
DES ose macesta shh a ta Cen BU nt Mastogentus
Mesosternummemarcinate ss 60... eo Be. 4
A-SOCULeM INGISUINGE eat 2) Re aes... Acmaeodera
Scucellum= plainly indicated ........... 220.002 ee Ptosima
5. Antennae not received in grooves on underside of
FORO GINO Ti casxet tee ka tae bid Pee es Ao oe eel Raa al oa ree aloe 6

Antennae received in grooves on underside of prothorax 7

157

10.
ble

12.

13.

158

First joint of hind tarsi scarcely pecan gall formers

Ont AIdeis4 ix uk. eonen |. Renee cine Eupristocerus
First joint of hind tarsi as long as 8 following joints
Agrilus

Tarsi over half as long as tibiae ... _... Paragrilus

Tarsi;zmuch/ shorter than tibiaet 4. 2.) re ees eee
Scutellum large; larvae leaf miners in certain leguminous

plants 1c. Datos. ae oro Pachyschelus
scutellum:smiall: aus)! ar} See ve Carre ee eee
Prosternum with posterior cavity; larvae leaf miners in

Various species of. oak =... e a eee Brachys
Prosternum without cavity; leaf miners in Scirpius and
Other: alliediplan ts pecs werimte renee ene iete tian neti Taphrocerus

Antennal pores scattered on both faces of serrate joints
Antennal pores concentrated in pits on these joints ...
Apical 1 of elytral margin entire, or finely serrate;

borers in pine stumps and logs _.................Chalcophora
Apical 14 of elytral margins strongly serrate; borers in
hardwood trunkse. = ee ... Chaleophorella

Front not narrowed at insertion of antennae; eyes
scarcely approaching, often distant on vertex .

Front narrowed at insertion of antennae; eyes oblique,
close together on vertex _..

Prosternal spine obtusely angulated behind prothracic
coxae; epimera of metathorax not covered by abdomen

Prosternal spine acutely angulated behind prothoracic
coxae; epimera of metathorax partly concealed by
abdomen ...... 5 ee

Prosternal spine obtusely rounded at apex.

Prosternal spine acute at apex; larvae heartwood borers

in oak and ironwood . : if | Cinyra
Mentum entirely corneous: |
Anterior edge of mentum coriaceous Buprestis
Elytra striate ...... es
Elytra not striate . Trachykele
Scutellum small, round, oval | se Dicerca
Scutellum large, trapezoidal; pronotum with smooth

median line _.. Poecilonota
Anterior edge of mentum coriaceous Melanophila

Mentum entirely corneous

Base of thorax trucate . |

Base of thorax sinuate Xenorhipis

Last ventral segment of abdomen punctured like the
segments preceding _.. Anthaxia

Last ventral segment of abdomen much more densely
punctured than the preceding segments Agrillaxia

Hind tarsi with lst joint considerably longer than 2nd;
3rd joint not prolonged at sides; sapwood and bark
borers Chrysobothris

Hind tarsi with 1st joint only slightly longer than 2nd;
38rd joint prolonged at sides; heartwood borers in
various hardwoods Actenodes

cl
12

13
21
14
18
15

16
bye

19
20

Key to the Larvae of the More Important Genera of Buprestidae

Leaf miners, larvae spindle- or wedge-shaped ........ ees
Twig or bark borers or occasionally gall makers; larval
form elongate, parallel, last segment ending in a pair
OL SPINS rye ee thes eho a he cle pe Td Oh (0 ae
Wood or bark borers; larvae hammer-headed or pestle-
Shaped HOTSPINES iP Ee es ieee Bk) ae ae | eae
Dorsal plate of prothorax marked with an inverted Y,
WAS CC URS sD M GN a mimes eek sue iai ers Wie gata eN nya sola real
Dorsal plate of prothorax bearing only a single median
impression or groove ..........
Dorsal plate of prothorax bearing chinous rugosities |
Dorsal plate of prothorax smooth |
Rugosities on prothoracic plates strongly developed form-
ing ridges; margins of plates distinct. _.............
Rugosities on plates not forming ridges, dispersed; plates
IMGIStINCT On MAnGINS. oe ar oie Li eee
Dorsal prothoracic plate marked with a distinct inverted
Ve breeds in conifers (es Chalcophora
Dorsal plate marked with an inverted V or U; breeds
INtMALEWOOUSIC= eek oe eee Chalcophor ella
Asperate areas of dorsal plate longer than wide; ventral
area completely bisected by median groove; first ab-
dominal segment smaller than second; mandible usually
bridentates = eke el ea Melanophila
Asperate areas of dorsal plate wider than long or of
approximately the same dimensions; ventral area not
completely bisected

Dorsal plate marked by an treerted ¥ ¢ or o pore eu dedi on
the anterior half by the asperities suggesting a hood;
ventral plate marked by a median groove extending
from posterior margin forward but not completely
bisecting plate; mandible usually tridentate, labrum
usually: tridentate 08 ies Bupr estis

Dorsal and ventral asperate plates nearly circular; ven-
tral bearing a median groove, which extends backward
from the anterior margin about 24 of the distance;
mandible usually bidentate; labrum entire

Chrysobothris

Metathorax bearing a dorsal and ventral pair of ampullae,

ventral groove not bisecting prothoracic plate
Anthaxia
Metathorax normal; ventral groove bisecting plate

Anterior end of dorsal inverted Y or V marking simple
Anterior end of both dorsal and ventral markings with
broad reticulated ends...

10. Dorsal marking an inverted Y; Brecde in ee _Cinyra

ie:

Dorsal marking an inverted V; breeds in willow and
DOD AR ies ce eee! | _. Poecilonota

Dorsal marking an “Eowertiad We oath a broad reticulated

apex; mandible tridentate; breeds en in hard-
WV OOUSist titre si: | We kee minnie Dicerca

13

14

or) On 00 > bo es) i)

10
11

159

Dorsal marking an inverted Y, the apex surrounded by

a depressed shiny diamond-shaped area; mandible tri-
dentate; breeds in conifers 0:08 Trachykele

12. First abdominal segment smaller than the rest; plates of
prothorax whitish opaque; prothoracic grooves dark

brown: breeds intredbudims:. 4a Serer Ptosima
First abdominal segment broader than the second
Acmaeodera

13. First segment as broad or slightly broader than the
others; body gradually tapering to the twelfth seg-
ment; slightly wedge-shaped 20... Brachys

First segment narrower than the others; body tapering
both ways from about the middle, more acute at the
posterior end; spindle-shaped ...................... Pachyschelus

14. Dorsal plate marked by 2 moderately separated dark-
brown lines which converge anteriorly; breeds in alder

Eupristocerus
Dorsal plate marked by a single median bisecting line;
attacks various hardwoods) 222-2 === Agrilus

Acmaeodera pulchella (Hbst.), the flatheaded bald cypress sap-
wood borer, breeds in bald cypress in the 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 prothoracic plates are marked by brownish median
grooves or lines. The sapwood of dead and dying cypress trees
and recently cut cypress logs is subject to severe attack and dam-
age. The removal of unseasoned logs from the woods before the
adults fly in the spring, or the girdling of trees in the fall that
are to be felled in the spring, should aid in the reduction of
damage.

The two-lined chestnut borer, Agrilus bilineatus (Web.), occurs
in southern Canada and throughout Eastern United States. It
breeds in various hardwoods, preferably chestnut and several of
the oaks. Trees weakened by drought, defoliation, or other ad-
verse factors are usually attacked. Adults are about 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 prepupal stage in cells constructed in the
outer layers of the sapwood and sometimes in the bark. 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. 47), 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

160

as the trees continue to weaken. There is a 2-year life cycle in the
North; in the South only one year is required.

Control under forested conditions is usually impracticable ex-
cept for such management practices as are helpful in maintaining
or promoting tree vigor. The watering or fertilizing of valuable
shade trees should be helpful in protecting them from attack.

The bronze birch borer, Agrilus anxius Gory, apparently occurs
throughout most of the range of birch in Canada and the United
States. It breeds in various birches, paper and yellow being pre-
ferred. The adult is deep green-bronze and about the same size
and shape as the adult of the two-lined 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 three weeks before egg laying begins. Eggs are deposited
singly or in small groups beneath loose flakes of bark and in

COURTESY OF DUKE UNIV. SCH. OF
FOREST.

FIGURE 47.—Galleries of the
two-lined chestnut borer.
Agrilus bilineatus, in the
inner bark of black oak.

cracks and crevices in the bark, mostly on unshaded parts of the
tree. Young larvae bore directly through the bark to the cam-
bium 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 sub-freezing tempera-
tures. In the North, 2 years are required to complete the life
cycle; in the South there is one generation per year. Barter (34)
reported on the biology of the bronze birch borer in New
Brunswick.

The bronze birch borer prefers weakened or injured trees
(471). Damage may be extremely severe in stands of such trees.
Weakened residual trees following logging and individual shade
and ornamental trees weakened by drought or other factors are
often seriously injured. During the past several years, enormous
volumes of birch suffering from ‘die back’’ were killed in the
Northeastern States and Canada. In Canada, at least 1 million
cords were killed during 1939 alone (108). The most effective
control measures are those designed to promote and maintain
host vigor.

The bronze poplar borer, Agrilus liragus B. & B., breeds in
poplar in Canada and southward in the United States to Pennsyl-
vania and Arizona. Adults are blackish with deep green reflec-
tions and are about 7 to 12 mm. long. Overmature or defective
trees and young trees suddenly released by the removal of domi-
nant trees are most often attacked; however, felled, topped, or
girdled trees and trees damaged by Saperda calcarata and Hy-
poxylon canker are also attractive. Infestations in standing trees
usually begin in the crown and move downward.

Adults are present during a period of several weeks during the
Summer and feed on the leaves of poplar. Eggs are deposited in
bark crevices and the larvae feed in the cambial region, excavat-
ing long tunnels that zigzag back and forth in a compact manner.
The larvae feed for two seasons in Ontario before becoming ma-
ture (35). Woodpeckers took heavy tolls of mature larvae, pupae,
and, presumably, adults in standing trees in Ontario.

Agrilus horni Kerr 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 which
encircle the roots and which may be continued up the trunk for
several inches. Pupation takes place in a cell in the center of the
stem, and the life cycle requires two years. Damage appears to

162

be most severe in sparsely stocked stands or in stands growing
on poor sites. (568).

Agrilus arcuatus var. torquatus LeC., the hickory spiral borer, has
been recorded from New York, Ohio, North Carollina, and Mis-
sissippi. It appears to favor hickory and pecan, but also infests
many other hardwoods. It, or a closely related species, has been
observed attacking young oaks and other hardwoods in the
Piedmont section of the South (40). Adults are about 8 mm. long.
Males have a greenish-bronze head and thorax, purplish-black
elytra, and the brassy underparts. Females are completely bronze.

Adults appear from May to July and feed on the leaves of
their host, eating 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 an inch or two and construct
pupal cells entirely within the pith. Two years are required to
complete the life cycle (104). This species is often very destruc-
tive of hickory seedlings in the South.

Agrilus acutipennis Mann. has been observed attacking overcup
oaks in Louisiana and Arkansas (539). Eggs are deposited on the
bark. The larvae bore through the bark and then excavate patches
of inner bark up to one-half inch 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.

Damage by Agrilus 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 18 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 one-half inch in di-
amter 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—honey locust; A.
leconteit Saund. and A. celtt Knull—hackberry; A. betulae Fisher
—river birch; A. cephalicus LeC.—dogwood; A. fuscipennis Gory
—persimmon; A. egenus Gory—black locust; and A. otiosus Say
—hickory, oak, maple, dogwood, red bud, walnut, and persimmon.

The genus Buprestis L. 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,

163

yellow, or orange. Often, there are many color variations and
patterns within a species.

The turpentine borer, Buprestis apricans Hbst., long considered
the most important eastern member of the genus, occurs through-
out the southern coastal regions from North Carolina to Texas,
and breeds in longleaf, slash, loblolly, shortleaf, and pitch pines.
The adult (fig. 48) is grayish-bronze with a greenish, metallic
lustre and is about 25 mm. long. It is elliptical, somewhat flat-
tened, and each elytron bears eight rows of large punctures.
Full-grown larvae have the prothoracic plates roughened and
Seepoe above by a dark brown Y, and they are up to 40 mm.
ong.

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 sur-
faces. The larvae tunnel in the sapwood and heartwood, construct-
ing long, narrowly oval, tortuous mines and filling 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 8 years are spent in the larval stage.

The turpentine borer used to be the most destructive insect in
the turpentine orchards of the South (38, 40). Borer riddled trees
were weakened so severely, they became subject to wind break-
age. The lumber value of such trees was virtually destroyed, and
gum production was seriously reduced. Attacks can be prevented
or reduced through the exercise of care in preventing the ex-

COURTESY OF DUKE UNIY. SCH. OF
FOREST.

FIGURE 48.—Adult of the
turpentine borer, Bupres-
tis apricans.

164

posure of dead, dry wood by fire, logging, or other forest opera-
tions. In recent years, 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 (Fab.)—adult brown with green-
ish reflections (breeds in dead branches of pine, hemlock, and
bald cypress) ; B. lineata (Fab.)—adult medium-sized and dark,
usually with brick-red to yellow longitudinal markings on the
elytra (attacks longleaf, loblolly, pitch, and scrub pines) ; B. ru-
fipes (Oliv.)—adult slender and dark green, with a long yellow
basal patch and two cross bars of yellow on the elytra (infesta-
tions occur around scars on elm, beech, hickory, oak, maple,
yellow poplar, and sour gum); B. salisburyensis (Hbst.)—adult
short, oval, and green with a tooth on the inner margin of each
elytron (breeds in pitch pine) ; and B. maculipennis Gory—adult
rather small and blackish, with a brassy tinge and scattered yel-
low spots or patches on the elytra (attacks pine and hemlock).

The genus Chrysobothris Esch. contains some of the most com-
mon 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-sized and are usually not con-
spicuously 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 strongly converge posteriorally. 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 in-
jurious, especially to young trees weakened by drought, defolia-
tion, or other adverse factors. The genus has been revised by
Fisher (253).

The flatheaded apple tree borer, Chrysobothris femorata (Oliv.),
one of the commonest and best known of the flatheaded borers
in America, occurs throughout most of Canada and the United
States. It attacks a wide variety of deciduous trees such as syca-
more, 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 several
inches long, especially in young trees. Tunnels in older trees are
confined mostly to 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 (242).

165

The flatheaded apple tree borer is especially destructive of
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 Australian-pine borer, Chrysobothris tranquebarica (Gmel-
in), also commonly known as the mangrove borer, occurs in south-
ern Florida and breeds in living mangrove and Casuarina trees.
The adult is greenish-bronze in color 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 orna-
mental or windbreak trees is often severe. The removal and de-
struction 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 flowing pitch to coagulate into large pitch masses
(fig. 49). Older larvae feed in the wood. The winter is spent in
the larval stage. In the spring of the third year, the larva con-
structs 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.

F-519916

FIGURE 49.—Pitch mass on red

pine caused by feeding of the
larva of Chrysobothris orono.

Several other species of Chrysobothris also occur fairly com-
monly in eastern forests. Some of these and some of their more
important hosts are as follows: C. pusilla L. & G.—pitch, short-
leaf, and white pines, white spruce, and hemlock; C. dentipes
(Germ.)—white, shortleaf, longleaf, and scrub pines, and larch
(this species is reported to be strongly attracted to sawmills) ; C.
floricola Gory—probaly all species of pines; C. scabripennis L. &
G.—pine, spruce, hemlock, and balsam fir; C. trinveria (Kby.)—
white pine and spruce; C. harrisi (Hentz) —Virginia, white, and
pitch pines; C. azurea LeC.—white oak, dogwood, maple, bass-
wood, birch, and willow; C. sexsignata (Say)—ash, red maple,
walnut, hickory, beech, yellow birch, white oak, hemlock, cypress,
and pitch pine; C. texana LeC.—red cedar; C. adelpha G. & H.—
hickory and pecan; C. veridiceps Melsh.—red maple, red oak, and
cherry; C. blanchardi Horn—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 eastern United States. Adults (fig. 50)
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 depres-
sions. 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 (Germ.) and C.

COURTESY OF DUKE UNIV. SCH. OF
FOREST.

FIGURE 50.—Adult of Chal-
cophora virginiensis, the
large flathead pine heart-
wood borer.

167

georgiana LeC. are often found in association with C. virginien-
sis. Adults of C. liberta are copper or brass-colored, while those
of C. georgiana are golden-bronze.

The Chalcophorella campestris Kerr., flatheaded sycamore-heart-
wood borer, breeds in injured, dying, or dead sycamore, beech,
oak, maple, yellow-poplar, and basswood, preferably in moist,
rotting logs. The larvae resemble those of Chalcophora 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 axe blazes and wounds and frequently riddles the wood beneath
with its tunnels.

The genus Dicerca Esch. contains many species that breed in
dead and dying trees and logs. The adults are dark-gray to brown,
medium-sized, 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 tail-like appendages. A few of the
more common eastern species and some of their hosts are as fol-
lows: D. divaricata (Say)—various dead, dying, or' injured hard-
woods; D. obscura (Fab.)—various dead hardwoods; D. punc-
tulata (Schon.)—various pines (often common around saw-
mills) ; D. lurida (Fab.)—hickory, blue beech, and basswood; D.
tenebrosa (Kirby)—pines and spruce; and D. tenebrica (Kby.)
poplar and willow.

The genus Melanophila Eschs. consists of small to medium-
sized 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 eco-
nomic importance. In the West, certain species are strongly at-
attracted to fires and are known as “fire bugs.” Sloop (653) pub-
lished a revision of the North American members of the genus.

The hemlock borer, Melanophila fulvoguttata (Harris), occurs
throughout eastern United States and eastern Canada. Hemlock
appears to be the preferred host, but it also occasionally attacks
several other conifers such as 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
equal-sized orange or yellow spots.

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 windthrown 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 here until they die. Under more favorable growth con-
ditions they penetrate to the cambium region and construct tor-
tuous, frass-filled galleries (fig. 51). Before becoming full-grown,
they construct cells in the outer bark in which they spend the
winter. The life cycle may be completed in one 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 dam-
age by the hemlock borer. Rapid salvage or cutting of heavily in-
fested trees is helpful in preventing population build-up in stands
weakened by windthrow or defoliation (470).

168

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 (Lap.)—pines; M. acuminata (DeG.)—
spruce, fir, pine, and eastern white cedar (adults are common
around forest fires and scorched timber) ; and M. aeneola Melsh.
—pines.

The genus Brachys Solier contains many species of leaf miners.
The larvae differ from the larvae of wood- or bark-boring species
in having the prothorax only slightly, if any, broader 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 is
very common on scrub oak in the sandhills of the southeastern
Coastal Plain. Heavy annual defoliation is not unusual. B. ovata
(Web.) also mines the leaves of oak. B. aeruginosa (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 red buds. 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
bald cypress in the 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;
Poecilonta cyanipes (Say) breeds beneath the bark at wounds

~< : i ae

F-514869

FIGURE 51.—Mature larvae

and galleries of the hem-

lock borer, Melanophila ful-

veoguttata, on the surface
of the sapwood.

on living poplars; P. thureuwra (Say) breeds beneath the bark at
wounds on willow; Agrillaxia flavimana (Gory) breeds in the
small branches of white oak; and Cinyra gracilipes (Melsh)
breeds in the dead branches of oak and ironwood.

FAMILY CERAMBYCIDAE
LONG-HORNED BEETLES OR ROUNDHEADED BORERS

The family Cerambycidae is one of the largest and most im-
portant of the families of wood-boring beetles. More than 1400
species have been recorded from the United States, about 450
of which occur east of the Mississippi River (221). Knull (431,
482) listed 262 species either known or believed to occur in Ohio
alone. Fattig (232) discussed the species occuring in Georgia. 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.

Th 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 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 de-
grade 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 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 seg-
ment. The beetles move rapidly when disturbed and are strong
fliers. Some species make squeaking noises when captured.

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 52.—Typical larva of a roundheaded borer.

170

Cerambycid larvae (fig. 52) are distinguished by a few promi-
nent characters. They are always fleshy, thin-skinned, white or
yellowish in color, and more or less cylindrical or depressed in
form. In some species, the body tapers somewhat posteriorally,
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 addominal
segment. There are two overlapping, circular bands of skin be-
tween 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 re-
tractile lobes (164, 165).

Keys to Cerambycid Larvae

Cerambycid larvae can be conveniently grouped as follows:
I. Girdlers of twigs or stems.
II. Gall formers on living plants.
III. Borers in the bark of living plants.
IV. Borers under the bark or in the wood of living trees.
V. Borers in the roots of living plants.
VI. Borers under the bark or in the wood of recently dead,
dying, or felled trees.
VII. Borers in older, moist wood in contact with the ground.
VIII. Borers in dry, seasoned wood or in cat faces and scars on
living trees.

I. TWIG AND STEM GIRDLERS.

Many species of cerambycids sever small twigs or branches
from living trees. These twigs may be seen hanging in the tree or
lying on the ground in late summer, fall, or winter. They are cut
either by the adult or the larva, the adult cutting from the outside
inward and the larvae from the inside outward.

1. Twigs or stems girdled from the outside. Stem never
hollowed out at point of severance 2
Twigs girdled from within. Part of stem always. hollow 8

2. Egg punctures and feeding scars on twigs or stems.
Severed twigs containing larvae .............. Oncideres spp.
No evidence of egg punctures or feeding scars. Severed
twigs never containing larvae ................... Oberea spp.
Se seomveerOund In PIrdled POLTION: 2) fe teeta 4
Larvae found in green portion of twig ee on plant.
Holes through bark of girdled portion .......... Hl
4. Twigs girdled by spiral cuts. Without holes in bark of :
TENRAIGR hs 4 Silaminacle Bee RA a MMR A HEMET ee en a Ec a oc OR ee
Twigs severed by complete hollowing. Holes through bark
MMeAnStbaAlOMe ING. eck Fc mere ate eee ax 6
5. Two ocelli on each side of head of larva; pronotum euhite
PATNA SOINLING os ee SSR RRO REA COE Elaphidionoides villosus
One ocellus on each side of head; pronotum brown and
CIV CU Vass ele he Pere elit lind Xylotrechus quadrimaculatus
6. Head of larva shining; pronotum wider than long
Aneflormorpha subpubescens

Luffa

7. In twigs of poplars; mandible pointed ...... Oberea schaumi

In shrubby plants; larvae boring down into base... 8
Si nAisassSaitras GaSe ei eee See ee Oberea ruficollis

In? SUMMA ee Sei SR: ED OR eae ee Oberea ocellata

In shadbush and related plants .....000000....... Oberea myops

Il. GALL MAKERS.

The larvae of several species of Cerambycidae cause galls or
swellings on the twigs or branches, along the trunk, or at the base
of trees or shrubs. They may be found either in the swelling
proper or in a mine above or below the gall.

1. Galls on twigs and branches:
On poplars and willows
Pronotum with 4 impressed lines, the inner being
darker* and obliques = Oberea ferruginea
Pronotum with 2 impressed, parallel lines
Saperda concolor; S. moesta

On Crataegus 2 ee eee Saperda fayt

On“Oakks * 3h Se Sarees aes hee aa enero rete Goes dibilis
2. Galls on main stem or trunk:

On“ willow. ee eee Oberea ferruginea

On red maple te se ee hn ener Xylotrechus aceris
3. Galls found at base:

On alderae grec een eee apes, eminem er Oar aes Saperda obliqua

On-elder,. 2s doa eae ..Desmocerus spp.

III. BORERS IN THE BARK OF LIVING TREES.

A few species of roundheaded borers are found in the bark
proper of living trees.

1. Larvae found only in the corky bark

Ofrelmiee ss .cs.4 oh el ee ee Physocnemum brevilneum
Ofte mulberry aise ek ee ce Elaphidion incertum
Ofwihitecoak 2.008 ie ee Encyclops coerulea

Of thick-barked rock oak or chestnut oak
Enaphalodes cortiphagus
2. Larvae bore into inner bark and often destroy the
cambium
In oaks and chestnut; at base or in crotches of
branches in damp situations ........ Anoplodera nitens
Trivashis. oh eae ee PET he Mea en Tylonotus bimaculatus

IV. BORERS UNDER THE BARK OR IN THE WOOD
OF LIVING TREES.

This group contains some of the most destructive roundheaded
borers. Some feed mostly beneath the bark, destroying the
cambium; the majority riddle the sapwood and heartwood, seri-
ously weakening the tree.

1. Larvae feed gregariously; mines overlap and meander 2

Larval mines essentiallyindividdal) ee). eee cme NSS
2. In heartwood at base of large trees
Pronotum shiny ...............Stenodontes and Archodontes
Pronotum /asperaterii). Avg seh Oo eee Seen Parandra
Beneath bark of firs, hemlocks, and larch ...... Tetropvum

172

In or beneath bark of ash; at base of privet

Tylonotus bimaculatus
AG baseqOr poplarsy: ser) e2ee. Xylotrechus obliteratus
Beneath bark and in wood of elm

EFutetrapha tridentata

Beneath bark and in wood of linden Saperda vestita

3. Boring beneath the bark and then directly up through the
wood; frass exuded through small hole in bark ................ 4

Boring in branches or main stems; long tunnel through
Centerson branch OF, Steers < Ge sie te eles i an i
4. In main trunks or branches ......... EMC eee irk! Acs Mindiayy 5
AMO ASC ten er me cm ee Pe er ee ag 6

5. Head of larva longer than wide; fibrous frass exuded
through holes in bark

In fig trees and alder ..... she Sh ache Re ae a Neoptychodes
In oak, beech, elm, and hickory _ PUM e Ls pel ted Deus 412) Goes
In poplar usc tua? Gan) Sins teint Seine er Lee __ Saper da calcarata
Head broader than HES granular frass exuded
nmOak se emrieee Enaphalodes rufulus
Iinwblack locust :2. sts ee be Megacyllene robiniae
IMNGmMaplerw ee ee Glycobius speciosus
6. In small oaks, Amelanchier, chestnut ......... Goes tessellatus

In apple, Amelanchier, Crataegus, ete.
Saperda candida
In poplar:
Pronotumeaspebate . 2.0 Saperda calcarata
Pronotum veivety pubescent and brownish
Plectrodera scalator
7. Last segment of body horny; in mesquite
Aneflus protensus

V. BORERS IN ROOTS.

The larvae of a few species of Cerambycidae are found in the
roots of living trees and shrubs. Some are true root feeders,
spending the entire larval stage in the roots; others spend only
the later instars in roots, having previously bored in the stems
above the ground.

1. Larvae feeding entirely in roots or directly at the base 2
Larvae feeding in roots, but stem above ground always
tunneled some distance; in shrubs 0...
2. In elder; sometimes causing swelling at ground surface;
basal portion of stem often hollowed ............ Desmocerus
In large hardwood trees or various shrubs... Prionus
3. Mandible rounded at apex; pronotum smooth
Aneflormorpha subpubescens
Mandible pointed at apex; pronotum roughened with fine

points

im@sassairas) 3... Reerdh havinge. oi eres ah Heck _Oberea ruficollis
IeSIINAC seer er tie sent, Het. tA Oberea ocellata
In rhododendron and related plants _...... Oberea myops

VI. BORERS BENEATH THE BARK OR IN THE WOOD
OF RECENTLY FELLED TREES

A number of species of cerambycids attack wood only when the
bark is present and still rather fresh and green. Trees dying

173

gradually are often subject to attack. After the wood has sea-
soned from 4 to 6 months it is nearly always immune to attack.
This group of borers causes heavy losses in rustic work and log
houses, and in lumbering operations when trees cut during the
winter are left lying on the ground throughout late spring and
early summer. Cordwood and pulpwood are also damaged.
The character of the frass, whether granular or fibrous or
shredded, distinguishes certain species.
1. Larvae found in coniferous trees _.0...... Ho ahs $5
Larvae found in hardwood trees _...... LS yFiaee AEE ens cs

2. Frass granular, closely packed behind larva
Frass composed largely of fibrous or shredded material
3. Frass pushed out through small hole...
Frass tightly packed; filling entire mine behind larva
4. Underside of head having four small tubercles
Callidium antennatum hesperum, C. texanum
Underside of head without tubercles... Oeme

5. [piey presen cs Geeaeey ie er oe Physocnemum andreae
In‘cedars, junipers; and Douelasir 33250 2 ee
In pines and SUEUICES) oe ee a ne

6. Larvae without ocelli and w jehout spines on lee sages
Semanotus ligneus
Larvae with 1 ocellus and with 2 small spines on last
Ssepment ‘aADOVE: 2) =e oe Aue: Atimia confusa
Head wider than long; Forni cylindrical
Larvae with 2 small dorsal spines on last segment of
body; mandible pointed at apex

Asemum, Arhopalus

Larvae without spines; mandible rounded at apex
— Xylotrechus
Head longer than wide; form depressed; ampullae velvety
pubescent Me eats eo ae Acanthocinus

8. Head wider than Wes 2 evel dorsal spines on last body
SQOWMION GE ont es eae ee Asemum, Arhopalus
Head wider than long and slightly keeled at sides: larvae
very depressed . Stenocorus inquisitor lineatus
Head longer than broad; last body ecement without dorsal
spines ...... PO bone an Rome UI, Se ee _.. Monochamus

9. Frass entirely ee Foe. Sete reat. er = 10
Frass containing much fibrous material _ f 11

10. Frass tightly packed behind larvae
Larval pronotum smooth and white. Larval mines
chiefly in the wood of ash, oak, and other hardwoods
Neoclytus
Larval pronotum covered with velvety, yellowish pu-
bescence; larvae without legs; larval mines under
the bark and in the wood Xylotrechus
Frass exuded through small hole

One ocellus on each side of the head; larvae yellowish

and rather elongate; chiefly in hickory
Knulliana cincta

Ory 00 09 CLO

~IS

~l

Two ocelli on each side of the head
Elaphidion mucronatum

174

Three ocelli on each side of the head

Houndinshickory |= one Megacyllene caryae
Found in mesquite ............ Megacyllene antennata
Found in black locust ............... Megacyllene robiniae

11. Pronotum brown, velvety pubescent; small irregular
dorsal and ventral tubercles on body segments
Leptostylus and related genera
Dorsum of prothorax and dorsal and ventral surfaces of
body segments armed with chitinous points

TCLS ete cet ee ee Leen Eutetrapha tridentata
invbasswood on inden 22. Saperda vestita
ime hickony aE Ne et aed Saperda discoidea

VII. BORERS IN OLDER WOOD IN MOIST CONDITION
OR IN CONTACT WITH THE GROUND

This group is composed mostly of large borers which occur in
large numbers in the same piece of wood until it is completely
riddled. Some are found in wood that has been cut a year or more
and that is in contact with the ground. They are rarely found in
recently dead or dying trees. Several of the species included here
are also considered in Group VI, as the larvae may continue to
bore for several years in stumps and logs lying on the ground.
Cross ties, mine props, fence posts, foundation timbers, and fire-
killed timber are subject to serious injury. A few species which
attack exposed heartwood at the bases of living trees cause
serious damage to shade trees.

1. Pronotum armed with small recurved chitinous points

Parandra

Pronotum brown, covered with very fine hairs or points;
two small dorsal spines on last body segments

Asemum, Arhopalus

Pronotum white, shiny, sometimes a little roughened or

TUNGULOSC es een) Moe cee er ee ee ee a 2
2. Dorsal and ventral surfaces of body covered with eral
Pullen elesin. <5. hc Wer aie re eee rena ee Leptura
Dorsal and ventral surfaces of body smooth; 2 transverse
impressions dorsally... NG riot OR OR OL as tele eg eI GA E 3
3. Front of head just above the mandibles proiertine in
FECUNEORALIIDCT CICS 2 ln cote es ee 4
Front of head projecting in transverse dull carina Cos
times divided in middle) | cee beens a 5)

4. Four rounded tubercles; body. an finely ried
Ergates (Western)
Four flat, sharp-edged teeth; body skin finely wrinkled
Tragosoma
Two flat, projecting teeth; body skin smooth, shiny
Orthosoma

5. Three ocelli on each side of head; body cylindrical
Stenodontes and Archodontes
One ocellus on each side of head; body wedge-shaped,
tapering posteriorally .__....................Prionus pocularis

175

VIII. BORERS IN DRY, SEASONED WOOD.

The larvae of a number of species of the family Cerambycidae,
belonging to several genera, are often referred to as “powder-post
borers” because of their common habit of boring for year after
year in the same wood, eventually leaving nothing but a thin
outer shell filled with a mass of powder. Certain species attack
only recently cut wood. Under normal conditions the life cycle of
the progeny would be completed in a year or two. But if the
wood, after being attacked, is placed in buildings or stored in
drier conditions, the larval period may be prolonged for several
years. Damage is often severe to stored lumber exposed to attack
before storing, to structural timbers in houses such as rafters and
beams, and to bridge timbers.

1. Three ocelli on each side of the head
Hylotrupes bajulus
One ocellus on each side of the head |. 2
2. Underside of mouth frame with 4 small projecting tu-
bercles; in coniferous wood
Callidium antennatum hesperum, C. texanum
Underside of mouth frame without tubercles
Tn ba nt OO re ee eae Chlorophorus annularis
ins NaTrgwo0dse fe (ano etc Ve ae a tie ne eee oles
3. Legs smaller than palpi.
A smooth white triangular are on venter of prothorax
Smodicum cucujiforme
Underside of prothorax not so marked; legs very
small; body dull, finely granulated above and below
Neoclytus caprea; N. acuminatus
ers larver thancpalpl «tcc a Eburia

The locust borer, Megacyllene robiniae (Forst.), 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 is about 18 mm. long. The jet-black
background is marked with bright yellow bands extending across
the thorax and wing covers, the third band on the wings being
W-shaped. The legs and antennae are moderately long and yel-
low. 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 crev-
ices and around wounds on the trunks of living trees. Newly-
hached larvae bore into the inner bark and construct small hiber-
nation cells in which they spend the winter. Activity is resumed
in the spring when the leaf buds begin to swell. At this time, 0oz-
ing 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 ex-
tensive tunnels throughout the heartwood (fig. 583A). As the larva
grows, it enlarges its tunnel to the exterior, through which it

176

pushes its granular frass to the outside and through which the
adult eventually emerges. There is one generation per year (280).

During the past several years, the locust borer has destroyed
thousands of acres of natural regeneration and plantations of
black locust. Enormous numbers of older trees, especially slowly
growing overtopped trees, have also been badly damaged or killed.
On the other hand, thrifty dominant trees over 10 years old have
seldom been killed. Infested trees are physically weakened by the
larval tunnels, the smaller ones often being restricted to shrub
form by repeated attacks (fig. 53B). Trees growing on poor sites
are especially susceptible to attack and suffer serious damage
during periods of prolonged drought.

F-501514, 501515
FIGURE 538.—Damage caused
by the locust borer, Mega-
cyllene robiniae: A, larval
damage in heartwood; B,
damage to small trees.

A number of practices have been suggested for the prevention
or reduction of damage by the locust borer. These include the
planting of 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 graz-
ing (318, 511, 189).

The painted hickory borer, Megacyllene caryae (Gahan), is
widely distributed in the eastern United States. Freshly-cut, hick-
ory logs are normally preferred for breeding, but dead trees of
several other hardwoods such as black and honey locust, oak,
hackberry, mulberry, walnut, butternut, and ash are also occa-
sionally attacked. Hickory wood cut during the winter may be
completely riddled by mid-summer. The adult closely resembles
the adult of the locust borer, but is slightly larger on the aver-
age. 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 antennata (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 de-
posited in crevices of the bark of recently cut wood. The larvae
feed first beneath the bark and then in the wood, excavating ex-
tensive mines and pushing quantities of frass to the outside
through a hole in the bark. This species is especially injurious to
mesquite cordwood. Fence posts also are greatly weakened or
destroyed. Craighead and Hofer (168) discussed methods of
control.

The banded hickory borer, Anulliana cincta (Drury), occurs
throughout much of eastern United States where it breeds in the
dead branches and trunks of a wide variety of hardwoods, such
as hickory, walnut, oak, American hornbeam, plum, and apple.
Hickory appears to be preferred. The adult (fig. 54) is dark
brown and about 22 mm. long. The body is clothed with grayish
fine hairs, and there is a short sharp spine on each lateral mar-
gin 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 sum-
mer, 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

178

F-480484
FIGURE 54.—Adult of the
banded hickory borer,
Knulliana cincta.

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.

The cottonwood borer, Plectrodera scalator (F.), breeds in the
bases of living cottonwoods and willows in the Southern States.
The adult is 25 to 35 mm. long and beautifully marked. The
ground color is black, but this is obscured by patches and cross
stripes of pure white, fine hairs that surround black, hairless
areas (fig. 55, left).

Adults appear in late spring or early summer and feed on the
tender shoots of young trees. These shoots often break, shrivel,

F-519918-9
FIGURE 55.—The cottonwood borer, Plectrodera scalator: Left, adults,
right, larva.

179

and turn black. Eggs are deposited in pits chewed in the bark
below the ground line at the base of the tree. Young saplings and
nursery stock are subject to attack, but larger trees are usually
selected. The larvae (fig. 55, right) feed in the phloem, mining
downward and commonly entering a large root by fall. The second
summer they continue feeding and excavating galleries and push-
ing out coarse frass through holes made near the egg slits. The
base of infested trees may be practically riddled by their tunnels.
Two years are required to complete the life cycle (533).

Cottonwoods planted on poor sites such as clay soils and sand
flats in Mississippi may be killed, or they may be broken off at
the ground by the wind. Damage may also be serious in natural
stands growing on poor sites (538).

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
about 27 mm. long. The head is clothed with yellow fine 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.

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, feed-
ing is resumed, with the larva cutting a larger gallery in the sap-
wood. The mature larva bores deep into the wood and constructs
a pupal cell at the end of its tunnel. Prior to entering the cell,
it cuts an exit hole through which the adult emerges. During this
activity, they push considerable quantities of sawdust to the out-
side. Pupation occurs in the spring, and the life cycle requires 2
years (703).

The presence of transverse ridges or elevations on the large
limbs or trunks of sugar maple, or of sawdust-like 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. These cracked, swollen areas often resemble cank-
ers or galls. Damage is generally most severe to shade trees or
to trees growing in open stands or along streams. Infestations
in the forest appear to be heaviest in open second growth stands,
or in heavily grazed, understocked stands. The growing of sugar
maple in well-stocked groups, the avoidance of overgrazing, and
the removal and burning of infested dead limbs and trees before
adult emergence in the spring should be helpful in reducing
losses (644). Borers in shade trees can be killed by the injection
of a fumigant into their tunnels or by piercing their bodies with
a wire pushed into their tunnels.

The red oak borer, Hnaphalodes rufulus (Hald.) (=Romaleum
rufulum), breeds in the trunks of living oaks in southern Canada
and throughout eastern United States, west to Minnesota, Iowa,
and Texas. In the Central States, red, scarlet, and black oaks are

180

especially subject to attack. Adults are light brown with spots of
lighter fine hairs, and are about 26 mm. long (fig. 56). 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.

~
~—-

BARS
B= 2 ae “<3
La ne F-520106
: FIGURE 56.—Adult of Enaph-
a ee alodes rufulus, the red oak
s borer.

Eggs are deposited singly on living trees over 2 inches in di-
amter in bark crevices or beneath lichen patches during the sum-
mer. Young larvae bore directly into the phloem, and they feed
there during the remainder of the summer, excavating cave-type
burrows about one-half inch 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 6 to 10 inches. The second winter is
also spent in the larval stage, with pupation occurring in the
spring. The life cycle requires 2 years. Granular frass pushed out
from points of attack and wet spots caused by sap leakage are
evidence of attack.

A large 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. Ants and fungi
entering the wounds add to the injury. Occasionally, branches
and entire trees are killed. Damage is severe in forest stands and
to valuable shade trees in parks and cities. 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 operations,
because the trees selected for removal in this work are the very
ones most likely to be heavily infested (326).

Einaphalodes cortiphagus (Craighead), the oak-bark scarrer,
breeds in the bark of living, mature oaks throughout eastern United
States and westward through the Ozark Mountains. Adults are
dark brown with patches of short, gray, fine hairs on the head,
pronotum, and elytra and are about 20 mm. long. Eggs are de-
posited in bark crevices in the spring, and the larvae feed in
the bark for upwards of three years. At the end of this period
they bore deeper into the bark and excavate a large pupal cell.

181

ee ee eT

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.. antomerius (Drury), breeds under the bark at the bases of dead
trees and on the stumps of oaks and chestnut in southern Canada
and throughout eastern United States.

The genus Goes LeC. contains a few species which 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
frass is expelled through openings maintained at the egg scar.
Life cycles vary from 2 to 5 years, depending on species.

Goes tigrinus (DeG.), the white oak borer, the largest species
in the genus, occurs throughout eastern United States. White
oak is its preferred host throughout most of its range, but vari-
ous other oaks, and hickory and walnut are attacked occasionally.
In Mississippi, overcup oak is heavily infested. The adult is
large, robust, dark brown, and from 25 to 30 mm. long. It is
irregularly covered with a dense coat of white fine hair, giving it
a white and brown mottled appearance (fig. 57). 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 a week or two
on the bark of tender twigs and leaves of oaks. Eggs are depos-
ited in the trunks of young trees, usually from 1 to 12 inches 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 one
inch in diameter and to 10 inches in length. Each borer bores two

Soap eNs

pane cas < > a
4 a> =

r
mR
e!

/ | F-519921
, FIGURE 57.—Adults of Goes tig-
lee re \ 3 rinus, the white oak borer.

182

separate holes in the tree—a small elongate entrance hole which
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.

The white oak borer is a major pest of overcup oak in the
bottom lands of Mississippi (679). Small trees down to one inch
in diameter are attacked and seriously damaged. Trees growing
on heavy clay soil with poor drainage, or where flooding is pro-
longed into the growing season, are frequently infested.

Goes pulcher (Hald.) the living-hickory borer, occurs in south-
ern Canada and throughout eastern United States and breeds in
the sapwood and heartwood of the trunk and branches of hickory.
| The adult is dark brown and about 18 to 25 mm. long. Its body
| is dark brown, covered with yellowish fine hair. The elytra are
clay-yellow with dark bands across the base and the middle. Eggs
| are laid in twig crotches and possibly at other places on the bark.
| The larvae feed for awhile beneath the bark and then bore into

the wood where they continue to feed for 2 to 4 years. Hickories
growing in the Piedmont area of North Carolina are particularly
prone to attack by this species.

The oak sapling borer, Goes tesselatus (Hald.), occurs from
Pennsylvania southward and in the Midwest where it usually
breeds in the base and roots of small oak saplings, preferably
white oak. Young chestnuts and shadbush are also attacked occa-
sionally. The adult is dull brown, about 25 mm. long, and its body
is covered with small patches of prostrate gray or yellowish fine
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 bulb-like swellings
around the wounds at the ground line. These trees are subject
to wind breakage.

Goes pulverulentus (Hald.), the living-beech borer, breeds in
the small trunks and branches of various hardwoods such as
beech, oak, elm, and sycamore throughout eastern United States.
The adult is brown, 18 to 25 mm. long, and its body is clothed with
short, white hairs. The thorax has a sharp spine on each side, and
the elytra are faintly barred at the middle and base with pale-
brown fine hairs. The life cycle requires 3 to 4 years.

Goes dibilis LeC., the oak branch borer, breeds in small, living
branches of oak, often causing gall-like swellings, in eastern
United States. Adults are brownish and from 11 to 19 mm. long.
The head, thorax, and apical half of the elytra are clothed with
reddish-yellow fine hairs. The elytra are crossed by two irregu-
lar, brown bands, and each basal half is mottled with grayish
hairs. The life cycle requires 2 to 3 years.

Many species of the genus Neoclytus Thom. occur in eastern
forests and attack both coniferous and deciduous trees. The larvae
feed first beneath the bark, then bore into the sapwood and heart-
wood, 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.

183

Se ee ee |

The pronotum bears transverse ridges and the body is marked
with transverse yellow lines.

The red-headed ash borer, Neoclytus acuminatus (F.), one of
our most common wood borers, occurs generally throughout the
eastern United States and in southeastern Canada. Its hosts in-
clude 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 about 15 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.

Neoclytus caprea (Say), the banded ash borer, occurs in east-
ern Canada and throughout much of the United States. Its hosts
are recorded as ash, hickory, elm, mesquite, and, rarely, white
oak. In eastern United States, it commonly breeds in ash logs.
Adults are dark brown to almost black and from 12 to 25 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 dur-
ing 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; how-
ever, 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 with the bark on may be literally honeycombed with
tunnels tightly packed with frass (fig. 58).

Other species of Neoclytus likely to be encountered in eastern
forests and some of their known hosts are as follows: N. mu-
cronatus (F.) (fig. 59)—the dead branches and trunks of hick-
ory; N. jouteli (Davis)—dead oak twigs; N. fulguratus Casey—
dead branches of oak; N. scutellaris (Oliv.)—elm, oak, hickory;
N. muricatulus Kirby—larch, spruce, pine (in southern Canada
and from Maine to Ohio) ; and N. muricatulus muricatulus Kirby
—small post oaks (in North Carolina).

The genus Saperda F. is represented in eastern forests by sev-
eral important species. Depending on the species, the larvae
either bore in (1) the large branches or trunks of living trees;
(2) in small branches and produce galls; or (3) in living or dead
wood of dying or recently killed trees. Adults are medium-sized
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

184

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 58.—Galleries of Neoclytus caprea, the banded ash borer.
Note that tunnels are tightly packed with granular frass.

clothed with a hairy covering. Felt (239) published a monograph
on the genus.

The poplar borer, Saperda calcarata Say, occurs throughout the
United States and Canada, wherever poplar grows. Willows are
also subject to attack. The adult is from 20 to 28 mm. long,
grayish-black or reddish-brown, and densely clothed with gray
and yellow fine hairs (fig. 60). There also are 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, usu-
ally 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 years in the North. In the Deep South it is shorter. In

185

186

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 59.—Adults of Neoclytus mucronatus.

F-519920

FIGURE 60.—Adults of the

poplar borer, Saperda cal-
carata.

i

| J. | 2 es is nen Sn

the North, the second winter is spent as a mature larva in a cell
at the end of its tunnel.

Small trees are occasionally killed by larvae girdling be-
neath the bark. Larger trees are seldom killed outright, but the
large larval tunnels make them susceptible to windbreakage. They
may be literally riddled with tunnels. Larval openings and tun-
nels serve aS openings for various rots, decays, and other in-
jurious insects which may kill the tree or degrade the lumber.
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
individual trees or small groups of trees unevenly distributed
throughout the stand. These trees, commonly known as brood
trees, are usually the larger, faster-growing trees in stands aver-
aging between 8 and 7 inches d.b.h. Lakes 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
clear cut them at maturity. The removal of “‘brood trees” should
also be helpful (234).

The roundheaded apple tree borer, Saperda candida F., occurs
in Canada and throughout eastern United States. It is most seri-
ous as a pest of apple orchards, but it also breeds in mountain-
ash, Crataegus, and shadbush. The adult is brilliantly white ex-
cept 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 (344).

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 one year and then bore into the wood,
making large excavations and riddling it. The presence of ten- .
drils of frass on the bark or at the base of the tree is evidence
of attack. Heavily infested trees may be killed in a single season.
Two or more years are required to complete the life cycle. Keep-
ing 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, Saperda vestita Say, occurs in the Northeast-
ern 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 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 kill-
ing the tips of infested branches. The larvae feed 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 sub-
ject to attack. Adults are 10 to 17 mm. long. The male is blackish
with reddish legs and lines of grayish fine hair on the thorax. The
female is reddish-brown or nearly black, clothed with yellowish

187

fine hairs, and has two spots separated by a curved bar on the
elytra. Eggs are deposited in bark crevices or in holes in the gal-
leries of bark beetle during late spring and summer. The larvae
feed gregariously beneath the bark, making extensive meandering
mines. Most of the larvae mature in 1 year.

Saperda concolor LeC. breeds in the living branches and main
stems of poplars and willows in the Northern and Central States.
Adults are black with fine gray hairs and are about 12 mm. long.
Eggs are laid in longitudinal slits in the bark. Callous tissue form-
ing over these slits causes the formation of globose galls. The
larvae mine around the stem and then bore into wood mak-
ing galleries about 1 inch long that run parallel to the axis. Usu-
ally the infested twigs are not killed. Branches or small stems
bearing numerous galls may stop growing, however, and become
subject to wind breakage.

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 several inches. 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 orna-
mental plantings are frequently killed.

Saperda fayi Bland, the thorn-limb borer, occurs in the North-
east 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
which extends upon 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 wind breakage. S. moesta LeC.
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 uniformly gray and about 10 mm. long. S. lateralis F.
breeds in dead hickory, elm, basswood, oak, and Prunus 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 and Joutel breeds in dead hickory, willow, and
basswood; S. cretata Newm. in living apple and the limbs of liv-
ing hawthorn; S. mutica Say in dead willow.

The elm borer, Hutetrapha (=Saperda) tridentata (Oliv.), oc-
curs in southeastern Canada and throughout 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. 61). 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 cross band at the base
of the wing covers followed by two oblique bands. Full-grown
larvae are about 12 to 15 mm. long.

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

188

COURTESY OF ILL. NAT. HIST. SURV.

FIGURE 61.—Adult of the elm

borer, Eutetrapha (= Saperda)
tridentata.

weakened trees. The larvae bore beneath the bark, filling their
mines with fibrous frass, and completely destroying the phloem
and cambium (587). When they reach maturity they bore into
the wood and construct cells in which to pupate. There is 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. Af-
fected trees tend to die very slowly, a branch at a time. The re-
moval of infested branches is sometimes helpful in control.

Smodicum cucujiforme (Say), the flat powder-post beetle, oc-
curs throughout eastern United States. The larvae excavate ex-
tensive 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 about 7 to
10 mm. long. Adults appear in July and August. Eggs are laid
in crevices of exposed wood. The larvae excavate tunnels in the
wood about 3 mm. in diameter, 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 Mulsant, 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, Oberea tripunctata (Swed.) breeds in
dogwood, elm, sourwood, laurel, azalea, Viburnum, and various
fruit trees in 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 cut-
ting off portions of the twig as it bores on into the green wood.
The winter is spent in the larval stage in the twig. Pupation oc-
curs in the spring between two wads of fibrous frass. The portion

189

of the twig containing the cell may have been girdled previously.
Occasionally dogwoods and elms are seriously damaged. The cut-
ting off and burning of infested twigs might be helpful in control.

Oberea schaumi LeC. 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 to black and is
marked by four round, smooth spots. The elytra are either yellow-
ish 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 6 inches
long, and boring holes to the outside for the expulsion of frass.
Black, necrotic areas develop around these holes. Infested twigs
die but are usually not sufficiently weakened to permit wind
breakage.

Oberea myops Hald., the rhododendron stem borer, breeds in
rhododendron, azalea, and 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 one-half inch apart. The larva
bores down the twig, to and 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 plant-
ings of rhododendron. Cutting off and burning of infested twigs
as soon as they are noticed is recommended.

Oberea ocellata Hald., the sumac stem borer, occasionally causes
serlous 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 proceeds to construct long tunnels and to
feed 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 sassa-
fras. Adults are pale reddish-yellow and about 18 to 20 mm. long.
The antennae, tibia, 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 Chitt.—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 Serville contains several species, the fe-
males of which deposit their eggs in previously girdled terminals,
twigs, or small branches of large trees, or in the stems of seed-
lings 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 wall-
ing off a portion of the mine with fibrous frass. The adults appear

190

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.

The twig girdler, Oncideres cingulata cingulata (Say), is the
most important eastern member of the genus. Its hosts include
hickory, persimmon, elm, oak, honey locust, hackberry, poplar,
basswood, dogwood, sourwood, and various fruit trees. Adults
(fig. 62A) are grayish-brown and about 15 mm. long. The front
of the head is clothed with fine golden hairs; the elytra are clothed
with gray, fine hairs and are marked with scattered yellow spots.
Full-grown larvae are up to 18 mm. long.

a

COURTESY OF DUKE UNIY. SCH. OF FOREST.
FIGURE 62.—The twig girdler, Oncideres cingulata 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 are also ragged
and unattractive. Hickory seedlings are especially prone to attack
and damage, often becoming distorted following attack (fig.
62B). Honey locust seedlings in nurseries are also subject to
serious injury. The collection and burning of infested twigs and

Oa:

branches during the fall or winter is an effective method of con-
trol, provided the trees to be protected are located at some dis-
tance from heavily infested stands.

Oncideres cingulata texanus Horn and O. pustulatus LeC. are
common in parts of eastern or southwestern United States. The
former is a pest of pecan in the South; the latter attacks hui-
sache, acacia, and mesquite in the Rio Grande River Valley and
other parts of the Southwest.

EHburia quadrigeminata (Say), the ivory-marked beetle, occurs
in both eastern and western United States and breeds in the dry
heartwood of various hardwoods. In the East, oak, hickory, ash,
maple, honey locust, elm, chestnut, and cypress are attacked.
Adults are light brown, with pairs of ivory spots at the base and
middle of each elytron, and are about 24 mm. long. The larvae are
wedge shaped, have tough, shiny skins, and are sparsely clothed
with golden hairs. Mature oaks having “cat faces” or scars
through which the larvae can gain access to the heartwood are
often badly damaged. Seasoning lumber is also subject to occa-
sional attack. The normal life cycle appears to be 2 years, but it
may be considerably longer. There are records of adults emerg-
ing from flooring, door sills, and furniture 15 years after it was
placed in use.

The twig pruner, H'laphidionoides villosus (F.) occurs through-
out eastern United States. It breeds in the twigs and branches of
living hardwoods such as the oaks, hickory, maple, locust, hack-
berry, walnut, elm, sweetgum, and pecan.Adults are slender, elon-
gate, light brown and about 23 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 bi-spinose.

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 including, the thin bark (fig. 63). These
branches, from 14-inch to 2 inches 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
they lose much of their esthetic value. The presence of numerous
dead twigs and branches hanging in the crown also detracts from
their appearance. The collection and burning of infested twigs
and branches in the spring before the adults emerge should be
helpful in control.

Elaphidion mucronatum (Say), the spined bark borer, also breeds
in the dead branches of various hardwoods in eastern United
States. Adults are dark brown, irregularly clothed with fine
brown hair and are about 19 mm. long. Eggs are deposited be-

192

F-504092
FIGURE 63.—Larva of the twig pruner, Elaphidionoides villosus, in an oak

twig. Note ends of severed twigs.

neath 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. EL’. incertum Newm., the mulberry bark borer, a species
that closely resembles the spined bark borer, breeds in the outer
bark of living mulberry trees in Eastern United States. Infesta-
tions have also been recorded in oak and hickory. It does not
cause serious damage.

Desmocerus palliatus (Forst.), the elder borer, breeds in elder
wherever it grows in eastern United States. Adults are bright,
metallic blue with nearly all of the basal half of the elytra yellow,
and are about 27 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.

Deoraschema wildii Uhler, the mulberry borer, breeds in the liv-
ing 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. 64). 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, destroying irregular patches of cam-
bium up to 2 square inches in area. After about 1 to 3 months
they bore into the wood, constructing galleries that angle upward
and inward for a couple of inches and then bend back towards the
surface. These galleries are close together and often overlap. The
winter is spent in the larval stage, and the life cycle varies from
1 to 2 years in length. Suppressed and fire-damaged trees are pre-
ferred, 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

193

on the trunk (678). The smaller but related species, D. alterna-
tum (Say) is commonly found in trees infested with the mulberry
borer.

Aneflormorpha subpuescens (LeC.) the oak-stem borer, occurs
in the Eastern, Southeastern and Central States. It breeds in
small living oak and chestnut seedlings from 14-inch to 1 inch
in diameter, and occasionally in the branches of larger trees. The
adult is narrow, elongate, light brown, clothed with semi-erect
fine brown hairs, and about 17 mm. long (fig. 65). There is a stout
spine on each of the third and fourth segments of female an-
tennae, and the tips of the elytra are notched and bi-spinose.

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 sec-
tion 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 generation per year. During certain years a high proportion
of the oak seedlings in the Southeast are killed by this species.

The black-horned pine borer, Callidium antennatum hesperum
Csy., occurs throughout the United States and breeds in dead or

COURTESY OF DUKE UNIV. SCH. OF FOREST. COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 64.—Doraschema wildii, the FIGURE 65.—Adult of Aneflormor-
mulberry borer: Top, adult; bot- pha subpubescens, the oak-stem
tom, damage by larvae. borer.

194

recently felled conifers, or in lumber with the bark on. It is espe-
cially 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. 66A). Male antennae are somewhat shorter than the body;
the thorax is rounded, with depressions on each side of the mid-
dle; and the legs are black with large femora.

F-519946, (B, COURTESY DUKE UNIV. SCH. OF FORESJ.)
FIGURE 66.—The black-horned pine borer, Callidium antennatum hesperum:
A, adult; B, larval galleries under the bark and in the wood.

Adults appear early in the spring and deposit their eggs be-
beneath bark scales on dead trees or on cut wood that has sea-
soned over winter. The larvae feed in the phloem and outer
sapwood making broad, wavy tunnels in the wood (fig. 66B) 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 sawn lumber,
and kiln drying are recommended methods of control.

Callidium texanum Schaeffer, the black-horned juniper borer,
breeds in various conifers, preferably juniper and cedar, through-
out the United States. Its habits are similar to those of the black-
horned pine borer. Other eastern species include: C. schotti,
Schffr. which breeds in the dead branches of cedar in the Midwest,
and C. violaceum (L.) which occurs in red spruce in New York.

The genus Acanthocinus Stephens contains a number of species
commonly known as pine-bark borers. The adults are elongate,
rather flattened, and from 12 to 28 mm. long. They are also usu-
ally black and mottled or striped with grayish-white or brown
pubescence. The basal joints of male antennae are fringed be-
neath with hairs, and the ovipositor of the female is characteris-
tically extended.

195

Acanthocinus nodosus (F.) is a common species in the South
where it breeds in the thick bark of pine logs and dying and re-
cently killed pines. The adult (fig. 67) 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 de-
stroy bark beetle broods. Pupation occurs in nestlike chambers
in the bark, near the surface. There is one generation per year.
Other eastern species include: (1) A. obsoletus (Oliv.)—adults
are about 10 mm. long. Hosts are recently killed pine and balsam
fir. (2) A. pusillus (Kirby) —adults are about 10 mm. long. It has
been reared from wind-thrown and fire-killed red and jack pines,
balsam fir, and spruce.

The genus Anoplodera Muls. contains many species of rather
robust and usually more or less triangular-shaped beetles that are
nearly always beautifully colored and hairy about the head and
thorax. They are also quick and active and are usually found on
flowers. Eggs are laid in crevices of bark or wood in moist loca-
tions. The larvae feed in the wood for 2 to 8 years, excavating
extensive galleries they then pack with fibrous frass. Infested
wood may be reattacked repeatedly until it is completely disinte-
grated. Poles, cross ties, and other material in contact with the
ground may be completely destroyed.

Anoplodera nitens (Forst.) 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.

Anoplodera canadensis (Oliv.) breeds in dead pines, spruces,
hemlock, and sometimes balsam fir in southern Canada and the
Northern States. Living trees are also 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

POS
% @ we ll “=
s : f y 1 ‘
- 5 f .
ee a 2s: de 7
j i { COURTESY OF DUKE UNIV. SCH. OF FOREST.
eee FIGURE 67.—Adult of Acanthocinus
aN he: Z| nodosus, a pine-bark borer.
‘: !
vee

196

is coarsely punctured, and the antennae are generally ringed with
yellow.

Additional eastern species of Anoplodera and their hosts include
the following: A. minnesotana (Casey)—hickory, elm, black
cherry, sourgum, and red spruce; A. vagans (Oliv.)—butternut,
hickory, birch and pine logs; A. proxima (Say)—maple, hickory,
basswood, and sourgum; A. nigrella (Say)—pines and spruce;
A. mutabilis (Newm.) and A. vittata Oliv.—red spruce; and A.
biforis (Newm.)—white pine.

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 about 14 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 re-
quired to complete the life cycle. Bark stored for tanning pur-
poses is subject to heavy damage. P. dimidiatus Kyb. breeds in
fir and spruce in the Northern States. Adults are about 11 mm.
long, and dark brown, with a light brown band across the base
of the elytra. P, varius (Fab.) breeds in or beneath the dead bark
of hickory and oak and possibly in various conifers. The adult is
light brown or dark brown to black and shiny. There are white
bands on the elytra, and the adult borer is about 10 mm. long.

Physocnemum andreae Hald., the cypress bark borer, breeds un-
der the bark of girdled, felled, or dead cypress 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, ex-
cavating large mines that deeply scar the wood. Winter is spent
in the larval stage, and pupation occurs in the spring. The spe-
cies often causes serious losses to cypress trees felled or girdled
during lumbering operations. Rustic work constructed from this
wood is also subject to serious damage. Rapid utilization of gir-
dled 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 (317). Adults are dark brown to
black and about 17 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 meander-
ing, frass-packed galleries. The bark over these galleries dies and
falls off. P. violareipenne Hamilton breeds in the small branches
of white oak in the Midwest. Adults are only about 9 mm. long;
otherwise, they are very similar to those of the elm bark borer.

Parandra brunnea (F.), the pole borer, occurs throughout much
of the United States, and attacks a wide variety of hardwoods
and conifers. Logs, poles, and other wood products in contact
with the ground, such as cross-ties and structural timbers, are
also infested. The adult is flat, shiny, mahogany-brown, and about
18 mm. long. Full-grown larvae taper slightly toward the rear
and are about 30 mm. long.

oT

Adults appear during mid-summer 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 8 or 4 years. Although the wood
may be completely honeycombed, a covering shell of sapwood is
always left intact. Pupation 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 considerable 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 cedar tree borer, Semanotus ligneus (F.), occurs through-
out the United States. Practically all species of conifers are sub-
ject to attack, but dying and recently felled cedars 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. The related species, Semanotus
litigiosa (Casey), has been recorded from 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 Esch. and Arhopalus Serville 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 ob-
soletus (Rand.) are common eastern species.

Tragosoma desparius (L.), the hairy pine borer, occurs from
coast to coast in southern Canada and the Northern States. It also
occurs southward through the Appalachians in the Eastern States
where it breeds in various dead conifers. 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. Occa-
sionally crossties, poles, and timbers in contact with the ground
are seriously damaged.

Atimia confusa (Say), the small cedar-bark borer, breeds in
dying cedars, junipers, and related trees throughout the Eastern
and Central States. The adult is small, stoutish, and about 9 mm.
long. The head, pronotum, elytra, and venter are black. The dor-
sum 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

198

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 cedar and juniper is subject to attack
and serious damage. The bark over damaged areas dries and
peels off.

Aneflus protensus LeC. attacks living mesquite in Texas and
other Southwestern 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 20 to 30 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
ue life cycle. Infested branches and small trees are sometimes
killed.

Tetropium cinnamopterum Kby., the eastern larch borer, breeds
beneath the bark of living and dead coniferous trees, especially
larch, and occurs from coast to coast in southern Canada and the
Northern States. 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. There
is one generation per year in the United States. This was the
most common cerambycid borer attacking windthrown spruce in
the Adirondacks following a severe windstorm in 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 ex-
cept for small reddish-yellow spots on the body and white stripes
on the elytra, and is about 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 Hald., 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 12 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 seales on living and dying ash trees during early summer.
Young larvae feed in the phloem; later they penetrate deeper and
sear 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 branch. Privet hedges are subject
to severe damage. The larval stage extends over a period of 2
years.

Eneyclops coerulea (Say), the oak-bark scaler, bores in the
outer bark of living white oaks, yellow-poplar, maple, sourgum,
black ash, and pignut hickory in the Eastern States. The adult is

199

slender, metallic blue or green, has light brown legs, and is about
9 mm. long. Dry scales over damaged areas of bark peel off.

Oeme rigida (Say) breeds in dead and dying cypress and juni-
per throughout eastern United States. The adult is light to dark
brown and between 12 and 22 mm. in length. Eggs are deposited
beneath bark scales and the larva feeds beneath the bark and in
the wood, exuding large quantities of granular frass. Deadened
cypress 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 3 inches in
diameter.

The genus Prionus Geoff. consists of a number of species of
robust, black or brownish-black, 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 eastern United States. Root rots, such as Armillaria, and
various species of secondary boring insects often attack the dam-
aged roots and kill the trees. Mature trees growing in open
stands, in well-drained gravelly soil and hillsides, in pastures, and
in heavily used recreation areas are especially subject to attack.
Small trees and shrubbery are occasionally killed by borers that
cut off their roots at the ground line.

The broad-necked root borer, Prionus laticollis (Drury), breeds
in the roots of a wide variety of trees and shrubs in eastern United
States, mostly hardwoods. Oak, poplar, chestnut, and basswood are
especially favored. Infestations have also been recorded in the
logs and stumps of all the above species. The adult is dark-
brown, shining, and up 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 3 to 5 inches of the soil surface in
the spring and form oval, compact cells in which to pupate. The
life cycle is believed to require 3 years. This species seldom causes
substantial damage in the forest.

The tile-horned prionus, Prionus imbricornis (L.), occurs
throughout much of eastern United States and breeds in the living
roots of oak, chestnut, pear, and various herbaceous plants.
Adults (fig. 68) are dark brown, shiny and up to 37 mm. long.
Male antennae are 18- to 22-jointed, and the segments overlap.
Eggs are deposited in groups in the soil around the base of the
tree. Larval habits are similar to those of the broad-necked root
borer. The life cycle requires at least 3 years.

Prionus pocularis Dalm. breeds in decaying coniferous logs and
stumps in the Central, Middle Atlantic and Southern States.
Adults are light brown, shiny, and about 42 mm. long. The elytra
are densely punctured.

200

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 68.—Adult of the tile-
horned prionus, Prionus imbri-
cornis.

Several species of Xylotrechus Chev. occur in eastern forests.
Adults are characterized by their moderate size, their short, fili-
form antennae, and a V-shaped callosity on the front of the head.

Aylotrechus quadrimaculatus (Hald.), the birch and beech
girdler, breeds in the branches of birch, beech, blue beech, maple,
and alder in the Eastern States. Adults are about 12 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 2 inches in diameter frequently fall to
the ground, with the larvae in them, during mid-summer. There
is one generation per year.

The gall-making maple borer, Xylotrechus aceris Fisher, breeds
in the trunks and branches of various maples, causing the forma-
tion of galls. The adults resemble those of the birch and beech
girdler, although they are somewhat smaller and have fainter
spots on the thorax and stronger markings on the elytra. Eggs
are deposited during mid-summer 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 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, be-
coming subject to breakage by the wind.

201

Aylotrechus obliteratus LeC., the poplar-butt borer, a serious
pest of aspen and other poplars in the Rocky Mountain region,
also occurs in many parts of 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 (Germ.) breeds in dead conifers in
eastern Canada and the Northern States. In areas where pines
predominate, it is especially common. Logs, slash and trees killed
by fire or bark beetles are particularly attractive. Adults (fig. 69)
are dark brown and about 18 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.

The rustic borer, Xylotrechus colonus (F.), one of the com-
monest of all cerambycids in eastern United States, feeds under
the bark of almost all dead hardwoods. It occurs also in southern

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 69.— Adult of
Xylotrechus sagittatus.

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 70.—Adult of the rustic
borer, Xylotrechus colonus.

202

Canada. The adult (fig. 70) is light to dark brown and about 14
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 ex-
clusively in the bark without scarring the wood (279). Recently
killed trees are preferred.

Xylotrechus annosus (Say) and X. undulatus (Say) both occur
in eastern United States. X. annosus, a western species, breeds in
poplar in the Lake States; X. wndulatus breeds in recently cut
copes fir, larch, hemlock, and spruce in the northern tier of

tates.

The genus Monochamus Serville is represented by several im-
portant 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,
felled, dying or recently dead trees are preferred. Young larvae
feed on the inner bark, cambium, and outer sapwood, forming
shallow excavations called surface galleries and filling them 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 re-
maining wood and bark. Full-grown larvae are often more than
50 mm. long. Members of this genus often cause heavy losses in
windthrown or fire-killed timber, in sawlogs left too long in the
woods before milling, and in improperly handled pulpwood. Hop-
ping (378) published a key to the adult of the genus.

The southern pine sawyer, Monochamus titillator (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 from 18 to 30
mm. long (fig. 71). Male antennae are often 2 to 8 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.

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 above for the genus. There are at
least two generations per year in the South, with overlapping
broods (750). 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 white-spotted sawyer, Monochamus scutellatus (Say), oc-
eurs from Newfoundland to North Carolina, Minnesota, and
Alaska. White pine appears to be its favorite host but it also at-
tacks many other conifers such as red and jack pines, balsam fir,
white, black, and red spruces, and larch. The adult (fig. 72) is
about 18 to 25 mm. long. The male is completely shiny black ex-
cept 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.

203

F-480483
FIGURE 71.—Adult of the southern pine sawyer, Monochamus titillator.

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 (773, 618).

The white-spotted sawyer causes heavy losses to sawlogs and
pulpwood in the Northern States and southern Canada. Damage
to sawlogs can be prevented by cutting the trees between Sep-
tember and early June and removing them from the woods before

F-496927

FIGURE %72.—Adults of the

white-spotted sawyer, Mon-
ochamus scultellatus.

204

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
1 to 2 feet thick.

The Northeastern sawyer, Monochamus notatus (Drury), oc-
curs in eastern Canada and in the Northeastern States, westward
to the Great Lakes region, and breeds in dead and dying white
pine and balsam fir and in windthrown red spruce. Adults are
dark brown and up to 80 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 inter-
rupted stripes. The female head is greatly flattened and elongated.

The balsam fir sawyer, Monochamus marmorator Kby., breeds
in balsam fir in eastern Canada and the Northeastern States, west
to the Great Lakes region and south to 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 (Oliv.) breeds in dead and
dying pines in the Southeastern 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. macu-
losus Hald., 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 grout 16 mm. long. The tip of each elytron is prolonged into
a tooth.

Orthosoma brunneum (Forst.), the brown prionid, breeds in
decaying coniferous and hardwood logs in southeastern Canada
and throughout much of 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.
Cross-ties, 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 (Say), the hardwood stump borer, oc-
eurs from Virginia southward and westward and breeds in the
heartwood of living hardwood trees such as various oaks, syca-
more, willow, and boxelder. Wood in contact with the ground is
also attacked. The adult is a large, somewhat flattened, reddish-
brown beetle from 30 to 45 mm. long. The head is 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 gregari-
ously for 3 or 4 years, completely honeycombing the wood. Shade
trees are sometimes weakened so badly that they break and fall
during storms. Cross-ties and other wood products in contact with
the ground are also subject to serious damage.

205

Archodontes melanopus (L.) breeds in the roots of live oaks
and possibly hackberry and pecan from Virginia to Florida and
westward along the Gulf Coast. The adult is broad, rather flat,
dark brown, and from about 438 to 56 mm. long. The head is dis-
tinct and rather large, and the edges of the prothorax are finely
toothed. Full-grown larvae are almost 90 mm. long and are about
as thick as one’s finger.

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 bush-like suckers. It has
been suggested that this species was largely responsible for the
creation of large areas of comparatively barren areas of scrub
oak in parts of southern Georgia and Florida.

Stenocorus inquisitor lineatus (Oliv.), 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 12 to 16 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
winter. Trees dead for only a short time and containing consider-
able amounts of moisture are preferred by the larvae. They feed
entirely beneath the bark, excavating irregular galleries and pack-
ing them with fibrous frass. When they become full-grown they con-
struct oval, fibrous-edged cells in which they pupate and spend
the winter as adults.

The old-house borer, Hylotrupes bajulus (L.), a European
species introduced into North America more than a century ago,
is now known to occur in the Atlantic Coastal States from Massa-
chusetts to Florida. Adults have also been collected in the Gulf
Coast States of Mississippi, Louisiana, and Texas. It breeds in
dry, seasoned, coniferous sapwood. Pine and spruce wood appear
to be preferred, but hemlock, true fir, and Douglas-fir wood are
also attacked. The adult (fig. 73 A) is a slightly flattened, brown-
ish-black beetle from 12 to 18 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 patches of gray
which fuse to form two cross bands or two whitish spots. Full-
grown larvae are wedge-shaped, deeply segmented, and up to 30
mm. long.

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 col-
lapse 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

206

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. 73 B), framing members, and
other wood parts are seriously weakened by its mining and tun-
neling. Air-dried pine floor joists, plates or sills, and sub-flooring
are apt to be damaged most severely. Other framing such as
studs, stair carriages, furring strips, and roof rafters are some-
times attacked.

When an old-house borer infestation is discovered, two types of
remedial action are necessary: (1) the repair or correction of
serious structural defects and (2) controlling the remaining in-
sects present in wood left in place (486).

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 are
about 10 mm. long. The first two pairs of legs and the inner parts
of the antennae are red. Well-seasoned bamboo is mined exten-
sively, and the mines are tightly packed with fine, powdery frass.
This species is a serious pest of bamboo in India and Japan.

F, 494430, 494432

FIGURE 73.—The old house

borer, Hylotrupes bajulus:
A, adults; B, damage.

207

Rhyncophora
(Snout Beetles)

Snout beetles differ from other Coleoptera in having the head
of the adult more or less prolonged to form a beak or snout, the
gular sutures united in a median line, and the joints of the palpi
usually rigid. The larvae are whitish, C-shaped, more or less
cylindrical, and legless. A great many species occur in eastern
North America, nearly all of which feed on trees and other types
of vegetation. The larvae usually are burrowing in habit, infest-
ing fruits, nuts, twigs, roots, and terminals. Trees of all ages are
attacked, and damage is often severe. Forest nurseries and plan-
tations are especially susceptible to serious injury (84).

FAMILY BRENTIDAE
BRENTID BEETLES

Only one tree-infesting species of this family occurs in eastern
United States. The remainder are confined almost entirely to the
tropics.

The oak timberworm, Arrhenodes minutus Drury, breeds in
hardwoods, chiefly oak, beech, and poplar in the Eastern States.
Adults are dark reddish-brown to black and up to 35 mm. long.
The snout of the female is straight, narrow, and much longer
than the head. In the male, it is broader and flatter and armed
with powerful mandibles (fig. 74). The larvae are elongate, cylin-
drical, and soft-skinned.

In Missouri, adults are active from early May to late August
and the female deposits her eggs in deep holes chewed into the
wood, often at fresh wounds. Living trees and trees dead up to 3
years are subject to attack. The larvae bore through the wood,
constructing pinhole galleries across the grain in all directions.
The life cycle usually requires 3 years (117).

F-519917

FIGURE 74.—Adults of the

oak timberworm, Arrhe-

nodes minutus. Male on
left; female on right.

208

The oak timberworm is capable of causing serious injury. Occa-
sionally, much of the timber in infested stands is so badly damaged
it is unfit for many uses. Control is difficult, but the removal of
dead and dying snags and fire-scarred trees and the prevention of
wounds in living trees should be helpful.

FAMILY ANTHRIBIDAE
FUNGUS WEEVILS

Fungus weevils are of little or no economic importance but are
frequently encountered in the wood of dead hickory, beech, and
maple trees. The adult is brown to whitish mottled and has a
broad, flat, and very short beak. The larvae feed by boring into
decaying wood directly beneath fungus sporophores and up into
the woody part of the sporophore itself.

FAMILY BELIDAE

BELID BEETLES

The New York weevil, Jthycerus noveboracensis (Forst.), is the
only known member of this family. It is rather widely distributed
in the Eastern and Central States, but seems to be most abundant
in the Mississippi River Valley (193). Adults have been collected
from oak, hickory, beech, and various fruit trees, and the larvae,
from within twigs of oak and hickory. The adult is robust, shiny
black, and from 12 to 18 mm. long. A covering of ash-gray and
pale-brown prostrate hairs gives it a mottled appearance.

FAMILY CURCULIONIDAE

WEEVILS

This family is reported to contain more species than any other

in the animal kingdom (391). Many species are extremely de-
structive pests of agricultural crops. Many others are serious
pests of forest, shade, and ornamental trees, also of seedlings and
young trees in nurseries and plantations. The adults of many
species drill holes in fruits, nuts, and other plant tissues to feed.
The larvae generally feed inside fruits, nuts, seeds, buds, and the
stems of their host. The adults of most species are typical snout
beetles, the head bending downward and forming a well-defined
and usually curved beak. The antennae are clavate and elbowed,
and are located at about the middle of the beak.

The white-pine weevil, Pissodes strobi (Peck), occurs through-
out the range of its most common host, eastern white pine, in
eastern North America. In descending order of preference, its
most commonly attacked hosts are: eastern white pine, Norway
spruce, jack pine, Scotch pine, pitch pine, and red pine. Most
other conifers, both native and introduced, occurring within its
geographic range also may be attacked. The adult (fig. 75 A) is a
small, brownish weevil, about 4 to 6 mm. long. The snout is
curved and about as long as the prothorax, and the elytra are

209

marked irregularly with brown and white scales. Mature larvae
are yellowish-white, legless, and slightly longer than the adults.
The pupae are creamy-white and about as long as the adults.

Winter is spent in the adult stage in the litter. Adults emerge
from March to May, depending on locality, and feed on the succu-
lent growth of the terminal leaders of their host, usually on the .
bark about 7 to 10 inches below the dormant buds. Eggs are de-
posited in small punctures in the bark of the leader and hatch in |
a week or 10 days. The young larvae bore downward side by side
in a ring, feeding on the inner bark and outer wood. By late July

F-482574, 482573, 482572
FIGURE 75.—White-pine weevil, Pissodes strobi: A, adult; B, infested )
terminal of a young white pine; C, white pines deformed by weevil attack,

210

or late summer, they become full-grown and pupate in chambers
formed in the wood or pith. Adults appear in 10 to 15 days and
feed until late fall on both old and mature new growth. Some-
times, 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 recoverable volume; (2) lumber degrade
in the remaining volume. Studies in New Hampshire showed an
estimated volume loss of 40 percent in the sawlog portion of saw-
timber trees and 70 percent loss in the portion above sawlog
limits of merchantability. The average volume loss in pole-size
trees was 13 percent (744).

The first evidence of attack in the spring is excessive pitch flow
from feeding punctures on the preceding year’s terminal shoots
(468). Later, the new growth appears stunted, and finally, the
needles wilt (fig. 75 B). Trees up to 3 feet tall may be killed.
Killed terminals on taller trees are replaced by one or more
branches of the topmost living whorl assuming vertical growth,
resulting in crooked or forked stems (fig. 75 C). Trees suffering
this type of damage for several years become multiple-stemmed,
cabbage-shaped, and worthless.

The white-pine weevil has been studied intensively during the
past several years, and much has been learned about its biology,
ecology, and control. As a result of these investigations, the fol-
lowing management practices have been recommended for reduc-
ing losses to white pine: (1) the planting of white pines with
hardwoods or under a hardwood cover; (2) the planting of white
pine on medium soils only where soil matting or hard pan does
not occur within 3 feet of the surface and where the trees will not
suffer from competition with hardwoods or jack or red pines;
(3) the selection and improvement by pruning of the least injured
pines for a final crop in heavily infested stands; and (4) the
removal of less desirable trees from white pine stands (467, 705,
706, 303, 56, 796, 507, 159, 598). Other types of indirect control,
such as that exerted by insect parasites and predators and birds,
are helpful in preventing excessive high weevil populations, but
are incapable of preventing intolerable levels of loss. Harman &
Kulman (321) published an annotated list of the parasites and
predators of the species.

The northern pine weevil, Pissodes approximatus Hopk., 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 spruce. The adult is very similar to the adult of
the white pine weevil but is somewhat larger, being about 5 to 8
mm. long. The beak is also somewhat larger, and the spots on the
elytra are uniformly smaller, the posterior ones rarely connected

24h).

Winter is spent mostly in the adult stage in the duff and top
soil beneath infested trees and under scales and crevices of the
rough outer bark of these trees. The remainder overwinter in the
larval and pupal stages in the tree. Overwintering adults emerge

211

in the spring and feed for several weeks on the inner bark of pine
branches and on the stems of seedlings and small trees. Adults
developing from the overwintering larval and pupal stages appear
in July and August. The life cycle may be completed in 1 or 2
years.

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 one-half inch 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. Normally, a pocket contains only one egg, but
frequently as many as four or five are found in one. The larvae
feed either upwara or downward or right or left from the site of
the egg, usually following the grain of the wood. In large, in-
fested material, they pupate in “chip cocoons” in the outer sur-
face of the wood; in smaller diameter material, they pupate at the
center of the stem.

Damage is often severe in regions where there are large quan-
tities of breeding material, such as fresh stumps in Christmas
tree plantations and in stands under intensive management. -Be-
cause 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 render-
ing them unsuitable for egg laying (250).

The deodar weevil, Pissodies nemorensis Germ., occurs in the
Southern States north to Pennsylvania and breeds in deodar
cedar, imported Atlas cedar, cedar of Lebanon, and various
species of pines. The adult is rusty-red to grayish-brown, has a
long snout, and is about 6 mm. long (fig. 76).

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 76.—Adult of the deodar weevil, Pissodes nemorensis.

Eggs are deposited in the inner bark of laterals and terminals
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 during March. Adults appear-
ing in April disappear immediately and aestivate until fall when
they again appear. Then they feed on the twigs and leading
shoots, puncturing the bark and eating large areas of the inner
bark and wood around the holes. Weakened deodar cedars are
often seriously injured or killed by the species. Damage to
healthy trees is less serious, consisting of the killing of leaders
only. Small pines may be girdled and killed.

Several other species of Pissodes also attack various coniferous
species in eastern forests. P. affinis Rand. 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 50 feet tall. They are dark brown
or black, from 5 to 8 mm. long, and are marked with white spots.
P. dubius Rand. breeds in windthrown, dying, or recently dead
balsam fir and red spruce. It is probably the most important in-
sect attacking dead and dying balsam fir following spruce bud-
worm defoliation (700). It also commonly attacks balsam fir
weakened or killed by the balsam woolly aphid. P. rotundatus
LeC. and P. fiskei Hopk. breed in windthrown red spruce.

The genus Hylobius is represented in North America by six
species, several of which are important pests. Millers et al. (532)
published a key to five of the species.

The pales weevil, Hylobius pales (Herbst.), a very destructive
pest of young pines, occurs in southeastern Canada from Nova
Scotia to southern Ontario, and throughout eastern and southern
United States west to the Great Plains. It breeds in all species of
pines within its range, and occasionally in spruce, fir, hemlock,
juniper, larch, cedar, and cypress. The adult is an oblong, robust,
black to reddish brown weevil, about 6 to 10 mm. long. The elytra.
are covered with small, scattered patches of yellowish hairs
(fig. 77 A).

In the north, winter is spent either as adults beneath the litter
or as larvae in the roots; in the South, it is spent as adults in the
soil. Overwintering adults emerge from April to June, depending
on location, and feed for a brief period on the tender bark of the
twigs of saplings and at the bases of seedlings. This feeding
occurs at night, the weevils hiding in the soil about the bases of
the seedlings during the day. After a few days of feeding, they fly
to recently cut over or damaged pine land. Here they feed, mate,
and lay eggs in the roots of pine stumps or weakened trees. Occa-
sionally, they burrow through the soil for distances of a foot or
more to reach a root. The larvae feed beneath the bark until early
fall. Then they form cells in the sapwood where they pupate.
Adults appearing during late September and October feed for
awhile on pine bark, twigs, and needles and then enter the soil to
hibernate. In the North, the life cycle usually requires 2 years; in
the South there are 1 and usually a partial second generations per
year.

213

Boo

F-514141, 514140

FIGURE 77.—The pine reproduction weevils: A, top, pitch-eating weevil (p.

217), middle, pales weevil (p. 218), and bottom, deodar weevil (p. 212);
B, adult pales weevil feeding on bark of pine seedling.

Prior to 1940 when it was first observed damaging pine repro-
duction in North Carolina (42), the pales weevil was a pest of
importance only in the Northeast. Since then, it has developed
into one of the most destructive insects in cutover pine lands in
the South. The first evidence of its attack is a series of small holes
or pits made in the bark by feeding adults. On lightly infested
trees these holes fill with oleoresin and heal over. Continued feed-
ing causes the damaged areas to merge (fig. 76 B). Heavy feeding
results in the girdling and death of seedlings up to one-half inch
in diameter. During fall feeding, the adults may feed on the
terminals and twigs. On large trees, damage is not serious, al-
though the ends of damaged branches may be killed. In contrast,
small trees in nurseries and plantations, as well as natural repro-
duction, are severely damaged or killed. Damage in Christmas
tree plantations is also severe. The girdling and killing of the
lower branches of residual trees makes them unfit for the Christ-
mas tree trade.

Damage to young pines planted in cut-over or disturbed pine
lands in the South can be reduced by planting only in areas cut
over or disturbed before July. In the North, the waiting period
should be 2 years. Where stands are to be established by direct
seeding in the South, seeding should be delayed for one year.
Before seedlings are planted on recently cut-over pine lands, on
areas adjacent to recent cutting or site preparation, or in places
where trees have been killed by fires or other disturbances, they
should first be chemically treated (6&3).

214

The pales weevel has been the subject of considerable interest
for many years, and several publications on its biology, ecology,
and control have been issued (245, 589, 682).

The pine root collar weevil, Hylobius radicis Buch., breeds in
the root collars of healthy pines in southern Canada from New-
foundland 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 mugho pines are also attacked. The adult (fig.
78) is dark reddish-brown to black and about 10 to 12 mm. long.
It is marked with irregular patches of white-to-yellow hairlike
scales, and the elytra have longitudinal rows of elongated
indentations.

In most localities, eggs are deposited from early May to late
September in cavities in the inner bark at the base of the tree, or
in the soil nearby (640). The larvae feed downward in the inner
bark of the base of the trunk and in the bases of large roots
below the ground line, widening their galleries as they develop.
Galleries are also formed in the soil around the base. The follow-
ing winter is spent in the larval stage in the Lake States. In
Southern Ontario, it is spent in the larval and adult stages and
occasionally in the pupal stage (246). 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 where they 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 (785).

Heavily infested trees may be severely injured (fig. 79). Small
trees, from 1 to 4 inches in diameter are most severely damaged—
smaller ones are seldom attacked. The presence of pitch flows at

F-488105
FIGURE 78.—Adult of the pine root collar weevil, Hylobius radicis.

215

ie

F-488104

FIGURE 79.—Damage by the
pine root collar weevil,
Hylobius radicis, at the
base of a pine tree.

the root collar and layers of pitch-infiltrated soil near damaged
areas are evidences of infestation. A number of silvicultural prac-
tices have been suggested for reducing populations (780). Finne-
gan and Stewart discussed methods of direct control (251).

Hylobius rhizophagus M. B. & W. is a newly-described species
recently discovered attacking jack pine in the Lake States (532).
The adult is black and about 14.5 mm. long. Dense patches of
coarse scales occur on the dorsum and the elytra bear longitudinal
rows of pits containing fine setae. The larvae feed in roots less
than one-half inch in diameter, tunneling from the smaller end
toward the base. Pupation occurs in cells in the roots. Pole-sized
pines in closed plantations on formerly cultivated land are most
frequently infested; reproduction in well-established infestations
and red pines growing in mixture with jack pine are also attacked
occasionally.

Hylobius warrent Wood attacks most species of conifers grow-
ing on moist to wet sites in southern Canada and south to North
Carolina in 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
1 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 are reported to have been killed by the species. H. pinicola
(Couper) occurs throughout the same range as H. warreni and
apparently breeds in the same species of trees. The adults of the
two species are similar in appearance, but differ in that the hind-
wings of this species are fully developed. Warren (741) described
H. warreni in detail.

216

Hylobius congener Dalla Torre, Schenkling and Marshall occurs
from the Northeastern Coast to Alaska and breeds in the inner
bark on logs and stumps of red, Scotch, and white pines. Adults
resemble the adults of the pales weevil but are slightly smaller.

Hylobius aliradicis Warner is a newly-described species (740)
recently discovered attacking the roots of healthy young slash
pine seedlings in south Georgia (219). The larvae hollow out the
smaller roots and bore extensively in the root collar area. Heavily
infested seedlings are killed. Infestations in plantations up to 4
years old on disturbed sites appear to be the most serious.

Pachylobius picivorus (Germ.), the pitch-eating weevil, occurs
throughout eastern United States, but is most common in the
South. It attacks various species of pines, the adults chewing on
the bark of the twigs and the larvae boring in the roots. Damage
is often serious in recently cut pine areas, especially where plans
call for the establishment of a new crop of seedlings during the
first year after the cutting of the old stand. Adults are dark
brown, robust, and clothed with patches of short, flattened, yel-
lowish or reddish-brown hairs. The tibiae are thick with the outer
part enlarged; the tarsi are densely hairy underneath with the
third segment bilobed. ;

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 3 feet.
Eggs are deposited in the cambial region of roots as small as 5
mm. in diameter. Burrowing may occur as far as 25 feet 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. Where populations are heavy, these cells are formed in
the wood. Adults emerge when stump root systems are 6 to 11
months old, depending on the season in which the tree was cut,
and they fly to fresh cutting areas where they feed and repeat
the cycle. Adult feeding occurs mainly at night and on cloudy
days, mostly on seedlings near fresh stumps (708).

Damage varies with the season, and with the size and vigor of
the host. It is severe early in the spring following planting. All
of the buds, bark, and foliage is frequently removed all the way
down to the ground line. Portions of root bark also may be re-
moved to depths of 8 inches. Newly established seedlings, if
attacked at all, are usually severed. Later in the season, damage
consists mainly of the removal of patches of bark and puncture
wounds. Very little feeding occurs in July and August and only a
limited amount during the fall. Heavy mortality has been incurred
among seedlings planted within 3 months after overstory pines
were cut. Heaviest attack and seedling mortality occurred in east
Texas between March and June. :

Pine seedlings may be planted safely during the winter in east
Texas on areas cut earlier than the previous July. Farther north,
a longer wait may be necessary. Before seedlings are planted on
areas cut within 6 months, they should be chemically treated.

217

Stands may be cut during seed years without danger of excessive
losses to regeneration.

The poplar-and-willow borer, Cryptorhynchus lapathi (L.), an
introduced species first recorded in North America in New York
City in 1882, now occurs from coast to coast in southern Canada,
in Washington and Oregon, and south to Virginia in eastern
United States. Its principal hosts are poplars and willows, but it
also attacks alder and birch. The adult (fig. 80) is brown to black
and about 8 to 10 mm. long. The snout is as long as the head and
thorax combined, and the sides of the thorax, parts of the legs,
and the posterior half of the elytra are clothed with pale, pinkish
scales intermixed with large, erect black bristles. Full-grown
larvae are thick, whitish, and about 12 mm. long; the pupae are
whitish to yellowish and armed with a pair of strong brown hooks
at the tip of the abdomen (509).

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, tunneling in all directions and pushing out their
borings through small holes. In late fall, they hollow out small
chambers in the inner bark in which they spend the winter. Feed-
ing 1s resumed in the spring, with the larvae usually boring
around the branch or stem and often causing girdling. When
ready to pupate, the larvae bore upward and inward and con-
struct 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
eracks 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 honey-
combed with larval tunnels. Branches or entire small trees may
be completely girdled or so badly weakened by tunnels that they
break. All poplars and willows over 1 inch in diameter are subject
to attack, recently planted trees and nursery stock especially. The
most serious damage usually occurs near the base.

The pine gall weevil, Podapion gallicola Riley, occurs through-
out eastern United States and southeastern Canada. It breeds in
red, pitch, Virginia and scrub 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. They first appear as slight swellings on one side of the
stem. By the time the adults emerge, these swellings are larger,
generally ovoid, and taper gradually toward the distal end (fig.
81). 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 (776).

The arborvitae weevil, Phyllobius intrusus Kono, an introduced
species first recorded on nursery stock in Rhode Island in 1947, is

218

ee

F-519950 F-501808

Figure 80.—Adult of the poplar- FIGURE 81.—Galls in pine produced

and-willow borer, Cryptorhyn- by the pine gall weevil, Podapion
chus lapathi. gallicola.

now known to occur also in Massachusetts, Maine, Connecticut,
Vermont, New York, and Pennsylvania. It feeds on a wide variety
of species in the genera Thuja, Chamaecyparis, and Juniperus.
Certain varieties of T. occidentalis and C. obtusa appear to be
particularly favored. The adult is light brown with grayish wings
and is about 6 mm. long. The beak is short, not quite as long as
broad, and there are light, metallic green hairs on the legs and
antennae.

Eggs are deposited in the soil during May and June, and the
larvae feed on the roots of the host plant for the remainder of
the season. Winter is spent in the late larval or pupal stage, and
the adults appear from early May to July (415).

The arborvitae weevil is destructive in both the larval and
adult stages, 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, an intro-
duced species first recorded in the United States at Rochester,
N. Y. in 1923, is now known to occur at least as far west as Ohio.
It feeds on a fairly wide variety of hardwoods including elm,
maple, willow, cottonwood, pear, apple, and plum. Adults feed on
the leaves and young shoots of their hosts; the larvae feed on the
roots of various plants.

The Japanese weevil, Pseudocneorhinus bifasciatus Roelofs, an
introduced species first recorded in the United States at Westville,
Conn., in 1920, is now rather widely distributed in the North-
eastern States. It feeds on a wide variety of trees and shrubs such
as flowering dogwood, hemlock, red bud, azalea, rhododendron,

219

and the sprouts of oak, sycamore, and walnut. The adult is stout,
sluggish, brownish-gray with two black bands across the elytra,
and about 4.5 to 7 mm. long. The snout is almost as wide as long,
and the elytra are apparently fused. Eggs are deposited in folds
at the margins of dead leaves or leaf fragments. The female then
seals the free edge of the fold to the basal portions of the leaf,
forming a pod in which the eggs are enclosed. When the eggs
hatch, the larvae enter the soil and feed on the roots of their
hosts. Adults feed at leaf margins, cutting out notches and caus-
ing a ragged appearance. Infested plants are occasionally heavily
defoliated by fall.

Polydrusus impressifrons Gyll., an introduced species first re-
corded in New York State in 1906, feeds on various hardwoods,
chiefly willow, poplar, birch, and plum in New York and Connecti-
cut, and possibly adjoining States. The adult is rather slender,
uniformly light metallic green, and from 4 to 5.5 mm. long. Birch,
poplar, apple, and pear appear to be preferred hosts. Adults occur
on a much wider variety of trees, however, and probably feed on
them to some extent. Eggs are deposited around scars and beneath
loose flakes of bark in roughened areas. The cut ends of pruned
branches and twigs of seedlings and young trees are especially
attractive as oviposition sites. Young larvae drop to the soil, enter
it and feed on the roots of their hosts or other plants. Pupation
occurs in the soil, and the 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 succulent stems may be girdled and killed. Damage is
usually most serious in nurseries (581).

Thylacites incanus (L.), an introduced species, has been re-
corded from Massachusetts and Long Island. Its hosts are various
species of pine where the adults feed on the needles and the
larvae, on the roots. Adults are brownish, with metallic reflec-
tions, 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.

The black vine weevil, Brachyrhinus sulcatus (Fab.), a prob-
able introduction from Europe, is widely distributed in the
Northern States. It feeds on a wide variety of plants, yew, arbor-
vitae, and rhododendron, in particular. The adult is a brownish-
black, flightless weevil, from 9 to 12 mm. long. The thorax is
densely covered with rounded tubercles, each bearing a short
hair, and the elytra are often speckled with white. The hindwings
are reduced to mere pads.

Winter is spent mostly in the larval stage, but also, to some
extent, as pupae and adults at depths of 6 to 18 inches in the soil.
Adults emerge from early May to late July and are present during
the remainder of the season. Generally, they hide during the day
beneath pieces of earth, stones, sticks, dried grass, or leaves on the
ground, or beneath loose flakes of bark of their host. They feed at
night by climbing up the stems and eating notches or holes in the
margins of the leaves. Eggs are deposited on the ground in trash
or among soil particles, under loose bark, or among the leaves of
their hosts. The larvae enter the soil and feed on the roots of
various plants. On woody plants, they feed first on the young

220

rootlets; later, on the cortex of larger roots. Infested roots may
be stripped entirely of their bark for several inches, or they may
be gouged out on one side only. Pupation occurs in cells formed
eae es surface of the ground. There is one generation per year

Damage may be severe in nurseries, especially to yews, azaleas,
and rhododendrons. Clean cultivation and the rotation of seedbeds
and transplant beds, and allowing infested areas to lie fallow and
be thoroughly cultivated in alternate years, are helpful in pre-
venting damage.

The strawberry root weevil, Brachyrhinus ovatus (L.), a close
relative of the black vine weevil, occurs throughout the Northern
States. Seedlings of hemlock, arborvitae, yew, juniper, and spruce
are especially favored as hosts. The adult is similar to the adult of
the black vine weevil except for its smaller size and the absence
of white specks on its wing covers. The life histories of the two
species are also similar. This species also frequently causes heavy
losses in nurseries and to ornamental plants. Hemlock and blue
spruce seedlings have been seriously damaged in nurseries in the
Northeast.

The Asiatic oak weevil, Crytepistomus 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 chestnut appear to be its pre-
ferred hosts but it also attacks hickory, beech, dogwood, hazelnut,
and azalea (720). 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 1 year life
cycle. Apparently, 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 6 inches 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 mar-
gins 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.

Odontopus calceatus Say occurs from New England to Florida
and Louisiana and west to Michigan, Iowa, and Missouri. It feeds
on the foliage of yellow poplar, magnolia, and sassafras. Adults
are mostly black and about 2.5 to 4 mm. long. The front legs are
strongly incurved, and the antennae, mouth parts, and tarsi are
usually dark brown.

Adults spend the winter in leaf litter. They emerge and begin
feeding on swelling buds and stipules on warm days in late April
and early May in the latitudes in Ohio and Pennsylvania. In Flo-
rida and along the Gulf Coast they may emerge as early as
February. Later, as the leaves unfold, they also are attacked.
Feeding damage consists of rice-shaped holes in the leaves about
3 mm. in diameter. Eggs are deposited in the midrib on the un-
derside of the leaves during May and early June. The midrib
usually breaks at the oviposition site. The larvae bore into the
leaf and mine the interior, commonly side by side in one or two
groups. Occasionally, there are two mines per leaf. As the larvae

221

increase in size they extend their mines and extrude filaments of
dark fecal material from them. Pupation occurs in spherical
silken cocoons in inflated portions of the mine. Newly-emerged
adults feed on the foliage, sometimes chewing through the leaves,
but more often leaving the upper surface intact (fig. 82). This
results in chlorotic spots and produces a ‘“‘burned”’ appearance on
severely attacked trees. Adults stop their feeding by mid-July,
aestivate, then hibernate until the following spring. There is one
generation per year (126).

F-519311
FIGURE 82.—Adult feeding damage to yellow-poplar leaf by
Odontopus calceatus.

This species has been particularly abundant on yellow-poplar
in eastern Kentucky since 1960. Heavy infestations have also oc-
curred 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. Late spring frosts have contributed
importantly to the control of the species in the northern portions
of its range.

The willow flea weevil, Rhynchaenus rufipes (LeC.), 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 white birch, cherry, shadbush, 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 cir-
cular holes in opening buds and new leaves. Eggs are laid in pits

222

on the underside of leaves, and the larvae feed almost entirely
within the tissues of the leaf, forming large blotch mines. Pupa-
tion 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 (553). The apple flea weevil,
Rhynchaenus pallicornis (Say), feeds on apple, hawthorn,
winged elm, hazelnut, and quince from New York to Illinois and
Missouri. Adults are black and about 2.5 mm. long.

Rhynchophorus cruentatus (F.), the palmetto pill bug, occurs
from North Carolina 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 yellowish hairs on the legs, and is 29 to 30 mm.
long. It feeds on bruised terminal buds or on sap exuding from
wounded or recently felled trees.

Several species of Attelabus L. feed as leaf-rollers on the foli-
age of various trees and shrubs in eastern United States. Adults
are robust and from 3 to 6 mm. long. The beak is flat and about
as long as the thorax; the color is usually dull red to black; and
the elytra have rounded tips, leaving the last abdominal segment
exposed from above. The female cuts a slit from each edge to the
midrib of a leaf and then rolls the portion beyond the slit into a
solid roll. One or more eggs are deposited in the roll. The female
then gnaws the petiole partly in two, and the leaf eventually falls
to the ground. The larvae feed on the inner parts of the roll and
eventually pupate in the ground. A. analis Ill. and A. bipustulatus
F. commonly occur on oaks, hickories, and walnut; A. nigripes
LeC., on sumac and hickory; and A. rhois Boh., on alder and
hazelnut.

Adults of the genera Cimberis and Diodyrhynchus feed on the
staminate flowers of various pines and other conifers from Can-
ada to Florida. They are usually dull red, have flat snouts which
are about as long as the thorax, and are usually less than 5 mm.
long. Several species of the genus Hugnamptus feed on the leaves
of various hardwods. Dryophthorus americanus Bedel occurs very
commonly in hickory killed by the hickory bark beetle. Stenoscelis
brevis (Boh.) is common in dead, dying, and rotting hardwoods.

The subfamily Cossoninae, the broad-nosed bark weevils, con-
tains 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 is killed. Anderson (10)
described the larvae of a number of species.

Several species in the genus Cossonus Clairville 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. impressus
Boh., a dull black species,—in the sapwood of both conifers and
hardwoods (reported from Florida) ; and C. cossonus Boh. and
C. impressifrons Boh.—under the bark of hardwoods.

223

Species of Cossoninae that attack and damage wood in build-
ings deposit their eggs in crevices in the wood. Adults of certain
species reattack the wood from which they emerge, thus intensify-
ing the damage they’ve already caused. The more important eas-
tern species are as follows: Hexarthrum ulket Horn—damages
woodwork in old buildings, often reducing the wood to powder;
Tomolips quericola (Boh.)—damages seasoned coniferous wood
such as pine flooring and pecky cypress paneling; and Pselactus
spadix (Hbst.)—occasionally damages damp wood beneath build-
ings and salt water piling above the high water mark.

A number of species of the genus Graphognathus, better known
as white-fringed beetles, have been introduced into the Southern
States in recent years. The adults feed on the foliage of hundreds
of species of plants, including such tree species as pecan, hack-
berry, black gum, poplar, black jack 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 the longest. The
forewings are fused together on the inner margins, and the hind-
wings are rudimentary, thus the beetle can’t fly. Eggs are de-
posited in masses covered and held together by a sticky, gelatinous
substance which 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 (800).

The black elm bark weevil, Magdalis barbita (Say), breeds in
the trunks and branches of unhealthy, weakened 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 114 inches
long. Pupation occurs in oval cells just beneath the bark. There
appears to be one generation per year. The bark of heavily in-
fested trees may be literally peppered with small, circular, emer-
gence holes.

The red elm bark weevil, Magdalis armicollis (Say), occurs in
the Eastern States and southern Canada and breeds in dying or
recently dead elms or occasionally in dead branches of living,
suppressed elms. The adult (fig. 83) 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, scar-
ring both the inner bark and the wood. Pupation occurs in cells at
the end of the tunnels. There is one generation per year.

Magdalis perforatas 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 at or near the tips of branches. The larvae bore to the
center of the stem and then tunnel through the pith toward its

224

F-519947
FIGURE 83.—Adult of the red elm bark weevil, Magdalis armicollis.

base. Trees under 12 feet 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. Deep pits are
chewed at the bases of needle fascicles.

Several other species of Magdalis are also encountered in eas-
tern forests. M. austera Fall. and M. hispoides LeC.—on the
needles of white pine in the Northeast (596). 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 substiga Fall.—on young Scotch pines; M. olyra (Hbst.)
—on weakened hickories; M. salicis Horn.—on willow and chest-
nut; and M. barbicornis (Latr.) and M. pandura (Say)—on elms.

The genus Curculio L. is represented in eastern United States
and eastern Canada by 15 species, many of which are important
pests of acorns, hickory nuts, chestnuts, and pecans, and the seeds
of several other nut-bearing trees. Damage is often severe. The
entire seed crop of certain tree species is destroyed in some years.
Injury is caused by the larvae which feed on the contents of the
acorns or nuts and destroy them.

Gibson (290) revised the species of Cwrculio occurring in
North America, north of Mexico. Many of the old names found in
current literature were changed and many species described by
previous workers were placed in synonomy. Up-to-date informa-
tion on the distribution and hosts of valid species was presented.
The taxonomy, biology, and ecology of several eastern species
were discussed by Brezner (98).

The life history and habits of all species of Cuwrculio are basic-
ally similar: Adults are found from early April to late November
in southern United States and from early June to late October in
the northernmost parts of the species’ range. They usually appear
2 to 3 weeks before acorns and nuts begin to ripen. The female
drills a hole through the shell of an acorn or nut and excavates
one or more small chambers near or in its inner surface, then

220

deposits one egg in each chamber. The larvae feed on the nut
meat until full grown, then they cut exit holes through the shell.
Most of the larvae vacate the nuts in the fall in northern areas,
but some may remain in them until the following spring. Full-
grown larvae enter the soil, usually within 2 or 3 inches of the
vacated nut, and burrow down to depths ranging from 14 inch to
1 foot, where they form cells in which to pupate. Here they usu-
ally remain in diapause for 1 to 2 years, depending upon the
species, before pupation begins. Newly-emerged adults fly im-
mediately to acorn-or nut-bearing trees.

Curculio adults are light tan to brown, or red and reddish-
brown to black; they are densely clothed with hairs, and range in
length from 4.2 to 13 mm. The beak is long and slender, some-
times longer than the body in the female, but always shorter than
the body in the male. The antennae are 11-segmented, elbowed,
long, and slender; they are inserted into the beak near its middle
in the male; in the female, they are inserted near the middle or
near the base.

The pecan weevil, Curculio caryae Horn., ranges from New
York to Iowa and south to Georgia, Oklahoma, and Texas. The
adult is dark reddish-brown and is from 7.5 to 12 mm. long. The
female beak is slightly recurved and longer than the body; the
male’s is only slightly more than one-half as long as the body.
Adults appear in late summer, and the female deposits several
eggs in a single nut. The larvae, only two or three of which sur-
vive, devour the entire contents of the nut within a month. Ma-
ture larvae vacate the nuts, enter the soil, and remain there until
the second summer after entry before they pupate. Heavy tree
losses are often incurred in commercial pecan and hickory nut
orchards. In some pecan orchards, up to 65 percent of the crop
has been destroyed.

The large chestnut weevil, Curculio caryatrypes Boheman
[=proboscideus (Fab.)], and the small chestnut weevil, C. sayz
(Gyll.) [=auriger (Casey) ], were important pests of American
chestnut before the latter disappeared from the American scene.
Now, they are troublesome only in plantations of Asiatic chest-
nuts (106, 732). C. neocarylus Gibson feeds exclusively in hazel-
nut. Several other species feed in the nuts of various hickories
and pecan.

Several other species of Curculio breed in the acorns of a large
number of oaks: C. sulcatulus (Casey) in nearly all species
within its range; C. pardalis (Chittenden) in 19; C. orthorhyn-
chus (Chitt.) in 18; C. longidens Chitt. in 11; and C. humeralis
(Casey) in 10.

The genus Conotrachelus Dejean also contains a number of
species important as pests of acorns, nuts, and fruit. The adults
resemble adults of the genus Curculio. They differ, however, in
having shorter and less curved beaks, in having the body scaled
on the upper surface, and in having the breast grooved for recep-
tion of the beak. Life histories are also similar to those of the
Curculios, except that Conotrachelus adults appear in the fall and
spend the winter in the ground litter (288). Gibson (289) pub-
lished keys to the larvae, pupae and adults of acorn-infesting
species. Schoof (642) discussed the species occurring in the
North Central States.

226

The most destructive species is the plum curculio, Conotrachelus
nenuphar (Herbst.), which breeds in plum, cherries, peaches,
apples, and other fruits. Adults are dark colored, have two promi-
nent tubercles on the back, and are about 8 mm. long.

Three acorn-infesting species occur in the United States: Con-
panes naso LeC., C. posticatus Bohemian, and C. carinifer

asey.

Conotrachelus naso LeC. occurs from Maine to Florida and
west to Minnesota in eastern United States. It has been bred
from acorns of 16 species of oaks. The adult is shiny black to
light reddish-brown, and from 4.8 to 6.6 mm. long. The beak is
moderately stout and curved, and 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 become full-grown in 2 to 3 weeks. Then they vacate the
nuts and enter the soil to pupate. New adults emerge about 114
to 8 months later.

Conotrachelus posticatus Boheman 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, and their
bodies are 414, to 5 mm. long. The beak is feebly curved, longer
than the prothorax, and of the same length in both sexes. 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.

Conotrachelus carinifer Casey 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, and the prothorax is
black. The beak is curved 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.

Other common eastern species of Conotrachelus include (1)
C. juglandis Le.C.—breeds in butternuts; (2) C. retentus (Say)
—breeds in walnuts; (3) C. hicoriae Schoof and C. affinis Boh.—
breed in hickory nuts; (4) the quince curculio, C. crataegi Walsh—
breeds in the fruit of hawthorn; (5) C. aratus (Germ)—feeds on
the young shoots and leaf petioles of hickory; and (6) C. anaglyp-
ticus (Say—breeds in the tissues around fresh wounds on various
deciduous trees such as hickory, birch, beech, maple, chestnut,
and oak (105).

FAMILY SCOLYTIDAE

BARK BEETLES

The family Scolytidae comprises three groups: (a) the true
bark beetles which mine between the bark and wood, usually
engraving both, of the twigs, branches, trunk, and roots of vari-
ous trees; (b) wood-boring bark beetles which mine directly into

227

the wood and feed on the woody material in both the larval and
adult stages; and (c) ambrosia beetles which bore directly and
deeply into the sapwood, and in some cases into the heartwood,
and feed entirely on fungi which grow on the tunnel walls. Sev-
eral publications dealing with eastern members of the family are
available (41, 197, 70, 698).

Bark beetles are typically soft and yellowish when first formed
but soon harden and turn reddish to dark brown or black.
They are small, cylindrical beetles, from about 0.9 to 9.5 mm.
long. The larvae are small, white, curved, legless grubs with en-
larged thoracic segments. The heads and mandibles are usually
strongly sclerotized and darker in color than the remainder of
the body. As a group, the larvae are so similar in appearance to
larvae of the family Curculionidae that it is next to impossible to
separate the two groups in the larval stages.

When they attack a tree, twig, branch, or log, the adults con-
struct entrance tunnels directly through the bark to the phloem
or into the wood, depending on the species. The entrance tunnel
is only long enough to reach a nuptial chamber or directly into an
egg gallery. Pitch and sap exuding from these holes harden at the
surface in various forms of pitch or resin tubes on conifers. In
the case of ambrosia beetles, boring dust is pushed out through
the holes. Egg tunnels of true bark beetles are constructed from
the entrance tunnels along the surface of the wood, cutting
through the inner bark and often scarring the wood. Egg tunnels
of wood-feeding species are completely within the wood, while
those of ambrosia beeties are deep in the sapwood. The latter may
be simple, branched, or compound. The females of many species
deposit their eggs in niches cut into the sides of the tunnel, cover
them with frass, and plug the opening toward the tunnel. Others
deposit their eggs in larger niches or in grooves or layers and
cover them with boring dust. Still others, particularly some of
the ambrosia beetles, place their eggs free in the tunnels.

When bark beetle larvae hatch they feed away from the egg
tunnel, more or less at right angles to it. The larvae of many
species construct galleries with a specific, recognizable pattern.
Ambrosia beetle larvae, in contrast, remain in the egg tunnel and
feed on fungi growing on its walls. A large number of species
pupate in cells at the ends of larval galleries, either in the
wood, in the outer bark, or between the wood and bark. The
adults remain in those cells until their exoskeletons harden, some-
times even longer. When first formed, they are soft and yellowish;
later, after hardening, they become reddish to dark brown or
black. While there, they feed on the phloem, often consuming all
that remains after the larvae finish feeding. When ready to
emerge, the beetles gnaw holes through the bark to escape. The
majority leave a given tree within a few days. The Ambrosia
beetle adults emerge through their parent’s entrance tunnels.

Bark beetles either fly to and attack new host trees as soon as
they emerge or congregate under the bark of the host tree and
hibernate or wait until sexually mature before flying. 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 in flight, or outside the bark, to
find new host material.

228

Winter is usually spent in the tree in the egg, larval, or adult
stage. Activity is resumed with the advent of warm weather in
the spring. Old adults continue their egg laying in the extended
portion of galleries started the year before or in newly con-
structed tunnels in 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.

The true bark beetles comprise one of the most destructive of
all the groups of forest insects in the country. During an average
year, they are credited with killing 414 billion board feet of
sawtimber, an amount equal to at least 90 percent of all of the
insect-caused mortality and 63 percent of the total growth impact
in the forests of the country (729). Losses caused by these insects
are not confined to damage resulting from their feeding activities
alone; they are intensified and often greatly increased by disease-
causing organisms which the beetles introduce into the affected
trees.

Many environmental conditions operate to control the abun-
dance of bark beetles. When favorable conditions prevail, many
species increase rapidly to epidemic proportions. At other
times, their numbers may be low enough to escape notice. Some
of the better known factors affecting abundance are: food supply,
weather conditions, prevalence of natural enemies, and associated
fungi (41). Many species increase most rapidly when there is an
abundance of overmature or decadent trees, windfalls, slash,
lightning-struck trees, or trees weakened by crowding, fire,
drought, excessive moisture conditions, disease, or other causes.
Factors tending to hold populations in check include natural
enemies, low winter temperatures, and stands of healthy, vigorous
trees. The most important natural enemies are insect parasites
and predators and other closely related arthropods. Thatcher
(707) presented a list of the parasites and predators of southern
bark beetles, and Bushing (128) published a synoptic list of the
parasites of Scolytidae in the United States and Canada.

Key to Injuries by Scolytidae and Platypodidae

1. Entrance gallery leading directly through the bark to the
surface of the wood, where it is elaborated into one of

VWarlous: types of bark DULTOWS 30k 2,
Entrance gallery extending through bark into the wood or

Sveneimtoutheypluly ere Ne. or ort les ere (eiece ie: 23

2 linestoesmner bark Of Coniferous trees 22..2)...4.2)h ee. 3

Inethnesnner bark of deciduous: trees 2. 20.060 20

jp aelneexposedsroots or basal part of trumk =... 4

In main trunk or thick barked, primary branches ................ 6

Enemedimm-rtossmmall branchesied 42 2 ar eee il

Hliineaeinryse See a ee ten ner cee he), hh Bont giaey eet Thee OMe elem 16

TRC OME Sete er ee hehe eS) eh ieyecel or sty aoe Mele ee vee LS

4. In fresh stumps or the bases of living pines and spruces;
the entrance hole often marked by large pitch tubes;

229

10.

1 aU

230

the egg gallery of irregular elongate type, extending

downward ‘from entrances 22) ee eee

In older stumps or the bases and roots of dying trees;
preferring fermenting bark

Hylastes spp., Hylurgops pinifex

Beetles reddish brown, 5 to 9 mm. long; common in all

parts of the United States 22. == Dendroctonus valens
Beetles nearly black, slightly smaller; Southern States,
rarer ins Northeast 2+). Dendroctonus terebrans

In:thicker:bark of pines* =o 4-2 - ee ie
In: thicker bark°of spruces7. =) -6e- = ee oe
In thicker bark of standing and felled balsam fir; burrows
radiate; beetle reddish brown, 2.5 mm. long
Pityokteines sparsus
In thicker bark of decadent and felled juniper, arborvitae,
cypress, etc.; also causing injury by feeding on the
twigs:
Arborvitae, Northeastern States
Phloeosinus canadensis
Juniper, Eastern and Southern States
Phloeosinus dentatus
Cypress, Southern States .............. Phloeosinus taxodiu
Brown to black beetles about 3.6 mm. long. In healthy
southern pines; S-shaped egg galleries in the inner bark
Dendroctonus frontalis
Reddish brown to black beetles, 3.5 to 6.5 mm. long; some-
times in smaller trees or the tops of larger trees, but
usually in dying or cut trees; burrows radiate
Ips calligraphus
Ips grandicollis, Ips pint
Beetles 3 to 6.5 mm. long, reddish brown to black ; in
radiate burrows in the inner bark of trunks of dying or
CU pINeSs =e. Ips longidens, Orthotomicus caelatus
Brown to black beetles, 5 to 6 mm. long; longitudinal egg
galleries in mature, living spruce....Dendroctonus obesus
Reddish brown to black beetles, 2 to 4 mm. long; in radi-
ate or forked burrows in the bark of the trunk or
larger limbs of dying or cut spruce «. 4. :
Beetles shining black, 3 mm. long; forked burrows
Scolytus piceae
Beetles dull black or brown, 2.4 mm. long; burrows

radiater =. Polygraphus rufipennis
Beetles reddish brown, 3 mm. . long; 3 teeth each side at
rear end; radiate burrows “ Orthotomicus caelatus

Dark reddish-brown beetles, 3.5 to 5 mm. long; straight
or wavy egg galleries in the bark of living, felled, or
decadent larch trees _.... Dendroctonus simplex
Small bark beetles in decadent or felled larch
Scolytus piceae, Polygraphus rufipennis
Orthotomicus caelatus
In medium to small branches of pine .
In branches of spruce
In branches of larch
Scolytus piceae, Polygraphus rufipennis

12
16

12.

13.

14,

15.

16.

17.

18.

IG).

20.

In branches of balsam fir
Pityokteimes sparsus, Taenioglyptes fraseri
In slash usually but also in decadent and weakened
branchesmamcatOpsyet oc. ee eee eee
Usually in slowly dying, shaded-out limbs, less commonly
IMbeslasiveteus oiatweihnencal slitee | Sed Meee
In smooth, thin bark of white pine, less common in other
DIMES sie rer seats 3 Pityogenes hopkinsi, P. plagiatus
Ips pint, Pityophthorus cariniceps
Iinvroughersbarkcot-pine.slash).<. 2s coees Atta
In southern pines ...00..... Ips grandicollis, Ips avulsus
Pityogenes plagiatus, Pityophthorus pulchellus
Pityophthorus pullus
Injnorethern: pinessi2 5 45 ee Ips grandicollis, Ips pini
In southern pines only
Pityogenes meridianus, Pityoborus comatus
Most commonly in eastern white pine but also in other
pines
Pityophthorus consimilis, Pityophthorus nudus
In medium-sized branches of spruce:

In longitudinal forked burrows ............... Scolytus piceae
Imtradiatecburrows" 220068 Polygraphus rufipennis
In transverse forked burrows ........... Phloeotribus piceae

In the smaller branches of spruce
Taenioglyptes ruficollis, Pityophthorus
cascoensis, P. shepardi, P. tonsus
P. pulchellus, P. consimilis
P. nudus
NE WASSZOLADINGS =. eRe: Behe oO ee Go ne ee
In twigs of spruce
Various species of Pityophthorus, Taenioglyptes spp.
Iintwieseorslanehi* 23.2.0. sean... Pityophthorus spp.
In twigs of balsam fir
Pityophthorus spp., Taentoglyptes spp.
In twigs of juniper, arborvitae, cypress, etc.; young
adults eat out the bases of twigs causing them to wilt

Ole tosbreakeolt 3... oes ea. Phloeosinus spp.
In living pine twigs, boring through the bark and wood

to the pith:

IMBECOLDINGry st: ths Paci rta ie. Tibia. Myeloborus fwazi

ingawihite: pie: 2.6... dale Myeloborus ramiperda

InesvaGlousspINes ic. 200520. Pityophthorus pulicarius

In twigs of decadent or severed branches; principally
in the inner bark but often affecting the wood
Carphoborus bifurcus, Pityophthorus spp.
In the cones of various pines:

Wihite-pine cones... Conophthorus coniperda
Redepine: cones 04.42. 4..f0L Conophthorus resinosae
Hoblotly:pine:cones 2.2... Conophthorus taedae
Virginia pine cones... Conophthorus virginianae

Under the bark of the main trunk and larger branches
Oiehardwood treese 2aee 2a ert tee ER EAS wri hs
In the inner bark of the smaller limbs and twigs of
hardwood trees! So) ae ae 2S posit ac .

13
15

14

18

21
22

231

21.

22.

23.
24,

232

Larvae forming centipedelike engravings on the inner
bark, killing the trees. Young adults feeding on the
small twigs of living hickory

Scolytus quadrispinosus
In the inner bark of decadent or felled elms:
Making short, simple, longitudinal egg galleries;
young adults injuring small twigs in feeding
Scolytus multistriatus, S. mala
Making branched, transverse galleries; young adults
often hibernating in bark of healthy elms
Hylurgopinus rufipes

Breeding in the inner bark of decadent and felled ash;

making regular, transverse, branched engravings
Leperisinus aculeatus, L. fasciatus

Breeding in the trunk and larger branches of decadent or
felled birch, beech, sweet gum, and wild cherry; ir-
regular radiate galleries in the inner bark

Dryocoetes betulae

Engraving the inner bark and sapwood of decadent and
felled mulberry; adults hibernating in inner bark of
livine® treesin...... Ne ee Phloeotribus frontalis

Breeding in the inner bark of injured wild plum and wild
cherry:

In short, unbranched, longitudinal galleries
Scolytus rugulosus
In forked, transverse galleries
Phloeotribus liminaris, P. mississippiensis
In the thin bark of sumac trunks
Pityophthorus rhois, P. crinalis, P. scriptor

Breeding in the smaller limbs and twigs of hickory:

Short, unbranched burrows in the smaller limbs
Chramesus hicoriae
Very slender, branched, longitudinal burrows in twigs
Cryptulocleptus dislocatus

Breeding in the small branches of hackberry:

Unbranched, longitudinal galleries
Scolytus muticus, Chramesus chapuisit
Transverse branched galleries
Phloeotribus dentifrons
Breeding in the small branches of oaks and chestnut
Pseudothysanoes lecontei, Pseudopityophthorus spp.
Breeding in the small branches of basswood
Pseudothysanoes rigidus
Breeding in the small branches of redbud
Pityophthorus natalis
Breeding in the small branches of sweet gum
Pityophthorus liquidambarus

In the bark, wood, or pith of small branches and twigs
Of various hardwoods 0...0. 4 Hypothenemus spp.

Unstained pinhole burrows in the W00d ooo...

Stained pinhole burrows extending into the wood ..........

Very small pinholes in the living twigs of pines, extend-
ing through the wood to the pith and continued there
as egg galleries; entrance covered by pitch tube:

24
28

25.
26.

Zk

28.

29.
30.

ol.

Tingriedigpineuees fe css cok vee wl ett Myeloborus fivazi

In White PING wireenes bose oly Myeloborus ramiperda
In various eastern and southern pines

Pityophthorus pulicarius

Pinhole burrows in the wood or pith of dying or dead

hardwoods, branching into several egg galleries in the

sapwood or pith

Micracis swainei

In redbud and other hardwoods ............ Micracis suturalis

In-willowsand-redbud 2... Micracis meridianus

In pith of oaks, maple, etc.

Micracisella opacicollis

Ingpithokioaks ete: Soci oe! Micracisella nanula

Small beetles with the first joint of antenna normal:
In honeylocust, hackberry, and other hardwoods

Hylocurus langstoni

inevhickony. maple. etes..) << tees. Hylocurus rudis
In hickory:
New York to North Carolina ........ Hylocurus biorbis
Mississippi ................ Hylocurus bicornus, H. harnedi

Pennsylvania to North Carolina

Hylocurus spadix
Small, dull, yellowish-brown to black beetles; the basal

joint of the antenna club-shaped with long hairs:
Yellowish-brown; in oaks, hickory, and other hard-
woods, from Pennsylvania to Florida and Texas
Thysanoes fimbricornis
Dark brown; in oaks and maples in Mississippi and

Geontiawe nner Thysanoes lobdelli
Dark brown; in elm and rattan vine from Virginia to
AROK AS ee eee. a cea ee as Thysanoes berschemiae

Small, shining, reddish-brown beetles; wing covers
coarsely, irregularly punctured; in maple and hickory
Lymantor decipiens
In stained ambrosial burrows usually confined entirely to
the sapwood; in both coniferous and broadleaved trees
In stained ambrosial burrows, in both sapwood and
heartwood; in both conifers and hardwoods ..
In hardwoods; burrows of various types 0000.
In decadent or felled conifers; burrows compound ..
In living hardwoods; compound burrows in yellow-
poplat, Maple, oak, etc..2-......2 Corthylus columbianus
In dying or dead hardwoods, especially in recently cut
EDINTING Veer te cn lee a ah Pal heme ese wee eer
Constructing simple, unbranched, burrows in sapwood
of various hardwoods:
Beetles 2 mm. or less in length; eastern part of the
Wimiteds StateStes meme oe! e Xyleborus saxesent
Constructing branched burrows in various hardwoods

25
27

29

36
30
35

ol

32

233

32.

30.

34,

30.

36.

234

Constructing compound burrows in various hardwoods ..
In the sapwood of dying and recently dead hickories;
cylindrical beetles 4 mm. or more in length
Xyleborus celsus
In the sapwood of various species of dying and dead
hardwoods; beetles from 1 to 4 mm. long:
In beech, birch, maple, oak, etc.; Northeastern States
Anisandrus obesus
In branches of apple, maple, etc.; Eastern States
Anisandrus pyrt
In small branches and twigs of maples and beech;
Northeastern States? =. Anisandrus minor
In stumps and logs of hardwoods in New Jersey, New
York, Connecticut, and the upper Ohio Valley
Xylosandrus germanus
In narrow, black-stained burrows in various hardwoods;
beetles small And Tslender te 2 oe es ee
In notably coarser, black-stained burrows in har dwoods;
beetles larger and much stouter:
In birch, beech, maple, etc.; beetles brown to black
Xyloterinus politus
In poplars; Northeastern States; beetle with smoky
yellow stripe on each wing cover
Trypodendron retusum
In various birches; Northeastern States; stripe of
yellow on each wing cover
Trypodendron betulae
Beetles very slender, more than 3 times as long as wide;
antennal funicle 2-jointed:
Beetle reddish brown, wing covers all one color; in
many Nard woods.) eee Monarthrum mali
Beetle dark brown, wing covers with cross band of
yellow; in many southern hardwoods
Monarthrum fasciatum
Beetles less than 3 times as long as wide; antennal
funicle 5-jointed; in various hardwoods
Xyleborus affinis, X. xylographus, X. fuscatus
In rather narrow, compound, ambrosial burrows in the
wood of decadent or felled conifers:
In pine; Pennsylvania and New York
Xyleborus howardi

In pines/South Dakota’. = = Gnathotrichus aciculatus
In pine, spruce, larch, balsam, etc.; in Eastern and
Southeastern States Gnathotrichus materiarius

In notably coarser, compound, ambrosial burrows in
decadent and felled conifers; beetles stout, dark brown
to black, marked with lighter brown or yellow;

In pine; wing covers often with indistinct stripe of
smoky brown or black
Trypodendron scabricollis

Flatheaded beetles making compound ambrosial burrows
in southern hardwoods; beetles less than 5 mm. long:
In hickories, oaks, gums, cypress, etc.

Platypus compositus

33

34

In southern oaks and chestnut
Platypus quadridentatus
Flatheaded beetles making compound, ambrosial bur-
rows in southern conifers, especially in pines; more
than So Mm slONS = 24 2e0 Platypus flavicornis

The genus Scolytus Geoff. is represented in eastern North
America by a number of species of true bark beetles, several of
which are of economic importance, all except one that breeds in
deciduous trees. The adults differ from other bark beetles in
having short, thick, brown or black bodies, in having the outer
angle of the fore tibia produced into a curved hook, and in having
the ventral surface of the abdomen ascend abruptly to the rear.
In some cases declivity is concave or excavated and ornamented
by spines, tubercles, ete. Blackman (74) revised the genus.

The smaller European elm bark beetle, Scolytus multistriatus
(Marsh.), the principal vector of the Dutch elm disease fungus,
Ceratocystis ulmi (Buisman) C. Moreau, in the United States, is
an introduced species, having been first observed in North
America at Boston, Mass., in 1909 (132). Since then, it has
spread over most of the United States and into southern Canada.
Its hosts include all native and introduced species of elms and the
related species, Zelkova serata. Adults are small, dark reddish-
brown, shiny beetles about 3 mm. long. The under-side of the
posterior is concave and armed with a noticeable projection or
spine on the undersurface of the abdomen (fig. 84 A). The larvae
are typical, legless grubs, and about 3 mm. long.

Winter is spent in the larval stage under the bark, and pupation
and transformation to adults occurs in the spring. The adults
begin to appear about mid-May through holes made in the bark.
On heavily infested trees or cut material the bark may be liter-
ally peppered with these small “shot-holes.”’ Soon after emergence
and before entering other dead, dying, or recently cut elms for
breeding purposes, they ordinarily fly to the smaller twigs of
nearby living, healthy elm trees to feed. Feeding occurs in the.
crotches of twigs, usually in the outer perimeter of the crown,
but also occasionally in its center. It is while the beetles are feed-
ing in these twig crotches that healthy trees are inoculated with
the Dutch elm disease fungus. Beetles emerging from trees killed
by the disease or from fungus-infected logs carry the spores of
the fungus on their bodies. When some of these spores rub off and
become lodged in the feeding wounds, infection can occur.
Chances of innoculation of a healthy elm are greatest in the spring
and early summer when the long vessels of the tree are open and
functioning and are near enough to the surface for the beetles to
cut them while feeding.

After having fed on healthy elms, the beetles seek out suitable
places for breeding purposes. Preferred breeding material con-
sists of living elms severely weakened by drought, trees rapidly
dying from disease or injury, broken limbs, firewood, or any
recently cut elm wood. None of this material is ever attacked,
however, if the bark has been removed or if it has dried to the
point of cracking. The beetles bore through the bark and con-
struct egg galleries 1 or 2 inches or more in length in the inner

235

236

F-519943-4

FIGURE 84.—The smaller European elm bark beetle,
Scolytus multistriatus: A, adult; B, gallery pattern.

bark, grooving the surface of the wood, parallel with the grain
(fig. 84 B). Eggs are deposited in niches along the sides of the
gallery. The maternal gallery is typically unbranched and runs
parallel to the grain. This distinguishes it from the maternal
gallery of the native elm bark beetle which is Y-shaped or bira-
mous. The larvae feed in the inner bark and the surface of the
wood, angling away from the gallery. When they become full
grown, they form cells in which to pupate in the bark. During the
spring and summer, the life cycle may be completed in 35 or 40
days. In the United States there usually are two generations per
year. Farther north in Canada, there are one and a partial second
generations per year.

Because of its role in the transmission of Dutch elm disease,
the smaller European elm bark beetle has been the subject of con-
siderable research, and huge expenditures have been made in
efforts to eradicate or control it. Efforts to eradicate it proved
futile, but much progress has been made in research on its con-
trol. So far, the most effective known method of reducing losses
is the destruction of beetle breeding places (762).

Scolytus mali (Bechst.) (=sulcatus Say) occurs in Connecti-
cut, New York and New Jersey, and probably in surrounding
states. It breeds in apple, cherry, and elm. The 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 on the
ventral side. The fifth sternite is longer than the third and fourth
combined and the posterior margin of the abdomen is lacking in
the male. The adult occasionally feeds in the twig crotches of elm
and is capable of transmitting the Dutch elm disease fungus.
Dying and weakened limbs and freshly cut wood are preferred for
breeding purposes. Winter is spent in the larval stage in the
bark, and there is one generation per year.

The hickory bark beetle, Scolytus quadrispinosus Say, a serious
pest of hickories, occurs from Quebec to Georgia, Alabama, and
Mississippi and west to Minnesota, Kansas, Oklahoma, and Texas.
It has also been recorded feeding on butternut and pecan. The —
adult is short, stout, black, almost hairless, and 4 to 5 mm. long.
There is a short curved spine or hook on the front tibia. The
venter of the male is deeply excavated. The third abdominal seg-
ment 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, and boring into the bark.
Here they construct rather short, longitudinal egg galleries be-
tween the bark and wood (fig. 85). 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 pockets
at each side of the gallery. The larvae feed in the phloem until
nearly full grown, and gradually angle away from the gallery.
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

237

BAN 6%,

*
a 3 . = A ees

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 85.—Galeries of the
hickory bark beetle, Scoly-
tus quadrispinosus, in
phloem of hickory. Note
short vertical egg galleries
and fan-shaped larval gal-

leries.

spring. There is one generation per year in northern areas. In the
south, there are normally two broods per year.

The hickory bark beetle is not only the most important insect
enemy of hickory, but also one of the most important insect pests
of hardwoods in eastern United States (41). During drought
periods, outbreaks often develop in the Southeast, and large tracts
of timber are killed. At other times, damage may be confined to
the killing of single trees or to portions of the tops of trees. The
foliage of heavily infested trees turns red within a few weeks
after attack, and the trees soon die. Control practices include the
felling of infested trees and destroying the bark during the
winter months or the storing of infested logs in ponds. To be
effective, this type of control should be conducted over large,
natural units.

The shot-hole borer, Scolytus rugulosus (Ratz.), an introduced
species, known to have been in the United States since 1878, now
occurs throughout most of the Eastern States and in several
Western States. It breeds in most of the common fruit trees and,
to a lesser extent, in mountain-ash, wild cherry, wild plum, haw-
thorn, 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 shallowly excavated at the rear
and unarmed. Because of its preference for broken, cut, or dying
material, this species is of minor economic importance as a forest
pest. There are one to three generations per year, depending on
locality.

Scolytus fagi Walsh breeds in beech and hackberry from Illi-
nois to Texas. Adults are 4.5 to 5 mm. long. The elytral striae are

238

distinctly impressed, and the strial punctures are much coarser
than those of the interspaces. The species is of slight economic
importance.

The hackberry engraver, Scolytus muticus Say, occurs from
New Jersey to Florida, Texas, Kansas, and Minnesota, and breeds
in dying and dead limbs of hackberry. 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 gal-
leries are similar to those of the hickory bark beetle. The larvae
feed first between the bark and wood; later they burrow into the
wood. Pupation occurs just beneath the surface of the wood.
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 Que-
bec to Manitoba, and from Maine to Colorado and Montana: Its
hosts are recorded as white, red, and Engelman spruces and
larch, and balsam fir. Adults are readily distinguished from other
eastern members of the genus by the presence of a tubercle or
spine extending backward from the center of the second ventral
abdominal segment. The burrows of the species also differ from
those of other eastern species in consisting typically of two and
sometimes three egg galleries extending longitudinally from a
central nuptial chamber. Broken limbs and tops are preferred
for breeding purposes. :

The genus Crypturgus Erichson is represented in eastern for-
ests by three common species. The adults are very small, brown
or black, about 1 mm. long, and are found in the inner bark of
dead or dying conifers. Their burrows usually originate from the
burrows of larger bark beetles, but sometimes from ventilation
holes made by Monochamus spp., and are very short. C. alutaceus
Schw., 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. atomus LeC. 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 Carphoborus Eichoff is represented in eastern for-
ests 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. bifwreus Kichhoff is a fairly common species
in the South. It breeds in dying, broken, and cut limbs of pines.

The genus Chramesus LeC. is represented in the East by two
species of stout, strongly convex, ‘humpbacked’ beetles, less than
2 mm. long. They are further distinguished by their large, long-
oval, unsegmented antennal clubs and the five-segmented anten-
nal funicle attached to the side of the club. Both species breed
in broken or dying twigs and small limbs. The adults construct
longitudinal, unbranched egg galleries, partly in the bark and
partly in the sapwood. C. hicoriae LeC. attacks various species of
hickory throughout the Eastern States and in eastern Canada.
C. chapuisi LeC. attacks hackberry from Pennsylvania to Florida
and Texas (75).

The genus Phloeotribus (—Phthorophloeus) is represented by
a number of eastern species, all but one of which breed in decidu-

239

ous 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 leaf-like 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 ab-
normal growths. These may show up as swellings on the trunks
of a badly infested tree. Infested trees are not killed, but they
may be seriously weakened.

Phloetribus frontalis Zimm. breeds in mulberry and is believed
to occur wherever its host grows in 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 female constructs a short transverse egg gal-
lery between the bark and wood, usually consisting of two short-
branched tunnels extending in either direction from the entrance
hole and deeply engraving the wood. The adults often cause seri-
ous damage when they bore into the bark and feed on the phloem
of living healthy trees. They tend to congregate in groups, and
several groups may occur in a single tree. When patches of bark
are killed they slough off.

Phloetribus dentifrons (Blackman) occurs in the South and
commonly in the Midwest. It breeds in hackberry. 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 over-wintering in their
burrows often destroy the gallery patterns during their prolonged
periods of feeding under the bark.

The peach bark beetle, Phloeotribus liminaris (Harris), occurs
in southern Canada and from New Hampshire to Michigan, Ten-
nessee, and North Carolina in eastern United States. It is pri-
marily a pest in peach orchards, but it also attacks elm, mul-
berry, wild cherry, wild plum, and mountain-ash. Breeding is
usually confined to weakened trees but the adults feed in the bark
of living trees. Adults are light brown to nearly black, feebly
shiny, sparsely clothed with long, whitish, fine hairs, and from
1.5 to 2.2 mm. in length. The galleries and habits of the species
are similar to those of P. frontalis, except that the galleries tend
to be somewhat more irregular. Damage in peach orchards may
be severe. In the forest, it is of minor importance.

The genus Dendroctonus Erichson contains many 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-seg-
mented; and the short antennal club is sutured toward the tip.
Hopkins (368) and Wood (793) published important papers on
the biology and taxonomy of the genus.

The southern pine beetle, Dendroctonus frontalis (Zimm.), the
most destructive of the eastern species of bark beetles, occurs

240

throughout the Southeastern and Southern States. From 1948 to
1966 over a billion board feet of pine timber was killed during
a series of outbreaks in the Southern States (fig. 86). It also
occurs as far west as Arizona and south to Honduras. During a
catastrophic outbreak in Honduras in the early 1960’s, about 10
billion board feet of pine was killed. It breeds in all species of
yellow pines in its range; also in white, red, and spruce pines,
and red and Norway spruce. Shortleaf, loblolly, Virginia, and pitch
pines appear to be most highly favored among the yellow pines.
Infestations in white pine are usually unsuccessful because of
heavy exudations of pitch. Attacks on red spruce are also unsuc-
cessful, the beetles dying in them after constructing short tun-
nels (438).

F-519566

FIGURE 86.—Stand of pines killed by the southern
pine beetle, Dendroctonus frontalis.

The southern pine beetle adult (fig. 87A) is chocolate brown
to black and from 2.2 to 4.2 mm. long. There is a prominent tu-
bercle on each side of a vertical median groove on the front of the
head; 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 legless grub with a glossy reddish-
brown head, and it is about 5 mm. long.

241

———

F-486343

FIGURE 87.—The southern

pine beetle, Dendroctonus

frontalis: A, adult; B, gal-

lery pattern. S-shaped

adult galleries are charac-
teristic of this species.

Winter is spent in the bark in all life stages—eggs, larvae,
pupae, and adults. Dates of adult appearance in the spring vary
with the overwintering stage 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 emerge is peppered with small, round holes (fig. 87B).
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 (367). Generally, they attack the middle
and upper trunk first, especially in the Middle Atlantic States.
Later they continue their attacks down the trunk to within 5
feet or less of the ground. In the Deep South they emerge in
March and may attack the lower trunk first.

The female bores directly into the cambium and constructs a
nuptial chamber. Points of attack are characterized by the pres-

242

ence of distinct pitch tubes, fine reddish boring dust, or white
resinous boring particles on the bark. 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 sur-
face of the wood faintly. The direction of the gallery eventually
is reversed thus creating a typical S-shaped or serpentine pattern
(fig. 87B). Eggs are deposited at intervals of one-eighth to one or
more inches 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 produce thread-like
mines. They are visible when the inner bark is exposed. As the
larvae develop, their mines become larger and are usually con-
cealed in the inner bark. As the larva approaches maturity, it
constructs a cell in the middle bark in which to pupate. Each
adult bores its own emergence hole. All of the adults in a given
brood may emerge during a period of 10 to 32 days. Three to five
generations per year have been recorded in western North Caro-
lina. In Virginia and West Virginia four and a partial fifth gen-
erations occur. In the Deep South there probably are five per
year.

Newly-emerged beetles may attack immediately adjacent trees
or they may fly to stands some distances away. Trees under 15
years of age and less than 2 inches in diameter are rarely at-
tacked. When outbreaks are developing, scattered groups of pines
in young dense stands are attacked. Once an epidemic is under-
way, stands of all age classes and densities are vulnerable. Spot
infestations, which may be anywhere from one-eighth to several
hundred acres in size, are characterized by a central zone of de-
foliated trees, a surrounding zone of red-topped trees, and a per-
iphery of infested trees with green or fading crowns. The needles
of pines infested during mid-summer 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 or from the effects of the blue stain
fungus, Ceratocystis minor, which the beetles introduce into the
tree.

Southern pine beetle outbreaks appear to be caused by condi-
tions that favor an increase in the vigor and size of beetle popu-
lations but which, at the same time, reduce the vigor of host trees.
Drought, overstocked stands, stand disturbances, and a reduction
in the abundance and effectiveness of natural enemies are prob-
ably responsible at times. In some parts of the region, especially
at high elevations and in the northern parts of the insect’s range,
low winter temperatures are often highly effective in terminating
outbreaks (39). A return to normal in conditions affecting the
size and vigor of populations as well as the vigor of trees also
appear to be helpful in the control of outbreaks.

Management practices designed to improve and maintain the
vigor of stands, and the removal of high-risk trees, such as those
struck by lightning or those attacked and weakened by other in-
sects, are helpful in preventing outbreaks. Once an outbreak is
underway, the salvage or chemical treatment of infested trees is
also helpful in suppressing populations. To be effective, these
practices must be applied before the beetles emerge from the
trees.

243

Several publications on the biology and control of the southern
pine beetle have been issued. This material is reviewed by Dixon
and Osgood (194) and Thatcher (707).

The black turpentine beetle, Dendroctonus terebrans (Oliv). oc-
curs from New Hampshire to Florida, Missouri, and Texas. All
species of southern pines and red spruce are attacked, but loblolly
and slash pines apparently are the most seriously injured. The
adult (fig. 88) 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 shal-
lowly punctate. Full-grown larvae are about 12 mm. long and are
marked with brown tubercles along each side of the body.

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 laid in the basal parts and large
roots of weakened and dying trees and in freshly cut stumps.
Green logs are also attacked occasionally. The female bores a hole
through the bark of the cambium. Here, she is joined by a male
and, working together, they excavate an egg gallery as large as
one inch wide and 20 inches long on the face of the sapwood,
usually in a downward direction. Eggs are deposited in a linear
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 cave-like galleries, usually somewhat fan-shaped, and occa-
sionally up to one foot across. Before they become full grown
they construct pupal cells either in 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.

ar D
SS Ay, Zi

5°

e: ae eo A,

Se nies

Bs
phe:
> Mag =
on pera cota

Rt

a re

F-486343-D
FIGURE 88.—Adult of the black turpentine beetle, Dendroctonus terebrans.

244

Freshly cut stumps are usually preferred for breeding; large
trees weakened by fire or other insects, or during naval stores
operations, are also highly susceptible. Occasionally, perfectly
healthy trees are infested. Attacks are usually confined to the
lower 6 feet of the trunks of standing trees. Sometimes, though,
they occur to a height of 12 feet. Infested trees are character-
ized by large, reddish to whitish pitch tubes on the bark surface
(fig. 89). They are also almost always invaded by ambrosia
beetles. The presence of the ambrosia beetles is signified by piles
of white sawdust around the base of the tree.

Prior to about 1949, black turpentine beetle damage was con-
fined largely to the killing of patches of bark on healthy pines.
The first evidence of its widespread killing of trees occurred dur-
ing an outbreak from 1949 to 1951 in Louisiana when several
million board feet and several thousand cords of pulpwood were
killed. Since then, severe infestations of slash, longleaf, and lob-
lolly pine have been reported from Florida, Georgia, Alabama,
Mississippi, and east 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 in
seed orchards also may be severe.

Some degree of natural control, but usually not very much, is
provided by other insects such as engraver beetles, borers, weev-
ils, and termites which compete with the larvae for food. In low-
lying areas, considerable brood mortality also occurs during peri-
ods of flooding. Losses may be prevented or reduced by the avoid-
ance of stand disturbance during logging operations or by the
rapid salvage of infested trees. Where the salvage of infested

F-519568
FIGURE 89.—Pine tree at-
tacked by the black tur-
pentine beetle, Dendroc-
tonus terebrans. Pitch
tubes are reddish to white
at first but soon assume a
grayish hue.

trees is not practicable, they may be sprayed to kill the broods
(668).

The red turpentine beetle, Dendroctonus valens LeC,, the larg-
est species of the genus, occurs in southern Canada and Mexico,
and in all the coniferous forests of continental United States ex-
cept in the Southern Atlantic and Gulf Coast States. It attacks
all species of pine within its range and, occasionally, spruce, true
fir, Douglas-fir, and larch. The adult is light reddish-brown to
dark brown and from 5.5 to 9 mm. long (fig. 90A). Full-grown
larvae are marked with brown tubercles on each side of the body
and are from 10 to 12 mm. long. 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 fre-
quently. 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 sus-
ceptible (664).

In the colder parts of the insect’s range, the winter is spent
chiefly in the adult stage, but also, to some extent, in the larval
stage. In the warmer parts of its range, adults fly intermittently
during the warmer, winter months. Attacks in standing trees are
initiated by females boring through the bark to the wood, usu-
ally in the basal 6 feet of the tree, but sometimes to a height
above 12 feet. Like those of the black turpentine beetle, these
attacks are characterized by the presence of pitch tubes on

F-494425, 494422, 494420

FIGURE 90.—Red turpentine

beetle, Dendroctonus val-

ens: A, adult; B, gallery

with mass of eggs along

the side; C, pitch tubes at
base of pine.

the bark (fig. 90C). 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. Eggs are laid in elongate masses along
the sides of the gallery (fig. 90B). The larvae feed gregariously
away from the gallery in the phloem. A fully-developed gallery
may be be from a few inches to more than a foot wide. Pupation
occurs in cells located between the bark and wood, either in the
gallery or a short distance forward in fresh phloem. Adults
emerge through holes chewed through the bark; sometimes sev-
eral use the same hole. In southern latitudes and at lower eleva-
tions, 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 two 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 not chopping into trees, digging up or
damaging roots, pushing deep earth fills over roots, or piling lum-
ber or green logs near trees. It is also helpful to debark or spray
freshly cut stumps and to cut and remove pines dying from other
causes. Watering or fertilizing of individual trees also increases.
their resistance.

The eastern spruce beetle, Dendroctonus obesus (Mann). (=
piceaperda Hopk.), occurs throughout the spruce forests of east-
ern America, and breeds in native, red, white, and black spruces.
The adult is dark brown to uniformly black and from 4.5 to 6.2
mm. long. Normally windfalls, prostrate dying green trees, and
overmature or weakened standing trees over 8 inches d.b.h. are
attacked. During epidemics, however, almost all trees are at-
tacked regardless of size or vigor. Attacks usually begin on the
lower third of the bole except for the first two or three feet
above the ground. Later in the season, they are continued upward |
and downward, exclusive of limbs and parts of the trunk less
than 8 inches 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 adults may
vacate their galleries and construct new ones in the same tree
or in nearby ones. They lay more eggs then die. The younger lar-
vae feed gregariously; older ones singly, in individual mines.
Pupation occurs in cells at the end of larval tunnels and the
winter is spent in the bark in either the larval or adult stages.
Signs of attack are red boring dust and pitch tubes on the bark,
the fading and dropping of needles, and the reddish appearance
of the twigs after the needles drop.

Several outbreaks have occurred in eastern forests during past
years. One, from 1897 to 1901, killed more than a billion board
feet of valuable spruce in Northern New England and eastern
Canada (368). The underlying causes of outbreaks are not well
understood, but piles of slash in mature stands are believed to

247

ee

produce heavy populations and trigger off outbreaks (699). 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, Dendroctonus simplex LeC., occurs in
the Northeastern States south to West Virginia and west to
Minnesota. It also occurs from coast to coast in Canada and north-
westward to Alaska. Its preferred host is eastern larch, Larix
laricina, 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 mostly as young adults in the brood gallery;
otherwise as larvae. Adults are active from May until late Au-
gust. Eggs are deposited in niches arranged in alternate groups |
of three to six eggs each along the sides of longitudinal, more or
less winding galleries. The larval mines are in the inner bark and
are quite short. Adults responsible for this brood may re-emerge
and construct several additional galleries during the season. Up
to three broods may be produced. The first of these reaches ma-
turity by mid-summer and the second, by mid-September. The
third brood spends the winter as larvae or young adults. |

The eastern larch beetle is not generally considered to be im- |
portant economically, because of its preference for dying or in-
jured trees. Living trees may be attacked and killed at times,
however, but are usually only after being weakened by other
causes, such as larch sawfly defoliation.

The genus Phloeosinus Chapuis is represented in North Amer-
ica by at least 40 species, five of which occur in eastern forests
(77). They breed preferably in cut, broken, or decadent conifers;
in short, longitudinal egg galleries constructed 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, goug-
ing out much sapwood. 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 red cedar, but it
also attacks arborvitae and Atlantic white-cedar. The adult is
piceous-brown to black, is clothed with rather abundant short,
gray hairs, and is 2.25 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. 91). Infestations are usually found in cut, broken,
or fire-damaged trees. Attacks have been reported on living over-
topped red cedars infested with the root rot fungus, F'omes an-
nosus, in North Carolina. Neither the insect nor the fungus
working alone usually killed the trees. Working together, how-
ever, they killed trees of all sizes (41). Cutting and burning of
infested twigs and keeping the trees in a healthy condition should
be helpful in control.

Phloeosinus taxodi Blackman, the southern cypress beetle,
breeds in bald cypress and probably occurs wherever its host
grows in the South. The adult is brownish-black to black, has

248

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, 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.

The other three species of Phloeosinus occurring in eastern
America are: (1) P, canadensis Swaine—on eastern red cedar,
arborvitae, and Atlantic-white cedar (in eastern Canada and
from Maine to the Lake States) ; (2) P. pint Swaine—on various
pines (in the Lake States and southern Canada); and (3) P.
texanus Blackman—on Mexican juniper (in Texas).

The genus Leperisinus Reitter contains a number of species
which breed by preference in various species of ash and are com-
monly known as ash bark beetles. The adults differ from most
other bark beetles in having variegated coloration, the markings
being produced by bands or spots of light colored scales alternat-
ing 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. Favor-
ite breeding material consists of recently cut or broken trees;
however, living trees weakened by mechanical injury, disease, or
fire may be attacked also. Parasites and predators, especially the
braconid, Coeloides scolytivorus (Cresson), and the clerid, Hno-
clerus quadriguttatus, provide a considerable degree of control
at times. :

Leperisinus aculeatus (Say), the eastern ash bark beetle, is the
most common eastern species. It occurs throughout the eastern

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COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 91.—Egg galleries and lar-
val mines of Phloeosinus denta-
tus, the eastern juniper bark
beetle, on trunk of red cedar.

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part of the United States and southeastern Canada and breeds in
various species of ash. The adult is grayish-brown with nearly
black markings and is about 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, the two arms being connected by a short tunnel or
nuptial chamber just below the entrance hole (fig. 92). Eggs are
laid in niches along the sides of the gallery. The larvae feed away
from the gallery, following the grain of the wood and deeply en-
graving it. Pupation occurs in deep oval cells between the bark
and wood. There are one to three generations per year, depending
on location. As the adults bore into the bark to hibernate, they
sometimes cause slight injury; otherwise, the species causes little
or no damage.

The genus Hylurgopinus Swaine is monotypic, only one species
occurring in North America. It is distinguished by its seven-
segmented antennal funicles, the strongly chitinized first two
sutures of the antennal club, and the widely separated fore coxae.

The native elm bark beetle, Hylurgopinus rufipes (Highh.), oc-
curs in southern Canada and throughout eastern United States
north of Alabama and Mississippi. It breeds in various species
of elm; also, reportedly, in basswood, ash, and wild cherry. The
adult is brownish-black, thinly clothed with short, stiff yellow
hairs, and from 2 to 3.5 mm. long (fig. 98). 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 abdomi-
nal sternites are red.

COURTESY OF
DUKE UNIV. SCH. OF FOREST.

FIGURE 92.—Transverse egg
galleries and vertical larval
tunnels of Leperisinus ac-
uleatus, the eastern ash
bark beetle, in white ash.

F-519939-40

FIGURE 93.—The native elm
bark beetle, Hylurgopinus
rufipes: top, adult; bottom,
gallery pattern.

Winter is spent in the bark of elm trees in either the larval or
adult stages. Overwintering adults emerge during May and fly
to living trees and feed in the bark. Later they fly to dead and
dying trees, broken limbs, or recently cut logs or limbs to breed.
Usually, dying or fairly moist dead limbs at least 2 inches 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. 98). The gallery may be constructed 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 gen-
erations per year in the southern portions of the insect’s range.
In the northern portions there may be only one or one and a
partial second (410).

When the adults emerge from elms dying from Dutch elm dis-
ease, they 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 parts
of the United States, the spread of the disease by the native elm
bark beetle is secondary to its spread by the much more abundant
smaller European elm bark beetle. Farther north in Canada,

251

where the latter species is either rare or absent, this species is
the primary vector of Dutch elm disease.

The genus Hylastes Erichson is represented by 21 species in
North America, several of which occur commonly in eastern
United States (76). 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 the roots of
stumps and in 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 current building operations. In the
Southeast, they fly from April to November.

Hylastes salebrosus Eichh. occurs commonly in the South At-
lantic States. It has been recorded breeding in loblolly, longleaf,
and shortleaf pines and spruces. It is also strongly attracted to
freshly sawn lumber. The adult is dark reddish-brown and about
4.5 mm. long.

Hylastes tenwis Kichh. occurs over most of eastern United
States from New York to Florida and Texas. Its hosts are listed
as shortleaf, longleaf, and loblolly 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. The adult is dark
reddish-brown and about 2.9 mm. long. Its hosts are not known,
but it has been collected while flying to downed pines.

The genus Hylurgops LeC. is represented in eastern forests by
only one species, H. pinifex (Fitch). It is widely distributed in
eastern United States and Canada and 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 Hylocurus Eichh. 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. Here they con-
struct slightly enlarged nuptial chambers. From each chamber,
one or 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.

Hylocurus rudis (LeC.) breeds in the twigs and branches of
hickory, walnut, maple, and hackberry throughout much of eas-
tern United States. The adult is dark brown to black and 2.2 to
3.0 mm. long. H. spadix Blackman, H. biorbis (Blackman), 4H.
bicornus (Blackman), and H. harnedi (Blackman) breed in
hickory.

252

Hylocurus langstont (Blackman) occurs in the South from the
East Coast to Texas. It breeds in the limbs and trunks of dying
or recently killed honey locust, hackberry, mulberry, and slippery
elm. Green poles and posts are also frequently attacked and dam-
aged. The damage resembles that caused by powder-post beetles.

The genus Micracis LeC. 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 red bud and willow. The adult is brown, and about
2.7 mm. long. The terminal hook of the anterior tibia is strongly
curved. M. suturalis LeC. breeds in the twigs and small branches
of red bud and walnut, and M. meridianus Blackman breeds in
the twigs and small branches of redbud and willow.

The genus Micracisella Blackman was described by Blackman
(78) to include certain species formerly included in the genus
Micracis. Female adults have the antennal scape flattened and
somewhat extended laterally and clothed with long hairs.

Micracisella opacicollis (LeC.) is widely distributed in the
Eastern and Southern States. It breeds in the pith of dead twigs,
sprouts, and shoots of oaks, maple, redbud, and cypress. The
adult is dark-brown to black and from 1.7 to 1.9 mm. long. M.
nanula (LeC.) breeds in oaks and redbud from South Carolina to
Florida and Texas.

The genus Thysanoes LeC. is closely allied to the genera Hylo-
curus, Micracis, and Micracisella, but differs in having the apex
of the elytra broadly rounded. T. fimbricornis LeC. breeds in the
twigs of red and black oaks, redbud, hornbeam, hackberry, hick-
ory, and acacia from Pennsylvania to Texas. The adult is yellow-
ish-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 gal- .
leries run parallel with the grain. The related species, T. lobdelli
Blackman has been observed breeding in oaks and maple in
Mississippi and Georgia and T. berschemiae Blackman, in elm
and rattan vine from Virginia to Florida and Texas.

Members of the genus Pseudothysanoes Blackman 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. leconter
Blackman attacks various oaks, hackberry, hop hornbeam, chest-
nut, and walnut from Maryland and West Virginia to North
Carolina. The adult is dark brown, shiny, and about 1.2 mm. long.
P. rigidus (LieC.) breeds in basswood from Canada to Michigan,
Ohio and West Virginia.

The genus Cryptulocleptus Blackman is represented by a single
species, C. dislocatus (Blackman), in eastern forests. The adult
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 unseg-
mented, 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 in West Virginia, the Carolinas, and Mis-
sissippi, and from acacia in Texas.

253

The genus Lymantor Lovendal is also represented by a single
species, L. decipiens (LeC.), in the United States. The adult is
reddish-brown and about 17 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
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
which are always present in the dead wood (698).

The genus Hypothenemus Westw. 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 Eichh. The latter is now recognized as a subgenus
by most American specialists in the family Scolytidae. Very few
species are economically important. However, a few of the seed-
infesting ones may be injurious at times.

Hypothenemus (Stephanoderes) dissimilis (Zimm.) is widely
distributed in eastern America from Quebec to Florida and west
to Michigan. It breeds in dying branches and dead twigs of vari-
ous hardwoods such as the hickories, oaks, honey locust, horn-
beam, and redbud. The adult is dark brown to black and from 1.2
to 1.8 mm. long. Deep entrance holes are bored 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.

Hypothenemus (Stephanoderes) rotundicollis (Eichh.) 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. (S.) quercus (Hopk.) attacks various oaks, hop
hornbeam, honey locust, and hickories in the Southeastern States.
The adult is dark brown and from 1.5 to 1.9 mm. long. H. (S.)
chapuisi (Eichh.) breeds in sassafras 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. (S.)
salicis (Hopk.) has been observed breeding in willow in North
Carolina, West Virginia and Pennsylvania. The adult is brown to
black and about 1.6 mm. long. H. (S.) interstitialis (Hopk.) at-
tacks oaks, hickory, and walnut. It has been recorded from Texas,
Mississippi, and North Carolina. 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. H. (S.) georgiae (Hopk.) has been
collected from the seeds of wild cherry, from pecan nuts, and
from the twigs of pine and several hardwoods in the Southeast.
The adult is dark brown and from 1.1 to 1.6 mm. long. H. (S)
obscurus (F.), an introduced species, has been recorded from a
wide variety of hosts, including black locust. H. eruditus Westw.
attacks dogwood, redbud, hickory, and black cherry from West
Virginia to Mississippi. The adult is dark brown to black and
from 1.1 to 1.3 mm. long.

The genus Pityophthorus Hichh., the largest of all the genera
of bark beetles, is represented by numerous species in eastern
forests. The majority of species breed in the inner bark of twigs
or small branches, but a few may be found in larger material.
Some species breed in the pith. Conifers, especially pines, are

254

usually preferred, but a few species confine their attacks to de-
ciduous trees. Attacks are usually confined to broken, cut, deca-
dent, or dying material, but perfectly healthy material may be
attacked. The adults are quite small, mostly from 1.3 to 1.75 mm.
long, and range in color from light brown to black. Females are
distinguished by the presence of long, yellowish hairs on the
front of the head (69, 70, 71, 698). A few common species of
Pityophthorus Eichh. are discussed here.

Pityophthorus natalis Blackman breeds in redbud from Mary-
land and West Virginia to Mississippi. The adult is reddish-
brown and about 1.5 mm. long. P. liquidambarus Blackman
breeds in sweetgum, probably wherever it grows. The adult is
reddish-brown and 1.8 mm. long. P. rhois Swaine, P. crinalis
Blackman, and P. scriptor Blackman breed in sumac.

Pityophthorus pulicarius (Zimm.) occurs throughout 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 killed by fire, in slash, and in one-year
old cones of felled pines. Scions of grafted slash pines being pre-
pared for seed orchard establishment in Florida have been seri-
ously injured. The adult is reddish-brown and 1.8 to 2.0 mm. long.

Additional eastern species of Pityophthorus and their hosts are
as follows: P. opaculus LeC.—white pine, larch, balsam fir, and
various spruces from Maine to West Virginia and South Dakota;
P. patchti Blackman—red spruce and balsam fir in New York;
P. biovalis Blackman—red spruce and red pine in New York and
Michigan; P. balsameus Blackman—balsam fir, red spruce, and
red pine in Maine and West Virginia; P. dentifrons Blackman—
red spruce from Maine to North Carolina; P. cariniceps LeC.—
white and red pines in the northern tier of states and southern
Canada; P. annectens LeC.—living trees and slash of various
pines from West Virginia to Florida and Texas; P. shepardi
Blackman—white and red spruce in Maine and New York; P. -
pulchellus Eichh.—probably all species of pines from Maine to
North Carolina and Texas; also red spruce and balsam fir; and
P. puberlus (LeC.)—all species of conifers from southern Can-
ada to North Carolina and westward to the Lake States and
Kansas.

The genus Pseudopityophthorus Swaine is closely allied to the
genus Pityophthorus. It differs in that the adults have a longer
and more acute prosternal process. The first segment of the an-
tennal 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. The genus was revised by
Blackman (72).

Pesudopityophthorus minutissimum (Zimm.), the oak bark
beetle, a common and widely distributed species from Quebec and
Massachusetts to Georgia and westward to Mississippi and Color-
ado, breeds in various species of oaks and occasionally in many
other hardwoods. The adult is dark reddish-brown and from 1.5
to 1.9 mm. long. In the southern portions of its range, it is active

255

throughout the year, and there are at least two generations per
year as far north as the Lake States. Eggs are laid in circum-
ferential galleries in branches from 1% to 4 inches 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 oak-wilted stands, this species is
strongly suspected of playing a primary role in the transmission
of oak wilt disease (116).

Other common eastern species of Pseudopityophthorus are as
follows: (1) P. pruinosus (Eichh.)—breeds primarily in dead
and dying oaks; also in beech, blue beech, hickory, maple, and
hop hornbeam. 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
1.8 to 2 mm. long. This species is also under suspicion as a pos-
sible vector of the oak wilt fungus (615). (2) P. pubescens Black-
man—breeds in various oaks, American hornbeam, and chestnut
in the Southeast. The adult is dark brown and about 1.8 mm. long.
(8) P. asperulus (LeC.)—breeds in various species of oaks,
chestnut, and gray birch from Maine to Florida and Texas. The
adult is dark reddish-brown and from 1 to 1.5 mm. long.

The genus Conophthorus Hopk. contains a number of species
which have the unique habit of breeding in the cones of conifers.
The adults are small, stout, dark brown to black, and about 1.25
to 4 mm. long. The female adult bores into a cone at the base and
then up through the axis, constructing a small tunnel and deposit-
ing eggs at intervals in small 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
reaching maturity.

The white-pine cone beetle, Conophthorus coniperda (Schwarz),
occurs throughout most of the natural range of its host, white
pine, in eastern America. 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 it and across the axis. Then she turns and bores along
the axis. Males join her at this time and the tunnel is extended
the full length of the cone (fig. 94). 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 a few
weeks. Some adults emerge during the fall but the majority do
not emerge 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 attack other in-
fested cones. There is one generation per year.

This is one of the most destructive insect pests of white pine
seed. During certain years entire seed crops in many stands in
New England are completely destroyed (572). Most of the dam-
age results from the killing of second-year cones; however, con-

256

F-505549

FIGURE 94.—Cone damage by

the white-pine cone beetle,
Conophthorus coniperda.

ey/
or ee a
—

siderable damage also results from attacks on first-year conelets,
shoots, and occasionally buds and male flowers.

The red-pine cone beetle, Conophthorus resinosae Hopk., occurs
in southeastern Canada and the northern tier of states from
Maine to Minnesota. It breeds by preference in second-year red
pine cones, but also attacks current-year red pine shoots and
occasionally second-year jack pine cones. The adult is shiny black,
with sparse short fine hairs and is about 3 to 3.5 mm. long (462).

Seasonal activity begins in May when the overwintering adults
emerge and attack current-year’s 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, the entrance tunnel often forming an
open groove at the cone base. The tunnel is extended to the pith
and is then continued 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, often near the
base. New adults remain inside the dead cones at least until they
become hard and black. During late summer, they 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 short,
current-year’s red pine shoots and then tunnel forward through
the pith into vegetative buds where they spend the winter.
Weakened at points of beetle entry, these shoots soon break off

207

and fall to the ground. There is one generation per year. Damage
by this species is often severe enough to make the commercial
collection of red pine seed impractical or impossible.

Conophthorus sp., a species closely related to C. resinosae, also
occurs in the Lake States and southern Canada. It breeds princi-
pally in the shoots of jack pine but also in Scotch pine. The adults
bore into the shoots about one inch below the bud then tunnel
toward the bud, either to feed or to deposit their eggs. The ovi-
position period lasts from late May into mid-July. Infested ter-
minals are often killed, leading to multiple branching and the
flat-topping of infested trees. Damage to natural jack pine repro-
duction and in jack pine plantations is often severe. Winter is
spent in infested buds, and there are two generations per year.

Conophthorus taedae Hopk. has been recorded damaging cones
of loblolly pine and Virginia pine in Virginia and C. virginianae
Hopk. in West Virginia.

The genus Pityogenes Bedel contains a number of species, all
of which breed in the twigs and thin-barked limbs 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. Occasion-
ally, young, healthy pines are attacked. 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.

Pityogenes hopkinsi Swaine occurs commonly in eastern Can-
ada and eastern United States. White pine appears to be the pre-
ferred host, but other pines such as jack and red, and red spruce
are also subject to attack. The adult is black with reddish-brown
elytra and is about 2 mm. long. The limbs and smooth bark of
recently killed trees may be heavily attacked. Shaded-out limbs on
large trees and weakened small trees in plantations may be at-
tacked and killed also. Destruction of infested slash or infested
trees by burning should be helpful in control.

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. Its burrows consist of two to five galleries originating
at and radiating away from a central nuptial chamber. The re-
lated species, P. plagiatus (LeC.), breeds in pines in the Atlantic
States. It is especially common in Pennsylvania and West Vir-
ginia. The adult of this species is somewhat similar to the adult
of P. hopkinsi. The male differs in having the dorsal tooth of the
declivity enlarged and hooked at the end.

The genus Pityoborus Blackman contains only one eastern
species, P. comatus (Zimm.). It is widely distributed in the South-
eastern 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.

258

Kgg galleries radiate in any direction, deeply engraving the
wood. The larvae construct wide, short, connecting galleries. The
adult is distinguished by the presence of a patch of fine, dense,
silky hair on each side of the pronotum of the female. The female
is also brown to black with yellowish appendages and is about
1.8 mm. long.

The genus Myeloborus Blackman is represented by only two
Species in eastern forests. The adults are very similar in appear-
ance to those of Pityophthorus, but differ in not having the
antennal club septate. They breed in the pith of living twigs of
pines. Entrance holes are bored directly through the bark and
sapwood to the pith. Here they are enlarged to form nuptial
chambers. Two egg galleries are then excavated away from the
chamber in opposite directions. The larvae feed on the wood, pith,
and bark, killing the twigs. Entrance holes are marked by small
but conspicuous white or cream-colored pitch tubes. These are
most frequently seen on the lower branches of the sides of trees
exposed to sunlight. Attacks on small trees may cause some dam-
age, but large trees are not seriously injured. M. ramiperda
Swaine breeds in white pine in eastern Canada and from Maine
i me Lake States. M. fivazi Blackman breeds in red pine in New

ork.

The genus Taentoglyptes (=Cryphalus) consist of small, dull,
dark brown to black beetles about 2 mm. or less in length. They
usually breed in the decadent bark of twigs or small limbs. T.
frasert (Hopk.) attacks Fraser fir in the Southern Appalachians
and balsam fir throughout the Northeastern States and in eastern
Canada. T. ruficollis (Hopk.) is found commonly in red and white
spruce in Maine and New York. T. rubentis (Hopk.) has been
collected from red spruce in West Virginia.

The genus Jps DeGeer ranks next in importance to the genus
Dendroctonus among the bark beetles in its destructiveness to —
forest trees, particularly the pines and spruces. Infestations nor-
mally occur in recently felled trees such as windfalls, snowbreak,
logging, and road slash. However, when heavy populations de-
velop 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 are characteristic of outbreaks.
Widespread outbreaks occur frequently during which losses may
be extremely severe. More than 500 million board feet of com-
mercial timber and more than 700 thousand cords of pulpwood
were killed in the South Atlantic and Coastal States during the
period of 1952-55 (707). It is estimated that annual losses of 300
thousand cords of pulpwood are incurred in Florida alone. Hop-
ping (369, 370, 371, 372, 378, 374, 375, 376, 377) arranged the
North American species of Ips into a number of natural groups.

The male initiates the attack by boring through the bark to the
wood and constructing a nuptial chamber. Here, he is joined by
three or more females, each of which excavates an egg gallery in
the phloem. These galleries radiate away in all directions from
the chamber through the phloem, but eventually tend to run par-

259

allel with the grain of the wood. The total pattern tends to form
a rough H or / shape. Eggs are deposited in small niches at irreg-
ular intervals along the sides of the gallery, and the larvae tunnel
in the phloem until full grown. Pupation occurs in cells hollowed
out in the inner bark. Young adults 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 either in the adult stage
under the bark of trees killed the previous year, or under bark
scales around the base of the tree. In the south, it may be spent
in the bark in all life stages.

Ips calligraphus (Germ.) occurs throughout eastern America
and breeds in practically all species of pines within its range. The
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 at times. Attacks on living trees
usually occur on the lower portions of trunks with diameters of
6 inches or more. In the south, this is one of the first species of
Ips to attack drought-ridden trees. It is also the first species to
attack thick-barked loblolly pines.

The adult is dark reddish-brown to black and from 3.5 to 6.5
mm. long. The declivity is deeply excavated and coarsely punc-
tured. 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. 95). 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 six or more
generations per year.

Ips grandicollis (Kichh.), the southern pine engraver, occurs in
eastern Canada and in eastern United States from Massachusetts
to Minnesota, Nebraska and Texas and south to Florida and Mis-
sissippi. Its hosts are shortleaf, loblolly, Scotch, Virginia, long-
leaf, pitch, jack, eastern white, and Caribbean pines. Recently
felled trees and slash are preferred, but the trunks and limbs of
apparently healthy trees are also subject to attack. Heaviest in-
festations 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 normally 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. 96). The larval mines are more or less
transverse. In the south, the life cycle requires from 20 to 25 days
and there are six or more generations per year.

Ips avulsus Hichh., the smallest of the eastern species of /ps,
breeds in all species of pines from Pennsylvania to Florida and
Texas. Thin-barked slash, such as the limbs and tops of pines, 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

260

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 95.—Gallery of Ips calli-
graphus on limb of a shortleaf
pine.

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 96.—Gallery of Ips grandi-
collis, the southern pine en-
graver, on limb of loblolly pine.
Note egg niches and incompleted
larval mines.

portions of the trunks by other species of Jps. Adults are attracted
to freshly-cut and injured trees. Any disturbance that causes
pitch flow may induce attack. Apparently healthy trees are also
attacked occasionally. Spot-killing occurs at times among pines
showing no evidence of previous injury or decreased vitality.

261

tl SEEPS SE 2 es EE ee

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. One to several long, winding egg galleries
originate from a central nuptial chamber (fig. 97). Larval gal-
leries 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.

COURTESY OF
DUKE UNIV. SCH. OF FOREST.

FIGURE 97.—Galleries of Ips
avulsus in bark of short-
leaf pine. Note radiate
tunnels of adults, short
larval mines, and pupal
chambers.

YE

i MEN. e

Tha iv u

The pine engraver, Jps pini (Say), occurs throughout the boreal
forests of North America and south to Tennessee in the Eastern
States. It breeds in several species of spruce and probably all
Species of pine within its range. Infestations usually develop in
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 oc-
curred in cut-over and burned-over areas in Canada.

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 eat irregular, meandering food tunnels, deeply en-
graving 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 (641).

Other less common eastern species of Jps include: (1) J. per-
turbatus (EHichh.)—breeds in white spruce in the Lake States
and Canada. (2) I. perrott Swaine.—breeds in red and jack pines
in Minnesota. (3) J. borealis Swaine.—breeds in spruce in the
Lake States. (4) J. latidens (LeC.).—breeds in white, red, jack,
and Scotch pines, white spruce, and hemlock in the Lake States.

The genus Orthotomicus Ferrari is represented by two species
in eastern forests. It is closely related to the genus /ps, but the
adults differ in having obliquely truncate antennae and feeble
teeth on the margin of a shallow concave declivity.

262

Orthotomicus caelatus (Eichh.) occurs throughout eastern
United 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, radiat-
ing 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 biological 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 (41). The western species, O. vicinus
LeC., breeds in black spruce in the Lake States.

The genus Pityokteines Fuchs 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, par-
ticularly firs and spruces. The eastern species, P. sparsus (LeC.),
is frequently injurious to balsam fir, killing large groups of trees.
Pines, spruce, and larch are also attacked. Infestations are found
in slash, in the limbs and tops of trees dying suddenly, in wind-
throws, 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 which radiate away from a central nuptial
chamber and scar the wood deeply. Larval tunnels are longitud-
inal and follow the grain of the wood.

The genus Dryocoetes Eichh. is represented in North America
by seven species, five of which occur in eastern forests (99). 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 (Mann.), the most common North American
species, occurs throughout the spruce forests of the continent
north of North Carolina and New Mexico. Spruces are preferred
hosts, but pines and larch 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 (Ratz.) 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 trees are
attacked including spruce, hemlock, Fraser fir and pines. 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, Dryocoetes betulae Hopk., occurs from
coast to coast in Canada and south to Florida and Mississippi in

263

the Eastern States. Its hosts are recorded as birch, beech, sweet-
gum, wild 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 Hopk. occurs throughout the northern conif-
erous 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 (LeC.), a rather rare species, breeds in
spruce from Quebec to North Carolina. The female adult is
reddish-brown and from 2.8 to 3 mm. long. D. piceae Hopk. is
often very abundant in windthrown spruce.

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 Appalachians. Its hosts, in addi-
tion 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 which radiate away from a central
nuptial chamber. The bark, but not the wood, is slightly engraved.

The Ambrosia Beetles
(Pin-hole Borers)

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. It was dis-
covered recently that female ambrosia beetles possess specialized
structures called mycetangia (291). They 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), it is found in both sexes (1). Since the discovery that
these specialized organs are possessed by ambrosia beetles, much
important knowledge regarding the relationship of beetles to
their specific microsymbiotic complexes has been gained (19, 37,
263).

About 36 genera of ambrosia beetles, some of which include up
to 200 species, have been recorded throughout the world. A
number of species breed in living trees, but decadent, dying, or
recently cut trees, logs, pulpwood or stumps are usually preferred.
All species require a considerable amount of moisture for de-
velopment. 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 (138).

Ambrosia beetles are important chiefly because of the degrade
of sawn timber that results from their invasion of trees or logs.

264

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 gum, cypress, and oak logs.

There are three types of ambrosia beetle tunnels; simple,
branched, and compound. Simple tunnels are unbranched, often
penetrating deeply into the wood. Branched tunnels penetrate
deeply into the wood and then break up into several branches,
which extend in various directions in the same plane. Compound
tunnels also branch off from a single entrance gallery but have egg
niches extending from the sides of the tunnel. As these tunnels
are excavated the beetles push the sawdust to the outside. The
larvae of certain species live and feed in so-called cradles which
branch off from the main galleries. Others live in the main gal-
leries. 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 (388, 698, 252, 734, 41).

FAMILY SCOLYTIDAE

This large family of bark beetles contains several species of
ambrosia beetles.

The Columbian timber beetle, Corthylus columbianus Hopk., oc-
curs from Michigan to Massachusetts south to Georgia and Ar-
kansas. It breeds in various living deciduous trees such as sugar,
silver, and red maple, sycamore, yellow poplar, boxelder, bass-
wood, beech, elm, yellow birch, and several species of oaks. Adults
are stoutish, black, and about 4 mm. long (fig. 98 A). The front
of the head of the female is convex and covered with short, stiff
hairs. 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.

Adults become active during May and June in Indiana and
West Virginia (407). They tend to reattack the tree in which
they develop, but some dispersal occurs. The wood is entered
through bark crevices, usually on the main trunk near the base.
Holes are bored straight into the sapwood until the tunnei nears
the heartwood, then it turns right or left (fig. 97 B). Entrance
holes are clean-cut and from 1/32 to 1/16 inch in diameter. Short
tunnels or chambers leading from the upper and lower surfaces
of the main tunnel 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 species Pichia (408). It is stored and trans-
mitted by prothoracic mycetangia possessed by the male beetle
(292). Winter is spent in both the pupal and adult stages in the
galleries of last attacks. 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

265

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 98.—The Columbian timber beetle, Corthylus columbianus: A, adult;

B, tunnels in the wood of a living white oak. Note that the entrance hole
had healed over several years before the tree was cut.

originating from the tunnels extend, often for considerable dis-
tances, 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 veneer, cooperage, or furniture. In
southern Indiana, red and silver maple wood, which is highly
valued in the furniture industry, is reduced in value by 38 percent
(499).

Corthylus punctatissimus (Zimm.), the pitted ambrosia beetle,
occurs from southern Canada to Georgia and westward to the
Great Plains. It breeds in a variety of trees and shrubs, such as
maple, dogwood, American hornbeam, hop hornbeam, sassafras,
rhododendron, and azalea. Young sugar maples are especially
subject to damage, destructive infestations in them having 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 about 4mm. 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 hood-
like over the head. The elytra are strongly punctured but not in

266

rows, are rounded behind, and are without furrows or teeth.

The adult bores into its host near the ground line, then exca-
vates a tunnel which may encircle the stem one or more times,
girdling it. Small stems, from 3 to 10 mm. in diameter, are usu-
ally attacked. Severe mortality of young sugar maples from 1 to
5 feet tall has occurred in dense, mature stands in Canada. In
some areas, almost ail of these young trees have been killed.
Finnegan discussed the biology of the species in Canada (249) ;
also its method of transfer of fungus spores (247).

Two species of the genus Monarthum, M. fasciatum (Say) and
M. mali (Fitch), occur in eastern North America. Adults of M.
fasciatum are marked with pale yellow bands across the elytra
and are about 2.5 to 3 mm. long. In this species, mycetangia are
possessed by the female (461). This species is most common in
the South. It breeds in most species of hardwoods, and has also
been observed in pine and hemlock. Adults of M. malt resemble
those of M. fasciatum except for their slightly smaller size and
their uniformly brown color. This species breeds in dying, in-
jured, or recently cut logs and stumps of practically all species of
hardwoods throughout the Eastern States. Both species are
highly destructive of green lumber and fresh logs cf gum in the
Gulf States.

Xyloterinus politus (Say), a widely distributed species in eas-
tern 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, ash, magnolia, black cherry, red spruce,
pine, and hemlock. The adult is dark brown to black and about
2.3 to 3.5 mm. long. The pronotum is almost square, rugose in
front, and has the anterior margin armed with two to four teeth.
The elytra are reddish-brown and covered with short, yellow
hairs. The galleries of this species differ from those of other am-
brosia beetles in that not only do they often fork and branch
secondarily, but they also possess four rows of larval cradles, two
above and two below the gallery (492, 493). Lumber cut from
infested wood may be severely degraded by adult entrance holes
and by associated stains.

Five species of the genus Trypodendron Stephens occur in
eastern America. Adults are distinguished by having divided eyes
and by the absence of distinct surtures in the antennal club. The
antennal funicle is four-segmented; the front of the male is
broadly excavated; the oral region of the head is visible from
above; and the corneous basal segment of the antennal club is
strongly angulate in front and produced toward the middle.

The striped ambrosia beetle, Trypodendron lineatum_ (Oliv.),
occurs throughout Canada and the Northern States and breeds in
a wide variety of conifers. It also occurs in the mountains of
western North Carolina. The adult is brown to black and from
3 to 3.5 mm. long. The elytra are dark brown to black, and are
usually marked with lighter colored stripes along the suture. The
main gallery of the species extends straight into the wood for 1 to
2 inches 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,
dying, or fire-scorched trees is often severe.

267

Trypodendron scabricollis (LeC.) occurs from New York to
North Carolina and Mississippi and breeds in various species of
pine and hemlock. The trunks (fig. 99) and larger limbs of
weakened and dying pines are preferred, but freshly cut lumber
is also subject to attack and serious damage. Adults are reddish-
brown and from 3 to 3.5 mm. long. The elytra are smooth and
finely striate.

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 99.—Tunnels of the ambrosia beetle, Trypodendron scabricollis, in
wood of the lower bole of a loblolly pine.

Trypodendron retusum (LeC.), the largest of the eastern
species, attacks poplars and paper birch throughout the Northern
States and southern Canada. It has also been recorded from West
Virginia and several Western States. Adults are from 3.8 to 4.6
mm. long; the pronotum is broadly emarginate in front; and
there is a broad, smoky yellow stripe on each elytron. T. betulae
Swaine occurs throughout the Northern States and eastern Can-
ada and breeds in birch. Adults are black and from 8 to 3.5 mm.
long. There is a faint yellow band toward the rear of the pro-
notum and a broad yellow stripe on each elytron. T. bivittatum
Kirby, the spruce timber beetle, occurs in eastern United States
and eastern Canada and breeds in pines, spruces, fir, arborvitae,
larch, and hemlock. Its color is similar to that of T. betulae, and
it is about 3 mm. long.

The genus Gnathotrichus Eichh. contains several species of
true ambrosia beetles. 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. Blackman (73) revised the genus.

Gnathotrichus materiarius (Fitch) occurs from eastern Canada
south to Florida and westward to Nebraska and Texas. It breeds
in the lower portions of the trunks of dead and dying pines,
spruce, balsam fir, larch, arborvitae, and other conifers. Adults
are dark brown to black and about 2.7 to 3.1 mm. long. The pro-
thorax is asperate in front; the disc is much longer than broad;
the elytra taper slightly at the rear end, and the declivity is
slightly grooved at the suture and covered with short, sparse
hairs.

268

Adults bore directly into the wood and their main galleries
may have several branches (fig. 100). Larval cradles are extended
both upward and downward from these galleries, and run parallel
to the grain of the wood. Pines killed by Ips bark beetles in the
South are often attacked by the species. The western species,
G. aciculatus Blkm., has been observed attacking ponderosa pine
in South Dakota.

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 100.—Tunnels of the am-
brosia beetle, Gnathotrichus ma-
teriarius, in the wood of a short-
leaf pine.

Ambrosiodmus lecontei Hopk. has been recorded from Florida
and North Carolina and has been collected from Asiatic Chestnut,
red maple, and walnut (41). The male adult is yellowish-brown
and 1.7 mm. long. The entrance galleries run straight into the
wood for a short distance and the egg galleries branch perpen-
dicularly and run generally parallel to the grain of the wood.
A. linderae Hopk. and A. tachygraphus (Zimm.) have been re-
corded from several eastern and southern States. A. linderae has
been collected from Asiatic Chestnut; A. tachygraphus from
yellow poplar, sugar maple, boxelder, beech, and birch.

Ansandrus pyri (Peck), the pear blight beetle, is widely dis-
tributed in southern Canada and the United States. It attacks all
of the common fruit trees as well as a great many other deciduous
trees. In North Carolina, it has been collected from yellow poplar,
chestnut, holly, honey locust, and hackberry (41). Dead, dying, or
weakened trees are preferred, but apparently healthy trees may
also be attacked. Female adults are black with yellow appendages
and are about 3 mm. long.

The genus Xyleborus Eichhoff is represented by 17 species in
the United States and Canada, 16 of which occur in eastern
United States. All species are ambrosia beetles. Many breed in
both coniferous 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. Distinguishing
characteristics of the adults are: a five-segmented antennal funicle
with the fifth joint short and broad; the outer face of the antennal
club obliquely truncate; and the eyes emarginate. The pronotum
is longer than broad and without serrations on the anterior mar-
gin. Beal and Massey (41) published keys to many of the species
commonly found in the eastern United States. Bright (100) pub-
lished keys to all species occurring in the United States and
Canada.

Xyleborus saxesenst (Ratz.) occurs commonly throughout
southern Canada and the United States and breeds in a wide

269

variety of trees. Some of its more important eastern hosts are
pecan, hickory, honey locust, walnut, sweetgum, yellow poplar,
dogwood, persimmon, holly, hemlock, cypress, and shortleaf and
loblolly pines. The adult is light brown to black and from 1.5 to
2.2 mm. long. The female scutellum is conical; the elytral declivity
steep and armed with rows of acute granules. There is also a
tooth-like tubercle on each side of the suture. The slope of the
declivity of the male begins so high on the elytra that the beetle
is humpbacked in appearance. The gallery of the species consists
of a single tunnel bored directly into the wood. Once inside it is
widened into a chamber in which eggs are laid and in which the
larvae live and feed. The life cycle is short and can be completed
in 2 months.

Xyleborus celsus Eichh., the largest member of the genus oc-
curring in the United States, breeds in dead, dying, and recently
felled trees and stumps of hickory in eastern United States west
to Kansas. Adults are light reddish to reddish-brown and about
2.3 to 4.45 mm. long. The declivity of the female is steep and
abrupt, the sides and upper margin are armed with several acute
granules, and there are four large teeth opposite the second inter-
space. The galleries of the species extend directly into the wood
to a depth of 12 to 18 mm., then branch one or several times.
Hickories killed by the hickory bark beetle are especially subject
to attack in the Southeast.

Xyleborus ferrugineus (F.) occurs from New York and Michi-
gan south to Florida and Texas. It breeds in dead, dying, or felled
trees of a wide range of species including oak, hickory, ash, cy-
press, walnut, pine, beech, and sweetgum. Adults are light reddish
to reddish-brown and from 2 to 3 mm. long. The elytral declivity
of the female is flat. The third interspace is armed with three
acute granules; the fourth and fifth interspaces are armed with
two to four granules each. Galleries resemble those formed by X.
celsus except that they are smaller; 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.

Xyleborus affinis Eichh. occurs in eastern United States east of
a line from Michigan to Texas and south of New York. It breeds
in dying trees and in green logs and lumber of various hardwood
trees such as oak, hickory, sweetgum, river birch, hackberry,
mimosa, persimmon, cypress, and black locust. Adults are light
reddish to reddish-brown and are from 1.8 to 2.8 mm. long. The
elytral declivity of the female is dull, opaque, and broadly slop-
ing, and the interspaces are armed with a few minute granules.
Galleries consist of short, transverse entrance holes and elongate
tunnels from which many transverse galleries often branch. This
species is very destructive of sweetgum in the Gulf Coast States.

Xyleborus xylographus (Say) occurs in eastern Canada and in
the United States west to Kansas. It reportedly attacks the lower
portions of dying trees and the stumps, roots, and slash of its
hosts. Oak appears to be favored, but hickory, chestnut, walnut,
and pine also are attacked. Favorite points of entry appear to be
the edges of wounds, deep crevices, or burrows made by other
insects. Adults are yellowish or reddish-brown to black and from

270

2.2 to 2.8 mm. long. The elytral declivity of the female is steep,
opaque, finely granulate, and marked by rather large striae punc-
tures. Galleries run obliquely across the grain of the wood at
depths of an inch or more (fig. 101). Eventually they branch and
the arms follow the grain.

Other eastern members of the genus and their hosts are as
follows: Xyleborus pint Eichh.—probably occurs in various hard-
woods and conifers; X. howardi Hopk.—pines; X. cavipennis
(Eichh.)—recorded from Rhizophora in southern Florida; X.
obliquus (LeC.)—birech, hickory, and chestnut; X. rubricollis
Eichh.—oak; X. lecontet (Hopkins)—hickory and palm in Flo-
rida; X. tachygraphus Zimm.—wide variety of deciduous trees;
X. obesus LeC.—wide variety of deciduous trees; X. sayz (Hop-
kins)—wide variety of deciduous trees; X. dispar (Fab.)—wide
variety deciduous trees and certain conifers; X. volvulus (Fab.)
—probably various deciduous trees and shrubs in southern Flo-
rida; and X. pecanis Hopk.—various hardwoods in Sauthern
States.

Xylosandrus zimmermant (Hopk.), an introduced species, has
been recorded in southern Florida. Its hosts are listed as Ardisia
sp., Ocotea catesbyana, and probably Chrysobalanus. X. germanus
(Blandford), also an introduced species, breeds in the branches,
logs, and stumps of elms in areas surrounding New York City
and in the Ohio River Valley. It also attacks many other species
of hardwoods. Heavy infestations have been found in elms killed

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 101.—Tunnels of the ambrosia beetle, Xyleborus xylographus,
in the wood of a shortleaf pine.

271

RN ___— eee

by the Dutch elm disease. Buchanan (115) demonstrated its
ability to transmit the Dutch elm disease fungus to healthy trees.

FAMILY PLATYPODIDAE

All members of this family are ambrosia beetles and they occur
principally in the tropics and subtropics. Only one genus has been
recorded from the United States. The adults differ from those of |
other ambrosia beetles in having longer and more slender bodies |
and wide heads flattened in front. The first segment of the tarsus |
is as long as all the other tarsal segments combined, and there are |
spine-like projections at the seam of the elytra of the males. |

Members of the family are usually more destructive than other .
ambrosia beetles. Their tunnels are more extensive, and they ex- |
tend deeper 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.

Eges are laid in small loose clusters in the tunnels. Larvae and
adults are also found in these tunnels.

Platypus flavicornis (Fab.) breeds commonly in various species |
of pines and occasionally in several hardwoods. Dead and dying |
trees, stumps, and logs cut or left in the woods during the summer
are preferred. 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 shal-
lowly punctured; and the elytra are elongate and striate, with the
third, fifth, seventh, and ninth interspaces produced into tooth-
like processes on the declivity of the male. The adult bores a
horizontal gallery in the sapwood (fig. 102). Here, it may branch
extensively and extend deep into the heartwood. The lower por-
tions of infested trees are sometimes literally riddled. In the
south, this species is so abundant that very few dying pines,

COURTESY OF DUKE UNIV. SCH. OF FOREST.
™ FicurRE 102.—Tunnel of the am-
brosia beetle, Platupus flavi-
cornis, in a chip from a recently
felled shortleaf pine.

272 |

stumps, or logs escape attack. Large amounts of white downy-
frass is evidence of attack.

Platypus quadridentatus (Oliv.) occurs throughout the South
and north to West Virginia and North Carolina. Various species
of hardwoods, especially the oaks, are most commonly attacked.
Magnolia seedlings have also been severely damaged in a nursery
in Florida. 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 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 de-
clivity of the male. Two large tuberosities also occur on the lower
edge of the declivity, and two hook-like spines are on the fourth
abdominal segment.

Platypus compositus (Say) occurs throughout the Southern
States northward to southern New York and southern Illinois. It
breeds in a wide variety of deciduous trees such as hickory, pecan,
birch, poplar, oak, chestnut, basswood, elm, beech, sweetgum,
sourgum, magnolia, persimmon, and cypress. Recently felled or
girdled cypress is often seriously damaged (fig. 103). 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 lo-
cated pits just behind the middle of the pronotum. The first,
third, and seventh interspaces of the elytra of the male are pro-
duced into small tubercles on the declivity. The declivity also bears
two large tridentate teeth at the outer apical angle.

Control of Ambrosia Beetles—Ambrosia beetle control is
largely a matter of prevention of damage to recently cut logs
through the regulation of woods practice, and in the proper hand-
ling of milled products (41). Control for several months has also ©
been obtained by spraying with an approved insecticide (404,
405).

COURTESY OF
DUKE UNIV. SCH. OF FOREST.

FIGURE 103.—Tunnels and
larval cradles of the am-
brosia beetle, Platypus
compositus, in the wood of
persimmon.

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 eastern
United States alone, and many are serious pests of forest, shade,
and ornamental trees. The adults differ considerably in appear-
ance from those in all other orders and are not difficult to recog-
nize. 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 which is carried coiled up like a watch spring beneath the
head. The wings are asually very broad, sub-triangular in form,
and 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
day time. (2) Moths usually have the wings wrapped around the
body, folded roof-like 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 hind-
wings at a different angle. (8) Moth antennae are usually
thread-like or feather-like; butterfly antennae are thread-like and
clubbed at the tip; the antennae of the skippers are usually re-
curved or hooked.

Lepvidopterous larvae are all very similar in structure. They are
usually cylindrical in shape, and, besides the head, the body is
composed of 13 segments, three 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 three to six and 10. Occasionally, some or all of
the prolegs are missing. A distinctive feature of the prolegs of
Microlepidoptera is the presence of fine hooks, known as crochets,
usually in a circle at the apex. The crochets usually 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 instar larvae 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 con-
siderable 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 parch-
ment-like cocoons, often with debris. The pupae of many others
are naked or are enclosed in slight cocoons of attached leaves.
Depending on the species, these 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

274

to leaves or twigs from which they hang head down. Some, such
as those of the families Papilionidae and Pieridae, are girdlers
and do not hang head down. Skippers pupate in cocoons made of
leaves fastened together with silk.

The order Lepidoptera contains several destructive forest and
shade tree insects. Several species such as the spruce budworm, the
forest tent caterpillar, and the gypsy moth often occur in out-
breaks covering tens of thousands of acres 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 be-
tween the upper and lower surfaces of leaves. The attractiveness
of shade trees, parks, and other recreational areas is often re-
duced or destroyed by these insects, fire danger is increased, and
wildlife habitats are impaired.

For a more extensive treatment of the order Lepidoptera the
reader is referred to the following publications: Borror and De-
Long (90), Dyar (213), Edwards (222), Forbes (258), Fracker
(262), Holland (364, 365), Klotz (422), McGugan (482), Pren-
tice (607, 608, 609), McDunnough (478), and Freeman (267).

Field Key to Some Common Lepidopterous Larvae Attacking
Eastern Forest and Shade Trees

LES PADD) Ste Vea OT Spe erect roe) er UNE OR RE RIEL SAVE TNL al ccccasvs 2
BOG CIS eae Be eek ae not a Rk Be ie ed EE IA ce. 15
2. Miners. ‘casebearers; bag worms 22 e i ee e. 3
Skeletonizers (leaves not webbed together) 00.0.0... 9
leeatrollerscandeleat tiers: =e A Re eccanee 10
Wielbworme cinodctent makers (ie ae da:
Hueco CCUei Sars aos Became ere EN EE ede a Se 12
3S Non=caseneatine leat MINers 266k 4
Casebearing leaf miners, other casebearers, webworms ..... i
At. Inthe rollage ot deciduous plants 2.72 ee. 5
Iinethesneedleswor Pinacede a. 25... JL Pe aan. 6
5. (1) Oak—
a. Very small, flat, and reddish brown or black.
SOMtAGye Lithocolletis hamadryadella.
b. Very small, flat, and reddish brown or black.
Gregarious. oe... Lithocolletis cincinnatiella.

(2) Tupelo—Head and cervical shield dark-brown; body
pale green and about 8 mm. long
Antispila nyssaefoliella.
6. (1) Arborvitae—

a. Head, cervical shield, and anal plate blackish;
body, reddish 3... Coleotechnites thuraella.
b. Head brown; body yellowish-white with shield-

shaped areas behind the head
Argyresthia thuiella.
ce. Head greenish or brownish; body yellowish-green

to green (also on red cedar)

Argyresthia freyella.

275

(2) Redcedar, juniper—Head, cervical shield and true
legs light brown; remainder of body pale green
with a pinkish tinge

(Colenteciites juniperella.

(3) Baldcypress, hemlock—Head, cervical shield, and
true legs pale brown; body greenish with brownish
CIC kt. of Serene Coleotechnites apictripunctella.

(4) Pines ‘Cane pitch, etc.) Head, anal plate, and true
legs dark brown; body pinkish. About 6 mm. long

Exoteleia pinifoliella.
(5) Spruce (Norway, white, Colorado blue)—
a. Head, cervical shield, thoracic legs, and anal plate
blackish; body dirty white to reddish. About 8
mm: long see ee eee eee Epinotia nanana.
b. Head and cervical shield light brown; body reddish
to light cinnamon brown
Coleotechnites piceaella.
ec. Head flattened and yellowish-brown; shields pale
greenish; body light greenish brown, semitrans-
parent, and sparsely clothed with hairs
Taniva albolineana.
i. “Case‘composediof partior parts otra leat ee eee
Caselitough oval. cilkent 72. =a eee Acrobasis spp.
Case baglike, silken, covered with bits of host plant
Family Psychidae.
8. (1) Sugar maple—Head amber brown; body dull white,
with a broad longitudinal stripe; about 6 mm. long
(case flat and circular)
Paraclemensia acerifoliella.

(2) Oak—Case ellipsoidal, portable, composed of bits of
leaves and silk; open at each end

Cincinnus melshermeri.

(3) Cases cigar- or pistol-shaped

Decl ae el ae, Teel” | sake Coleophora limosipinella.
bsvAliderwbinrchies: tc Moras Coleophora fuscedinella.
Cr ickonya pecan. ee Coleophora caryaefoliella.
dseAppleshaw thorny 2. eee Coleophora serratella.
6: archi shes wie x ce Se cose Coleophora laricella.
9. Skeletonizers (leaves not webbed together)
(Ui) i.O cake Meatvtneet eit te toned. EEE eee Bucculatrix ainslella.
(2) eBitchind: codecs Pie iene al Bucculatrix canadensisella.
(3) Apple, Hawthorn ................. Bucculatrix pomifoliella.
(4) Paper and river birches 2. Acleris logiana.

10. Leaf rollers and leaf tiers (some are skeletonizers)
(1) Basswood—Head black; body green. A leaf roller
Pantographa limata.

(2) Beech—Head brown; body pinkish. A leaf tier
Psilocorsis faginella.
(8) Elm—Body green; margins of segments tinged with

yellow. In silken web between 2 leaves
Canarsia ulmiarrosorella.
(4) Hickory—
a. Head, shield, tubercles black; body dull green with
black, prominent tubercles

Archips infumatanus

276

cea

b. Head pale green, tinged with brown; body pale,
tRanslucentse =. 2226. 5 x. Argyrotaenia juglandana.
(5) Black locust—

a. Head and cervical shield blackish; body green with
yellowish lines. ................ Nephopteryx subcaesiella.

b. Head light brown; body light green
Nephopteryx virgatella.
c. Head dull yellow; 2 yellow spots on lower part of
face; neck and sides of first thoracic segment
red; cervical shield black; body light green,

marked with fine black rings

Epargyreus clarus.

(6) Maple—
a. Head yellowish; body light green (In a trumpet-
HIKE GUIDE) pen ele tele ce etd le eo Epinotia aceriella.
b. Head and cervical shield reddish brown; body dull,
yellowish-green.................... Sparganothis pettitana.

c. Head tan; body yellowish-green
Sparganothis acerivorana.
(7) Oak—
a. Head and cervical shield dark brown to black;
bodyalightsoreenc.....- Archips argyrospilus.
b. Head amber yellow; body light green
Argyrotaenia quericifoliana.
c. Body greenish (between two leaves tied together
WG Ke te ee so ee Psilocorsis spp.
d. Head pale; thoracic legs brown to black; body
dirty white to light green. About 12 mm. long
Croesia albicomana.
(8) Pine— Head greenish-brown, with dark patches on
the sides; body greenish-brown
Argyrotaenia pinatubana.
(9) Poplar—
a. Head and cervical shield brown; body translucent ;
legs and tubercles black
Anacampsis innocuella.
b. Head blackish; shields mostly black; body dull

Oliveroreen ie Choristoneura conflictana.
c. Head large, dull brownish or reddish-brown ; body
Daleeenreenes te we eS ee Erynnis icelus.

d. Head brown; body light green
Amorbia humerosana.
(10) Redbud—Body white with black markings
Fascista cercerisella.
(11) Sassafras—Body largest at 3rd thoracic segment;
dorsum pea green and sides yellowish; black line
across prothorax; eyelike spots with blackish
centers on enlarged segment; pair of spots on Ist
apdominalV@segment. 22.2 5..0.........5 Papilio troilus.
(12) Spruce, fir—
a. Head black; body brown, with conspicuous pale
tubercles! wxif)..ih ccs Choristoneura fumiferana.

b. Head brownish or black; body bright green _
Acleris varianda.

277

ec. Head brownish yellow; body yellowish to grayish-
green (Opening buds and terminal needles
webbed together in the spring)
Zeiraphera ratzeburgiana.
(18) Sumac—
a. Body green, more or less tinged with red
Episimus argutanus.
b. Head yellow and brown; body yellow green, with
brick*redalines#a2ee= Nephopteryx sugfuscella.
(14) Sycamore—Body light green; tapers toward each end
Adoxophyes furcatana.
(15) Tupelo—Head yellowish; body black
Actrix nyssaecolella.
(16) Willow—
a. Head large, dull brownish or reddish brown; body
dull*pale=oreen: te % ee ee Erynnis icelus.
b. Head light brown; body light green
Amorbia humerosana.
(17) Various hosts—
a. Head brownish; body pale green
Choristoneura rosaceana.
b. Head light to dark brown; body dull green
Archips rosanus.
ec. Head dark brown or black; body light green
Archips argyrospilus.
d. Head black; body dark brown
Sptlonota ocellana.
e. Head yellow brown; body translucent, greenish
with 4 whitish, longitudinal lines
Dichomeris ligulellus.
f. Head large; body green and tapering
Machimia tentoriferella.

11. Webworms and tent makers

278

(1) Ailanthus—Body dark olive brown, marked with

WIC O IMCS oc a a anaes Atteva aurea.
(2) Wild cherry, apple—Body dark colored with a white
stripe on the dorsum ...... Malacosoma americanum.

(3) Apple, wild plum, cherry—Head light brown; body
dark brown to almost black, with a broken white
line on each side and 2 conspicuous reddish spots
on the back near the rear end

Nygmia phaeorrhoea.

(4) Cherry, elm, apple, persimmon, and many other species
—Body generally pale yellowish or greenish, with
a broad, dusky longitudinal stripe on the back and
a yellowish stripe on the sides; clothed with whit-
ish to reddish hairs arising from black and orange
warts ....... ; r Hyphantria cunea.

(5) Beech—Body yellowish green with faint stripes

Tetralopha spp.

(6) Birch, Willow—Body dark brown to black, with black
dots on the sides below which are spots that vary
from white to red FBulype hastata.

|

12. Larvae sluglike
13.

(7) Cherry—
a. Looper; body black and yellow (In nest of leaves
near end of the branch ......... Calocalpe undulata.
b. Head black; body yellow (in a dense web)
Archips cerasivoranus.
(8) Juniper—
a. Body brown and black
Dichomeris marginellus.
b. Head dark; body brownish yellow
Phalonia ruttilana.
(9) Maple—Body brown with yellow and black stripes
(ime smallenest) nc. ee ears. ce. 8 Tetralopha sp.
(10) Oak—
a. Head black; body gray green
Archips fervidanus.
b. Body brown with yellow stripes; legs and spiracles
Dacha ete acces Tetralopha asperatella.
(11) Pine—Body yellowish brown with darker stripes
(In silken tubes in globular masses of frass and
il Neo) seeps Sater ease sete Tee Tetralopha robustella.
(12) Poplar—
a. Head black; body dark brown, with yellowish lines
(nest of leaves drawn together and lined with

SKG) ay eet eee 2 ET Ichthyura inclusa.
b. Body brown and yellow striped; spiracles and legs
loksekepe = eat ee Tetralopha spp.

(183) Mimosa, honey locust—Body pale green to dark
brown with 5 longitudinal white lines. About 12
TMM elOne cat ee Homadula anisocentra

iypicalscaterpiilars: (not sluelike) 03 See.

Densely hairy larvae

(1) Oak, elm, hackberry, maple, sycamore, ete.—Densely
clothed with long, yellowish and reddish brown or
mouse gray hairs, with those at the rear end form-
We eaSOrtor tail 8... Megalopyge opercularis.

(2) Oak and many other hardwoods—Body almost oval
and covered with long silky hairs, with those on
the back raised to a ridge and sloping off rooflike
Onreachsiders. ss Pe ek Lagoa crispata.

Larvae smooth or armed with spines

(1) Oak, apple, dogwood, and many other hardwoods—
Body brownish in color, with a green patch on
middle of the back resembling a saddlecloth; in
middle of patch an oval purplish-brown saddle-like
spot; sides armed with fascicles of spines, and a
pair of spiny tubercles at each end

Sibine stimulea.

(2) Norway and sycamore maples plus a wide variety of
other hardwoods—Body marked with yellow, blue,
green, and purple; upper surface marked with a
dumbell-shaped purple area; long, shiny tubercles

13
14

2719

arise from each end and shorter ones along the
Sidesics:)...5e). qe ee tere Cnidocampa flavescens.
(3) Various hardwoods and shrubs—Body with 9 pairs of
lateral brown processes, the 3rd, 5th, and 7th pairs
curved, twisted, and longest. These processes
clothed with stinging hairs
Phobetron pithecium.
14. Typical caterpillars
Clothed with ‘harrs7or“bristles/2) 5 eee eee
Row of branched spines across top of each body
SEOMENE iLA.SIOROe Le Oe ee eee
Body naked or sparsely hairy; horny or wart-like
tubercles or spines on some segments 00...
Body naked; a prominent horn or eyelike tubercle on
Sthiabdominalseoment.) .<cen So) nee 2 eee
Body naked; one or more prominent dorsal humps on
eachsabdominalysecment 2) see eee
Body naked; anal projections ending in stemapoda
or modified anal legs, or head armed with antlers
Body naked (loopers or measuring worms) ...................
Body without prominent tubercles or filaments on
dorsumiss5-pairs-abdominal, lees) 225 eb2 2a) ae
15. Borers
Ins budsStemss Shoots: twits: sce eee ee
Inj fruit; nuts; cones Seeds) (hon -laiew ai eens eee ee
Insbark wood, roots 2. = eb 5 ee eee ee
16. Hairy, gregarious, raise front and rear ends when
disturb@d) sess: ostielen dart oe oe ea eater eet ee
Hairy or bristly; solitary or gregarious; body prominently
marked but with hair pencils, brush-like tufts, or
TUSSOCKSic:. 24.2 =. Shatin ieee Rares Tete Rican odice . Aiee Does
Hairy or bristly; with long hair pencils, brushlike tufts, or
GUSSOCK Sy ain) sete ie Letra eas eee eee ole
17. (1) Apple, birch, basswood, elm, oak, ete—Head black;
neck with yellow ring; cervical shield waxy yellow;
body black with narrow, yellow stripe
Datana ministra.
(2) Oak, beech, gray birch, hickory, butternut—Similar to
ministra except for an entirely black cervical
STS hag ie ates fey ius aks ie ay one Datana angusi.
(3) Oak and sycamore—Head and neck black; cervical
shield waxy orange-yellow; body black, with white
or yellow stripes; clothed with longish hairs
Datana contracta.
(4) Walnut, butternut, hickory—Head and body black;
clothed with long, dirty white hairs
Datana integerrima.
(5) Apple, azalea—Head, cervical shield, legs mahogany
red; body black with a broken white line
Datana major.
(6) Sumac—Head dark red or black; shield reddish-
brown; anal plate blackish; body yellow with
reddish to blackish stripe ...........Datana perspicua.

280

iw)
(o6)

18. (1)

iD:

(2)

(3)

(4)

(5)

(1)

(2)

(3)

(4)

(5)

(6)

(7)

Oak, birch, poplar, ete—Head yellow and_ black;
body dusky with 5 pairs of blue spots followed by
6 pairs of red spots on the back
Porthetria dispar.
Poplar and willow—Head black with bluish tinge;
body blackish, with a row of white blotches on top
and a narrow, broken, subdorsal line; sides mottled
with black and «whites. Stilpnotia salicts.
Poplar, sugar maple, tupelo, ete.—Head and body pale
bluish; a row of keyhole-shaped white spots on
the “dorsum: Gs. ee eee Malacosoma disstria.
Persimmon, palmetto, ground oak—Head orange
brown; front of prothorax, legs, anal segment, and
undersurface of body yellowish green to orange;
tops of other body segments crossed by 2 yellowish
stripes and a row of orange-colored warts
Seirarctia echo.
Ash, basswood, elm, maple, oak—Body flattened,
gray, and with faint longitudinal lines; lateral lap-
pets with long hairs, forming a fringe around the
body; a pair of warts bordered by a black band on
eMC OT Ax eer ei he aa lia ee Tolype velleda.
Elm, maple, poplar, sycamore, willow, ete.—Head red;
a pair of long, black hair pencils on prothorax and
another pair on the 8th abdominal segment; red-
dish dots on top of 6th and 7th abdominal seg-
ments; tufts whitish
Hemerocampa leucostigma.
Various hardwoods—Head black; body dark gray;
tufts whitish; a pair of black hair pencils on top
of prothorax and one on each side of the 2nd
abdominall.seg@ment =... 8. Orgyia antiqua.

Sycamore—Head yellowish brown; body yellowish,
clothed in whitish to yellowish hairs; long orange-
Colored: -nairpencils |. 2 Halisidota harrisit.

Various hardwoods—Head black; body blackish,
clothed in gray to yellowish hairs with an olive
tinge; hair pencils brownish to black with some
MEIC pe ee eee egies Halisidota tessellaris.

Butternut, hickory, walnut—Head black; body grayish-
white with white markings, clothed in grayish-
white hairs; a row of black tufts on the dorsum,
and 2 long black hair pencils on 2nd thru 6th
apaominal SE@MENtS sxxecceccks. Halisidota caryae.

Apple, birch, maple, oak, aspen, ete.—Head black;
body blackish; row of mostly black tufts on dor-
sum; yellow tufts on sides of abdominal segments
2 to 6; others black; tufts on thoracic and 8th
abdominal segments longest and containing some
yellow or white hairs .................... Halisidota maculata.

Apple, maple, etc.—Head shiny black; body greenish
white above and blackish below; clothed with fine
yellowish hairs; 2 long black hair pencils on each

281

of 1st and 3rd abdominal segments and a single 1
onthe Sthis..065 Se mele ee Acronicta americana.
(8) Poplar and willow—Head, shield, and true legs black;
body dull, whitish, clothed in long, yellow hairs; a
single, long hair pencil on each of Ist, 3rd, 4th, 5th
and 8th abdominal segments

Acronicta lepusculina. |
(9) Poplar, birch, willow—Head greenish-white or marked |
with black; body clothed with long, fine, curved, .

white or yellow hairs; a few bristly black hairs at

ends of the body ............ Acronicta leporina vulpina.

(10) Maple, cherry—Head rounded and yellowish; body
whitish or yellowish on the back, except for a row

of black spots; blackish beneathi; densely clothed

with long, fine white or yellow hairs; long hair

pencils on 2nd and 38rd thoracic segments and

8th abdominal segment ...... Apatelodes torrefacta.
20. Caterpillars with transverse rows of prominent, branched
Spines on the dorsum. |

(1) Elm—Head dark red; body brownish, mottled with
yellow; spines light colored, some tipped with
blacket.ck catno. cee 8 Polygonia interrogationis.

(2) Bireh—Head black; body reddish to blackish on top,
dotted with green; spines above spiracles black

Nymphalis j-album.

(8) Elm, poplar, willow—Head black, bilobed; body black,
sprinkled with whitish dots; black dorsal line
broken by row of 7 or 8 reddish dots; spines black;
abdominal legs reddish ............... Nymphalis antiopa.

(4) Various hardwoods—Head pea green and glossy; body |
pea green with a broad, reddish spiracular stripe
below which is a white stripe and another reddish |
line; spiracles yellowish ringed with black

~ Automeris 10.

(5) Oak—Head reddish brown; body dull brown to black-
ish, covered with small, pale yellowish dots; spir-
acles pale; clothed with rusty, brown-tipped bris-
tles and black or white, branched compound spines

Hemileuca mara.

21. Thorax with long, prominent SPINGS: «eee ace 22
Thorax with short, blunt spines or oy enlarged tu-

bercles 2 Prone cle) SME aS Geena re

22. Thorax with 1 pair of spines | De eS de Se SO Sie Lat Mart 23

Thorax ‘with’ 2-or more pairs Of Splines: sees. ee 24

23. (1) Head bilobed and covered with small, sharp tu-
bercles; a pair of large, barbed, club-shaped spines
On Zndsthoraciessesment eee Limenitis spp.
(2) A pair of slender, stiff spines, about as long as body
is thick, on 2nd thoracic segment; other spines
Short: ehiiuy. ise. Ree te he eee Anisota spp.
24. (1) Elm, poplar, cherry, willow, etc—Head and hump on
back of lst abdominal segment red, body black,
with black and yellow lines; a double row of short,

282

(2)

(3)

25. (1)

(2)

(3)

(4)

(5)

black spines on the dorsum, with the largest pair
on the 1st abdominal segment ... Schizura concinna.
Various hosts—Head orange yellow with the sides
green; body greenish, sometimes tinged with red
or brown; clothed with long whitish hairs; 2 pairs
thorny tubercles on top of 2nd and 8rd thoracic
segments long and backward curving; long, caudal
HORN CRE cer ON bis ne Taree es Eacles imperialis.
Hickory, butternut, walnut, persimmon, etc.—Body, 4 to
5 inches long and about 34-inch thick; smooth,
green or bluish, sometimes reddish brown; trans-
verse row of spines on each segment, some of
which, especially those on the thorax and the
caudal horn, are long and spiny; 2 large, black
patches between horns of 2nd and 8rd thoracic
SESIMEMUS oer tes ee es Citheronia regalis.
Ash, cherry, sassafras, yellow poplar—Head small
and yellow; body bluish or greenish white; 4 large,
coral-red tubercles on tops of 2nd and 3rd thor-
acic segments and a large, yellow one on top of
the 8th abdominal segment; other tubercles rudi-
Mentalbys ee oe Callosamia promethea.
Apple, ash, birch, maple, cherry, willow, ete.—Head
green with 2 black spots on each side; body pea
green with bluish tinge; 4 tubercular knobs on
thorax; all other turbercles somewhat rounded,
those on dorsum red and yellow, the others bluish,
and all bearing short, black spines
Hyalophora cecropia.
Ailanthus, wild cherry, plum—Head yellowish green;
prothoracic segment, anal plate, legs and top of
9th abdominal segment lemon yellow; rest of body
bluish green to yellow and dotted with black;
3 dorsal and 2 lateral long, bluish tubercles with
short-sradiatine bristles’ = oye Samia cynthia.
Hickory, walnut, beech, birch, persimmon, sweetgum,
willow—Head bluish green shaded with brown
above and at sides; body green to yellowish green,
each segment with 6 pinkish or greenish tubercles
bearing yellowish bristles; anal plate amber col-
ored, true legs brown and spiracles reddish brown;
2 pairs tubercles on thorax and one on 8th ab-
dominal segment larger than others
Actias luna.
Basswood, beech, birch, elm, hickory, maple, oak, etc.—
Head reddish brown, front of prothoracic segment
yellow, body apple green; body segments convex
to angular on back, each segment with 6 orange or
golden tubercles, each bearing 3 straw-colored
bristles; anal shield edged with purple
Antheraea polyphemus.

26. Poplar, willow—Glossy, leaden or tinged with purple;

head rounded and bilobed; anal plate coarsely
granular and rust red; 8th abdominal segment

283

swollen dorsally and bearing a well-developed horn
Pheosia rimosa.
Not glossy, but smooth and granulated ............. Sphingidae.
Kighth abdominal segment bearing a well developed horn
(6) Walnut, butternut, pecan—Head truncate in front,
apex with 2 rough brownish projections and a yel-
lowish stripe on each side; body light green to
reddish with yellowish stripes, sometimes bordered

above with reddish; caudal horn brownish
Cressonia jublandis.

Larvae without 7 oblique stripes on each side

(1) Catalpa—Head and caudal horn black; body either
pale yellow, with black markings and patches on
the back or black on top and pale yellow with
black spots on the sides and a stripe of black near
thesbase,Orscnewlecse se ees Ceratomia catalpae.
(2) Viburnum—Head bluish green, rounded; body yellow-
ish or sometimes purplish green; green dorsal
stripe bordered with whitish; horn bluish with
blacks oranulatiOnS. sesso) on eeeees Hemaris thysbe.

. Larvae with 7 oblique stripes on each side of the body

(1) Apple, black cherry, plum—head with reddish band on
each side; body yellowish green with broad, car-
mine stripes; horn and true feet dark reddish to
laches yee. tees ees ee ee Sphinx drupiferarum.

(2) Ash and lilac—Head bluish with broad pale band on
each side; body bluish green with light yellow
stripes; caudal horn reddish and curved down-
Walker tate ably ee. cae Ceratomia undulosa.

(3) Ash and lilac—Head bluish with pale band on each
side; body light green with light yellowish stripes
edged above with bluish-green; caudal horn blue

Sphinx chersis.

(4) Ash, lilac, laurel, rhododendron, etec—Head with black
band; body yellowish-green, with black stripes
edged below with yellow; anal plate dotted with
black; caudal horn blue marked with black

Sphinx kalmiae.

(5) Elm, basswood. birch—Body pale green to reddish
brown, with whitish, granulated stripes; pair of
tuberculated horns on each of 2nd and 3rd thoracic
segments; strong caudal horn on &th segment

Ceratomia amyntor.

(4) Wild cherry ete.—Head dotted with black; body pale

with chain-like black markings on the sides
Cingilia catenaria.

(5) Wild cherry, basswood, oak—Head pale green, rather
flat, sometimes tinged with light brown; body
smooth, greenish, with a broken, purplish brown
line on top, a broad stripe on each side, and a yel-
lowish, stigmatal stripe Melanolophia canadaria.

(6) Ash, hickory, walnut, basswood, beech, etc.—Head
large and usually bright rusty; body a dull, slate

284

27

Eee, EEE

black, sometimes uniformly green or yellow, or
mottled brown, tan, or rose
Ennomos subsignarius.
(7) Hemlock, balsam fir, arborvitae, spruce, larch, etc.— Y el-
lowish green to grayish or even darker; head and
body flecked with black .......... Lambdina fiscellaria.
(8) Hemlock—Head yellowish, with brownish or black-
ish spots; body yellowish with the top lighter than
the sides; sides marked with wavy, dark reddish
brown lines, interrupted with dashes of white
Lambdina athasaria athasaria.
(9) Pitch, red, and shortleaf pines—Straw to greenish yel-
low in color; head freckled; body with blackish
dots and wavy lines
Lambdina athasaria pellucidaria.
(10) Oak, red maple—Head flat, quadrangular, mottled
with black; body flesh colored with many wavy
blackish lines; thoracic segments thick; hairy tu-
bercles on all body segments ............. Phigalia titea.
(11) Basswood, elm, hickory, maple, oak, ete.—Head rusty
brown; body bright yellow with 10 wavy black
lines on top; legs yellow ................... Eranus tilaria.
(12) Oak—Head angular, bilobed; first 2 thoracic seg-
ments mottled with reddish brown and black;
body stout and ‘‘armed” with brownish tubercles
Phaeoura quernaria.
(18) Sassafras, tulip poplar—Head small and _ reddish
brown; body stout, yellowish to dark brown with
many fine, irregular, wavy lines
Epimecis virginaria.
(14) Pine, fir, spruce, hemlock, larch—Head and legs
flecked with brown; body pale yellow, marked
with a wide, orange-red dorsal stripe and thin,
longitudinal, black and white lines
Nepytia semiclusaria.
(15) Hemlock, spruce, fir, larch—Head pale whitish or
reddish brown, with 5 or 6 large and a few small,
| black dots; body whitish tinged with yellow or
| red, marked with black dots, yellowish lateral
stripe and subdorsal rows of white spots
Nepytia canosaria.

Measuring worms more than 50 mm. long.

(1) Hickory, basswood, ash, maple, etc.—Yellowish green,
tinged with red; head bilobed; body smooth with
swollen areas on top of abdominal segments 2 to
5 and conical tubercles on top of the 8th

Deuteronomos magnarius.

(2) Willow, poplar, wild cherry, etc.—Head deeply cleft
and flat in front; body greenish to reddish brown;
anal claspers large ou... Biston cognataria.

28. Body naked; 1 or more dorsal humps on abdomen
(1) Elm—Head pale green with 4 broad white bands in
front and on sides; body polished, bluish green;

285

thoracic segments unarmed; first 8 abdominal seg-
ments each have a large forward projecting promi-
nence ending in a bifid ridge; a pair of small, dor-
sal tubercles on 9th abdominal segment
Nerice bidentata.
(2) Oak, beech, elm, maple—Head orange red; body
smooth, naked, shining and increasing in diameter
to orange-red enlargement on 8th abdominal seg-
ment; a yellow, subdorsal and a yellow, stigmatal
stripe and several fine blackish lines
Symmerista spp.
(3) Oak and beech—Head bilobed and pale, with dark
brownish, branched band on each side outlining
whitish patches on the face; large, slightly cleft
dorsal tubercle on 1st abdominal segment; smaller
brownish tubercle on 8th abdominal segment; body
color pea green with shades of brown
Dicentria lignicolor.
(4) Poplar and willow—Head oval, flattened in front, and
vertex slightly lobed; body pearl gray, somewhat
marbled with brown and marked with a reddish
brown, dorsal line from head to 2nd tubercle;
conical tubercles directed posteriorly on 2nd and
3rd segments; prominent hump on 8th abdominal
Seoment 2h) i et. hee Hyperaeschra stragula.
29. Body naked, with two anal projections
(1) Poplar, willow, wild cherry—Prothorax with lateral
tubercles wider than the head; body green and
brown, tinged reddish or purplish, and tapering
gradually to the rear; anal legs filamental and
BVOVOD NY Wes TOONS MOVIL” soars sa msccresrserncnes Cerura spp.
(2) Beech, maple, poplar, oak, etc. —Head conical, pinkish
on sides; thickest in middle, a yellowish dorsal
stripe; filamentous and legs not quite as long as
body aismthichksiic.. a eee Fentonia marthesia.
(8) Hackberry—Head broad and armed with branched
antlers; body greenish often marked with yellow
stripes and spots, widest in middle and tapering
toward each end. Two anal projections
Asterocampa spp.
30. Measuring worms—not over 1 inch long
(1) Birch (gray and paper)—Body green to yellowish
brown, with fine yellow ten lines and a broad sub-
spiracular SUPIPC Wels ee. | Brephos infans.
(2) Elm, apple, etc.—Light green to dark brownish green ;
darker dorsal stripe; rudimentary prolegs on 5th
abdominal segment .................. Alsophila pometaria.
(3) Elm, apple, etc.—Reddish or yellowish brown _ to
greenish; body lines numerous and _ irregular,
sometimes almost invisible; prolegs on 6th and
anal segments only ...Paleacrita vernata.
(4) Hemlock—Head broader than thorax, brown, and bi-
lobed; body greenish tinged with redish brown;

286

ol.

swelling usually on each side of 2nd abdominal seg-
TMETG ees wae Fy ese ta Anacamptodes ephyraria.

(5) Hemlock, fir, larch, spruce—Head pale whitish, some-
times reddish brown, with 5 or 6 large and some
small dots; body whitish with yellow or reddish
tinge; dotted with black and with 4 dark, wavy,
hair lines below a yellowish, lateral stripe

Neptyia canosaria.

(6) Sugar maple, poplar, beech, etc.—Body light green
with 3 narrow yellowish-white stripes on each side
of the body; 2 pairs of prolegs

Operophtera bruceata.

(7) Red maple, ete.—Head pale green, slightly bilobed;
body green, skin much wrinkled, a double, whitish,
dorsal line bordered by yellowish white lines;
Spaces between segments yellowish
lobed; body greenish tinged with reddish brown;
swelling usually on each side of 2nd abdominal seg-

Physostegania pustularia.

(8) White pine—Head slightly brownish; body deep green
with greenish-white line on the dorsum and a nar-
row subdorsal and stigmatal white stripe

Eufidonia notataria.

(9) White pine, spruce, fir, larch—Head brownish; body
light green, sometimes brownish above; 2 longi-
tudinal, light stripes on dorsum between which
are 2 lighter stripes. ................. Semiothisa granitata.

(10) Black locust—Body green with many obscure, wavy
lines giving a reddish tinge

Semiothisa ocellinata.

(11) Black and choke cherry—Head and thoracic shield
dark amber; body blackish above, with 4 longi-
tudinal lines, and straw yellow beneath; about 20
MOM Goi yt ee Calocalpe undulata.

Measuring worms—25 to 50 mm. long

(1) Alder, beech, yellow birch—Head rather small and
rather flattened in front; body green to brownish
with whitish spots on dorsum; spiracles black

Hyperetis amicaria.

(2) Birch, maple, oak, walnut, etc—Head brownish, some-
what mottled; body reddish to chocolate brown,
with pair of blunt tubercles on top of 8th abdomi-
Male seoment 5 a ee Ectropis crepuscularia.

(3) Yellow birch, maple—Head bilobed; body dull brown,
with lighter and darker blotches; prominent swol-
len area on top of 6th abdominal segment

Plagodis serinaria.

Naked, without prominent tubercles, and with 5 pairs

abdominal legs.

(1) Beech, sugar maple, etc.—Head large with broad red-
ish, lateral band; body yellow green, or light green
with bluish cast; usually a reddish-brown or pur-
plish saddle-shaped patch on the back

Heterocampa guttivitta.

287

(2) Beech, oak, basswood, elm, etc.—Head large, blackish
with whitish band on each side; body smooth yel-
lowish green with a pale median line on each side

Heterocampa manteo.

(3) Hemlock, larch, spruce—Head yellowish; body light
green; pale median stripe, and a subdorsal and
stigmatal stripe on each side; the stigmatal stripe
bordered int red 3. Feralia jocosa.

(4) Maple, ash, ete-—Pale green; head with yellowish
tint, body with broad stripe on top; skin dotted
with white; a narrow subdorsal and broad stig-
matal stripe on each side .... Lithophane antennata.

(5) Oaks mostly—Pale pea green; head large and round-
ed; body with yellowish, subdorsal stripes; spir-
ACIES: “PEG Pi Aw: een Serene | ..Nadata gibbosa.

(6) White, pitch, and jack pines—Head triangular, coni-
cal, with a pale, yellow stripe on each side; body
grass green, with 3 longitudinal white stripes on
each side; broad, brick-red, median dorsal stripe
on dorsum of abdomen ........... Lapara bombycoides.

(7) Spruce—Head somewhat mottled; body dull brown
with a dark dorsal stripe; spiracles and tubercles
black pet: ess lo ee ae: Epizeuxis aemula.

(8) Willow, elm, maple, poplar, ete—Body wine-colored,
with four large whitish spots on the front of the
prothorax and marked with many irregular, wavy,
longitudinal lines and creamy-white spots

Lycia ursaria.
32. Borers in buds, stems, shoots, twigs.

(1) Pines (red, Scotch, Austrian, etc..—Head and thoracic
shield black; body brownish; almost 12 mm. long.
In buds and expanding shoots

Rhyacionia buoliana.

(2) Pines (Pitch and other hard pines)—Head and cervical
shield blackish; body yellowish to pale brown;
about 9 mm. long. In buds and succulent growth
at tips of twigs ... Rhyacionia frustrana frustrana.

(3) Pines (Red, Scotch, jack)—Head dark brown to black;
prothoracic shield black; anal plate dark brown;
remainder of body yellowish-brown to reddish-
brown; about 8 mm. long. In buds and new shoots

Rhyacionia adana.

(4) Pines (Red, Scotch, Austrian, longleaf, etc.)—Head, cer-
vical shield, anal plate brown to dark brown; body
yellowish; about 6 mm. long. In buds

Battaristis vittella.

(5) Pines White, red, Scotch, jack, etc.).—Head yellowish-
brown, with a round, blackish spot at the hind
margin; most of remainder of the body a dirty

white: sinwlateralsn. +. oes. Eucosma gloriola.
(6) Jack pine—Larva brownish eray ‘and about 12 mm.
long. In new shoots 0.0.0.0... Hucosma sonomana.

(7) Pines (pitch, etc..—Head and cervical shield dark
brown; body bale brown; about 12 mm. long. In
CWIESE ie ee eet rene Petrova comstockiana.

288

(8) Pines (Jack, ete..—Head and cervical shield light
brown; body reddish with small, shiny tubercles.
invvyounebranches, 2.0.26. Petrova albicapitana.

(9) Pines, fir, spruce—Head dark reddish brown with dark
brown maculations; body moderately slender,
somewhat fusiform, the dorsum usually reddish
purple, the venter pale reddish purple, tinged
with green; about 20 mm. long. In _ shoots,
branches, and the trunk ........... Dioryctria abretella.

(10) Pines (Longleaf, loblolly)—Head shiny red-brown;

thoracic legs and thoracic and anal shields brown;

body greenish-white with brownish markings

above; about 25 mm. long. In terminals and trunk

OO TS tite Aes ete cney lea deod c3 Dioryctria amatella.

(11) Pines (red, Scotch, white, jack, shortleaf, loblolly, etc.)

—In the North: head brown, body pink to green-

ish and covered with small, black dots; about 19

mm. long. In the South: uniformly bluish-black

above, excepting the top of the thorax, and

greenish-blue with a violaceous tint, below: 18

COMZosmMmMs ONS, os... Dioryctria zimmermant.

(12) Pines (Scotch, red, jack, longleaf, loblolly, scrub)—IJn

the North: slender, olive-green to buff and from

14 to 18 mm. long. In the South: moderately slen-

der, grayish-buff, and 20 to 25 mm. long. In
staminate flowers and cones

Dioryctria disclusa.

(13) Spruces, firs, larch, and jack pine—Head and cervical
shield reddish-brown; body brown to amber and |
ornamented with piliferous warts; about 18 mm.
long. In needles, buds, staminate flowers, and
CONMCS ir, Bo eee els ies | Dioryctria reniculella.

(14) Black locust—Head brownish-black; body greenish
with broken stripes of dark brown; about 16 mm.
long. Girdling near ground line

Elasmopalpus lignosellus.

(15) Spruce—Head brownish-yellow; cervical shield
paler; body yellowish or grayish-green; about 18
mme lone. In buds Zeiraphera ratzeburgiana.

(16) Gray birch, black cherry, maple, oak, ete.—Head
slightly bilobed and rusty red; body greenish-
brown; pair of prominent tubercles on lst ab-
dominal segment; a pair of long, fleshy brownish
filaments on each of 2nd and 3rd segments; broad
white stripe on back from prothorax to lst pair
of filaments; about 18 mm. long

Nematocampa limbata.
33. Borers in fruit, nuts, seeds, and cones.

(1) Oaks, beech, hazel—Full-grown larvae pink or whit-
ish; 18 mm. long. In acorns, hazelnuts, and beech-
NULLES Hea eek clea enue aly: no er: Melissopus latiferreanus.

(2) Oak, hickory—Body grayish white or yellowish with
black spots on top; cervical and anal shields brown-
ish. In acorns and hickory nuts

Valentinia glandulella.

289

(3) Hickory and pecan—Head light brown; body creamy
white and about 9 mm. long. In hickory nuts and
PECANS 812A eee ee ae eee Laspeyresia caryand.

(4) Pines (Red, Scotch, Austrian, longleaf, etc.)—See 32 (4)
for description. In cones and buds

Battaristis vittella.

(5) Pines (Various species, particularly Southern pines)—See

32(11) for description. In seeds and cones
Dioryctria zimmermant.

(6) Pines and spruce—See 32(9) for description. In cones
ANG: SCCUSi Ail is ee Dioryctria abietella.

(7) Pines (Scotch, red, jack, longleaf, loblolly, scrub)—See
32(12) for description. In cones and flowers

Dioryctria disclusa

(8) Pines (longleaf, loblolly)—See 32(10) for description.

I :CONCSH Ae ee ee Cee Dioryctria amatella.
(9) Spruce, fir, ete—See 32(138) for description. In flow-
ers. cand conest i. 225 a eee Dioryctria reniculella.

(10) Red pine—Head and pronotum brown; remainder
of body grayish-white; about 12 mm. long. In

CONICS). SA ee et Eucosma monitorana.

(11) Pines (Slash, loblolly, longleaf, shortleaf)—Head red-
dish-brown or nearly black, with dark markings;
thoracic shield brownish; anal shield brownish-
yellow; about 18 mm. long. In conelets and cones
Dioryctria clariorellia.

34. Borers in branches, trunk, or roots.

290

(1) Ash, lilac, privet-—Whitish or yellowish. In main stem
Podosesia syringae syringae.

(2) Flowering dogwood—Whitish with brown heads. Un-
der the bark? =2 0) oe Synanthedon scitula

(3) Elm, locust,oaks, green ash, ete.—Greenish-white; head
shiny brown, with powerful, nearly black mandi-
bles; thoracic legs yellowish, with pointed, curved
tarsal claw; 50 to 75 mm. long

Prionoxystus robinae.

(4) Elm, maple, ete—Head and thoracic and anal plates
brownish-black; body pale yellow, frequently with
a pinkish tinge, sparsely hairy and spotted with
brown or black tubercles; about 50 mm. long. In
twigs, limbs, and trunk. Twigs and branches fre-
CUE MIE yaa aC Clee ater te Zeuzera pyrina.
(5) Hickory, oak, pecan—Head, cervical shield, and anal
plate shiny dark-brown; body pinkish, and sparsely
clothed in fine hairs; about 37 mm. long. In the
pith of twigs at first; later in large galleries sev-

eral inches long in larger limbs and the trunk
Cossula magnifica.
(6) Linden (European)—Head brown; body whitish with
contents of alimentary tract visible through the
integument. About 6 mm. long. In the bark of
limbs and trunk .. Chrysoclista lineella.
(7) Maples—Head brownish; body white; about 12 mm.

long. In bark and sapwood. usually around wounds
Sylvora acerni.
(8) Maples (red and silver)—In small branches causing
eallelikerswellings 4455 ce Carmento corni.
(9) Persimmon—In solid wood of main stem or taproot,

sometimes to a depth of 18 inches in the ground
Sannia uroceriformis.
(10) Poplar and willow——Head brownish; body white and
about 12 mm. long. In roots, trunks, or large

limbs. Extensive burrows and swellings

Aegeria apiformis.
(11) Rhododendron, laurel, azalea—Yellow-white and
about 12 mm. long. Under the bark of twigs,
branches, and stems ...Synanthedon rhododendri.

(12) Pines (various species)—See 32(11) for description.
In branches and stems .....Dioryctria zimmermani.
(13) Pines (White, pitch) and spruce—White to pink and
about 62 mm. long when full-grown. In the inner
bark or sapwood. Pitch masses 75 to 100 mm. in
diameter at entrance hole............... Vespamima pint.

FAMILY PAPILIONIDAE
SWALLOWTAIL BUTTERFLIES

Swallowtail butterflies are of considerable interest to many
people because of their large size and striking appearance; other-
wise, they are of minor importance. The adults are distinguished
by the wavy margins and tail-like prolongations of the hindwings;
the larvae, by the protrusive, bright-colored forked processes
rising from the first thoracic segment. They also emit a disagree-
able odor when disturbed.

The tiger swallowtail, Papilio glaucus L., occurs commonly in
eastern North America. The caterpillars feed on various decidu-
ous 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 which is also edged with black. The posterior part of
the first abdominal segment bears a transverse, yellowish ridge,
edged posteriorally 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. Trans-
formation to the chrysalis usually takes place on some object
above the ground.

The spice-bush swallowtail, Papilio 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 in diameter 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 segments each bear
two orange spots. Those on the thorax have black centers. Six

291

small spots are on abdominal segments two to seven and four on
the eighth. P. cresphontes Cramer feeds on prickly ash in the
Northern States.

Graphium marcellus Cramer, the zebra swallowtail, feeds on
pawpaw. /

FAMILY NYMPHALIDAE
BRUSH-FOOTED 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-sign feeds on elm,
oak, and hackberry in eastern America, especially on sprout
growth along roadsides. Full-grown caterpillars (fig. 104) 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 (Harr.),
the comma butterfly, occurs on elm, nettle, and hop from Canada
to the Carolinas and Texas. Full-grown caterpillars are yellowish
white, and each body segment bears a transverse row of branched
spines.

Nymphalis j-album Bdv. and LeC. 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

F-519524

FicurRE 104.—Larvae and chrysalis of the butterfly,
Polygonia interrogationis, the question-sign.

292

dots of light green, and each segment bears a transverse row of
branched spines.

The mourning-cloak butterfly, Nymphalis antiopa (L.), is a
widespread species, occuring throughout the subarctic regions of
North America. The larvae, commonly known as spiny-elm cater-
pillars, feed on elm, willow, poplar, and hackberry, and are some-
times abundant locally, especially on shade and ornamental trees
and along fence rows. Adults are black-bodied and have wing-
spreads of 60 to 80 mm. The upper wing surface is dark red-
dish-brown except for a broad, creamy-yellow border, which
contains a row of blue spots. Full-grown caterpillars (fig. 105)
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 CONN. AGR. EXPT. STA.
FIGURE 105.—Larvae of the

mourning-cloak butterfly.

Nymphalis antiopa.

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. Then they pupate and the adults emerge to hi-
bernate. There are one or two generations per year, depending on
location. —

The mourning-cloak butterfly is of minor importance in the
forest, but is sometimes injurious to shade and ornamental trees.
Infestations can be controlled by cutting and burning infested
twigs and small branches.

The viceroy, Limenitis (=Basilarchia) archippus (Cramer),
occurs over most of the United States and feeds on poplar, willow,
and plum. The adult resembles the well-known monarch butterfly,
Danaus plexippus (L.), but differs by being slightly smaller, hav-
ing a narrow black line across the hindwings and only a single
row of black 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

293

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 others. There are two generations per year. The
two related species, L. arthemis (Drury) and L. astyanaz (F.),
are found on poplar, willow, birch, black cherry, apple, and bass-
wood. The caterpillars are similar in appearance to those of
L. archippus.

Asterocampa clyton (Bdv. & LeC.) occasionally seriously de-
foliates hackberry in the Lake States. Full-grown caterpillars are
25 to 37 mm. long. The body is greenish except for a yellow stripe
down the back and a deep blue, yellow-bordered stripe on each
side. The head is armed with branched, antler-like spines, and
there are two projections at the posterior end of the body. A.
celtis (Bdv. & LeC.) is also abundant occasionally 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.
Otherwise, it resembles the caterpillar of A. clyton. Langlois &
Langlois (449) discussed its life history.

FAMILY HESPERIIDAE
SKIPPERS

Members of this family are commonly known as skippers be-
cause of the way they flit or dart from place to place. They are
distinguished by the head which is nearly as wide or wider than
the thorax, and the antennal club which usually has two pairs of
spurs. The larvae usually have large heads and strongly con-
stricted necks. They are also usually solitary, each one concealing
itself under part of a leaf which it cuts and folds over.

The silver-spotted skipper, HE pargyreus 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, ground nut, and wisteria. Adults are brown except
for yellow and white triangular spots on the forewings. The fore-
wings 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 five
black rings.

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 or one and a partial
second.

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. They have stout, spindle-shaped bodies and

294

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 usu-
ally 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 either 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 abdominal
legs, holds the front part of the body aloft, and bends its head
downward.

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 (Hbn.), feeds on basswood,
birch, and elm throughout 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 abdominal segments; and have one
caudal horn. They are about 75 mm. long. C. wndulosa (Wlkr.)
feeds on white ash and lilac from eastern Canada and Maine to
the Carolinas and westward to the Mississippi River Valley. Full-
grown larvae are about 62 mm. long. The head is bluish-green
marked 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-colored; and the caudal horn is
reddish and curved towards the tip.

The catalpa sphinx, Ceratomia catalpae (Bdv.), occurs from
New Jersey 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 de-
foliating them. The adult is heavy-bodied and has a wingspan of
about 75 mm. The front wings and body are gray 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.
106). There are two color forms of large larvae—dark and light
(385). The dark form is black on top and pale yellow underneath ;

COURTESY OF ILL. NAT. HIST. SURV.
FIGURE 106.—Larvae of the catalpa sphinx, Ceratomia catalpae.

295

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 gregar-
iously; older ones, singly. In the Deep South, all life history
stages may be present during the summer, and there may be three
or four generations per year. In the North there is only one gen-
eration per year. Larvae are often heavily parasitized by the
hymenopteron, Apanteles congregatus (Say) (fig. 107).

F-519572

FIGURE 107.—Larvae of the

ecatalpa sphinx bearing co-

ecoons of Apanteles congre-
gatus.

The great ash sphinx, Sphinx chersis (Hbn.), feeds on lilac and
white ash throughout the United States. Full-grown larvae are
usually light green with 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 de-
foliation in such areas.

Sphinx kalmiae J. E. Smith feeds on white ash, fringe tree,
mountain laurel, rhododendron, and lilac from southern Canada
through the Atlantic States to Georgia. Full-grown larvae are
yellowish-green and about 75 mm. long. The body is marked by
seven oblique, black stripes edged below with yellow; the caudal
horn is arcuate and blue with black, raised markings. S. luscitiosa
Clem. feeds on poplar and willow in the Atlantic Coast States.

296

S. drupiferarum J. E. Smith, a widely distributed species, feeds
on apple, beach plum, wild cherry, hackberry, plum, and peach.

White, pitch, red, and jack pines are fed on by the larvae of
Lapara bombycoides Wlk. in eastern Canada and from the At-
lantic Coast to the Mississippi River Valley. The larvae are
present from July to September. Full-grown specimens are green
except for a broad, brick-red, median, dorsal stripe; 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 pine in the same general territory.

Aspen, willow, and sometimes apple and birch are fed on in the
Atlantic States by Paonias excaecatus (J. E. Smith). 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. P. myops (J. E. Smith) larvae feed on
black cherry in eastern United States. They have rose-colored
spiracles; otherwise, they closely resemble the larvae of P. ex-
caecatus. The larvae of Smerinthus jamaicensis (Drury) (=gem-
inatus Say), feed on aspen and willow. They also resemble the
larvae of P. excaecatus. They differ mainly in having a bluish-
purple caudal horn, violet thoracic legs, and sometimes sub-dorsal
rows of reddish spots on each side of the body.

The walnut sphinx, Cressonia juglandis (J. EK. 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 hop hornbeam. 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 bordered above
with reddish, on each side of the body. The caudal horn is brown-
ish and very granulated. There are two generations per year in
the South.

FAMILY SATURNIIDAE
GIANT SILKWORM MOTHS

This family contains some of the largest and most colorful
moths in eastern United States. The larvae, also large and con-
spicuous, are more or less armed with tubercles and spines. Be-
cause 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 injuri-
ous. Some of the most common species are discussed briefly below.

The cynthia moth, Samia (=Philosamia) cynthia (Drury), was
introduced into this country from Asia about 100 years ago and
now occurs from Connecticut to Virginia. Its preferred host is

297

ailanthus, but wild cherry and plum are also infested. The adult
is 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 lemon-yellow
prothoracic segments. The rest of the body is light bluish-green
to yellowish, dotted with black, and there are long, bluish tu-
bercles with short bristles on each body segment. Moths are pres-
ent from June to September; larvae from July to October. Winter
is spent as pupae in cocoons on the tree or on the ground.

The cecropia moth, Hyalophora (=sami) cecropia L., occurs
throughout eastern United States and in southern Canada. The
larvae feed on a wide variety of hardwoods such as ash, birch,
cherry, hawthorn, walnut, maple, sassafras, pussy willow, elm,
poplar, and basswood. Feeding damage is often rather severe in
shelterbelt plantings in the Northern Great Plains. The moth has
a wing expanse of 125 to 150 mm. There is a white, crescent-
shaped spot near the center of each wing, a red-bordered cross-
band on each wing, and a dark spot near the end of each forewing.
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
four, large, coral-red tubercles on the top of 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 tachinid, Lespesia ciliata (Mac.), is a common
parasite in the Northeast.

The promethea moth, Callosamia promethea (Drury), occurs
throughout most of eastern United States and southern Canada,
and feeds on a variety of hosts such as ash, wild cherry, lilac,
sassafras, spicebush, yellow-poplar, maple, and birch. The adult
has a wing expanse of about 75 mm. The wings of the female are
reddish-purple to brown with light brown borders, 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 is small and yellow; the body bluish or greenish white.
The second and third thoracic segments bear four large, coral-red
tubercles or horns; the eighth abdominal segment, bears a large
yellow one. Each segment is also ornamented by a deep blue wart
or button. Winter is spent in a tough, light, colored cocoon en-
closed 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 (=Tropaea) 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, ironwood, and butternut. The adult moth, a very beautiful
insect, has delicate green wings which expand up to 100 mm.
Each forewing bears a conspicuous eyelike spot, and is edged
with a purplish-brown band. The hindwings extend into long,
curved swallow-tails. 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

298

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 segments are red; the remainder, 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 genera-
tions per year in the South.

The polyphemus moth, Antheraea (=Telea) polyphemus
(Cram.), 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 brownish-yellow and has a wing expanse of 100 to 150
mm. There is an eyelike transparent spot on each wing and a
sooty, transverse stripe outwardly edged with light pink near the
outer margin of the wings. Full-grown larvae are apple green in
color and about 75 mm. long. The head is reddish-brown, and the
thoracic shield is sometimes margined along the front with
yellowish-green. 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 on the ground. Larvae may be found from June
to October. The tachinid, Lespesia frenchii (Williston), is a
common parasite in the northeast.

The io moth, Automeris io (F.), occurs throughout eastern
United States and attacks a wide variety of trees including paper
birch, wild cherry, black locust, aspen, willow, beech, apple,
maple, oak, hickory, elm, mulberry, dogwood, and sycamore. Fe-
male moths are purplish-red and have a wing expanse 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. Full-grown larvae are about
75 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 June to July; larvae from July to September.
Winter is spent in the pupal stage, in a tough, brown, oval cocoon
often covered with bits of dead leaves and other debris, usually
on the ground.

The buck moth, Hemileuca maia (Drury), occurs from New
Hampshire to Georgia, Louisiana, and Oklahoma, and it appar-
ently feeds almost exclusively on various species of oak. Adults
have black, thin-scaled, sometimes semi-transparent wings. A
common white band crosses both the forewing and hindwing near
the middle. 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 in the Northeast is the tachinid,

299

Leschenaultia fulvipes (Bigot). H. lucina Hy. Edw. feeds princi-
pally on Spiraea latifolia, but also on wild black cherry, gray
birch, and oak in the Northeast. H. nevadensis Stretch, the Ne-
vada buck moth, has been reported feeding on poplar and willow
in Nebraska and Oklahoma.

FAMILY CITHERONIIDAE

ROYAL MOTHS

Royal moths are medium-sized to large, with stout bodies and
large, strong wings. The head is generally sunken in the pro-
thorax, 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 thinly hairy. The horns or spines on the second and
sometimes third 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
hazelnut from southern Canada and New England to Georgia
and Kansas. The adult has a wing expanse of about 50 mm. Its
forewings resemble those of the orange-striped oakworm, but
differ in being darker, more speckled, and having the outer por-
tion often tinged with lilac. Full-grown larvae are about 387 to
50 mm. long. The body is tawny, often tinged with rose or pink.
It is covered with tiny ivory-white specks, denticles, or granules,
and is marked with single dorsal and lateral stripes. There are
two long, curved spines on the second thoracic segment. The
remaining segments bear backward-pointing spines.

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 outbreaks have been recorded, some of which
covered hundreds of acres.

The orange-striped oakworm, Avnisota senatoria (J. EK. Smith),
feeds on various oaks from eastern Canada to Georgia and west-
ward to Minnesota, Iowa, and Texas. The adult has a wingspread
of 37 to 62 mm., and its thick body is covered with yellowish-red
hairs. The forewings are orange-purple and are marked with an
oblique band, a white spot, and numerous black dots. The hind-
wings of the male are distinctly triangular and only about two-
thirds as long as the body. Full-grown catrpillars are black, with
eight longitudinal orange-yellow stripes on the dorsum and sides,
and the caterpillars are about 50 mm. long. 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 (fig. 108).

Adults appear during June or July and deposit their eggs in
clusters of several hundred eggs 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

300

tertile an amend meme aaiainiiieal

COURTESY CONN. AGR. EXPT. STA.
FIGURE 108.—Adult, eggs, and larva of the orange-striped
oakworm, Anisota senatoria.

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 3 to 4 inches. There is one and
possibly two generations per year, depending on location. During
recent years this species has been responsible for a considerable
amount of defoliation in oak stands in Connecticut, New Jersey,
and Pennsylvania.

The pink-striped oakworm, Anisota virginiensis (Drury), oc-
curs in southern Canada and throughout the eastern half of the
United States. The larvae feed principally on various oaks, but
also on many other hardwoods such as chestnut, hazel, maple,
and birch. There are two generations per year in the southern
parts of its range.Adults (fig. 109) are brownish-red, often with
a purplish cast. Females have a wingspread of about 37 mm.
Males are considerably smaller. The forewings are thinner, less
speckled, and more transparent beyond the discal dot than those
of the orange-striped oakworm or the spiny oakworm. Full-
grown larvae are about 50 mm. long. The body is greenish to
brownish-yellow with two dorsolateral, rose-colored stripes and
a similarly colored stripe along each side. Two spines on the
second thoracic segment are slightly curved and are conspicu-
ously longer than the others.

The green-striped mapleworm, Anisota rubicunda (F.), is found
throughout most of 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

501

COURTESY OF DUKE UNIV. oa FOREST.
FIGURE 109.—Adult of the pink-striped oakworm, Anisota virginiensis.

most likely to occur in the South. The moth (fig. 110A) hasva
woolly body and a wingspan 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 hind-
wings are either pure yellow, or yellow with rose-pink streaks.
Full-grown larvae ( fig. 110B), have cherry-red heads, pale yel-
low-green bodies and are about 87 mm. long. The body also has
seven dark green or nearly black 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.

There are one and two generations per year in the North and
South, respectively. Eggs are laid on the undersides of leaves

F-500801, 500799

FIGURE 110.—The green-striped mapleworm,
Anisota rubicunda: A, adult; B, larva.

302

during May and June and hatch in about 10 days. Larvae feed
singly, devouring entire leaves, and become full-grown in about
a month. In the South, where two generations per year occur,
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 green-striped mapleworm
have been recorded, but never in any great abundance. Birds
devour some of the larvae but are probably not very effective
in control (774).

The regal moth, Citheronia regalis (F.), occurs throughout the
Southern States, northward to Illinois 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, wal-
nut, butternut, persimmon, sweet gum, sycamore, ash, and sour-
wood. Adults are quite large, males having wingspreads of 100
to 125 mm., and females 125 to 150 mm. The head and body are
orange-colored with pale yellow markings; the forewings 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. 111).

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 species is
of little or no economic importance. It is of interest mostly be-
cause of the frightening appearance of the larvae.

Citheronia sepulchralis G. & R. is occasionaly 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 the wingspread is about 75 to 100 mm.
Full-grown larvae are dull brown, armed with short, orange -
colored horns, and about 100 mm. long.

The imperial moth, Hacles imperialis (Drury), occurs in east-
ern United States and southern Canada. The larvae feed on
the foliage of a wide variety of trees including pines, red cedar,
oaks, sweetgum, elm, persimmon, hickory, maple, beech, honey
locust, and cypress. The moth is sulfur yellow with lilac mark-

COURTESY CONN. AGR. EXPT. STA.

FIGURE 111.—The hickory

horned devil, larva of Cit-
heronia regalis.

303

Ce

ings 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, pale green to dark green or
reddish purple and covered with long, whitish hairs. Adults ap-
pear during June and July and. larvae are present from July to
October. Winter is spent in the pupal stage in the ground, and
there is one generation per year.

Eacles imperialis pini Michener feeds on 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. Markings on the surfaces of the wing |
are also heavier than those of the imperial moth. Adults are
present from mid-June to mid-July. |

FAMILY CTENUCHIDAE

Lymire edwardsu (Grote) has been recorded feeding as larvae
on the foliage of banyan and Fcus trees in Florida. The adult is
a Sluggish, bluish gray to purplish gray moth, with blue and .
plumose antennae. The thorax is orange-red beneath, the abdo-
men white beneath and blue above. Larvae are somewhat whitish
with a dark tuft of hairs on the thorax. When touched, they usu-
ally flip from the leaf to drop to the ground on silken threads.
This species is occasionally destructive of shade or ornamental
trees (283). |

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 roof-like 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 Lyman-
triidae. The hairs of certain species are irritating. The majority
of species prefer the foliage of low growing plants. A few feed
on the foliage of trees and shrubs.

The hickory tussock moth, Halisidota caryae (Harr.), occurs in
southern Canada and south in the Eastern States to North Caro-
lina. 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 aspen. Walnut, butternut, and hick-
ory appear to be preferred. Adults are light brown or buff in
color, with numerous silvery white spots on the forewings, and
they have wingspreads of about 50 mm. Full-grown larvae are
grayish-white and 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

304

long black hair pencils on the first and seventh abdominal seg-
ments (fig. 112).

Adults appear from late May to early July. The female deposits
her eggs in batches of 50 to 400 eggs each in a single layer on the
undersides of the 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, Halisidota maculata (Harr.), occurs
from coast to coast 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, wild black
cherry, maple, and willow. The oaks, willow, and poplar are par-
ticularly attractive. The adult is pale yellow, with long, somewhat
pointed, brown-spotted forewings and has a wingspread of 37 to
50 mm. Full-grown larvae are about 30 mm. long, dull black
above, 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 ab-
dominal segments are longest and bear an intermixture of white,
yellowish, and black hairs, with those cn the thorax overhanging
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 spe-
cies is occasionally abundant enough to damage shade trees.

The pale tussock moth, Halisidota tesselaris (J. E. Smith), oc-
curs in southern Canada and throughout the eastern part of the
United States. The larvae feed on practically all common, decidu-
ous trees and shrubs. The adult is pale yellow and has a wing-
spread of about 50 mm. The forewings are translucent and
crossed by five broad, darkish bands. Full-grown larvae are dark-
ish and densely clothed with compact tufts of light yellow or |
dirty white fine hairs. Black pencils rise in pairs from the second
and third thoracic and eighth abdominal segments (fig. 118).
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 gen-
eration per year. This species is frequently abundant in the for-
est and along roadsides in the South Central States, but is usually
of minor importance.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 112.—Larvae of the

hickory tussock moth,
Halisidota caryae.

COURTESY CONN. AGR. EXPT. STA.
FIGURE 113.—Larva of the pale
moth, Halisidota tessellaris.

The sycamore tussock moth, Halisidota harrisii (Walsh), feeds
on sycamore and London plane trees, and it probably occurs
wherever sycamore grows in this country. Adults are indis-
tinguishable from those of the pale tussock moth, and the larvae
of the two species differ only in color. Those of this species have
yellowish bodies clothed in whitish to yellow hairs and their long
hair pencils are orange-colored. Infestations are often heavy on
shade and ornamental sycamore in the Northeast.

The fall webworm, Hyphantria cunea (Drury), occurs through-
out 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 red. 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 is often 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. The larvae of the textor form are dark. Their
bodies are covered with long silky, gray hairs arising in tufts
from orange-yellow or black tubercles.

Adults appear mostly from May to July and deposit their eggs
in hair-covered masses of several hundred eggs each, usually on
the undersides of leaves. Newly-hatched larvae immediately begin
to spin a silken web over the foliage on which they feed; and,
as they grow, they enlarge the web to enclose more and more
foliage (fig. 114). On heavily infested trees several branches may
be enclosed in webs. Small trees are often enclosed entirely. The

306

F-519570

FIGURE 114.—Defoliation and

webbing caused by _ the

fall webworm, Hyphantria
cuned.

larvae are gregarious until the last instar. During the early in-
stars they feed on the upper surface of the leaves; later they
devour entire leaves excepting the larger veins and midribs. As
they approach maturity, some of them 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 or two
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 occur, however, sometimes en-
compassing tracts several miles in extent. It is often a serious
pest of shade trees and ornamentals. These trees may not only be
heavily or completely defoliated, but the presence of numerous, ~
unsightly webs detracts greatly from their esthetic value. Oliver!
discussed the ecology of the species in the Deep South, and (575)
its natural and biological control in Louisiana. Tothill (715) dis-
cussed its natural and biological control in Canada.

Seirarctica echo (J. E. Smith) larvae feed on the foliage of
persimmon, ground oak, and sabal palmettos 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-colored warts that cross
each segment.

Other species of arctiids likeiy to be encountered in eastern
forests include: Haploa clymene (Brown)—on maple, hickory,
and apple; H. lecontei (Guer.)—on maple, birch, oak, and cherry;
Apantesis radians (Wlk.)—on slash pine seedlings in Georgia;
and Lexis bicolor Grt—on balsam fir and spruce.

1! Oliver, A.D. 1963. An ecological study of the fall webworm, Hyphantria
cunea (Drury), in Louisiana. PhD Thesis, Louisiana state Univ.

307

FAMILY NOCTUIDAE

UNDERWING MOTHS

This is the largest of all the families of Lepidoptera in North
America. The adults are mostly nocturnal in habit. The majority
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. Many are
found on the foliage of forest and shade trees and shrubs, but
they usually do not cause serious damage. Other species attack
seedlings in nurseries and young trees in plantations and occa-
sionally cause serious injury. This family has received extensive
treatment by Smith and Dyar (658), Crumb (176), and Knutson
(433).

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 dagger-
like mark near the anal angle of the forewing. |

The American dagger moth, Acronicta americana (Harr.), oc-
curs throughout 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, white and
yellow birch, 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 winter is spent in the pupal
stage in dense silken cocoons. During an outbreak in Maine in
1945, larvae were so abundant that they literally swarmed all
over buildings and gardens.

The cottonwood dagger moth, Acronicta lepusculina Guen., oc-
curs from coast to coast in southern Canada and the Northern
States. Its favorite hosts appear to be trembling aspen and
willow, 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 Guen.—hickory and birch;
A. morula Grote & Rob.—elm and basswood; A. interrupta Guen.
elm, cherry, birch, and apple; A. lithospila Grote—hickory and
oak; A. funeralis Grote & Rob.—hickory, elm, and birch; A.
modica Wlk.—oak; A. retardata (Wlk.)—maple; A. leporina vul-
pina (Grote)—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 hebaceous vege-
tation but also occasionally poplar, willow, boxelder, wild cherry,
alder, and apple.

The genus Catocala contains many large, conspicuous species,
the larvae of which feed on the foliage of various forest trees

308

SS

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 when they are
resting with their wings folded on the trunks of trees. The hind-
wings, in contrast, are beautifully marked with bright red, yel-
low, 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. Barnes
and McDunnough (33) discussed the life histories of many
species.

Many other species of noctuids’ are commonly known as cut-
worms. The larvae are usually stout, naked, and dull grayish or
brownish in color. 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 (Guen.)—hemlock, spruce, larch; Epizeuxis sem-
ula (Hbn.)—spruce (particularly in ornamental plantings, the
larvae commonly being found in webbed masses of dried needles
and frass on the branches) ; Panthea furcilla (Pack.)—pines; P.
acronyctoides (Wlk.)—spruce; Papaipema furcata (Sm.)—ash
seedlings in nurseries in the Lake States; Colocasia propinqui-
linea (Grote)—birch, beech, maple, walnut; C. flavicornis (Sm.)
—hickory; Charadra deridens (Guen.)—oak, maple, elm, birch;
Raphia frater Grote—poplar, willow; Euxoa scandens (Riley) —
young white and overcup oaks (serious defoliation reported in
Minnesota) ; the zebra caterpillar, Ceramica picta (Harr.)—usu-
ally most abundant on herbacious vegetation, but occasionally
damages pine seedlings in nurseries in the Northern Great
Plains; Orthosia hibisci (Guen.)—the opening buds and fruit of
many deciduous trees and shrubs and a few conifers; Lithophane
latincerea Grt.—many species of deciduous trees (severe defolia-
tion of red maples on islands in Penobscot River, Maine, since
1935) ; the green fruitworm, L. (=Grapholitha) antennata (W1k.)
—ash, boxelder (has occurred in large numbers locally in New
York and Vermont); Anomogyna eliminata Gn—various conl-
fers, especially balsam fir and jack pine (one of the most com-
mon of the cutworm larvae on conifers in the Northeast) ; and
Palthis angulalis Hbn.—a wide variety of conifers, especially
balsam fir and white spruce,also occasionally on various deciduous
trees.

Clean cultivation is recommended for preventing the develop-
ment of damaging populations of cutworms in forest nurseries.

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

309

refers to the fact that in some species there are backward-pro-
jecting tufts on the hind margin of the wings which protrude
when the wings are folded. The larvae are usually solitary, but
some are nest builders. When disturbed, they often elevate each
end of the body and remain attached by the four pairs of pro-
legs at the middle of the body, Many feed exposed on the foliage;
others feed from within folded leaves or tents.

The poplar tent maker, [chthyura inclusa Hbn., 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.

Adults appear from March to July and from July to August,
depending on location. Eggs are laid in clusters on the undersides
of leaves. The larvae are gregarious and live in tents or webs
which they construct by pulling together the edges of one or more
leaves and lining them with silk (fig. 115). They feed from May
to October, then crawl to the ground and pupate in loose cocoons
to spend the winter. Old, abandoned nests often remain on trees
throughout the winter. There are one and a partial second or
two generations per year. This species often seriously defoliates

F-519526
FIGURE 115.—Nest of the poplar tent maker,
Ichthyura inclusa, torn open to show the larvae.

310

small groups of trees, especially trees growing more or less in the
open. Twigs supporting large webs sag badly.

Other species of Jchthyura frequently encountered are J. albo-
sigma Fitch on aspen; and I. apicalis Wlkr., I. brucei Hy. Edw.,
and I. strigosa Grote on aspen and willow. None are very impor-
tant.

The yellow-necked caterpillar, Datana ministra (Drury), occurs
in southern Canada and throughout most of eastern United States.
Its food plants include many species of fruit, shade, and forest
trees. Important forest and shade tree hosts include white and
yellow birch, basswood, elm, oak, maple, butternut, walnut,
mountain ash, hop hornbeam, and honey locust. The adult has a
wingspread of about 50 mm., and its forewings are cinnamon-
brown, 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 is marked longitudinally with alternate
yellow or whitish and black stripes.

Adults appear during June and July. Eggs are laid in masses
of 100 or more each on the undersides of the leaves. The larvae
feed in colonies (fig. 116.) 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 2 to 4 inches where
they pupate and spend the winter. There is one generation per
year.

Damage is seldom serious in the forest, although heavily in-
fested trees may be completely defoliated. Fruit, shade, and orna-
mental trees are injured most severely. Two species of tachinid
parasites, Compsilura concinnata Meig. and Winthemia datanae
Town. are important natural enemies.

Datana angusi G. & R. feeds on hickories, oaks, beech, gray
birch, and butternut and occurs throughout eastern United States,
west to Illinois, and along the north shore of Lake Erie in On- -
tario, Canada. Full-grown larvae resemble those of D. ministra,
but differ in having an entirely black cervical shield.

COURTESY ILL. NAT. HIST. SURV.
FIGURE 116.—Cluster of lar-
vae of the yellow-necked
caterpillar, Datana min-
istra,

oll

Datana perspicua G. & R., the sumac datana, feeds on sumac
throughout most of eastern United States and in southern Can-
ada. Full-grown larvae are moderately hairy and about 50 mm.
long. The head is dark reddish to black, the cervical shield red-
dish brown, and the anal plate, blackish. The body is deep straw
or lemon yellow, with 11 longitudinal, dark reddish-brown to
blackish stripes.

Datana contracta Wlkr. feeds cn oak and sycamore in 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 drexeli Hy. Edw. 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, Datana integerrima G. & R., occurs
commonly in southern Ontario and throughout eastern United
States where it feeds on a wide variety of deciduous trees, pre-
ferably 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 are up to
50 mm. long. The body is black with longitudinal, yellowish
stripes and is covered with long white, or dirty gray hairs.

Adults are present during the spring and summer. Egg laying
begins in early June, the eggs being deposited in masses on the
undersides of leaves. The larvae feed in colonies until almost full
grown (fig. 117). They are often found in masses on the trunk

COURTESY ILL. NAT. HIST. SURV.
FIGURE 117.—Colony of larvae of the walnut caterpillar, Datan integerrima.

312

and larger limbs where they congregate to molt. Later, they re-
turn 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

eee) are one or two generations per year, depending on locality
3828).

The walnut caterpillar is frequently a serious pest of walnut.
Trees heavily defoliated 2 or more years in succession are seri-
ously 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.

Datana major G. & R. 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 south-
ern Canada and the Northern States, where it feeds on willow
and aspen. Full-grown larvae are about 37 mm. long. The head is
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 promi-
nent, pale rust hump on the eighth abdominal segment. Larvae
are found from June to October and the winter is spent as a pupa
in the ground. There are one and sometimes a second generation
per year.

Pheosia rimosa Pack. Occurs rarely in eastern Canada and from
Coast to Coast in northern United States. Its hosts are recorded
as poplars, especially trembling aspen, and willow. Full-grown
larvae are lead-colored with a 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 Pack. occurs on
paper birch in the Northeastern States and southeastern Canada.

Nadata gibbosa (J. E. Smith) 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

313

is spent as a pupa in the ground. There may be two generations
per year as far north as New England.

Nerice bidentata Wlkr. 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 abdomi-
nal 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 are one or one and a partial second
generations per year.

Symmerista canicosta Franclemont [=albicosta (Hbn.)], the
red-humped oakworm, occurs in southeastern Canada and
throughout much of northeastern United States. Its hosts are
various oaks, preferably white and bur, and several other decidu-
ous trees such as basswood, sugar maple, paper birch, beech, and
elm. An outbreak covering several thousands of acres of oak type
occurred in Michigan in 1957 and 1958. 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 yel-
lowish body that increases in width back to an orange-red en-
largement on the eighth abdominal segment. The body is also
marked with fine black dorsal lines (fig. 118).

Adults appear from May to July and the female deposits her
eggs in masses on the undersides of leaves. The larva feed gre-
gariously 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. Some-
times though, they are heavy enough to cause severe defoliation
in isolated spots.

Symmerista albifrons (J. E. Smith) occurs in southern United
States. Its hosts are much the same as those of the red-humped
oakworm, a species with which it is easily confused both in ap-
pearance and habits. At least 12,000 acres of northern hardwoods
were defoliated in Upper Michigan and Wisconsin during an out-
break in 1961. There are two generations per year. The orange-
humped mapleworm, S. leucitys Franclemont, occurs in the

F-519531

FIGURE 118—Larvae of Symmer-

ista canicosta, the red-humped
oakworm.

Northern States and southern Canada where it feeds principally
on sugar maple. Its life history and habits are similar to those of
the red-humped oakworm.

The variable oak leaf caterpillar, Heterocampa manteo(Dbldy.),
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 (771). The adult is
ashy gray and has a wingspread of about 37 to 42 mm. (fig.
119 A). Full-grown larvae are yellowish green and about 37 mm.
long. The body is variously marked, usually with a broad band
down the back (fig. 119 B).

Pupation occurs in early spring, and the adults appear from
early in 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 eggs. Young larvae skeletonize the lower surfaces of the
leaves and older ones eat entire leaves, excepting the larger veins.
Mature larvae move to the ground and spin cocoons in the litter
or top soil. 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 millions of acres
and extending for hundreds of miles. 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.

F-496932, 496931
FIGURE 119.—The variable oak leaf caterpillar,
Heterocampa manteo: A, adults; B, larvae.

315

The saddled prominent, Heterocampa guttivitta (Wlkr.), occurs
in southeastern Canada and throughout eastern United States.
Beech, paper birch, and sugar maple are its preferred hosts, but
practically all other species of deciduous trees are fed upon in
heavily infested stands, particularly the oaks and poplar. 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 green with purple, gold, and brown markings on the
back, and they are about 30 mm. long (fig. 120).

Adults emerge in late May or early June and the female de-
posits up to 500 eggs singly on the leaves. Young larvae skeleton-
ize 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 leafmold 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 cen-
tury (148, 582), during which heavy defoliation occurred over
large forested areas. In areas suffering 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.

Other species of Heterocampa likely to be encountered in eas-
tern forests include: H. umbrata Wlkr.—oak and maple; H. biwn-
data Wlkr.—beech, paper birch, wild black cherry, and maple;
and H. bilineata (Pack.)—elm and birch.

Fentonia marthesia (Cram.) occurs from Maine to Florida and
Texas and feeds on the leaves of beech, maple, 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 longi-
tudinal, 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 to pupate and spend the winter.

Dicentria lignicolor (Wlkr.) feeds on various species of oaks
and beech throughout eastern United States. The larva is about
37 mm. long and resembles larvae of the genus Schizura. Its
prominent characters 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, parchment-like
cocoon on the ground.

The red-humped 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, trembling aspen, willow, hickory,
black locust, dogwood, sweetgum, persimmon, and paper birch.
The adult is grayish brown and has a wingspread of about 30
mm. Full-grown larvae are about 25 mm. long. The head and a
hump on the eighth abdominal segment are red; the body is

316

a

marked with black and yellowish lines and bears a double row of
short, stout, black lines on top (fig. 121). When at rest, the larva
holds the rear end in an elevated position, and, when handled, it
gives off a pungent, disagreeable odor.

Adults appear from late May to August depending on locality.
| Eggs are laid in masses containing up to 100 eggs 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 com-
pletely defoliate one branch before moving on to another. Mature
larvae move to the ground and construct parchment-like cocoons
in the duff in which to spend the winter. There is one generation
per year.

Populations are often heavy in unsprayed apple orchards, along
roadsides and fence rows, and on ornamentals. In light infesta-
tions 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. wnicornis (J. E. Smith), S. lep-
tinoides (Grote), and S. zpomaeae Dbldy. Each species feeds on a
wide variety of hosts, such as apple, wild cherry, elm, aspen,
hickory, beech, paper birch, and willow.

Gluphisia septentrionis Wlkr. occurs in southeastern Canada
and the northern portions of eastern United States. It feeds on
various poplars, especially trembling 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 the pupal stage in cocoons on the ground.
As a rule, there is one generation per year, but in some localities
there may be two.

F-519533

FIGURE 120.—Dorsal and _ lateral

views of larvae of the saddled .

prominent, Heterocampa gutti-
vitta.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 121.—Larva of the red-

humped caterpillar, Schizura
concinna.

317

The genus Cerura is represented in eastern forests by C. bore-
alis (Bdv.), C. occidentalis Lint., C. cinerea Wlkr., and C. multi-
scripta Riley. The first feeds on aspen, willow, and black cherry;
the last three on aspen and willow. Notodonta simplaria Graef.
also feeds on aspen and willow. Misogada wnicolor (Pack.) occurs
on sycamore, and Hyparpax aurora (J. E. Smith) and H. pero-
phoroides (Stecker) are found on oak.

FAMILY LYMANTRIIDAE
TussockK MOTHS

This family includes some of the most serious tree defoliators
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 which hardens and forms a protec-
tive covering. The hairs of certain species are poisonous in all
stages to man when they come into 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 (=Notolophus) antiqua (Linn.),
an introduced species, 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. The male adult
is rusty colored. The forewings are crossed by two darker bands
and each bears a conspicuous 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 pro-
thorax and abdomen are similar to those on the larvae of the
white-marked 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 north to
New England. This species is sometimes abundant locally. Con-
spicuous defoliation has been recorded in Canada.

The white-marked tussock moth, Hemerocampa leucostiqma (J.
E. Smith), occurs commonly throughout eastern United States
and eastern Canada and feeds on a wide variety of deciduous and
coniferous trees. Preferred species appear to include the follow-
ing: apple, basswood, elm, and poplar; Norway, silver, and syca-
more maples; sycamore, paper, and yellow birches; larch; and bal-
sam 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, spanning about 30 mm. The forewing has a con-
spicuous, white spot near the anal angle, and is marked with dark
wavy bands. Full-grown larvae (fig. 122) 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 brush-like tufts of white or

318

COURTESY CONN. AGR. EXPT. STA.
FIGURE 122.—Larvae of the
white-marked tussock moth,
Hemerocampa leucostigma.

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 consider-
able 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 the body. The
pupal stage lasts about 2 weeks, and the number of generations
per year varies from one in the northern parts of the insect’s
range to three in the South.

The white-marked tussock moth is a pest of shade trees, par-
ticularly American elm, in cities and towns. It also occurs in
forested areas but usually causes minor damage there. Howard
(382, 383) discussed the life history, habits, and parasites of the
species.

Hemerocampa definita (Pack.), the definite-marked tussock moth,
occurs in southern Ontario and several of the Eastern States and
feeds on quite a wide variety of deciduous trees such as willow,
apple, wild cherry, elm, paper birch, red oak, red maple, and ash.
The adults and larvae closely resemble those of the white-marked
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 the Northeastern States these egg masses are often
mistaken for those of the gypsy moth. In addition to the con-
spicuous 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

319

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 (Pack.), the dark tussock moth, is some-
times 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 infesta-
tions on valuable shade and ornamental trees may cause serious
damage.

The pine tussock moth, Dosychira (=Olene) plagiata (Walker),
occurs in the Northeastern States west to the Lake States and in
southeastern Canada. The larvae feed on various conifers such as
jack pine, red pine, eastern white pine, spruce, and fir. Jack pine
is especially favored. The moth is gray-brown with lighter and
darker stripes across the forewings and has a wing expanse of
25 to 87 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.

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 mid-
crown but also on the trunk, dead twigs, and similar vegetation.
Young larvae feed on the flat surfaces of needles. During August,
second or third instar larvae svin 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 (736).

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 tens of
thousands of acres of jack pine, red pine, and white spruce were
recorded in the region.

Other eastern species of Dasychira are as follows: D. atomaria
(Wlkr.), and D. vagans (B. & McD.)—sometimes fairly abundant
on oaks in the Northeast; D. meridionalis (B. & McD.)—on oak
in the Southeast; D. cinnamomea (G. & R.)—on elm from New
England to the Lake States; and D. tephra Hbn.—on oak in
Maine and southern Canada.

The gypsy moth, Porthetria 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 silk worm. During the course of his work,
some of the eggs were spilled or lost or some of the larvae es-
caped, and the species became established. About 20 years later, it
had spread over an area of about 350 square miles around Boston,

320

and shade and fruit trees were being completely defoliated. Dur-
ing the next 5 years, it spread rapidly, and populations increased
enormously. Since that time, it has continued to spread, but at a
far less rapid rate. It now occurs throughout the New England
States, except northern New Hampshire, Maine, and northeastern
Vermont. It also occurs in eastern New York, northern New
Jersey, northeastern Pennsylvania, southeastern Quebec, and in
the vicinity of Lansing, Michigan. Its hosts include most species
of hardwoods, the oaks, gray birch, and poplar being most highly
favored. Several conifers are also attacked, usually when growing
in mixture with the more highly favored hardwoods. Mosher
(548) published a list of host plants, dividing them into groups of
favorability.

The male of the gypsy moth (fig. 123 A) is dark brown, with
blackish bands across the forewings and has a wingspread of
about 37 mm. Females (fig. 123 B) are almost white and have

F-489191-5

FiGurE 123.—Life stages of the gypsy moth, Porthetria dispar: A, adult

male moth; B, adult female moth; C, female pupa and cast larval skin;
D, full- -grown larva; E, egg mass ‘laid on bark of a white oak tree.

321

eh ee

wingspreads of about 50 mm. The female abdomen is clothed in
yellowish hairs and is too large and heavy for her to fly. Full-
grown larvae (fig. 123 D) 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 5 pairs of blue spots, followed
by a double row of 6 pairs of red spots, on the dorsum. The pupa
(fig. 123 C) is reddish brown with a sprinkling of reddish hairs.

Adults appear and mate in late July and August. The males fly
vigorously in a zig-zag manner, often within a few feet of the
ground on warm days. 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. 123 E)
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. 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 towards the
tops of trees in search of foliage. During this period they often
spin down on silken threads, especially when disturbed, and some
may be transported considerable distances by the wind (147).
Newly-hatched larvae feed first on leaf bases, then on the leaf sur-
faces, chewing small holes in the leaves. Older larvae feed al-
most 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 (259). Large larvae consume
entire leaves, except the larger veins and midribs. In heavily
infested stands the entire crop of foliage may be consumed be-
fore the larvae reach maturity. When this happens, tne larvae
usually vacate the trees and often migrate considerable distances
in search of food. In heavily defoliated stands when the food
supply is depleted but not completely consumed before the larvae
reach maturity, many succeed in pupating although they are
smaller than normal in size. 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, forest debris, and the like. In neavy 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 im-
portant forest insects in the United States. Enormous sums of
money, probably in excess of 100 million dollars, have been ex-
pended since the turn of the century by the Federal Government
and the States involved to suppress it and to prevent its further
spread. In 1923, a “barrier zone” from 25 to 30 miles wide and
about 250 miles long, was established from Long Island to the
Canadian border to prevent further western spread of the spe-
cies by intensive eradication efforts (121). Despite these efforts,
outbreaks have continued to occur in the older infested portions
of New England, some of which covered tremendous areas, and
extensive infested areas occur west of the barrier zone. In 1945,

322

an estimated 1,500,000 acres were defoliated in the New England
area. Losses through mortality of defoliated trees have been
great. From 1933 to 1952, they amounted to an estimated 2,279,-
819 cords and 128,951,000 board-feet of merchantable timber, hav-
ing a total estimated value of $4,223,556 (592).

Gypsy moth damage is not confined to the killing of trees.
Heavy losses in reduced growth of surviving trees are also in-
curred (592). Continued defoliation also causes loss of tree vigor,
leading to attacks by secondary insects and fungi. Surviving trees
are often stunted, ill-formed, and unmerchantable. Further losses
result from the reduction in quality of forest sites in stands
suffering repeated defoliation.!* in increased fire damage, in the
impairment of watersheds, and in reductions in esthetic, recrea-
tional, and wildlife values.

When the gypsy moth was introduced, the insect parasites,
predators, and pathogens affecting it in its native habitats abroad
were left behind, and it encountered none here 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 continued intensively for about 25 years (128, 387).
Collections representing 45 different species of the parasites and
predators were imported, colonies of which were released in in-
fested stands in this country. Nine species of the parasites and
two of the predators became established. Two are hymenopterous
egg parasites—Dencyrtus kuwanat (How.) and Anastatus dis-
paris Ruschka; four are tachinid larval parasites—Compsilura
concinnata (Meig), E'xorista larvarum (L.), Parasetigena agilis
(R.-D.), and Blepharipa scutellata (R.-D.) ; three are hymenop-
terous larval parasites—Phobocampe disparis (Vier.), Apanteles
melanoscelus (Ratz.), and Monodontomerus aerus Wlkr.; and two
are coleopterous predators—Calosoma sycophanta (L.) and Cara-
bus auratus (L.). The current status of these enemies of the gypsy
moth and effectiveness in its control are discussed by Dowden
(205).

One of the most important factors affecting heavy gypsy moth
populations is the so-called ‘‘wilt’’ disease, caused by a nuclear-
polyhedral virus, Borrelinavirus reprimens. This pathogen is also
a native of Europe. It is believed to have entered this country in
gypsy moth larvae imported for the recovery of parasites. Dur-
ing moth epidemics it increases tremendously and _ practically
wipes out populations over large areas. It occurs only rarely in
light infestations; thus, is ineffective in preventing outbreaks.
Other natural control factors include low winter temperatures—
exposed eggs are unable to withstand temperatures lower than
—?25° F (697). Late spring frosts kill newly hatched larvae, and
rodents devour large larvae and pupae found on the forest floor.
Many larvae also die of starvation in woodlands entirely stripped
of foliage before the larvae reach maturity. Bess (63) and Camp-
bell (133) discuss the population dynamics of the species.

Silvicultural practices designed to promote the health and
vigor of stands is helpful in increasing tree resistance to gypsy

12 House, W. P. 1952. Appraisal of damage by the gypsy moth in New
England, 1933-52. Unpublished report, U. S. Dep. Agr., Bur. of Entomol.
and Plant Quar., Greenfield, Mass., 22 p.

323

moth damage (64). A reduction in the proportion of favored host
species also decreases the chances of infestation and limits the
degree of population build-up (532).

Insecticidal spraying by aircraft has been most widely used
in gypsy moth control. Attempts to secure control by spraying
an infested stand with the nuclear-polyhedral virus, Borrelina-
virus reprimens appeared to be successful (617). Spraying with
the microbial insecticide, Bacillus thuringiensis thuringiensis
Berliner (455) has given promising results. Other promising
methods of control include male sterilization by gamma irradia-
tion (300), sterilization through the use of chemo-sterilants
(146), and the use of natural and synthetic sex attractants. The
natural female attractant known as gyplure has been isolated,
identified, and synthesized (397).

The satin moth, Stilpnotia salicis (L.) (fig. 124), an introduced
species first discovered in North America near Boston, Mass., in
1920 (120) 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 Ore-
gon. The larvae feed on most species of poplar and willow. Adults
of both sexes are pure white with a satiny color and have a wing-
spread 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 is 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, ir-
regular holes in the leaf, and full-grown larvae devour the entire
leaf except the large veins. Feeding is completed by June. Pupa-
tion 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. Mostly in July, eggs are deposited on leaves, branches, and
trunks of trees or on other surfaces in masses of 100 to 400 eggs
each. Each mass is oval, about 9 mm. long, and covered with a
glistening white secretion. Hatching occurs in about two 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 craw] to limbs or the trunk
and spin hibernaculae in bark crevices or under loose bark where
they remain until the following spring (123).

The satin moth is not very important as a forest insect in the
Eastern States, although heavy infestations are occasionally re-
ported. From time to time, it seriously defoliates poplars in orna-
mental plantings. Introduced parasites are generally credited
with holding populations to harmless levels (205) with some as-
sistance from low winter temperatures, disease, and birds. The
hymenopteron, Apanteles solitarius (Ratz.), appears to be the
most effective parasite; however, the dipteron, Compsilura con-

324

$

. <f
Sos ao — = eS “
: eee @ &
> ef FS ‘ i
Ce a mes, >
ES Be = = = oye ‘.
ey. x e “ Aaa re
Bi

aah ee

F-519523

FIGURE 124.—The satin moth, Stilpnotia salicis: A, male adult; B, female

adult; C, larva; D, male pupa; E, female pupa; F, egg cluster; G, cocoon;
H, hibernating web on bark.

connata Meig. and the hymenopteron, Eupteromalus nidulans
(Thomson) are also important.

The brown-tai! moth, Nygmia phaeorrhoea (Donov), (fig. 125)
an introduced species, was first recorded in North America in
Somerville, Mass., in 1897. During the next few years, it in-
creased enormously and spread rapidly. By 1905, it was extremely
abundant throughout Rhode Island, eastern Massachusetts, south-
ern New Hampshire, and southwestern Maine, and had been re-
corded 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 heav-
ily infested. Infestations also occurred in Vermont and west of
the Connecticut River in Massachusetts and Connecticut. Since

329

; F-519522

FIGURE 125.—The brown-tail moth, Nygmia phaeorrhoea: A, 2 winter nest;
B, cocoon in leaves; C, mature larva; D, female depositing egg mass; E,
egg mass on leaf; F, male adult; G, female adult.

then, the area and intensity of infestations have declined greatly.
Since the late fifties, infestations have been recorded from only a
few coastal counties in Maine and New Hampshire and from
Cape Cod. During the first several years following its introduc-
tion, its hosts were recorded as several species of deciduous trees,
principally apple, pear, plum, oak, willow, elm, and maple. At
present, infestations occur almost entirely in old, abandoned
apple orchards and on beach plum.

Brown-tail 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

326

about 37 mm. 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 nu-
merous tubercles with long, barbed hairs and many short, brown
hairs on the dorsum and sides. These hairs are poisonous to man
and cause a severe rash when they come in contact with the 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 gre-
gariously 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 2 to 6
inches long where they spend the winter. Feeding is resumed in
the spring, and the larvae become full-grown by mid-June. Pu-
pation occurs in silken cocoons spun usually among the leaves at
the tips of twigs (121).

The brown-tail moth, although one of the better known forest
and shade tree insects in eastern United States, is no longer of
economic importance. For some undetermined reason, it has al-
most disappeared. Introduced parasites, many of which became
established (205), and a fungus disease probably helped toward
its control, but it seems unlikely that they were primarily respon-
sible for its decline.

FAMILY LASIOCAMPIDAE
TENT CATERPILLAR MOTHS AND ALLIES
The family Lasiocampidae is represented in North America by
less than 30 species, several of which are important economic

pests of trees. The moths are medium-sized 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 Mala-
cosoma Hubner, and are commonly known as tent caterpillars.
Four and a possible fifth species occur in eastern United States
and eastern Canada. The remaining eleven North American spe-
cies have a western distribution. The genus was recently revised
by Stehr and Cook (687). They include keys to the adults, ma-
ture larvae, and egg masses of all species. They also discussed
their distribution, their hosts, and some of their habits.

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 de-
posited, they are held in place by a frothy substance called spu-
maline (358). The majority of species also 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
ege shells, then on the buds and young leaves. Those of tent-
building species also begin immediately to construct a tent on a
branch or in a nearby crotch. They do not feed from within the

By A|

tent, but on the leaves of neighboring branches. As they crawl
to these branches, they spin strands of silk which 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 feed-
ings. 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 in which
to pupate. The cocoons are about 1-inch long, fairly loosely con-
structed, and white or yellow because of a powdery material dis-
persed 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, Malacosoma americanum (F.) (fig.
126), is generally distributed throughout the eastern half of the
United States and southern Canada. Its preferred hosts are wild
cherry and apple, but it also attacks a wide variety of other
forest, shade, and fruit trees. The adults are light to dark choco-
late-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 appar-
ently continuous mid-dorsal light stripe, bordered on each side
with longitudinal reddish-brown and black wavy lines. The sub-
dorsal area is marked with a central black area on each segment,
crossed by a vertical blue mark posteriorally.

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 hatch-
ing 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 crotch, and continue to enlarge it as they
grow. From this tent, the larvae craw! 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, t
or on the trunks of small trees. In the Lake States, eggs are often |
found on the trunks of very small trees about 6 inches above the
ground. There is one generation per year (102, 600). :
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 |

328

COURTESY CONN. AGR. EXPT. STA.
FIGURE 126.—Eastern tent caterpillar, Malacosoma americanum: A, Adults;
B, egg masses encircling twigs; C, larvae; D, a typical tent.

black cherry whose wood is of value for furniture (444). The spe-
cies is primarily a nuisance pest. Infested trees in parks, recrea-
tional 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. Periodically, however, popu-
lations reach outbreak proportions. An effective method of control

329

on isolated trees is to prune off and burn twigs containing egg
masses. The destruction of tents on cold, cloudy days when the
tents are still small is also effective.

The prairie tent caterpillar, Malacosoma californicum lutescens
(Neumoegen & Dyar) occurs throughout the Great Plains area
east of the Rocky Mountains to central Texas. Its hosts are re-
corded as choke cherry, Rosa, willow, wild plum, and Ribes. Male
adults range from dark reddish-brown to very light yellow; fe-
males are yellowish to medium reddish-orange brown. The fore-
wings are crossed by light yellowish lines, and the wingspread is
about 87 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 mid-dorsal area of each ab-
dominal segment is marked with an elongate, somewhat pointed,
blue-white dash. These dashes combine to form a broken mid-
dorsal 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 defoli-
ated. Because of the low value of its hosts, the species is of
minor economic importance.

Malacosoma tigris (Dyar), the Sonoran tent caterpillar, 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. They are usually formed near the end of
each larval instar. This species does not appear to be of much
economic importance, unless it occurs in association with other
defoliating species. During recent years, it has combined with
the forest tent caterpillar in heavily defoliating oaks in south
central Texas. :

The western tent caterpillar, Malacosoma californicum pluviale
(Dyar), 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 (687). Alder, shadbush, willow, choke
cherry, birch, apple, plum, cherry, and trembling aspen are known
to be attacked. Full-grown larvae are predominantly black and yel-
low 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 blue, 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.

The forest tent caterpillar, Malacosoma disstria Hubner, occurs
throughout most of the United States and Canada and feeds on

300

a wide variety of hardwoods. In the North and West, trembling
aspen is preferred; in the South, tupelo gum, black gum, sweet
gum 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. 127A). 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 yel-
lowish-buff, keyhole-shaped spot which may be divided to form an
anterior spot and a smaller posterior spot (fig. 127B). The venter
is blue-gray to dark gray, usually with a median spot on each
segment, and often with a dark-gray area running full length of
the body between the bases of the legs.

Winter is spent in the egg shape 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
three instars, the larvae are gregarious. At first, all of those from

A AND B COURTESY OF CONN.
AGR. EXPT. STA.. C, F-506692

grown caterpillar, showing

FIGURE 127.—Forest tent
caterpillar, Malacosoma
disstria: A, adults; B, full-
grown caterpillar, showing
key-hole spots along the
dorsum; C, caterpillars at
rest on trunk and branches
of an aspen.

one egg mass often cluster on one leaf or one group of small ex-
panding leaves. Later, they become more widely dispersed on sur-
rounding foliage. Although this species is referred to as a tent
caterpillar, it does not construct tents. The larvae do lay down
strands of silk along which they travel, however. They also form
silken mats on the trunks or branches on which they congregate
in masses to rest or to molt (fig. 127C). As they approach maturity,
they tend to wander individually over the trees and other vege-
tation 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 which encircle twigs up to 37 mm. in diameter. The eggs are
cemented together and are coated with dark brown spumaline.
There is one generation 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 1940, five general outbreaks, some of which
covered thousands of square miles and lasted for several years,
occurred in northwestern United States and eastern Canada.
Since the 1930’s several outbreaks have also occurred in Virginia,
South Carolina, Mississippi and Louisiana (356). In general,
tree mortality has not been severe, but losses in reduced incre-
ment following defoliation has been great. Studies in Minnesota
indicate that about 70 percent reduction in basal area growth
of aspen occurs 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 (211). Heavy defoliation of bottomland gums in the
South results in substantial mortality and dieback, as well as
severe reduction in annual increment. 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 busi-
ness because of the nuisance created by migrating caterpillars
and the midwinter appearance of defoliated trees during the
tourist season. Rose (619) and Hildahl and Reeks (353) discuss
the effects of defoliation on trembling aspen in Canada.

Outbreaks usually subside after 3 or 4 consecutive years of
defoliation. Several adverse environmental factors are responsi-
ble for population declines. Freezing weather shortly after the
eggs hatch may kill large numbers of larvae. Excessively high
temperatures later in the spring may kill large numbers of adults
and seriously reduce the viability of newly-laid eggs. Mortality
in the late larval instars may be severe or complete as a result
of starvation in heavily or completely defoliated stands. A poly-
hedrosis virus disease sometimes kills enormous numbers of lar-
vae in the late stages of outbreaks. The sarcophagid parasite,
Sarcophaga aldrichi Park., often becomes extremely abundant
during the late stages of outbreaks in the Lake States and greatly
aids in their termination (355). The tachinid parasites, Zenilla

332

protuberans A. & W. and Compsilura concinnata Meig. are also
helpful in control, especially in the Northeastern States.

Tolype velleda (Stoll.), 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 in-
cluding ash, 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 an expanse 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 posteriorally 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, parchment-like cocoons
on the bark. There is one generation per year. This species seldom
causes serious injury. The related but smaller species, T. laricis
(Fitch), feeds on larch. Its range and life cycle are similar to
those of T. velleda.

The lappet moth, E'picnaptera americana (Harr.), is generally
distributed, but not often common, through the Eastern States
and southern Canada. Its hosts are aspen, wild cherry, hickory,
birch, maple, oak, and various other hardwoods. Adults are red-
dish 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 ZANOLIDAE

This family is represented in North America by only three
species, two of which are encountered fairly often in eastern
forests.

Apatelodes torrefacta (J. E. Smith) occurs throughout the
eastern part of the United States and feeds on maple, wild 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 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.

333

Apatelodes 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 net-
work of fine wavy black lines, except for the top of the thorax
which bears two broad black transverse bands. The tops of seg-
ments one and seven bear yellowish-green spots; long brown
and white hairs project forward over the head fom the pro-
thorax; 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 GEOMETRIDAE
GEOMETRID MOTHS

This is one of the largest families in the order Lepidoptera,
with some 1200 species occurring in the United States and Can-
ada alone. The larvae are all 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 ex-
tending the whole body forward. This characteristic, which re-
sults from the absence of 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 in-
teresting habit of standing nearly erect on the posterior prolegs
when disturbed. Motionless in this position, they are almost in-
distinguishable from small twigs or spurs. All members of the
family are foliage feeders, and the majority inhabit forests and
areas bordering woodlands.

Brephos infans (Moeschler) occurs from Coast to Coast in
southern Canada and in the Northern States south to Pennsyl-
vania. Its hosts are listed as white and gray birches and poplar.
The adult is brown-colored and has a wingspan 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 found from May to July, and winter is spent
in the pupal stage.

The fall cankerworm, Alsophila pometaria (Harr) (fig.128),
occurs from the Maritime Provinces to Alberta in southern Can-
ada, and throughout the Eastern States south to North Carolina
and west to Missouri and Montana. Its preferred hosts appear
to be elm and apple, but it also attacks many other hardwoods
such as hickory, maple, ash, beech, boxelder, basswood, cherry,
and the oaks. The male moth is brownish-gray and has a wing-
spread 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.

334

F-519582

FIGURE 128.—The fall cankerworm, Alsophila pometaria: A, larva; B, eggs;

C, pupae; D, male adult; E, female adult laying eggs; F, pupae in
cocoons; G, female moths.

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 sometimes
mottled. A median, longitudinal darker stripe runs down the

330

=

back. Green larvae have longitudinal white lines. There are three
pairs of prolegs—a very small vestigial pair on the fifth ab.
dominal segment and larger pairs on the sixth and anal segments.

The fall cankerworm spends the winter in the egg stage, and
hatching occurs in late April or early May. Young larvae skele-
tonize the young leaves at the tips of branches; older ones devour
all but the midribs or larger veins of the leaves. They become
mature in 5 to 6 weeks and drop to and enter the soil where
they make cocoons in which to pupate. Adults emerge in Novem-
ber or December, usually following some freezing weather. Fe-
males climb the trees and deposit about 100 eggs in compact,
uniform rows of single layers on the smaller twigs and branches.
There is one generation per year.

The fall cankerworm is an important pest of forest and shade
trees. Outbreaks occur periodically, sometimes covering large,
forested areas. Shade trees in urban areas are subject to heavy
infestation and may be seriously damaged. Heavy infestations
in recreational areas are particularly annoying.

Synchlora aerata (F.) feeds on the foliage of willow and apple
in New York State. Full-grown larvae are brownish, have curved
appendages on the dorsum, and are about 20 mm. long. Winter
is spent in the pupal stage, and there may be two generations
per year.

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,
trembling aspen, willow, and beech, but many other hardwoods
such as white birch, red oak, pin and choke cherries, alder, and
shadbush are also attacked. Serious outbreaks have occurred in
both the United States and Canada.

Female moths are light brownish-gray in color and practically
wingless; males are light brown and have fully-developed wings.
The forewings are semi-transparent, banded with brown or gray,
and have an expanse of 25 to 30 mm. Full-grown larvae are bright
green except for three narrow yellow stripes on each side; larvae
are about 18 mm. long.

In Canada, the 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 cocoons in the soil
or duff. Adults appear in the fall. The female climbs trees and
deposits eggs in bark crevices, under loose bark, or in other hid-
ing places on the tree (111). A polyhedrosis virus disease has
occurred commonly in New Brunswick infestations.

The winter moth, Operophtera brumata (L.), an introduced spe-
cies known to have been present in Nova Scotia since about 1930,
now occurs also on Cape Breton and Prince Edward Islands and
in New Brunswick. So far, it is not known to occur in the United
States. If it continues its recent rate of spread, however, it may
soon cross the border. The female moth has a dusky brown to
silver gray body with occasional irregular black spots and is prac-
tically wingless. Males have fully developed wings with an ex-
panse of 27 to 30 mm. The body is dusky brown with black spots

336

and the forewings are dusky brown with obscure markings.

The most common hosts in Canada are apple, red oak, American
elm, red maple, basswood, and eastern hop hornbeam, but a num-
ber 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 de-
foliation 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 they become mature, they drop to and
enter the ground where they construct cells in which to pupate.
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 eggs in bark crevices, under lichens on
the trunks or branches, or in other places affording suitable
shelter (178). Embree (227) discussed the population dynamics
of the species.

Two imported parasites, the tachinid, Cyzenis albicans (Fall.),
and the ichneumonid, Argypon flaveolatum (Grav.), are provid-
ing a considerable degree of control in Nova Scotia (228).

The genus Hupethecia contains a large number of species, four
of which occur in forests of the Eastern States and eastern Can-
ada. EH. palpatata Pack. occurs on various species of pine, espe-
cially white and jack; E. luteata Pack., feeds on various spruces,
especially white, and other conifers, especially balsam fir and
larch; EH. filmata Pears is found on a wide variety of conifers,
especially white spruce and balsam fir; and EF. transcanadata
Mack. which also feeds on a wide variety of conifers, preferably
white and black spruce and balsam fir.

Calocalpe undulata (L.), the cherry scallop shell moth, occurs on
wild and choke cherries throughout much of eastern North Amer-
ica. The adult has a wingspread of about 37 mm. The forewings
are marked with 12 whitish, scalloped, parallel] lines and a black
discal dot; the hindwings are marked with six 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 which they construct by webbing
together the leaves toward the end of a branch. Heavily infested
trees occasionally are completely defoliated.

Eulype hastata (L.), the spear-marked black moth, feeds on
birch, willow, and alder from Coast to Coast and north to Alaska.
Full-grown larvae are about 25 mm. long. The head is shiny
black; the body, 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.

Bapta semiclarata (Walker) occurs on wild cherry in southern
Canada and the Atlantic Coastal States. Full-grown larvae are
light green and about 18 mm. long. The related species, Bb. ves-
taliata (Guen.), is known to occur on birch in Maine.

Deilinea erythemaria (Guen) feeds on willow and poplar in the

337

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, D. variolaria (Guen.), is found on willow and trembling
aspen in the Northeastern States and southwestern Canada.

Physostegania pustularia (Guen,) occurs on red maple in
Maine where it is occasionally abundant. It has also been reported
from widely scattered localities in southern Canada. The adult
has a wingspread of about 25 mm. and is pure white except for
four brownish spots on the margin of the forewing. Full-grown
larvae are green and about 15 mm. long. There are two whitish
lines down the back, bordered by yellowish-white lines; the skin
is quite wrinkled. Larvae are present during May and June.

Semiothisa granitata (Guen.), the green spruce looper, feeds on
the foliage of various conifers such as white pine, spruce, fir,
and hemlock in the Northeastern States and southeastern Canada.
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. The winter is spent in the pupal stage in
the duff. S. ocellinata (Guen.) occurs in the Northeastern and
Lake States and feeds on honey and black locusts. Full-grown
larvae are green except for faint wavy lines of red and are
about 25 mm. long. S. bisignata (Wlk.) occurs in southern Can-
ada and the Northeastern States and feeds principally on white
pine. S. sexamaculata Pack., a widely distributed species, feeds
on larch.

Eufidonia notataria (Walker) occurs on white pine in the At-
lantic Coastal States. In southern Canada, it has been observed
mainly on balsam fir and spruce. Full-grown larvae are deep
green and about 25 mm. long. There is a light, colored line on
the dorsum and two stripes along each side. Larvae are present
from July to September, and winter is spent in the pupal stage.

Melanolophia canadaria (Guen.) feeds on the foliage of various
hardwoods and conifers such as basswood, paper birch, ash, hem-
lock, larch, pine, and spruce in the United States and Canada.
Full-grown larvae are about 30 mm. long. The body has a broken,
purplish line on the top and yellowish stripe suffused with pur-
plish red on each side. Larvae are present from June to early
August, and winter is spent in the larval stage in the ground.

Anacamptodes ephyraria (Walker) has been observed feeding
on hemlock in the Northeastern States and on several hardwoods
in Canada. Full-grown larvae are greenish with reddish-brown
tinges and are about 20 mm. long. The head is bilobed and wider
than the thorax, and the second abdominal segment is usually
swollen at the sides. Winter is probably spent in the egg stage.
A. larvaria (Guen.) feeds on aspen, paper birch, and alder in the
Northeastern States and southern Canada. Full-grown larvae are
yellowish-green and about 30 mm. long. There is a reddish band
on the second abdominal segment, several reddish blotches on
the dorsum, and a pair of blunt tubercles on the eighth abdominal
segment. A. pergacilis (Hulst) feeds on cypress in the South.
During a recent outbreak in Arkansas, several thousand acres
were defoliated.

338

Ectropis crepuscularia (Denis & Schiff.), the saddle-back
looper, feeds on the foliage of a wide variety of trees in the north-
ern tier of States and from Coast to Coast in southern Canada.
South of Canada, it is usually found on hardwoods such as birch,
maple, oak, poplar, and walnut. In Canada, it appears to prefer
conifers. Full-grown larvae are reddish to chocolate brown and
about 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.

Hpimecis virginiaria (Cram.) occurs on sassafras and yellow
poplar in the Atlantic Coastal 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
50 mm. Full-grown larvae are yellowish to dark brown with 5
pale yellowish longitudinal lines and are about 37 mm. long.
Larvae are present during June and July, and they feed at night.
There are one and a possible second generation per year.

Phigalia tites (Cram.) occurs in southern Canada westward to
Saskatchewan and in several Eastern States. Its hosts include
various hardwoods, especially red oak, red maple, basswood,
hickory, and elm. Numerous outbreaks have been recorded in
mixed oak and maple stands in the Northeastern States. 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 func-
tionless. Full-grown larvae are pinkish with many blackish longi-
tudinal lines and are about 37 mm. long. The thoracic segments
are thick, and there are hairy tubercles on all body segments.
Eggs are laid beneath the bark on dead branches. Larvae are
present from May to July, and winter is spent in the pupal stage
in the soil.

The spring cankerworm, Paleacrita vernata (Peck) (fig. 129),
occurs throughout the same general range as the fall cankerworm,
also farther southwest and west to Texas, Colorado, and Cali-
fornia. Its hosts are about the same as those of the fall canker-
worm. Apple and elm are especially favored. Male moths have
a wing expanse 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 posteriorly. Female moths are wingless and gen-
erally 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 promi-
nent 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-

339

F-519581

FIGURE 129.—Spring cankerworm, Paleacrita vernata. A, eggs on

inner surface of a piece of bark; B, male moth; C, female moths;
D, partly grown larva; E, pupae.

340

yellow stripe down the middle of the venter. There is a pair 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 cells in the soil. Pupation occurs in late win-
ter, and the adults appear about the time frost leaves the ground
in early spring. Female moths crawl up the trunks of trees 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. Then
they drop to and enter the soil where they remain until late
winter or the following spring. Like the fall cankerworm, this
species is an important pest of shade trees and in 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 so as to prevent female
moths from climbing them to lay their eggs. (724).

The linden looper, Hrannis tiliaria Harr., occurs in southeast-
ern Canada and throughout eastern United States. 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 12 mm. long, and
is marked with black spots on the sides and back. Male moths
have fully developed wings with an expanse of about 42 mm. The
forewings are buff-colored and marked with two transverses,
wavy brown bands and a sprinkling of brownish dots. Full-grown
larvae (fig. 130) are bright yellow, have rusty brown heads, and
10 wavy black lines running down the dorsum. They are about
Semin One.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 130.—Larvae of the

linden looper, Erannis tili-
aria.

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 ave attracted to
street lights in nearby towns in considerable numbers. Two dip-
terous parasites, Pseudotachinomya webberi Sm. and Zenillia
vulgaris Fall., commonly attack the larvae. A virus epizootic
occurred in an outbreak in Quebec in 1961.

Lycia ursaria (Walker) larvae feed on the foliage of a number
of hardwoods such as paper birch, willow, elm, maple, poplar,
basswood, ash, wild cherry, and alder in eastern Canada and the
Atlantic Coastal States. Willow and paper birch appear to be
preferred. The adult is gray, stout-bodied, and has a wingspread
of about 50 mm. Both pairs of wings are crossed by diffused,
blackish lines. Full-grown larvae are about 50 mm. long. There
are four large whitish spots on the front of the prothorax, and
the body is marked with numerous longitudinal wavy lines and
creamy-white spots. Larvae are present from May to July, and
winter is spent as pupae in the ground. This species is occa-
sionally quite common in New England.

Phaeoura quernaria J. E. Smith occurs in oaks in the Atlantic
Coastal States. 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 is spent in the pupal
stage.

Plagodis serinaria (H.-S.) feeds on various hardwoods such as
red maple, yellow and white birch, beech, and aspen in the North-
eastern 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 promi-
nent 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. phlogosaria keutzingaria Pack. feeds on ash
in the Northeastern States.

Hyperetis amicaria (H.-S.) feeds on beech, yellow and paper
birch, willow, and alder throughout much of eastern United
States and from coast to coast in southern Canada. There are
also records of its feeding on such conifers as larch, white spruce,
and balsam fir. Full-grown larvae are green to brownish and
about 80 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 first thoracic. Larvae are present from July
to early September, and winter is spent in the pupal stage.

The filament bearer, Nematocampa limbata (Haw.), feeds on
the foliage of hemlock, fir, and such hardwoods as maple, oak,
gray birch, wild cherry, horse chestnut, and apple in eastern
United States and eastern Canada. Full-grown larvae are green-
ish-brown with reddish-brown heads and are about 18 mm. long.
There is a pair of tubercles on the first abdominal segment; a

342

LT TT aii aed

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 on the dorsum from the pro-
thorax to the first pair of filaments. Larvae are active from May
to July; adults, from late June to August.

The elm spanworm, E'nnomos subsignarius (Hbn.) (fig. 131),
occurs occasionally in southern Ontario and throughout eastern
United States, west to Michigan, Colorado, and Texas. Its pre-
ferred hosts are recorded as hickory, oak, and ash, but it also
attacks a large number of other hardwoods (140). About 100
years ago, it was best known as a shade tree pest in the larger
cities of eastern United States (343). In recent years it has been
predominantly a forest pest, with widespread severe outbreaks
occurring in the southern Appalachians (238).

The adult is a powdery white moth with a wingspread of 30
to 87 mm. Full-grown larvae are usually dull, slate black in color
except for rusty head capsules; they are slightly more than 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,
producing shot-hole effects. Older ones eat the entire leaf with
the exception of the midrib and petiole. To pupate, mature larvae
spin coarse, net-like cocoons of silken threads, often on partly

bs
y

iu 3

C D

F-500617, 500618, 500615, 500619
FIGURE 131.—Elm spanworm, Ennomos subsignarius:
A, adults; B, eggs; C, larvae; D, pupae.

343

eaten leaves in which to pupate. In completely defoliated stands,
cocoons may be spun on exposed branch tips, in leaf axils, in
bark crevices, or on stumps or undergrowth.

The elm spanworm is capable of completely defoliating large
areas of mixed hardwood forests during outbreaks. This has hap-
pened during recent years in the lower Appalachians. During
intervals between outbreaks, populations often exist at very low
levels. The recent spate of outbreaks in the South was unex-
pected. 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
hymenopteron, Telenomous alsophilae Vier., parasitized and de-
stroyed over 80 percent of the eggs in certain areas (141).
The predacious beetle, Calosoma scrutator (F.), a voracious
feeder on lepidopterous larvae, was also abundant in outbreak
areas.

Pero honestarius (Wlkr.) occurs on black locust, wild cherry,
and larch in the Eastern States. Full-grown larvae are dark
brown, with lighter and darker markings; increase in girth to-
ward the rear; and are about 37 mm. long. There appears to be
two generations per year. P. morrisonarius Hy. Edw. occurs on
a wide variety of trees, including fir, spruce, larch, pine, willow,
aspen and other trees.

Neptyia semiclusaria (Walker) occurs on fir, larch, pine,
spruce, and hemlock in the Northeastern States and on sand pines
in Florida (347). Full-grown larvae are pale yellow with a wide,
orange-red dorsal stripe and thin, longitudinal black and white
lines; they are about 50 mm. long. Eggs are deposited in masses
under bark scales, and the larvae feed on the old needles. Pupae
are found among these needles, some of which are tied together
with strands of silk.

The false hemlock looper, Neptyia canosaria (Walker), occurs
on hemlock, fir, white and black spruce, and larch in the North-
eastern States and southern Canada. Full-grown larvae are about
25 mm. long. The head is pale whitish or reddish-brown and bears
a few biack dots; the body is whitish 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 hemlock looper, Lambdina fiscellaria (Guen.), occurs from
Newfoundland 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 out-
breaks such as larch, red and black spruce, cedar, arborvitae,
jack pine, basswood, maple, paper and yellow birch, elm, and wild
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 Massachuetts, New Hampshire and Vermont from 1949 to
1952. The adult is creamy-tan with grayish-brown with a tinge
of purple and has a wingspread of about 62 mm. Two irregular
purplish-brown lines cross the forewings, and a dot of the same
color lies between near the costal margin. Full-grown larvae
are usually grayish-green to grayish and are about 30 mm. long.
The head and body are flecked with black dots.

344

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
shoots; older ones, on old needles. Needles are often chewed off
at the base and drop; others, incompletely chewed off, usually dry
out, turn brown, then drop. Pupation takes place in bark crev-
ices, or in masses of lichens on or near the bark (186).

Hemlock looper outbreaks may develop very suddenly, the most
serious ones occurring in mature and over-mature hemlock and
balsam fir stands.

Lambdina athasaria pellucidaria (G. & R.) occurs in several
Atlantic Coastal States and feeds on pitch, red, shortleaf, and
possibly other pines. Several widespread outbreaks have been
recorded during the past 60-odd years. The adult is ash gray to
smoky in color and has a wingspread of about 37 mm. The fore-
wing 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 larvae are pale straw to yellow with black
markings and are 25 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 athasaria athasaria (Walker) occurs on hemlock in
several Eastern States. Local outbreaks have occurred in Massa-
chusetts, Connecticut, and Ohio. The adult resembles the adult of -
L. athasaria pellucidaria except for a slightly narrower wing-
spread. 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. L. endopiaria (G. & R.) has been
observed on oak in the Northeastern States.

The chain-spotted geometer, Cingilla catenaria (Drury), occurs
in eastern Canada and the Northeastern States. It has many hosts
including gray and paper birch, oak, poplar, willow, wild cherry,
balsam fir, larch, and white spruce. Young white and red pines
growing in mixture with the above hosts are also subject to heavy
defoliation. Blueberries, huckleberries, and small trees growing
in pastures and cutover areas are especially subject to infesta-
tion. Local outbreaks have been recorded frequently in the North-
eastern 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 mark-
ings. 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
producing chain-like effects. There also are three or four thin
lines below these rows of dots. Winter is spent in the egg stage.
Larvae are present from June to August.

345

Deuteronomous magnarius (Gn.), 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, bass-
wood, maple, aspen, white 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 con-
spicuous lobe near the middle, and is shaded with brown on the
outer margin. Mature larvae are yellowish-green and about 50
mm. long. Reddish areas occur on the tops of segments two and
five and on the venter of segment three. Adults deposit eggs dur-
ing August and September. 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.

Tetracis lorata Grote occurs on wild 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
with white markings and a black line running down the middle
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.

Abbottana clemataria (J. E. Smith) occurs on basswood,
maple, trembling aspen, paper birch, wild cherry, and hemlock
in southeastern Canada and the Middle Atlantic 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 abdminal segments are swollen on top; and
there are prominent tubercles on the fifth and ninth abdominal
segments. Larvae feed from June to August, and winter is spent
as pupae in cocoons in leaves on the ground.

Prochoerodes transversata (Drury) occurs on various hard-
woods such as maple, oak, willow, trembling aspen, paper birch,
and mulberry in southern Canada and 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 thor-
acic segment is swollen and streaked with red; and the eighth
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 (=Amphidasis) cognataria
(Guen.), occurs in the Northeastern States where it feeds on elm,
willow, poplar, black locust, wild cherry, and apple. The adult is
dark brown to black and has a wingspread of about 55 mm. Full-
grown larvae are about 50 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.
Larvae feed from July to October, and winter is spent in the
pupal stage in the ground.

Campaea perlata Gn. 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. Protobarmia por-
celaria Wlk. is also a common species in Northeastern forests,
where the larvae feed on a wide variety of trees including balsam
fir, white spruce, larch, jack pine, birch, aspen, and willow.

346

LS ea ae Whoa le ee Ge ie

FAMILY LACOSOMIDAE

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 en-
larged, and the frenulum is rudimentary. Two species are known
to occur in eastern United States, neither of which is of economic
importance. However, they both often attract attention.

Cincinnus melsheimeri (Harr.), Melsheimer’s sack bearer, occurs
fairly commonly on oak, especially scrub 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 is 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. Later on it constructs an ellipsoidal portable
case of 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 by which it is anchored and trans-
ports it to the new location. When it is at rest the larva anchors
the case with silk and stops the openings 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 the Melsheimer sack bearer. Adults are dark yel-
lowish-brown, with deeply scalloped forewings and have wing-
spreads of 25 to 30 mm.

FAMILY LIMACODIDAE
SLUG CATERPILLAR MOTHS

Limacodid larvae are sluglike in appearance. The head is con-
cealed in the thorax, the thoracic legs are small, and the prolegs
are replaced by sucking disks. Pupation occurs in dense, brown-
ish, 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 (Clem.), is widely
distributed in 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 on the middle of the back which
resembles a saddlecloth. In the middle of the back 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. 182).

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 (Wlkr.) (fig. 183),
an introduced species first recorded in this country near Boston,

347

COURTESY CONN. AGR. EXPT. STA.

FIGURE 132.—Larvae of the
saddleback caterpillar, S7-
bine stimulea,

: SX. AP ail be

Mass. in 1906 (149), 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 tees, including
Norway and sycamore maples, black birch, cherry, apple, pear,
plum, oak, aspen, willow, honeylocust, hickory, and hackberry.

Adults deposit their eggs either singly or in groups on 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 which fills the spaces between
the threads and hardens. Winter is spent as a prepupa in the
cocoon. People are severely irritated when they come into contact
with the larva’s poisonous spines. The tachinid parasite, Chae-
texorista javana B. & G., was imported from Japan against this
species in 1929 and 1930 and has exerted a considerable degree
of control.

Prolimacodes badia Hbn. 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 Harr. feeds on wild cherry and black
gum in Massachuetts and New Jersey. Sisyrosea textula (H.-S.)
feeds on Norway maple and oak; Packardia geminata (Pack.) on
wild cherry; and Tortricidia flexuosa (Grt.) on oak, gray birch,
and wild cherry in the New England States.

FAMILY MEGALOPYGIDAE
FLANNEL MOTHS
The bodies of flannel moths are covered with dense coats of
scales and long crinkly hairs. The larvae are also covered rather

densely with long soft hairs, with an intermingling of poisonous

348

|

F-519528

FIGURE 133.—The oriental moth, Cnidocampa flavescens: Upper, adults and
cocoons; middle, a defoliated Norway maple; lower, newly hatched and
full-grown larvae.

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), oc-
curs throughout the Southern States where it feeds on various
deciduous trees and shrubs. Forest and shade trees commonly in-
fested 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 with hairs at the rear end tail-like.

349

Young larvae feed gregariously on the surface of the leaf,
skeletonizing it. Older larvae devour the entire leaf. Serious in-
festations have been recorded in Florida and Texas. Several thou-
sand acres of turkey oak were defoliated during an outbreak in
Florida in 1966. Generally speaking, however, the species is most
important as a pest of people, because of its poisonous spines
(68). 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. Hand picking of larvae,
using gloves, is a common control practice. The tachinid para-
site, Carcelia lagoa (Towns.), is sometimes abundant in Texas
infestations.

The crinkled flannel moth, Lagoa crispata (Pack.), occurs
throughout the eastern half of the United States and feeds on a
wide variety of plants, including oak, black locust, birch, cherry,
and apple. According to some reports, it occurs most commonly
in the northern parts of its range; however, it is known to have
comyletely defoliated shin oak over several hundreds of acres of
range land in Texas. The adult is cream-colored, with black wavy
lines and brownish, crinkled hairs on the forewings. Full-grown
larvae are oval-shaped and about 25 mm. long. The body is cov-
ered with long, silky brown hairs which meet in the form of a
ridge along the back, and then slope off roof-like 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 which is lifted as the moth emerges. The stings produced by
the spines on the larvae of this species apparently are less severe
than those produced by related species (68).

Norape ovina Sepp feeds on red bud, mimosa, and beech from
New Jersey and southern Pennsylvania southward. The adult is
a pure white moth with a small amount of crinkly hair. The lar-
vae are spotted and sparsely clothed in tufts of hair.

FAMILY PYRAUSTIDAE

The grape leaf folder, Desmia funeralis (Hbn.), an important
pest of grape, both cultivated and wild, occurs in eastern America
and along the West Coast. It also feeds on Virginia creeper and
red bud. The adult is black except for a white band on the ab-
domen of the male and two white bands on the abdomen of the
female. The forewings are trimmed with white 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 become full-grown, larvae are covered with
sparse, fine, yellow hairs, are translucent yellow green on the
sides, 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 leaf roller, Pantographa limata G. & R., occurs
on basswood in southern Canada and throughout eastern United
States. The adult is white, except for shadings of pale yellow and
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

300

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. 184). 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.

Phylctaenia coronata (Hufn.) [=tertialis (Cren.)], the elder
leaf tier, larvae feed as leaf rollers on the leaves of 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-growr larvae are translucent, whitish or pinkish, and
about 18 mm. long. Winter is spent as a prepupa in a hiberna-
culum usually spun in the hollow stems or pith of elder. There
may be two generations per year. This species occasionally causes
serious defoliation.

FAMILY EPIPASCHIDAE

The pine webworm, Tetralopha robustella Zell., occurs in south-
ern Canada and throughout most of the eastern half of the
United States. Its food plants include several species of pines:
jack, red, white, Scotch, pitch, Virginia, shortleaf, longleaf, lob-
lolly, and slash. Jack pine is preferred in the Lake States and
adjacent parts of Canada. In the Northeast pitch pine is pre-
ferred. The adult has a wingspread 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 which extend through globular masses of brown,
coarse frass webbed together by strands of silk (fig. 1385). These
masses, which are found on the twigs, enclose the needles upon
which the larvae feed and range in length from about 8 to 5
inches. 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 (738).

The pine webworm is often troublesome in pine plantations.
Young seedlings up to 2 feet tall are sometimes completely de-
foliated and killed by the larvae in a 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 (Clem.) occurs in southeastern Canada
and throughout eastern United States. The larvae feed on the
foliage of various hardwoods such as sugar, rea, and mountain
maples, oak, elm, beech, trembling 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 skele-
tonizers; older larvae web together groups of leaves, sometimes

Bol

COURTESY CONN. AGR. EXPT. STA.
FIGURE 135.—Web nest of the pine
webworm, Tetralopha robustella.

F-519529
FIGURE 134.—Apical portion of leaf

rolled into a tube by a larva of
the basswood leaf roller, Pan-
tographa limata.

F-519532

FIGURE 136.—Characteristic nest of Tetralopha asperatella on oak.

352

including all of the leaves on a branch in a web (fig. 136). Heavi-
est infestations apparently occur in the more open portions of
the crown 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 ap-
pears to be only one generation per year (294).

This species had never been considered economically important
until it was shown to be a major factor leading to the develop-
ment of “maple blight,’’ a condition responsible for the killing of
thousands of valuable sugar maples in Wisconsin during the late
1950’s.

Other species of Tetralopha occurring in eastern United States
are T. militella Zell., feeding on sycamore; and T. melanogrammos
Zell., 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.

FAMILY PHYCITIDAE

This family contains many important tree-infesting species.
The larvae differ considerably in their habits. Some feed on
rolled or folded leaves; some construct silken cases 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. Mem-
bers of the family are rather difficult to classify because of the
variability of many of their distinguishing characters.

The pecan leaf casebearer, Acrobasis juglandis (LeBaron), oc-
curs from southern Canada to Florida and Texas, and the larva
feeds on the buds, flowers, and leaves of hickory, walnut, butter-
nut, and pecan. The adult is a grayish moth with a wingspread
of 14 to 17 mm. The forewing is blackish at the middle of the
costa and reddish near the middle of its inner margin. Full-grown
larvae are 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. Finally, before becoming mature, they move
to the leaves to feed. Here, they often chew into the petioles and
cause the leaves to break off and fall. This, plus the injury to the
buds, often results in serious damage to infested areas (295).

Acrobasis caryivorella Ragonot occurs in southern Canada and
throughout eastern United States and the larva feed on the folli-
age of hickory, walnut, and pecan. The adult has dark bluish-
gray to nearly black forewings and a wing expanse of about 20
mm. The larvae are dark grayish-green and about 19 mm. long.
In the spring, they feed by boring into new shoots which they tie
together with silken threads. Full-grown larvae construct oval
cocoons in which they pupate. Two generations per year have
been recorded in Texas. In Florida, there are probably three or
four generations per year. This species has seriously damaged
pecan seedlings in nurseries in Florida and Texas.

308

The pecan nut casebearer, Acrobasis caryae Grote, occurs from
southern Canada to Florida and westward to Illinois and Texas.
Its preferred host appears to be pecan, but it also feeds on hick-
ories. The adult is dark gray, with a ridge or tuft of long dark
scales extending across each forewing, and has a wingspread of
18 to 20 mm. Mature larvae are a dirty, olive green and are about
12 mm. long. Winter is spent in the larval stage in a small case
near the base of a bud. Later, they bore into tender shoots, caus-
ing them 4p become stunted and distorted. Larvae of the second
generation appear in May and bore into newly-set nuts, destroy-
ing from two to five nuts each. Larvae of later generations usu-
ally feed on the shucks only and cause little or no damage. There
are three or four generations per year in the South.

Several other species of Acrobasis also occur on forest and
shade trees in eastern United States. The birch tube maker, A. bet-
ulella Hulst is common on various species of birch, white birch in
particular (in the Northern States and southern Canada) ; A. ru-
brifasciella Pack.—on alder; A. indiginella (Zeller)—on hawthorn
(in the Northeast); A. demotella Grote and A. septentrionella
Dyar—on pecan buds (in Texas) ; and A. feltella Dyar—on hick-
ory (in southern Canada and from New England to Illinois).

The genus Dioryctria Zeller has a world-wide distribution in
the Northern Hemisphere, and the larvae of all species are borers
in conifers. Many are highly injurious to seeds and cones. The
larval habits of some of the more widely distributed species are
quite variable. This suggests that more than one species is in-
volved in certain cases. This can be established only through
further taxanomic and biological investigations. The genus has
been discussed by Ebel (217), Heinrich (336), Neunzig and Mer-
kel (565), Munroe (551), and Neunzig, et al. (566).

Dioryctria clarioralis (Walker) occurs throughout the South
and infests the vegetative buds, male bud clusters, conelets, and
cones of longleaf, slash, loblolly, and shortleaf pines. Moths have
forewings marked with wide and nearly black transverse bands
near the base, and have wingspreads of 22 to 29 mm. (fie. 137).
Larvae are usually yellow-brown to orange-brown, often heavily
suffused with gray; they are about 18 mm. long. This was the
most common of the moths infesting the cones of shortleaf and
loblolly pines in Arkansas in 1960 (799). Cone damage is almost
identical to that caused by D. zimmermant.

Winter is probably spent in the larval stage in conelets. During
the spring and summer, the larvae bore into buds, conelets, and
cones to feed. A characteristic of this species is the presence of
resin-coated silk over the entrance hole in which ejected frass

F-494452
FIGURE 137.—Adult of Dioryctria clarioralis.

354

is accumulated. Pupaticn takes place inside hollowed-out cones
or conelets or in silken cocoons on uninfested twigs or cone stalks.
One generation per year has been recorded in North Carolina.
Farther South there may be as many as three per year.
Dioryctria abietella (D & 8.) [=abietivorella (Grote) ] attacks
the cones, shoots, and bark of many different conifers, particu-
larly in the genus Pinus, and apparently occurs throughout the
range of these trees in the Northern Hemisphere. The larvae are
among the most destructive pests of slash and longleaf pine cones
in the Southern States. Cones infected with the rust Cronartium
strobilinum are especially attractive (513). In the Lake States
and southern Canada, red pine cones also are occasionally seri-
ously damaged. The adult (fig. 188) is dark gray with white,
zigzag lines bordered by black on the forewings, and has a wing-
spread of about 20 to 30 mm. Mature larvae are usually reddish-
purple with a greenish tinge below, and are about 20 mm. long.

F-519534 ~
FIGURE 138.—Adult of
Dioryctria abietella.

There appears to be one generation per year in southern Can-
ada; in Florida there may be five or six. Infestations in rust-
infected cones are noticeable as early as April in Florida. By mid-
June attacks on second-year cones are also apparent.

The spruce coneworm, Doryctria reniculella (Grote), occurs
throughout most or all of the forested regions of Canada and as |
far west as the Lake States in northeastern United States. Its
preferred hosts appear to be white spruce and balsam fir, but
many other conifers including Sitka, Engelmann, red, black,
Norway, and Colorado blue spruces, Douglas fir, larch, and jack
pine are also attacked. The adult is usually silvery-gray and has
a wing expanse of 22 to 26 mm. The forewings are marked with
zig-zag white lines and a white discal spot. Mature larvae are
reddish or amber brown, have hairy warts on each body segment,
and are about 18 mm. long.

In Canada, adults emerge from late June to early August and
deposit their eggs singly or in small groups in such places as
cracks or fissures in the bark of stems and twigs, in lichens,
within the axils of shoots, or between the scales on cones. The
eggs hatch in about 10 days, and the young larvae, without feed-
ing, spin hibernaculae in which they spend the winter. When they
become active again in the spring, they usually mine one or two
needles and then move to and bore into buds, staminate flowers,
or cones. During certain years, this species takes a heavy toll
of white spruce seed (495).

355

The Zimmerman pine moth, Dioryctria zimmermani (Grote),
as presently identified, is widely distributed in the United States
and southern Canada. In the northern part of its range, the
larvae feed in the cambial region and outer xylem under the bark
of limbs and trunks of all commonly grown species of pines. In
the Southeastern States, it feeds almost exclusively within dam-
aged cones of loblolly pine. The adult is gray and has a wing-
spread of about 25 to 37 mm. The forewing is mottled with zig-
zag lines of red and shades of gray. Mature larvae from the
Northern States are pink to greenish and covered with small
black dots, each with a black seta; they are about 18 mm. long.
Larvae from the Southeast differ by having the abdominal seg-
ments bluish-black above. These differences in habits and ap-
pearance of the larvae suggest that not one but two closely re-
lated species are involved (565).

Adults in the Northern States 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 hibernaculae. Here they remain until the
following spring. Feeding begins in May or June, first on 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. 139), and burl-like growths on trunks above gir-
dled whorls and pitch masses (fig. 140) are evidence of attack.
Infested trees tend to be reattacked again and again, thus be-
coming so-called brood trees (614).

In the Southwest, the winter is spent as young larvae in ter-
minals or second-year cones. Older larvae feed in second-year
cones only. Entrance holes in cones are usually filled with !oose
frass but are not surrounded or covered with resin or webbing.
Pupation occurs deep within hollowed out cones and the adults
appear in September and October. Infested cones stop growing
and turn brown when about half to three-quarters grown.

Zimmerman 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 in recent years.

Dioryctria amatella (Hulst.) occurs from Virginia to Florida
and westward through the Gulf Coast States to Texas and Arkan-
sas. It appears to be most closely associated with longleaf pine,
the larvae attacking both the terminals and second-year cones;
however, it also attacks second-year cones of loblolly, slash, and
shortleaf pines, and stem and branch galls of southern fusiform
rust on loblolly and slash pines. The adult (fig. 141) has a wing-
spread of 27 to 32 mm. and the forewing is dark-brown to black,
with prominent patches and zigzag lines of white. Mature larvae
are greenish-white with brownish markings above; they are
about 25 mm. long.

Winter is spent mostly as young larvae at the base of first-
year cones or under expanded terminal needles—occasionally as

356

|

F-492960

FIGURE 140.—Burl - like

growths on trunk above

gridled whorl, a sign of

attack by Zimmerman pine
moth.

F-492959
FIGURE 139.—Damage to ycung pine
by Dioryctria zimmermani.

older larvae in damaged second-year cones. In the Southeast, a
high proportion of the overwintering population attacking elon-
gating terminals of longleaf pine in the spring. The tips of these
terminals soon turn brown and die. Most of the larvae move to
and attack second-year cones by late June, entering them usually
near the base. Entrance holes become filled with frass and pitch,
and the pitch tends to spread over the outer surface of the cone.
By late July, the cones are almost completely browned. If feed-
ing continues in them to the end of the season, they dry out
and remain either partially or entirely closed. Pupation occurs
either in infested terminals or in hollowed-out cones, and there
are several generations per year. This is one of the most abundant
species of the genus in the Southeastern States where it occa-
sionally causes heavy seed losses.

F-519535
FIGURE 141.Adult of
Dioryctria amatella.

3507

Dioryctria cambiicola (Dyar) has been found damaging sec-
ond-year cones and shoots of red pine in southern Canada and the
Lake States. The adult has a wing expanse of about 20 to 30 mm.,
and its forewing is dark purple-brown. Mature larvae are gray to
green and about 15mm. long. The larvae apparently feed pri-
marily in the pith of large current-year’s red pine shoots and
attack cones only when mature (464). D. pygmaeela Ragonot
occurs in eastern United States. It infests the cones of cypress
from Maryland to Florida.

Dioryctria disclusa Heinrich larvae feed in the cones of Scotch,
red, and jack pines in the Northern States and southern Canada
and in the cones of longleaf, loblolly, and scrub pines in the
Southeastern States. The adult is bright white, orange, and red
and has a wingspread of 17 to 29 mm. In the North, a mature
larva is slender, olive-green to buff in color, and about 14 to
18 mm. long. Southern larvae are similar in appearance but are
somewhat larger, from 20 to 25 mm. in length.

In the Northern States and Canada, winter is spent as a first-
instar larvae in hibernaculae spun beneath bark scales on the
branches of red pine. During the spring, the larvae feed first in
staminate flowers. Later, they tunnel in second-year cones (464).
In the Southeast, winter is spent as partly-grown larvae in newly-
formed cones. In the spring, the larvae continue their feeding,
sometimes destroying two or three cones each. Infested cones
have silk attached to their outer surfaces around larval entry
holes and to nearby needles. Pupation occurs in damaged cones
from mid-May to mid-June, and adults are present from late May
to July. There is one veneration per year.

The locust leaf roller, Nephopteryx (=Salebria) subcaesiella
(Clemens), occurs from southern Canada and Maine to West Vir-
ginia and westward to Colorado. The larvae feed inside rolled
leaves of honey locust and wisteria. The adult is gray and has a
wingspread of 25 mm. The forewing is reddish near the base and
has a broken, black terminal line. Mature larvae are about 25
mm. long. The head is blackish and the body green, except for a
dark line down the middle of the dorsum and for five faint yel-
low lines on each side. The winter is spent as pupae in silken
cocoons among leaves on the ground. This species is quite common
at times, but is seldom injurious. N. virgatella (Clem.) is a leaf
roller on black locust. The larvae resemble those of the locust
leaf roller except for their brown heads. N. subfuscella (Ragonot)
is a leaf roller on sumac from Maine to Texas. The adult is ash
gray and has a wingspread of 22 mm. Mature larvae are yellow-
ish-green except for brick-red lines on the dorsum and sides and
are about 25 mm. long.

The lesser cornstalk borer, Vlasmopalpus lignosellus (Zeller), a
widely distributed species, has damaged black locust seedlings in
nurseries in several Southern States. Loblolly pine seedlings in
nursery beds in Virginia and Rhode Island have also been dam-
aged. 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 ochre yellow to light brown and
has a wingspread of 16 to 24 mm. The forewings are long and
narrow and marked with several black spots. Mature larvae are

358

greenish-white with interrupted longitudinal stripes of dark
brown. The winter is spent either in the larval or pupal stage
and there are up to four generations per year in the Deep South.

The American plum borer, Huzophera semifuneralis (Walker),
is widespread and attacks a wide variety of hardwoods, includ-
ing plum, wild cherry, apple, London plane, mountain ash, wal-
nut, persimmon, mulberry, basswood, poplar, sweetgum, and
gingko. 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.

Euzophera ostricolorella Hulst has been recorded from Long
Island to South Carolina and westward to Ohio, Kentucky, Ten-
nessee, 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 10 inches in
diameter (325). Attacks above a height of 2 feet 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 genera-
tion 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 puprlish-brown,
with black terminal dots. Full-grown larvae are white and flat-
tened, they taper toward the rear and are about 28 mm. long.
Attacks occur at the base, the larvae tunneling in the lower inches
of the trunk and in the larger roots entering the crown. Heavily
infested trees may be girdled and killed( 134).

Canarsia ulmiarrosorella (Clem.) occurs in southern Canada
and throughout eastern United States. The larvae feed on the
leaves of elm which they web together with silk. The adult has
a wingspread of 15 to 20 mm. The forewings are dusty white
and crossed by dark gray or blackish bands. 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
southeasten Canada and 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 pine cones in Ar-
kansas; in sumac heads in New York, Pennsylvania, and Texas;
and in fusiform rust cankers on slash and longleaf pines in Flor-
ida. Full-grown larvae vary from yellow with a greenish tinge
to bright yellow orange or orange, and are about 8 to 10 mm.
long.

Actrix nyssaecolella Dyar is a leaf folder or leaf roller of
tupelo from western Pennsylvania to New Jersey and Massachu-
setts. The adult is powdery gray and has a wingspread of 15 mm.
Full-grown larvae are black with yellow heads. Winter is spent
in the pupal stage, and larvae are present during July and
August.

3509

Laetilia coccidivora Comstock larvae feed on various scale in-
sects. According to some reports, it has aided materially in the
control of outbreaks of the pine tortoise scale. It has been re-
corded from Pennsylvania and Ohio to Florida and Texas.

FAMILY OLETHREUTIDAE
OLETHREUTID MOTHS

This family contains a large number of economically important
species of forest insects. 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 feed on the
foliage as leaf miners, from within folded leaves, or on the ex-
posed surface. MacKay (487, 488) described the known larvae of
North American species.

Episimus argutanus (Clem.) is widely distributed and feeds as
a leaf roller on sumac, poison ivy, witch hazel, and various other
shrubs. The adult has a wing expanse of about 12 mm. and is
dull reddish or grayish-brown, mottled with darker 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 needle miner, Taniva albolineana (Kft.), occurs from
coast to coast in southern Canada and from Maine to North Caro-
lina, Colorado, and Idaho in the United States. Its hosts are
white, Norway, Engelmann, Colorado, and black spruce. The
adult is dark brown and has a wingspread of about 12 mm. The
forewings have three irregular, transverse, broken, grayish-white
bands. ;

Adults are present from mid-May to mid-June. Eggs are de-
posited so that they overlap in a single row on the undersides of
needles in groups of two to twelve 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 is 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 needle miner is most important as a pest of orna-
mental 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 handpick-
ing 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 (Zell.) feeds as a leaf roller 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 12 to 16 mm. Full-grown larvae are dark green to almost
black, sparsely hairy, with light-colored tubercles and light-brown

360

oe and are about 18 mm. long. Larvae are present from May
to July.

The European pine shoot moth, Rhyacionia buolinana (Schiff.)
(fig. 142), an introduced species first recorded in North Amer-
ica on Long Island, N.Y., in 1914 (127), is now widely dis-
tributed in the Northeastern States and southern Canada. Out-
lying infestations also occur in Washington, Oregon, and British
Columbia. Its hosts include Scotch, red, Austrian, mugho, Japa-
nese red, Japanese black, ponderosa, eastern white, jack, pitch,
longleaf, and Virginia pines (527). Red, mugho, Scotch and Aus-
trian pines are most heavily attacked in eastern United States,
especially red and mugho. The adult moth is rusty orange-red
and has a wingspread of about 18 mm. The forewings are marked
with several irregular, forked, silvery cross lines; the hindwings
are plain dark brown, and the legs are whitish.

Adults appear in late spring and fly at night. They lay eggs
singly or in groups of two to ten on the bases of buds, on needle
fascicles and twig tips, or on the bark of new and old shoots.

F-493841, 493467 |)

FIGURE 142.—European pine §
shoot moth, Rhyacionia bu-
oliana: 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.

Newly-hatched larvae spin resin coated, tentlike webs between
needle sheaths and the stems of current year’s growth, then they
bore through the sheaths and mine the bases of the needles. About
mid-summer, the larvae move to buds and construct new resin
coated webs. At first, these webs glisten brightly; then they
solidify into yellowish-white masses. Feeding ceases in August.
Winter is spent in the larval stage in a feeding tunnel in a bud.
When activity is resumed in April, the larvae move to undamaged
buds and new shoots, construct new tents, and resume feedings.
During this period, a single larva may feed on more than one
bud or shoot. Larvae reach maturity in May, and pupation occurs
inside the burrow or tent in late May or June. Two or three weeks
later, the larva works its way out of its chamber. There is one
generation per year.

The most important and permanent damage to trees results
from spring feeding. Shoots weakened by larval tunneling fall
over, but continue to grow. This results in the formation of
crooked trunks and branches or “post horns.” The killing of ter-
minal 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. All open-
grown young trees of susceptible species below a height of 20 to
25 feet are subject to attack and damage. Taller trees or trees
growing in closed stands are usually not seriously damaged.

Climate is an important factor affecting the distribution and
abundance of the European pine shoot moth. Overwintering lar-
vae are killed by temperatures colder than about —20° F. Warm,
dry summers followed by mild winters permit maximum 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.
Five introduced species have become established: Orgilus obscur-
ator (Nees), Temelucha interruptor (Grav.), Hulimneria rufife-
mur Thom., Pimpla turionellae (L.), and Tetrastichus turionum
Htg. O. obscurator appears to offer the most promise. Yates
(797) published a key to the nearctic parasites of the genus.

The planting of susceptible pines on good sites, “snow-depth
pruning,” and Christmas tree shearing are helpful in reducing
damage (526). Fumigation in the spring has provided a degree
of control on seedlings and larger ornamental specimens (135).
The reader should consult Agricultural Handbook 331 for details
on registered control methods.

Many publications have been issued on the biology, ecology,
and control of the European pine shoot moth. In addition to those
cited in preceding paragraphs, there are several others that are
of ‘special interest (273, -328, 529, 5380, 599, 755). Articles by
Heikkenen !3 and Torgerson !* are also pertinent.

Rhyacionia adana Heinrich has been recorded from Massachu-
setts, Pennsylvania, Virginia, Michigan, Wisconsin, and Ontario.

13 Heikkenen, H. J. 1963. Influence of site and other factors on damage
by the European pine shoot moth. PhD thesis, Univ. of Mich.

14Torgerson, T. P. 1964. The binomics of the European pine shoot moth
in Wisconsin. PhD thesis, Univ. of Wis.

362

Red, jack, and Scotch pine seedlings usually under 3 feet tall in
nurseries, plantations, and natural stands and the lower half of
trees up to 25 feet tall are attacked. The adult has a wingspread
of 15 to 17 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
th 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 (506). Serious damage has been recorded in
red and Scotch pine plantations in Canada and the Lake States.

Rhyacionia rigidana (Fern.) occurs from Georgia to Texas
and north to Missouri, New York and Maine. 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. Its life history is not too well understood; how-
ever, there appears to be at least three generations per year in
the South. In the North there is probably only one. Damage is
also similar to that caused by the Nantucket pine tip moth but
apparently differs in the ability of this species to damage trees
of large size. Pitch pines up to 15 inches in diameter may be in-
fested, and terminals and laterals of slash pines up to 50 feet
tall have been killed.

The Nantucket pine tip moth, Rhyocionia frustrana (Comst.)
(fig. 148), is widely distributed in the Eastern, Central, and
Southern States. Its hosts include nearly all species of pines
growing within its range, the only exceptions are longleaf
and eastern white pines (798). Slash pine is also somewhat re-
sistant, but is occasionally attacked. In the South and Southeast,
loblolly and shortleaf pines are preferred; in the Northeast and
Mid-Atlantic States pitch, Virginia, and Scotch appear to be
favored; and in the Central States, shortleaf is attacked most
heavily. Red pine has been infested along the coast of Maine.
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 wingspread is 9 to 15 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 de-
veloping tips, or on buds. Newly-hatched larvae wander about the
shoots looking for suitable feeding sights. 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 instar larvae spin new and larger

363

F-494206, 494207

FIGURE 143.—Adult and lar-
vae of the Nantucket pine
tip moth, Rhyacionia frus-
trana.

\. A) |

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 hav-
ing consumed the bud, it bores down the center of the stem. The
larval period lasts for 2 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 five or six generations per
year. In the North Carolina, Virginia area, there are three per
year. Still farther north, in Pennsylvania and Ohio, there are only
two per year.

The Nantucket pine tip moth is a major pest of young plants
in the eastern United States. During recent years, it has become
increasingly abundant and destructive as a result of the estab-
lishment of large areas of pine plantations and seed orchards.
Damage consists of the retardation of height growth, the crook-
ing or forking of main stems, the reduction of cone crops, and,
occasionally, the death of the tree. Attacks are generally re-
stricted to trees less than 15 feet in height and are most severe in
young plantations, but severe attacks on commercial-size trees
have also been reported.

364

At least 60 species of parasites of the Nantucket pine tip moth
have been recorded (798), but they are rarely abundant enough
to provide satisfactory control. Low winter temperatures in the
northern parts of the infested region provide some degree of
control by killing overwintering pupae. Damage can also be re-
duced 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 frustrana bushnelli (Busck), the western pine tip
moth, occurs from the Dakotas and Nebraska south to New Mex-
ico. Its preferred hosts are ponderosa, red, jack, and Scotch pines.
Infestations have been particularly severe in pine plantations in
Nebraska (308). The adult is indistinguishable from those of the
Nantucket pine tip moth, although it tends to be larger. Heinrich
(335) suggested that it and the true R. f. frustrana are one and
the same, but Miller (528) believes that bushnelli may be a dis-
tinct species because of its overwintering habit and size. There is
one generation per year in the Dakotas and two per year in
Nebraska.

In 1925, Campoplex frustranae Cush., a common parasite of the
Nantucket pine tip moth in the East, was liberated in infested
plantations in the Nebraska National Forest. It became estab-
lished immediately and increased rapidly. By 1930, parasitism
at the original liberation point had reached 80%. Unfortunately,
the related tip moth, R. frustrana neomexicana, which the para-
site also attacks but in which it is unable to develop, increased
very rapidly in the area about this time, and parasitism of R. f.
bushnellt declined. This may have happened because the parasite
deposited too many of its eggs on R. f. neomexicana.

Rhyacionia subtropica Miller, a newly-described species (524),
occurs throughout the range of slash pine in the South. Typical
slash pine, Pinus elliott var. elliotti, is the preferred host, but
longleaf, loblolly, and tropical pine are also attacked. South Flor-
ida slash pine, P. elliottw var. densa, is fairly resistant. Heavy in-
infestations have been recorded in slash pine plantations on poor
sites in Florida; whereas naturally regenerated slash pine seed-
lings have rarely been infested. Serious losses of grafted slash
pine scions in tree improvement programs have been incurred.

Rhyacionia sonia Miller, a newly-described species (528), has
been recorded from Maine west to southern Manitoba and from
South Carolina. Its hosts are jack and pitch pines in the North,
and slash and loblolly pines in the South. Superficially, the adult
is indistinguishable from those of R. f. frustrana and R. f. bush-
nellt. Winter is spent in the pupal stage and there is one genera-
tion per year.

The pitch twig moth, Petrova comstockiana (Fern.), 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 14 to 20 mm.

In Ohio, eggs are deposited on twigs during May and June. The
larvae bore into and downward in the twigs for distances of 3 or
4 inches, and pitch masses form over the entry holes (fig. 144).
Winter is spent as a larva under the pitch mass. Development is

365

F-519519

FIGURE 144.—Larva of the
pitch twig moth, Petrova
comstockiana, in the twig
of red pine. Pitch mass
covers the opening to the
larval gallery.

resumed in May, and pupation occurs under the mass. There is
one generation per year. Damaged twigs break off, leading to de-
formation of infested trees. The braconid, Agathis pint (Mues.),
has been a major factor in control (531).

Petrova virginiana McD. feeds on jack pine in the Lake States
and southern Canada. The larvae feed in and hollow out the ter-
minals 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.

Petrova albicapitana (Busck), the pitch nodule maker, 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 wing-
spread of 16 to 23 mm.

Young pines from 1 to 5 feet 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 30 feet tall.
Larvae feed singly under masses of pitch, some of which may be
up to 115 inches 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 cycle, winter
being 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

366

7 ee Ae se i ee ee Lee

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 (7232).

Petrova houseri Miller attacks shortleaf pine in Ohio, West
Virginia, and Georgia. The adult is dark gray with light and dark
brown areas, has brown, fringed hindwings, and a wingspread of
about 12 mm. Prior to its description as a new species (525), 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 forming over the entrance holes,
average about one-half inch 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 1 inch. 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 pint (Muese-
beck), is an important parasite of the species.

The eye-spotted bud roth, Spilonota ocellana (D. & S.), an in-
troduced 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 12 to 16
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 hibernaculae in which to spend the winter. Feeding is re-
sumed in the spring on opening buds and unfolding leaves. These
are bound together with silk. Pupation occurs in June in silk-
lined cocoons. There is one generation per year (571). This spe-
cies is most important as an orchard pest.

Eucosma gloriola Heinrich, the white-pine shoot borer, is widely
distributed in the Northeastern and Lake States and southern
Canada. Its hosts are eastern white, jack, red, Scotch, Austrian,
and mugho pines. The adult is coppery-red, with two shining gray
transverse bands on the forewings, and has a wingspread of
15 to 18 mm.

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 inches above the node of the new shoot and construct
tunnels several inches long toward the tip. Infested shoots are
eventually girdled from the inside (209). 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 (132).

The white-pine shoot borer is a serious pest of young jack pines
in the Lake States and Ontario and in white, red, and Scotch pine
plantations in New York and Pennsylvania. Infested terminals
bend over or break off. Damaged laterals turn yellow, then red,

367

and die gradually as the season advances. Infested trees become
bushy after repeated attacks.

EKucosma monitorana Heinrich attacks the cones of various
pines in the Northeastern States and southern Canada. The adult
is reddish-brown with tan and silver markings on the forewings
and has a wingspread of 13 to 16 mm.

In southern Canada, 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. Last instar larvae sometimes migrate to and enter rela-
tively 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. There is one generation per year (465).

EKucosma sonomana Kft., the jack-pine shoot moth, occurs in the
Lake States and attacks jack pine. The adult has coppery-red
forewings and a wingspread of about 18 mm. The larvae are
brownish-gray and about 12 mm. long. Young plantation pines
from 3 to 12 feet tall are usually attacked, but trees up to 30 feet
tall may also be infested. The larvae bore into the pith, 6 to 8
inches above the branch whorl of the previous season, and mine
downward for a few inches. Then they move back up the shoot for
a few inches, girdle the shoots, bore to the surface and drop to the
ground to pupate. By this time, the shoot has wilted and has bent
over or broken off (fig. 145) (473).

Fucosma cocana Kft., has been recorded as occurring in Mas-
sachusetts, Connecticut, Virginia, North Carolina, and Florida,
and as probably feeding on the cones of loblolly pine. EF. tocullio-
nana Heinrich has been recorded from Ontario and New Bruns-
wick, Canada, and from Wisconsin, Massachusetts, New York,
Connecticut, Pennsylvania, Virginia, North Carolina, and Ten-

F-492898

FIGURE 145.—Damage to pine

terminal by Hucosma sono-
mana.

368

et eS

nessee. The larvae feed on the cones of various conifers, such as
white pine, spruce, balsam fir, and hemlock (6032).

Gypsonoma haimbachiana (Kft.), has been recorded from On-
tario, from the northern tier of States from New York to Michi-
gan 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 18 to 17 mm. The basal portion of the forewing
is darker than the apical portion. Full-grown larvae are pale and
from 13 to 17 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 either 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-instar larvae 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 10 inches of the terminal. There appears to be four or
five generations per year in the Mississippi Delta region (540).

This is one of the most destructive of the insects that damage
young trees of eastern cottonwood. Damaged trees are stunted,
have crooked trunks, and produce too many limbs. This leads to a
great reduction in the quality and quantity of merchantable pulp-
wood, sawlogs, or veneer from these trees.

Proteoteras aesculana Riley, the maple seed caterpillar, occurs
throughout southern Canada and the Northern States, south to
Tennessee. The adult is olive-green with yellow, gray, and black
markings, and has a wingspread of 11 to 18 mm. The larvae bore
in the seeds, leaf stalks, and terminal twigs of horse chestnut and
maple and sometimes cause serious injury. Seedlings in nurseries
have been heavily attacked. Adults emerge in July and August.

The boxelder twig borer, Proteoteras willingana Kft., 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 11 to 18 mm. The larva destroys
dormant leaf buds in the fall and early spring. Later in the spring,
it burrows in succulent twigs (fig. 146), causing the formation of
spindle-shaped galls (594). 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.

The spruce bud moth, Zeiraphera ratzeburgiana (Ratz.), an
apparent introduction from Europe late in the 19th century, now
occurs throughout the range of spruce in northern United States
and Canada (595). In addition to spruce, it feeds on several other
conifers, especially fir. The adult is light brown and has a wing-
spread of about 12 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.

369

F-500805

FIGURE 146-——Harva) vot “the

boxelder twig borer, Protec-

teras willingana, in a box-
elder stem.

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 immedi-
ately below the ground surface. Winter is passed in the egg stage,
and there is one generation per year. Zeiraphera diniana Gn.
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. Z. fortunana (Kft.)
occurs commonly from coast to coast in southern Canada. Infes-

370

tations also have been found in Maine. It apparently prefers
white spruce.

The maple trumpet skeletonizer, E'pinotia aceriella (Clem.), oc-
curs 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 trumpet-like tube of silk and frass on the underside of a leaf,
causing the leaf to 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. E. solandriana
(L.) feeds as a leaf roller on trembling aspen, white birch, and
various other hardwoods. It is sometimes abundant in the North-
east and Ontario.

Epinotia nanana (Treit.), an introduced species first recorded
in North America in Massachusetts in the early days of this cen-
tury, now occurs from Maine to Ohio and Michigan and in On-
tario, Quebec, and British Columbia, Canada. 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 dis-
tinct 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 the 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
bunchs 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 (183).
Webs washed from the trees in the spring with a strong stream
of water before the buds break has been suggested for control on
ornamentals.

Anchylopera plantanana 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 undersur-
face 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 ones feed beneath these webs and skele-
tonize the leaf. The winter is spent in the larval stage (190).

The spruce seed moth, Laspeyresia youngana (Kft.), is rather
widely distributed in the United States and Canada. It attacks the
cones of white, red, black, Colorado, Sitka, and Engelmann
spruces. The larvae make tortuous mines near the cone axis, de-
stroying both scales and seeds. White spruce cones are especially
susceptible. The adult is smoky brown with four cross bars of

371

shining silver and four shining costal spots on the forewings and
has a wingspread of 8 to 11 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
1 to 2 or more years (721). Female moths are reluctant to fly, and
as a result, populations tend to build up on old cones bearing
trees.

The hickory shuckworm, Laspeyresia 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 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, de-
pending on locality. Heavy infestations may seriously reduce
hickory nut and pecan crops. Gathering and destroying infested
nuts during the winter is helpful in control.

Laspeyresia toreuta (Grote) occurs in southern Canada and
south in the Eastern States to South Carolina and Texas. It at-
tacks 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 13 to 15 mm. The mature
larva is creamy white and about 10 mm. long.

Adults emerge during late May and early June in the mid-South
and during late June in Ontario and deposit their eggs in crevices
over the cone surface. Young larvae bore into the upper part of
second-year cones and enter the seeds to feed; older ones bore
around the cone through the bases of scales, feeding on available
seeds. Infestations appear to be heaviest on open-grown trees
with branches to the ground or on trees in low density stands
(439). 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 (465).

Laspeyresia anaranjada Miller occurs throughout the range of
typical slash pine, Pinus elliotti Engelmann, and its south Florida
variety, P. elliotti var. densa Little and Dorman, in the South.
The adult (fig. 147 B) has a wingspan of 14 to 16 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 cross bands. The species has been reared principally from
mature cones of slash pine, occasionally from longleaf cones, and
rarely from loblolly cones.

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 con-
sumes from five to seven seeds. Eventually, the full-grown larva
bores into the woody cone axis where it spends the winter. Pupa-
tion occurs in the spring, and the adults emerge during April and

372

F-519536

FIGURE 147.—Adults of seedworm moths:
A, Laspeyresia ingens; B, Laspeyresia anaranjada.

May. There is one generation per year. According to estimates,
this species destroys from 2 to 10 percent of the seed in open-
grown slash pine stands in northeast Florida each year (514).

Laspeyresia ingeus Heinrich has been recorded from South
Carolina, Georgia, Florida, Alabama, and Mississippi and prob-
ably occurs throughout the range of its favored host, longleaf
pine. It also infests slash pine to minor extent and may occur
occasionally on other southern pines. The adult (fig. 147 A) is
grayish-brown and has a wingspread of 17 to 20 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. Larva feeds during the first
two instars, each in a seed, vacating it and tunnelling 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 1 to 2 inches. 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 1 to 2
weeks later. There is one generation per year, except for occa-
sional individuals that enter diapause (163). This species causes ~
an estimated loss of 21 percent of longleaf pine seed in the Gulf
States region. The related species, L. erotella (Heinrich) attacks
loblolly pine cones in the Southeastern States.

The filbertworm, Melissopus latiferreanus (Wlsm.), occurs
throughout most of the United States and in southern Canada.
The larvae feed in hazelnuts, 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.

Paralobesia liriodendrana (Kft.) 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
mid-vein 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 togther. Pupation
occurs undr the web (593).

The locust twig borer, Ecdytolopha insiticiana Zell., attacks

373

black locust throughout eastern United States and in parts of
southern Canada. It also occurs in Colorado, Arizona, and Cali-
fornia. The adult has a wingspread of about 20 to 25 mm. The |
forewings are dark, ashy-brown with large, dull, pinkish-white |
patches on their outer parts. There are also 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 |
12 to 20 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 3 inches 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 olethreutids likely to be encountered in eastern forests
and their hosts are as follows: Hedi« chionosema (Zell.)—haw-
thorn, occasionally red oak and mountain-ash; Exartema per- .
mundanum Clem.—hickory; FE. quadrifidwm—wild cherry;
Sciaphila duplex (Wlshm.)—poplar; Epinotia similana (Hbn.)—
gray birch; E. lindana (Fern.)—alternate-leaved and flowering
dogwood; and Griselda radicana Heinrich—various conifers,
principally white spruce and balsam fir.

FAMILY PHALONIIDAE

The moths of this family resemble those of the families Oleth-
reutidae and Tortricidae. Only one eastern species is worthy of
mention.

Phalonia rutilana (Hbn.) is an introduced species first re-
corded 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
Juniperus communis on which it is often abundant. The adult is
yellowish and has a wingspread of 10 mm. There are red markings
on the head, thorax and forewings, those on the wings occurring
as four broad cross bands. 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 TORTRICIDAE
LEAF ROLLER MOTHS

Members of this family 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. The wing expanse is usu-
ally 25 mm. or less and appear bell-shaped while folded. The
larvae are usually some shade of green, seldom more than 25 mm.
long, and they pupate in flimsy silken cocoons. Just before the

374

a

pupa transforms to the adult, it works its way partly out of its
cocoon.

Adoxophyes furcatana (WI1k.), a leaf roller on sycamore, oc-
curs 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.

Amorbia humerosana Clem. occurs from coast to coast in
southern Canada and in the Northeastern States south to Penn-
sylvania. Its hosts include a wide variety of tree species, both
coniferous and deciduous. Adults are light gray and have wing-
spreads of about 25 mm. Larvae are light green except for light
brown heads. |

Sparganothis acerivorana (MacKay) occurs in southern Can-
ada and the Lake States. In Canada it feeds on several hard-
woods, 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 during recent years (294).

Sparganothis pettitana (Rob.) occurs in southeastern Canada
and south and west to Florida and the Mississippi River Valley.
The larvae are solitary leaf rollers on various hardwoods, es-
pecially basswood and maples. The adult is plain lemon yellow
and has a wingspread of about 22 mm. The forewings are some-
times 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 (Clem.) has been observed feeding in
the tips and shoots of small red pines in Canada and in the ter-
minals of loblolly pine seedlings in Georgia. Willow, honey locust,
apple, and cherry are also attacked. S. folgidipenna (Blanchard)
occurs over about the same range as S. sulfureana. It feeds on
white ash, maple, large tooth aspen, and paper birch. S. diluto-
costana (Wlshm.) 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. tristriata Kft. occurs fairly com-
monly on jack pine in southern Ontario. It probably occurs in the
Lake States also.

The genus Pandemis Hbn. contains several species which attack
a wide variety of hardwoods in eastern America. P. lamprosana
(Rob.) 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. P. limitata (Rob.)

375

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 wild cherry.

Archips rosanus (L.), an introduced species, occurs in south-
eastern Canada, British Columbia and from New England to the
Lake States. The larvae are general feeders on a number of de-
ciduous growths. Privet appears to be especially attractive. Adults
are dull light-brown to olive-brown and have wingspreads of 17
to 22 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 ugly-nest caterpillar, Archips cerasivoranus (Fitch), occurs
from coast to coast in the Northern States and southern Canada.
Its preferred hosts are choke 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 18 to 25 mm. The forewing is
irregularly speckled with dark reddish-brown and has three
patches of the same color. The hindwing is solid, bright orange.
Full-grown larvae are yellowish or greenish yellow except for
black heads and cervical shields and are about 20 to 23 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. 148). In heavily infested areas, these nests

F-519520
FIGURE 148.—Nest of the ugly-nest caterpillar,
Archips cerasivoranus, on wild cherry.

are often numerous. Some may be large enough to enclose entire
small trees.

The ugly-nest 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. The cutting and
removal of nests provides adequate control in most situations.

The oak webworm, Archips fervidanus (Clem.), 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 18 to 25 mm. The forewing is yellowish brown with
dark patches. The hindwing is uniformly smoky except for a light
colored fringe marked with a fine, fasal line. Full-grown larvae
are grayish-green and about 20 mm. long. Larvae live together in
webs, some of which may be several inches wide and up to 114
feet long (fig. 149). Occasionally, they are large enough to en-
close 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.

The fruit-tree leaf roller, Archips argyrospilus (Wlkr.), 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 horse chestnut. The adult is pale yellow to
orange red and has a wingspread of 18 to 25 mm. The forewings

COURTESY CONN. AGR. EXPT. STA.

FIGURE 149.—Nest of larvae
of the oak webworm, Ar-
chips fervidanus.

377

are mottled with golden scales, their tips triangular. The hind-
Wings are fuscous and have dirty white fringes. Full-grown
larvae are light green and about 20 to 25 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 acres, have occurred in oak
stands in Eastern and Lake States in recent years.

Many other species of Archips are also encountered in eastern
forests (266). A few of these and some of their hosts are as fol-
lows: A. infumatanus (Zell.)—hickory and pecan; A. semiferanus
(Wlk.)—oak and witch-hazel; A. negundanus (Dyar)—boxelder ;
A. rileyanus (Grote)—hickory and walnut; A. georgianus (WIlk.)
—oak; A. griseus (Rob.)—oak and hickory; A. magnolianus
(Fern.)—yellow cucumber tree; and A. purpuranus (Clem.)—
basswood, paper birch, trembling aspen, willow, and black cherry.

The spruce budworm, Choristoneura fumiferana (Clemens)
(fig. 150), occurs in the region of North America extending from
Virginia to Labrador and westward to the MacKenzie River Val-
ley, Yukon Territory, Canada (688). Until recently, it had been
considered present throughout the ranges of spruce and fir in
North America. Forms occurring in western Canada and western
United States are now considered different species (269). Balsam
fir is the preferred host of the spruce budworm. To a lesser
degree, it also feeds on white, red, and black spruce, and on larch,
pine, and hemlock (47). |

Spruce budworm adults are smaller than the adults of other
spruce-fir feeding species. In series, they are mostly gray with a
few ochreous-gray specimens. The male has a wingspread of 21
to 26 mm; the female, 22 to 30 mm. Full-grown larvae are about
20 to 28 mm. long. The head is usually almost entirely dark
brown; the prothoracic shield is brownish-yellow with some dif-
fusion of brown pigment or is entirely dark brown; and the anal
shield is brownish-yellow. Pupae are light to reddish-brown ex-
cept 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 two to sixty eggs each, the eggs overlapping
like shingles on a roof. A large proportion of the masses are usu-
ally found on needles near the periphery of the crown. The eggs
hatch in about 10 days under normal conditions. After a period
of 1 to 2 days during which they are dispersed throughout the
tree and stand, the larvae spin hibernaculae in suitable sites and
molt to the second-instar. While the majority of hibernaculae 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

378

ss F-493456, 495235, 501800
FIGURE 150.—Spruce budworm, Choristoneura fumiferana: A, adult;
B, eggs; C, larvae on twig of balsam fir.

and drop down on silken threads. At this time, they may be
blown considerable distances by the wind, often into uninfested
stands.

379

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
some occurs 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 (5432).

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 needles on webbed branch
tips turn reddish-brown by mid-summer. In heavy, persistent in-
festations, all of the new foliage may be consumed (fig. 151) for
several successive years, and opening vegetative buds and de-
veloping shoots may be killed in their formative stages. Top
killing usually occurs after about 3 years of severe infestation,

F-492901

FIGURE 151.—Balsam fir defoliated
by the spruce budworm, Choris-
toneura fumiferana.

ee we ee ae

and tree mortality, after about five years. In sustained outbreaks,
nearly complete mortality of the merchantable volume of balsam
fir may occur by the 8th year (55). During widespread outbreaks,
the magnitude of these losses is great: It is estimated that 225
million cords of pulpwood were destroyed in an outbreak which
occurred in Quebec, New Brunswick, Maine, and northern Min-
nesota from 1910 to 1920.

Much has been learned about the factors or conditions condu-
cive to the development and decline of spruce budworm outbreaks.
For example, it was determined many years ago (700) that out-
breaks 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
(313). Much has also been learned about factors tending to hold
populations in check during intervals between outbreaks or which
assist in bringing them under control once they are underway
(543).

The spruce budworm has many natural enemies, including para-
sites and predators, 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 starva-
tion, has reduced the budworm population considerably (521). A
number of authors have evaluated the control effectiveness of
natural enemies in specific outbreaks (206, 207, 400, 483, 79, 80).

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; (38) loss of large larvae through starvation following
complete defoliation; and (4) loss of adults through dispersal.

A number of management practices have been suggested for
reducing the chances of spruce budworm outbreaks. These in-
clude: the utilization of balsam fir (700); the regulation of age
classes to prevent the occurrence of large areas of overmature fir
(310); and the favoring of less susceptible species, such as
spruce. The removal of overstory mature balsam fir was suggested
for preventing outbreaks (757, 496) ; management plans and risk-
rating systems for selective cutting were presented (497, 498,
758); and risk-rating systems for the Lake States were devised
(45, 306). The budworm and forest management in the Maritime
Provinces of Canada was discussed by Balch (23).

For further information on the spruce budworm, the reader is
referred to McKnight’s review of the literature (491) which
covers 370 citations on the species and the two related species,
Choristoneura occidentalis Freeman and C. biennis Freeman.

The jack pine budworm, Choristoneura pinus Freeman, is
known to occur in Nova Scotia, Ontario, Manitoba, and Michigan.
Its hosts include various species of pine, especially jack and red.
The adult has a wingspread of 15 to 28 mm. The head, thorax, and
forewing are ochreous-tawny; the forewings, distinctly mascu-
late; and the hindwings, entirely 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

381

the head by a narrow white band, a reddish brown bcady with
yellowish sides and two rows of white dots along the back, and it
is about 20 to 22 mm. long (fig. 152). Pupae are pale green when
just formed; later, they become dark reddish brown, and they
are about 12 mm. long. This species was considered to be a racial
form of the spruce budworm until it was described as a new
species in 1958 (265).

Jack pine budworm adults are present from early July to early
August and lay their eggs in clusters of about 40 eggs each.
Hatching occurs in about 10 days. A few days later, the young
larvae, without feeding, spin hibernaculae 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
remainder 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 killing and stag-
headedness. During outbreaks, however, heavy losses in poles,
saplings, and reproduction may result. In heavily infested stands,
young understory red and white pines also are often severely

F-506744

f FIGURE 1538.—Pupae of the large

. aspen tortrix, Choristoneura
conflictana, in webbed leaves.

F-501801

FIGURE 152.—Larva of the jack

pine budworm, Choristoneura

pinus, feeding on needles of jack
pine.

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 (359). These trees are usually coarsely branched and
large crowned, or suppressed and slow growing. The growing of
hard pines in fully stocked stands or in groups; the elimination
of large-crowned, ‘wolf’ trees; the utilization of trees before
they become mature; and the encouragement of species suited to
the site are also recommended (309). 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 new subspecies, Choristoneura 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 (269).

The large aspen tortrix, Choristoneura conflictana (Wlk.), oc-
curs throughout the range of trembling aspen in Canada and
eastern United States south to New Jersey and Ohio. Several
other hardwoods also serve as hosts such as balsam poplar, large-
tooth aspen, paper birch, willow, and alder. The adult is dull,
light gray in color and has a wingspread of 25 to 35 mm. Full-
grown larvae are usually dark green, sometimes almost black, and
are 12 to 21 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. First instar larvae feed gre-
gariously on leaf surfaces during July, spinning much silk and
webbing the surfaces together (fig. 1538). Later they move to the
trunk in search of hibernation sites in rough bark or under moss.
Here they molt and spend the winter in the second instar. 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 (606).
Many outbreaks have occurred in aspen stands in Canada. Serious
defoliation has also been recorded in New England, New York,
and Michigan.

The oblique-banded leaf roller, Choristoneura rosaceana
(Harr.), occurs in southern Canada 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 trembling aspen, paper birch, and willow but has also been
observed feeding on Scotch pine. In New York, it has seriously
damaged coniferous seedlings in nursery seedbeds (652). Seed-
lings most seriously affected were those of white, Scotch, and red
pines. The infestation is believed to have resulted from an inva-
sion of the nursery from larvae developing on weeds or other
vegetation surrounding the nursery. The adult is reddish-brown
and has a wingspread of 18 to 20 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,

383

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 (Clem.) feeds on sugar maple,
beech, paper birch, red maple, elm, and red oak in southern Can-
ada and from Massachusetts to Wisconsin and Colorado. It is
sometimes mistaken for C. rosaceana. C. houstonana (Grote) at-
tacks various species of Juniperus, especially red cedar, and is a
pest in windbreak and ornamental plantings in western Kansas
(OG.

The red-banded leaf roller, Argyrotaenia velutinana (Wlk.),
occurs in southern Canada, mostly in the southeastern part, and
throughout eastern United States, westward to Iowa, 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, it occurs on a wide variety of de-
ciduous trees. Damage is often serious in apple orchards. The
adult has a wingspread of 12 to 16 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 15 to 18 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
which tends to retard larval development has been reported in
Virginia. Eggs are sometimes heavily parasitized by Tvricho-
gramma minutum Riley.

The pine tube moth, Argyrotaenia pinatubana (Kft.), feeds on
various conifers, mostly white pine, in southern Canada, the
Northeastern States, Florida, and Louisiana. The adult is small,
slender, and grayish; it has a wingspread of 14 mm. The fore-
wings 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 och-
reous, apical tufts. Full-grown larvae are 10 to 14 mm. long.

The larva lives within the tube it constructs by drawing from
five to 20 needles together and fastening them with silk. It feeds
on the tips of these needles (fig. 154). 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 objectionable where valuable ornamentals are involved.

The hickory leaf roller, Argyrotaenia juglandana (Fern.), oc-
curs 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 18 to 25 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 16 to 24

384

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.

Several other species of Argyrotaenia are recorded as feeding
on eastern trees (266): A. occultana Freeman—on spruce and
occasionally balsam fir and larch (in Canada and New York) ;
A. mariana (Fern.)—on paper birch, willow, choke cherry, elm,
pear, Vaccinium, and possibly oak (from eastern Canada to
Florida); A. quadrifasciana (Fern.)—on hawthorn, shadbush,
plum, and pear (in southeastern Canada and from Maine to Mis-
souri) ; and A. alisellana (Rob.)—on oak (from southern Canada
to Florida).

Croesia albicomna (Clem.) [=semipurpurana (Kft.)] 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 a week or two later (51).

Serious outbreaks have occurred in the Northeastern and
Middle Atlantic States in recent years. During 1964 and 1965,
approximately a half million acres of red oak were severely de-
foliated in Pennsylvania alone, resulting in considerable tree
mortality.

The black-headed budworm, Acleris (=Peronea) variana
(Fern.), occurs from coast to coast in Canada and southward into
the United States. In eastern United States and Canada, balsam
fir is its preferred host, but during epidemics, white and some-
times red and black spruces and hemlock may also be defoliated
(475). 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 is about 19 mm. Full-grown larvae are bright
green and about 12 to 16 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 expanding
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.

In eastern America, extensive outbreaks tend to occur at in-
tervals of 10 to 15 years in stands where maturing balsam fir is
dominant, but they usually subside before many trees are killed.

Acleris chalybeana (Fern.) has been recorded from Ontario,
Quebec, Wisconsin, Maine, New York, and Pennsylvania. The

385

larvae feed on the foliage of sugar, red, and mountain maples,
yellow birch, beech, and ironwood. The adult is grayish and has a
wingspread of 21 mm. Full-grown larvae are light green and
from 17 to 21 mm. long. In Wisconsin, winter is apparently spent
as first or second instar larvae in hibernaculae on the twigs.
During June, the majority of them were found in rolled leaves
previously occupied by larvae of Sparganothis 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 to the development of maple blight, a condition lead-
ing to the deterioration and death of large quantities of sugar
maple trees and saplings during recent years (294). 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 (Hbn.) feeds on paper
birch.

Aphelia alleniana (Fern.) normally feeds on weeds and clovers,
but may also attack and injure small seedlings in coniferous plan-
tations. Seedling losses have been severe in the Lake States and
southern Canada. The larvae tie the shoots of the seedlings
togther 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 (Rob.) also normally feeds on weeds and
clover, but it also occasionally damages seedlings in young con-
iferous plantations in the Lake States and southern Canada. It is
widely distributed in eastern United States, occurring from New
England to the Lake States, Missouri, and Texas. The larvae draw
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 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
hibernaculae spun within folded leaves (503).

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 Massa-
chusetts. Its favored hosts are 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 also sparsely hairy and dotted with brown or black
tubercles.

586

Adults are present from late spring until fall. Eggs are de-
posited 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 (386).

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. 155) 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.

The pecan carpenterworm, Cossula magnifica (Strecker), occurs
throughout the Southern States. Its hosts are pecan, oak, and

COURTESY CONN. AGR. EXPT. STA.
FIGURE 155.—Elm branch nearly severed by
larvae of the leopard moth, Zeuzera pyrina.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 154.—Larval tube and lar-

val feeding damage by the pine

tube moth, Argyrotaenia pina-
tubana.

387

hickory. The adult is grayish with brown markings and has a
Wwingspread 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
several inches 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.

Acossus centerensis (Lint.) and A. populi (Wlk.), the aspen
carpenterworm, also occur in eastern United States. A. centerensis
bores in poplars from New Jersey to Illinois. A. populi is found
in poplars and cottonwoods from coast to coast.

The carpenterworm, Prionoxystus robiniae (Peck.) (fig. 156),
is widely distributed in the United States and southern Canada
where it breeds in various hardwoods. In the Eastern and

apt - Baer

Pas"
SS

* “i es
ete - AB. p ty? =.

F-498365

FIGURE 156.—Egegs, larva, pupal case, and adults
of the carpenter-worm, Prionoxystus robiniae.

388

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, cot-
tonwood, and occasionally fruit trees and ornamental shrubs
(327). The adult is dark, slightly mottled, stout-bodied, and has
orange and 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, are 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.

Carpenterworm moths begin to appear in late 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 upwards of
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, and feed there until about
half-grown. Then they bore into the wood, their tunnels angling
upward in the sapwood and straight upward in the heartwood.
Feeding may be finished in one year in the Deep South but may
continue for 8 or 4 years in the North. Tunnels are kept open and
enlarged as needed by the growing larvae; eventually they may
reach a diameter of 18 mm. and a length of 375 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. 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. Open-grown trees, or trees growing on
poor sites such as dry ridge tops or ridge slopes, are especially
subject to attack and damage.

Little is known about natural control of the carpenterworm. It
is believed, however, that predation by woodpeckers on larvae and
kingbirds on adults frequently accounts for the destruction of
fairly large numbers. Insect parasitism appears to be inconse-
quential. No satisfactory method of control in the forest is
known, although the removal or killing of heavily infested weak,
deformed, cull trees should, however, be helpful in reducing the
intensity of infestations. 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.

The little carpenterworm, Prionoxystus macmurtret (Guer.), a
species very similar to the carpenterworm, is widely distributed
in eastern Canada and eastern United States. The larvae are
borers in oaks, and their habits are similar to those of the car-
penterworm. They spend their first summer in the outer layers of
bark and the second in the sapwood. During the third summer,

389

yw

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 over 1 inch in diameter, and trees infested continuously be-
come badly honeycombed. Fortunately, infestations are usually
very local.

FAMILY COSMOPTERYGIDAE

Chrysoclista linneella (Clerck), an introduced European
species, was first reported in this country in 1928 when it was
found infesting linden trees near New York City. Other infesta-
tions were later found in surrounding parts of New York and
New Jersey, and near Boston, Massachusetts. Its current distri-
bution 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
aoe in cells formed in the galleries close to the surface of the

ark.

The palm leaf skeletonizer, Homaledra sabalella (Chamb.),
feeds on many varieties of palms, particularly Canary Island date
palm and sabal palmetto, in the Southern States. It is often quite
injurious in Florida. The larvae feed in groups of 35 to 100 larvae
each under webs of silk on both the upper and lower surfaces of
the leaf (173). 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 of all interleaf
husks and infested fronds is helpful in control.

FAMILY WALSHIIDAE

Aenea ostryaealla Chmb., a leaf miner 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 (458).

FAMILY GELECHIIDAE
Wax MOTHS

Wax 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 leaf miners; still others feed in buds, seeds and cones, and roots.
A few species are quite destructive.

Coleotechnites (—Recurvaria) apicitripunctella (Clem.) occurs
in Quebec and in the Northeastern States. Its known hosts are
hemlock and cypress. The adult is buff yellow to whitish and has

390

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. 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 (=—Recurvaria) thujaella (Kft.), occurs from
New Jersey to New Brunswick and westward to Saskatchewan,
in Canada. The larvae are leaf miners on arborvitae. 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 scale-like leaves on growing tips. The
larvae bore into the tips and mine along the twigs, causing the
foliage to turn brown. Winter is spent in the larval stage in a
mine, and there is one generation per year. Ornamental arbor-
vitae is subject to serious injury.

Coleotechnites (=Recurvaria) piceaella (Kft.) occurs com-
monly from Maine to Colorado and from the Maritime Provinces
to Alberta in Canada. Its hosts are white, Colorado blue, Engel-
mann, Norway, red, and black spruces. The adult is light gray
and has a wingspread of 10 to 18 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’s needles,
in insect-damaged or mechanically-damaged needles, in insect- |
damaged cones, or in spent staminate flowers. Some also may be
deposited 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 (494). 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 (Kft.) which mines the needles of red cedar and
common juniper in the Northeastern States; C. dorsovittella
(Zell.) which feeds on sweetgum; and C. variella (Chambers)
which feeds on cypress. Heavy infestations of the latter species
have killed the top 2 or 3 feet of cypresses up to 20 feet tall in
Ohio.

The pine needle miner, EH'xoteleia pinifoliella (Chambers), oc-
curs in southeastern Canada and south to Georgia and Texas.

391

P

Jack, pitch, and shortleaf pines are preferred hosts, but it has
also been observed feeding on Virginia, Scotch, scrub, 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 (248). Forest-grown trees are
occasionally infested heavily, but are seldom injured seriously. On
the other hand, heavy infestations on ornamentals or in planta-
tions may be serious. The mite, Pediculoides ventricosus, appears
to be an effective predator in some areas. It is reported to have
destroyed over 75 percent of late instar larvae in infestations in
North Carolina (40).

Exotelleia dodecella (L.), the pine bud moth, was discovered in
the Niagara Peninsula, Ontario, in 1928 and now occurs through-
out southern Ontario. So far as known, it has not yet crossed the
border into the United States. Its preferred hosts appear to be
Scotch and mugho pines, but several other pines such as eastern
white, red, jack, and Austrian may be attacked in heavily infested
areas. Old Scotch pines, particularly those growing along road-
sides, are highly subject to attack. Infestations on such open-
grown trees may persist for years. Heavy infestations cause a
thickening or browning of the needles, followed by branch mal-
formation. Eventually, the needles thin out. A high proportion of
the buds on young trees may be killed.

Eggs are laid on current-year and 11-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 (504).

Exoteleia chillcotti Freeman is a common needle miner of long-
leaf pine in Louisiana. Bennett (60) discussed the morphology of
the pupa.

Exoteleia 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, Mugho 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 either re-enter the same needles, adjacent needles, or
swelling buds. Larval growth is completed in May, and pupation

392

occurs in the flowers or shoots of the tree. There is one generation
per year (459).

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 10 to 25 feet tall.

Anacampsis nnocuella (Zeller) occurs in southern Canada and
from Massachusetts to Colorado and Texas. The larva is a leaf-
roller 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 leaf rolling begin in Ohio about the time
the leaves of bigtooth aspen begin to develop. Prior to 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 is believed that winter is
spent in the egg stage on twigs of the host (523).

Anacampsis rhoifructella (Clem.) feeds on Viburnum and
sumac in the Northeastern States. The adult is grayish-brown
and has a wingspread of 15 mm. Markings on the forewings re-
semble 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 ligulellus (Hbn.), occurs in Can-
ada 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 greenish 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.

Eggs are laid on the undersides of leaves in early spring. The
larvae feed on and skeletonize the leaves, either in exposed posi-
tions or within folded or rolled leaves. Pupation occurs either in
these rolls or in ground litter. Adults appear during July or
August and apparently 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, Dichomeris marginellus D. & S., an in-
troduced 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 red cedar. 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 needle miners, entering the upper surface of the needle near
the axil. Mined needles turn brown and die. Dead needles are in-
corporated in webbing constructed between branchlets. Larvae
continue to use mined needles as protective retreats while de-
veloping. During July, larvae crawl] from primary mined needles
and feed on adjacent needles, eventually constructing silken tubes

393

ee

from holes in needles to other nearby needles. Webbing area is
expanded from July to September. Entire trees up to 8 feet tall
may be completely webbed (fig. 157). 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
(569). Trees grown as ornamentals or in permanent plantings
may be seriously damaged by this species. The cutting out and
removal of webbed masses of foliage is a helpful control practice.

Fascista cercerisella (Chambers) feeds on redbud from Dela-
ware 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 (Buseck) is widely distributed throughout
eastern United States and southern Canada. It has been reared
from the buds of mugho pine; from the cones of Scotch, Austrian,

wa F-519583

<= FIGURE 157.—Defoliation by the
juniper webworm, Dichomeris
marginellus.

394

and longleaf pines; and from cecidomyid galls 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.

FAMILY OEKCOPHORIDAE

This family contains a large number of moderately small
moths, only a few are ever very injurious to trees in eastern
United States. The larvae of most species either roll, tie, or web
together the leaves on which they feed. Gates-Clarke (282) re-
vised the family several years ago.

Machimia tentoriferella Clem. occurs rather commonly in the
Northeastern States and southeastern Canada. Its hosts are
various hardwoods such as birch, ash, maple, oak, honey locust,
mountain ash, hickory, elm, and wild 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 faginella (Chmb.) occurs in eastern Canada and the
Northeastern States. The larva feeds principally as a leaf tier on
American beech. Other hosts include the birches, maple, and red
oak. Embree discussed its biology in Nova Scotia (226).

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 larva 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. Psilocorsis reflexella Clem., P. obsoletella
(Zell.), and P. quercicella Clem. occur on oaks in the Eastern
States. P. fletcherella Gibson has been observed feeding on tremb-
ling aspen in eastern Canada. It probably occurs in the North-
eastern States also.

Other tree-infesting species of oecophorids in eastern United
States are as follows: Agonopterix pteleae Barnes & Busck—on
the common hop tree; A. robiniella (Pack.)—on black locust; A.
argillaceae (Wlsm.)—on willow; A. nigrinotella (Busck)—on
prickly ash; Bibarrambla allenella (Wls.)—on white birch and
oaks; and Depressaria betulella (Bsk.)—on white birch and
hophornbeam.

FAMILY BLASTOBASIDAE

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
either in the cones, nuts, and seeds of various trees; as scavengers

395

ee ee)

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.
ayes are grayish-white or yellowish, with blackish marks on
Op.

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 wing-
spread of 11 to 17 mm. The head is ochreous-white, with slight
infuscation posteriorly; the antennae are ochreous-white with
narrow, dark annulations; and the forewings are ochreous-buft.
The larvae of Zenodochium coccivorella Chambers are internal
parasites of female scale insects of the genus Kermes in Florida.

FAMILY GLYPHIPTERYGIDAE

The apple-and-thorn skeletonizer, Anthophila 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, paie 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 Sep-
tember. 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 generations per year.

The mimosa webworm, Homadula anisocentra Merrick (=al-
bizzae Clark), 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, Ala-
bama, and Mississippi, and westward to Kansas and Nebraska.
Its hosts are mimosa and honeylocust trees. The adult has a wing-
spread 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 in color, are marked with five longi-
tudinal white lines, and are about 12 mm. long. Pupae are
yellowish-brown and about 6 mm. long. They are found in whitish,
silken cocoons.

Winter is spent in the pupal stage in the soil or in other pro-
tected 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

396

second instar larvae spin webs around flowers and leaves, within
which they live and feed (fig. 158). Adjacent surfaces of webbed
leaves may be skeletonized, turn brown, and die. Older larvae feed
on the tender, terminal leaves. Pupae of this generation 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 (756).

The mimosa webworm is a serious defoliator of ornamental
plantings of mimosa and honey locust. The thornless variety of
honey locust is heavily attacked in nurseries. Season-long protec-
tion against attack has been obtained by the use of systematic
insecticides (198).

FAMILY AEGERIIDAE
CLEARWING MOTHS

Members of this family are known as clearwing moths because
the greater parts of one or both pairs of wings are without scales,
thus leaving them clear or transparent. The forewings are long
and narrow, with the outer margins short and the anal veins
reduced. The hindwings are somewhat broader than the fore-
wings, 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 and shrubs, and vines, or in the stems
and roots of herbaceous plants. A few form galls, others are in-
quilines in galls, and some inhabit injured areas on the trunks or
branches of their hosts. A number of species are important pests

COURTESY OF ILL. NAT. HIST. SURV.

FIGURE 158.—Webs formed
by the mimosa webworm,
Homadula anisocentra.

of forest and shade trees and ornamentals. The family has been
treated by Engelhardt (2303) and MacKay (489).

The hornet moth, Aegeria apiformis (Clerck), an introduced
species first observed in North America around 1880, is now
known to occur in Massachusetts, Connecticut, New York, New
Jersey, Pennsylvania, and California. Its 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.

Aegeria tibialis (Harr.) attacks poplar and willow from New
York to Nova Scotia and in the Midwestern States. Adults are
distinguished by their black abdominal segments, all but the
second and fourth of which have narrow posterior yellow margins.
Full-grown larvae are 40 mm. or more long. Infestations are lo-
cated well down on the trunk or in the roots.

The dogwood borer, Synanthedon (=Thamnosphecia) scitula
(Harris), occurs in southeastern Canada 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, and 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 Andricus cornigerus on black
and pin oaks, sometimes occur in the thousands on a single tree,
and nearly every one will be infested by this borer (230).

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 wing
spread 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 mid-summer 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
2 feet or more in length and may contain up to 50 larvae each on
the larger trees. A single larva can kill a dogwood 4 inches in
diameter in one year. There are one and possibly two generations
per year, depending on locality. Keeping the bark smooth, es-
pecially at the base of branches has been recommended for control
on dogwood (737).

The rhododendron borer, Synanthedon (=Ramosia) rhodo-
dendri Beutenmuller, attacks rhododendron and sometimes laurel
and azalea in the Atlantic Coastal States. The adult is blackish,
except for various white and yellow markings, and has a wing-

598

|

spread of 10 to 15 mm. Mature larvae are up to 15 mm. long. The
winter is 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 of infested
parts of plants is a recommended control practice.

Several other species of Synanthedon (=Thamnosphecia) also
occur in eastern America. S. geliformis (Wlk.) is a bark borer in
dogwood and pecan. It also infests Andricus galls on oaks and
diseased and injured tissues of various other plants. S. pyri
(Harr.) attacks apple, hawthorn, shadbush, and mountain-ash.
The larvae excavate shallow, tortuous tunnels in and beneath the
bark of trees usually injured by storms and disease, causing the
bark to blister and peel. S. pictipes (Grote & Robinson) breeds in
peach, wild plum, and in fungus growths on wild plum. It is a
major pest of peach. S. viburni Eng. larvae bore under the bark
of Viburnum, preferably in injured areas or galls. S. acerrubri 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. castanea (Beutenmuller) bores in the trunks of
American chestnut, preferably in bruised areas.

Thamnosphecia rubrofascia (Hy. Edw.) larvae excavate long,
sinuous tunnels in the bark of sour gum from Maryland to
Florida. Injuries and healing wounds on well matured trees are
preferred. T. sigmoidea (Beut.) attacks low-growing willows in
bays, along streams, and in depressions in sand dunes of coastal
and lake regions. Infestations are often heavy.

The maple callus borer, Sylvora (=Conopia) acerni (Clem.), is
widely distributed in southern Canada and throughout 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 in to 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 of
roughened bark areas, removal of borers from under the bark in
the spring, and painting of wounds are recommended control
practices. The southern form, S. a. buscki Eng., attacks silver
maple in Florida and Georgia.

The pitch mass borer, Vespamima (=Parkarmonia) pini (Kel-
licott), occurs in eastern Canada and eastern United States south-
ward to Georgia and Tennessee and westward through the Great
Lakes region. Its preferred host appears to be white pine but it
also attacks Austrian and Scotch pines and spruce. The forewings
are blue-black, with a metallic green luster, and have an expanse

399

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.

Kiggs 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 3 to 4 inches 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.

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 tap roots 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 cut over areas and in hedge rows are particularly sub-
ject to infestation. Young trees in nurseries are also damaged
seriously at times. The life cycle requires 2 or 8 years.

The lilac borer, Podosesia syringiae syringiae (Harris) (fig.
159), occurs throughout the eastern half of the United States and
Canada. It attacks lilac and privet and other trees and shrubs of
Oleaceae. The adult has a wingspread of 26 to 38 mm. Southern
adults have the thorax and abdominal segments two and three
almest entirely chestnut-red, only narrowly edged with black.
Northern ones have these parts mostly brown-black with edgings
of yellow or chestnut-red. The larvae bore into the main stems of
lilac and privet, causing them to wilt, take on an unhealthy ap-
pearance, or break. Winter is spent in the larval stage, and there
is one generation per year. Borers in lilacs can be killed by
inserting a flexible wire or injecting a fumigant into their
burrows.

The ash borer, Podosesia syringiae fraxini (Lugger) (fig. 160),
occurs from Texas north through the Midwestern States to Mani-
toba and Saskatchewan. It attacks green, white, red, and Euro-

COURTESY CONN. AGR. EXPT. STA.
FIGURE 159.—Adult, pupal
case, and larval tunnel of
the lilac borer, Podosesia
syringiae syringiae.

400

COURTESY OF DUKE UNIV. SCH. OF FOREST.
FIGURE 160.—Adult and larval galleries of the
ash borer, Podosesia syringiae fraxini.

pean ash. The adult is blackish and has a wingspread of 28 to 34
mm. The forewing is opaque with hyaline streaks between the
veins to the wing base, and the fringe is dark brown. The hind-
wing is transparent, with yellowish-brown veins and a sordid
brown fringe darkening toward the base. The larvae tunnel in
branches and trunks, excavating galleries several inches long.
Winter is spent as a larva in the tunnel. Young shade trees and
trees growing in shelterbelts may be seriously damaged. Control
is difficult, but the cutting and burning of infested trees should
be helpful.

Paranthrene palmu (Hy. Edw.) (fig. 161) occurs from the
New York area to Florida and along the Gulf Coast at least to
Mississippi. Its hosts are various species of oaks. In Mississippi,
it shows a preference for red oaks, especially Nuttail and cherry-
bark oaks (680). Eggs are deposited in bark crevices at the base
of the tree, preferably in trees over 12 inches d.b.h. The larvae

F-519923-22 “%
FIGURE 161.—Adult and larva
of Paranthrene palmii.

bore through the bark and into the wood. Here they excavate
relatively uniform galleries up to 9 mm. in diameter and 4 inches
long. The first inch or two the gallery slopes upward, then it con-
tinues straight up. The life cycle in Mississippi requires 2 years.
The major hazard associated with the work of this borer is the
possibility that rot and stain producing fungi will enter the larval
galleries and degrade the butt logs.

Paranthrene simulans simulans (Grote) occurs from Nova
Scotia to Maryland, Illinois and the Lake States. It has also been
recorded from black, red, and pin oaks. The larvae bore into solid
wood, constructing galleries up to 2 inches long. It, or a closely
related species, has also been found infesting natural oak growth
and 2- to 5-inch nursery stock in Long Island. The larvae, gener-
ally one to four per tree, bore into the wood of the trunk from
1- to 2-feet above the ground and then tunnel upwards 2- to
3-inches in the center of the trunk.

Paranthrene dollu dollii (Newm.) 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 cross-bands
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, con-
structing open tunnels up to 6 inches long. Heavily infested trees
are weakened and subject to wind breakage. Damage is often
severe in cottonwood nurseries and plantations in Mississippi
where there are two generations per year. The closely related
species, P. d. castanea (Beut.) also occurs in the Deep South. Its
hosts, habits, and damage are much the same as those of P. dollu
dollii.

Paranthrene tricincta (Harris) larvae bore in the branches and
terminals of young cottonwoods in the Deep South, weakening
them and causing some to break. There are two generations per
year. P. tabaniformis (Rott.), an introduced species with similar
hosts, habits, and life cycle, occurs in the Northern States.

Conopia proxima (Hy. Edw.) larvae bore in the cones,
branches, and exposed roots of low-growing willows in moist or
shady locations. C. bolteri (Hy. Edw.) also attacks low-growing
willows. C. acerrubri (Engelhardt) attacks various poplars. The
larvae feed under the bark and cause slight swellings. Local out-
breaks have been recorded.

FAMILY YPONOMEUTIDAE

Moths of this family are small, usually brightly patterned, and
have rather broad wings.

The genus Argyresthia contains a number of leaf mining
species. The adults have wingspreads of about 25 mm. While at
rest, their wings are folded close to the body, their front legs are
extended forward, and their hind legs are slanted upwards at 45°
angles.

Argyresthia freyella Wlshm. is generally distributed in eastern
Canada and south and west to the Middle Atlantic States and
Missouri. Its hosts are red cedar and arborvitae. 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

402

of each. The larvae feed in mines which 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 cocoons spun on
leaves or branches in the spring. Damage to ornamentals and
nursery seedlings appears to be less than that caused by the re-
lated species, A. thuiella. A. auveoargentella Brower also attacks
arborvitae. It resembles A. freyella except that its cocoons are
whiter and larger.

The arborvitae leaf miner, Argyresthia thuiella (Pack.), feeds
on arborvitae throughout much of the same areas as that occu-
pied 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 branch-
lets or along the edges of leaves. Newly-hatched larvae bore into
th 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 occurs in the spring. Outbreaks in Maine have
severely damaged forest stands of arborvitae. Damage to orna-
mentals and nursery seedlings is often serious. Heavily defoliated
trees may be killed (107). Silver (647) discussed arborvitae leaf
miners in New Brunswick.

Larvae of the apple fruit moth, Argyresthia conjugella Zell.,
bore in the fruit of mountain ash, shadbush, Crataegus, 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 Kft.
larvae bore in the shoots of larch in southern Canada and the
Lake States.

The ailanthus webworm, Atteva aurea (Fitch), feeds on the fo-
liage 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. This species is often common in the South. Zelleria
haimbachi Busck feeds rather commonly on jack pine in southern
Ontario. It may occur in the Lake States.

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 leaf miner, Antispila nyssaefoliella Clem., feeds on
tupelo throughout the eastern part of the United States. The adult
is dark brown and has a wingspread of 7 to 8 mm. The pale green

403

larvae feed within leaf tissues, forming blotch-like mines. A full-
grown larva spins a cocoon within the mine and then cuts 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 mid-summer.

Coptodisca splendoriferella (Clem.) also occurs in eastern
United States. Larvae mine the leaves of apple, cherry, and re-
lated hosts. They pupate in cases attached to the limbs and trunks
of their hosts.

FAMILY COLEOPHORIDAE
CASEBEARER MOTHS

Nearly 100 species of casebearer moths occur in North Amer-
ica. The adults are plain, little moths with markings limited to
dustings of lighter or darker colored scales. The wings are nar-
row and have an expanse 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 constructed 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 se-
curely to a twig or branch with silk.

Coleophora tilaefoliella Clem. constructs a black case on bass-
wood. C. atromarginata Braun occurs on southern red and
swamp white oaks. Its case is black with white markings on the
underside. C. querciella Clem. occurs on white and swamp chest-
nut oaks. Its case is grayish with a black patch on the top near
the rear end. C. alniella Hein. has been observed on beech in New
York. C. ulmifoliella McD. feeds on various species of elm in
southern Ontario and possibly occurs in the Lake States.

The pecan cigar casebearer, Coleophora caryaefoliella Clem., oc-
curs from New England 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 (295).

The elm casebearer, Coleophora limosipennella (Dup.), an in-
troduced 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.

Adults appear in late July and lay their eggs on the leaves of
their hosts. The larvae feed within mines in the leaves at first.

404

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 be-
tween the larger veins as far as it can reach. The case is en-
larged as needed to accommodate the growing larva, eventually
reaching a length of 9 to 10 mm. This species is primarily a pest
of shade and ornamental trees. The mined parts of laves turn
brown and, where numerous, are unattractive. Fortunately, out-
breaks are usually limited.

The larch casebearer, Coleophora laricella (Hbn)., an intro-
duced species, was first recorded in North America in Massachu-
setts in 1886. It now occurs throughout most of the range of
tamarack, Larix laricina, in North America, west to central
Minnesota and northwestern Ontario. In 1957, an infestation was
also discovered on western larch, L. occidentalis, in Idaho (191).
The adult is silvery to grayish-brown, has narrow wings fringed
with long hairs, and a wingspread of about 9 mm. Full-grown
larvae are about 6 mm. long. The pupa is colored brown. This
species is one of the most serious defoliators of larch in North
America. Outbreaks have been occurring at about 8 year inter-
vals in eastern Canada. One was also recorded in the Lake States
in the late 1940’s and early 1950’s. Heaviest losses result from
reduced growth. Trees completely defoliated for 2 or more years
in a row 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 re-
maining 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 are particularly
effective in control. Two introduced hymenopterous parasites,
Agathis pumilus (Ratz.) and E'pilampsis laricinellae (Ratz.) are
now widely distributed and are believed to be quite helpful in
control.

The birch casebearer, Coleophora fuscedinella (Zell.) (=salmant
Heinr), an introduced species first observed in North America
in Maine in 1927, is now known to occur throughout the North-
eastern States and from Newfoundland and New Brunswick to
southern Ontario, Canada. Its favored hosts are paper, gray, and
European white birches. Eggs are laid along the mid-ribs and
larger veins on the undersides of leaves in July. Young larvae
enter the leaves and feed as miners for several days; then they

405

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. During recent years, this
species has been abundant in Quebec, New Brunswick and north-
ern Maine. Many stands of white birch in New Brunswick were
completely defoliated in 1968.

The cigar casebearer, Coleophora serratella (L.) (=fletcherella
Fern.), is principally a pest of apple in the Northern States, but
it also attacks cherry, hawthorn, plum, quince, and pear. It is
most injurious in the spring when the larvae feed on expanding
foliage, flowers, fruit, and fruit stems. The adult is dark gray to
grayish-brown and has a wingspread of about 12 mm. The larval
case is brown, shaped like a cigar, and about 8 mm. long. The
ae cycle and habits are similar to those of the pecan cigar case-

earer.

FAMILY GRACILLARIIDAE
LEAF MINERS

The family Gracillariidae, the largest of the leaf-mining fami-
lies of Lepitoptera, is represented in North America by more than
200 species (225, 559). The adults are tiny and beautifully ar-
rayed in shining scales and plumes, and their more or less lanceo-
late wings are overlaid with glistening scales of silver or bur-
nished gold. While an adult is at rest, the front part of the body
is raised and the wing tips touch the surface on which it sits.
Early instar larvae 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, usu-
ally within feeding mines or shelters, in which to pupate. Winter
is spent either as larvae, pupae, or adults, depending on the
species.

The solitary oak leaf miner, Lithocolletis (=Cameraria) hama-
dryadella (Clem.), occurs throughout much of 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 ones are cylindrical and about 5 mm. long. The larvae
feed singly, forming irregular blotch-like mines just below the
upper leaf surface. A single leaf may contains several contiguous
mines (fig. 162). 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 leaf miner, Lithocolletis (—Cameraria) cin-
cinnatiella (Chamb.), occurs throughout the same range as that
of the solitary oak leaf miner, and it feeds on the same hosts. The

406

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 and feed together, 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 Lithocolletis attack a wide variety of
deciduous trees in eastern America. L. salicifoliella Chambers
occurs in Canada and probably throughout most of the United
States. Its preferred host appears to be trembling aspen. Martin
(502) discussed its biology in Canada. L. crataegella Clem. mines
the leaves of apple, hawthorn, wild cherry, plum, and quince.
Heavily infested trees become ragged and scorched in appearance.
L. lucetiella Clem. mines the leaves of basswood. The mine is
nearly square. L. hamameliella Buseck mines the upper surface
of witch hazel leaves. It probably occurs wherever witch hazel
grows. The mines are circular or somewhat irregular. L. bethu-
nella Chamb. mines the leaves of oak in Delaware. L. trinotella
Braun mines the undersides of the leaves of red and Norway
maples; L. aceriella Clem. produces large white mines in the
upper sides of red and sugar maple leaves. L. robiniella Clem.
produces digitate mines in the upper surfaces of black locust
leaves. L. ostensackenella Fitch forms blotch mines in black locust
leaves. The aspen blotch miner, L. tremuloidiella Braun, con-

F-504082

FIGURE 162.—Oak leaves with the blotchy mines characteristic of attack by
the solitary oak leaf miner, Lithocolletis hamadryadella.

407

structs oval mines in the leaves of aspen. It is occasionally quite
abundant in the Lake States and Maine. L. corylisella Chamb.
mines the leaves of hazel in Maine.

Marmara fasciella Chamb. is widely distributed, probably oc-
curring throughout the range of its host, 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 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 (258).

Phyllocnistis populiella Chamb., the aspen leaf miner, is widely
distributed in southern Canada and the Northern States. It feeds
on various species of poplar, especially trembling aspen. In the
Eastern States, it has been recorded as far south as West Vir-
ginia. Eggs are laid near the tips of young leaves in the spring.
The larvae form tortuous mines in the leaves (fig. 163), mostly
on one side of the midrib (154). Noticeable infestations have been
observed in Maine.

Phyllocnistis liriodendrella Clem. larvae mine the undersides of
young leaves of yellow poplar and P. magnoliella Chamb., 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 Chamb. mines the leaves
of sweet gum.

The larvae of Parectopa robiniella Clem. feed in the leaves of
black locust, forming triangular, blister-like mines.

Callisto geminatella (Pack.), the unspotted leaf miner, occurs
throughout the Northern States, west and south to Colorado, Vir-
ginia, and Arkansas. Its hosts are apple, quince, pear, sour cherry,
wild 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.

Gracillaria negundella Chamb. occurs throughout eastern
America west to the Prairie Provinces of Canada and feeds on
boxelder. Early instar larvae 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

F-506745

FIGURE 163.—Small - track

mines of Phyllocnistis pop-

uliella, the aspen leaf
miner.

408

per year as far north as Canada. Heavily infested trees may be
seriously damaged.

The lilac leaf miner, Gracillaria syringella (F.), an introduced
species, occurs in many of the Northeastern States and south-
eastern 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 ir-
regular transverse patches of yellow, and the hindwings are
grayish-brown. Full-grown larvae are pale yellowish, translucent,
and about 8 mm. long. Early instar larvae feed gregariously in-
side the leaves and produce blotch-like mines. Later, they abandon
their mines and roll or web several leaves together which they
feed on from within this shelter. There are two generations per
year. Injured portions of infested leaves dry up and become un-
sightly. Raking and destruction of fallen leaves is a good control
practice.

Gracillaria sassafrasella Chamb., the sassafras leaf miner, 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, capsule-like cases on the undersides of
other leaves in which to pupate. G. azaleella Brants. larvae bend
the tips of azalea leaves downward and feed from inside the folds.
Injured leaves wilt and die. G. packardella Chamb. 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 G. bimacula-
talla Ely on maple; G. purpuriella Chamb. on poplar and willow;
G. quercinigrella Ely on oak; and G. pulchella Chamb. on yellow
birch and alder.

FAMILY LYONETIIDAE

This family contains numerous species of tiny moths, the ma-
jority of which belong to the genus Bucculatrix Zeller. At least
18 species are known to feed on trees in eastern America (97).

Adults of the genus Bucculatrix have the vertex of the head
rough or tufted, and the face smooth and produced into a point
extending 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 Clem., the apple bucculatrix, is widely
distributed in southern Canada and eastern United States. Its
hosts are apple, black cherry, shadbush, 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 shadbush, reddish-brown
on black cherry, and dark brown on hawthorn. There is one gen-
eration per year in the North and two in the South.

The birch skeletonizer, Bucculatrix canadensisella Chambers, is
a common species in southern Canada and throughout the birch-

409

growing regions of the eastern States from Maine to North Caro-
lina 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 hair. Full-grown larvae are yel-
lowish-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 two weeks and the young larvae enter the leaves to feed,
forming serpentine mines. Three or four 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 skele-
tonizers 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 intevals, often over large areas,
and last for 2 or 8 years. Defoliation may be severe but it seldom
causes much tree mortality. The defoliated trees may be so seri-
ously weakened, however, that they are attacked and killed by the
bronze birch borer.

The oak skeletonizer, Bucculatrix ainslhiella Murt., 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 7 to 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-instar larvae enter the
leaves to feed, forming serpentine or blotch mines. Older larvae
feed externally on the lower surface, often completely skeletoniz-
ing the leaves (fig. 164) (287). Two generations per year have
been recorded in Michigan and Massachusetts. Outbreaks occa-
sionally occur over large areas.

Other widely distributed species of Bucculatrix and their hosts
are as follows: B. packardella Chamb.—various oaks and occa-
sionally beech; B. quinquenotella Chamb.—oaks; B. luteella
Chamb.—white oak; B. recognita Braun—various white oaks,
preferably bur, and B. coronatella Clem.—black birch.

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

410

F-519521
FIGURE 164.—Red oak leaves
skeletonized by larvae of
the oak skeletonizer, Buc-
culatrix ainsliella.

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 en-
larging its case. There is also a smaller opening in the bottom end
of the bag, through which excrement drops out. About 24 species
have been recorded from eastern United States (185). Jones and
Parks (406) discussed the species occurring in Texas.

Oiketicus abbotwu 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 cypress, live oak, bayberry, syca-
more, elm, hackberry, sweetgum, and willow. It is noted for the
rather large bag, from 69 to 70 mm. long, which the larva con-
structs. Small twigs used in its construction are placed in a cir-
cular pattern around it. Fumaria casta (Fall.) feeds on lichens,
mosses, and the beech scale, Cryptococcus fagi, in Massachusetts.
The larvae occasionally climb up on the sides of houses in such
large numbers that they are a nuisance. PBasicladus 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 (Haw.), is
widely distributed in eastern United States, and attacks a wide
variety of trees. Arborvitae and red cedar appear to be preferred,
but many other conifers and hardwoods such as pine, spruce,
black locust, sycamore, willow, maple, elm, basswood, poplar, oak,
cypress, and persimmon are also attacked. The male moth is sooty
black, densely hairy, and has a wingspread of about 25 mm. Fe-
males are wingless, have no functional legs, eyes, or antennae,
and are almost maggot-like in appearance. The body is soft, yel-
lowish-white, and practically naked except for a circle of woolly

41

hairs at the posterior end of the abdomen (284). Full-grown lar-
vae 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 either in or outside the pupal case. 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. 165) 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 arborvitae and other conifers are lost each
year as a result of complete defoliation. Many others only partly
defoliated are weakened and rendered unsightly. Damage to forest
trees is usually not very severe (444); however, heavy infesta-
tions do occur occasionally in stands of cedar and black locust,
especially in the Deep South.

Low winter temperatures, bird predation, and parasitism by
the ichneumonid, /toplectis conquisitor (Say), are often particu-
larly effective in bagworm control. Handpicking and burning of
overwintering bags is also helpful in control, especially on small
trees and ornamentals ( 790).

F-508522
FIGURE 165.—Larval cases of the bagworm, Thyridopteryx ephemeraeformis,
attached to the twigs of an infested tree.

412

Cryptothelea gloverii (Packard) occurs from South Carolina
to Florida and westward along the Gulf Coast. Its hosts are re-
corded as two species of scale insects, and persimmon, oak, hick-
ory, Acacia sp., Crataegus sp., and Citrus aurantium. 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.

FAMILY NEPTICULIDAE

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 leaf miners;
a few construct mines in the bark of their hosts. Many of the
more common deciduous trees in 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 (96).

The genus Nepticula Von Heyden contains many leaf mining
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 either side; and some mine different
sides at different periods of larval life. The mines may be linear
and gradully 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. Heavy outbreaks of N. sericopeza Zell., 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 prematurely. There are two and perhaps three genera-
tions per year.

The genus E'ctodemia Busck 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) forms pea-sized globu-
lar galls on the petioles of poplar leaves. EF’. heinrichi Busck larvae
excavate flattened oval, spiral mines in the bark of young branches
of pin oak.

Obrussa ochrefasciella Chamb., the hard maple bud borer, 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. Kulman (446) discussed the biology and importance of the
species.

FAMILY INCURVARIIDAE

The maple leaf cutter, 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

413

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 814 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 hairs. There is a dense tuft of bright orange-yellow hairs
on the top of the head, the thorax is steel blue, the legs whitish,
and the abdomen dark brown. First instar larvae are flattened,
about 114 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 yellow-ish 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 under-
sides 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 dise 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 14 inch 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 gener-
ation per year (620).

The maple leaf cutter 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 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 wingspreads 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 5 pairs of prolegs, and normally live as root borers. Sthen-
opis argenteomaculatus (Harr.) breeds in the base of the trunk
and roots of alder in the Northeastern States; S. thule (Stkr.)
breeds in the roots of willow.

FAMILY ERIOCRANIIDAE

Larvae of the very small species, Mnemonica auricyonia
(Wlsm.), mine the leaves of oak, chestnut, and chinquapin in the
Eastern States. The mines are often blotch-like and may encom-
pass up to one-fourth of the leaf area. Damage is sometimes a
matter of concern to owners of Asiatic chestnut orchards.

414

Order DIPTERA
Flies

The Diptera constitute 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, stone flies, and dragonflies,
by the fact that they possess only one pair of wings, the fore-
wings. 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 im-
mobile 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 importance. Blood-sucking forms such as the mosquitoes,
black flies, punkies, horse flies, and others are serious pests of
man and animals. Some of these as well as some of the scavanger
species, such as the house fly, are important vectors of the causa-
tive 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 para-
sites of some of our most destructive tree-defoliating insects, espe-
cially of the order Lepidoptera. A few aquatic species, some of
which are nuisance pests in the adult stage are economically im-
portant as fish food in the larval stage.

Several comprehensive treatments of the order Diptera have
been published. Curran’s revision (181) of Williston’s Manual
(769) is especially useful in the classification of adults. Brues,
Melander and Carpenter (114) published a key to families. Stone, |
et al (695) issued a catalog of the species occurring in North
America north of Mexico.

Family CULICIDAE
MOSQUITOES

Mosquitoes are important pests of man and animals. Not only
are their bites extremely annoying, but they transmit many of the
most serious diseases of man and animals, such as malaria,
dengue, yellow fever, and encephalitis. As nuisances, they often
seriously interfere with public enjoyment of parks, vacation sites,
and other recreational areas. Woods workers, fishermen, hunters,
vacationers, and hikers are also hounded by them. The health of
domestic animals and wildlife suffers with serious losses of
weight and disease.

A total of 22 genera and 117 species of true mosquitoes occur in
the United States and Canada. (693). The larvae and pupae live
in standing or slowly moving bodies of water that range in size

415

from small accumulations held by plants to vast salt marshes.
Many of the most important species belong to the genera Ano-
pheles, Aedes, Culex, and Psorophora. King, et al. (417) discussed
the mosquitoes of the Southeastern States.

FAMILY CERATOPOGONIDAE
SANDFLIES, PUNKIES, ‘‘NO-SEE-UMS”

This family contains a number of species of very tiny flies,
usually 1 to 4 mm. in length, that feed on the blood of man and
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, fishermen, picnickers, and
others frequenting these areas often find their presence almost
intolerable because of their very burning and painful bites. Popu-
lations are heaviest during late summer. At this time, these flies
bite chiefly in the evening and very early in the morning. Wirth
(786) records 27 genera and some 300 species as occurring in
America north of Mexico.

FAMILY CHIRONOMIDAE
MIDGES

Midges are small, slender flies, rarely over 10 mm. in length.
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 fresh water fishes and other aquatic
animals. Sublette and Sublette (696) published a list of the spe-
cles occurring in America north of Mexico.

FAMILY SIMULIDAE
BLACK FLIES

Black flies occur in nearly all parts of the United States and
Canada. In different parts of their range they have various com-
mon names such as buffalo gnats and turkey gnats, as well as
black flies. The females feed on the blood of man and animals and
their bites cause swellings, 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
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
vild and domestic birds.

416

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 are
usually encountered in large numbers not too far from streams.
Sometimes, however, the wind blows them a considerable distance.
pone (694) published a list of species occurring in America north
of Mexico.

FAMILY BIBIONIDAE
MARCH FLIES

March flies are slender-bodied, stout-legged, rather hairy, and
have short, many jointed antennae. They are usually dark colored,
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 group because of the frequent appearance of the
species, Bibio albipennis albipennis Say, in large numbers in
March. The larvae usually feed on decaying vegetable matter, but
a few feed on the roots of grass and other plants.

FAMILY MYCETOPHILIDAE

FUNGUS GNATS

Fungus gnats are moderately small, slender, delicate, mosquito-
like 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 mushrooms or
on fungi growing on trees and logs. A number of others are pre-
dacious and are found under bark or in the galleries of wood
boring insects.

FAMILY CECIDOMYIIDAE

GALL MIDGES

There are about 1200 species of Nearctic cecidomyids, of which
several hundred are known to attack various trees and shrubs
(256). The oaks are especially favored as hosts but several other
species of trees are also infested, such as willow, elm, maple, wal-
nut, and the pines. The larvae of about two-thirds of the Nearctic
species cause the formation of galls or pronounced swellings by
their feeding (241). Others feed in such places as patches of pitch
exuding from injured limbs, in the excrement of insects, birds,
and mammals, and in galls produced by other insects. A few are
predators on small insects such as asphids, 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

A417

by the presence of a sclerotized structure, commonly known as a
spatula, on the underside of the front end.

The balsam gall midge, Dasineura balsamicola (Lint.), produces
swollen, oval galls about 3 mm. in diameter near the bases of
needles of balsam fir and Fraser fir. It probably occurs wherever
its hosts grow in North America (293). Damage in Christmas
tree plantations may be severe. D. gleditchiae (O. S.) produces
oblong, padlike galls on the new leaflets of honeylocust. Damage is
sometimes rather severe to thornless honeylocust seedlings in
nurseries in the Midwest. D. pseudacaciae (Fitch) attacks the
young leaves of black locust and causes them to fold. D. communis
Felt produces greenish or reddish pouch galls on the veins of red
and sugar maple leaves.

The boxwood leaf miner, Monarthropalpus buxi (Laboulbene),
an introduced species, occurs from Rhode Island south to Dela-
ware 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 yellow-
ish-brown and often drop prematurely.

Contarinia juniperina Felt, the juniper midge, is a pest of red
cedar and other junipers in the Midwest. Adults are very small,
only about 1.5 mm. long, and have bright red abdomens. Eggs are
laid on the needles of new growth, near the tips of twigs. Larvae
bore into the twigs at the bases of the needles and kill the portion
beyond the entrance hole (324). 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 out-
break 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.
Wilson (779) discussed its biology in Michigan. Other fairly com-
mon species of Contarinia include C. virginianiae (Felt), which
feeds on the fruit of choke cherry, causing it to become swollen
and deformed; and C. canadensis Felt, which produces galls on
the leaves of ash.

Asphondylia ilicicola Foote larvae feed in the berries of Ameri-
can holly, and the infested berries remain green all winter. It has
been recorded from New Jersey, Maryland, Virginia, and West
Virginia (352). The related species, A. azalea 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 (411).
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 grow-
ing trees.

Cecidomyia pintinopsis O. 8., the gouty pitch midge, causes
swellings and malformations on the twigs of Virginia and pitch
pines in eastern America. Dead needles, dead or dying twigs, and

418

distorted, twisted terminals are evidence of its attack. Eaton and
Yuill (215) discussed its biology and control. The larvae of C.
resinicola (O. 8S.) (fig. 166) are found in small patches of fluid
resin exuding from wounds caused by their feeding in Virginia
pine. C. reeksi Vockeroth larvae are found in similar situations in
jack pine. C. cupressi (Schweinitz) feeds in the seeds and causes
a leaf gall on bald cypress.

Several other species of Cecidomyia also attack a variety of
eastern trees. C. catalpae Comst. larvae feed on the young leaves
and seed pods of Catalpa. Damaged leaves wilt and fall prema-
turely, and the terminals of heavily infested trees may be dwarfed
and stunted. C. poculum O. S. produces clusters of pale or red
saucerlike galls that are attached by slender stalks to the under-
surfaces of oak leaves. C. pellex O. S. produces reddish-brown bul-
let galls on ash. C. ocellaris (O. S.) causes the yellow galls mar-
gined with red that are often seen on the upper surface of red
maple leaves. C. verrucicola O. S. produces wartlike galls on the
leaves of basswood. C. foliora Rssl. and Hkr. produces marginal
fold galls on the leaves of black, red, and pin oaks. Finally, C.
cerasiserotinae O. S. produces bright red or yellow irregular bud
or terminal galls on wild cherry.

Oligotrophus papyrifera Gagné larvae feed in the buds of paper
birch in the Lake States, causing the formation of galls (fig. 167).
The galls are comprised of the basal portions or all of one or two

F-501805
FIGURE 166.—Resin masses |
caused by feeding activity
of larvae of Cecidomyia
resinicola.

leaf petioles and either a bud or a part of the stem near the bases
of the petioles (782). Damaged leaves fall prematurely and buds
and branches are killed. Trees from 2 to 30 feet tall are attacked.
The younger ones are often badly deformed. The related species,
O. betulae (Winn.) feeds on and destroys the seeds of beech.

The willow beaked-gall midge, Mayetiola rigidae (O.S.), pro-
duces apical, beaked galls on the lower branches of many species
of willow (fig. 168). 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 larval 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 (781).

Many other species of gall midges have been recorded attacking
various species of eastern trees. A few of the more common ones
are as follows: Parallelodiplosis florida (Felt)—produces elon-
gate, pocketlike swellings on the veins and midribs of scrub and
pin oak leaves; Obolodiplosis robiniae (Hald.)—causes the mar-
gins of black locust leaves to fold; Caryomyia holotricha (O. S.),
C. sanguinolenta (O. S.), and C. tubicola (O. S.)—produce galls
on hickory; Cincticornia pilulae (Beut.)—produces subglobose or
globose, irregular, wrinkled, reddish galls on red oak leaves;
Janetiella coloradensis Felt—causes swellings at the bases of
serub pine needles; Prodiplosis morrist Gagne—feeds on the

F-519914

FIGURE 168.—Gall produced on

willow by the willow beaked-gall
midge, Mayetiola rigidae.

F-519915

FIGURE 167.—Damage to paper

birch by the cecidomyid, Oligo-
trophus papyrifera.

420

young leaves of cottonwood in the South, causing them to turn
black, unfold improperly, and drop off frequently. Terminal
growth is sometimes severely stunted. Mycodiplosis alternata Felt
—produces clublike swellings on small twigs of dogwood. This
sometimes kills several inches of the damaged twigs.

Aphidoletes thompson Mohn, a European predator of the bal-
sam woolly aphid, has been imported and is established in aphid-
infested stands of Fraser fir in the Southern Appalachians.

FAMILY XYLOPHAGIDAE
ICHNEUMON FLIES

These flies, as their common name 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 decay-
ing vegetable matter, under the bark of trees, or in decaying logs.
Erinna lugen Loew larvae may occur in large numbers under the
bark of elm in association with the elm borer. The larvae of
E. 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 some-
times aquatic and act mostly as scavengers. Zabrachia polita Coq.
has been reared from decaying pine logs and from beetle infested
elm logs.

FAMILY TABANIDAE
HORSE F'LIES AND DEER FLIES

Many members of this family are important blood-sucking
pests of livestock and wild animals. Some species also attack man
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
Tabanus. These flies breed mostly in marshes, swamps, bogs, and
ponds. The largest and best known species is the black horse fly,
T. atratus Fab. The adult is up to 25 mm. long and its bite can
result in the loss of a considerable quantity of blood. In addition

421

to the amount swallowed by the insect, as much as 100 ce. of
blood may ooze from the puncture after the fly has left. Other
important species include: the striped horse fly, 7. lineola Fab.,
T. costalis Wied., T. abactor Philip, T. quinquevittatus Wied.,
T. sulcifrons Macq., and T. nigrovittatus Macq.

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 eastern United States. Diachlorus ferrugatus (Fab.) is
a common 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 Symphoro-
myta fly persistently about the head and are very annoying be-
cause their bites are painful.

FAMILY ASILIDAE
ROBBER FLIES

Robber flies are mostly large to very large (fig. 169). Some
have long, tapering abdomens; others are stout-bodied and re-
semble bumblebees. All are rather hairy. They are predacious on
other insects and occasionally on their own kind. The larvae of
Tolmerus notatus (Wield.), 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 grass-
hopper eggs and white grubs.

COURTESY CONN. AGR. EXPT. STA.
FiguRE 169.—Adult of a robber fly,
Asilus sericeus.

422

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, es-
pecially 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 be-
cause of their habit of flying up and down, up and down, in
Swarms in the woods, along streams, or on the shores of ponds
and lakes. The fiies range in length from 1 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. They are probably all predators.
Tachydromia sp. has been reared from white pine leaders infested
with the white pine weevil.

FAMILY DOLICHOPODIDAE
LONG-LEGGED FLIES

Members of this family are very small, long-legged 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. Doli- .
chopus vittatus Loew has been reared from beetle infested hick-
ory, and Medetera sp. from beetle infested larch.

FAMILY PIPUNCULIDAE
BIG-HEADED FLIES

Pipunculids are small, dark flies with large heads composed
mostly of large approximated eyes. The larvae are small, ellip-
tical, 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 to man (516). The adults are strongly

attracted to flowers and play important roles in the cross-pollina-
tion of plants and trees. The larvae of many species feed on and

423

aid in the 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 bumblebees. The males 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. Wirth,
Sedman, and Weems (787) published a list of the syrphids occur-
ring in America north of Mexico.

FAMILY CONOPIDAE
THICK-HEADED 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 3-seg-
mented, the third segment bearing a dorsal arista. Adults fly
slowly and are usually seen around flowers. The larvae are soli-
tary 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 the larvae feed in various parts of plants. Cer-
tain species produce root and stem galls and a few are leaf
miners. 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 occur-
ring in the woodlands or forests of eastern United States are
discussed below.

The apple maggot, Rhagoletis pomonella (Walsh), an important
pest of apple, also breeds in the fruit of hawthorn, choke cherry,
plum, and dogwood. The adult is dark colored and a little smaller
than the house fly. Each wing is crossed by four dark bars which
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 wild cherry. Adults are black except for yellow
margins on the thorax, two white cross bands on the abdomen,
and a dark band on each wing. They are smaller than house flies.
The black cherry fruit fly, R. fausta (O. S.), breeds in wild 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

424

by the larvae produces a slimy condition which 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 cross-bars. Toxotry-
pana curvicauda Gerstacker breeds in papaya in Florida and
Texas.

FAMILY CHAMAEMYIIDAE

The larvae of a number of species in several genera of these
small, grayish flies are predators of aphids, scale insects, and
mealybugs. One species, Leucopis obscura Hal., a native of
Europe, was introduced into Canada against the balsam woolly
aphid in the early thirties. It quickly became established and, fol-
lowing additional colonizations, spread over most of the infested
areas in eastern Canada and into northern New England. Since
1954, colonies have been released in aphid-infested stands in Ver-
mont, New Hampshire, New York, and 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 Cz., another European species, also
imported into eastern Canada and the Pacific Northwest against
the balsam woolly aphid, is now established. A fairly large
number have developed on stem-infested trees, but their spread
has been very slow.

FAMILY LONCHAEIDAE

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 in-
sects. Lonchaea polita Say has been reared from bark-beetle in-
fested wood. L. corticis Taylor has been recorded as an important
parasite of the white pine weevil (467).

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 mark-
ings 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,
Chalara quercina (315).

FAMILY CHLOROPIDAE
CHLOROPID FLIES
These flies are small to very small and bare to nearly bare in-
sects. Some species are brightly colored with yellow and black.

The larvae of certain species parasitize or prey on other insects;

425

others have been found feeding in the seeds and cones of conifers.
Adults of the genus Hippelates are attracted to the eyes of man
and animals and are very annoying. Certain species are also re-
ported to transmit yaws and pinkeye. Gauraz apicalis Mall., Osci-
nella coxendix (Fitch), Madiza glabra Fall., and Hippelates sp.
have been reared from the leaders of white pine infested by the
white pine weevil (467).

FAMILY AGROMYZIDAE
LEAF MINER FLIES

These flies are very small and light or dark colored. 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. Frick (270) published a synopsis of North American
species.

Larvae of the genus Phytobia Lioy feed in the cambium of
living trees, making long, thin, gradually widening mines (fig.
170). 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 pith-ray flecks. Heavily infested logs may be
rendered unfit for some uses.

Phytobia setosa (Loew) attacks red and sugar maples (320).
Adults are small and dark colored. The larvae are opaque-white
and about 16 mm. long. P. pruinosa (Coq.) infests wild cherry,
maple, and black 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) attacks shadbush and P. pruni (Gross.), wild cherry.

The holly leaf miner, Phytomyza ilicicola Loew, is a serious pest
of American holly in eastern United States. The adult is a small,
grayish-black fly about 25 mm. long. The female punctures leaves
with her ovipositor and feeds on the juices exuding from the
wounds. She also deposits eggs in the undersurfaces of leaves in

F-506078, F-506069
FIGURE 170.—Cambium miner work: Left, streaklike, longitudinal mines on
trunk and roots of infested trees; right, “pitch-fieck’”’ defects in the wood.

426

punctures made near the midrib. The larvae mine the tissues
between the leaf surfaces. The mine is hairlike at first but gradu-
ally widens as the larva continues to grow. Eventually it becomes
blotchlike (fig. 171). Heavily infested leaves become unsightly
and usually drop prematurely. Leaves damaged by feeding punc-
tures become roughened, twisted, and stunted. The winter is spent
in the larval stage in the mine. 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. AGR. EXPT. STA.
FIGURE 171.—Injury by the holly leaf miner, 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 (Coq.)—
produces blotch mines in the leaves of red oak; Melanagromyza
schineri (Gir.)—causes the formation of slight swellings on the
smaller twigs of poplar; M. tiliae (Couden)—produces swellings
about half an inch long on the twigs of basswood; Phytobia posti-
cata (Meigen) —produces blotch mines in the leaves of sweetgum;
and Phytomyza clara (Melander)—produces blotch mines on Ca-
talpa bungit.

FAMILY ANTHOMYIIDAE

Members of this family are quite similar in appearance to those
of the family Muscidae. The maggots vary in habits, some feeding
on the roots of plants, some as scavengers, and others as parasites
of other insects. The seed corn maggot, Hylema platura (Meigen),
damaged red cedar seedlings in a forest nursery in Tennessee
(766). 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.

427

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 in color, but the majority are gray or brown.
There are several economically important species, such as the
well-known house fly, Musca domestica L., the stable fiy, Stomoxys
Haemetobia irritans (F.), and the tsetse flies. Fortunately,
tsetse flies, which transmit the organisms responsible for sleeping
sickness and other diseases, are native to Africa and do not occur
in this country. A number of svecies 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 Lynchia americana
(Leach) —parasitizes grouse, hawks, and owls; Olfersia fumi-
pennis (Sahl.)—attacks the bald eagle; species of Ornithoica and
Ornithomyia—feed on various small birds; and species of Lipop-
tena—parasitize deer.

FAMILY CALLIPHORIDAE
THE 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, or
larvae, the posterior spiracles are flush. A few species are eco-
nomic 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 screw-worm, 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 eradica-
tion from the region through the mass release of sexually sterile
male flies (457), it caused tremendous losses to livestock. Big
game, such as deer, and smaller animals, such as racoons, 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 addi-
tional flies and compounds the infestation. Heavily infested ani-
mals may die within a few weeks unless the maggots are killed
and the wounds treated.

428

Other blow flies occasionally attacking living animals include
Phaenicia sericata (Meigen), the greenbottle fly, the black blow fly,
Phormia regina (Meigen), and the secondary screw-worm, Coch-
liomyia macellaria (Fab.). They also oviposit on wounds or sores.
Larvae of the genus Protocalliphora are blood-sucking parasites
of nestlings.

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 3 black stripes on the thorax; and the
abdomen is checkered. The larvae have their posterior spiracles
located in a pit.

Sarcophaga aldrichi Park., an important parasite of the forest
tent caterpillar, occurs in southern Canada and from New Eng-
land to the Lake States and Southern Appalachians. It suppresses
significantly 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
(355). Full-grown larvae are about 138 mm. long. The adults are
strong, active fliers and during tentcaterpillar outbreaks may
occur in enormous numbers, swarming over everything, 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 para-
sites of many kinds of insects, especially Lepidoptera, but also
various species of Coleoptera, Hemiptera, Orthoptera, and
Hymenoptera (629). .

Tachinid flies look very much like overgrown house flies but
differ in having an entirely bare bristle on each antennae. 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 host, on
leaves and other parts of plants frequented by their hosts, or on
the ground. The number of generations per year varies from 1 to
10, depending on species and climate.

A large number of native species and a few introduced species

A429

a a

parasitize various native and introduced species of eastern forest
insects. More than 125 species have been recorded attacking lepi-
dopterous larvae in this country (638, 639). A few of the more
important introduced species are discussed here.

Compsilura concinnata (Meigen) (fig. 172) was imported
against the gypsy and brown-tail moths in the early days of 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 much of the infested area.
Even higher percentages are recorded in brown-tail and satin
moth infestations (205, 177, 751, 124).

Blepharipa scutellata (R.-D.) (fig. 178), 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. Burgess and Crossman (124)
discussed its life history and habits.

Parasetigena agilis (R.-D.) and Exzorista larvarum (L.) were
imported against the gypsy moth from 1924 to 19338. P. agilis, a
single brooded species, is probably the most important parasite of
the nun moth in Europe. EF. larvarum, a multi-brooded species,
attacks about 45 different hosts in Europe. Both species are now
important parasites of the gypsy moth throughout the generally
infested area in the United States. E. larvarum is also a parasite
of the satin moth and brown-tail moth in this country.

Two species, Townsendiellomyia nidicola (Tns.) and Carcelia
laxifrons Vill., were imported against the brown-tail moth during

wis
l oe 3

F-519565

FIGURE 173.—Adult of Blepharipa
scutellata, a parasite of the
gypsy moth and other defoliators.

F-519564

FIGURE 172.—Adult and puparium of Comp-

silura concinnata, a parasite of the gypsy
moth and other defoliators.

430

the early days of this century. T. nidicola quickly spread through-
out the infested area. During recent years it has frequently para-
sitized an average of 17 percent of overwintering brown-tail
moth larvae. C. laxifrons also occurs throughout the infested
area. Unfortunately, it parasitizes a very low percentage of the
population.

Chaetexorista javana B. & B., 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, Massachusetts
averaged 60 percent. Hrynniopsis rondanii Towns., 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.
Drino bohemica Mesn., a native of Europe, was successfully in-
troduced 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 Sturmia
inconspicua (Meig.) against the gypsy moth, brown-tail moth, and
red-headed pine sawfly (205). 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.

FAMILY OESTRIDAE
WARBLES OR BOT FLIES

Warble or bot flies are all parasitic on animals, and several are
serious pests. They are best known as enemies of domestic ani-
mals, but many species also attack various species of wild animals.
The adults are medium- to large-sized flies and resemble small
bees. Cephenomyia pratti Hunt. and C. phobifer (Clark) are
serious enemies of deer. Various species of the genus Pseudo-
bogeris infest squirrels, rabbits, cats, and meadow mice. Cattle
are subject to serious infestations by the northern cattle grub, Hy-
poderma bovis (L.) and the common cattle grub, H. lineatum
(Villers).

Order HYMENOPTERA

The order Hymenopt:ra is one of the largest and most impor-
tant orders of insects. Besides a large number of harmful species,
the order contains a far greater number of beneficial species.
Bees, wasps, and ants are among the better known groups. Bene-
ficial forms include the well-known honeybee, plus a large number
of less familiar forms. Most of the species are parasites of other
insects, and many are important pollinators of plants. Among
harmful species are the sawflies. Many of them are important
pests of forest and shade trees and young trees in nurseries and
plantations.

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

A431

hooks along the anterior margin by which they are attached to
the front wing. Some forms, such as the common ant, are wing-
less. The ovipositor is usually well-developed and in the higher
forms is modified into a sting.

The order is divided into two suborders—Anocrita (=—Clisto-
gastra) and Symphata (=Chalastogastra) (550). Each of these,
in turn, is divided into a number of superfamilies.

Suborder SYMPHATA (— CHALASTOGASTRA)

Members of the suborder Symphata are distinguished by hav-
ing the abdomen of the adult broadly joined to the thorax—the
first abdominal segment is not modified 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 saw-like. 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 to feed. Orussid
larvae are parasitic on wood-boring coleopterous larvae. While
external leaf-feeding larvae look like lepidopterous larvae, they
have only one ocellus on each side of the head and have six or
more pairs of prolegs on the abdomen, none of which bear hooks
or “crochets.”

The suborder Symphata is represented in the United States
and Canada by approximately 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 |

Members of the family Xyelidae are medium-sized to small
sawflies, most less than 19 mm. long. They differ from all other
sawflies in having the third antennal segment longer than all of
the remaining segments combined and in having three marginal
cells in the forewing. Unlike all other sawflies, except the Pam-
philiidae, they have the costal cell divided by a longitudinal vein.

The genus Xyela Dalman contains several species which feed
as larvae on the developing pollen of the male strobili of various
species of pines. The adults are usually found on the catkins of
willow and alder and occasionally on the pollen of pines. Only 16
species are known in the world (119). X. baker1 Konow—reared
or collected from longleaf, slash, and jack pine (known to occur
from Maryland to North Carolina as well as in Florida and
Texas) ; X. minor Norton—known hosts are Virginia, slash, and
longleaf pines (in Virginia and from Florida to Texas) ; X. ob-
scura (Strobl)—known hosts are jack, Virginia, slash, and long-
leaf pines from southern Canada to Florida and Texas); X.
alpigena (Strobl) —recorded from white pine and spruce (south-
ern Canada, New York, Maryland, and Illinois) ; X. styrax Bur-
dick—recorded from Virginia pine (Maryland and Virginia).

432

Ebel (218) discussed the species occurring on slash and longleaf
pines.

Pleroneura brunneicornis Rohwer occurs in eastern Canada and
in the Northeastern States from Maine to Michigan. Its hosts are
listed as balsam and white firs. Full-grown larvae are white and
from 4 to 7 mm. long. The true legs are weakly developed and the
prolegs consist of nothing more than slight swellings. In New
Brunswick, Canada, adults are found during late May. Eggs are
deposited in tightly packed needle clusters on expanding buds,
and the larvae feed in tunnels excavated in the center of new
shoots. These shoots stop growing, turn brown, and drop. Later
on, they resemble frost-killed shoots. Winter is spent in the pupal
state in oval cocoons in the soil, and there is one generation per
year (749).

An unidentified species of Pleroneura (or possibly Xyela) has
been recorded attacking loblolly, shortleaf, and slash pine in
Georgia. Eggs are deposited on new shoots and the larvae feed on
the tissues of the shoots, causing the formation of spindle-shaped
galls. Full-grown larvae vacate the galls and drop to the ground.
Pupation occurs in cells in the soil. The curculionid, Conotrachelus
caroliniensis, breeds in and hollows out the vacated galls. This
weakens the shoots and causes them to break in the wind.

Several species of Megaxyela Ashmead have been observed
feeding as larvae on hickory, and the larvae of one species of the
genus Macroxyela Kirby feeds on elm.

FAMILY PAMPHILIIDAE
WEB-SPINNING SAWFLIES

The family Pamphiliidae is represented in North America
north of Mexico by four genera and 85 or more species, better
than half of which occur in eastern America. The adults are
moderately large with long, many-segmented antennae. The ab-
domen is fiattened 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, but
abdominal legs are absent. The larvae of certain species are gre-
garious and feed together in nests which they prepare by webbing
together leaves of their hosts. Others roll the edges of leaves or
spin silken tubes in which to live (519).

The pine false webworm, Acantholyda erythrocephala (Linn.),
an introduced species first recorded in Pennsylvania in 1925, now
occurs from Connecticut to New Jersey and Pennsylvania, in the
Lake States, and in New Brunswick. Its preferred hosts are white
and red pines, but it also attacks several other pines, including
Scotch, Austrian, mugho, Swiss Mountain red, 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.

Winter is spent in the larval stage; pupation occurs in early
spring, and adults appear from about mid-April to mid-May. Eggs

433

are deposited in short rows of three to 10 eggs each on the pre-
vious year’s needles. The larvae spin loose webbing about them-
selves 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 (fig. 174). Full-grown larvae
drop to and enter the ground for hibernation. There is one gen-
eration per year. Heavy infestations sometimes develop locally,
causing severe defoliation.

COURTESY CONN. AGR. EXPT. STA.
FIGURE 174.—Damage and
webbing by the pine false
webworm, Acantholyda er-
ythrocephala.

Acantholyda zappei (Rohwer), the nesting-pine sawfly, occurs
in southern Canada and the Northeastern, Central, and Lake
States. 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

434

develop and may reach a length of 5 inches by the time the larvae
are full-grown. Winter is spent as mature larvae in cells in the
soil, and pupation occurs in the spring. There is one generation
per year (802).

Acantholyda_ circumcincta (Klug) has been recorded from
Georgia and Florida. During 1968, it defoliated approximately
100 acres of sand pine in Florida.

Additional eastern species of Acantholyda, along with known
hosts and known areas of distribution, include the following:
A. apicalis Westw.—loblolly pine (in North Carolina, Mississippi,
and Florida) ; A. pint Rohwer—red pine (in southern Canada and
from New York to North Carolina) ; A. angulata (MacGillivray)
—white, pitch, jack, Austrian, and Japanese red pines (from
southeastern Canada and Maine to Pennsylvania and Minnesota) ;
A. luteomaculata (Cresson)—white and jack pines, preferably
white (in southeastern Canada and from New Hampshire and
Massachusetts to Ohio and Minnesota); A. maculiventris (Nor-
ton) —balsam fir and white spruce (in southern Canada and from
Maine to North Carolina).

Cephalcia fascipennis (Cresson) occurs in Canada, Maine, New
Hampshire, and West Virginia. Its hosts are listed as blue and
white spruce. 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 pres-
ence of larval nests (224). 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 Quebec, New York, Massachusetts, Connecticut, Penn-
sylvania, Virginia, and West Virginia. Daviault (182) discussed
its life history.

The plum web-spinning sawfly, Neurotoma inconspicua (Nor-
ton), occurs in southeastern Canada and from New England to
Montana in the United States. Its hosts are listed as hawthorn
and various wild plums 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 be-
yond the coxae. The larvae are gregarious and web together the
foliage (the webs are somewhat similar to those of the fall web-
worm). Heavily infested trees no more than 6 feet tall may
support up to 25 webs, some of which may enclose entire branches.
Such trees may be completely defoliated.

Neurotoma fasciata (Norton) occurs in southern Canada, and
in the Eastern States from New Hampshire to Florida, westward
to Illinois and Michigan. Its hosts are listed as wild black and pin
cherries. The full-grown larva has a shiny black head and pro-
thoracic plates and a deep green body, and is about 18 mm. long.
The larvae are gregarious and construct dirty, brown nests on
the branches and shoots of young trees especially. Larvae over-
winter in cells in the soil. Some of these larvae pupate in the
spring. The remainder do not pupate until late fall or the follow-
ing spring. The related species, N. crataegi Middlekauff, feeds on
hawthorn from Massachusetts to Michigan and Illinois.

435

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 pro-
tecting sheaths. The upper piece is a rigid lance with grooves
along which the other two pieces slide. The last two consist of
thin plates with their flat inner surfaces together. Each plate is
usually shaped like a long acute triangle with the edge of one long
side along a groove of the lance. The edge of the other long side
is saw-toothed. The narrow base of the lance is the point from
which springs the rod that attaches the blade to the abdomen and
controls its movements.

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 distinguish-
ing sawfly larvae from lepidopterous larvae is the presence of
only one 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 webbed
nests; 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. Leaf mining
species sometimes form their cocoons within their mines.

Sawfly adults frequently resemble small bees or wasps, except
for their antennae, which differ in the number of joints. The an-
tennae may also be feathered, clubbed, threadlike, forked, or may
bear spurs on some of the joints. It is rather difficult to separate
the adults into species because the females are variable, with the
variations in related species overlapping, while the males are
monotonously alike. It is often much easier to separate the larvae
into species.

Sawflies comprise 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. During the past several years, outbreaks of an in-
creasing number of species have been reported, possibly as a
result of the establishment of more and more pine stands in
planting programs. Ross (622) published a list of the species in
the superfamily occurring in America north of Mexico.

FAMILY PERGIDAE

The family Pergidae contains only one genus, Acordulecera
Say, and 138 species, all of which occur in eastern United States.

436

Full-grown larvae are greenish with light or blackish heads and
distinct lateral lobes, and measure less than 12 mm. long. Each
body 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 winter is
spent in cocoons in the ground. There is usually one generation a
year, but some species occasionally have a partial second.

FAMILY ARGIDAE
ARGID SAWFLIES

The family Argidae is represented by eight genera and 32
species in the United States and Canada, the majority of which
occur in the Eastern States. The adults are medium- to small-
sized, stout-bodied sawflies. They can be recognized by their
three-segmented antenna, the third segment of which is very long
and sometimes U-shaped or Y-shaped. The more common species
are also usually either 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 rather thick-set, widest on ab-
dominal segments one to three, tapering toward the rear end,
and the venter is flattened.

The birch sawfly, Avge pectoralis (Leach), is widely distributed
in the Northeastern States and southern Canada. Its preferred
hosts are various birches, especially gray and paper. It also has
been recorded feeding on willow. Full-grown larvae are about 18
mm. long. The head is reddish-yellow with a spot on each side;
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 full-grown larvae
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 States are as follows: Arge clavi-
cornis (Fab.)—willow, birch, and azalea; A. scapularis (Klug) —
elm and birch; A. coccinea (Fab.)—sumac; Sterictiphora pruni-
vora (Dyar)—cherry; and Arge sp.—elm and oak.

FAMILY CIMBICIDAE
CIMBICID SAWFLIES

The family Cimbicidae is represented in the United States and
Canada by 12 species, some of which have been recorded feeding
on trees in the Eastern States. The adults are large; the antennae
are clavate with an abrupt enlargement at the tip. 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. During life they are covered with a
waxy bloom.

437

The elm sawfly, Cumbex 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 Northeas-
tern States west to Colorado. 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 the
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 and about 48 mm. long, with a pebbly
skin and a black dorsal stripe. While feeding, the larva usually
coils its rear end around a stem or twig; at rest it usually lies
coiled up like a snail fig. 175).

COURTESY CONN. AGR. EXPT. STA.
FIGURE 175.—Larva of elm
sawfly, Cimbex americana.

Adults appear from about mid-May to mid-August, and the
eges are deposited in pockets cut into leaf tissues. Larvae occur
from June until October, depending on location. When they be-
come 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 larval 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 the 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.

Trichiosoma triangulum Kirby occurs fairly commonly in the
Eastern States. The larvae are solitary feeders on the foliage of
birch, poplar, willow, and wild cherry. At maturity, they are
about 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.

438

FAMILY DIPRIONIDAE

CONIFER SAWFLIES

The family Diprionidae includes many of the most serious de-
foliators of conifers. 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 Dipri-
oninae, and five genera (550). Adults are described as follows:
Antennae composed of 18 or more segments, serrate in the female,
and pectinate or bipectinate in the male. Mesosterno-plural su-
tures atrophied, mesoscutellum with anterior margin V-shaped,
posterior margin with an exremely narrow and cordlike poster-
gite (621). 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 (801).

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 (11, 160).

The genus Monoctenus Hartig is represented in the United
States and Canada by four species, all of which occur in eastern
America on Juniperus. None are considered serious pests of forest
or shade trees. M. melliceps (Cresson) occurs in southern Canada
and in the Northeastern States west to Kansas. Its hosts are red
cedar, arborvitae, 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 Rohwer contains many of the most de-
structive species of sawflies. During recent years outbreaks of
several species have occurred, resulting in serious damage to
young pines on thousands of acres of pine plantations in eastern
United States. Some outbreaks spread over millions of acres of
forested areas before subsiding. Atwood and Peck (12) and Ross
(623) discussed the taxonomy of the genus and Krombein (442)
published a check list of North American species.

The red-headed pine sawfly, Neodiprion leconter (Fitch), occurs
in southeastern Canada and throughout eastern United States. Its
preferred hosts are jack, red, shortleaf, loblolly, slash, longleaf,
pitch, and Swiss Mountain pines. 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 22 to 25 mm. long. The head is reddish and the body is
yellowish-white, with six rows of black spots (fig. 176).

Winter is spent in the prepupal stage. 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 100 eggs. The larvae feed gregariously on new
and old needles and usually completely defoliate one branch be-

439

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COURTESY ARS

FIGURE 176.—Red-headed pine sawfly.

440

fore moving to another. They also feed frequently on the tender
bark of young twigs. Sometimes they completely defoliate a tree,
with defoliation progressing from the top downward, before they
reach maturity. When this happens, they may abandon the tree
and migrate for distances of several yards in search of new foli-
age. Full-grown larvae drop to the ground, enter the soil, and
spin tough, reddish-brown cocoons in which they spend the
winter. In the South there may be five generations per year; in
the Northern States and Canada there is only one.

The red-headed pine sawfly is one of the most widespread and
destructive of the pine sawflies. It usually feeds on young trees,
preferably on trees from 1 to 15 feet tall (474). In the southern
portions of its range, it seems to prefer trees growing in shaded
areas. In Canada, eggs are typically laid on trees in full sunlight
and often on the leaders. Damage is most severe in young planta-
tions and nurseries and to reproduction and ornamentals. Out-
breaks occur frequently through the area of infestation. From
1935 to 1953, a total of 73 were recorded throughout eastern
United States, 36 of which were widespread (58).

Among the more important natural control factors affecting
the species are parasites, a polyhedrosis virus disease, and tem-
perature extremes during the larval stage. The egg parasite,
Closterocerus cinctipennis Ashm., and the larval parasite, Spathi-
meigenia aurifrons Curran, are often quite effective. The Euro-
pean parasite, Dahlbominus fuscipennis (Zett.), was liberated in
infested stands in Tennessee, Alabama, New York, and Michigan
in the late thirties and early forties. It has been recovered only
in Michigan.

A number of management practices have been suggested for
preventing damage in plantations by the red-headed pine sawfly:
(1) Do not plant hard pines under hardwoods or closer than 25
feet to hardwood borders; (2) remove hardwood overstories be-
fore the planting of pines; (8) replant pines in areas of poor
survival; and (4) promote early closure of plantations by plant-
ing pines with spacing not greater than 6 by 6 feet (in the North
Central States).

Neodriprion prattt pratti, the Virginia pine sawfly, has been
recorded from New Jersey and Maryland to North Carolina and
westward to Illinois. Its hosts are listed as Virginia, pitch, jack,
shortleaf, loblolly, and red pines. Newly-hatched larvae are pale
green, with black head capsules, and are about 3 mm. long. Full-
grown larvae (fig. 177) are spotted or marked with longitudinal
black stripes and are from 16 to 23 mm. long.

Winter is spent in the egg stage, and hatching occurs in early
spring. Newly-hatched larvae feed gregariously on previous year’s
needles, beginning about one-half inch below the tip. Feeding in
groups of 2 to fifteen 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 por-
tions 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.

44]

F-519912

FIGURE 177.—Larvae of Neo-

diprion pratti pratti, the
Virginia pine sawfly.

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 (634, 536).

Prior to the development of an outbreak in Maryland, Virginia,
and North Carolina during the late fifties, this species was not
considered an important pest in the Middle Atlantic States. Sur-
prisingly, the outbreak occurring at that time swept over more
than 14 million acres 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 hymenopteron, Hxen-
terus canadensis Provancher, is also an effective parasite of un-
cocooned prepupae on the ground (86). Dahlbominus fuscipennis
(Zett.), an introduced parasite of sawfly cocoons, has been liber-
ated 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 (485).

The jack-pine sawfly, Neodiprion pratti banksianae Rohwer,
occurs in southeastern 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 by larvae if they are grow-
ing with heavily infested jack pine. Full-grown larvae are yellow-
ish-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.

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

442

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 compliment of eggs on
the needles of one twig. There is one generation per year.

All sized trees are attacked, but small trees with open crowns
are more susceptible to damage than trees in closed stands. Even
aged stands and plantations are especially attractive. Mortality
from a single defoliation seldom occurs, but it may result from
several successive defoliations. Generally, the most serious dam-
age 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 by Ewan (232).

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. Supraspir-
acular spots are usually partially or entirely atrophied in the
middle of the body. Widespread outbreaks have occurred in pitch
pine and shortleaf pine stands in New Jersey during recent years.

Neodiprion taedae taedae Ross [=americanum (Leach) ], the
loblolly pine sawfly, has been recorded from Maine and eastern Vir-
ginia. 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
which extends from the second thoracic to the ninth abdominal
segments, 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. In late September, they drop to the
ground and tunnel in for several inches, where they spin cocoons.
Adults appear about mid-October and lay their eggs, in rows in
the sides of needles. There is one generation per year (345).

Since the new foliage is not eaten, infested trees are not com-
pletely defoliated or killed. Infestations occur more commonly in
understocked pine stands where the trees retain their lower
branches.

Neodiprion taedae linearis Ross., the Arkansas pine sawfly, long
recognized as a pest of loblolly pine in Arkansas, is now known to
occur also in Louisiana, southeast Texas, Mississippi, South Caro-
lina, Missouri, Ohio, and Illinois. As far as known, loblolly and
shortleaf pines are its only hosts, loblolly 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
eregariously in groups, often encircling the needles about half
way from end to end, and partially girdling them. Infested ter-
minals soon take on a reddened appearance. Older larvae feed
singly or in pairs and consume the entire needle, leaving short

443

stubs on the branch. They still retain their gregarious habit,
however, and move in a group from branch to branch (fig. 178).
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 (162). Full-grown larvae drop to the
ground and spin mahogany-colored cocoons in the litter or top-
soil. Pupation occurs in October or November and the adults
appear immediately. Eggs are laid in slits cut into the needles,
usually two to ten per needle. Each female lays from 90 to 120
eggs, often all in the needles of one twig (742).

This sawfly is found chiefly on medium-sized or large trees in
forest stands. Several outbreaks have been recorded. One, which
lasted from 1945 to 1948, spread over a gross area of about 3
million acres in Arkansas before it subsided. Defoliation causes a
reduction in growth, especially if the tree is defoliated during
successive years, and tree mortality may occur (fig. 179). Impor-
tant natural control factors are a polyhedrosis virus disease, cold,
rainy weather in the spring, and two larval parasites, the dip-
teron, Anthrax sinuosa (Wied.), and the hymenopteron, EH xente-
rus canadensis Provancher.

The Swaine jack-pine sawfly, Neodiprion swainet 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 fifties. It is now widely distributed in the Upper Penin-
sula, Michigan, and in north-central Minnesota and Wisconsin.
Jack pine is its favored host, but red, Scotch, and white pines
growing in close proximity to heavily infested jack pines are also
subject to defoliation by migrating larvae. 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 also occasionally
found.

In the Lake States, winter is spent as full grown larvae in
cocoons in the litter or top soil, and pupation and adult emergence
occurs in early to late spring. Eggs are laid in current year’s

F-386627
FIGURE 178.—Larvae of Neodiprion taedae linearis, the Arkansas pine sawfly.

444

F-486626

FIGURE 179.—Large pines defoli-
ated and killed by Neodiprion
taedae linearis, the Arkansas
pine sawfly.

needles, one to three eggs per needle, during June or July. The
larvae are gregarious and feed primarily on old foliage, usually
on exposed trees during July and August, until they become ma-
ture and drop to the ground. There is one generation per year
(48).

Several outbreaks have occurred in jack pine stands in Ontario
and Quebec, and heavy tree mortality has occurred. Furthermore,
practically all surviving mature trees over many square miles
remained stag-headed for several years. Some of the most severe
infestations in the Lake States have occurred in jack pine planta-
tions and windbreaks. Aerial application of a polyhedral virus
spray provided satisfactory control for at least one year in tests
conducted in Quebec (654).

The red-pine sawfly, Neodriprion nanulus nanulus Schedl, oc-
curs in many Northeastern States west to Wisconsin and in
southeastern Canada west to Saskatchewan. Its favored hosts are
red and jack pines, but it also feeds on white, Japanese red, and
mugho 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.

Winter is spent in the egg stage. Hatching occurs in early May,
and larvae are present until July or August, depending on loca-
tion. They are gregarious and usually consume all of the mature
needles from one branch before migrating to another. Mature
larvae drop to the ground and spin cocoons in the duff. Adults

445

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 does not lay her entire complement of eggs
in the needles of a single twig—sometimes only one needle per
twig is chosen. There is one generation per year (409).

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 percentage of the affected trees may be
killed. Overmature trees are particularly susceptible to damage.

The white-pine sawfly, Neodiprion pinetum (Norton), occurs
throughout the range of eastern white pine, its preferred host, in
eastern United States and southern Canada. Pitch, shortleaf, red,
and mugho pines are also attacked at times. 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 her eggs in needles, three to
four 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 and
spin cocoons in the soil. Winter is spent as a mature larva in a
cocoon spun in the soil and pupation occurs in the spring. There
is one, and sometimes a partial second, generation per year, de-
pending on location.

The white-pine sawfly attacks trees of all sizes both in planta-
tions 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 balsam-fir sawfly, Neodiprion abietis (Harris), occurs from
New England to the Lake States and Missouri and from coast to
coast in southern Canada. 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, eat-
ing 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 ap-
pear 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 lake shore
lines 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 have also been damaged. The introduced parasite, Dahl-
bominus fuscipennis (Zett.), has destroyed up to 40 percent of

446

cocoons in the duff in Maine. Cocoons spun among the needles on
the trees in the same stands escaped parasitization.

Neodiprion abbotu (Leach) has been recorded in Ontario, Wis-
consin, Virginia, North Carolina, South Carolina, Georgia, and
Florida. Its hosts are listed as loblolly, slash, longleaf, shortleaf,
red, and Carribean pines. The full-grown larva has a brown to
black head with a light spot on the frons and the thorax and
abdomen are pale green. There are also four dark green to black
lateral stripes.

Winter is spent as a mature larva in a cocoon and pupation
occurs in the spring. Adults emerge as early as Jate March in the
South. Eggs are laid in single rows of 15 to 20 eggs each mostly
on 2-year-old needles, the eggs of a row almost touching each
other. Newly-hatched larvae feed gregariously; older ones tend
to feed singly. Mature larvae spin cocoons on the needles of the
tree. There appears to be at least three generations per year in
the South. First generation larvae are found in May; second gen-
eration larvae during July and August; and third generation
larvae from October to December (346).

Light defoliation by this species may not be noticeable because
of the tendency of the large larvae to wander in search of needles
on which to feed.

Neodiprion rugifrons Middleton occurs in southern Canada and
the Lake States and feeds on jack pine. Full-grown larvae have
orange-brown to bright orange heads with black eyespots. The
prothorax is yellowish-green to light olive-green with dark mark-
ings. A dark lateral stripe extends backward along each side from
the prothorax to the ninth abdominal segment. The question as to
whether this is a true species, or whether it is a member of a
group known as the ‘“Virginianius complex” is not settled. The
complex consists of three species, N. virginianus, N. rugifrons,
and N. dubiosus.

Winter is spent in cocoons in the soil. Adults emerge in May
and June and lay their eggs in l-year-old or older needles. There
are one or two generations per year. The larvae of the first gener-
ation feed in colonies on previous year’s needles, while those of
the second generation feed on foliage of all ages.

In heavy infestations, jack pine may be completely defoliated
by September or October. Planted pines from 3 to 20 feet tall may
be killed. Trees in isolated stands and windbreaks are also subject
to severe damage (767).

The European pine sawfly, Neodiprion sertifer (Geoff.), an in-
troduced species first recorded in North America at Somerville,
New Jersey, in 1925 (636), is now widely distributed. It has been
recorded from New England to Michigan and southwestern On-
tario, and south and southwest to southern Ohio, southern Illinois,
and southwestern Iowa. It has many hosts, particularly Scotch,
red, jack, Japanese red, Table-Mountain, and mugho pines. Eas-
tern white, Austrian, ponderosa, shortleaf, and pitch pines also
are fed on to some extent when they occur in mixture with more
favored species.

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.

447

There are also two light and one intense green or black stripes
on each side.

Winter is spent in the egg stage and hatching occurs from
early April to mid-May. The larvae feed gregariously on previous
year’s foliage (fig. 180) and devour all of the 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 (777). When an entire tree
is defoliated before the larvae reach maturity, it is vacated and
the larvae craw! 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. Six to eight 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.

F-519945

FIGURE 180.—Colony of lar-

vae of the European pine

sawfly, Neodiprion serti-

fer. Larvae hanging head

downward were killed by
a polyhedral virus.

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 mor-
tality but it is not serious. Losses in height and diameter growth
may be severe however, especially following repeated defoliations.
Damage is most severe to Scotch pines grown as Christmas trees
in plantations.

During the late thirties and early forties, colonies of the two
European parasites, Dahlbominus fuscipennis (Zett.) and Hwxen-
terus abruptorius (Thunberg), were received from Canada for
release in eastern infestations of this sawfly. D. fuscipennis be-
came established in New Jersey and within a few years became
rather abundant locally (299). E. abruptorius also became estab-
lished in New Jersey. So far, neither species has given adequate
control. A supply of a European polyhedrosis virus of the sawfly
was also received from Canada for use in its control (66). It has
proved to be very effective when applied as a spray either by
aircraft or by knapsack or hydraulic sprayers. Lyons (466) re-

448

viewed the literature on the biology, ecology, and control of the
European pine sawfly.

Neodiprion excitans Rohwer occurs from Virginia to Florida
and west to Texas. Loblolly and shortleaf pines are its preferred
hosts. Slash, longleaf, pond, and Sondregger pines are also at-
tacked but to a much lesser extent (709). The full-grown larva is
olive-green and about 25 mm. long. (fig. 181). 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.

Winter is spent mostly in cocoons, but occasionally in the egg
stage or as older larvae. Adult appearance begins in early spring
and continues until it reaches a peak in late summer. Egg laying
begins in March by the female inserting her eggs in slits cut near
the bases of needles. Needles closest to the tips of exposed shoots
are chosen during spring and fall, whereas needles on shaded
shoots are chosen during hot weather. The peak of egg laying
occurs in July or August in east Texas and from late August to
October in the Southeast. Newly-hatched larvae are gregarious,
often with several encircling a single needle. Older larvae feed
singly or in pairs on a single needle, preferably older needles.
They usually eat most of the needle, leaving only a stub. Some-
times they attack the needle at the base first, causing the re-
mainder of the needle to fall. When a branch is completely
defoliated, the larvae migrate in groups to other branches or
trees. Full-grown larvae either drop to the ground and spin
tough, light to dark brown cocoons in the duff or loose topsoil, or
remain on the tree and spin their cocoons on twigs, needles, or

F-514368
FIGURE 181.—Larva of
Neodiprion excitans.

449

beneath loose bark scales. There are four or five generations per
year (347, 346, 709).

Infestations usually occur on medium to large trees in moder-
ately to densely stocked stands, but they are also occasionally
found on seedlings and saplings. Because of the lateness of the
season when the larvae are most active, heavily infested trees
may remain without needles throughout the winter. Heavily de-
foliated trees are weakened and rendered susceptible to attack by
bark beetles, especially [ps spp., and wood borers. Tree mortality
as a result of defoliation has not been reported. Several outbreaks
have been recorded in Florida and Texas in recent years, most of
which subsided after one season. A polyhedral virus disease and
several parasites and predators apparently are highly effective in
bringing outbreaks to an early end.

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. Full-grown non-feeding larvae are gray-
pink with a tinge of yellow on the lateral margins (49). N. piniri-
gidae (Norton) feeds on pitch and shortleaf pines from New
Jersey to Maine. The full-grown larva is dull green, has a double
longitudinal black line on the dorsum and a broken black stripe
on each side. Sometimes a double row of black dots occur below
each lateral stripe. N. warreni Ross occurs on shortleaf pine in
Arkansas. Full-grown larvae have almost entirely black heads and
broad black longitudinal stripes on the dorsum. An outbreak was
recorded in 1958. N. merkelit Ross attacks slash pine in southern
Florida. The head of a full grown larva is reddish above the eyes
and sooty black below, with a reddish infusion. N. hetricki Ross
has been observed feeding on loblolly pine in Virginia and pond
pine in South Carolina. The head of the larva is black, subdorsal
lines are solid, and supradorsal lines are composed of separate
spots or beaded lines. N. compar (Leach), feeds on red and jack
pines in southeastern Canada, Virginia, North Carolina, and
Georgia. N. maurus Rohwer occurs on jack pine in southeastern
Canada and the Lake States.

The genus Diprion Schrank is represented by only three species
in the United States and Canada and all three are of foreign
origin.

The introduced pine sawfly, Diprion similis Hartig, was first
recorded in North America at New Haven, Connecticut, in 1914.
It now occurs from Maine to Virginia, in the Central and Lake
States, and in southern Ontario and Quebec. White pine appears
to be its favored host but Scotch, red, jack, and mugho pines are
also commonly attacked. Full-grown larvae (fig. 182) are about
20 mm. long. The head is shiny black and the body is dark brown
or black with a double black stripe down the dorsum and black
and yellow spots on the sides.

Winter is spent in the prepupal stage. Pupation occurs in early
spring and the adults appear from May to mid-June. The females
are reported to possess a potent sex attractant (161). Eggs are
laid in rows of about 10 eggs each and deposited in slits cut in the
edge of old needles. Young larvae feed gregariously; older ones,
singly. First generation larvae feed on old needles only, while the
larvae of later generations eat both new and old needles. Young

450

larvae eat only the outer, tender parts of needles; older ones con-
sume the entire needle and nibble the bark. First generation
cocoons are usually spun among the needles, at the bases of small
branches, or in bark crevices on the trunk, and pupation occurs
in late July. Second generation larvae feed until September and
then spin cocoons on the tree or in the duff. Under favorable con-
ditions some pupate and emerge in late fall to produce a partial
third generation. The remainder do not pupate until the following
spring. Because of some overlap between the first and second
generations, all stages of the sawfly may be found during the
summer (778, 520).

Heavily infested trees may be completely defoliated in one
season. When this happens late in the season after the buds are
formed, many branches and occasionally trees may be killed. The
introduced parasite, Monodontomerus dentipes (Dalm.), is occa-
sionally very effective in control. Up to 75 percent parasitization
has been recorded in New Jersey. Second generation larvae
and cocoons located above the snow line on trees are particularly
susceptible to low winter temperatures.

Diprion frutetorum (Fab.) was discovered in North America
in Massachusetts and Rhode Island in 1932, and now occurs
throughout many of the Northeastern States and in southeastern

| Whe

COURTESY CONN. AGR. EXPT. STA.
FIGURE 182.—Larvae of the ss.

introduced pine sawfly, B\ Saw

Diprion similis. ,

—

451

Canada. 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. 183).

Winter is spent as a full-grown, cocooned larva 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. Usually,
the first sign of infestation is the presence of frass and green
needle fragments on the ground beneath the tree. Heavy infesta-
tions occur occasionally in pine plantations (637).

The European spruce sawfly, Diprion hercyniae Hartig, was first
recorded in North America near Ottawa, Canada, in 1922. In
1929 it was found in the United States on Mount Washington,
New Hampshire. By the late thirties it had spread throughout
the eastern spruce forests. White spruce appears to be the
favored host; however, red, black, and Norway spruces also are
attacked. Young larvae are yellowish-green; older larvae are
darker green, marked with five longitudinal white lines, and
about 20 mm. long (fig. 184). During the last instar, the white
lines are absent and the body is somewhat shortened.

In the Northeastern States there may be 1 or 2 or a partial
third generations per year, depending on location. Farther north
there is only one. In the Northeastern States, adults of the first
generation appear from early May to early June. 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

F-519578
FIGURE 183.—Larvae of
Diprion frutetorum.

F-519573
FIGURE 184.—Larva of European
spruce sawfly, Diprion hercyniae.

452

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 (604).

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 2,000 square miles 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 6,000 square miles, and
tree mortality in some areas had reached serious proportions.
Conditions continued to worsen through 1937, but in 1938 a gen-
eral reduction in population levels was reported from nearly all
regions. The downward trend in populations continued and by
1942, they had declined to medium or light intensity in all but a
few restricted localities. The collapse of the outbreak was caused
by an accidentally introduced polyhedral virus disease (26). The
virus was first observed in Canadian infestations in 19380, but it
was not until 1988 that numerous diseased larvae became notice-
able in parts of New Brunswick. Four years later it had brought
the widespread outbreak under control. Since then populations
have been fluctuating at very low levels and the sawfly is no
longer considered an economic pest (561).

During the thirties and forties, the Canadian Government im-
ported and liberated approximately 20 species of parasites in
infested areas (484). 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
(205).

FAMILY TENTHREDINIDAE

The family Tenthredinidae is represented by more than 350
species in the United States and Canada, the majority of which
occur in eastern America. The larvae are all leaf feeders, leaf
miners, gall formers, or fruit borers. Many species are important
pests of forest and shade trees and forest plantations.

The antennae of the adults have seven to ten segments and
range in shape from setaceous and filiform to clavate. The meso-
thorax 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 pre-apical spurs, and
the apical spurs of the front tibia 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
(621, 801).

Heterarthrus nemoratus (Fall.) (=Phyllotoma nemorata
[Fall.]), the birch leaf-mining sawfly, an introduced species first
recorded from North America in 1905, is now widely distributed

453

in the Northeastern States and southeastern Canada. 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 brown-
ish, and is about 10 mm. long.

In Maine, winter is spent in the larval 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 blister-like or blotch
mines free of frass (fig. 185). Each full-grown larva constructs a
cocoon or hibernaculum within its mine. The leaf then falls to the
ground and the larva remains in its hibernaculum throughout the
winter. There is one generation per year (591).

Heavy infestations of the birch leaf-mining sawfly occurred in
Maine during the 1920’s and 30’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 (204), two of which, Kratochviliana laricinellae (Ratze-
burg) and Phanomeris phyllotomae Muesebeck, became
established.

The birch leaf miner, Fenusa pusilla (Lepeletier), an introduced
species first recorded from North America in Connecticut in 1928,
now occurs in southeastern Canada and from Maine to New Jer-
sey and west to Ohio and the Lake States. 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.

F-519525
FIGURE 185.—Mines and hi-
ee? bernaculae of Heterarthrus
‘ie, ot, nemoratus, the birch leaf-
ee mining sawfly, in leaf of
paper birch.

454

Winter is spent as full-grown larvae in cocoons in small earthen
cells 1 or 2 inches below the soil surface. 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. Then, as they increase
in size, the mines of different larvae coalesce and form large,
hollowed-out brown areas in the leaf. These areas (fig. 186)
wrinkle and turn brown. Full-grown larvae chew their way out
of the leaf and drop to and enter the ground where they form
earthen cells in which to pupate. There are three or four genera-
tions per year in the southern portions of the infested region
(271).

Outbreaks occur frequently in the Northeastern States and re-
sult in the browning of birch stands over wide areas. Some tree
killing may occur, but the greatest damage is the weakening of
affected trees, which leads to attack by other insects.

The elm leaf miner, Fenusa ulmi Sundevall, an introduced
species, occurs in southeastern Canada and the Northeastern
States west to the Lake States. Its preferred hosts are English,
Seotch, 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 full-grown larvae 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 be-
tween the leaf surfaces, causing large blotch or blister-like 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. 187). These leaves soon wither and fall. Where only a
small portion of a leaf is mined, the surfaces dry out and crack,

COURTESY CONN. AGR. EXPT. STA.

FIGURE 186.—Mines of the
birch leaf miner, Fenusa
pusilla, in leaves of gray
birch.

455

COURTESY CONN. AGR. EXPT. STA.
= FIGURE 187.—Mines of the elm leaf
Le. miner, Fenusa ulmi, in elm leaves.

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.
This species appears to be most injurious to small trees in nur-
series and ornamental plantings.

The European alder leaf miner, /‘enusa dohrnii (Tischbein), an
introduced species, occurs in southeastern Canada and through-
out the Northeastern States. Its hosts are listed as alders, es-
pecially the introduced European alders. Winter is spent in the
larval stage in cocoons in the ground. Pupation occurs in the
spring, and the adults appear and lay eggs during late May and
early June. The larvae feed in the tissues of the leaf, forming
blotch mines. 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 to spend the winter.

Profenusa thomsoni (Konow), possibly an introduced species
from the Far East, is widely distributed in southeastern Canada
and from Maine to Illinois and Wisconsin. Its hosts are gray,
paper, and yellow birches. In Ontario, female adults were ob-
served in late July and early August and laid eggs in the tissues
of leaves. The larvae mined the tissues, forming light-colored,
blotch mines. Up to 40 larvae were found in a single mine. When
all of the tissues of a leaf were consumed, all of the larvae, re-
gardless of age, vacated the mine and dropped to the ground. Of
these, only those in the latter part of the fifth instar were able to
enter the soil and survive. In light infestations, sucker growth up
to about 4 feet tall in shaded locations was preferred. In contrast,
trees up to 35 feet tall in all types of habitats were attacked in
heavily infested areas. Damage was not considered severe because
defoliation occurred late in the season (505).

Profenusa mainensis Smith, a newly-described species (655),
has been observed mining the leaves of oak, primarily red oak, in
Maine during recent years. Full-grown larvae are about 6 mm.
long and have prognathous, octagonal heads more than twice as
wide as long. Winter is spent as mature larvae in cells in the duff.

456

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.

Profenusa canadensis (Marlatt) has been recorded mining the
leaves of hawthorn and cultivated cherry in Massachusetts, New
York, and Wisconsin. Serious infestations have occurred in Mas-
sachusetts and New York. P. lucifex (Ross) attacks white oak in
Maine and bur oak in Ontario.

Messa populifoliella (Townsend) mines the leaves of poplar
from New Brunswick southwestward through the United States
to Arizona.

The pear-slug, Caliroa cerasi (L.), an introduced species, occurs
from coast to coast in southern Canada and 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 shadbush. Full-grown
larvae are tadpole-shaped, slug-like, 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 or in cocoons composed of
grains of earth and a substance secreted by the larvae. Pupation
occurs in June. Eggs are deposited singly in small semi-circular
slits cut in the leaf tissue. The larvae feed mostly from 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. Adults appear in 2 or 3 weeks and lay
eggs. Larvae of this generation are present in August and Sep-
tember. When they become mature, they also drop to the ground.
There are two generations per year throughout most of the in-
fested portions of the United States.

Caliroa lineata MacGillivary, the pin oak sawfly, has been re-
corded in New Jersey, North Carolina, and Missouri. Its hosts
are pin, white, and various red oaks, preferably pin. Full-grown
larvae are slug-like, yellowish-green, and about 12 mm. long. The
head and thoracic legs are shiny black. There appears to be at
least two generations per year in New Jersey (319). Both larvae
and adults may be found almost any time from late May or early
June until late September. The larvae feed on the lower surfaces
of the leaves, leaving only the upper surface and a fine network
of veins. Defoliation is usually noticed first in the tops of trees.
Late in the season heavily infested trees may be completely de-
foliated.

Caliroa (near) quercuscoccineae (Dyar) has been recorded
feeding on pin oak in Massachusetts and North Carolina. 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 epi-
dermis of the leaf, leaving it almost colorless and transparent. Evi-
dence of feeding is apparent during late summer. There may be

A457

two generations per year, and the winter may be spent in the lar-
val stage (40).

The larch sawfly, Pristiphora erichsonii (Hartig), probably an
introduced species, was first recorded in North America at Bos-
ton, Massachusetts, in 1880. It now occurs in all Canadian Prov-
inces, in Alaska, and all of the northern tier of States except
Ohio, Indiana, and Idaho. Its hosts are listed as tamarack and
Western, European, Japanese, and Siberian larches. In eastern
America, tamarack is most seriously infested. Full-grown larvae
(fig. 188), 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 band, tapers sharply posteriorly, and is keeled longitudi-
nally 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 loca-
tion. Eggs are laid in rows under the bark of current terminal,
lateral twigs. Hatching occurs in about 8 days, and the larvae
move back to the foliage on older twigs, where they usually feed
in groups. Feeding is completed in about 20 days and the mature
larvae drop to the ground, enter the duff, and spin tough, papery,
brown cocoons. Because of the long period of adult emergence,
feeding larvae, cocooned larvae, pupae, and adults all may be
found at the same time in early summer. Normally, there is one
generation per year. Occasionally, however, small numbers of
second-generation adults are produced. A small number of larvae
enter dianause and require 2 years to complete one generation
(210, 307).

Larch sawfly larvae eat the needles on older twigs, and where
the attack is heavy, the entire tree may be defoliated. These trees
do not die readily from defoliation, however, because of their
ability to refoliate within the same season. Normally, during out-
breaks the affected trees are characterized by thin foliage, re-
duced radial and terminal increment, reduced production of nor-
mal shoots with a tendency toward adventitious growth, and
branch mortality. Marked loss of radial increment occurs after
4 to 6 years of outbreak, and after 6 to 9 years of moderate to
heavy defoliation, tree mortality occurs. Many widespread out-
breaks have been recorded since 1880 and losses have been severe.
Since the middle fifties, an estimated volume loss of 40 percent
in valuable sawtimber and pulp stands has occurred in managed
forests in Minnesota. Between 1910 and 1926 in Minensota an
estimated one billion board feet was killed.

Overwintering cocooned sawfly larvae are destroyed in enor-
mous numbers by small mammals, especially shrews and voles.
High surface water in tamarack bogs also destroys large numbers
at times. Many species of parasites occur but only two, the im-
ported ichneumon, Mesoleius tenthredinis Morley, and the tachi-
nid, Bessa harveyi (Townsend), have been prevalent during the
current outbreak which began in about 1938. Initially, M. tenth-
redinis was quite effective in control; however, the sawfly de-
veloped an immunity to it in central Canada, Minnesota, and
Wisconsin (208).

458

The mountain ash sawfly, Pristiphora geniculata (Hartig), a
probable introduction from Europe, was first recorded in North
America at Haines Falls, New York, in 1926 (635). It is now
known to occur in New Jersey, New York, Pennsylvania, Michi-
gan, all of the New England States except Rhode Island, and in
all eastern provinces of Canada west to Ottawa, Ontario. Its prin-
cipal hosts are American and European ash. It also occurs occa-
sionally on showy mountain ash and the hybrid, Sorbaronia hy-
brida. 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. 189).
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. Pu-
pation 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 lar-
vae straddle the edge of the leaf and feed around the periphery.
When disturbed they raise their abdomens in the form an “’S”.
Larvae of the first two instars are gregarious. Later, especially
in the fourth and fifth instars, they often feed singly. They usu-
ally 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 top soil. About 20 percent of
these pupate and appear as second generation adults in July
(257).

The mountainash sawfly is primarily a pest of shade and orna-
mental trees. The esthetic values of these trees is seriously re-
duced by defoliation, but they usually survive even when com-
pletely defoliated.

Other species of Pristiphora occurring in eastern forests in-
clude P. chlorea (Norton) on oak, P. siskiyouensis Marlatt on
birch, and P. sycophanta Walsh on willow.

The yellow-headed spruce sawfly, Pikonema alaskensis (Roh-
wer), is widespread, occurring from Maine and New Brunswick
to Idaho, British Columbia and Alaska. Its hosts are white, black,
red, blue, Norway, and Engelmann spruces (772). Full-grown
larvae (fig. 190) are olive-green above and lighter green below
and are about 18 mm. long. The head is chestnut-brown or red-
dish-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.

In Maine, winter is spent as full-grown 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 on tender bark of
the stem between needles. Young larvae feed on the new needles.
Older ones move on to and devour the old needles. While feed-
ing, their rear ends curve downward or upward. The larvae be-
come full-grown in July and drop to the ground. Here they spin

459

Ps 4 5 i ae ; ox ne te,
iE a ‘an « ‘ A! 3 ee — > —
en We

COURTESY CONN. AGR. EXPT. STA.

FIGURE 188.—Larvae and cocoons of

the larch sawfly, Pristiphora erich-
Sonii.

‘

F-495242

FIGURE 190.—Dorsal and lateral

views of larvae of the yellow-

headed spruce sawfly, Pikonema
alaskensis.

F-519576

FIGURE 189.—Larvae of the moun-

tain ash sawfly, Pristiphora
geniculata.

cocoons in the duff or top soil. There is one generation per year
(554).

The yellow-headed spruce sawfly is most injurious to shade
trees and ornamentals, to nursery stock, and to young trees in
plantations, as well as naturally regenerated cutover areas, in
shelterbreaks, and windbreaks. Open-grown trees older than 5
years are especially susceptible to injury, the majority being
killed by 3 or 4 consecutive years of moderate to heavy defolia-
tion. Trees in natural or closed stands are practically immune to
attack and damage.

The green-headed spruce sawfly, Pikonema dimmockii (Cres-
son), also occurs from coast to coast in southern Canada and
from Maine to Idaho, and feeds on various species of spruce.
Populations are usually low and apparently cause little injury.

460

The willow sawfly, Nematus (=Pteronidia) ventralis Say, oc-
curs in southeastern Canada and south and west in the Eastern
States to Georgia 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
top soil beneath the trees. Pupation and adult emergence occurs
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 are usually one
to five generations per year depnding on location. In the northern
areas some individuals may remain in diapause up to 20 months
before completing their development.

The willow sawfly occasionally heavily defoliates willows in or-
namental plantings and along streams. Basket willows in the
South have been damaged severely.

Nematus salicisodoratus Dyar occurs in the Eastern States
west to Illinois. 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 longitudinal 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 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 North-
eastern States.

Approximately 40 additional species in the genus Nematus
have been recorded from eastern forests (Muesebeck, et al.).
Known hosts of a few species are as follows: black locust—N.
abbotti (Kirby) and N. tibialis Newman, white birch—WN. hya-
linus (Marlatt) and N. pinguidorsum Dyar, poplar—N. fulvicrus
Provancher and N. populi (Marlatt), hophornbeam—WN. ostryae
(Marlatt), and willow—N. limbatus Cresson.

Trichiocampus irregularis (Dyar) occurs in southeastern Can-
ada and southward through the New England States, and feeds
on willow. Full-grown larvae are yellow, sparsely clothed with
hairs, and about 18 mm. long. The head is almost entirely black
and there are two rows of black spots on each side of the body.
The related species, T. gregarius Dyar feeds on poplar in the
Eastern States.

Trichiocampus viminalis (Fallen), an introduced species, oc-
curs across southern Canada and the Northern States. Its hosts
are various species of poplar and 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
smaller (fig. 191).

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.

461

; . : d COURTESY CONN. AGR. EXPT. STA.
FIGURE 191.—Larvae of Trichiocampus viminalis on leaves of poplar.

Older ones tend to scatter and eat all but the midribs and larger
veins. They become full grown in June or early July. Some pu-
pate and give rise to a second generation of adults in August;
the remainder do not complete their development until the follow-
ing spring. Carolina and Lombardy poplars grown for ornamental
purposes may be seriously defoliated.

The dusky birch sawfly, Croesus latitarsus Norton, occurs in
eastern Canada and south through the Eastern States to Florida.
Its hosts are various species of birch, preferably gray birch. It
has also been found feeding on alder and cherry in Canada. 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 subspirac-
ular area.

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 overlapping generations per year and lar-
vae may be found from spring until fall. The larvae are gre-
garious and feed along the edges of the leaf (fig. 192). Colonies
are often found defoliating small saplings, but this usually does
not occur over wide areas.

COURTESY CONN. AGR. EXPT. STA.

FIGURE 192.—Larvae of the
dusky birch sawfly, Croe-
sus latitarsus.

462

ae ee

The striped alder sawfly, Hemichroa crocea (Fourcroy), prob-
ably an introduced species, occurs from coast to coast in southern
Canada and northern United States. Its hosts include various
species of alder and occasionally birch. 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
and sand. 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 mid-
rib and larger veins of the leaf. They become full grown in July,
and, during late July and August, adults of a second generation
appear. Larvae of this generation are found during August and
September. There are two generations per year. This species
occasionally severely defoilates alder in the Lake States, North-
astern States, and Canada.

The maple petiole borer, Caulocampus acericaulis (MacGilliv-
ray), an introduced species, is known to occur in the Northeastern
States south to New Jersey. The larvae feed by boring into and
tunneling the petioles of leaf stems of maple. Full-grown larvae
are light yellow with light brown heads and are about 8 mm. long.

Winter is spent in the larval stage in a cell 2 or 3 inches below
the surface of the soil. Adults appear early in May and deposit
their eggs near the bases of the petioles of maple leaves. The
larvae tunnel in the petioles until they break, usually near the
leaf blade, and the leaves fall (fig. 198). After the petioles break,
the larvae continue to feed for about a 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.

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 on them
during the summer.

The brown-headed ash sawfly, Tomostethus multicinctus (Roh-
wer), occurs in southern Canada and throughout eastern United
States west to the Great Plains. Its hosts 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 cocoon-like cells in the
top soil. 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 vora-
cious feeders that they may completely defoliate a heavily infested

463

P.

COURTESY CONN. AGR. EXPT. STA.
FIGURE 193.—Petiole of maple leaf severed by larvae of the
maple petiole borer, Caulocampus acericaulis.

tree in a week. Shade trees are especially subject to serious de-
foliation (448).

The black-headed ash sawfly, Tethida cordigera (Beauvois), oc-
curs throughout much the same area as the brown-headed ash
sawfly. The habits, food plants, and life histories of the two spe-
cies 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 blackish-brown. This species is
occasionally a pest of shade trees.

Eriocampa juglandis (Fitch) (previously misidentified as
Blennocampa caryae (Norton), the butternut woolly worm, occurs
in southern Canada and throughout the Northeastern States. Its
hosts are butternut, black walnut, and hickory. Full-grown larvae
are green, with indistinct black spots on the sides and are about
18 mm. long. The body is covered with flocculent white tufts
which rub off when touched, and the head is white with black eye
spots. The larvae feed gregariously, often causing considerable
defoliation locally. When they become full grown they move to
and enter the ground, where they form cocoons composed of par-
ticles of sand and earth cemented together. Winter is spent in
the cocoons and there is one generation per year.

The genus Hoplocampa Hartig is represented in eastern Amer-
ica by a number of species, all of which presumably feed as lar-
vae 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 eastern species and their known hosts are as
follows: H. oskina Ross—hawthorn in the Central States, H.

464

halcyon (Norton)—shadbush in the Northeastern States and
southeastern Canada, H. pallipes MacGillivray—shadbush in the
Lake States, H. lactetpennis Rohwer—choke cherry in southeast-
ern Canada and the Northern States, and H. montanicola Rohwer
—choke cherry in the Northern States and southern Canada.

The genus Anoplonyx Marlatt contains only eight species, two
of which occur in eastern 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 and A. lu-
teipes (Cresson), are green in color and range in length from
about 9 to 15 mm. The thorax is larger than the abdomen, pro-
ducing a humpbacked appearance and causing the body to taper
posteriorally (932).

The genus Huura Newman is represented in eastern United
States and Canada by 16 species, and all are gall makers on
various species of willow. Larvae are usually yellowish or green-
ish-white with black eye spots, and the head is often tinted
brown. All species apparently have one generation per year. Win-
ter is spent as full-grown larvae 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 en-
veloped in galls. When they become full grown, some of them
vacate the galls and drop to the ground to pupate; others remain
inside their galls. Galls usually consist of a somewhat woody
swelling of the twig. Sometimes an entire shoot is enlarged. In
other cases, the gall appears as an abrupt swelling on one side
of the shoot. These galls may cause economic loss when they occur
where normal twig development of willow is important.

The genus Macremphytus MacGillivray is represented in east-
ern forests by five species all of which feed as larvae on dogwood.
Full-grown larvae 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 pow-
dery secretion.

SUPERFAMILY SIRICOIDEA
FAMILY SIRICIDAE
HORNTAILS

Members of the family Siricidae are commonly known as horn-
tails 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 ovi-
positor of the female is long and fitted for boring. In this respect
it differs from the ovipositor of sawflies which consist of saw-
like 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
badly weakened condition. Horntail larvae are parasitized by sev-
eral species of ichneumon wasps of the genus Megarhyssa Ashm.
Two of these, M. atrata atrata and M. macrurus macrurus, are
of special interest because of their striking appearance. Using

465

their extremely long ovipositors, the females bore deep holes into
the wood and deposit their eggs on or near horntail larvae in
their galleries.

Horntail adults are medium to large in size and are usually
metallic blue or black in color. Some are vari-colored with com-
binations of black, red, and yellow. The head, thorax, and ab-
domen are of equal width; the wings are well developed; the an-
tennae 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 the prompt utilization or submersion in water of in-
fested logs and by kiln drying of green lumber sawed from in-
fested logs.

The genus Sirex L. 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.
S. abbottw Kirby has been recorded breeding in larch in New
York and Georgia. The blue horntail, S. cyaneus F., attacks spruce
and pine in southern Canada and the northern tier of eastern
States. S. edwardsi Brulle has been taken from pitch pine in the
Atlantic Coast States. S. nigricornis F. attacks shortleaf pine
from New York to Ohio and Florida.

The genus Urocerus Geoffrey is represented by three eastern
species. U. albicornis (F.), the white-horned 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 spe-
cies has also been observed attacking freshly sawed lumber. U.
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. U. cressont Norton,
the black and red horntail, occurs from eastern Canada to Georgia
and breeds in poplar.

The pigeon tremex, Tremex columba (L.), is the most common
of the horntails. It occurs throughout nearctic 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
an expanse 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 to a depth of about 12 mm.
in the wood to deposit her eggs. Although laid singly, several
eggs may be found near each other in a limited area. The larvae
feed by excavating tunnels entirely in the wood. This frequently
weakens the tree and leads to wind breakage. Pupation occurs at

466

the end of the larval tunnel and the adults emerge through circu-
lar holes about 8 mm. In diameter. There appears to be one
generation per year.

FAMILY XIPHYDRIIDAE

The family Xiphydriidae is represented in the United States
and Canada by only six species, all of which occur in eastern
United States. 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. Few, if any, are ever very abundant.

Eastern species and their known hosts and distribution are as
follows: Xiphydria abdominalis Say—basswood, maple, and elm
from southern Canada to North Carolina and Jowa; X. maculata
Say—maple in Canada and most of the United States except the
Gulf States; X. tibialis Say—elm, birch, beech, American horn-
beam, and hawthorn in southeastern Canada and the Northeast-
ern States; X. hicoriae Rohwer—hickory and elm in southeastern
Canada and from Massachusetts and New Jersey to Illinois; X.
mellipes Harris—beech, birch, and alder across southern Canada
and the Northern States.

FAMILY ORUSSIDAE

Eight species of the family Orussidae have been recorded from
the United States and Canada, five of which occur in eastern
United States. The adults are 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 of the coleopterous family Buprestidae.

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 an-
tennae 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 and from New England to Virginia and South
Dakota. Its hosts are willow and poplar. Full-grown larvae are

467

tte. thet eee

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 cocoon-like structures.

The genus Periclista Konow is represented in eastern forests
by eight species which feed as larvae on the leaves of oaks and
hickory. The larvae are usually light green or have the dorsum
grayish, and are armed with rows of small, single, two-pointed
spines.

Suborder APOCRITA (—CLISTOGASTRA)

Members of the suborder Apocrita have the base of the abdo-
men constricted into a slender petiole or ‘‘waist.’’ The constricted
portion is comprised of the first abdominal segment 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;
others 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 are injurious to tree seed and cone
crops. Some species of ants are destructive of young trees in nur-
series, plantations, and natural regeneration areas, while others
are destructive of finished wood products.

SUPERFAMILY ICHNEUMONOIDEA

This superfamily constitutes one of the largest groups of para-
sitic insects and from the point of view of effectiveness in holding
in check the numerous pests that infest plants, it probably takes
first rank (144). It is comprised of three families (Stephanidae,
Braconidae, and Icheneumonidae), and these have been broken
down into more than 30 subfamilies, hundreds of genera, and
thousands of species. Only a small portion of the important species
parasitic on forest insects are mentioned. A complete listing of
the species known to occur in the United States and Canada has
been published by Meusebeck, et al. (550) and Krombein (442).

FAMILY STEPHANIDAE
This very small family of rather rare insects is represented in

eastern forests by only one genus and three species. They are
usually collected on dead trees and all are presumed to be para-

468

sitic on wood-boring Coleoptera. The adult is odd looking, having
a spherical head situated at the end of a long neck, bearing
crowns of teeth.

FAMILY BRACONIDAE

The family Braconidae comprises one of the major groups of
insect parasites. The majority of species are parasitic in the
larvae of Lepidoptera, but a large number are also parasitic 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 (144).

In addition to the large number of native species attacking
forest insects, several species have been imported from abroad
against important introduced pests. A few of these are discussed
briefly below.

Agathis (=Bassus) pumilis (Ratz.) was imported into the
United States and Canada against the larch casebearer during the

thirties. It is now widely distributed throughout eastern Canada

and Northeastern United States and is providing a high degree
of control in some areas. It has also been established in Idaho
where the casebearer was discovered on western larch in 1957.
Winter is spent in the larval stage within the host and there is
one generation per year.

Meteorus versicolor (Wesm.), a parasite of the brown-tail
moth and various other species of Lepidoptera, was introduced
into the United States early in the century, and is known to have
been established since 1909. It is now widely distributed through-
out the range of the brown-tail moth in New England, but ap-
pears to be of little value in its control. 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 (205).

Apanteles lacteicolor Vier., a parasite of the brown-tail moth
in Europe, was introduced and established in the United States in
1908, and is now generally distributed throughout the range of
its host. Studies made several years after its establishment
showed a parasitization of 7 percent of overwintering larvae over
a wide area. Winter is spent within young brown-tail moth larvae.
The adults appear in the spring, and two generations may de-
velop in alternate hosts (549, 124).

Apanteles melanoscelus (Ratz.), a European parasite of the
gypsy moth, was introduced into New England in 1911 and 1912
and became established very quickly. Later, it was recolonized
widely and is now generally established over the infested area of
New England. Parasitization is sometimes fairly high in spots.
Its abundance is greatly reduced in the spring by hyperparasites
which attack overwintering cocoons.

469

Apanteles solitarius (Ratz.) was introduced into New England
against the satin moth in 1927 and soon became established
throughout the infested area. Parasitization as high as 67 per-
cent has been recorded. The winter is spent either as a first-instar
larva in a satin moth larva or as a prepupal larva in a satin moth
cocoon. Thus, two separate broods of adults arise from the over-
wintering generation, each of which produces a second genera-
tion during the summer (577).

Phanomeris phyllotomae Mues. was imported into New Eng-
land and New York in the early thirties against the birch leaf-
mining sawfly. It became established but its effectiveness in
control has not been determined. There is one generation a year
(204).

Orgilus obscurator (Nees) was imported by the United States
and Canada in the thirties against the European pine shoot moth
and is now widely distributed in most shoot moth infested areas.
Percentage parasitization is quite variable but often high in the
United States, ranging 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 instar larva within the
hibernating host larva and there is one generation per year.

FAMILY ICHNEUMONIDAE

ICHNEUMONS

There are approximately 2500 described species of ichneumons
in the United States and Canada, and probably at least 5,000
undescribed ones (716). The family has been divided into 14 sub-
families and 53 tribes and subtribes (550). All members of the
family are parasites of the larvae and pupae of holometabilous
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 several
species of Coleoptera are also 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 tro-
chanter two-segmented, in having the female ovipositor perma-
nently extruded, and in lacking a costal cell in the front wing.
They also resemble adult braconids but differ in having two
recurrent veins in the front wing instead of just one.

Forest and shade tree insects are parasitized by a great many
species of ichneumons, far too many to mention here (550). 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 (205).

Aptesis basizonia (Grav.), a European parasite of sawfly co-
coons, was introduced originally into Canada against the Euro-
pean spruce sawfly during the thirties, and several colonies were
made available for liberation in the United States as well as
Canada against both the European spruce sawfly and the Euro-
pean pine sawfly. It became established on the European pine

470

sawfly, which it parasitizes heavily at times. It has not been
recovered from the spruce sawfly in the United States.

Exenterus abruptorius (Thnb.), an important parasite of the
European pine sawfly in Europe, was imported into Canada during
the thirties against the European spruce sawfly and European
pine sawfly. Colonies were later made available for release in the
United States. Recoveries were made from the pine sawfly sev-
eral years after liberation in New Jersey.

Mesoleius tenthredinis Morley was imported into Canada from
England in 1910-11 and liberated against the larch sawfly. A
colony was also released against the sawfly in the Great Lakes
region. It became established and for many years effected con-
siderable control. Gradually though, the host developed an im-
munity to the parasite in many portions of its range, thereby
greatly reducing its effectiveness. So far, this immunity does not
seem to have developed in the more easterly infestations into
which the parasite has spread.

Pimpla turionellae (L.), a European parasite of the pupae of
many Lepidoptera, was imported against the European pine shoot
moth during the thirties. It has not been recovered in the United
States but is apparently established in southern Ontario.

Temelucha interruptor (Grav.) was imported from England
and Europe against the European pine shoot moth during the
thirties. 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.

Eastern forest insects also are attacked by numerous native
species of ichneumonid parasites. Itoplectis conquisitor (Say)
parasitizes a tremendous number of species, and is especially
important. I. 4-cingulatus (Prov.) and I. viduata (Grav.) also
attack a great many species. Megarhyssa lunator (F.) is a con-
spicuous parasite of horntail larvae. The female is equinped with
an extremely long ovipositor which is used as a drill for laying
eggs in horntail larval galleries deep in infested wood. The ovi-
positor often becomes caught in the wood and the female, unable
to escape, dies.

SUPERFAMILY CHALCIDOIDEA

This superfamily contains more species than any other suver-
family in the order Hymenoptera. It contains, among its families,
probably a majority of all entomophagous insects, with an ex-
tremely wide range in form, habits, host preferences, and host
relationships. These comprise the majority and are mostly bene-
ficial, but there also are a number of phytophagous species, many
of which are economic pests. The species occurring in the United
States and Canada are divided into 21 families and more than
460 genera (588).

Practically all of the more common orders of insects serve as
hosts for the parasitic and predacious members of the Chalci-
doidea, with the Lepidoptera, Diptera, Coleoptera, and Homoptera
being preferred. The majority of the injurious tree-infesting

A471

ae in this country are those which destroy the seeds of their
osts.

Chalcids are mostly small to minute in size—some are less than
one-fourth of a millimeter 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, and the ovipositor issues some distance before the apex of
the abdomen.

FAMILY MYMARIDAE

This family 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 yellowish in color and are extremely minute in size,
usually less than 1 mm. long.

Polynema striaticorne Gir. is an important parasite of several
species of membracids. It also attacks various aphids and other
insects. Balduf (30) discussed its bionomics in Ohio and Illinois.
Acmopolynema bifasciatipennis bifasciatipennis (Girault) para-
sitizes the eggs of several species of tree crickets. Ooctonus aphro-
phora Milliron attacks the Saratoga spittlebug.

FAMILY TRICHOGRAMMATIDAE

The family Trichogrammatidae consists of extremely small in-
sects all of which are internal parasites in the eggs of other in-
sects. Hosts have been recorded from the orders Lepidoptera,
Coleoptera, Hymenoptera, Neuroptera, Diptera, and Hemiptera,
but the Lepidoptera are preferred. Muesebeck, et al. (550) list 40
species as occurring in the United States and Canada.

Trichogramma minutum Riley parasitizes the eggs of a great
many species of insects, including many important enemies of
trees. The following is only a partial list of important eastern
hosts: locust leaf miner, gypsy moth, brown-tail moth, orange-
striped oak worm, the saddled prominent, walnut caterpillar,
satin moth, hickory shuckworm, European pine shoot moth, Nan-
tucket pine tip moth, forest tent caterpillar, spruce budworm (as
many as 75 percent of spruce budworm eggs may be attacked),
black-headed budworm, bagworm, elm sawfly, fall webworm,
yellow-headed spruce sawfly, and the fall cankerworm (588).

The adult is less than 14 mm. long, and it inserts its eggs
directly into host eggs. During warm weather, the life cycle may
be completed within 9 to 16 days and there may be a dozen or
more generations per year.

FAMILY EULOPHIDAE

Adults of this family are very small, ranging in length from 1
to 3 mm. Well over 100 species are known to parasitize tree-
infesting insects, several of which are important pests (588). A
number of foreign species have been imported into the United
States and Canada in efforts to control several species of intro-
duced pests.

A472

i 8 eee

Kratochviliana (=Chrysocharis) laricinellae (Ratz.) (fig.
194), a parasite of the larch casebearer and the birch leaf-mining
sawfly was introduced into New England and Canada in the late
twenties and thirties. It now appears to be widely established.
Adults are bright, metallic green with pale yellow legs, and are
about 2 to 3 mm. long. There may be 8 generations per year in
the casebearer, but there are only one and a partial second in the
sawfly. This species is not an effective parasite of the casebearer
because it sometimes also acts as a secondary parasite and attacks
the more effective introduced parasite, Agathis pumilis (204).
Its effectiveness in controlling the birch leaf-mining sawfly is
unknown.

Dahlbominus fuscipennis (Zett.), a parasite of several species
of sawflies in Europe, was introduced into 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 red-headed pine sawfly in Michigan, the Virginia pine sawfly
in Virginia, Neodiprion pratti paradoxicus and N. pinirigidae in
New Jersey, the red-pine sawfly in Wisconsin, the balsam-fir saw-
fly in Maine, and Diprion frutetorum in Connecticut (205).

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 saw-
fly pupae within their cocoons. There are from two to seven gen-
erations per year, depending on location. It has never become a
very effective parasite of the spruce sawfly. Yet, almost 50 per-
cent parasitization of the European pine sawfly has been recorded
locally in New Jersey.

Coccophagus insidiator (Dalman), a European parasite of the
introduced European elm scale was discovered at Ithaca, New
York, in 1924 where it was parasitizing the scale quite heavily.
It had apparently been introduced into the area by accident. The
female is an endoparasite of the scale; the male an ectoparasite

F-519579

FIGURE 194.—Adult of the

parasite Kratochviliana
laricinellae.

473

of the larval stage of the female parasite. There may be three to
five generations per year.

The genus Tetrastichus contains a large number of important
parasites of important forest insect pests. Descriptions of most
North American species and information on their distribution
and hosts were published by Burks (125).

Tetrastichus turionum (Htg.) was imported during the thirties
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 is known to occur in Massachusetts, New
Jersey, on Long Island, 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 control of the shoot moth in
this country.

Tetrastichus brevistigma Gahan is a pupal parasite of the elm
leaf beetle. Adults are black with a slight, metallic, greenish tinge
and are only about .5 to 1.5 mm. long. An average of 12 parasite
larvae develop in each pupa and there are three or four genera-
tions per year. Parasitization as high as 50 to 80 percent occurs
commonly in the vicinity of Boston, Massachusetts, where it was
introduced from California. T. holbeini Gir. and T. rugglest Roh.
attack several species of Chrysobothris and Agrilus, respectively.
Dimmockia incongrus (Ashm.) parasitizes the gypsy moth and
many other species of Lepidoptera. Elachertus cocoeciae (How.)
parasitizes various species of Lepidoptera. Members of the genus
Hyssopus Gir. parasitize several species of shoot and tip moths
and seed and cone infesting insects.

FAMILY ELASMIDAE

This family is represented in North America by one genus and
17 species, all of which occur in eastern United States. They are
parasitic on lepidopterous larvae or hyperparasitic on ichneu-
monid or braconid parasites of lepidopterous larvae.

FAMILY THYSANIDAE

This family is also represented by one genus and 17 species in
the United States and Canada. As far as known, they either at-
tack various species of scale insects, whiteflies, and other Homop-
tera, or they attack other chalcid parasites of Homoptera.

FAMILY ENCYRTIDAE

This is a large family of parasites. Insect hosts are widely dis-
tributed among the various orders, but the majority of species are
parasitic on aphids, scales, and whiteflies. Adults are 1 to 2 mm.
long.

Ooencyrtus kuwanai (Howard), a parasite of gypsy moth eggs,
was introduced into the United States in 1908 and 1909 from
Japan (175). The adult is black and about 1 mm. long. Winter is
spent in the adult stage, during which mortality may be severe.

474

Surviving adults appear during April and lay their eggs in the
overwintering eggs of the host. There may be one or two spring
generations; for the entire year there may be four or five genera-
tions. This species is an important parasite of the gypsy moth in
the southern portion of its range. Parasitization of 40 to 50 per-
cent frequently occurs in Massachusetts and Connecticut.

Habrolepis dalmani (Westw.) is sometimes fairly common as a
parasite of the golden oak scale in this country. It has been intro-
duced into New Zealand from New England and is credited with
having saved the oaks in that country. Kermes pubescens, another
oak scale, is attacked by at least five other species of encyrtids.
The European fruit lecanium is attacked by more than 20 dif-
ferent species. The San Jose scale and oystershell scale each are
also attacked by several species.

FAMILY KUPELMIDAE

Members of the family Eupelmidae parasitize a wide range of
insects including Coleoptera, Orthoptera, Diptera, Lepidoptera,
Hymenoptera, Hemiptera, and Homoptera. 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 into New England in 1906,
and quickly became established (175). It is now generally dis-
tributed throughout the infested parts of New England. Adult
females are marked with green and brown, have two broad fus-
cous 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 full-grown 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 de-
velopment is rapid, the hibernating stage being reached within
about 2 weeks. Yet, there is only one generation per year. This
species is usually scarce in areas where the egg parasite, Ooen-
cyrtus kuwanat, also occurs. Interspecific competition may be the
cause.

A few other species of eupelmid parasites and some of their
hosts are as follows: Eupelmella vesicularis (Retzius)—gypsy
moth, satin moth, forest tent caterpillar, European pine shoot
moth, and several species of pine sawflies; Ewpelmus cyaniceps
amicus Girault—Dioryctria disclusa, Nantucket pine tip moth,
bagworm, and the smaller European elm bark beetle; H’. cyani-
ceps cyaniceps Ashm.—European pine shoot moth, Nantucket pine
tip moth, and the bagworm; E. pini Taylor—white pine weevil;
E. allynit (French)—various species of tree crickets; and Meta-
palma spectabile Westwood—buprestid and cerambycid borers.

FAMILY EUCHARITIDAE

The family Eucharitidae is comprised of only six genera and
27 species. As far as known, all species are parasitic on the pupae
of ants. Adults are distinguished by the configuration of the scu-
tellum, which is frequently produced backward in the form of
powerful spines.

A75

FAMILY PERILAMPIDAE

This is a small family with only two genera and 31 species in
the United States and Canada. Its members are secondary para-
sites of various Diptera, Hymenoptera, Orthoptera, and Neurop-
tera. The pupae of tachinid, braconid, and ichneumon parasites
are especially attractive.

Perilampus hyalinus Say, a common species throughout the
United States, is 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 bright, metallic,
bluish-green and is from 2 to 4 mm. long. The thorax is large and
the abdomen triangular. Peck (588) lists more than 20 species of
Orthoptera, 16 species of Lepidoptera, 12 species of sawflies, and
many species of ichneumonid, sarcophagid, and tachinid parasites
as hosts. Smith (657) discussed its biology as a secondary parasite
of the fall webworm.

FAMILY TORYMIDAE

Most members of this fairly large family are parasites of gall-
forming insects. A number of others are parasitic on various
lepidopterous larvae; quite a few feed in the seeds of various
plants; and some act as hyperparasites.

The genus Torymus Dalman contains a large number of species
that parasitize the immature stages of gall-forming cynipids and
cecidomyids. T. rugglesi Milliron has been collected from the
seeds of American holly in Delaware.

Members of the genus Megastigmus Dalman are all phytopha-
gous, developing in the seeds of plants. Eastern species include
M. amelanchieris Cush.—on shadbush; 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.

The genus Monodontomerus contains several important para-
sites of various species of Lepidoptera and sawflies. M. dentipes
(Dalman), a European species which probably entered this coun-
try with its European host, the introduced pine sawfly, is now
widely distributed in southeastern Canada and in northern States
from Maine to the Lake States. It is not only an effective parasite
of the introduced pine sawfly, but also an important parasite of
several other sawflies attacking conifers. 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.

Monodontomerus aerus Walker was introduced into New Eng-
land from 1906 to 1910 against the gypsy and brown-tail moths.
It was released originally as a primary parasite but is much more
common as a hyperparasite, attacking both hymenopterous co-
coons and tachinid puparia. It is also parasitic on the white-
marked tussock moth and the eastern tent caterpillar.

476

Other species of Monodontomerus recorded as parasites of im-
portant forest insects in the Eastern States include M. indiscretus
Gahan—on the birch leaf-mining sawfly; M. montivagus Ashm.—
on the spruce budworm; M. minor (Ratz.)—on the eastern tent
cateripillar, the cecropia moth, and the spruce budworm; and
M. japonicus Ashm., an introduced species—on the European
spruce sawlly.

FAMILY PTEROMALIDAE

The family Pteromalidae is the largest in the superfamily and
its members act as parasites or hyperparasites of almost all
orders of insects. The adults are minute, black or metallic green
or bronze insects. Many have a more or less triangularly shaped
abdomen.

Schizonotus sieboldi (Ratz.), a primary parasite of the im-
ported willow leaf beetle and several allied species of Chrysomel-
idae, is widely distributed in eastern United States. The larva
feeds externally on the pupa of its host. High percentages of par-
asitization were recorded over a 8-year period in the vicinity of
Boston, Massachusetts, but host populations were not materially
reduced (203).

Dibrachys cavus (Walker), a hyperparasite of many primary
parasites, occasionally acts as a primary parasite. It is extremely
destructive of 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

This family contains a number of both phytophagous and en-
tomophagous species. 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 para-
sites 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 full-grown
weevil larvae and 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 the
following: Eurytoma verticillata (F.)—on the European pine
shoot moth; E. pini Bugbee—on the European pine shoot moth,
Nantucket pine tip moth, bagworm, and Rhyacionia rigidana,; E.
tylodermatis Ashm.—on the hickory shuckworm, European pine
shoot moth, and Nantucket pine tip moth; and BE. magdalidis
Ashm.—on the southern pine beetle and Pityophthorus

liquidambarus.
ATT

FAMILY CHALCIDIDAE

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;
a are solitary in habit; and practically all develop inside their

osts.

A few of the more important species and their hosts are: Halti-
chella rhyacioniae Gahan—the Nantucket pine tip moth, Rhya-
cionia rigidana, and the western pine tip moth; H. wanticles
(Walk.)—the European pine shoot moth and the oak skeletonizer ;
Phasgonophora sulcata Westw.—the bronze birch and the flat-
headed apple tree borers; Trigonura elegans Prov.—several
species of woodborers including the flatheaded apple tree borer,
the red elm bark and black elm bark weevils; Spilochalcis albi-
frons (Walsh)—numerous hosts including the locust leaf and
arborvitae leaf miners, and the larch casebearer; S. flavopicta
(Cresson)—the Nantucket pine tip moth; S. mariae (Riley) —
the bagworm and several species of Saturnid moths; Brachymeria
compsilurae (Cwfd.)—tachinid flies, especially Compsilura con-
cinnata and Blepharipa scutellata (202); and B. ovata (Say)—a
wide variety of lepidopterous hosts including the white-marked
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. Weld (754) divides
the superfamily into four families—Ibaliidae, Liopteridae, Figiti-
dae, and Cynipidae. The liopterids are mostly exotic, and their
habits are unknown.

FAMILY IBALIIDAE

Members of this family are all parasitic on horntails of the
family Siricidae. Four species are recorded from eastern United
States—Ibalia anceps Say, I. ensiger Norton, I. maculipennis
Haldeman, and I. scalpellator Westw.

Ibalia maculipennis Hald., a common species in eastern United
States and southeastern Canada, is a parasite of the pigeon tre-
mex. 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
This family is represented in eastern America by 30 species.

The majority are parasitic in dipterous puparae; a few are para-
sitic in the cocoons of chrysopids.

A478

FAMILY CYNIPIDAE
THE GALL WASPS

This large family of several hundred species consists largely of
species which produce galls on their plant hosts in which they live
and feed during the larval stage. Many others live in galls pro-
duced by other insects, and a few are parasitic on other insects.
The adults are small to extremely small and usually black. They
are distinguished by the abdomen which is oval, shining, some-
what compressed, and almost covered by the first tergite. Many
gall-forming species produce two quite different generations per
year. One generation develops during the summer in one type of
gall. Adults appear in the fall and consist entirely of partheno-
genetic females. Eggs laid by these females give rise to larvae
which produce 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 produce galls on oaks and are confined to them (419).
There are 717 species listed as occurring in the United States and
Canada, 76 percent of which form galls on oaks (550). |

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 which produce large irregular galls on the smaller
branches are capable of causing injury (fig. 195). Infested
branches may be disfigured or even killed. Occasionally, entire
trees are killed. On the other hand, galls produced by certain other
Species are economically valuable: Some have long been used in
the manufacture of ink and in dyeing and tanning; others serve
as a source of winter food for bees. One, the deciduous oak gall, is
occasionally abundant enough on black oak in Missouri to be used
as food for hogs, cattle, sheep, turkeys, and chickens (418, 420).
Some of the more common and important gall-forming species are
discussed below.

Callirhytis floridana (Ashm.) occurs from Virginia to Florida
and Missouri and Arkansas. It produces slender, elongate swell-
ings from 12 to 75 mm. long on branches close to the ground of
Chapman, post, and dwarf post oaks. Large acreages of dwarf
post oaks were nearly all killed during an outbreak in eastern
North Carolina in 19385 (420).

Callirhytis punctata (O. 8.), the gouty oak gall, occurs from
southern Canada to North Carolina and Illinois. It produces galls
(fig. 196) on the twigs and smaller limbs of scarlet, pin, and
black oaks. These galls are about 12 to 38 mm. long and they fre-
quently occur so close together that they form practically con-
tinuous masses. This species has alternate generations. The first
produces small blisterlike galls on the leaves near the veins in the
spring. The second produces 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.

A479

OO ee |

Callirhytis cornigera (O. 8.), the horned oak gall, occurs from
southern Canada to Georgia and Iowa. It produces 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.

Calliirhytis (=Andricus) gemmaria (Ashm.), the ribbed bud
gall, occurs from Massachusetts to Florida, Illinois, and Texas. It
produces 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.

Callirhytis operator (O. 8S.) occurs from New England to Vir-
ginia and Texas and 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. C. futilis (O. 8.) produces globose,
grayish galls up to 414 inches long on the main roots of young
white oaks just below the ground line.

Neuroterus quercusbatatus (=batatus) (Fitch), the oak potato
gall (fig. 197), occurs from Ontario and Rhode Island to Florida

F-520107

FIGURE 195.—Heavy infesta-

tion of galls on twigs of

Nuttall oak in the Missis-
sippi River Delta region.

480

and 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 (Bass.), the oak flake gall, produces 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 (Bass.), 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.

Amphibolips confluenta (Harr.), the large oak-apple gall (fig.
198), 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 quite large, from 12 to 50 mm.
in diameter, and greenish to brownish in color, depending on age.
The related species, A. fulginosa Ashm., produces globose galls
on the sides of willow and laurel acorn cups in Florida.

COURTESY ILL. NAT. HIST. SURV.
FIGURE 196.—Gouty oak galls
caused by Callirhytis punctata.

COURTESY AMER. MUS. NAT. HIST.

FIGURE 197.—Oak potato gall on

white oak caused by Neuroterus
quercusbatatus.

481

COURTESY ILL. NAT. HIST. SURV.

FIGURE 198.—Large oak-ap-
ple gall caused by Amphi-
bolips confluenta.

Dryocosmus quercuspalustris (O. 8.) (=Callirhytis palustris) ,
the succulent oak gall, occurs from southern Canada and New
England to Iowa and Florida. It produces somewhat circular, suc-
culent 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.

Xanthoteras forticorne (O. 8.) (=Biorhiza forticornis), the
oak fig gall, produces galls on the leaves, twigs, and stems of
white oaks in the Eastern States north of Virginia. Early in the
season they are reddish and bladderlike. Sometimes they occur in
dense clusters several inches long around a twig. When they do,
the irregular masses look like pressed figs.

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 rather poorly 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 EVANIIDAE

Members of this small family of spider-like wasps are parasitic
in the egg capsules of cockroaches. Adults are about 10 to 15 mm.
long and are distinguished by very small, oval abdomens attached
by petioles to the propodeum considerably above the base of the
hind coxae. The majority of described species occur in eastern
United States.

482

FAMILY AULACIDAE

The family Aulacidae consists of slender ichneumon-like wasps,
with the female 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. Pris-
taulacus rufitarsis (Cress.) is a parasite of the hemlock borer and
poplar borer in eastern America; P. bilobatus (Prov.) is also
parasitic on the hemlock borer. Aulacus burquei (Prov.), A. digi-
talis Townes, A. lovei (Ashm.), and A. pallipes Cress are para-
sitic on various species of Xiphydria. Several species in the genera
Gasteruption and Rhydinofoenus are parasitic in bee’s and wasp’s
nests in twigs or wood.

FAMILY PELECINIDAE

This family appears to be represented in North America by
only one species, Pelecinus polyturator (Drury), a parasite of
May 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.

FAMILY CERAPHRONIDAE

A number of species in this family have been taken from colo-
nies of ants. Various others are known to be hyperparasitic on
various hymenopterous and dipterous parasites. One of the latter,
Conostigmus virginicus (Ashm.), is a parasite of Blepharipa
scutellata (P.-D.), an introduced parasite of the gypsy moth.

FAMILY DIAPRITIDAE

This is a large family of very small black wasps, most of which
appear to be parasitic on immature Diptera. Adults of most
species can be recognized by the location of the antennae, which
arise on a shelflike protuberance in about the middle of the face.
Psilus politus (Say) has been recorded as parasitic on the cherry
fruit fly. Trichopria tabanivora Fouts is a parasite of horseflies.
Apparently, only a small percentage of the species in the family
have been described.

FAMILY SCELIONIDAE

As far as known, all members of this large family of small in-
sects are parasitic in the eggs of other insects. Species attacking
some of the more important forest and shade tree insects are as
follows: Telenomus dalmani (Ratz.) and T. hemerocampae Wil-
cox—the white-marked tussock moth, T. californicus Ashm.—the
satin moth, T. geometriae Ashm. and T. alsophilae Vier.—the fall
cankerworm, T. bifidus Riley—the fall webworm, T. catalpae
Mues.—the catalpa sphinx, 7. dalmani—the hemlock looper, 7.

483

alsophilae—the elm spanworm, and T. clisiocampanae Riley—
forest tent and eastern tent caterpillars.

FAMILY PLATYGASTERIDAE

The majority of species in this family are parasites of Diptera,
especially of the families Cecidomyiidae and Tipulidae. A number
of others are also important as enemies of mealybugs and white-
flies. The introduced species, Allotropa burrelli Mues., an impor-
tant parasite of the Comstock mealybug, has been widely released
in eastern United States. Two native species, A. ashmeadi Mues.
and A. convexifrons Mues., also are important parasites of the
Comstock mealybug.

SUPERFAMILY CHRYSIDOIDEA
FAMILY CHRYSIDIDAE

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 either 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 into Massa-
chusetts from Japan in 1917-18 against the oriental moth, also an
introduced species. Recoveries were made the year following its
release (576), but it has not been recorded since that time.

FAMILY CLEPTIDAE

This family is represented in North America by two genera and
seven species. The genus Cleptes Latrielle contains six species, all
of which are western in distribution and all of which apparently
parasitize various sawflies. The genus Mesitiopterus Ashm. is
represented in eastern United States by the single species, M.
kahli Ashm., an egg parasite of the walkingstick.

SUPERFAMILY BETHYLOIDEA
FAMILY BETHYLIDAE

Members of the family Bethylidae are parasitic almost exclu-
sively on the larva 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 Scleroderma Latreille are
often parasitic on larvae of the family Cerambycidae. Additional
species of importance as parasites of forest insects and their hosts
are: Pseudobrachium mandibulare (Ashm.)—the black carpenter

484

ant; Perisierola punctaticeps (Kieffer)—the hickory shuckworm;
Goniozus electus Fouts—Nantucket pine tip moth; and G. longi-
nervis Fouts—the western pine tip moth.

SUPERFAMILY SCOLIOIDEA
FAMILY TIPHIIDAE
TIPHIID WASPS

The family Tiphiidae contains a number of important parasites
of scarabaeid larvae in the soil. As far as known, all species are
external parasites, usually feeding on the final larval 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 tnornata Say is one of the commonest and most impor-
tant parasites of white grubs in the United States. Infestations
are heaviest during the years when full-grown white grubs are
present. At other times, they are usually very low (788).

Tiphia vernalis Rohwer was introduced against the Japanese
beetle from Japan and Korea during the period 1924-1933. It be-
came established and now plays an important role in control of
the beetle in certain parts of the East, especially in Pennsylvania
(143). T. popilliavora Rohwer, also introduced against the Japa-
nese beetle during the twenties and thirties, is now well estab-
lished from New Hampshire to Virginia and Ohio. T. asericae
A. & J. was introduced against the Asiatic garden beetle and
Serica peregrine during the twenties. It is well established in
Pennsylvania.

FAMILY MUTILLIDAE
VELVET ANTS

Members of the family Mutillidae are commonly known as vel-
vet ants because 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-killer
ants.’ Adults are usually brightly colored with red, orange, or
yellow markings, and some of the females are 25 mm. or more
long. The majority of species are parasitic on ground-nesting bees
and wasps (518). A common species in the South, where members
of the family are most common, is Dasymutilla occidentalis occt-
dentalis (L.). Because of their large size, velvet ants are capable
of inflicting painful stings.

FAMILY SCOLIDAE

The family Scolidae is represented by about a dozen species in
eastern United States and, as far as known, all species are ex-
ternal parasites of Scarabaeid larvae in the soil. White grubs are
especially susceptible to attack. The adults are large, hairy, and
usually black except for spots or bands of yellow or red, and

A485

their wings are often dark brown with purple or green irides-
cence. Campsomeris annulata (Fab.), a native of China and
Japan, was introduced against the Japanese beetle in the twenties
but did not become established.

FAMILY SAPYGIDAE

As far as known, all members of this family are parasitic on
the larvae of leaf-cutting bees. Less than a dozen species are re-
corded from eastern United States. Adults are moderate in size,
short legged and usually spotted or banded with yellow.

FAMILY FORMICIDAE
THE ANTS

Ants are among the most abundant and widespread of all in-
sects and they are found in practically all terrestrial habitats.
The majority of species nest in the soil, while many others build
their nests in wood, in 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 leaf-cutting 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; and
in having the antennae elbowed, with the first segment greatly
elongated in the females and workers. They also differ from ter-
mites, 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 containing
from a few to several thousands of individuals each. A cvolony
consists of three castes—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 are usually smaller
than the females or males.

With the exception of the carpenter ants and certain leaf-
cutting 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 (277, 632, 649, 660).

The black carpenter ant, Camponotus pennsylvanicus (DeGeer),
nests in standing live and dead trees, rotting logs, and stumps,
telephone and telegraph poles, and the wood of houses and other
buildings. It is widely distributed in eastern United States and
Canada, occurring from North Dakota to Quebec and Ontario and
south to Texas and Florida. These are large ants (fig. 199), the
workers ranging in length from 6 to 13 mm. The body color is
typically black, but in some individuals the pleuron, petiole, and
legs are reddish. The gaster is covered with dense, long, appressed,

486

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.

Live trees are occasionally infested but usually only when the
ants are able to enter them through cracks, scars, knot holes, and
decayed or other faulty places. Once inside the tree, they remove
the faulty wood and often extend their galleries into adjacent
soundwood. A wide variety of trees such as poplar, cherry, white
and pitch pine, balsam fir, elm, willow, and red, white, scarlet,
black, and post oaks have been found infested. Infestations fre-
quently 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 wind-
breakage. The wood also may be rendered worthless for lumber or
pulpwood (fig. 200).

Houses also are often invaded by carpenter ants coming from
nests located nearby outdoors. Possibilities of this happening are
greatest where houses are located in the vicinity of trees, logs, or
stumps. Entry is usually gained through openings around the
foundation 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. They
differ in that they run across the grain, are sandpaper smooth,

COURTESY CONN. AGR. EXPT. STA.

FIGURE 199.—Black carpen-
ter ant, Camponotus penn-
sylvanicus: A, Adult
winged female; B, adult
winged male.

A487

F-504087

FIGURE 200.—Cross-section of an oak log showing galleries made by
carpenter ants.

and are kept free of frass. The presence of large black ants run-
ning about the house; swarms of large black, winged ants about
the house in the spring; piles of sawdust-like borings; slit-like
holes in woodwork such as window and door casings; and faint
rustling sounds in walls, floors, woodwork, and flush-panel doors
are all evidences of infestation in a house. 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. 201) (272).

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,

COURTESY CONN. AGR. EXPT. STA.

FIGURE 201.—Colony of carpenter

ants in a 2-foot section of a tele-

phone pole in winter. Arrow in-
dicates ground level.

488

|
y
[)
1

|

|

log, or stump and seal themselves in. Here they rear their first
broods of workers to maturity on salivary secretions. These
workers, being inadequately fed, are smaller than normal. Subse-
quent 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. The removal of all wood from near or under
a house site prior to construction, making certain that infested
wood is not brought into the house, and the cutting back of
branches in contact with the house are also helpful in preventing
infestations in buildings.

The red carpenter ant, Camponotus ferrugineus (Fab.), occurs
over much of the same portion of the United States as the black
carpenter ant, but it is apparently less common. It appears to
prefer wooded areas where it normally nests in or beneath well-
rotted logs and stumps. Under these conditions, it often extends
its galleries for considerable distances in the soil. Nests are also
found in dead, standing trees and apparently, rarely in houses.
The workers are about the same size as those of the black car-
penter 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 white cedar in Minnesota
(305).

The Florida carpenter ant, Camponotus 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 yel-
lowish hairs. This species builds its nests 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 wood-
work of porches, roofs, kitchen sinks, and paneling. Outdoors, the
ants feed largely on living and dead small insects, and on honey-
dew which they secure by tending aphids, mealybugs, and scales.
Indoors, they feed on such items as molasses, honey, and liver.

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 rasilis Wheeler occurs from South Carolina to
Florida and Texas, 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;

489

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 1 inch to over 5 feet 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 C. rasilis,
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 logs 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 south-
ern half of Florida and apparently nests in small colonies in rot-
ting 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 Missis-
Ssippi. Galleries are apparently limited to one year’s growth.
C. pylartes fraxinicola M. R. Smith nests in the dead branches of
various hardwods. It has been recorded from Mississippi.

The Texas leaf-cutting ant, Atta texana (Buckley), is a serious
pest of pine seedlings in east Texas and west-central Louisiana.
It also defoliates and damages a wide variety of other plants, in-
cluding orchard trees. Injury to pines is especially severe during
the winter when there is a dearth of other green foliage.

Texas leaf-cutting ants are rusty-red in color. The head is
strongly bi-lobed; 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 extra-
ordinarily long. The queen is about 18 mm. long; workers, from
1.5 to 12 mm. long.

Nests of the Texas leaf-cutting ant are constructed in the
ground, usually in well-drained stand or loamy soils and com-
monly on slopes facing the south or west (59). The interior of the
nest may reach a depth of 20 feet and its outer surface may
occupy up to one-tenth of an acre. It may contain a thousand or
more entrance holes. Nest areas are usually marked by many
crescent-shaped mounds, up to 5 to 14 inches high and a foot in
diameter. Each mound surrounds an entrance hole (fig. 202). The
nest consists of many cavities connected by narrow tunnels. There
are vertical tunnels that extend to mound openings and lateral
tunnels that lead outwards, sometimes for a hundred yards or

F-482847

FIGURE 202.—Mounds of the

Texas leaf-cutting ant,
Atta texana.

more. Above ground, sharply defined foraging trails sometimes
extend hundreds of feet 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.

The ants do not eat the foliage which they remove from plants.
Instead, they cut it up into small fragments, shape it into small
pellets, and carry it into their underground 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 which serves as the food of
the colony. During summer most of the foliage is brought into the
nest during the night; whereas, during fall, winter and spring
most of it is brought in during the day, unless it is too cold or
wet.

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.

A pelleted bait has given good control when scattered over por-
tions of nests where excavation mounds are concentrated (567).
The reader should consult Agricultural Handbook 331 for details
on the control method.

The Allegheny mound ant, Formica exsectoides Forel, is a
serious pest of young white pines, red pines, Scotch pines, red
cedars, and spruces in the Eastern and Northeastern States. It
nests in the ground and constructs mounds which may be up to
4 feet in height and 6 feet across. In forested areas, these mounds
are most often found in openings or along the edges of stands.
All vegetation, excepting large trees, may be destroyed in an area
40 to 50 feet 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 is reddish brown and about as wide as 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 vege-
tation. 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 shad-
ing 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 downward movement of foods
in the inner bark (590).

The life history of the Allegheny mound ant is not too well
understood, but it is known to forage for food from April to Sep-
tember 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,

A491

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
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, Jridomyrmex 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 (660). 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 honey-
dew produced by aphids, mealybugs, and scale insects. By foster-
ing and protecting the latter insects from many of their enemies,
they encourage the development of heavy infestations which 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
in color; 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 and when freshly crushed emit a stale, greasy,
musty odor. Nests occur in all sorts of places—in the soil, in rot-
ten wood, in cavities in trees, and in refuse piles, bird nests, bee-
hives, and other places. The number of ants present in well-estab-
lished infested areas is beyond comprehension. 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 some-
times a nuisance in houses. The odor of freshly crushed workers
resembles the odor of rotten coconut.

The genus Solenopsis Westw. is represented by seven species in
the United States, six of which occur in the Southeastern States.
Nests are usually constructed in the soil, but also sometimes in
rotten wood and in houses. The workers are aggressive and prac-
tically omnivorous. Several species are serious economic pests.
They may remove seeds from seed beds; kill young quail and
poultry and even small game; gnaw into vegetables, fruits, and
flowers; make large ugly mounds in fields, parks, playgrounds,
and recreational areas; and damage young nursery trees.

The imported fire ant, Solenopsis saevissima richteri Forel, an
introduced species, first officially recorded in the United States at
Mobile, Alabama, in 1930, now occurs from North Carolina to
Florida, Texas and Arkansas, exclusive of Tennessee. Workers
are about 2.8 to 6 mm. long and are usually reddish in color, ex-
cept for a blackish gaster. Queens are about 9 mm. long.

492

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 1 foot high and 2 feet across; some are up to 3
feet 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 60 nests per acre. Nests also are found
in rotting logs, around tree trunks, and occasionally under build-
ings. Green (312) discussed the biology of this species.

For more comprehensive treatments of ants, the reader is re-
ferred to publications by Wheeler (759), Smith (659), and
Creighton (174).

SUPERFAMILY VESPOIDEA

This superfamily is comprised of 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 gener-
ally feed on nectar, sap, or similar materials, whereas the larvae
feed on other insects or spiders. A few species are of some impor-
tance as pests of forest or shade trees; quite a number of others
may create problems when they occur in abundance in picnicking
areas, campgrounds, or other recreational areas.

FAMILY VESPIDAE
HORNETS, YELLOW JACKETS, AND POTTER WASPS

The family Vespidae contains the well known stinging wasps,
hornets, and yellow jackets. More than 350 species have been re-
corded in North America north of Mexico, about one-third of
which occur in 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 considerably in size and appearance,
but all are distinguished by the very long discoidal cell in the
forewing, 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. Bohart (87) published a synoptic cata-
log of species occurring in America north of Mexico.

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
in shape and are often quite large, measuring several inches in
diameter. Yellow jackets 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 at-
tached to a support by a slender stalk. They may be found in all
sorts of places such as under the eaves of roofs, under porches, in

493

open sheds, and on bushes, shrubs, and trees. The adults of all
members of the family feed commonly on nectar, ripe fruits, sap,
ee honeydew. The larvae feed on other insects provided by the
adults.

The giant hornet, Vespa crabro germana (Christ), an intro-
duced species, has been recorded from Quebec and Massachusetts
to South Carolina. According to reports, hollow trees, hollow
posts, sheds, barns, porches, and even attics are preferred as nest-
ing sites. These nests occasionally become very large, reaching a
length of 3 feet and a diameter of 20 inches. There are many
reports of damage to trees and shrubs by the adults (643). They
may girdle small twigs, and they gnaw holes in the bark of larger
branches, possibly in search of nest-building materials and pos-
sibly in search of sap. Injuries have been reported to lilac, birch,
ash, and horsechestnut. The adult possesses a long sting and a
large poison sac. It is probably capable of inflicting painful stings,
but does not appear to be as likely to attack when disturbed as is
the bald-faced hornet.

The bald-faced hornet, Vespula maculata (Linn.), a well known
member of the family, is widely distributed in the United States
and Canada. Adults are 12 to 19 mm. long and largely black, with
white or yellow 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 15
inches and contain up to five thousand hornets each. Bald-faced

COURTESY CONN. AGR. EXPT. STA.
FIGURE 203.—Nest of the yellow jacket, Vespula arenaria arenaria.

494

hornets attack at once when their nests are disturbed and ave
capable of inflicting extremely painful stings.

Vespula arenaria arenaria (Fab.) is probably the most common
species of yellow jacket in eastern United State It also occurs
throughout the remainder of the United States, and in Canada,
and Alaska north to the Arctic Circle. Adults a: 2 black and yel-
low with black predominating. The legs are yellow with the
femora mostly black, and the length is about 12 mm. Nests (fig.
203) are usually built in the ground, with an exit hole leading to
the outside. This species also inflicts severe stings and it often
attacks in force when its ne is endangered. It is a common
species around picnic tables where it is often a nuisance.

The genus Polistes Latreille contains the familiar wasps whose
nests are so frequently encountered hanging under the eaves of
buildings and in various and sundry other places. They differ in
appearance from the hornets and yellow jackets 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 some may be several inches in
diameter. The adults of a number of species collect lepidopterous
larvae as food for their young and, where abundant, are appar-
ently capable of exercising a considerable degree of control of
their hosts. Like hornets and yellow jackets, these insects are
vicious stingers, and their nests should be approached with
caution.

FAMILY POMPILIDAE

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 some 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 in-
sects. 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 which serves as food for their progeny. Host preferences are
varied, including spiders and most of the more common orders of
insects. Adults generally can be recognized by the structure of the
pronotum which does not extend back to the tegulae, by their un-
jointed trochanters, by the absence of dilation in the hind tarsus,
and by the simple pubescence of the head and thorax.

FAMILY AMPULICIDAE

This family is represented in North America by only two gen-
era and three species. As far as known, they nest in twigs, under
bark, and under litter on the ground. Their prey consists of im-
mature cockroaches.

FAMILY SPHECIDAE

This family is represented by 86 genera and scores of species in
the United States and Canada. The majority are solitary nest-
building wasps which provision their nests with other insects or
spiders. Many species nest in the ground; some construct nests of
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 presence
of an anal lobe and several closed cells in the hindwing, the
presence of seven exposed tergites on the abdomen of the male,
the failure of the hypopygium to enclose the sting in the female,
and the lack of dilation of the hind tarsus. In a few species, such
as the familiar dirt-daubers, the petiole of the abdomen is greatly
lengthened.

The cicada killer, Sphecius speciosus (Drury), which provisions
its nest with adult cicadas, is the most conspicuous member 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 and the ovipositor or stinger is long and curved.
Adults are usually present from midsummer to early fall and they
burrow into the ground for nesting purposes. The female para-
lyzes a cicada by stinging it and then carts 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 nests with a wide
variety of wood-boring buprestids. Species of the genus Psen
Latrielle prey on various membracids, cereopids, and leafhoppers.
Crossocerus ambiguus (Dahl) preys on various species of tree-
infesting cicadellids (184) ; Stictia carolina (Fab.) is frequently
seen hovering over livestock in search of horseflies.

FAMILY DRYINIDAE

This is a small family of rare insects. As far as known, all
species are parasitic on the nymphs of Homoptera, especially the
membracids, cicadellids, flatids, and cercopids. The females of
certain species 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 internal parasites of
the abdomen 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. Aphelopus theliae Gahan, a parasite of the membracid,
Thelia bimaculata, which feeds on young black locust, lays a
single egg in a nymph of its host. Polyembronic 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 (278).

496

SUPERFAMILY APOIDEA
BEES

It has been estimated that more than 2,500 species of bees occur
in North America, many of which, such as the honeybee and
bumble bee, are so common as to require no description. Fortu-
nately, none of these insects are particularly injurious to forest
or shade trees. On the contrary, as a group they are highly bene-
ficial 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 tarsi are dialated or thickened; and the head
and thorax are covered with feathery hairs.

FAMILY MEGACHILIDAE
LEAF-CUTTING BEES

Leaf-cutting bees are small to medium 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 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 non-injurious.

FAMILY APIDAE
BUMBLE, CARPENTER, AND HONEY BEES

The family Apidae is represented by more than a thousand
species in the United States and Canada. It has been divided into
three subfamilies—Anthrophorinae, Xylocopinae, and Apinae.

The subfamily Anthrophorinae 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, are parasitic in the nests of other bees. The adults are usu-
ally wasp-like in appearance. Members of the subfamily Xyloco-
pinae construct their nests in wood or plant stems and are
commonly known as carpenter bees. The genus Xylocopa Latreille
contains the carpenter bees. The adults look like bumble bees but
differ in having the dorsum of the abdomen largely bare. They
nest in solid wood. The subfamily Apinae contains the well-known
bumble bees and honey bees.

Considered as a whole, members of the family are far more
beneficial than harmful. Many species are highly efficient plant

497

|

pollinators, and the importance of the honey-producing honeybee
is too well-known for discussion. The only destructive members of
the family are the carpenter bees, several species of which attack
and damage wood in use.

The carpenter bee, Xylocopa virginica virginica (=virginica)
(Linnaeus), is an important pest because of its habit of tunneling
into the solid wood of beams, rafters, telephone poles, or struc-
tural timbers. This may lead to structural damage, especially
when the same piece of timber is attacked for several years. Dead
but sound wood of cypress, cedar, white and hard pines, and Cali-
fornia redwood that has been softened by weathering, 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 dis-
tance, then makes a right angle turn and follows the grain of the
wood for a distance of 6 to 8 inches. 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. Fe-
males spend the night 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.

Two other species of carpenter bees, Xylocopa virginica texana
Cresson and X. micans Lepeletier, also occur in eastern United
States. The former is found from Maine to Michigan and south to
Florida and the Gulf of Mexico. The latter occurs from North
Carolina to Florida and westward along the Gulf Coast.

INSECTS AND TREE DISEASES

Insects are involved in or are directly responsible for the trans-
mission of many serious diseases of forest and shade trees in
eastern United States. Some of these diseases, such as Dutch elm
disease and elm phloem necrosis, are among the most destructive
tree diseases known. The majority are caused by various species
of fungi. One is of virus origin. A few, such as Dutch elm disease
and elm phloem necrosis, attack living, healthy trees and kill
them quickly. Many stain the sapwood of coniferous trees, usually
of felled timber. They also occur occasionally in living trees
weakened by environmental conditions and hasten their death.
Others cause the decay of sapwood or heartwood of dead, dying,
and felled conifers.

Fungus diseases are transmited by insects through the direct
transfer of spores from infected to healthy trees or wood. The
bodies of insects emerging from diseased material become con-
taminated with the spores and when the insect attacks healthy
material spores are rubbed into the wounds. Wind-borne spores
also gain entry into healthy trees through insect-produced wounds.
Viruses, such as the one responsible for elm phloem necrosis, are
most often transmitted by species of sucking insects.

498

Dutch elm disease.—This disease is caused by the introduced
fungus, Ceratocystis ulmi (Buisman) C. Moreau. It was first re-
corded in North America in Ohio in 1930. Three years later it
was found in the New York City area. It is now known to occur
on the east coast from New England south to the Carolinas, Ala-
bama, and Mississippi and west to the Rocky Mountains (763).
Its hosts are listed as both native and exotic species of elm,
American elm especially. Its principal insect vectors are the
smaller European elm bark beetle and the native elm bark beetle
(150). Root grafting may also account for local tree-to-tree
spread, particularly among closely spaced street trees.

The first symptoms of Dutch elm disease are the wilting and
yellowing or drying of the foliage. This is followed immediately
by defoliation and death of the affected branches. Diseased trees
commonly die within a single growing season, but some may die
gradually, branch by branch, over a period of several years. Dis-
eased trees develop a brown discoloration in the water-conducting
vessels in the wood. In early spring, this occurs as brown streaks
in the wood just under the bark of diseased branches (fig. 204).
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.

Dutch elm disease is one of the most destructive tree diseases
known. It kills both wild and planted elms but is best known as a
killer of valuable shade tree elms. Since its discovery, it has killed
too many of these trees to count. Currently, the value of trees
killed and the cost of their removal is estimated at $100 million
each year. A considerable amount of research has been devoted to
the control of the bark beetles responsible for its spread (763).

F-519563
FIGURE 204.—Cross-section of an elm branch affected with Dutch elm
disease, showing brownish discoloration in outer ring of sapwood.

499

Elm phloem necrosis.—This is a virus disease of American and
winged elms (702). It occurs in the Midwest in an area bounded
by latitudes 30° and 40° North, and by longitudes 80° and 100°
West. The virus is transmitted by the white-banded elm leaf-
hopper, or through root grafts of trees growing close together.
The insect sucks up the virus while feeding on the sap of a dis-
eased tree. Later, the virus is transmitted when the insect feeds
on a healthy tree. This leafhopper is widely distributed in eastern
United States.

Foliar symptoms of elm phloem necrosis vary but are some-
what 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; usually, there is a gradual decline
of the tree. In large trees, there is usually a slight scarcity of
foliage in the top or at the outer tips of branches. This is followed
by a scarcity of foliage throughout the crown. The leaves turn
yellowish-green, then yellow, and some on the lower suckers be-
come dry and brown. Defoliation follows soon thereafter, and the
tree dies. This occasionally happens within 3 to 4 weeks but it
usually requires a year or a year and a half. Before the tree dies,
a typical discoloration of the phloem develops. 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 occa-
sionally in branches. The discoloration (fig. 205) is at first yellow.
Later, it turns to “butterscotch” and often contains scattered
dark flecks. Still later, the phloem becomes brown and necrotic.
Moderately discolored phloem has a faint odor of wintergreen, a

F-520108 F-519951
FIGURE 205.—Typical discolo- FIGURE 206.—Shade tree elms killed by
ration of the phloem of an phloem necrosis virus.

elm tree infected with elm
phloem necrosis.

500

distinguishing characteristic of the disease. Death follows the
dying of the roots.

Elm phloem necrosis is an extremely destructive disease. Dur-
ing epiphytotics such as occurred in the Central States in the late
thirties and forties, it has killed enormous numbers of valuable
shade tree elms in many cities and towns (fig. 206). More re-
cently, however, it has been at a low ebb.

Beech bark disease.—A destructive disease of American and
European beeches and all. their varieties is caused by the
fungus, Nectria coccinea var. faginata Lohman, Watson, and
Ayres. The pathogen gains entry into the tree through tiny rup-
tures in the bark caused by the feeding of the beech scale. The
scale, and probably the fungus, is of foreign origin. The disease,
which was first recorded at Halifax, Nova Scotia in 1920, now
occurs throughout the Maritime Provinces of Canada and south
through the beech-growing areas of New England to Central
New York and eastern Pennsylvania.

The fungus may infect large areas on scale-infested trees, com-
pletely girdling and killing them. Trees only partially girdled may
remain alive in a weakened state for many years, or they may be
broken by the wind. On some trees, the fungus may be confined to
strips or spots on the bole where it produces cankers. Parts of the
crowns of these trees become chlorotic and die.

The beech bark disease is destructive. In many stands it has
killed more than half the beech, and the commercial value of many
of the survivors has probably been seriously reduced. Shigo
(646) discussed possible methods of control.

Oak wilt.—This disease is caused by the fungus, Ceratocystis
fagacearum (Bretz). It occurs over a wide area, from Nebraska
and Kansas to the Carolinas and Pennsylvania, and, as far as
known, attacks all native species of oak regardless of size, age,
or vigor. The fungus spreads from infected trees to adjacent
healthy trees through natural root grafts, and there is consider-
able evidence that insects are responsible for spread over longer
distances: fungus spores have been recovered from the bodies
of several species of insects after they fed on fungal mats, and
some of these are known to feed on sap at fresh wounds on healthy
trees (116, 170, 314).

Oak wilt is potentially a very serious disease (fig. 207). Given
the proper conditions for an epiphytotic, it could cause heavy
losses in the vast oak forests of eastern United States. It has al-
ready caused considerable damage in many oak woodlots and
forest stands in Wisconsin, Minnesota, and Iowa. Methods of con-
trol are discussed by Fowler (260).

Persimmon wilt.—This fast-killing disease of common persim-
mon, Diospyros virginiana L. in the Southeastern States, 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
cambium region and are released when the bark begins to disin-
tegrate or is removed or broken off.

d01

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 contaminated when
they emerge through spore masses on dying trees. Feeding in-
juries on the terminals and twigs by adults of the twig girdler,
also serve as entry courts for wind-borne spores (172).

Blue stains.—In 1928, it was suggested that blue-staining fungi
were probably introduced into pines by bark beetles and that they
played important roles in the killing of trees (166). This was
later confirmed (564). Blue-staining occurs mostly in felled
timber but also may be found in beetle-infested trees weakened by
fire, drought, and other adverse factors.

Ceratocystis minor (Hedgec.) Hunt one of the most important of
the blue-staining fungi in eastern United States, is introduced into
pine trees by the southern pine beetle. It does not cause decay but
stains the wood which may greatly reduce its market value. In
living trees, it also interferes with transpiration, and the trees die
from the top downward.

Ceratocystis ips (Rumbold) C. Moreau is introduced into south-
ern pines by the engraver beetles, /ps calligraphus and I. grandi-
collis, and it produces a stain that spreads inward from the beetle
galleries toward the heartwood (fig. 208). This fungus has also
been found in living and felled red pines in the Lake States at-
tacked by J. pini and I. grandicollis (450).

: 5
ye
ca

3

COURTESY OF DUKE UNIV. SCH. OF FOREST.

FIGURE 208.—Cross-section of bole
of shortleaf pine killed by Ips
calligraphus, showing develop-
ment of blue-stain fungus, Cera-
tostomella ips, in sapwood.

F-519913

FIGURE 207.—Oak tree killed by

the oak wilt fungus, Ceratocystis
fagacearum.

Rumbold (626) discussed the interrelationships of bark
beetles and blue-staining fungi. Bramble and Holst (95) discussed
the fungi associated with the southern pine beetle in killing short-
leaf pines. Nelson (563) reported the effect of blue-staining fungi
on southern pines attacked by bark beetles.

Several species of ambrosia beetles carry staining fungi on
their bodies and, since they attack hardwood logs and lumber in
ee are considered to be important stain disseminators

733).

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Steinhaus, E. A.

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Stevens, R. E. and Stark, R. W.

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Stone, A.

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INDEX TO INSECTS BY HOST PLANTS

This index should enable foresters and others not trained in
entomology to identify many of the insects 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; otherwise, the scientific name is given.
Where more than one page reference is given for an insect, the
page on which the insect is discussed is in boldface type.

Acacia:
borer, twig and bark—
Cryptulocleptus dislocatus, 253
borer, wood—
mesquite borer, -178
Oncideres pustulus, 192
Thysanoes fimbricornis, 253
leaf feeders—
Cryptothelea gloverii, 413
sucking insects—
oleander scale, 101
Ailanthus:
ailanthus webworm, 403
Asiatic garden beetle, 149, 485
eynthia moth, 297
Alder (Alnus spp.):
borer, root and trunk (base)—
Sthenopis argenteomaculatus,
413
borer, wood—
alder borer, 188
birch and beech girdler, 201
fig tree borer, 199
horntail, Xiphydria
467
Oberea pallida, 190
poplar and willow borer, 218
Sinodendron rugosum, 145
flower feeder—
Xyela sp., 432
leaf feeders—
beetles:
alder flea beetle, 141
Attelabus rhois, 223
Chrysomela crotchi, 22, 123
anterrupta, Papa

mellipes,

138

cottonwood leaf beetle, 138
Dichelonyx elongata, 149

moths and butterflies:
Acrobasis rubrifasciella, 354
Acronicta distans, 308
Anacamptodes larvaria, 338
birch skeletonizer, 409

Alder—Continued
moths and butterflies—Con.
Bruce spanworm, 336
Eulype hastata, 337
Evora hemidesma, 360
Gracillaria pulchella, 409
Hyperetis amicaria, 342
large aspen tortrix, 3838
Lycia ursaria, 342
Pandemis limitata, 375
smeared dagger moth, 308
western tent caterpillar, 330
sawflies:
dusky birch sawfly, 462
elm sawfly, 438, 472
European alder leaf miner, 456
striped alder sawfly, 463
sucking insects—
Clastoptera obtusa, 74
Corythucha pergandei, 64
Myzocallis alnifoliae, 84
Psylla floccosa, 77
galeaformis, 77
woolly alder aphid, 85
Apple:
borers, bark and phloem—
flat-headed apple tree
165, 478
Lymantor decipiens, 254
Scolytus mali, 237
Synanthedon pyri, 399
borers, wood—
beetles:
apple twig borer, 136
flat-headed apple tree borer,
165, 478
Kulliana cincta, 178
Lichenophanes bicornis, 136
Lymantor decipiens, 254
Pseudolucanus capreolus, 145
round-headed apple tree
borer, 187
Saperda cretata, 188
stag beetle, 145

borer,

561

Apple—Continued
borers, wood—Continued
horntail, pigeon tremex, 466
moths and butterflies:
American plum borer, 359
leopard moth, 386
Synanthedon pyri, 399
fruit feeders—
apple fruit moth, 403
apple maggot, 424
cigar casebearer, 406
plum ecurculio, 227
leaf feeders—
beetles:
apple flea weevil, 223
basswood leaf miner, 141
European snout beetle, 219
flat-headed apple tree borer,
165, 478
locust leaf miner, 1438, 472,
478
Polydrusus impressifrons, 238,
220
round-headed apple tree
borer, 187
willow flea weevil, 222
mites, Eriophyes pyri, 7
moths and butterflies:
Acronicta distans, 308
interrupta, 308
Amorbia humerosana, 374
apple-and-thorn skeletonizer,
396
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
Bucculatrix pomifoliella, 409
cecropia moth, 298, 477
Coptodisca splendoriferella,
404
crinkled flannel moth, 350
Datana major, 313
eastern tent caterpillar, 328,
476, 477, 483
eye-spotted bud moth, 367
fall cankerworm, 21, 334, 339,
472, 483
filament bearer, 342
gypsy moth, 12, 18, 19, 24, 25,
2871025" Gos Cin lao, eo20,
430, 481, 469, 472, 474, 475,
476, 483
Haploa clymene, 307
Hemerocampa definita, 319
io moth, 299
Limenitis arthemis, 294
astyanax, 294
linden looper, 341
Lithocolletis crataegella, 407
Oriental moth, 347, 349, 431,
484
palmerworm, 393
Paonias excaecatus, 297
pepper and salt moth, 346
red-banded leaf roller, 384
red-humped caterpillar, 316
saddled prominent, 316, 472
Schizura tpomaeae, 317
leptinoides, 317

562

Apple—Continued
leaf feeders—Continued
moths and butterflies—Con.
smeared dagger moth, 308
Sparganothis sulfureana, 375
Sphinx drupiferarum, 297
spring cankerworm, 339
Synchlora aerata, 336
Tolype velleda, 333
unicorn caterpillar, 317
unspotted leaf miner, 408
variable oak leaf caterpillar,
SIS
western tent caterpillar, 330
white-marked tussock moth,
318, 476, 478, 483
winter moth, 336
yellow-necked caterpillar, 311
sucking insects—
Comstock mealybug, 100
periodical cicada, 74
woolly apple aphid, 84
Arborvitae:
borers, bark and phloem—
Phloesinus canadensis, 249
dentatus, 248
borers, wood—
Gnathotrichus materiarius, 268
spruce timber beetle, 268
leaf feeders—
arborvitae leaf miner, 403, 478
Argyresthia auveoargentella,
403
freyella, 402
bagworm, 21, 411, 472, 475, 477,
478
black vine weevil, 220
Coleotechnites thujaella, 391
hemlock looper, 344, 478, 483
Monoctenus melliceps, 439
Platytetranychus thujae, 7
root feeders—
arborvitae weevil, 218
black vine weevil, 220
strawberry root weevil, 221
sucking insects—
Cinara tujafilina, 83
European fruit lecanium, 105,
475
Fletcher scale, 106
juniper scale, 111
Ash (Fraxinus spp.) :
bark, under—
ash and privet borer, 199
oak-bark scaler, 199
borers, bark and phloem—
ash and privet borer, 199
Cucujus clavipes, 124
eastern ash bark beetle, 249
borers, wood—
beetles:
ash borer, 400
banded ash borer, 184
Chrysobothris sexsignata, 167
ivory-marked beetle, 192
Lyctus spp., 21, 127
painted hickory borer, 178

Ash—Continued
borers, wood—Continued
beetles—Continued
powder-post beetles, 17, 21,
126, 128, 155
red-headed ash borer, 184
Xyleborus ferrugineus, 270
Xyloterinus politus, 264, 267
moths and butterflies:
carpenterworm, 388
leopard moth, 386
gall insects—
Cecidomyia pellex, 419
Contarinia canadensis, 418
leaf feeders—
beetle:
Pachybrachys othonus, 145
mites:
Eriophyes fraxiniflora, 7
Tetranychus homorus, 7
moths and butterflies:
Agonopterix nigrinotella, 395
American dagger moth, 308
Apatelodes angelica, 334
ececropia moth, 298, 477
elm spanworm, 343, 484
fall cankerworm, 21, 334, 339,
472, 483
forest tent caterpillar, 19, 330,
AZO PAG 2 CATS
fruit-tree leaf roller, 377
green fruitworm, 309
Hemerocampa definita, 319
hickory horned devil, 303
lilac leaf miner, 409
Lycia ursaria, 342
Machimia tentoriferella, 395
Melanophia canadaria, 338
Papilio cresphontes, 292
Papaipema furcata, 309
Plagodis phlogosaria kuentz-
ingaria, 342
polyphemus moth, 299
promethea moth, 299
regal moth, 303
spring cankerworm, 339
tiger swallowtail, 291
Tolype velleda, 333
sawflies:
black-headed ash sawfly, 464
brown-headed ash sawfly, 463
mountain ash sawfly, 459
walkingstick, Diapheromera fe-
morata, 55
sap feeder—
Eastern hercules beetle, 152
sucking insects—
boxelder bug, 66
buffalo tree hopper, 70
English ivy scale, 101
Rune pear fruit lecanium, 105,
47
oystershell scale, 109, 110, 475
periodical cicada, 74
Prociphilus corrugatana, 85
Putnam scale, 108
San Jose scale, 106, 475
scurfy scale, 108

Ash—Continued
sucking insects—Continued
sycamore lacebug, 63
terrapin scale, 105
Tropidosteptes amoenus, 65
walnut scale, 107
white peach scale, 112
Ash, white:
borer, wood—
ash borer, 400
leaf feeder—
black-headed ash sawfly, 464
brown-headed ash sawfly, 463
Ceratomia undulosa, 295
Deuteronomos magnarius, 346
great ash sphinx, 296
Sparganothis dilutocostana, 375
fulgidipenna, 375
Sphinx kalmiae, 296
sucking insects—
cottony maple scale, 101
oystershell scale, 109, 110, 475
Aspen, see Poplar (Populus spp.)
Azalea:
borer, root—
rhododendron stem borer, 190
borer, wood—
dogwood twig borer, 189
pitted ambrosia beetle, 266
rhododendron borer, 398
rhododendron stem borer, 190
leaf feeders—
Arge clavicornis, 437
Asiatic oak weevil, 221
black vine weevil, 220
Datana major, 313
Gracillaria azaleella, 409
Japanese weevil, 219
Oligonychus ilicis, 6
root feeders—
Asiatic garden beetle, 149
Asiatic oak weevil, 221
black vine weevil, 220
Japanese weevil, 219
sucking insects—
azalea lacebug, 64
azalea whitefly, 92
mulberry whitefly, 92
striped mealybug, 100
rhododendron lacebug, 64

Bamboo (spp.):
borers, wood—
Chlorophorus annularis, 22, 207
false powder-post beetle, 134
sucking insects—
Asterolecanium miliaris mili-

aris, 101
miliaris ro-
bustum, 101

bamboo scale, 101
Banyon:
leaf feeder—
Lymire edwardsui, 304
Basswood:
borer, bark and phloem—
Chrysoclista linneella, 238, 390
linden borer, 187

563

Basswood—Continued

borers, bark and phloem—Con.
native elm bark beetle, 250, 499
Pseudothysanoes rigidus, 253
borers, wood—

beetles:
Anoplodera proxima, 197
Chalcophorella campestria,
168

Chrysobothris azurea, 167
Columbian timber beetle, 265
Dicerca lurida, 168
linden borer, 187
Platypus compositus, 2738
Polycaon stoutii, 136
Saperda imitans, 188
lateralis, 188
vestita, 185
sapwood timberworm, 126
twig girdler, 191, 502
horntail, Xiphydria abdomin-
alis, 467
moths and butterflies:
American plum borer, 359
flower feeder—
Japanese beetle, 19, 151, 485,
486
fruit feeder—
fruit tree leaf roller, 377
gall insects—
Cecidomyia verrucicola, 419
Melanagromyza tiliae, 427
leaf feeders—
beetles:
basswood leaf miner, 141
Japanese beetle, 19, 151, 485,
486
linden borer, 187
Phyllophaga crenulata, 148
drakei, 146
implicta, 148
prunina, 148
Plagiometriona clavata, 145
mite, Tetranychus canadensis, 7
moths and butterflies:
Abbottana clemataria, 346
Acronicta morula, 3¢8
American dagger moth, 308
Archips purpuranus, 378
bagworm, 21, 411, 472, 475,
477, 478
basswood leaf roller, 350
Ceratomia amyntor, 295
cecropia moth, 298, 477
Coleophora tilaefoliella, 404
Datana drexeli, 312
Deuteronomos magnarius, 346
elm sphinx, 295
fall cankerworm, 21, 334, 339,
472, 483
fruit tree leaf roller, 377
hemlock looper, 344, 478, 483
hickory tussock moth, 304
Limenitis arthemis, 294
astyanax, 294
linden looper, 341
Lithocolletis lucetiella, 407
Lycia ursaria, 342

064

Basswood—Continued

leaf feeders—Continued
moths and butterflies—Con.
Melanolophia canadaria, 338
palmerworm, 393
Pandemis lamprosana, 375
limitata, 375
Phigalia titea, 25, 339
polyphemus moth, 299
red-humped oakworm, 314
Sparganothis pettitana, 375
spring cankerworm, 339
Symmerista albifrons, 314
tiger swallowtail, 291
Tolype velleda, 333
variable oak leaf caterpillar,
315
white-marked tussock moth,
318, 476, 478, 483
winter moth, 336
yellow-necked caterpillar, 311
sawfly, elm sawfly, 4388, 472
walkingstick, 55
root feeders—
broad-necked root borer, 200
Japanese beetle, 19, 151, 485, 486
Phyllophaga crenulata, 148
drakei, 146
implicta, 148
prunina, 148
sucking insects—
basswood lacebug, 64
Corythucha juglandis, 64
cottony maple scale, 101
European fruit lecanium, 105,
475
giant bark aphid, 82
mulberry whitefly, 92
Myzocallis tiliae, 84
Phenacoccus acericola, 22, 99
Putnam scale, 108
San Jose scale, 106, 475
Telamona reclivata, 71
terrapin scale, 105
tuliptree scale, 103
walnut scale, 107

Beech (Fagus spp.):

bark, under dead—
Eucrada humeralis, 133
borers, bark and phloem—
Dryocoetes betulae, 263
flat-headed apple tree _ borer,
165, 478
Pseudopityophthorus pruinosus,
256
Scolytus fagi, 238
borer, nut—
filbertworm, 373
borer, wood—
beetles:
Ambrosiodmus tachygraphus,
Actenodes acornis, 169
269
Anisandrus spp., 269
birch and beech girdler, 201
Buprestis rufipes, 165
Chalcophorella campestris,
168

Beech—Continued
borer, wood—Continued

beetles—Continued
Chrysobothris sexsignata, 167
Columbian timber beetle, 265
Conotrachelus anaglypticus,
227
Eucrada humeralis, 133
flat-headed apple tree borer,
165, 478
fungus weevils, 209
Lichenophanes bicornis, 136
living-beech borer, 183
New York weevil, 209
oak timberworm, 208
Platypus compositus, 273
Ptilinus ruficornis, 133
Ayleborus ferrugineus, 270
XAyloterinus politus, 264, 267
horntails:
pigeon tremex, 466
Xiphydria mellipes, 467
tibialis, 467
moths and butterflies:
leopard moth, 386

gall insects—

Oligotrophus betulae, 23, 420

leaf feeders—

beetles:
Asiatic oak weevil, 221
flat-headed apple tree borer,
165, 478
locust leaf miner, 143, 472,
478
Polydrusus impressifrons, 238,

220
Phyllophaga drakei, 146
forsteri, 148
implicta, 148
luctuosa, 147
prunina, 148
Aanthonia decimnotata, 145
mite, Oligonychus bicolor, 6
moths and butterflies:
Acleris chalybeana, 385
Bruce spanworm, 336
Bucculatrix packardella, 410
Choristoneura fractivittana,
384
Coleophora alniella, 404
Colocasia propinquilinea, 309
dark tussock moth. 320
Datana angusi, 311
Deuteronomos magnarius, 346
Dicentria lignicolor, 316
fall cankerworm, 21, 334, 339,
472, 483
Fentonia marthesia, 316
gypsy moth, 12, 18, 19, 24, 25,
Ase oe bu. oh 23d.. 320:
430, 481, 469, 472, 474, 475,
476, 483
Heterocampa biundata, 316
Hyperetis amicaria, 342
imperial moth, 303
io moth, 299
luna moth, 298
maple leaf cutter, 413

Beech—Continued
leaf feeders—Continued
moths and butterflies—Con.
Nadata gibbosa, 313
Norape ovina, 350
Pandemis lamprosana, 375
Plagodis serinaria, 342
polyphemus moth, 299
Prolimacodes badia, 348
Psilocorsis faginella, 395
red-humped oakworm, 314
saddled prominent, 316, 472
Schizura ipomaeae, 317
leptinoides, 317
spotted tussock moth, 305
spring cankerworm, 339
Symmerista albifrons, 314
Tetralopha asperatella, 62,
351, 352
sp. near aspera-
tella, 351
unicorn caterpillar, 317
variable oak leaf caterpillar,
315
walnut sphinx, 297
root feeders—
Asiatic oak weevil, 221
Phyllophaga drakei, 146
implicta, 148
forsteri, 148
luctuosa, 147
prunina, 148
Polydrusus impressifrons, 28,
220
sucking insects—
beech blight aphid, 84
beech scale, 97, 501
Calaphis betulella, 84
Corythucha pallipes, 63
cottony maple scale, 101
European fruit lecanium, 105,
A75
giant bark aphid, 82
Lecanium caryae, 106
oystershell scale, 109, 110, 475
Phyllaphis sp., 80
Putnam scale, 108
Xylococculus betulae, 96
Beech, blue: See Hornbeam, Ameri-
can
Birch (Betula spp.)—
borers, bark and phloem—
bronze birch borer, 29, 161, 478
Conotrachelus anaglypticus, 227
Dryocoetes betulae, 2638
borer, cambium—
Phytobia pruinosa, 426
borer, seed—
Oligotrophus betulae, 23, 420
borers, wood—
beetles:
Actenodes acornis, 169
alder borer, 188
Ambrosiodmus tachygraphus,
269
Anoplodera vagans, 197
birch and beech girdler, 201
Chrysobothris azurea, 167

565

Birch—Continued
borers, wood—Continued

beetles—Continued

Conotrachelus anaglypticus,
227

Platypus compositus, 273
poplar-and-willow borer, 218
sapwood timberworm, 126
Trypodendron betulae, 268
Xyleborus obliquus, 271
Xyloterinus politus, 264, 267

horntails—

pigeon tremex, 466
Xiphydria mellipes, 467
tibialis, 467

fruit feeder—

fruit-tree leaf roller, 377

leaf feeders—

566

beetles:

basswood leaf miner, 141
Dichelonyx elongata, 149
locust leaf miner, 143, 472,
478
Pachybrachys peccans, 145
Phyllophaga crenulata, 148
drakei, 146
forsteri, 148
luctuosa, 147
Polydrusus impressifrons, 238,
220
Syneta ferruginea, 145

moths and butterflies:

Acronicta distans, 308
funeralis, 308
innotata, 308
interrupta, 308
leporina vulpina,
308
Bapta vestaliata, 337
birch sketonizer, 409
Bucculatrix coronatella, 410
cecropia moth, 298, 477
Charadra deridens, 309
Coleophora fuscedinella, 28.
405
Colocasia propingiilinea, 309
crinkled flannel moth, 350
Ectropis crepuscularia, 339
elm sphinx, 285
Eulype hastata, 337
fall cankerworm, 21, 334, 339,
472, 483
forest tent caterpillar, 19,
330, 429, 472, 475
fruit-tree leaf roller, 377
gypsy moth, 12, 18, 19, 24, 25,
28°62) 6a 7. 123-320; 430:
431, 469, 472, 474, 476, 483
Haploa lecontei, 307
Heterocampa bilineata, 316
hickory tussock moth, 304
lappet moth, 333
large aspen tortrix, 383
Limenitis arthemis, 294
astyanax, 294
linden looper, 341
luna moth, 298
Machimia tentoriferella, 395
maple leaf cutter. 413

Birch—Continued
leaf feeders—Continued
moths and butterflies—Con.
Oriental moth, 347, 349, 431,
484
Paonias excaecatus, 297
pink-striped oakworm, 301
polyphemus moth, 299
promethea moth, 298
Protobarmia porcelaria, 346
Psilocorsis faginella, 395
spotted tussock moth, 305
western tent caterpillar, 330
sawflies:
Arge clavicornis, 437
scapularis, 437
birch leaf miner, 454
birch leaf-mining sawfly, 453,
470, 473
dusky birch sawfly, 462
elm sawfly, 438, 472
Nematus pinguidorsum, 461
Pristiphora siskiyouensis, 459
striped alder sawfly, 463
: Trichiosoma triangulum, 438
sucking insects—
Calaphis betulaecolens, 84
betulella, 84
Chionaspis lintneri, 109
Clastoptera obtusa, 74
Corythucha pallipes, 63
pergandei, 64
Euceraphis betulae, 84
giant bark aphid, 82
Leeanium caryae, 106
oystershell scale, 109, 110, 475
Phenacoccus seratus, 99
Psylla carpinicola, 77
Birch, gray:
borers, bark and phloem—
bronze birch borer, 29, 161, 478
Pseudopityophthorus asperulus,
256
leaf feeders—
beetles:
Japanese beetle, 19, 151, 485,
486
willow flea weevil, 222
moths and butterflies:
Brephos intans, 334
birch skeletonizer, 409
chain-spotted geometer, 345
Coleophora fuscedinella, 23,
405
Datana angusi, 311
Epinotia similana, 374
filament bearer, 342
gypsy moth, 12, 18, 19, 24, 25,
28. 62, . G5. TA. ais eae:
430, 481, 469, 472, 474, 476,
483
Hemileuca lucina, 300
Nymphalis j-album, 292
Tortricidia flexuosa, 348
sawflies:
birch leaf-mining sawfly, 453,
470, 473
birch sawfly, 462

Birch, gray—Continued
leaf feeders—Continued
sawflies—-Continued
dusky birch sawfly, 462
elm sawfly, 438, 472
Profenusa thomsoni, 456
root feeders—
Japanese beetle, 19, 151, 485,
486

sucking insects—-
Euceraphis lineata, 84
Kleidocerys resedae geminatus,
6

ff
Lygaeid bugs, 67
Birch, paper:

borer, bark and phloem—

bronze birch borer, 29, 161, 478
borer, wood—

Trypodendron retusum, 268
gall insect—

Oligotrophus papyrifera, 419
leaf feeders—

beetles:

Birch, paper—Continued
leaf feeders—Continued
moths and butterflies—Con.

saddled prominent, 316, 472
Schizura inpomaeae, 317
leptinoides, 317
Sparganothis diliutocostana,
375
fulgidipenna, 375
Symmerista albifrons, 314
unicorn caterpillar, 317
variable oakleaf caterpillar,
315
white-marked tussock moth,
318, 476, 478, 483
yellow-necked caterpillar, 311

sawflies:

birch leaf-mining sawfly, 453,
470, 473

birch sawfly, 462

Profenusa thomsoni, 456

walkingstick, Diapheromera fe-

morata, 55, 57

bronze birch borer, 29, 161,
478
willow flea weevil, 222

moths and butterflies:

Abbottana clemataria, 346
Acleris logiana, 386
tripunctana, 386
American dagger moth, 308
Anacamptodes larvaria, 338
Archips purpuranus, 378
Argyrotaenia mariana, 385
Bibarrambla allenella, 395
birch skeletonizer, 409
birch tube maker, 354
Brephos infans, 334
Bruce spanworm, 336
chain-spotted geometer, 345
Choristoneura fractivittana,
384
Coleophora fuscedinella, 23,
405
Depressaria betulella, 395
Deuteronomos magnarius, 346
Epinotia solandriana, 371
gypsy moth, 12, 18, 19, 24, 25,
28202 65,1 lif, 12308 .320,
430, 431, 469, 472, 474, 476,
483
Hemerocampa definita, 319
hemlock looper, 344, 478, 483
Heterocampa biundata, 316
Hyperetis amicaria, 342
io moth, 299
Lophodonta ferruginea, 313
Lycia ursaria, 342
Melanolophia canadaria, 338
Nadata gibbosa, 313
Nymphalis j-album, 292
oblique-banded leaf roller, 383
Pandemis lamprosana, 375
limitata, 375
Prochoerodes transversata,
346
red-humped caterpillar, 316
red-humped oakworm, 314

sucking insects—
Corythucha pallipes, 63
Psylla annulata, 77
Xylococculus betulae, 96
Birch, river:
borer, bark and phloem—
Agrilus betulae, 163
borer, cambium—
Phytobia pruinosa, 426
borer, wood—
Xyleborus affinis, 270
leaf feeders—
Acleris logiana, 386
birch leaf-mining sawfly, 453,
470, 473
dusky birch sawfly, 462
SVPSV Mote wlZ lero. 24. 25.
28, 62, 65, 117, 123, 320, 430,
431, 469, 472, 474, 476, 483
larger elm leaf beetle, 141
sucking insects—
Euceraphis mucidas, 84
Birch, white, see Birch (Betula
spp.)
Birch, yellow:
borer, bark and phloem—
bronze birch borer, 29, 161, 478
borers, wood—
Chrysobothris sexsignata, 167
Columbian timber beetle, 265
leaf feeders—
beetle:
bronze birch borer, 29, 161,
478
moths and butterflies:
Acleris chalybeana, 385
American dagger moth, 308
birch skeletonizer, 409
elm spanworm, 343, 484
Gracillaria pulchella, 409
gypsy moth, 12, 18, 19, 24, 25,
28) (62.5 Ob. wleliian 23 7320,
430, 431, 469, 472, 474, 476,
483
hemlock looper, 344, 478, 483

567

Birch, yellow—Continued
leaf feeders—Continued
moths and butterflies—Con.
Hyperetis amicaria, 342
Plagodis serinaria, 342
white-marked tussock moth,
318, 476, 478, 483
yellow-necked caterpillar, 311
sawflies:
birch leaf-mining sawfly, 453,
470, 473
dusky birch sawfly, 462
Profenusa thomsoni, 456
sucking insects—
Carynota stupida, 71
Corythucha pallipes, 63
Elasmuche lateralis, 62
Euceraphis betulae, 84
Kleidocerys resedae geminatus,

i
Xylococculus betulae, 96
Boxelder:
borer, bark and phloem—
flat-headed apple tree borer,
165, 478
borer, seed—
Proteoteras aesculana, 369
borers, wood—
Ambrosiodmus
269
boxelder twig borer, 369
Columbian timber beetle, 265
flat-headed apple tree borer,
165, 478
hardwood stump borer, 205
gall insect:
boxelder gall midge, 418
leaf feeders—
beetles—
Asiatic garden beetle, 149, 485
flat-headed apple tree borer,
165, 473
moths and butterflies—
American dagger moth, 308
Archips negundanus, 378
fall cankerworm, 21, 334, 339,
472
Gracillaria negundella, 408
green fruitworm, 309
green-striped mapleworm, 301
Pandemis limitata, 375
smeared dagger moth, 308
spotted tussock moth, 305
spring cankerworm, 339
variable oak leaf caterpillar,
315
root feeders—
Asiatic garden beetle, 149, 485
sucking insects—
boxelder aphid, 83
boxelder bug, 66
cottony maple scale, 101
gloomy scale, 108
terrapin scale, 105
walnut scale, 107
Boxwood (spp.):
boxwood leaf miner, 418
boxwood psyllid, 77

tachygraphus,

568

Boxwood—Continued
Comstock mealybug, 100
Eurytetranychus buxi, 7
Buckeye:
borer, wood—
Petalium bistriatum bistriatum,

sapwood timberworm, 126
leaf feeder—
Derocrepis aesculi, 145
sucking insects—
Corythucha aesculi, 64
walnut scale, 107
Butternut:
borers, bark and phloem—
Agrilus juglandis, 163
Scolytus quadrispinosus, 235
borer, nut—
Conotrachelus juglandis, 227
borers, wood—
Anoplodera vagans, 197
Lyctus spp., 127
painted hickory borer, 178
Saperda discoidea, 187
leaf feeders—
Datana angusi, 311
Eriocampa juglandis, 464
hickory horned devil, 303
luna moth, 298
pecan leaf casebearer, 353
Pergidae spp., 436
polyphemus moth, 299
regal moth, 303
walnut caterpillar, 312, 472
walnut sphinx, 297
yellow-necked caterpillar, 311
sucking insects—
Corythucha juglandis, 64
Enchenopa binotata, 71
European fruit lecanium, 105,
475

Camellia:
Fiorinia theae, 112
Oligonychus ilicis, 6
Camphor tree:
camphor scale, 111
Caranga:
ash gray blister beetle, 120
Epicauta subglabra, 121
Lytta muttalli, 121
Casuarina, see Australian Pine
Catalpa:
catalpa sphinx, 295, 483
Cecidomyia catalpae, 419
Comstock mealybug, 100
tuliptree scale, 103
Catalpa bungii:
Phytobia clara, 427
Cedar spp.:
borers, bark and phloem—
Deodar weevil, 212
Melanophila acuminata, 169
pales weevil, 213
Phloesinus canadensis, 249
dentatus, 248
small cedar-bark borer, 198

Cedar—Continued
borers, root—
pales weevil, 213
borers, wood—
black-horned juniper borer, 195
Callidium schotti, 195
carpenter bee, 498
cedar tree borer, 198
Melanophila acuminata, 169
Pityophthorus pulicarius, 255
small cedar-bark borer, 198
Urocerus albicornis, 466
leaf feeders—
bagworm, 21, 411, 472, 475, 477,
478
Coleotechnites junipcrella, 391
hemlock looper, 344, 478, 483
juniper scale, 111
red carpenter ant, 489
red-headed pine sawfly, 19, 431,
439, 473
spruce spider mite, 6
Cedar, red:
ant, Allegheny mound, 491
bark feeder—
pales weevil, 213
borer, wood—
Chrysobothris texana, 167
gall insect—
juniper midge, 418
leaf feeders—
Argyresthia freyellu, 402
Pee Orn 21, 411, 472, 475, 477,
47
Choristoneura houstonana, 384
Coleotechnites juniperella, 391
imperial moth, 303
juniper webworm, 393
Monoctenus melliceps, 439
Paria sexnotata, 145
Trisetaceus cupressi
root feeder—
seed corn maggot, 427
sucking insects—
Cinara canadensis, 83
juniperivora, 83
sabinae, 83
Spilococcus juniper?, 99
Chamaecyparis:
arborvitae weevil, 218
Chamaecyparis obtusa, see Chamae-
cyparis
Cherry:
borer, bark and phloem—
Scolytus mali, 237
borers, wood—
Chrysobothris verdiceps, 167
powder-post beetles, 17, 21, 126,
128, 155
Pseudolucanus capreolus, 145
Trypopitys sericeus, 133
fruit feeders—
apple fruit moth, 403
Coleophora serratella, 406
plum curculio, 227
leaf feeders—
beetles:
basswood leaf miner, 141
Pyrrhalta cavicollis, 140

Cherry—Continued
leaf feeders—Continued
beetles—Continued
willow flea weevil, 222
moths and butterflies:
Acronicta distans, 308
interrupta, 308
Anacampsis innocuella, 393
Bruce spanworm, 336
cecropia moth, 298, 477
Coleophora serratella, 406
Coptodisca splendoriferella,
404
crinkled flannel moth, 350
eastern tent caterpillar, 328,
476, 477, 483
fall cankerworm, 21, 334, 339,
472, 483
Haploa lecontei, 307
oriental moth, 347, 349, 481,
484
palmerworm, 393
Sparganothis sulfureana, 375
spring cankerworm, 339
tiger swallowtail, 291
Tolype velleda, 333
unspotted leaf miner, 408
western tent caterpillar, 330
sawflies:
dusky birch sawfly, 462
Neurotoma fasciata, 435
pear slug, 457
Profenusa canadensis, 457
Sterictiphora prunivora, 437
sucking insect—
Quadraspidiotus forbesi, 107
Cherry, black:
borers, bark and phloem—
Hypothenemus eruditus, 254
borers, wood—
Anoplodera minnesotana, 197
Oberea ulmicola, 190
Xyloterinus politus, 264, 267
leaf feeders—
Archips purpuranus, 378
Bucculatrix pomifoliella, 409
Cerura borealis, 318
Limenitis arthemis, 294
astyanax, 294
Paonias myops, 297
pear slug, 457
Prolimacodes badia, 348
ugly-nest caterpillar, 376
Cherry, choke:
fruit feeders—
apple maggot, 424
Contarinia virginianiae, 418
Hoplocampa lacteipernnis, 465
montanicola, 465
leaf feeders—
Argyrotaenia mariana, 385
Bruce spanworm, 336
cherry scallop shell moth, 337
prairie tent caterpillar, 330
ugly-nest caterpillar, 376
western tent caterpillar, 330
Cherry, wild:
borers, bark and phloem—
Dryocoetes betulae, 263

569

Cherry, wild—Continued
borers, bark and phloem—Con.
native elm bark beetle, 250, 499
peach bark beetle, 240
shot-hole borer, 238
borers, cambium—
Phytobia prutnosa, 426
pruni, 426
borers, wood—
American plum borer, 359
powder-post beetle, 17, 21, 126,
128, 155
flower feeders—
Trichiotinus spp., 153
fruit feeders—
black cherry fruit fiy, 424
cherry fruit fly, 424
Hypothenemus georgiae, 254
gall insect—
Cecidomyia cerasiserotinae, 419
leaf feeders—
beetle:
eset leaf miner, 143, 472,
(
moths and butterflies:
Abbottana clemataria, 346
Apatelodes torrefacta, 3338
Bapta semiclarata, 337
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
chain-spotted geometer, 345
cherry scallop shell moth, 337
eynthia moth, 297
eastern tent caterpillar, 328,
476, 477, 483
Exartema quadrifidum, 374
fall cankerworm, 21, 334
filament bearer, 342
gypsy moth, 12, 18. 19, 24, 25,
25 OS sobs oluigeslla are 320;
430, 431, 469, 472, 474, 475,
476, 483
Hemerocampa definita, 319
Hemiluca lucina, 300
hemlock looper, 344, 478, 483
Heterocampa biundata, 316
io moth, 299
lappet moth, 333
Lithocolletis crataegella, 407
Lycia ursaria, 342
Machimia tentoriferella, 395
Packardia geminata, 348
Pandemis limitata, 375
pepper and salt moth, 346
Pero honestarius, 344
Prolimacodes scapha, 348
promethea moth, 298
red-humped caterpillar, 316
Schizura ipomaeae, 317
leptinoides, 317
smeared dagger moth, 308
Sphinx drupiferarum, 297
spotted tussock moth, 305
spring cankerworm, 339
Tetracis lorata, 346
Tortircidia flexuosa, 348
unicorn caterpillar, 317
unspotted leaf miner, 408

570

Cherry, wild—Continued
leaf feeders—Continued
moths and butterflies—Con.
yellow-necked caterpillar, 311
sawflies:
Neurotoma fasciata, 435
inconspicua, 435
Trichiosoma triangulum, 438
walkingstick:
Diapheromera femorata, 55,
57
sucking insects—
Corythucha associata, 64
pruni, 64
Prosapia bicincta, 73
Psylla trimaculata, 77
Cherry, wild black, see Cherry, wild
Chestnut:
borers, bark and phloem—
broad-necked root borer, 200
chestnut bark borer, 196
Conotrachelus anaglypticus, 227
Enaphalodes antomerius, 182
Pseudopityophthorus asperulus,
256
pubescens, 256
Pseudothysanoes lecontei, 253
two-lined chestnut borer, 160
borers, nut—
filbertworm, 373
large chestnut weevil, 226
small chestnut weevil, 226
Valentinia glandulella, 396
borers, root—
Asiatic oak weevil, 221
broad-necked root borer, 200
tile-horned prionus, 200
borers, wood—
beetles:
Ambrosiodmus lecontei, 269
linderae, 269
chestnut timberworm, 126
Cupes concolor, 118
ivory-marked beetle, 192
Magdalis salicis, 225
oak sapling borer, 183
oak stem borer, 194
pear blight beetle, 269
Platypus compositus, 273
Scobicia bidentata, 136
Xyleborus obliquus, 271
xylographus, 270
moths and butterflies:
leopard moth, 386
Synanthedon castanea, 399
leaf feeders—
Mnemonica auricyonia, 414
eak skeletonizer, 410, 411, 478
pink-striped oakworm, 301
Chestnut, horse:
borer, twig and wood—
Proteoteras aesculana, 369
flower or fruit feeder—
Japanese beetle, 19, 151, 485,
486
leaf feeders—
Asiatic garden beetle, 149, 485
Eotetranychus hicoriae, 7

Chestnut, horse—Continued
leaf feeders—Continued
flament bearer, 342
fruit-tree leaf roller, 377
Japanese beetle, 19, 151, 485,
486
Phyllophaga crenulata, 148
Tetranychus canadensis, 7
white-marked tussock moth, 318,
476, 478, 483
root feeders—
Asiatic garden beetle, 149, 485
Japanese beetle, 19, 151, 485,
486
Phyllophaga crenulata, 148
sucking insects—
Comstock mealybug, 100
Diaspidiotus aesculi, 108
Phenacoccus acericola, 22, 99
scurfy scale, 108
walnut scale, 107
Chinaberry:
Aspidiotus hederae, 112
citrus whitefly, 92
white peach scale, 112
Chinquapin:
Mnemonica auricyonia, 414
Conifers (spp.):
bark feeders—
Acanthocinus spp., 196
deodar weevil, 212
pales weevil, 213
pitch-eating weevil, 217
borers, bark and phloem—
Acanthocinus spp., 196
Crypturgus borealis, 239
deodar weevil, 212
Dryocoetes autographus, 263
eastern larch borer, 199
Hylobius pinicola, 216
warreni, 216
Ips spp., 260
pales weevil, 213
Phymatodes varius, 197
pitch-eating weevil, 217
Pityophthorus spp., 255
ribbed pine borer, 206
small cedar-bark borer, 198
white-pine weevil, 209, 423, 426,
475, 477
borers, cone and seeds—
Conophthorus spp., 256
Dioryctria abietella, 23, 355
Eucosma tocullionana, 368
Megastigmus spp., 476
spruce coneworm, 355
borers, wood—
beetles:
Anoplodera spp., 196
Asemum spp., 198
black-horned juniper borer,
195
black-horned pine borer, 194
brown prionid, 205
Buprestis spp., 164
cedar-tree borer, 198
Cossonus impressus, 223
Dicerca, spp., 168

Conifers—Continued
borers, wood—Continued
beetles—Continued

Dryophthorus americanus,

223

eastern larch borer, 199

Ernobius mollis, 1338

Eustrophus tomentosus, 125

Gnathotrichus materiarius,

268

hairy pine borer, 198

Hexarthrum ulkei, 224

Magdalis spp., 224

old house borer, 206

Pityophthorus, spp., 255

Platypus flavicornis, 272

pole borer, 197

Prionus pocularis, 200

ribbed pine borer, 206

Serropalpus barbatus, 125

striped ambrosia beetle, 267

Tomolips quercicola, 224

Trypodendron spp., 267

white-spotted sawyer, 203

Xyleborus dispar, 271

pini, 271

Xyletinus peltatus, 132

Aylotrechus sagittatus, 202
horntails—

Urocerus albicornis, 466

flavicornis, 466
flower feeders—
Cimberis spp., 223
Diodyrhynchus spp., 223
leaf feeders—
beetle:

white-spotted sawyer, 203
grasshopper:

Melanoplus bruneri, 58
mite, spruce spider mite, 6
moths and butterflies:

Anomogyna eliminata, 309

Aphelia alleniana, 386

bagworm, 411, 472, 475, 477,

478
Ectropis crepuscularia, 339
Eupethecia filmata, 337
luteata, 337
transcanadata,
337
Griselda radicana, 374
gypsy moth, 12, 18, 19, 24, 25,
ZOmsOZe OOK Ligseel23. 1320:
430, 431, 469, 472, 474, 475,
476, 483

Hyperetis amicaria, 342

Melanolophia canadaria, 338

oblique-banded leaf roller,

383

Orthosia hibisci, 309

Palthis angulalis, 309

pine tube moth, 384

pine tussock moth, 320

red-banded leaf roller, 384

rusty tussock moth, 318

Semiothisa granitata, 338

spruce bud moth, 369

spruce budworm, 25, 378, 472,

AT7

571

Conifers—Continued
leaf feeders—Continued
moths and butterfiies—Con.
white-marked tussock moth,
318, 476, 478, 483
Xenotenna pallorana, 386
Zeiraphara diniana, 370
root feeders—
Dichelonyx spp., 149
Diplotaxis spp., 149
goldsmith beetle, 152
Hylobius pinicola, 216
warreni, 216
Phyllophaga crenulata, 148
prununculina, 148
rugosa, 148
Polyphylla occidentalis, 149
variolosa, 149
Serica spp., 149
Cottonwood, see Poplar
Crapemyrtle:
citrus whitefiy, 92
Myzocallis kahawaluokalani, 84
Crataegus, see Hawthorn
Cypress:
borers, bark and phloem—
eypress bark borer, 197
southern cypress bark beetle,
248
borers, cone—
Cecidomyia cupressi, 419
Dioryctria pygmaeela, 358
borers, root—
pales weevil, 213
borers, wood—
bee, carpenter, 497, 498
beetles:
Acmaeodera pulchella, 160
ambrosia beetles, 12, 21, 120,
124, 264
Buprestis striata, 165
Chrysobothris sexrsignata, 167
cypress barker borer, 197
ivory-marked beetle, 192
Micracisella opacicollis, 253
Oeme rigida, 200
Platypus compositus, 273
Tomolips quercicola, 224
Trachykele lecontei, 169
Ayleborus affinis, 270
ferrugineus, 270
saxeseni, 269
termite, Formosan _ subterra-
nean, 50
gall insect—
Cecidomyia cupressi, 419
leaf feeders—
Anacamptodes pergacilis, 338
bagworm, 21, 411, 472, 475, 477,
478
Coleotechnites apictripunctella,
390
variella, 391
imperial moth, 303
Oiketicus abbotti, 411
Oligonychus boudreausxi, 6
pine colaspis, 142
Platytetranychus thujae, 7

O72

Cypress—Continued
leaf feeders—Continued
Systena marginalis, 144
sucking insects—
Cinara tujafilina, 83
juniper scale, 111

Deciduous trees and shrubs, see
Hardwoods spp.
Dogwood (Cornus spp.):
borers, bark and phloem—
Agrilus cephalicus, 163
dogwood borer, 398
Hypothenemus eruditus, 254
pecan borer, 398
Sunanthedon geliformis, 399
borers, fruit—
apple maggot, 424
borers, wood—
Chrysobothris azurea, 167
dogwood borer, 398
dogwood twig borer, 189
Oberea ulmicola, 190
Petalium bistriatum bistriatum,
133
pitted ambrosia beetle, 266
twig girdler, 191
Xyleborus saxeseni, 269
gall insect—
Mycodiplosis alternata, 421
leaf feeders—
Asiatic oak weevil, 221
dark tussock moth, 320
Diapheromera femorata, 55, 57
Epinotia lindana, 374
io moth, 299
Japanese weevil, 219
Macremphytus spp., 465
Phyllophaga drakei, 146
implicta, 148
red-humped caterpillar, 316
root feeders—
Japanese weevil, 219
Phyllophaga drakei, 146
implicta, 148
sucking insects—
Chionaspis corni, 109
lintneri, 109
salicisnigrae, 109
cottony maple scale, 101
mulberry whitefly, 92
obscure scale, 108
periodical cicada, 74
Putnam scale, 108
white peach scale, 112
Dogwood, flowering and ornamental,
see Dogwood
Douglas-fir:
borer, bark and phloem—
red turpentine beetle, 246
borer, root—
European chafer, 150
borers, twig, shoot and seed—
Dioryctria abietella, 23, 355
Megastimus spp., 476
spruce coneworm, 355
white-pine weevil, 209, 423, 426,
475, 477

Elm—Continued
borers, wood—Continued
horntails—Continued
Xiphydra abdominalis, 467
hicoriae, 467
tibialis, 467

Douglas-fir—Continued

borers, wood—
old house borer, 206
Semanotus litigiosus, 198
Stephanopachys substriatus, 136

leaf feeders—
European chafer, 150 moths and butterflies:
spruce budworm, 25, 378, 472, carpenterworm, 388

477 leopard moth, 386
sucking insect— leaf feeders—

: beetles:
Cooley spruce gall aphid, 87, 91 Mien Gir

Anomoea laticlavia, 145
‘ Brachys aeriguinosa, 169
Elder: elm calligrapha, 140
elm leaf beetle, 138, 431, 474
elder borer, 193 European snout beetle, 219

Phlyctaenia coronata, 351
Elm (Ulmus spp.) : ea apple tree borer,

bark feeders—

Desmocerus spp., 193

elm bark borer, 197
elm sawfly, 438, 472

borers, bark and phloem—

black elm bark weevil, 224, 478

elm borer, 188

flat-headed apple tree borer, 165,
478

native elm bark beetle, 250, 499

Neoclytus scutellaris, 184

peach bark beetle, 240

red elm bark weevil, 224, 478

Scolytus mali, 237

shot-hole borer, 238

smaller European elm bark bee-
Clee O One 4e 20 era 235,
475, 499

borers, wood—

beetles:
Anoplodera minnesotana, 197
banded-ash borer, 184
Buprestis rufipes, 165
Columbian timber beetle, 265
dogwood twig borer, 189
elm borer, 188
flat-headed apple tree borer,
165, 478
Goes spp., 182
Hylocurus langstoni, 253
ivory-marked beetle, 192
Lichenophanes bicornis, 136
living-beech borer, 183
Magdalis barbicornis, 225
pandura, 225
Neoclytus scutellaris, 184
Oberea tripunctata, 189
ulmicola, 190
Platybregmus canadensis, 134
Platypus compositus, 273
powder-post beetle, 17, 21,
126, 128, 155
Saperda lateralis, 188
Scobicia bidentata, 136
Thysanoes berschemiae, 253
twig girdler, 191, 502
twig pruner, 192
Xylosandrus germanus, 271
horntails:
pigeon tremex, 466

Japanese beetle, 19, 151, 485,
486
larger elm leaf beetle, 141
locust leaf miner, 143, 472,
A478
Pachybrachys othonus, 145
Phyllophaga drakei, 146
forsteri, 148
implicta, 148
prunina, 147
tristis, 148
willow flea weevil, 222
Xanthonia decimnotata, 145
mites:
Eotetranychus matthyssei, 7
Tetranychus canadensis, 7
schoenei, 7
moths and butterflies:
Acronicta funeralis, 308
interrupta, 308
morula, 308
American dagger moth, 308
Argyrotaenia mariana, 385
bagworm, 21, 411, 472, 475,
477, 478
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
Canarsia ulmiarrosorella, 359
cecropia moth, 298, 477
Charadra deridens, 309
Choristoneura fractivittana,
384
Coleophora ulmifoliella, 404
dark tussock moth, 320
Deuteronomos magnarius, 346
elm casebearer, 404
elm spanworm, 343, 484
elm sphinx, 295
fall cankerworm, 21, 334, 339,
472, 483
fruit-tree leaf roller, 377
gypsy moth, 12, 18, 19, 24, 25,
Pow, (Ova (oy Ie Be ey Thy
430, 431, 469, 472, 474, 475,
476, 483
Hemerocampa definita, 319
hemlock looper, 344, 478, 483
Heterocampa bilinecta, 316
hickory tussock moth, 304

573

Elm—Continued

leaf feeders—Continued
moths and butterflies—Con.
imperial moth, 303
i0 moth, 299
linden looper, 341
Lycia ursaria, 342
Machimia tentoriferella, 395
maple leaf cutter, 413
mourning cloak butterfly, 293
Nerice bidentata, 314
Pandemis lamprosana, 375
limitata, 375
pepper and salt moth, 346
Phigalia titea, 25, 339
Polygonia comma, 292
interrogationis, 292
polyphemus moth, 299
puss caterpillar, 349
red-humped oakworm, 314
Schizura ipomaeae, 317
leptinoides, 317
spring cankerworm, 339
Symmerista albifrons, 314
Tetralopha asperatella, 62,
351, 352
Tolype velleda, 333
unicorn caterpillar. 317
variable oak leaf caterpillar,
315
white-marked tussock moth,
318, 476, 478, 483 ~-
winter moth, 336
yellow-necked caterpillar, 311
sawflies:
Arge spp., 437
scapularis, 437
elm leaf miner, 455
elm sawfly, 438, 472
Macroxyela sp., 433
walkingstick:
Diapheromera femorata, 55,
57
nesting insect—
black carpenter ant, 484
root feeders—
Phyllophaga dralcei, 146
forsteri, 148
implicta, 148
prunina, 148
tristis, 148
sucking insects—
Brochymena quadripustulata, 62
buffalo treehopper, 70
Colopha ulmisacculi, 85
Corythucha pergandei, 64
cottony maple scale, 101
elm coxcomb gall aphid, 85
elm lacebug, 63
elm scurfy scale, 108
European elm scale. 108
European fruit lecanium, 105,
A475
giant bark aphid, 82
gloomy scale, 108
Gobaishia ulmifusa, 85
Gyponana spp., 69
Lecanium caryae, 106

O74

Elm—Continued

sucking insects—Continued
Myzocallis ulmifolii, 84
Neolygus invitus, 65
obscure scale, 108
oystershell scale, 109, 110, 475
periodical cicada, 74
Ponana spp., 69
Putnam scale, 108
San Jose scale, 106
Scaphoideus spp., 69
scurfy scale, 108
white-banded elm leafhopper, 69,
500
woolly apple aphid, 84
woolly elm bark aphid, 84
termite, Formosan subterranean,
50

Ficus spp.:

fig-tree borer, 199
Lymire edwardsii, 304

Fig tree, see Ficus
Fir, balsam:
- bark feeders—

Acanthocinus obsoletus, 196
pusillus, 196
Hylastes tenuis, 252
pales weevil, 213
white-spotted sawyer, 208
borers, bark and phloem—
beetles:
balsam fir sawyer, 205
Crypturgus atomus, 239
eastern larch borer, 199
four-eyed spruce bark beetle,
264
Hylastes tenuis, 252
Melanophila acuminata, 169
drummondi, 169
Northeastern sawyer, 205
Orthotomicus caelatus, 263
Pissodes dubius, 213
Pityokteines sparsus, 263
Pityophthorus balsameus, 255
opaculus, 255
patchi, 255
pulchellus, 255
spruce scolytus, 239
Taeneoglyptes fraseri, 259
moths and butterflies:
Marmara spp., 408
borers, cone—
Eucosma tocullionana, 368
spruce coneworm, 355
borers, wood—
beetles:
Acanthocinus pusillus, 196
Anoplodera canadensis, 196
balsam fir sawyer, 205
Chrysobothris neopusilla, 167
scabripennis,
167
Gnathotrichus materiarius,
268
hemlock borer, 168, 483
Melanophila acuminata, 169
drummondi, 169

|

Fir, balsam—Continued
borers, wood—Continued
beetles—Continued
northeastern sawyer, 205
old house borer, 206
Pityokteines sparsus, 263
Semanotus litigiosus, 198
spruce timber beetle, 268
Stephanopachys substriatus,
136
white-spotted sawyer, 203
Xylotrechus undulatus, 203
horntails, Sirex juvencus, 466
sawflies, Pleroneura brunneicor-
nis, 433
bud and flower feeders—
spruce bud moth, 369
spruce budworm, 25, 378, 472,
ATT
spruce coneworm, 355
gall insect, balsam gall midge, 418
leaf feeders—
beetles:
white-spotted sawyer, 203
moths and butterflies:
Anomogyna eliminata, 309
Argyrotaenia occultana, 385
black-headed budworm, 385,
472
Campaea perlata, 346
chain-spotted geometer, 345
Eufidonia notataria, 338
Eupethecia filmata, 337
luteata, 337
transcanadata, 337
false hemlock looper, 344
filament bearer, 342
Griselda radicana, 374
hemlock looper, 344, 478, 483
Hyperetis amicaria, 342
Lexis bicolor, 307
Palthis angulalis, 309
Pero morrisonarius, 344
pine tussock moth, 320
Protobarmia porcelaria, 346
red-banded leaf roller, 384
Semiothisa granitata, 338
spruce budworm, 25, 378, 472,
ATT
white-marked tussock moth,
318, 476, 478, 483
sawflies:
Acantholyda maculiventris,
435
balsam fir sawfly, 446, 473
Pleroneura brunneicornis, 433
nesting insects—
black carpenter ant, 484, 486
seed insects—
Megastigmus specularis, 476
sucking insects—
balsam twig aphid, 83
balsam woolly aphid, 12, 24, 88,
| 90123124
| Cinara curvipes, 80
/ Fiorinia externa, 111
hemlock scale, 108
pine spittlebug, 71

Fir, balsam—Continued
sucking insects—Continued
Saratoga spittlebug, 72, 472
Fir, Fraser:
balsam gall midge, 418
baisam twig aphid, 83
balsam woolly aphid, 12, 24, 88,
90, 1238; 124
Dryocoetes autographus, 263
Taenioglyptes fraseri, 259
Fir, Siberian, see Balsam Fir
Fringe tree:
Sphinx kalmiae, 296
Fruit trees, various:
borer, bark and phloem—
shot-hole borer, 238
borer, root—
Prionus spp., 200
borers, wood—
apple twig borer, 186
carpenterworm, 388
dogwood twig borer, 189
New York weevil, 209
pear blight beetle, 269
twig girdler, 191, 502
bud and fiower feeders—
eye-spotted budmoth, 365
fruit-tree leaf roller, 377
fruit feeders—
fruit-tree leaf roller, 377
plum curculio, 227
leaf feeders—
Datana ministra, 311
eastern tent caterpillar, 328,
476, 477, 483
eye-spotted budmoth, 365
fruit-tree leaf roller, 377
Oriental moth, 347, 349, 431, 484
red-humped caterpillar, 316

Gingko:

American plum borer, 359
Gum, black or sour, see Tupelo
Gum, red, see Sweetgum

Hackberry:

borers, bark and phloem—

Agrilus celti, 163
lecontei, 163
Chramesus chapuisii, 239
flat-headed apple tree borer,
165, 478

hackberry engraver, 239
Phloeotribus dentifrons, 240
Pseudothysanoes lecontei, 253
Scolytus fagi, 238

borer, root—
Archodontes melanopus, 206

borers, wood—
flat-headed apple tree borer,

165, 478
Hylocurus langstoni, 253
rudis, 252

Lichenophanes bicornis, 136
painted hickory borer, 178
pear blight beetle, 269
red-headed ash borer, 184
Scobicia bidentata, 136

575

Hackberry—Continued
borers, wood—Continued
twig girdler, 191, 502
twig pruner, 192
Thysanoes fimbricornis, 253
Xyleborus affinis, 270
gall insects—
blistergall psyllid, 76
bud-gall psyllid, 76
hackberry stargall, 76
hackberry-nipple-gall maker, 76
Pachypsylla celtidisinteneris, 77
petiole-gall psyllid, 77
leaf feeders—
Asterocampa celtis, 294
clyton, 294
mourning cloak butterfly, 293
Oiketicus abbotti, 411
oriental moth, 347, 349, 4381, 484
Polygonia interrogationis, 292
puss caterpillar, 349
Sphinx drupiferarum, 297
sucking insects—
Corythucha celtidis, 63
cottony maple scale, 101
elm scurfy scale, 108
European fruit lecanium, 105,
A475
gloomy scale, 108
mulberry whitefly, 92
psyllids, 76
obscure scale, 108
Putnam scale, 108
termite, Formosan subterranean,
50
Hardwoods:
bark feeders—
rustic borer, 202
borers, bark and phloem—
broad-necked root borer, 200
dogwood borer, 398
hickory bark beetle, 237
pecan borer, 398
Pseudopityophthorus minutissi-
mus, 255
two-lined chestnut borer, 160
borers, buds and shoots—
fruit-tree leaf roller, 377
Orthosia hibisci, 309
Polydrusus impressifrons, 28,
220
spined bark borer, 192
borers, root—
broad-necked root borer, 200
borers, wood—
beetles:
ambrosia beetle, 17, 21, 120,
124, 264
ash and privet borer, 199
banded hickory borer, 178
brown prionid, 205
Columbian timber beetle, 265
Cossonus cossonus, 223
impressifrons, 223
impressus, 223
platalea, 223
Dendrobiella aspera, 136
sericans, 136

576

Hardwoods—Continued

borers, wood—Continued
beetles—Continued
Dicerca divaricata, 168
obscura, 168
Dircacea quadrimaculata, 125
flat-headed apple tree borer,
165, 478
heartwood stump borer, 205
hickory spiral borer, 163
Holostrophus bifasciatus, 125
Hypothenemus dissimilis, 254
georgiae, 254
ivory-marked beetle, 192
Lichenophanes armiger, 136
living-beech borer, 183
Melandrya striata, 125
Monarthrum fasciatum, 267
mali, 267
oak timberworm, 208
painted hickory borer, 178
pear blight beetle, 269
Petalium bistriatum bistria-
tum, 133
Platypus flavicornis, 272
quadridentatus, 273
pole borer, 197
powder-post beetles, 17, 21,
126, 128, 155
Pseudopityophthorus minutis-
simus, 255
red-headed ash borer, 184
rustic borer, 202
sapwood timberworm, 126
snout beetles, 208
spined bark borer, 192
Stenoscelis brevis, 2238
twig pruner, 192
two-lined chestnut borer, 160
Xyleborus affinis, 270
pecanis, 271
saxeseni, 269
Xyletinus peltatus, 132
Xylobiops basilaris, 134, 502
Xyloblaptus quadrispinosus,
136
Xylosandrus germanus, 271
horntails, 465
moths and butterflies:
American plum borer, 359
carpenterworm, 388
leopard moth, 386
ten eastern subterranean,
4

fruit and seed borers and
feeders—
fruit flies, 424
Hypothenemus georgiae, 254
Japanese beetle, 19, 151, 485,
486
rose chafter, 150
vinegar flies, 425
leaf feeders—
ant, Texas leafcutting, 490
beetles:
Asiatic garden beetle, 149,
485
Brachys spp., 169

Hardwoods—Continued
leaf feeders—Continued
moths and butterflies—Con.

Hardwoods—Continued
leaf feeders—Continued
beetles—Continued

Eugnamptus spp., 223
European chafer, 150
European snout beetle, 219
flat-headed apple tree borer,
165, 478
green June beetle, 152
goldsmith beetle, 152
hickory spiral borer, 163
Japanese beetle, 19, 151, 485,
486
Phyllophaga crenulata, 148
forsteri, 148
prunina, 148
rugosa, 148
Polydrusus impressifrons, 238,
220
rose chafer, 150
Serica intermixta, 149
sericae, 149
tristis, 149
vespertina, 149
willow flea weevil, 222

crickets:

Mormon cricket, 59
snowy tree cricket, 60

hickory horned devil, 303

hickory tussock moth, 304

Hyphantria textor, 306

imperial moth, 303

io moth, 299

lappet moth, 333

leopard moth, 386

linden looper, 341

Lithophane latinicerea, 309

Lycia ursaria, 342

Machimia tentoriferella, 395

Melanolophia canadaria, 388

Nadata gibbosa, 313

oblique-banded leaf roller, 383

oriental moth, 347, 349, 481,
484

pale tussock moth, 305, 306

palmerworm, 393

Palthis angulalis, 309

Pero morrisonarius, 344

Phigalia titea, 25, 339

pink steed oakworm, 301
lagodis serinaria, 342

promethea moth, 298

Prochoerodes transversata,

grasshoppers, 57 342

katydids, 59
mae EO Ct ii oc: Prolimacodes badia, 348

American dagger moth, 308
Amorbia humerosana, 374
Anacamptodes ephyraria, 338
Apatelodes torrefacta, 333
Archips rosanus, 23, 376
bagworm, 21, 411, 472, 475,
477, 478
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
Bruce spanworm, 336
Campaea perlata, 346
Catocala spp., 308
cecropia moth, 298, 477
chain-spotted geometer, 345
cutworms, 309
dark tussock moth, 320
Deuteronomos magnarius, 346
Dichelonyx elongata, 149
eastern tent caterpillar, 328,
476, 477, 483

puss caterpillar, 349
red-banded leaf roller, 384
red-humped oakworm, 314
rusty tussock moth, 318
saddleback caterpillar, 347
Schizura ipomaeae, 317
leptinoides, 317
Sparganothis acerivorana,
375, 386
pettitana, 875
spring cankerworm, 339
Tetralopha asperatella, 62,
351 2352
Tolype velleda, 333
ugly-nest caterpillar, 376
unicorn caterpillar, 317
variable oak caterpillar, 315
white-marked tussock moth,
318, 476, 478, 483
winter moth, 3386
yellow-necked caterpillar, 311

Ectropis crepuscuiaria, 339
elm spanworm, 343, 484 : :
Epinotia solandriana, 371 nesting insects—
fall cankerworm, 21, 334, 339, carpenter ants, 486

A72, 483 Crematogaster ashmeadi, 49

walkingstick, giant, 57

fall webworm, 64, 306, 472,
476, 483

filament bearer, 342

forest tent caterpillar, 19, 330,

cerasi, 491
clara, 492
laerinsecula. 492
lineolata, 492

429, 472, 475
fruit-tree leaf roller, 377
Sypsyemoths 12, 18, 19, 24, 25,
2oO2 OD. tl], 123, 320, 430,
431, 469, 472, 474, 475, 476,

root feeders—
European chafer, 150
European snout beetle, 219
goldsmith beetle, 152
Phyllophaga crenulata, 148

486 forsteri, 148
hag moth, 347 prunina, 148
Hemerocampa definita, 319 rugosa, 148

O77

Hardwoods—Continued

root feeders—Continued
Polydrusus impressifrons, 28,
220
Serica intermixta, 149
serica, 149
tristis, 149
vespertina, 149
sap feeders—
ants, 486
Eastern hercules beetle, 152
hornets, 493
sap beetles, 121
springtails, 41
wasps, gall, 479
yellow jackets, 43
sucking insects—
Anormenis septentrionalis, 68
Claatoptera obtusa, 74
Emposca spp., 69
Erythroneura spp., 69
European fruit lecanium, 105,
475
Metcalfa pruinosa, 68
oystershell scale, 109, 110, 475
periodical cicada, 74
plant bugs, 65
Psylla spp., 77
Putnam scale, 108
San Jose scale, 106, 475
scurfy scale, 108
terrapin scale, 105
Typhlocyba spp., 69
walnut scale, 107

Hawthorn (Crataegus spp.):

borers, bark and phloem—
round-headed apple tree borer,
187
shot-hole borer, 238
Synanthedon pyri, 399
borers, fruit—
apple fruit moth, 403
apple maggot, 424
Coleophora serratella, 406
Hoplocampa oskina, 464
quince curculio, 227
borers, wood—
nound headed apple tree borer,
187
Saperda cretata, 188
Synanthedon pyri, 399
thorn-limb borer, 188
Aiphydria tibialis, 467
flower feeder—
Coleophora serratella, 406
leaf feeders—
beetles:
apple flea weevil, 223
Graphognathus spp., 224
larger elm leaf beetle, 141
locust leaf miner, 143, 472,
478
round-headed apple tree
borer, 187
moths and butterflies:
Acrobasis indiginella, 354
ep pleandsine skeletonizer,

578

Hawthorn—Continued

leaf feeders—Continued
moths and butterflies—Con.
Argyrotaenia quadrifasciana,
385
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
Bucculatrix pomifoliella, 409
cecropia moth, 298, 477
Coleophora serratella, 406
Cryptothelea gloveru, 413
eastern tent caterpillar, 328,
476, 477, 483
eye-spotted bud moth, 367
gypsy moth, 12, 18, 19, 24, 25,
285625 65, elie 2 3en 320,
430, 431, 469, 472, 474, 475,
476, 483
Hedia chionosema, 374
Lithocolletis crataegella, 407
maple trumpet skeletonizer,
371
polyphemus moth, 299
unspotted leaf miner, 408
sawflies:
Neurotoma crataegi, 485
inconspicua, 435
pear slug, 457
Pergidae spp., 436
Profenusa canadensis, 457
sucking insects—
Corythucha bellula, 64
cydoniae, 64
Eriosoma crataegi, 84
Frankliniella tritici, 42
Phenacoccus acericola, 22, 99
terrapin scale, 105
woolly apple aphid, 84

Hazel (Corylus spp.):

borers, nut—
Curculio neocorylus, 226
filbertworm, 373

leaf feeders—
apple flea weevil, 223
Asiatie oak weevil. 221
Attelabus rhois, 223
Dichelonyx submttata, 149
larger elm leaf beetle, 141
Lithocolletis corylisella, 408
palmerworm, 393
pink-strined oakworm, 301
spiny oakworm, 300

sucking insects—
Corythucha pergandei, 64

Hemlock (T7suga spp.):

bark feeders—
pales weevil, 213
tanbark borer, 197

borers, bark and phloem—
Dryocoetes autographus, 2638
eastern larch borer, 199
hemlock borer, 168, 483
Ips latidens, 262
Melanophila drummondi, 169
Microbreqma emarginatum

emarginatum, 133

pales weevil, 213

Hickory (Carya spp.):
bark feeders—
Cryptulocleptus dislocatus, 253
Hypothenemus spp.
New York weevil, 209
borers, bark and phloem—
Chramesus hicoriae, 239
196 Conotrachelus anaglypticus, 227
flat-headed apple tree borer,

Hemlock—Continued
borers, cone—
Eucosma tocullionana, 368
borers, root—
pales weevil, 2138
strawberry root weevil, 221
borers, wood—
Anoplodera canadensis,

black-horned juniper borer, 195

black-horned pine borer, 194

Buprestis maculipennis, 165

striata, 165
Chrysobothris pusilla, 167
scabripennis, 167
sexsignata, 167

hemlock borer, 168, 483

Melanophila drummondi, 169

Monarthrum fasciatum, 267
mahi, 267

old house borer, 206

pales weevil, 213

spruce timber beetle, 268

Stephanopachys substriatus, 136

striped ambrosia beetle, 267

Trypodendron scabricollis, 268

Xyleborus saxeseni, 269

Xyloterinus politus, 264, 267

Xylotrechus undulatus, 203

gall formers—

Nalepella tsugifoliae, 8

leaf feeders—

beetles:

Asiatic garden beetle, 149,
485

Japanese weevil, 219

pales weevil, 213

moths and butterflies:

Abbottana clemataria, 346

Anacamptodes ephyraria, 338

black-headed budworm, 385,
472

Campaea perlata, 346

Coleotechnites apicitripunc-
tella, 390

false hemlock looper, 344

Feralia jocosa, 309

filament bearer, 342

gypsy moth, 12, 18, 19, 24, 25,
28, 62, 65, 117, 123, 320, 4380,
431, 469, 472, 474, 475, 476,
483

hemlock looper, 344, 478, 483

Lambdina athasaria atha-
saria, 345

Melanolophia canadaria, 338

Neptyia semiclusaria, 344

Semiothisa granitata, 338

spruce budworm, 25, 378, 472,
U7

root feeders—

Japanese weevil, 219
strawberry root weevil, 221

sucking insects—

Fiorinia externa, 111
hemlock scale, 108
pine spittlebug, 71

165, 478
hickory bark beetle, 237, 270
hickory spiral borer, 163
mulberry bark borer, 193
Neoclytus mucronatus, 184, 186
scutellaris, 184
oak bark scaler, 199
Phymatodes varius, 197
Pseudopityophthorus pruinosus,
256

borers, nut—

Conotrachelus affinis, 227
hicoriae, 227
hickory shuckworm, 372, 472,
477, 485
pecan nut casebearer, 354
pecan weevil, 226
Valentinia glandulella, 396

borers, shoot and leaf petiole—

Acrobasis caryivorella, 353
apple twig borer, 1386
Chramesus hicoriae, 239
Conotrachelus aratus, 227
hickory bark beetle, 237, 270
pecan carpenterworm, 387
pecan nut casebearer, 354

borers, wood—

beetles:
Actenodes acornis, 169
Agrilus otiosus, 163
Anoplodera minnesotana, 197
proxima, 197
vagans, 197
banded-ash borer, 184
banded-hickory borer, 178
Buprestis rufipes, 165
Chramesus hicoriae, 239
Chrysobothris adelpha, 167
sexsignata, 167
Cryptulocleptus dislocatus,
253

Dicerca lurida, 168

Dryophthorus americanus,
223

flat-headed apple tree borer,
165, 478

flat powder-post beetle, 199
hickory spiral borer, 163
Hylocurus bicornus, 252

biorbis, 252

harnedi, 252

rudis, 252

spadix, 252
Hypothenemus dissimilis, 254

eruditus, 254

inmterstitialis, 254

quercus, 254

rotundicollis, 254
ivory-marked beetle, 192
Lichenophanes bicornis, 1386

579

Hickory—Continued
borers, wood—Continued

beetles—Continued
living-hickory borer, 183
Lyctus spp., 127
Lymantor decipiens. 254
Magdalis olyra, 225
Neoclytus mucronatus, 184,
18

scutellaris, 184
New York weevil, 209
Oberea ulmicola, 190
Orchesia castanea, 125
painted hickory borer, 178
Platypus compositus, 273
Polycaon stouti, 136
powder-post beetle, 17, 21, 126,
2S ali
red-headed ash borer, 184
Saperda discoidea, 187
imitans, 188
lateralis, 181
Scobicia bidentata, 1386
Thysanoes fimbricornis, 253
Trypopitys sericeus, 133
twig girdler, 191, 502
twig pruner, 192
white-oak borer, 182
Xyleborus affinis, 270
celsus, 270
ferrugineus, 270
lecontei, 271
obliquus, 271
saxeseni, 269
xylographus, 270
Xylobiops spp., 134
Xyloterinus politus, 264, 267
horntail, Xiphydria hicoriae,
467
moths and butterflies:
pecan carpenterworm, 387
sawflies, pigeon tremex, 466

gall insects—

Caryomyia holotricha, 420
sanguinolenta, 420
tubicola, 420

hickory gall aphid, 92

leaf feeders—

beetles:
Asiatie oak weevil, 221
Attelabus analis, 223
bipustulatus, 223
nigripes, 223
Bassareus lituratus, 145
flat-headed apple tree borer,
165, 478
Glyptoscelis barbata, 145
hickory spiral borer, 163
Pachybrachys peccans, 145
tridens, 145
Phyllophaga crenulata, 148
tristis, 148
mites:
Eotetranychus hicoriae, 7
Tetranychus homorus, 7
moths and butterflies:
Acrobasis caryivorella, 353
feltella, 354

Hickory—Continued
leaf feeders—Continued

moths and _ butterflies—Con.
Acronicta funeralis, 308
innotata, 308
lithospila, 308
American dagger moth, 308
Archips griseus, 378
infumatanus, 378
rileyanus, 378
Catocala spp., 308
Colocasia flavicornis, 309
Cryptothelea gloverii, 413
dark tussock moth, 320
Datana angusi, 311
elm spanworm, 343
Exartema permundanum, 374
fall cankerworm, 21, 334, 339,
472, 483
fruit-tree leaf roller, 377
gypsy moth, 12, 18, 19, 24, 25,
28> 6257 6os slippers 20)
430, 481, 469, 472, 474, 475,
476, 483
Haploa clymene, 307
hickory horned devil, 303
hickory leaf roller, 384
hickory tussock moth, 304
imperial moth, 303
io moth, 299
lappet moth, 333
linden looper, 341
luna moth, 298
Machimia tentoriferella, 395
oriental moth, 347, 349, 431,
484
pecan cigar casebearer, 404
pecan leaf casebearer, 353
Phigalia titea, 25, 339
polyphemus moth, 299
red-humped caterpillar, 316
regal moth, 3038
Schizura tpomaeae, 317
leptinoides, 317
spring cankerworm, 339
unicorn caterpillar, 317
walnut caterpillar, 312, 472
walnut sphinx, 297
sawflies:
Eriocampa juglandis, 464
Megaxyela spp., 433
Pergidae spp., 436
Periclista spp., 468
walkingstick: 55

root feeders—

Asiatic oak weevil, 221
Phyllophaga crenulata, 148
tristis, 148

sucking insects—

Clastoptera obtusa, 74
European fruit lecanium, 105,
A475
hickory gall aphid, 92
Lecanium caryae, 106
Monellia caryella, 84
costalis, 84
microsetosa, 84
Monelliopsis nigropunctata, 84

Hickory—Continued
sucking insects—Continued
Myzocallis caryaefoliae, 84
obscure scale, 108
periodical cicada, 74
Phylloxera spp., 91
plant bugs, 65
scurfy scale, 108
sycamore lacebug, 63
Holly (llex spp.):
borers, wood—
pear blight beetle, 269
Xyleborus saxeseni, 269
gall insect—
Asphondylia ilicicola, 418
leaf feeders—
Epicauta torsa, 121
holly leaf miner, 426
Tolype velleda, 333
sucking insects—
Chrysomphalus
alt?
Comstock mealybug, 100
Fiorinia theae, 112
holly scale, 101
mulberry whitefly, 92
Prosapia bicincta, 73
Honeylocust:
borers, bark and phloem—
Agrilus difficilis, 163
Hypothenemus dissimilis, 254
quercus, 254

dictyospermi,

borers, seed—
Amblycerus robiniae, 137
borers, wood—
Hylocurus langstoni, 253
Hypothenemus dissimilis, 254
quercus, 254
ivory-marked beetle, 192
painted hickory borer, 178
pear blight beetle, 269
twig girdler, 191, 502
Xyleborus saxeseni, 269
gall insects—
Dasineura gleditchiae, 418
leaf feeders—
beetles:
Anomoea laticlavia, 145
Epicauta cinerea, 121
mites:
Platytetranychus
tuli, 7
Tetranychus ellipticus, 7
moths and butterflies:
imperial moth, 3038
locust leaf roller, 358
Machimia tentoriferella, 395
mimosa webworm, 396
oriental moth, 347, 349, 481,
484
Semiothisa ocellinata, 338
Sparganothis sulfureana, 375
yellow-necked caterpillar, 311
sucking insects—
cottony maple scale, 101

European fruit lecanium, 105,
475

multidigi-

Honeylocust—Continued
sucking insects—Continued
Macropsis spp., 69
Micrutalis calva, 71
periodical cicada, 74
Hophornbeam:
borers, bark and phloem—
Pseudopityophthorus pruinosus,
256

Pseudothysanoes lecontei, 253
borers, wood—
Hypothenemus dissimilis, 254
quercus, 254
pitted ambrosia beetle, 266
leaf feeders—
beetles:
basswood leaf miner, 141
Epicauta cinerea, 121
Thysanoes fimbricornis, 253
mite:
Oligonychus letchworthi, 6
moths and butterflies:
Depressaria betulella, 395
gypsy moth, 12, 18, 19, 24, 25,
28762 165 r line 123s 320,
430, 4381, 469, 472, 474, 475,
476, 483
maple leaf cutter, 143
walnut sphinx, 297
winter moth, 336
yellow-necked caterpillar, 311
sawfly, Nematus ostryae, 461
sucking insects—
Corythucha pallipes, 63
Phenacoccus acericola, 22, 99
Hornbeam, see Hophornbeam
Hornbeam, American (blue beech) :
borers, bark and phloem—
Pseudopityophthorus pruinosus,
256
pubescens, 256
two-lined chestnut borer, 160
borers, wood—
banded hickory borer, 178
birch and beech girdler, 201
Dicerica lurida, 168
pitted ambrosia beetle, 266
Xiphydria tibialis, 467
Xylotrechus quadrmaculatus,
201
Huisache:
Oncideres pustulatus, 192

Incense-cedar, see Pine, incense-
cedar
Ironwood, (see also Hophornbeam;
Hornbeam, American) :
borers, wood—
Cinyra gracilipes, 170
two-lined chestnut borer, 160
leaf feeders—
Acleris chalybeana, 385
Aenea ostryaealla, 390
luna moth, 298
Pandemis lamprosana, 375
Tymnes tricolor, 145

581

Juniper (Juniperus spp.):

bark feeder—
pales weevil, 213
borers, bark and phloem—
Phloeosinus dentatus, 248
texanus, 249
borers, wood—
black-horned juniper borer, 195
black-horned pine borer, 194
cedar tree borer, 198
Oeme rigida, 200
pales weevil, 213
gall insect—
juniper midge, 418
leaf feeders—
arborvitae weevil, 218
Choristoneura houstonana, 384
Coleotechnites juniperella, 391
juniper webworm, 393
Monoctenus melliceps, 439
Phalonia rutilania, 23, 374
Platytetranychus thujae, 7
strawberry root weevil, 221
root feeders—
arborvitae weevil, 218
pales weevil, 213
strawberry root weevil, 221
sucking insects—
balsam twig aphid, 83
European fruit lecanium, 105,
475
Fletcher scale, 106
juniper scale, 111

Larch (Larix spp.):

bark feeders—
Hylurgops pinifex, 252
pales weevil, 213
borers, bark and phloem—
Crypturgus atomus, 239
Dryocoetes affaber, 263
eastern larch beetle, 248
eastern larch borer, 199
four-eyed spruce bark beetle,
264
hemlock borer, 168, 483
Melanophila drummondi, 169
Neoclytus muricatulus, 184
Orthotomicus caelatus, 263
Pityokteines sparsus, 263
red turpentine beetle, 246
spruce scolytus, 239
white-spotted sawyer, 203
borers, cone, bud and staminate
flower—
Spruce coneworm, 355
borers, root—
pales weevil, 213
borers, seed—
Megastigmus laricis, 476
borers, shoot and twig—
Argyresthia laricella, 403
Pityophthorus opaculus, 255
borers, wood—
Chrysobothris blanchardi, 167
dentipes, 167
sexsignata, 164

582

Larch—Continued

borers, wood—Continued
Gnathotrichus materiarius, 268
hemlock borer, 168, 483
horntail, Sirex abbottwu, 466
Melanophila drummondi, 169
Neoclytus muricatulus, 184
pales weevil, 213
Pityokteines sparsus, 263
Semanotus litigiosus, 198
Trypodendron lineatum, 267
spruce timber beetle, 268
white-spotted sawyer, 203
Xylotrechus undulatus, 203
bud feeders—
eye-spotted bud moth, 367
spruce budworm, 25, 378, 472,
AT7
leaf feeders—
beetles:
Anomala oblivia, 150
pine chafer, 150
white-spotted sawyer, 203
moths and butterflies:
Argyrotaenia occultana, 385
chain-spotted geometer, 345
Eupethecia luteata, 337
eye-spotted bud moth, 367
false hemlock looper, 344
Feralia jocosa, 309
gypsy moth, 12, 18, 19, 24, 25,
PAspmliy4s Ao) JU, IWR, YO.
430, 431, 469, 472, 474, 475,
476, 483
hemlock looper, 344, 478, 483
Hyperetis amicaria, 342
larch casebearer, 24, 405, 469,
473, 478
Melanolophia canadaria, 338
Neptyia semiclusaria, 344
Pero honestarius, 344
morrisonarius, 344
Protobarmia porcelaria, 346
red-banded leaf roller, 384
Semiothisa sexmaculata, 338
spruce budworm, 25, 378, 472,
477
spruce coneworm, 355
Tolype laricis, 333
white-marked tussock moth,
318, 476, 478, 483
Zeiraphara diniana, 370
sawflies:
Anoplonyx canadensis, 465
luteipes, 465
larch sawfly, 458, 471
red-headed pine sawfly, 19,
431, 439, 473
sucking insects—
Adelges lariciatus, 89
strobilobius, 22, 89

Larch, eastern, see Tamarack
Lilac:

Apatelodes angelica, 334
Ceratomia undulosa, 295
great ash sphinx, 296
lilac borer, 400

lilac leaf miner, 409

~

Lilac—Continued
promethea moth, 298
Sphinx kalmiae, 296
Linden, see Basswood
Loblolly Bay:
tuliptree scale, 103
Locust spp.:
twig pruner, 192
Locust, black:
borers, bark and phloem—
Agrilus egenus, 163
Hypothenemus obscurus, 254
locust borer, 17, 29, 64, 176
borers, root—
lesser cornstalk borer, 358
borers, wood—
carpenterworm, 388
Hypothenemus obscurus, 254
Lichenophanes bicornis, 136
locust borer, 17, 29, 64, 176
Lyctus spp., 127
painted hickory borer, 178
powder-post beetles, 17, 21, 126,
128555
XAyleborus affinis, 270
gall insects—
Dasineura pseudacaciae, 418
lesser cornstalk borer, 358
locust twig borer, 3738
Obolodiplosis robinae, 420
leaf feeders—
beetles:
Anomoea laticlavia, 145
Epicauta cinerea, 121
locust leaf miner, 143, 472, 478
mite, Tetranychus schoenei, 7
moths and butterflies:
Agonopterix robiniella, 395
bagworm, 21, 411, 472, 475,
477, 478
crinkled flannel moth, 350
hickory tussock moth, 304
io moth, 299
Lithocolletis ostensackenella,
407
robiniella, 407
Nephopteryx virgatella, 358
Parectopa robiniella, 408
pepper and salt moth, 346
Pero honestarius, 344
red-humped caterpillar, 316
Semiothisa ocellinata, 338
silver-spotted skipper, 294
spotted tussock moth, 305
walkingstick, Diapheromera fe-
morata, 55, 57
sucking insects—
Aphis craccivora, 84
cottony maple scale, 101
European fruit lecanium, 105,
475
Putnam scale, 108
Spissistilus festinus, 71
Thelia bimaculata, 496
Vanduzea arquata, 71

Magnolia:

borers, bark and phloem—
Euzophera ostricolorella, 359
borers, root—
Euzophera magnolialis, 359
borers, wood—
Euzophera magnolialis, 359
Platypus compositus, 273
quadridentatus, 273
Xyloterinus politus, 264, 267
leaf feeders—
Odontopus calceatus, 221
Paralobesia liriodendrana, 373
Phyllocnistis magnoliella, 408
Phyllophaga forsteri, 148
Tetranychus magnoliae, 7
root feeders—
Phyllophaga forsteri, 148
sucking insects—
Diaspidiotus liquidambaris, 108
European fruit lecanium, 105,
A475
magnolia scale, 103, 104
oleander scale, 101
striped mealybug, 100
San Jose scale, 106, 475
Trioza magnoliae, 77
tuliptree scale, 103

Mangrove:

Australian pine borer, 166

Maple (Acer spp.):

borers, bark and phloem—
Conotrachelus anaglypticus, 227
oak bark scaler, 199
Pseudopityophthorus pruinosus,
25

Synanthedon acerrubi, 399
borers, cambium—
Phytobia pruinosa, 426
borers, leaf petiole, twig and
shoot—
birch and beech girdler, 201
boxelder twig borer, 369
Nepticula sericopeza, 28, 413
Pityophthorus spp., 255
Proteoteras aesculana, 369
borers, root—
apple twig borer, 136
borers, seed—
Nepticula sericepeza, 23, 413
Proteoteras aesculana, 369
borers, wood—
beetles:
Agrilus otiosus, 163
Anisandrus obesus, 284
minor, 234
Anoplodera proxima, 197
Anthribidae, 116, 209
birch and beech girdler, 201
Buprestis rufipes, 165
Chalcophorella campestria,
168
Chrysobothris azurea, 167
gall-making maple borer, 201
Hylocurus rudis, 252
ivory-marked beetle, 192
Lymantor decipiens, 254
Micracisella opacicollis, 253

583

Maple—Continued
borers, wood—Continued
beetles—Continued
Orchesia castanea, 125
pitted ambrosia beetle, 266
Platybregmus canadensis, 134
Platypus compositus, 278
powder-post beetles, 17, 21,
1264 128 rl 55
Ptilinus ruficornis, 133
Thysanoes lobdelli, 253
twig pruner, 192
horntails:
pigeon tremex, 466
Xiphydria abdominalis, 467
maculata, 467
moths and butterflies:
carpenterworm, 388
leopard moth, 386
bud, blossom and fruit feeders—
fruit-tree leaf roller, 377
gall insects—
Cecidomyia ocellaris, 419
gall-making maple borer, 201
leaf-feeders—
bee, leaf-cutting, 486, 497
beetles:
Asiatic garden beetle, 149, 485
basswood leaf miner, 141
European snout beetle, 219
Japanese beetle, 19, 151, 485,
486
Phyllophaga drakei, 146
forsteri, 148
implicta, 148
tristis, 148
mite, Oligonychus aceris, 6
moths and butterflies:
Abbotana clemataria, 346
Acronicta retardata, 308
Apatelodes torrefacta, 333
bagworm, 21, 411, 472, 475,
477, 478
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
cecropia moth, 298, 477
Charadra deridens, 309
Colocasia propinquilinea, 309
Deuteronomos magnarius, 346
Ectropis crepuscularia, 339
fall ean’erworm, 21, 334, 339,
472, 483
Fentonia marthesia, 316
filament bearer, 342
forest tent caterpillar, 19,
330, 429, 472, 475
fruit-tree leaf roller, 377
Gracillaria bimaculatella, 49
green-striped mapleworm, 301
gypsy moth, 12, 18, 19, 24, 25,
28562 Gb pelleieel2o. 6320;
430, 431, 469, 472, 474, 475,
476, 483
Haploa clymene, 307
lecontei, 307
hemlock looper, 344, 478, 483
Heterocampa biundata, 316
umbrata, 316

584

Maple—Continued
leaf feeders—Continued
moths and butterflies—Con.
imperiai moth, 303
io moth, 299
lappet moth, 333
linden looper, 341
Lycia ursaria, 342
Machimia tentoriferella, 395
Nadata gibbosa, 313
oriental moth, 347, 349, 431,
484
pink-striped oakworm, 301
Plagodis serinaria, 342
polyphemus moth, 299
promethea moth, 298
Psilocorsis faginella, 395
puss caterpillar, 349
red-humped oakworm, 314
saddled prominent, 316, 472
Sparganothis pettitana, 375
spotted tussock moth, 305
spring cankerworm, 339
Tolype velleda
white-marked tussock moth,
318, 476, 478, 483
yellow-necked caterpillar, 311
root feeders—
Asiatic garden beetle, 149, 485
European snout beetle, 219
Phyllophaga drakei, 146
forsteri, 148
implicta, 148
tristis, 148
sucking insects—
Aleurochiton forbesi, 92
boxelder bug, 66
Comstock mealybug, 100
Corythucha pallipes, 63
cottony maple scale, 101
Drepanaphis acerifoliae, 84
carolinensis, 84
nigricans, 84
sabrinae, 84
Drepanosiphum platanoides, 84
European fruit lecanium, 105,
A475
giant bark aphid, 82
mulberry whitefly, 92
obscure scale, 108
oystershell scale, 109
Periphyllus negundinis, 83
Psylla annulata, 77
scurfy scale, 108
terrapin scale, 105
walnut scale, 107
Maple, hard, see Maple, sugar
Maple, mountain:
Acleris chalybeana, 385
Tetralopha asperatella, 62, 351,
352
Maple, Norway:
borers, leaf petiole and seed pod—
Nepticula sericopeza, 23, 413
flower feeder—
Japanese beetle, 19, 151, 485, 486
leaf feeders—
Gracillaria packardella

Maple, red—Continued
leaf feeders—
flat-headed apple tree borer,
165, 478

Maple, Norway——Continued
leaf feeders—Continued
sie ace beetle, 19, 151, 485,
486

Lithocolletis trinotella, 407

oriental moth, 347, 349, 481, 484

Sisyrosea textula, 348

white-marked tussock moth, 318,
476, 478, 483

fruit-tree leaf roller, 377
white-marked tussock moth, 318,
476, 478, 483

sucking insects—

cottony maple scale, 101

gloomy scale, 108
maple leaf scale, 102
oystershell scale, 109, 110, 475
Putnam scale, 108
woolly alder aphid, 85
Maple, soft, see Maple, silver
Maple, sugar:
borer, bud—
hard maple bud borer, 413
borer, cambium—
Phytobia setosa, 426
borers, wood—
Ambrosiodmus tachygraphus,

sucking insects—

Drepanosiphum platanoides, 84
Maple, red:

borer, cambium—
Phytobia setosa, 426

borers, wood—
Actenodes acornis, 169
Ambrosiodmus lecontei, 269
Chrysobothris sexsignata, 167

veridiceps, 167

Columbian timber beetle, 265
gall-making maple borer, 201
maple callus borer, 399

pitted ambrosia beetle, 266
gall insects—
Cecidomyia ocellaris, 419
Dasineura communis, 418
leaf feeders—
beetle:
willow flea weevil, 222
moths and butterflies:
Acleris chalybeana, 385
American dagger moth, 308

Choristoneura fractivittana,

384
elm spanworm, 343, 484
Hemerocampa definita, 319
Lithocolletis aceriella, 407
trinotella, 407
Lithophane latinicerea, 309
maple leaf cutter, 413

maple trumpet skeletonizer,

371
Nadata gibbosa, 313
Pandemis lamprosana, 375
Phigalia titea, 25, 339

Physostegama pustularia, 338

Plagodis serinaria, 342

Sparganothis acerivorana,

375, 386

Tetralopha asperatella, 62,

S5tnsoD2
winter moth, 336
sucking insect—

Diaspidiotus liquidambaris, 108

269
Columbian timber beetle, 265
maple callus borer, 399
pitted ambrosia beetle, 266
Ptilinus pectinicornis, 133
sugar-maple borer, 180
Xyloterinus politus, 264, 267

gall insects—

Dasineura communis, 418
Vasates aceris-crummena, 7

leaf feeders—

Acleris chalybeana, 385
American dagger moth, 308
Bruce spanworm, 336
Choristoneura fractivittana, 384
fall cankerworm, 21, 334, 339,
472, 483
forest tent caterpillar, 19, 330,
A293. AT2 tA 00
Gracillaria packardella, 409
green-striped maple worm, 301
Lithocolletis aceriella, 407
maple leaf cutter, 413
maple trumpet skeletonizer, 371
Nadata gibbosa, 313
orange-humped mapleworm, 314
Pandemis lamprosana, 375
red-humped oakworm, 314
saddled prominent, 316, 472
Sparganothis acerivorana, 375,
386
spring cankerworm, 339

Maple, silver: Symmerista albifrons, 314
borer, bark and phloem— Tetralopha asperatella, 62, 351,
flat-headed apple tree _ borer, 352
165, 478 sucking insects—
borers, wood— gloomy scale, 108
Columbian timber beetle, 265 maple leaf scale, 102

flat-headed apple tree borer, Norway maple aphid, 84
165, 478 Periphyllus americanus, 84

y : Be ot lyropictus, 84
. Sees Beer Ut busch, 3899 Phenacoccus acericola, 22, 99
yloterinus politus, 264, 267 Mesquite:
rf gall insects— ; Aneflus protensus, 199
-_ maple bladder gall mite, 7 banded-ash borer, 184
Vasates aceris-crummena, 7 mesquite borer, 193

585

Mesquite—Continued
Oncideres pustulatus, 192
Ptilinus ruficornis, 133

Mimosa:

Anomoea laticlavia, 145
Epicauta torsa, 121
mimosa webworm, 396
Norape ovina, 350
Xyleborus affinis, 270

Mountain-ash:

borers, bark and phloem—
peach bark beetle, 240
shot-hole borer, 238
borers, fruit—
apple fruit moth, 403
borers, wood—
American plum borer, 359
ash borer, 400
rOUne eae apple tree borer,
187
Synanthedon pyri, 399
flower or fruit feeder—
Japanese beetle, 19, 151, 485,
486

leaf feeders—
Eriophyes pyri, 7
Hedia chionosema, 374
Japanese beetle, 19, 151, 485,
486
Machimia tentoriferella, 395
mountain-ash sawfly, 459
Paria quadrinotata, 145
pear slug, 457
Foun ance apple tree borer,
7
yellow-necked caterpillar, 311
root feeders—
ea beetle, 19, 151, 485,
4

sucking insects—
Corythucha pallides, 63
San Jose scale, 106, 475
scurfy scale, 108
Mulberry:
borer, bark—
mulberry bark borer, 193
borers, bark and/or phloem—
peach bark beetle, 240
Phloeotribus frontalis, 240
borers, wood—
American plum borer, 359
Doraschema alternatum, 194
Hylocurus langstoni, 253
mulberry borer, 193
painted hickory borer, 178
leaf feeders—
io moth, 299
Phyllophaga luctuosa, 147
Prochoerodes transversata, 346
root feeders—
Phyllophaga luctuosa, 147
sucking insects—
Comstock mealybug, 100
cottony maple scale, 101
European fruit lecanium, 105
mulberry whitefly, 92
oleander scale, 101

086

Mulberry—Continued
sucking insects—Continued
sycamore lacebug, 63
terrapin scale, 105

Oak (Quercus spp.) :
borers, acorn—
beetles:
Conotrachelus carinifer, 227
naso, 227
posticatus, 227
Curculio caryatrypes, 226
humeralis, 226
longidens, 226
orthorhynchus, 226
pardalis, 226
sulcatulus, 226
moths and butterflies:
filbertworm, 373
Valentinia glandulella, 396
borers, bark and/or phloem—
beetles:
broad-necked root borer, 200
chestnut-bark borer, 196
Conotrachelus anaglypticus,
22
Enaphalodes antomerius, 182
Eucrada humeralis, 133
hickory spiral borer, 163
mulberry bark borer, 193
Neoclytus fulguratus, 184
jouteli, 184
scutellaris, 184
Nitidulidae spp., 115, 121
oak-bark scarrer, 181
Phymatodes varius, 197
Pseudopityophthorus asperu-
lus, 256
minutissimus, 255
pruinosus, 256
pubescens, 256
Pseudothysanoes lecontei, 253
red oak borer, 180
Stephanopachys substriatus,
136

tanbark borer, 197
two-lined chestnut borer, 160
moths and butterflies:
dogwood borer, 398
Marmara spp., 408
pecan borer, 398
borers, gall—
Synanthedon geliformis, 399
sapygaeformis, 399
borers, root—
broad-necked root borer, 200
oak sapling borer, 183
tile-horned prionus, 200
borers, wood—
bee, carpenter, 497, 498
beetles:
Atenodes acornis, 169
Agrilus otiosus, 163
ambrosia beetles, 17, 21, 120,
124, 264
apple twig borer, 136
banded hickory borer, 178
Buprestis rufipes, 165

Oak—Continued
borers, wood—Continued

Oak—Continued
borers, wood—Continued
beetles—Continued moths and butterflies—Con.
Chalcophorella campestria, leopard moth, 386
168 little carpenterworm, 389
Cinyra gracilipes, 170 Paranthrene palmu, 401
Coelostethus notatus, 133 pecan carpenterworm, 387
Columbian timber beetle, 265 bud and blossom feeders—
Cupes concolor, 118 eye-spotted bud moth, 367
Enaphalodes antomerius, 182 fruit-tree leaf roller, 377
Eucrada humeralis, 133 Pseudopityophthorus minutissi-
flat powder-post beetle, 134 mus, 255
hardwood stump borer, 205 gall insects—
hickory spiral borer, 163 Cecidomyiidae spp., 418
Hypothenemus (Stephano- Cynipidae spp., 479
deres) dissimilis, 254 leaf feeders—

interstitialis, 254 beetles:

quercus, 254
rotundicollis, 254
ivory-marked beetle, 192
Lichenophanes armiger, 1386
bicornis, 136
living-beech borer, 183
Micraciseila nanula, 253
opacicollis, 253
Neoclytus fulguratus, 184
jouteli, 184
scutellaris, 184
New York weevil, 209
oak branch borer, 183
oak sapling borer, 183
oak stem borer, 194
oak timberworm, 208
Oberea ulmicola, 190
Orchesia castanea, 125
painted hickory borer, 178
Petalium bistriatum bistri-
atum, 1338
seriatum, 133
Platypus compositus, 273
quadridentatus, 273
powder-post beetles, 17, 21,
126, 128, 155
Pseudolucanus capreolus, 145
Pseudopityophthorus minutis-
simus, 255
Ptilinus ruficornis, 133
red-headed ash borer, 184
red oak borer, 180
Saperda lateralis, 188
Scobicia bidentata, 136
Thysanoes lobdelli, 253
tile-horned prionus, 200
Trypopitys sericeus, 133
twig girdler, 191, 502
twig pruner, 192
two-lined chestnut borer, 160
white-marked spider beetle,
137
white oak borer, 182
Xyleborus affinis, 270
ferrugineus, 270
rubricollis, 271
xylographus, 270
Xyletinus lugubris, 132

horntail, pigeon tremex, 466
moths and butterflies:

carpenterworm, 388

Asiatic oak weevil, 221
Attelabus analis, 223
bipustulatus, 223
basswood leaf miner, 141
Brachys ovata, 169
Dichelonyx subvittata, 149
goldsmith beetle, 152
hickory spiral borer, 163
Japanese weevil, 219
locust leaf miner, 143, 472,
478
Pachybrachys carbonarius,
145

Piatti prununculina,
148

tristis, 148
Plagiometriona clavata, 145
Syneta ferruginea, 145
Systena marginalis, 144
Tymnes tricolor, 145
Xanthonia decimnotata, 145

grasshopper and katydid:

post oak locust, 58
Scudderia curvicauda, 59

mites:

Eotetranychus hicoriae, 7
Oligonychus bicolor, 6
propetes, 6

moths and butterflies:

Acronicta lithospila, 308
modica, 308
American dagger moth, 308
Archips georgianus, 378
griseus, 378
semiferanus, 378
Argyrotaenia alisellana, 385
mariana, 385
bagworm, 21, 411, 472, 475,
477, 478
Basicladus celibatus, 411
Bibarrambla allenella, 395
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
buck moth, 299
Bucculatrix packardella, 410
quinquenotella,
410
Catocala spp., 308
chain-spotted geometer, 345
Charadra deridens, 309
crinkled flannel moth, 350

587

Oak—Continued
leaf feeders—Continued

588

moths and butterflies—Con.
Croesia albicomma, 385
Cryptothelea gloverii, 413
Dasychira atomaria, 320
meridionalia, 320
tephra, 320
vagans, 320
Datana angusi, 311
contracta, 312
ministra, 311
Dicentria lignicolor, 316
Ectropis crepuscularia, 339
elm spanworm, 343, 484
eye-spotted bud moth, 367
fall cankerworm, 21, 334, 339,
472, 483
Fentonia marthesia, 316
filament bearer, 342
forest tent caterpillar, 19, 330,
429, 472, 475
fruit-tree leaf roller, 377
Gracillaria quercinigrella, 409
green-striped mapleworm, 301
gregarious oak leaf miner,
406
gypsy moth, 12, 18, 19, 24, 25,
285 62.2 OD eoliivan eh, 320)
430, 431, 469, 472, 474, 475,
476, 483
Haploa lecontei, 307
Hemileuca lucina, 300
hemlock looper, 344, 478, 483
Heterocampa umbrata, 316
Hyparpax aurora, 317
perophoroides, 317
imperial moth, 303
io moth, 299
Lacosoma chiridota, 347
Lambdina endopiaria, 345
lappet moth, 333
linden looper, 341
Iithocolletis bethunella, 407
Lophodonta angulosa, 313
luna moth, 298
Machimia tentoriferella, 395
Malacosoma tigris, 330
Melanolophia canadaria, 338
Melsheimer’s sack bearer, 347
Mnemonica auricyonia, 414
Nadata gibbosa, 313
oak skeletonizer, 410, 411, 478
oak webworm, 377
orange-striped oakworm, 300,
472
oriental moth, 347, 349, 431,
484
palmerworm, 393
Pandemis lamprosana, 875
limitata, 375
pecan carpenterworm, 387
Phaeoura quernaria, 342
Phigalia titea, 25, 339
pink-striped oakworm, 301
Polygonia interrogationis, 292
polyphemus moth, 299

Oak—Continued

leaf feeders—Continued
moths and butterflies—Con.
Prochoerodes transversata,
346
Prolimacodes badia, 348
Psilocorsis obsoletella, 395
quercicella, 395
reflexella, 395
puss caterpillar, 349
red-humped oakworm, 314
saddled prominent, 316, 472
Sisyrosea textula, 348
solitary oak leaf miner, 406
Sparganothis dilutocostana,
375
spiny oakworm, 300
spotted tussock moth, 305
spring cankerworm, 339
Symmerista albifrons, 314
Tetralopha asperatella, 62,
35132
Tolype velleda, 3338
Tortricidia flexuosa, 348

variable oak leaf caterpillar,

315
yellow-necked caterpillar, 311
sawflies:
Arge sp., 437

Pergidae spp., 436
Periclista spp., 468
Pristiphora chlorea, 459
Profenusa mainensis, 456
walkingstick, Anisomorpha
buprestoides, 57
root feeders—
Asiatic oak weevil, 221
Japanese weevil, 219
Phyllophaga prununculina, 148
tristis, 148
sap feeders—
Nitidulidae spp., 115, 121
sucking insects—
Alebra spp., 69
Asterolecanium minus, 101

Brochymena quadripustulata, 62

European fruit lecanium, 105,
475
Eutettix spp., 69
giant bark aphid, 82
Kermes galliformis, 97
kingui, 97
pettiti, 97
pubescens, 97, 475
trinotatus, 97
Lecanium caryae, 106
quercitronis, 106
oak lecanium, 106
obscure scale, 108
Myzocallis alhambra, 84
bella, 84
discolor, 84
melanocera, 84
Penthimia spp., 69
periodical cicada, 74
plant bugs, 65
Platycotis vittata, 71
yellow oak scale, 101

Oak, black:
borers, bark and phloem—
flat-headed apple tree borer, 165,
478
red oak borer, 180
borers, wood—
flat-headed apple tree borer, 165,
478
Paranthrene simulans simulans,
402
red oak borer, 180
Thysanoes fimbricornis, 253
gall insects—
Amphibolips confluenta, 481
Callirhytis operator, 480
Cecidomyia foliora, 419
gouty oak gall, 480, 481
horned oak gall, 479
large oak-apple gall, 481
ribbed bud gall, 480
leaf feeders—
Argyrotaenia quercifoliana, 384
Dichelonyx elongata, 149
flat-headed apple tree borer, 165,
478
Py psyaamothyl2. 185 519,.24.. 25,
DSO 60, Uli 23.5320.-450,
431, 469, 472, 474, 475, 476,
483
oak skeletonizer, 410, 411, 478
oak webworm, 377
walkingsticks, 55
nesting insects—
black carpenter ant. 484, 486
sucking insects—
European fruit lecanium, 105,
475
Oak, blackjack:
Callirhytis cornigera, 480
Oak, bur:
Bucculatrix recognita, 410
oak lacebug, 63
oak webworm, 377
Profenusa lucifex, 457
red-humped oakworm, 314
Symmerista albifrons, 314
Oak, Chapman:
Callirhytis floridana, 479
Oak, cherrybark:
Paranthrene palmii, 401
Oak, chestnut:
Asterolecanium quercicola, 101
golden oak scale, 101, 474
Iiothrips umbripennis, 42
oak bark scarrer, 199
oak lacebug, 63
obscure scale, 108
Oak, English:
Asterolecanium quercicola, 101
golden oak scale, 101, 474
Oak, ground:
Seirarctia echo, 307
Oak, live:
Anomoea laticlavia, 145
Archodontes melanopus, 206
Oiketicus abbott, 411
terrapin scale, 105

Oak, Nuttall:
Paranthrene palmii, 401
Oak, overcup:
Agrilus acutipennis, 163
Euxoa scandens, 309
white oak borer, 182
Oak, pin:
Argyrotaenia quercifoliana, 384
Caliroa (near) quercuscoccineae,
457
Cecidomyia foliora, 419
Eotetranychus querci, 7
horned oak gall, 479
obscure scale, 108
Parallelodiplosis florida, 420
pin oak sawfly, 457
ribbed bud gall, 480
Oak, post and dwarf post:
black carpenter ant, 484, 486
Callirhytis floridana, 479
Neoclytus muricatulus muricatu-
lus, 184
Phylloxera rileyi, 92
Oak, red:
borers, bark and phloem—
red oak borer, 180
borers, wood—
apple twig borer, 136
carpenterworm, 388
Chrysobothris veridiceps, 167
Paranthrene palmii, 401
simulars simulans,
402
red oak borer, 180
Thysanoes fimbricornis, 253
gall insects—
Cecidomyia foliora, 419
Cincticornia pilulae, 420
large oak-apple gall, 481, 482
succulent oak gall, 482
leaf feeders—
beetle, willow flea weevil, 222
fly, Japanagromyza viridula, 427
mite, Oligonychus bicolor, 6
moths and butterflies:
Argyrotaenia quercifoliana,
384
Bruce spanworm, 336
Choristoneura fractivittana,
384
gypsy moth, 12, 18, 19, 24, 25,
2851025 .0058 Whip ato uss 20"
430, 431, 469, 472, 474, 475,
476, 483
Hedia chionosema, 374
Hemerocampa definita, 319
oak skeletonizer, 410, 411, 478
oak webworm, 377
Phigalia titea, 25, 339
Psilocorsis faginella, 395
winter moth, 336
sawflies:
pin oak sawfly, 457
Profenusa mainensis, 456
nesting insects—
black carpenter ant, 484, 486
sucking insects—
Asterolecanium quercicola, 101

589

Oak, red—Continued
sucking insects—Continued
Cyrtolobus discoidalis, 71
Oak, scarlet:
Amphibolips confluenta, 481
black carpenter ant, 484, 486
Callirhytis punctata, 479
gypsy moth, 12, 18, 19, 24, 25, 28,
62) 05; ial 23.7320, 430843
469, 472, 474, 475, 476, 483
oak webworm, 377
red oak borer, 180
Symmerista albifrons, 314
Oak, scrub:
Argyrotaenia quercifoliana, 384
Brachys tesselatus, 169
Callirhytis operator, 480
Cincinnus melsheimeri, 347
horned oak gall, 479
oak fig gall, 482
oak webworm, 377
Oecanthus latipennis, 60
Parallelodiplosis florida, 420
Oak, southern red:
Coleophora atromarginata, 404
Oak, swamp chestnut:
Coleophora querciella, 404
Oak, swamp white:
Asterolecanium quercicola, 101
Coleophora atromarginata, 404
Neuroterus noxiosus, 481
Oak, water:
horned oak gall, 479
Oak, white:
borers, bark and/or phloem—
flat-headed apple tree borer, 479
oak-bark scaler, 199
oak-bark searrer, 181
borers, root—
oak sapling borer, 183
borers, wood—
Agriliaxia flavimana, 170
banded-ash borer, 184
chestnut timberworm, 126
Chrysobothris azurea, 167
sexsignata, 167
flat-headed apple tree borer, 165,
478
oak timberworm, 208
Physocnemum violareipenne,

white oak borer, 182
gall insects—
Callirhytis futilis, 480
Neuroterus floccosus, 481
quercusbatatus, 480
Xanthoteras forticorne, 482
leaf feeders—
beetle, flat-headed apple tree
borer, 165, 478
moths and butterflies:
Argyrotaenia quercifoliana,
384
Bucculatrix luteella, 410
recognita, 410
Coleophora querciella, 404
dark tussock moth, 320

590

Oak, white—Continued
leaf feeders—Continued
moths and butterflies—Con.
Euxoa scandens, 509
gregarious oak leaf miner,
406

gypsy moth, 12, 18, 19, 24, 25,
G2, nD lelu( eee o20)
ASO VAST ACO we Aii2er AlAs
476, 483
oak skeletonizer, 410, 411, 478
red-humped oakworm, 314
solitary oak leaf miner, 406
Symmerista albifrons, 314
variable oak leaf caterpillar,
315
sawflies:
Periclista spp., 468
pin oak sawfly, 457
Profenusa lucifex, 455
walkingsticks, 55
Orange, sour:
Cryptothelea gloverti, 413
Osage-orange:
Comstock mealybug, 100
Doraschema alternatum, 194
European fruit lecanium, 105, 475
Lyctus spp., 127
mulberry borer, 193
painted hickory borer, 178
Tetranychus canadensis, 7

Palm:
palm leaf skeletonizer, 390
palmetto pill bug, 223
royal palm bug, 67
Xyleborus lecontei, 271
Palm, Canary Island date:
palm leaf. skeletonizer, 390
Palmetto:
Caryobruchus gleditsiae, 137
Palmetto, cabbage:
palmetto pill bug, 223
Palmetto, Sabal:
palm leaf skeletonizer, 390
Seirarctia echo, 307
Papaya:
Toxotrypana curvicauda, 425
Peach:
borers, bark and phloem—
peach bark beetle, 240
borers, wood—
Synanthedon pictipes, 399
borers, fruit—
plum curculio, 227
leaf feeders—
Pyrrhalta cavicollis, 140
Sphinx drupiferarum, 297
sucking’ insects—
Comstock mealybug, 100
Cuerna costalis, 70
Graphocephala versuta, 70
Homalodisca coagulata, 70
insolita, 70
Oncometopia orbona, 70
periodical cicada, 74
white peach scale, 112

Pear:
borers, bark and phloem—
Dryocoetes betulae, 263
pacoded apple tree borer, 165,
47
borers, root—
tile-horned prionus, 200
borers, wood—
flat-headed apple tree borer, 165,
478
leopard moth, 386
tile-horned prionus, 200
flower and fruit feeder—
Coleophora serratella, 406
gall insect—
Eriophyes pyri, 7
leaf feeders—
beetles:
European snout beetle, 219
flat-headed apple tree borer,
165, 478
Polydrusus impressifrons, 23,
220
moths and butterflies:
apple-and-thorn skeletonizer,
396
Argyrotaenia mariana, 385
quadrifasciana, 385
brown-tail moth, 24, 325, 430,
431, 469, 472, 476
Coleophora serratella, 406
eye-spotted budmoth, 367
hickory tussock moth, 304
oriental moth, 347, 349, 431,
484
palmerworm, 393
unspotted leaf miner, 408
sawfly, pear slug, 457
root feeder—
European snout beetle, 219
Polydrusus impressifrons, 28,
220
sucking insects—
woolly apple aphid, 84
Pecan (see also Hickory):
borers, bark and phloem—
dogwood borer, 398
hickory bark beetle, 237, 270
pecan borer, 398
Synanthedon geliformis, 399
borers, nut—
hickory shuckworm, 372, 472,
477, 485
Hypothenemus (Stephanoderes)
georgiae, 254
pecan nut casebearer, 354
pecan weevil, 226
borers, shoot—
apple twig borer, 136
pecan nut casebearer, 354
twig girdler, 191, 502
borers, wood—
beetles:
Archodontes melanopus, 206
Chrysobothris adelpha, 167
hickory spiral borer, 163
Lichenophanes bicornis, 136
Lyctus spp., 127

Pecan—Continued
borers, wood—Continued
beetles—Continued

Oncideres cingulata texanus,
192

Platypus compositus, 273

twig girdier, 191, 502

Xyleborus saxeseni, 269

Xylobiops basilaris, 134, 502

moth, pecan carpenterworm, 387
bud and flower feeders—
Acrobasis demotella, 354
septentrionella, 354
pecan leaf casebearer, 354
leaf feeders—
beetles:

hickory spiral borer, 163

larger elm leaf beetle, 141

Phyllophaga spp., 146

white-fringed beetle, 224

mite, Hotetranychus hicoriae, 7
moths and butterflies:

Acrobasis caryivorella, 353

Archips infumatanus, 378

pecan cigar casebearer, 404

pecan leaf casebearer, 353

walnut caterpillar, 312, 472

walnut sphinx, 297

sawflies:
Megaxyela spp., 433
Pergidae spp., 436
sucking insects—
Clastoptera achatina, 74
giant bark aphid, 82
Myzocallis caryaefoliae, 84
obscure scale, 108
pecan leaf phylloxera, 92
pecan phylloxera, 92
Persimmon:
borer, bark and phloem—
Agrilus fuscipennis, 163
borer, root—
persimmon borer, 400
borers, wood—
beetles:

Agrilus otiosus, 168

Dicerca obscura, 168

Platypus compositus, 273

powder-post beetles, 17, 21,
126, 128, 155

red-headed ash borer, 184

twig girdler, 191, 502

Xyleborus affinis, 270

saxeseni, 269

Xylobiops basilaris, 134, 502

moths and butterflies:

American plum borer, 359

persimmon borer, 400

leaf feeders—
beetles:

Phyllophaga crenulata, 148
luctuosa, 147
prununculina, 148
tristis, 148

moths and butterflies:
bagworm, 21, 411, 475, 478
Cryptothelea gloverti, 413

591

Persimmon—Continued
leaf feeders—Continued
moths and butterflies—Con.
fall webworm, 64, 306, 472,
476, 483
hickory horned devil, 303
imperial moth, 303
luna moth, 298
red-humped caterpillar, 316
regal moth, 303
Seirarctia echo, 307
variable oak leaf caterpillar,
315
root feeders—
Phyllophaga crenulata, 148
luctuosa, 147
prununculina, 148
tristis, 148
sucking insects—
persimmon scale, 112
Trioza diospyri, 77
tuliptree scale, 103
Pines (Pinus spp.):
bark feeders—
beetles:
Acanthocinus spp., 196
Hylastes exilis, 252
porculus, 252
Hylurgops pinifex, 252
pales weevil, 213
Pissodes approximatus, 211
pitch-eating weevil, 217
white-pine weevil, 209, 423,
426, 475, 477
cricket, Oecanthus pini, 60
borers, bark and phloem—
beetles:
Acanthocinus nodosus, 196
obsoletus, 196
black turpentine beetle, 244
Carphoborus bifurcus, 239
Cossonus corticola, 223
impressus, 223
Crypturgus alutaceus, 239
atomus, 239
deodar weevil, 212
Dryocoetes affaber, 263
autographus, 263
four-eyed spruce bark beetle,
264
Hylastes exilis, 252
porculus, 252
Hylurgops pinifex, 252
Ips spp., 260
Ips avulsus, 260
calligraphus, 260, 502
Melanophila acuminata, 169
aeneola, 169
notata, 169
Neoclytus muricatulus, 184
Orthotomicus caelatus, 263
pales weevil, 213
pecan borer, 398
Phloeosinus pini, 249
pine engraver, 262
pitch-eating weevil, 217

592

Pines—Continued
borers, bark and phloem—Con.
beetles—Continued
Pityogenes meridianus, 258
plagiatus, 258
Pityokteines sparsus, 263
Pityophthorus annectens, 255
pulchellus, 255
pulicarius, 255
red turpentine beetle, 246
ribbed pine borer, 206
southern pine beetle, 240, 477,
502
southern pine engraver, 260
southern pine sawyer, 203
spotted pine sawyer, 205
Stephanopachys substriatus,
136
moth, dogwood borer, 398
borers, cone and seed—
beetles:
Conophthorus spp., 256
Ernobius granulatus, 133
Pityophthorus pulicarius, 255
Tricorynus sp., 134
moths and butterflies:
Dioryctria spp., 355
Eucosma monitorama, 368
Laspeyresia toreuta, 372
Moodna ostrinella, 359
borers and chewers, root—
beetles:
Dichelonyx spp., 149
Diplotaxis spp., 149
pales weevil, 213
Phyllophaga, spp., 146
pine root collar weevil, 213
pitch-eating weevil, 217
Polyphylla occidentalis, 149
Thylacites incanus, 23, 220
borers, wood—
beetles:
Anoplodera canadensis, 196
vagans, 197
Asemum striatum, 198
black horned juniper borer,
195
black-horned pine borer, 194
brown spider beetle, 137
Buprestis maculipennis, 165
striata, 165
Chalcophora georgiana, 168
liberta, 167
Chrysobothris floricola, 165
scabripennis, 167
Cupes concolor, 415
deodar weevil, 212
Dicerca punctulata, 168
tenebrosa, 168
Ernobius granulatus, 133
mollis, 133
furniture beetle, 132
Gnathotrichus materiarius,
268
Hexarthrum ulkei, 224
Hypothenemus (Stephano-
deres) georgiae, 254

.

Pines—Continued
borers, wood—-Continued
beetles—-Continued
Hypulus concolor, 125
Magdalis perforatas, 224
Melanophila acuminata, 169
aeneola, 169
notata, 169
Micracisella opacicollis, 253
Monarthrum fasciatum, 267
Monochamus carolinensis, 205
Myeloborus sp., 259
Neoclytus muricatulus, 184
old house borer, 206
Orthotomicus caelatus, 263
pales weevil, 213
Petalium seriatum, 133
Pityophthorus puberlus, 255
pulicarius, 255
Platypus flavicornis, 272
Prionus pocularis, 200
southern pine sawyer, 2038
spotted pine sawyer, 205
spruce timber beetle, 268
Stephanopachys cribratus, 136
densus, 136
hispidulus, 136
substriatus, 136
striped ambrosia beetle, 267
Tomolips quercicola, 224
Trypodendron scabricollis, 268
white pine weevil, 209, 423,
426, 475, 477
Xyleborus ferrugineous, 270
howardi, 271
xylographus, 270
Xyloterinus politus, 264, 267
Xylotrechus sagittatus, 202
horntails:
Sirex cyaneus, 466
juvenecus, 23, 466
Urocerus albicornis, 466
moths and butterflies:
Dioryctria abietella, 23, 355
amatella, 356
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
pitch-twig moth, 365
Rhyacionia spp., 361
Zimmerman pine moth, 356
flower feeders—
Cimberis spp., 223
Diodyrhynchus spp., 223
Gnophothrips fuscus, 42
Oecanthus pini, 60
Xyela spp., 432
leaf cutters—
Texas leaf cutting ant, 490
leaf feeders—
beetles:
Anomala oblivia, 150
Asiatic garden beetle, 149, 485
Dichelonyx albicollis, 149
subvittata, 149
Diplotaxis liberata, 149
Glyptoscelis pubescens, 145

Pines—-Continued
leaf feeders-——Continued
beetles—-Continued
Phyllophaga forsteri, 148
prunina, 148
prununculina, 148
pine colaspis, 142
Thylacites incanus, 238, 220
crickets, grasshoppers and katy-
dids:
Hubbellia marginifera, 59
Oecanthus pini, 60
post-oak locust, 58
short-tailed cricket, 60
mites:
Oligonychus milleri, 6
Phytoptus pini, 8
moths and butterflies:
Aphelia alleniana, 386
bagworm, 21, 411, 472, 475,
ATA EATS.
Basicladus celibatus, 411
Citheronia sepulchralis, 303
Eupethecia palpatata, 337
eypsy moth; 12, 18.19; 24) 25,
298; AOZ OD cell aol 23. “320;
430, 431, 469, 472, 474, 475,
476, 483
imperial moth, 303
jack pine budworm, 381
Lambdina athasaria_ pelluci-
daria, 345
Lapara coniferarum, 297
Melanolophia canadaria, 338
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
Neptyia semiclusaria, 344
Panthea furcilla, 309
Pero morrisonarius, 344
spruce budworm, 25, 378, 472,
AT7
zebra caterpillar, 309
sawflies:
Diprion frutetorum, 238, 451,
473
introduced pine sawfly, 28,
450, 476
Neodiprion spp., 441
pine false webworm, 433
nesting insects—
Camponotus nearcticus, 489
carpenter bee, 497, 498
root feeders—
Dichelonyx spp., 149
Diplotaxis spp., 149
Phyllophaga forsteri, 148
prunina, 148
prununculina, 148
Polyphylla occidentalis, 149
variolosa, 149
stem cutter—
short-tailed cricket, 60
sucking’ insects—
Aspidiotus abietis, 112
black pine leaf scale, 111
Cinara atlantica, 83
Clastoptera proteus, 74
hemlock scale, 108

593

Pines—Continued
sucking insects—Continued
Matsucoccus alabamue, 96
pine bark aphid, 89
pine leaf chermid, 91
pine needle scale, 109
pine tortoise scale, 103, 123
pine twig gall scale, 65, 96
Pineus floccus, 91
Toumeyella parvicornis, 105
Trioza tripunctata, 77
woolly pine scale, 106
Pine, Australian:
Australian-pine or mangrove bor-
er, 166
Clastoptera undulata, 73
Pine, Austrian:
bark feeders—
northern pine weevil, 211
borers, bark and phloem—
pine root collar weevil, 213
pitch mass borer, 399
borers, bud and shoot—
Battaristis vittella, 394
European pine shoot moth, 12,
DAP ZA ee SONA Osea lena ion Aran
475, 477, 478
pine bud moth, 392
white-pine shoot borer, 367
Zimmerman pine moth, 356
borers, cone—
Battaristis vittella, 394
borers, root—
pine root collar weevil, 213
borers, wood——
northern pine weevil, 211
pine root collar weevil, 213
pitch mass borer, 399
Zimmerman pine moth, 356
leaf feeders—
Acantholyda angulata, 435
zappei, 434
European pine sawfly, 12, 19,
24, 447, 470, 471, 473
pine bud moth, 382
pine webworm, 65, 351
sucking’ insects—
Leptoglossus occidentalis, 66
pine bark aphid, 89
pine needle scale. 109
pine tortoise scale, 103, 123
Pine, Caribbean:
Neodiprion abbotii, 447
southern pine engraver, 260
Pine, eastern white:
ant, mound-building—
Allegheny mound, 491
bark feeders—
pales weevil, 213
borers, bark and phloem—
beetles:
Hylobius congener, 217
Ips spp., 260
Myeloborus ramiperda, 259
pales weevil, 213
pine root collar weevil, 213
Pityogenes hopkinsi, 258
Pityophthorus spp., 255

594

Pine, eastern white—Continued
borers, bark and phloem—Con.
beetles—Continued
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
white-pine weevil, 209, 423,
426, 475, 477
white-spotted sawyer, 203
moths and butterflies:
Marmara fasciella, 408
pitch mass borer, 399
borers, bud, twig and shoot—
beetles:
Myeloborus ramiperda, 259
Pissodes affinis, 213
Pityophthorus spp., 255
white-pine weevil, 209, 423,
426, 475, 477
moths and butterflies:
Dioryctria abietella, 23, 355
European pine shoot moth, 12,
224.6 36l A Os ATA AT 2.
474, 475, 477, 478
pine bud moth, 392
white-pine shoot borer, 367
Zimmerman pine moth, 356
borers, cone—
Dioryctria abietella, 23, 355
Eucosma tocullionana, 368
white-pine cone beetle, 256
borers, wood—
beetles:
Anoplodera biforis, 197
Chrysobothris blanchardi, 167
dentipes, 167
harrisi, 167
pusilla, 167
trinervia, 167
hemlock borer, 168, 483
Hylobius congener, 217
northeastern sawyer, 205
northern pine weevil, 211
pales weevil, 213
Pityogenes hopkinsi, 258
Pityophthorus cariiceps, 255
opaculus, 255
white-pine weevil, 209, 423,
426, 475, 477
white-spotted sawyer, 203
moths and butterflies:
Dioryctria abietelia, 28, 355
pitch mass borer, 399
Zimmerman pine moth, 356
leaf feeders—
ant, Allegheny mound, 491
beetles:
Glyptoscelis pubescens, 145
Magdalis austera, 225
hispoides, 225
Anomala oblivia, 150
grasshopper:
Melanoplus punctulatus, 58
moths and butterflies:
chain-spotted geometer, 345
Eacles imperialis pini, 304
Eufidonia notataria, 338
Eupethecia palpatata, 337

Pine, eastern white—Continued
leaf feeders—Continued
moths and butterflies—Con.
gypsy moth, 12, 18, 19, 24, 25,
Z2oeaoOl OD elias 235 320)
430, 431, 469, 472, 474, 475,
476, 483
jack-pine budworm, 381
Lapara bombycoides, 297
oblique-banded leafroller, 383
pine bud moth, 392
pine tube moth, 384
pine webwormn, 65, 351
Semiothisa bisignata, 338
granitata, 388
Xenotemna pallorana, 386
sawflies:
Acantholyda angulata, 485
luteomaculata, 435
European pine sawfly, 12, 19,
24, 447, 470, 471. 473
introduced pine sawfly, 28,
450, 476
nesting-pine sawfly, 434
pine-false webworm, 433
red-headed pine sawfly, 19,
431, 439, 478
red-pine sawfly, 445, 473
Swaine juck-pine sawfly, 24,
444
white-pine sawfly, 446
nesting insects—
carpenter ants, 484, 486
carpenter bee, 497, 498
pollen feeders—
jack-pine budworm, 381
Ayela alpigena, 4382
root feeders—
pine root collar weevil, 213
sucking insects—
Eulachnus rileyi, 83
pine bark aphid, 89
pine leaf chermid, 91
pine needle scale, 109
pine spittlebug, 71
Pineus floccus, 91
Pitedia uhleri, 62
Prociphilus bumeliae, 83
Saratoga spittlebug, 72, 472
white-pine aphid, 83
Pine, incense-cedar:
juniper scale, 111
Pine, jack:
borers, bark and phloem—
beetles:
Ips latidens, 262
perroti, 262
northern pine weevil, 211
pine engraver, 262
pine root collar weevil, 213
Pityogenes hopkinsi, 258
Pityophthorus spp., 255
red turpentine beetle, 246
southern pine engraver, 260
white-pine weevil, 209, 423,
426, 475, 477
white-spotted sawyer, 203

Pine, jack—Continued
borers, bark and phloem—Con.
moth:

pitch nodule maker, 366

borers, bud and shoot—
beetles:

Conophthorus spp., 256
white-pine weevil, 209, 423,
426, 475, 477

moths and butterflies:

European pine shoot moth, 12,
Zile 2A S861, Avi OsrAjolerea tics
474, 475, 477, 478

jack-pine shoot borer, 368

Petrova virginiana, 365

pine bud moth, 392

pitch twig moth, 365

Rhyacionia adana, 362

sonia, 365

spruce coneworm, 355

western-pine tip moth, 365,
478, 485

white-pine shoot borer, 367

borers and feeders, root:
Dichelonyx spp., 149
Diplotaxis spp., 149
Hylobius rhizophagus, 216
pine root collar weevil, 213
borers, wood—
beetles:

Acanthocinus pusillus, 196
Chrysobothris orono, 166
northern pine weevil, 211
Pissodes affinis, 213
Pityophthorus spp., 255
white-pine weevil, 209, 423,
426, 475, 477
white-spotted sawyer, 203

moths and butterflies:

Petrova virginiana, 365
pitch nodule maker, 366

cone, seed, and flower insects—
Dioryctria disclusa, 358, 475
Gnophethrips fuscus, 42
red-pine cone beetle, 257
spruce coneworm, 355

gall insects—
Cecidomyia reeksi, 419

resinicola, 419

leaf feeders—
beetles:

Dichelonyx albicollis, 149
Diplotaxis sordida, 149
pine chafer, 150
white-spotted sawyer, 203

moths and butterflies:

Eacles imperialis pini, 304

Eupethecia palpatata, 337

European pine shoot moth, 12,
PAL PE SYA MAD), Gals oes
474, 475, 477, 478

hemlock looper, 344, 478, 483

jack-pine budworm, 381

Lapara bombycoides, 297

pine bud moth, 392

pine needle miner, 391

pine tussock moth, 320

pine webworm, 65, 351

595

Pine, jack—Continued

leaf feeders—Continued

moths and butterflies—Con.
Protobarmia porceiaria, 346
Rhyacionia adana, 362
Sparganothis tristriata, 375
spruce coneworm, 355
western-pine tip moth, 365,

478, 485

Xenotemna pallorana, 386
Zelleria haimbachi, 403

sawflies:

Acantholyda angulata, 435
luteomaculata, 4385
Cephalcia fulviceps, 435
Diprion similis, 23, 450
European pine sawfly, 12, 19,

24, 447, 470, 471, 4738
jack-pine sawfly, 19, 442
Neodiprion compar, 450
dubiosus, 447
maurus, 450
nigroscutum, 450
pratti paradoxicus, 443, 473
pratti pratti, 441
rugifrons, 447
red-headed pine sawfly, 19,
431, 439, 473
red-pine sawfly, 445, 473
Swaine jack-pine sawfly, 24,
444

pollen feeders—
jack pine budworm, 381
Xyela bakeri, 432

sucking insects—

Gnophothrips fuscus, 42
pine spittlebug, 71
pine tortoise scale, 103, 123
Saratoga spittlebug, 72, 472
Schizolachnus piniradiatae, 83
Pine, Japanese:
pine false webworm, 433
pine spittlebug, 71
Pine, Japanese black:

European pine shoot moth, 12, 21,
243612 4A Te Ale ate. £15,
A477, 478

red pine scale, 65, 95, 123

Pine, Japanese red:

Acantholyda angulata, 435

zappei, 434

European pine sawfly, 12, 19, 24,
447, 470, 471, 473

European pine shoot moth, 12, 21,
24, 361, 470, 471, 472, 474; 475,
477, 478

pine false webworm, 433

red-pine sawfly, 445, 473

red-pine scale, 65, 95, 123

Pine, loblolly:
ant, Texas leaf-cutting, 490
bark feeders—
Hylastes salebrosus, 252
tenuis, 252
pales weevil, 213
pitch-eating weevil, 217

borers, bark and phloem—

black turpentine beetle, 244

596

Pine, loblolly——Continued
borers, bark and phloem—Con.
deodar weevil, 212
Hylastes salebrosus, 252
tenuis, 252
Ips avulsus, 260
calligraphus, 260, 502
pales weevil, 213
pitch-eating weevil, 217
Pityoborus comatus, 258
Pityogenes meridianus, 258
Pityophthorus spp., 255
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
southern pine engraver, 260
borers, bud, twig and shoot—
beetles:
Pityophthorus spp., 255
pitch-eating weevil, 217
moths and butterflies:
Diorcyctria clarioralis, 354
European pine shcot moth, 12,
2 2A AGley Ae te Ae
474, 475, 477, 478
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
Rhyacionia rigidana, 363, 477,
478
subtropica, 365
Sparganothis sulfureana, 375
borers, cone and seed—
beetle, Conophthorus taedae, 258
moths and butterflies:
Dioryctria amatella, 356
clarioralis, 354
disclusa, 358, 475
Eucosma cocana, 368
Laspeyresia anaranjada, 372
erotella, 373
Zimmerman pine moth, 356
borers, gall—
Battaristis vittella, 394
Dioryctria amatella, 23, 355
borers, root—
black turpentine beetle, 244
lesser cornstalk borer, 358
borers, wood—
Buprestis lineata, 165
Ernobius tenuicornis, 133
Gnathotrichus materiarius, 268
Monochamus spp., 203
pales weevil, 213
turpentine borer, 164
Xyleborus saxeseni, 269
gall insects—
Pleroneura sp., 433
Xyela sp., 432
leaf feeders—
beetles:
Phyllophaga lurtuosa, 147
prununculina, 148
pine chafer, 150
pine colapsis, 142
turpentine borer, 164
moths and butterflies:
pine needle miner, 391
pine webworm, 65, 351

Pine, loblolly—Contiihued
leaf feeders—Continued
sawflies:
Acantholyda apicalis, 435
Neodiprion abbotii, 447
excitans, 19, 449
hetricki, 450
pratti pratti, 441
taedae linearis, 443, 473
taedae taedae, 443
red-headed pine sawfly, 19,
431, 439, 473
root feeders—
Phyllophaga luctuosa, 147
prununculina, 148
shoot feeders—
Pleroneura sp., 433
xyela sp., 432
sucking insects—
Cinara carolina, 83
taedae, 83
watsoni, 83
Dysmicoccus obesus, 99
Essigella pini, 83
Oracella acuta, 100
pine needle scale, 109
pine spittlebug, 71
pine twig gall scale, 65, 96
Saratoga spittlebug, 72, 472
Schizolachnus lanosus, 83
woolly pine scale, 106
terminal feeders—
Sparganothis sulfureana, 375
Pine, longleaf:
Ant, Texas leaf-cutting, 490
bark feeders—
Hylastes salebrosus, 252
tenuis, 252
pales weevil, 213
borers, bark and phloem—
beetles:
black turpentine beetle, 244
Hylastes salebrosus, 252
tenuis, 252
Ips avulsus, 260
calligraphus, 260, 502
pales weevil, 213
pitch-eating weevil, 217
Pityoborus comatus, 258
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
southern pine engraver, 260
moth, Dioryctria abietella, 25,
355
borers, bud and shoot—
beetles:
| Gnathotrichus
| 268
Pityophthorus spp., 255
| moths and butterflies:
Dioryctria abietella, 23, 355
| amatella, 356
) clarioralis, 354
European pine shoot moth, 12,
QZ A e361, 0 470.471, 472,
474, 475, 477, 478
Holcocera lepidophaga, 396

materiarius,

sisi rat

Pine, longleaf—Continued
borers, bud and shoot—Continued
moths and butterflies—Con.
Rhyacionia subtropica, 365
borers, cone—
Battaristis vittella, 394
Dioryctria abietella, 23, 355
amatella, 356
clarioralis, 354
disclusa, 358, 475
Holcocera lepidophaga, 396
Laspeyresia anaranjada, 372
ingens, 373
borers, gall—
Moodna ostrinella, 359
borers, wood—
Buprestis lineata, 165
Chrysobothris dentipes, 167
pales weevil, 213
southern pine sawyer, 203
turpentine borer, 164
flower feeders—
Holcocera lepidophaga, 396
leaf feeders—
beetles:
Phyllophaga luctuosa, 147
prununculina, 148
turpentine borer, 164
mite, Oligonychus cunliffei, 6
moths and butterflies:
Exotelia chilleotti, 392
pine needle miner, 391
pine webworm, 65, 351
sawflies:
Neodiprion abbott, 447
excitans, 19, 449
red-headed pine sawfly, 19,
431, 439, 473
Xyela bakeri, 432
minor, 4382
obscura, 482
root feeders—
Phyllophaga luctuosa, 147
prununculina, 148
sucking insects—
Brochymena carolinensis, 62
Cinara carolina, 83
Leptoglossus corculus, 66
Tetrya bipunctata, 62
woolly pine scale, 106
Pine, Mugho:
bark and phloem feeders—
pine root collar weevil, 213
borers, bud and shoot—
Battaristis vittella, 394
European pine shoot moth, 12,
21 2A e326 ATO ardent 2. 474,
475, 477, 478
Exotelia nepheos, 392
pine bud moth, 392
white-pine shoot borer, 367
white-pine weevil, 209, 423, 426,
475, 477
Zimmerman pine moth, 356
leaf feeders—
beetle, Anomala oblivia, 150
moths and butterflies:

597

Pine, Mugho—Continued
leaf feeders—Continued
moths and butterflies:
European pine shoot moth, 12,
Zi? CSG Ape Auman
474, 475, 477, 478
Exoteleia nepheos, 392
pine bud moth, 392
sawflies:
Diprion similis, 23, 450
European pine sawfly, 12, 19,
24, 447, 470, 471, 473
pine false webworm, 433
red-pine sawfly, 445, 473
white-pine sawfly, 446
root feeders—
pine root collar weevil, 213
sucking insects—
pine needle scale, 109
Toumeyella pini, 105
Pine, pitch:
Ant, black carpenter, 484, 486
borers, bark and phloem—
Ips spp., 260
pine root collar weevil, 213
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
southern pine engraver, 260
borers, bud and shoot—
beetles:
pine gall weevil, 218
Pityophthorus spp., 255
moths and butterflies:
Choristoneura pinus
tima, 383
Dioryltria spp., 355
European pine shoot moth, 12,
21, 2436 AOA 8472,
474, 475, 477, 478
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
Rhyacionia rigidana, 363, 477,
A478
sonia, 365
Zimmerman pine moth, 356
gall insects—
gouty pitch midge, 418
pine gall weevil, 218
leaf feeders—
beetles:
Anomala oblivia, 150
turpentine borer, 164
moths and butterflies:
Choristoneura pinus
tima, 383
Lambdina athasaria_ pelluci-
daria, 345
Lapara bombycoides, 297
pine needle miner, 391
pine tussock moth, 320
pine webworm, 65, 351
sawflies:
Acantholyda angulata, 435
zappei, 434
European pine sawfly, 12, 19,
HA W447 AAO Ale vA7S

mari-

mari-

098

Pine, piteh—Continued
leaf feeders—Continued
sawflies—-Continued
Neodiprion pinirigidae, 450,
473
pratti paradoxicus, 443, 473
pratti pratti, 441
red-headed pine sawfly, 19,
431, 439, 473
white-pine sawfly, 446
sucking insects—
Cinara watsoni, 83
Eulachnus riley, 83
Matsucoccus matsumurae, 96
pine spittlebug, 71
pine twig gall scale, 65, 96
Pineus coloradensis, 91
Saratoga spittlebug, 72, 472
Pine, pond:
Cinara carolina, 83
watson, 83
Essigella pini, 83
Neodiprion excitans, 19, 449
hetricki, 450
Pine, ponderosa:
European pine sawfly, 12, 19, 24,
447, 470, 471, 473
European pine shoot moth, 12, 21,
ZA 361 AOA Ac AA Arion
477, 478
Gnathotrichus aciculatus, 269
pine twig gall scale, 65, 96
western-pine tip moth, 365, 478,
485
Pine, red:
Ant, Allegheny mound, 491
bark feeders—
Northern pine weevil. 211
pales weevil, 213
pine root collar weevil, 213
Pissodes affinis, 213
white-pine weevil, 209, 423, 426,
475, 477
white-spotted sawyer, 203
borers, bark and phloem—
Chrysobothris orono, 166
Dioryctria abietella, 23, 355
Hylobius congener, 217
Ips latidens, 262
perroti, 262
Myeloborus fivazi, 259
Pityophthorus balsameus, 255
biovalis, 255
cariniceps, 255
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
white-pine weevil, 209, 423, 426,
ATS, 477
borers, bud, twig and shoot—
beetles:
Conophthorus vesinosa, 257
Myeloborus fivazi, 259
pine gall weevil, 218
Pityophthorus spp., 255
white-pine weevil, 209, 423,
426, 475, 477

Pine, red—Continued
borers, bud, twig, and shoot—Con.

Pine, red—Continued
leaf feeders—Continued

moths and butterflies:
Dioryctria abietella, 23, 355
cambuicola, 358
European pine shoot moth,
22h, 24 SOle a0. aa
472, 474, 475, 477, 478
Exoteleia nepheos, 392
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
pine bud moth, 392
Rhyacionia adana, 362
rigidiana, 363,
477, 478
Sparganothis sufureana, 375
western-pine tip moth, 365,
478, 485
Zimmerman pine moth, 356

borers, cone and seeds—

Conophthorus resinosae, 257
Dioryctria abietella, 23, 355
cambiicola, 358
disclusa, 358, 475

borers, root—

Hylobius rhizophagus, 216
pine root collar weevil, 213

borers, wood—

Acanthocinus pusillus, 196
Chrysobothris orono, 166
Myeloborus fivazi, 259
northern pine weevil, 211
pales weevil, 213
pine root collar weevil, 2138
Pissodes affinis, 2138
Pityogenes hopkinsi, 258
Pityophthorus balsameus, 255
biovalis, 255
cariniceps, 255
white-pine weevil, 209, 423, 426,
475, 477
white-spotted sawyer, 203
Zimmerman pine moth, 356

moths and butterflies—Con.
pine webworm, 65, 351
Rhyacionia adana, 362
rigidiana, 363,
AT7, 478
Sparganothis acerivorana,
375, 386
Xenotemna pallorana, 386
sawflies—
Acantholyda pini, 4385
zappei, 434
Cephalcia fulviceps, 435
marginata, 435
Diprion frutetorum, 28, 451,
473
similis, 28, 450
European pine sawfly, 12, 19,
24, 447, 470, 471, 478
jack-pine sawfly, 19, 442
Neodiprion abbott, 447
compar, 450
pratti pratti, 441
pine false webworm, 433
red-headed pine sawfly, 19,
431, 439, 473
red-pine sawfly, 445, 473
Swaine jack-pine sawfly, 24,
444
white-pine sawfly, 446

pollen feeders—

jack pine budworm, 381

root feeders—

Diplotaxis sordida, 149
Phyllophaga spp., 146

sucking insects—

Cinara watsoni, 83
Eulachnus agilis, 83

rileyi, 83
pine needle scale, 109
pine tortoise scale, 103, 123
Pineas coloradensis, 91

Bee eeu 218 red-pine scale, 65, 95, 123
Thecodiplosis piniresinosae, 418 Saratoga spittlebug, 72, 472
leat fecders—— Schizolachnus piniradiatae, 83

beetles: Pine, sand:
Anomala oblivia. 150 Acantholyda circumcincta, 4385
Diplotaxis sordida, 149 Neptyia semiclusaria, 344
white-spotted sawyer, 203 Pine, Scotch:

moth and butterflies: ant, Allegheny mound, 491

chain-spotted geometer, 345 bark feeders—

European pine shoot moth,
ae 24-361, 470, 471;
472, 474, 475, 477, 478

Exoteleia nepheos, 392

jack-pine budworm, 381

Lambdina athasaria_ pelluci-
daria, 345

Lapara bombycoides, 297

Nantucket pine tip moth, 363,
472, 475, 477, 478, 485

oblique-banded leaf roller,
383

pine bud moth, 392

pine needle miner, 391

pine tussock moth, 320

pales weevil, 213
white-pine weevil, 209, 423, 426,
475, 477

borers, bark and phloem—

Hylobius congener, 217
warren, 216

Ips latidens, 262

northern pine weevil, 211

pales weevil, 213

pine root collar weevil, 213

pitch mass borer, 399

red turpentine beetle, 246

southern pine engraver, 260

white-pine weevil, 209, 423, 426,

475, 477

599

Pine, Scotch—Continued
borers, bud and shoot—
beetle, white-pine weevil, 209,
423, 426, 475, 477
moths and butterflies—
European pine shoot moth, 12,
Zee 24s 361s e400, 4 Aue,
474, 475, 477, 478
Exoteleia nepheos, 392
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
pine bud moth, 392
Rhyacionia adana, 362
rigidana, 363, 477,
478
western-pine tip moth, 365,
478, 485
white-pine shoot borer, 367
Zimmerman pine moth, 356
borers, cone—
Battaristis vittella, 394
Dioryctria disclusa, 358, 475
borers, wood—
beetles:
Hylobius congener, 217
warreni, 216
northern pine weevil, 211
pine root collar weevil, 213
Pissodes affinis, 213
white-pine weevil, 209, 423,
A226 54,0414
moths and butterflies:
pitch mass borer, 399
pitch nodule maker, 366
Zimmerman pine moth, 356
leaf feeders—
beetle:
Magdalis austera substiga,
225
moths and butterflies:
European pine shoot moth, 12,
Pa Pa EY YA Nal AW
474, 475, 477, 478
Exoteleia nenheos, 392
jack-pine budworm, 381
Nantucket pine tip moth, 363,
Aaa Als A Us Alor 400
oblique-banded leaf roller, 383
pine bud moth, 392
pine needle miner, 391
pine webworm, 65, 351
Rhyacionia adana, 362
rigidana, 363, 477,

478

Sparganothis acerivorana,
375, 386

western-pine tip moth, 365,
478, 485

Xenotemna pallorana, 386

sawflies:

Diprion frutetorum, 28, 451,

473

similis, 23, 450
European pine sawfly, 12, 19,
24, 447, 470, 471, 473
jack-pine sawfly, 19, 442
Neodiprion pratti paradoxi-
cus, 443, 445

600

Pine, Scotch—Continued
leaf feeders—Continued
sawflies—Continued
pine false webworm, 433
Swaine jack-pine sawfly, 24,
444
root feeders—
pine root collar weevil, 213
sucking insects—
Cinara pini, 83
watsoni, 83
Eulachnus agilis, 83
riley, 83
meadow spittlebug, 74
pine bark aphid, 89
pine needle scale, 109
pine spittlebug, 71
pine tortoise scale, 103, 123
Toumeyella pine, 105
Pine, scrub:
Buprestis lineata, 165
_Chrysobothris dentipes, 167
Dioryctria disclusa, 358, 475
Janetiella coloradensis, 420
pine gall weevil, 218
pine needle miner, 391
red turpentine beetle, 246
Pine, shortleaf:
bark feeders
deodar weevil, 212
Hylastes salebrosus, 252
tenuis, 252
pales weevil, 213
pitch-eating weevil, 217
borers, bark and phloem—
beetles:
black turpentine beetle, 244
Hylastes porculus, 252
salebrosus, 252
tenuis, 252
Ips avulsus, 260
calligraphus, 260, 502
northern pine weevil, 211
Orthotomicus calceatus, 2638
pales weevil, 213
Pityogenes hopkinsi, 258
Pityophthorus puberlus, 255
pulicarius, 255
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
southern pine engraver, 260
moth:
Petrova houseri, 367
borers, bud, twig and shoot—
beetle:
Pityophthorus spp., 255
moths and butterflies:
Dioryctria clarioralis, 354
Nantucket pine tip moth,
363, 472, 475, 477, 478, 485
Petrova houseri, 367 )
borers, cone—
Dioryctria amatella, 356
clarioralis, 354

Pine, shortleaf—Continued
borers, wood—
beetles:
Chrysobothris dentipes, 167
pusilla, 167
Gnathotrichus materiarius,
268
Monochamus titillator, 203
northern pine weevil, 211
pales weevil, 213
Pityoborus comatus, 258
Trypodendron scabricollis,
268
turpentine borer, 164
Xyleborus saxeseni, 269
horntail:
Sirex nigricornis, 466
gall insects—
Pleuroneura sp., 433
AXyela sp., 432
leaf feeders—
beetles:
Glyptoscelis pubescens, 145
pine colaspis, 142
turpentine borer, 164
moths and butterflies:
Lambdina athasaria_ pelluci-
daria, 345
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
pine needle miner, 391
pine webworm, 65, 351
sawflies:
European pine sawfly, 12, 19,
24, 447, 470, 471, 473
Neodiprion abbotu, 447
excitans, 19, 449
pinirigidae, 450, 473
pratti paradoxicus, 443, 473
pratti pratti, 441
taedae linearis, 443, 445
warren, 450
red-headed pine sawfly, 446
white-pine sawfly, 446
pollen feeder—
Pomphopoea polita, 121
shoot feeder—
Pleuroneura sp., 433
Xyela sp., 432
sucking insects—
Cinara carolina, 83
pini, 83
watsoni, 83
Eulachnus rileyi, 83
pine twig gall scale, 65, 96
Pine, slash:
bark feeders—
pales weevil, 213
borers, bark and phloem—

beetles:
black turpentine beetle, 244
Ips spp., 260

pales weevil, 213

Pityoborus comatus, 258

Pityophthorus spp., 255

red turpentine beetle, 246

southern pine beetle, 240, 477,
502

Pine, slash—Continued
borers, bark and phloem—Con.
moth, Dioryctria abietella, 28,

5
borers, bud and shoot—
beetle:
Pityophthorus spp., 255
moths and butterflies:
Dioryctria abietella, 28, 355
clarioralis, 354
Holcocera lepidophaga, 396
Nantucket pine tip moth,
363, 472, 475, 477, 478, 485
Rhyacionia rigidana, 363, 477,
478
subtropica, 365
borers, cone—
Dioryctria abietella, 23, 355
amatella, 356
clarioralis, 354
Holcocera lepidophaga, 396
Laspeyresia anaranjada, 372
ingens, 378
borers, gall—
Dioryctria amatella, 356
Moodna ostrinella, 359
borers, root—
Hylobius aliradicis, 217
borers, wood—
pales weevil, 213
Pityoborus comatus, 258
southern pine sawyer, 263
turpentine borer, 164
cone feeders—
Gnopothrips fuscus, 42
Holcocera lepidophuga, 396
flower feeders—
Holcocera lepidophaga, 396
thrips spp., 41, 43
Xyela bakeri, 432
minor, 432
obscura, 432
gall insects—
Pleroneura sp., 433
AYela Sp: u4dz
leaf feeders—
ant, Texas leaf-cutting, 490
beetles:
pine colaspis, 142
cricket, short-tailed cricket, 60
moths and butterflies:
Apantesis radians, 307
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
pine webworm, 65, 351
sawflies:
Neodiprion obboti, 447
excitans, 19, 449
merkeli, 450
red-headed pine sawfly, 19,
431, 439, 473
shoot feeder—
Pleroneura sp., 433
Ayela sp., 432
sucking insects—
Brochymena carolinensis, 62

601

Pine, slash—Continued
sucking insects—Continued
Cinara carolina, 83
watsoni, 83
Essigella pini, 83
Leptoglossus corculus, 66
pine needle scale, 109
pine spittlebug, 71
Tetrya bipunctata, 62
thrips sp., 41, 43
Pine, Sondregger:
Neodiprion excitans, 19, 449
Pine, Tablemountain:
European pine sawfly, 12, 19, 24,
447, 470, 471, 4738
northern pine weevil, 211
pine twig gall scale, 65, 96
southern pine beetle, 240, 477, 502
Pine, Virginia:
bark feeders—
pales weevil, 213
borers, bark and phloem—
black turpentine beetle, 244
Crypturgus alutaceous, 239
pine gall weevil, 218
Pityophthorus spp., 255
red turpentine beetle, 246
southern pine beetle, 240, 477,
502
southern pine engraver, 260
borers, bud and shoot—
Choristoneura pinus maritima,
383
European pine shoot moth, 12,
QU AZ4 S61 v4 ads aia 44.
475, 477, 478
Nantucket pine tip moth, 363,
472, 475, 477, 478, 485
Petrova albicapitana, 366
Pityophthorus spp., 255
Rhyacionia rigidana, 363, 477,
478
borers, cone—
Conophthorus virginianae, 258
borers, wood—
Chrysobothris blanchardi, 167
harrisi, 167
northern pine weevil, 211
Platypus flavicornis, 272
gall insects—
Cecidomyia reeksi, 419
resinicola, 419
gouty pitch midge, 418
pine gall weevil, 218
leaf feeders—
beetle:
Glyptoscelis pubescens, 145
grasshopper, Dendrotettix aus-

tralis, 58
moths and butterflies:
Choristoneura pinus mari-
tima, 383

European pine shoot moth,
V2 a 24a alee a Omar.
475, 477, 478

Nantucket pine tip moth, 363,
Alle, 41D MA Aton 4oD

pine needle miner, 391

602

Pine, Virginia—Continued
leaf feeders—Continued
moths and butterflies—Con.
pine webworm, 65, 351
Rhyacionia rigidana, 363, 477,
478
sawfly, Neodiprion pratti pratti,
441

pollen feeders—
Xyela minor, 432
obscura, 432
styrax, 432
sucking insects—
Cinara carolina, 83
pini, 83
taedae, 83
watsoni, 83
Eulachnus rileyi, 83
Matsucoccus matsumurae, 96
pine needle scale, 109
pine spittlebug, 71
pine swig gall scale, 65, 96
Saratoga spittlebug, 72, 472
Toumeyella sp. near numismati-
cum, 105
Plane tree, London:
American plum borer, 359
sycamore tussock moth, 306
Plum:
borers, bark and phloem—
peach bark beetle, 240
Scolytus mali, 237
shot hole borer, 238
borers, fruit—
apple fruit moth, 403
apple maggot, 424
plum curculio, 227
borers, wood—
American plum borer, 359
banded hickory borer, 178
leopard moth, 386
Synanthedon pictipes, 399
flower and fruit feeders—
Coleophora serratella, 406
gall insect—
Aceria phloceoptes, 8
leaf feeders—
beetles:
European snout beetle, 219
Polydrusus impressifrons, 28,
220
Pyrrhalta cavicollis, 140
moths and butterflies:
apple-and-thorn skeletonizer,
396
Argyrotaenia quadrifasciana,
38

brown-tail moth, 24, 325, 4380,
431, 469, 472, 476

Coleophora serrateila, 406

eynthia moth, 297

eastern tent caterpillar, 328,
476, 477, 483

eye-spotted but moth, 367

hickory leaf roller, 384

Lithocolletis crataegella, 407

oriental moth, 347, 349, 431,
484

Plum—Continued
leaf feeders—Continued
moths and butterflies—Con.
palmerworm, 393
prairie tent caterpillar, 330
Sphinx drupiferarum, 297
unspotted leaf miner, 408
viceroy, 293
western tent caterpillar, 330
sawflies:
Neurotoma inconspicua, 485
pear slug, 457
root feeders—
European snout beetle, 219
Polydrusus impressijrons, 28,
220
Plum, beach:
brown-tail moth, 24, 325, 430, 431,
469, 472, 476
Sphinx drupiferarum, 297
Poplar (Populus spp):
ant, black carpenter, 484, 486
bark feeder—
poplar borer, 185, 483
borers, bark and phloem—
broad-necked root borer, 200
bronze poplar borer, 162
Cucujus clavipes, 124
Micracis swainei, 253
poplar borer, 185, 483
borers, shoot and twig—
Gypsonoma haimbachiana, 369
Micrasis swainei, 253
borers, root—
beetles:
Agrilus horni, 162
broad-necked root borer, 200
Polydrusus impressifrons, 23,
220
moths:
Aegeria tibialis, 398
hornet moth, 398
borers, wood—
beetles:
Agrilus horni, 162
Dicerca tenebrica, 168
Lyctus spp., 127
Micracis swainei, 253
oak timberworm, 208
Oberea schaumi, 190
Platypus compositus, 273
Poecilonota cyanipes, 169
poplar borer, 185, 483
poplar-butt borer, 202
poplar and willow borer, 218
Ptilinus pruinosus, 133
Saperda concolor, 188
moesta, 188
sapwood timberworm, 126
Sinodendron rugosum, 145
twig girdler, 192
Trypodendron retusum, 268
Xylotrechus annosus, 203
moths and butterflies:
Acossus centerensis, 388
populi, 388
Aegeria tibialis, 398
American plum borer, 359

22

Poplar—Continued
borers, wood—Continued
moths and butterflies—Con.
carpenterworm, 388
Conopia acerrubri, 402
hornet moth, 398
leopard moth, 386
Paranthrene dollii castanea,
402
dollui dollu, 402
tabaniformis, 402
tricincta, 402
sawflies:
Urocerus cressoni, 466
willow shoot sawfly, 467
bud and blossom feeders—
Eriophyes parapopuli, 7
fruit-tree leaf roller, 377
gall insects—
Ectodemia populella, 4138
Eriophyes parapopuli, 7
Melanogromyza schineri, 427
Pemphigus spp., 79
poplar vagabond aphid, 85
Prodiplosis morrisi, 420
Saperda concolor, 188
moesta, 188
leaf feeders—
beetles:
bronze poplar borer, 162
Chrysomela crotchi, 22, 123
knabi, 138
cottonwood leaf beetle, 138
European snout beetle, 219
goldsmith beetle, 152
Graphognathus spp., 224
imported willow leaf beetle,
143, 477
Japanese beetle, 19, 151, 485,
486
Polydrusus impressifrons, 23,
220
Zengophora scutellaris, 28, 144
butterflies:
mourning-cloak butterfly, 293
Nymphalis j-album, 292
tiger swallowtail, 291
viceroy, 293
mites:
Eotetranychus populi, 7
weldoni, 7
moths:
Abbottana clementaria, 346
Acronicta distans, 308
leporina vulpina,
308
Anacampsis innocuella, 393
Anacamptodes larvaria, 338
Archips purpuranus, 378
aspen blotch miner, 407
aspen leaf miner, 408
bagworm. 21. 41) e4v25 410;
477, 478
Brephos infans, 334
Bruce spanworm, 336
cecropia moth, 298, 477
Cerura spp., 318
chain-spotted geometer, 345

603

Poplar—Continued
leaf feeders—Continued

604

moths—Continued
cottonwood dagger moth, 308
Deilinea erythemaria, 337
variolaria, 338
Deuteronomos magnarius, 346
Ectropis crepuscularia, 339
Epinotia solandriana, 371
Evora hemidesma, 360
Fentonia marthesia, 316
forest tent caterpillar, 19, 330,
429, 472, 475
fruit-tree leaf roller, 377
Gluphisia septentrionis, 317
Gracillaria purpuriella, 409
gypsy moth, 12, 18, 19, 24, 25,
28) 2G2e MOO ML Leena dees 2 OF
430, 481, 469, 472, 474, 475,
476, 483
Hemileuca nevadensis, 300
hickory tussock moth, 304
Hyperaeschra stragula, 313
Ichthyura albosigma, 311
apicalis, 311
brucei, 311
strigosa, 311
io moth, 299
lappet moth, 333
large aspen tortrix, 383
Limenitis arthemis, 294
astyanax, 294
Lithocolletis tremuloidiella,
407
Lycia ursaria, 342
Notodonta simplaria, 318
oblique-banded leaf roller, 383
oriental moth, 347, 349, 431,
484
Paonias excaecatus, 297
pepper and salt moth, 346
Pero morrisonarius, 344
Pheosia rimosa, 313
Plagodis serinaria, 342
poplar tent maker, 310
Prochoerodes transversata,
346
Protobarmia porcelaria, 346
Psilocorsis fletcherella, 395
Raphia frater, 309
red-humped caterpillar, 316
saddled prominent, 316, 472
satin moth, 24, 324, 430, 470,
472, 475, 483
Schizura ipomaeae, 317
leptinoides, 317
smeared dagger moth, 308
Smerinthus jamaicensis, 297
Sparganothis fulgidipenna,
375
Sphinx luscitiosa, 296
spotted tussock moth, 305
Tetralopha asperatella, 62,
351, ab52
Tolype velleda, 333
unicorn caterpillar, 317
western tent caterpillar, 330

Poplar—Continued
leaf feeders—Continued
moths—Continued
white-marked tussock moth,
318, 476, 478, 483
winter moth, 336
sawflies:
elm sawfly, 438, 472
Messa populifoliella, 457
Nematus fulvicrus, 461
populi, 461
salicisodoratus, 461

Trichiocampus gregarius, 461
viminalis, 23,

461

Trichiosoma triangulum, 438

willow sawfly, 461
walkingsticks, 55
sucking insects—

Chaitophorus populicola, 84
steveusis, 84

Chionaspis salicisnigrae, 109

Comstock mealybug, 100

Corythucha elegans, 64

cottony maple scale, 101

European fruit lecanium, 105,

475

Idiocerus, spp., 69

Lachnus spp., 82

Macropsis spp., 69

oystershell scale, 109, 110, 475

Pemphigus spp., 79

poplar vagabond aphid, 85

Pterocomma populifoliae, 84
smithae, 84

Putnam scale, 108

Quadraspidiotus gigas, 107

San Jose scale, 106, 475

Saratoga spittlebug, 72, 472

scurfy scale, 108

Telamona decorata, 71

terrapin scale, 105

walnut scale, 107

Poplar, tulip, see Yellow-poplar

Redbud:
borers, bark and phloem—
Hypothenemus chapuisi, 254
dissimilis, 254
eruditus, 254
Pityophthorus natalis, 255
borers, seed—
Gibbobruchus mimus, 137
borers, wood—
Agrilus otiosus, 163
Hypothenemus chapuisi, 254
dissimilis, 254
eruditus, 254
Micracis meridianus, 2538
suturalis, 2538
swainet, 253
Micracisella nanula, 258
opacicollis, 253
Pityophthorus natalis, 255
Ptosima gibbicollis, 169
Thysanoes fimbricornis, 2538
leaf feeders—
Fascista cercerisella, 394

Redbud—Continued

leaf feeders—Continued
grape leaf folder, 350
Japanese weevil, 219
Norape ovina, 350

sucking insects—
European fruit lecanium, 105,

475
periodical cicada, 74
Prosapia bicincta, 73
San Jose scale, 106, 475
terrapin scale, 105
Rhododendron:

borers, root—
rhododendron stem borer, 190

borers, wood—
pitted ambrosia beetle, 266
rhododendron borer, 398
rhododendron stem borer, 190

leaf feeders—
black vine weevil, 220
Japanese weevil, 219
Orthosia hibisci, 309
Sphinx kalmiae, 296

root feeders—
Asiatic garden beetle, 149, 485
black vine weevil, 220
Japanese weevil, 219

sucking insects—

Kleidocerys resedae geminatus,
67

rhododendron lacebug, 64
rhododendron whitefly, 92

Sassafras:
borers, bark and phloem—
Hypothenemus chapuisi, 254
borers, root—
Oberea ruficollis, 190
borers, wood—
Hypothenemus chapuisi, 254
Oberea ruficollis, 190
pitted ambrosia beetle, 266
Scobicia bidentata, 136
leaf feeders—
beetles:
Graphognathus spp., 224
Odontopus calceatus, 221
white-fringed beetles, 224
moths:
cecropia moth, 298, 477
Datana drexeli, 312
Epimecis virginiara, 339
Gracillaria sassafrasella, 409

gypsy moth, 12, 21, 24, 361,
ATO471, 472, 474, 475, 477,

478
Pandemis lamprosana, 375
polyphemus moth, 299
promethea moth, 298
spice-bush swallowtail, 291
sucking insect—
European fruit lecanium, 105,
475
Seedlings, Various:
leaf feeders—
ants, Texas leaf-cutting, 490

Seedlings, various—Continued
leaf feeders—Continued
beetles:
blister beetles, 120
Epicauta pestifera, 120
vittata, 120
European chafer, 150
flatheaded borer, 156
Macrobasis fabricii, 120
unicolor, 120
pine chafer, 150
Pomphopoea aenea, 120
sayi, 120
Serica vespertina, 149
white-fringed beetles, 224
grasshoppers, 57
moths and butterflies:
Nantucket pine tip moth, 363,
AV2. A475, ATT, 478,483
cutworms, 309
sawfly:
introduced pine sawfly, 28,
450, 476
tip or twig feeders—
juniper midge, 418
root feeders—
beetles:
Anomala spp., 150
Asiatic garden beetle, 149,
485
Diplotaxis spp., 149
Graphognathus, spp., 224
green June beetle, 152
Japanese beetle, 19, 151, 485,
486
Phyllophaga spp.. 147
Polyphylla spp., 149
Serica spp., 149
white- fringed beetles, 224
wireworms, 153
cricket, European mole, 60
stem feeders—
Geopinus incrassatus, 118
Harpalus spp., 118
Prionus spp., 200
Scarites subterraneus, 118
seed-corn maggot, 427
termites, 45
stem and twig girdlers—
cutworms, 309
grasshoppers, 57
lesser cornstalk borer, 358
sucking’ insects—
aphids; lL92043 G45 967, ise va,
424, 472
seales, 67, 121, 424
spider mites, 6
Shadbush:
bark feeder—
roundheaded apple tree borer,
187
borers, bark and phloem—
Synanthedon pyri, 399
borers, cambium—
Phytobia amelanchieris, 426
borers, fruit—
apple fruit moth, 403

605

Shadbush—Continued
borers, fruit—Continued
Hoplocampa halcyon, 464
pallipes, 465
borers, root—
oak sapling borer, 183
borers, seed—
Megastigmus amelanchieris, 476
borers, wood—
oak sapling borer, 183
roundheaded apple tree borer,
187
Synanthedon pyri, 399
gall insects—
Eriophyes pyri, 7
leaf feeders—
beetles:
Phyllophaga spp., 146
roundheaded apple tree borer,
187
willow flea weevil, 222
mite:
Oligonychus newcomeri, 6
moths and butterflies:
Argyrotaenia quadrifasciana,
385
Bruce spanworm, 330
Bucculatrix pomifoliella, 409
western tent caterpillar, 330
sawfly:
pear slug, 457
sucking insects—
Chionaspis salicisnigrae, 109
Prociphilus fraxinifolii, 85
woolly elm aphid, 84
Sourwood:
dogwood twig borer, 189
twig girdler, 191, 502
hickory horned devil, 303
regal moth, 303
Spruce (Picea spp.):
ant, Allegheny mound, 491
bark feeders—
Acanthocinus pusillus, 196
Hylastes salebrosus, 252
tenuis, 252
Hylurgops pinifex, 252
pales weevil, 213
white-pine weevil, 208, 4238, 426,
475, 477
borers, bark and phloem—
Acanthocinus pusillus, 196
Crypturgus atomus, 239
Dryocoetes affaber, 263
autographus, 263
granicolhis, 264
eastern spruce beetle, 247
four-eyed spruce bark beetle,
264
Hylastes salebrosus, 252
tenuis. 252
Hylurgops pinifex, 252
Ips borealis, 262
Melanophila acuminata. 169
drummondi, 169
Neoclytus muricatulus, 184
Orthotomicus caelatus, 263
pales weevil, 213

606

Spruce—Continued
borers, bark and phloem—Con.
Phymatodes dimidiatus, 197
pine engraver, 262
pitch mass borer, 399
Pityokteines sparsus, 263
Pityophthorus opaculus, 255
red turpentine beetle, 246
spruce scolytus, 239
borers, cone and seed—
Dioryctria abietella, 28, 355
Eucosoma tocullionana, 368
spruce coneworm, 355
spruce seed moth, 371
borers, root—
pales weevil, 213
Thylacites incanus, 238, 220
borers, wood—
beetles:
Anoplodera canadensis, 196
peer juniper borer,
19
black-horned pine borer, 194
Chrysobothris scabripennis,
167
trinervia, 167
Dicerca tenebrosa, 168
Ernobius mollis, 1338
Gnathotrichus materiarius,
268
hemlock borer, 168, 483
Hylotrupes, bajulus, 22, 206
Melanophila acuminata, 169
drummondi, 169
Monochamus spp., 203
Neoclytus muricatulus, 198
old house borer, 206
Semanotus litigiosus, 198
spruce timber beetle, 268
striped ambrosia beetle, 267
white-spotted swayer, 203
Xylotrechus undulatus, 203
horntails:
Sirex cyaneus, 466
juvencus, 23, 466
Urocerus albicornis, 466
flavicornis, 466
moths:
Dioryctria abietella, 28, 355
pitch mass borer, 399
bud feeder—
spruce bud moth, 369
gall insects—
Cooley spruce gall aphid, 87, 91
Pineus floccus, 91
similis, 91
leaf feeders—
beetles:
European chafer, 150
pales weevil, 213
mite, spruce spider
moths:
Argyrotaenia occultana, 385
bagworm, 21, 411, 472, 475,
477, 478
black-headed budworm, 385,
472
Epinotia nanana, 23, 371

__ ae

Spruce—Continued
leaf feeders—Continued
moths—Continued
Epizeuxis aemula, 309
Eufidonia notataria, 338
Feralia jocosa, 309
gypsy moth, 12, 18, 19, 24, 25,
Z287O2 805) 112343205 430),
431, 469, 472, 474, 475, 476,
483
hemlock looper, 344, 478, 483
Lexis bicolor, 307
Melanolophia canadaria, 338
Neptyia semiclusaria, 344
Panthea acronyctoides, 309
Pero morrisonarius, 344
pine tussock moth, 320
Semiothisa granitata, 338
spruce bud moth, 369
Zeiraphera diniana, 370
sawflies:
balsam-fir sawfly, 446, 473
green-headed spruce sawfly,
460
yellow-headed spruce sawfly,
459, 472
pollen feeder—
Xyela alpigena, 432
root feeders—
European chafer, 150
Serica tristis, 149
strawberry root weevil, 221
sucking insects—
Adelges strobilobius, 22, 89
balsam twig aphid, 83
Cooley spruce gall aphid, 87, 91
Fiorinia externa, 111
hemlock scale, 108
pine needle scale, 109
Pineus floccus, 91
Spruce, bigcone, see Douglas-fir
Spruce, black:
bark feeder—
white-spotted sawyer, 203
borers, bark and phloem —
Crypturgus alutaceus, 239
eastern spruce beetle, 247
hemlock borer, 168, 483
northern pine weevil, 211
Orthotomicus vicinus, 263
borers, cone and seed—
Coleotechnites piceaella, 391
spruce coneworm, 355
spruce seed moth, 371
borers, twig and shoot—
white-pine weevil, 209, 423, 426,
475, 477
borers, wood—
hemlock borer, 168, 483
northern pine weevil, 211
Orthotomicus vicinus, 263
white-spotted sawyer, 203
leaf feeders—
balsam-fir sawfly, 446, 473
black-headed budworm, 385, 472
Coleotechnites piceaella, 391
European spruce sawfly, 18, 19,
24, 431, 452, 470, 471, 4738, 477

Spruce, black—Continued
leaf feeders—Continued
Eupethecia transcanadata, 337
false hemlock looper, 344
hemlock looper, 344, 478, 483
red-banded leaf roller, 384
spruce budworm, 25, 378, 472,
AT7
spruce coneworm, 355
spruce needleminer, 360
white-spotted sawyer, 203
yellow-headed spruce sawfly,
459, 472
sucking insects—
Adelges strobilobius, 22, 89
Cinara abietis, 83
pine leaf chermid, 91
Pineus floccus, 91
similis, 91
Spruce, blue (Colorado spruce) :
borers, cone and seed—
Coleotechnites piceaella, 351
spruce coneworm, 355
spruce seed moth, 371
borers, twig and shoot—
white-pine weevil, 209, 423, 426,
ATD 4
leaf feeders—
Cephalcia fascipennis, 485
Coleotechnites piceaeilla, 351
Epinotia nanana, 23, 371
gypsy moth; 12) a8) 19.724" 125.
280 02snGow 17 612351320.0 430)
431, 469, 472, 474, 475, 476,
483
spruce needleminer, 360
yellow-headed spruce sawfly,
459, 472
root feeders—
strawberry root weevil, 221
sucking insects—
Cooley spruce gall aphid, 87, 91
eastern spruce gall aphid, 86, 89
Pineus similis, 91
Spruce, Engelmann:
borers, bark and phloem—
Dryocoetes caryi, 264
Scolytus piceae, 239
borers, cone and seed—
Coleotechnites piceaella, 351
spruce coneworm, 355
spruce seed moth, 371
leaf feeders—
Coleotechnites piceaella, 351
spruce coneworm, 355
spruce needleminer, 360
yellow-headed spruce sawfly,
459, 472
sucking insects—
Cooley spruce gall aphid, 87, 91
Pineus similis
Spruce, Norway:
borers, bark and phloem—
Crypturgus alutaceus, 239
southern pine beetle, 240, 477,
502
white-pine weevil, 209, 423, 426,
ATO ALT

607

Spruce, Norway—Continued
borers, cone and seed—
Coleotechnites piceaella, 351
spruce coneworm, 355
borers, twig and shoot—
white-pine weevil, 209, 423, 426,
A475, 477
leaf feeders—
Coleotechnites piceaella, 351
Epinotia nanana, 23, 371
European spruce sawfly, 18, 19,
24, 431, 452, 470, 471, 473, 477
red-headed pine sawfly, 19, 431,
439, 473
spruce coneworm, 355
spruce needleminer, 360
yellow-headed spruce sawfly,
459, 472
sucking insects—
Adelges lariciatus, 89
eastern spruce gall aphid, 86, 89
pine spittlebug, 70
Pineus similis, 91
Spruce, ornamental:
Cephalcia fascipennis, 435
pine colaspis, 142
Spruce, red:
bark feeder—
white-spotted sawyer, 203
borers, bark and phloem—
black turpentine beetle, 244
Dryocoetes caryi, 264
eastern larch beetle, 248
eastern spruce beetle, 247
four-eyed spruce bark beetle,
264
hemlock borer, 168, 483
northeastern sawyer, 205
northern pine weevil, 211
Pissodes dubius, 2138
fiskei, 213
rotundatus, 213
Pityogenes hopkinsi, 258
Pityophthorus balsameus, 255
biovalis, 255
dentifrons, 255
patchi, 255
pulchellus, 255
shepardi, 255
southern pine beetle, 240, 477,
502
spruce scolytus, 239
Taenioglyptes rubentis, 259
ruficollis, 259
white-spotted sawyer, 2038
borers, cone and seed—
Coleotechnites piceaella, 391
spruce coneworm, 355
spruce seed moth, 371
borers, root—
black turpentine beetle, 244
borers, twig and shoot—
Pityophthorus spp., 255
white-pine weevil, 209, 423, 426,
AT OAT,

608

Spruce, red—Continued
borers, wood—
Anoplodera minnesotana, 195
mutabilis, 197
vittata, 197
Callidium violaceum, 195
hemlock borer, 168, 483
northeastern sawyer, 205
northern pine weevil, 211
Pityophthorus balsameus, 255
biovalis, 255
dentifrons, 255
patchi, 255
pulchellus, 255
shepardi, 255
white-spotted sawyer, 203
Xyloterinus politus, 264, 267
leaf feeders—
black-headed budworm, 385, 472
Coleotechnites piceaella, 391
Epinotia nanana, 238, 371
European spruce sawfly, 18, 19,
24, 431, 452, 470, 471, 473, 477
hemlock looper, 344, 478, 483
red-banded leaf roller, 384
spruce budworm, 25, 378, 472,
AT7
spruce coneworm, 355
white-spotted sawyer, 203
yellow-headed spruce _ sawfly,
459, 472
sucking insects—
Adelges strobilobius, 22, 89
eastern spruce gall aphid, 86, 89
pine leaf chermid, 91
pine spittlebug, 71
Pineus simiiis, 91
Spruce, Sitka:
Cooley spruce gall aphid, 87, 91
spruce coneworm, 355
spruce seed moth, 371
Spruce, white:
bark feeder—
Taenioglyptes ruficollis, 259
borers, bark and phloem—
Dryocoetes caryi, 264
eastern spruce beetle, 247
hemlock borer, 168, 483
Ips latidens, 262
perturbatus, 262
northern pine weevil, 211
Pityophthorus shepardi, 231
spruce scolytus, 239
Taenioglyptes rufitcollis, 259
white-spotted sawyer, 203
borers, cone and seed—
Coleotechnites piceaella, 391
spruce coneworm, 355
spruce seed moth, 371
borers, twig and shoot—
white-pine weevil, 209, 423, 426,
AS 8 407
borers, wood—
Chrysobothris pusilla, 167
hemlock borer, 168, 483
northern pine weevil, 211
white-spotted sawyer, 203

Spruce, white—Continued
leaf feeders—

beetle:
white-spotted sawyer, 203
moths:
chain-spotted geometer, 345
Coleotechnites piceaella, 391
Epinotia nanana, 23, 371
Eupethecia filmata, 337
luteata, 337
transcanadata, 337
false hemlock looper, 344
Griselda radicana, 374
hemlock looper, 344, 478, 483
Hyperetis amicaria, 342
Palthis angulalis, 309
red-banded leaf roller, 384
spruce budworm, 25, 378, 472,
A477
spruce coneworm, 355
spruce needleminer, 360
Zeiraphara fortunana, 370
sawflies:
Acantholyda maculiventris,
435
balsam-fir sawfly, 446, 473
Cephalcia fascipennis, 4385
European spruce sawfly, 18,
LOMS2A ASI 4452-54704 471.
473, 477
yellow-headed spruce sawfly,
459, 472

sucking insects—

Adelges lariciatus, 89

balsam twig aphid, 83

Cooley spruce gall aphid, 87, 91
eastern spruce gall aphid, 86, 89
pine spittlebug, 71

Sweetgum—Continued
borers, bark and phloem—Con.

Pityophthorus liquidambarus,
ey PALE

borers, wood—

ambrosia beetles, 17, 21, 120,
124, 264
American plum borer, 359
Columbian timber beetle, 265
Lyctus spp., 127
Platypus compositus, 273
Ptilinus ruficornis, 133
Trichodesma gibbosa, 1338
Tricorynus spp., 184
twig pruner, 192
Xyleborus affinis, 270
ferrugineus, 270
saxeseni, 269

leaf feeders—

bagworm, 21, 411, 472, 475, 477,
478

Coleotechnites dorsovittella, 391

forest tent caterpillar, 19, 330,
AZO NM AUZ 8415

hickory horned devil, 303

imperial moth, 303

luna moth, 298

Oiketicus abbott, 411

Phyllocnistis liquidambarisella,
408

Phytobia posticata, 427

red-humped caterpillar, 316

regal moth, 303

Tetralopha melanogrammos, 358

sucking insects—

Diaspidiotus liquidambaris, 108
periodical cicada, 74
Stictocephala militaris, 71

Saratoga spittlebug, 72, 472 Sycamore:
Tibicen canicularis, 75 borers, bark and phloem—
Sumac: flat-headed apple tree borer, 165,
borers, bark and phloem— 478
borers, wood—

Pityophthorus crinalis, 255
rhois, 255
seriptor, 255
borers, fruit spike—

Anacampsis rhoifructella, 3938

borers, root—
sumac stem borer, 190
borers, wood—
Pityophthorus crinalis, 255
rhois, 255
scriptor, 255
sumac stem borer, 190
leaf feeders—
Arge coccinea, 437
Attelabus nigripes, 223
Episimus argutanus, 360
Nephopteryx subfuscella, 358
regal moth, 3038
Sumac datana, 312
seed feeders—
Moodna ostrinella, 359

Sweetgum:

borers, bark and phloem—
Dryocoetes betulae, 263

Chalcophorella campestris, 168

Columbian timber beetle, 265

flat-headed apple tree borer, 165,
478

hardwood stump borer, 205

living-beech borer, 183

Lichenophanes bicornis, 136

Orchesia castanea, 125

pigeon tremex, 466

powder-post beetle, 17, 21, 126,
123501155

leaf feeders—

Adoxophyes furcatana, 375

American dagger moth, 308

Anchylopera plantanana, 371

bagworm, 21, 411, 472, 475, 477,
478

Chlamisus platani, 145

Datana contracta, 312

Fentonia marthesia, 316

flat-headed apple tree borer, 165,
478

hickory horned devil, 303

io moth, 299

609

Sycamore—Continued

leaf feeders—Continued
Japanese beetle, 19, 151, 485, 486
Japanese weevil, 219
Misogada unicolor, 318
Oiketicus abbotii, 411
Phyllophaga implicta, 148
Plagiometriona clavata, 145
polyphemus moth, 299
puss caterpillar, 349
regal moth, 303
sycamore tussock moth, 306
Tetralopha militella, 353
white-marked tussock moth, 318,

476, 478, 488

root feeders—
Japanese beetle, 19, 151, 485, 486
Japanese weevil, 219

sucking insects—
cottony maple scale, 101
European fruit lecanium, 105,

A475

giant bark aphid, 82
grape mealybug, 100
mulberry whitefly, 92
periodical cicada, 74
sycamore lacebug, 63
terrapin scale, 105

Taxus, see Yew
Thuja, see Arbovritae
Tulip-poplar, see Yellow-poplar
Tuliptree, see Yellow-poplar
Tupelo (gum, black or sour):
borers, bark and phloem—
oak bark scaler, 199
Thamnosphecia rubrofascia, 399
borers, wood—
Anoplodera minnesotana, 197
proxima, 197
Buprestis rufipes, 165
Platypus compositus, 273
leaf feeders—
Actrix nyssaecolella, 359
forest tent caterpillar, 19, 330,
429, 472, 475
gypsy moth, 12, 18, 19, 24, 25,
28.902; 005. lsum ca 3205-430)
AZ ACO Sage AT AN. 475, 410,
483
Phyllophaga forsteri, 148
implicta, 148
luctuosa, 147
Prolimacodes scapha, 348
tupelo leaf miner, 403
root feeders—
Phyllophaga forsteri, 148
implicta, 148
luctuosa, 147
sucking insect—
Phylloxera nyssae, 97

Viburnum:

borers, wood—
Oberea tripunctata, 189

610

Viburnum—Continued
borers, wood—Continued
Synanthedon viburni, 39S
leaf feeders—
Anacampsis rhoifructella, 393
hickory leaf roller, 384

Walnut, black:

borers, bark and/or phloem—

flat-headed apple tree borer, 165,
478

Hypothenemus interstitialis, 254
Pseudothysanoes lecontei, 253

borers, nut—
Conotrachelus retentus, 227
walnut husk fly, 424

borers, twig, shoot, or bud—
Conotrachelus juglandis, 227

retentus, 227

pecan leaf casebearer, 353

borers, wood—
Agrilus otiosus, 163
Ambrosiodmus lecontei, 269
American plum borer, 359
banded hickory borer, 178
Chrysobothris sexsianata, 167
flat-headed apple tree borer, 165,

478

Hylocurus rudis, 252
Hypothenemus interstitialis, 254
leopard moth, 386
Lyctus spp., 127
Micracis suturalis, 2538
painted hickory borer, 178
Petalium bistriatum bistriatum,

133
powder-post beetle, 17, 21, 126,
123 lo

sapwood timberworm, 126
twig pruner, 192
white oak borer, 182
Xyleborus ferrugineus, 270
saxeseni, 269
eylographus, 270
leaf feeders—
beetles:
Attelabus analis, 223
bipustulatus, 223
flat-headed apple tree borer,
165, 478
Japanese beetle, 19, 151, 485,
486
Japanese weevil, 219
Paria quadrinotata, 145
Phyllophaga forsteri, 148
implicta, 148
luctuosa, 147
prunina, 148
Tymnes tricolor, 145
mite:
Eriophyes caulis, 7
moths and butterflies:
Acrobasis caryivorella, 353
Archips rileyanus, 378
cecropia moth, 298, 477
Colocasia propinquilinea, 309
Datana drexeli, 312

Walnut, black—Continued
leaf feeders—Continued
moths and butterflies—Con.
Ectropis crepuscularia, 339
fruit-tree leaf roller, 377
hickory horned devil, 303
hickory tussock moth, 304
luna moth, 298
pecan cigar casebearer, 404
pecan leaf casebearer, 353
polyphemus moth, 299
regal moth, 303
variable oak leaf caterpillar,
315
walnut caterpillar, 312, 472
walnut sphinx, 297
yellow-necked caterpillar, 311
sawfly:
Eriocampa juglandis, 464
root feeders—
Japanese beetle, 19, 151, 485, 486
Japanese weevil, 219
Phyllophaga forsteri, 148
implicta, 148
luctuosa, 147
prunina, 148
sucking insects—
Corythucha juglandis, 64
European fruit lecanium, 105,
475
giant bark aphid, 82
Lecanium caryae, 106
periodical cicada, 74
scurfy scale, 108
tuliptree scale, 103
walnut scale, 107
Willow:
ant, black carpenter, 484, 486
bark feeders—
elm sawfly, 438, 472
poplar borer, 185, 483
borers, bark and phloem—
cottonwood borer, 179
flat-headed apple tree borer, 165,
478
Hypothenemus salicis, 254
Marmara spp., 408
poplar borer, 185, 483
Thamnosphecia sigmoidea, 399
borers, cone—
Conopia proxima, 402
borers, root—
Aegeria tibialis, 398
Conopia proxima, 402
cottonwood borer, 179
hornet moth, 398
Polydrusus impressifrons, 23,

220
Sthenopis thule, 413
borers, wood—
beetles:
Chrysobothris azurea, 167
cottonwood borer, 179
Dicerca tenebrica, 168
flat-headed apple tree borer,
165, 478
hardwood stump borer, 205

Willow—Continued
borers, wood—Continued

beetles—Continued
Hypothenemus salicis, 254
Magdalis salicis, 225
Micracis meridianus, 253
swainei, 253
Oberea ferruginea, 190
Platypus compositus, 273
Poecilonota thrueura, 170
poplar borer, 185, 483
poplar-and-willow borer, 218
Pseudolucanus capreolus, 145
Saperda concolor, 188
imitans, 188
moesta, 188
mutica, 188
Sinodendron rugosum, 145
moths and butterflies:
Aegeria tibialis, 398
carpenter worm, 388
Conopia proxima, 402
hornet moth, 398
leopard moth, 386
Paranthrene dollii castanea,
402
d. dollii, 402

catkin feeder, Xyela spp., 432
gall insects—

Amphibolips fulginosa, 481
Euura spp., 465
Oberea ferruginea, 190
Rhabdophaga sp., 23
Saperda concolor, 188

moesta, 188
willow-beaked gall midge, 420

leaf feeders—

beetles:
Altica suplicata, 142
Asiatic garden beetle, 149, 485
basswood leaf miner, 141
Calligrapha multipunctata
bigsbyana, 140
Chrysomela knabi, 138
cottonwood leaf beetle, 138
elm cailigrapha, 140
European snout beetle, 219
flat-headed apple tree borer,
165, 478
goldsmith beetle, 152
imported willow leaf beetle,
143, 477
Pachybrachys tridens, 145
Phyllophaga crenulata, 148
drakei, 146
forsteri, 148
wmplicta, 148
luctuosa, 147
prunina, 148
tristis, 148
Polydrusus impressifrons, 23,

220
Pyrrhalta decora decora, 140
tuberculata, 140
willow flea weevil, 222
grasshopper, Conocephalus bre-
vipennis, 59

611

Willow—Continued
leaf feeders—Continued

612

mites, Hotetranychus populi, 7

weldoni, 7

moths and butterflies:

Acronicta distans, 308
leporina vulpina, 308
oblinita, 308

Agonopterix argillacea, 395

American dagger moth, 308

Amorbia humerosana, 374

Anacampsis innocuella, 393

Archips purpuranus, 378

Argyrotaenia mariana, 385

bagworms 921-3411, e472, 475,
477, 478

brown-tail moth, 24, 325, 4380,
431, 469, 472, 476

Bruce spanworm, 336

buck moth, 299

cecropia moth, 298, 477

Cerura borealis, 318
cincerea, 318
multiscripta, 318
occidentalis, 318

chain-spotted geometer, 345

cottonwood dagger moth, 308

Deilinea erythemaria, 337

variolaria, 338

Deuteronomos magnarius, 346

Eulype hastata, 337

Evora hemidesma, 360

fruit-tree leaf roller, 377

Gracillaria purpuriella, 409

gypsy moth, 12, 18, 19, 24, 25,
287-625 Come lu el Zone 320,
430, 481, 469, 472, 474, 476,
483

Hemerocampa definita, 319

Hemileuca nevadensis, 300

hickory tussock moth, 304

Hyperaeschra stragula, 313

Hyperetis amicaria, 342

Ichthyura apicalis, 311
brucei, 311
strigosa, 311

io moth, 299

large aspen tortrix, 383

Limenitis arthemis, 294

astyanax, 294

luna moth, 298

Lycia ursaria, 342

mourning-cloak butterfly, 293

Nadata gibbosa, 313

Notodonta simplaria, 318

Nymphalis j-album, 292

oblique-banded leaf roller, 383

Oiketicus abbotii, 411

oriental moth, 347, 349, 431,
484

Orthosia hibisci, 309

Pandemis limitata, 375

Paonias excaecatus, 297

pepper and salt moth, 346

Pero morrisonarius, 344

Pheosia rimosa, 313

polyphemus moth, 299

poplar tent maker, 310

Willow—Continued
leaf feeders—Continued
moths and butterflies—Con.
prairie tent caterpillar, 330
Prochoerodes transversata,
346
Protobarmia porcelaria, 346
Raphia frater, 309
red-humped catervillar, 316
satin moth, 24, 324, 4380, 470,
472, 475, 483
Schizura itpomaeae, 317
leptinoides, 317
Smerinthus jamaicensis, 297
Sparganothis sulfureana, 375
Sphinx luscitiosa, 296
spiny-elm caterpillar, 293
spotted tussock moth, 305
Synchlora aerata, 336
Tetralopha asperatella, 62,
351, 352
unicorn caterpillar, 317
viceroy butterfly, 293
western tent caterpillar, 330
white-marked tussock moth,
318, 476, 478, 483
winter moth, 3386
sawflies:
Arge clavicornis, 437
birch sawfly, 437
elm sawfly, 438, 472
Nematus limbatus, 461
salicisodoratus, 461
Pristiphora sycophanta, 459
Trichiocampus irregularis,
461
viminalis, 23,
461
Trichiosoma triangulum, 488
willow sawfly, 461
willow shoot sawfly, 467
root feeders—
Asiatic garden beetle, 149, 485
European snout beetle, 219
Phyllophaga crenulata, 148
drakei, 146
forsteri, 148
implicta, 148
luctuosa, 147
prunina, 148
tristis, 148
Polydrusus impressifrons, 23,
220
sucking insects—
Brochymena quadripustulata, 62
Chionaspis lintneri, 109
salicisnigrae, 109
Corythucha elegans, 64
mollicula, 63
pallipes, 63
cottony maple scale, 101
giant bark aphid, 82
Idiocerus spp., 69
Lecanium caryae, 106
Macropsis spp., 69
obscure scale, 108
Omalophora salicis, 74
oystershell scale, 109, 110, 475

Willow—Continued
sucking insects—Continued
Pterocomma salicis, 84
smithae, 84
Quadraspidiotus gigas, 107
San Jose scale, 106, 475
Saratoga spittlebug, 72, 472
scurfy scale, 108
Witch-hazel:
gall insects—
Hormaphis hamamelidis, 85
spinosus, 85
leaf feeders—
Archips semiferanus, 378
Datana drexeli, 312
Episimus argutanus, 360
Lithocolletis hamameliella, 407
sucking insects—
Hormaphis hamamelidis, 85
spinosus, 85
Wood products insects:
borers infesting lumber, piling,
poles, woodwork, and manufac-
tured products—
ants:
black carpenter, 484, 486
Camponotus caryae discolor,
490
nearcticus, 489
rasilis, 486
tortuganus, 490
Crematogaster cerasi, 491
lineolata, 492
Florida carpenter, 489
Iridomyrmex pruinosus, 492
red carpenter, 489

bee:
carpenter bees, 497, 498
beetles:
ambrosia beetle, 17, 21, 120,
124, 264

Anobiidae, 31, 126, 131
Anobium punctatum, 22, 132
Asemum spp., 198
panipoe powder-post beetle,
it
banded-ash borer, 184
Bere ned juniper borer,
5
black-horned pine borer, 194
Bostrichidae, 33, 39, 116, 126,
127, 134
brown prionid, 205
brown spider beetle, 137
chestnut timberworm, 126
Chlorophorus annularis, 22,
207
Cossonus spp., 223
Cupes concolor, 118
cypress bark borer, 197
Dermestidae, 115, 156
flat powder-post beetle, 189
hairy pine borer, 198
hardwood stump borer, 205
Hemicoelus carinatus, 132
gibbicollis, 1382
umbrosus, 132

Wood products insects—Continued
borers—Continued
beetles—Continued
Heterobostrychus aequalis, 22,
136
Hexarthrum ulkei, 224
Hylocurus langstoni, 253
ivory-marked beetle, 192
Lichenophanes bicornis, 136
Lyctus powder-post beetle, 17,
21, 126, 128, 129, 130, 155
Nicobium castaneum, 134
oak timberworm, 208
Oeme rigida, 200
old house borer, 206
Orthosoma brunneum, 205
Parandra brunnea, 197
pole borer, 197
Polycaon stoutii, 1386
Prionus pocularis, 200
Pselactus spadix, 224
Ptilinus ruficornis, 1338
Ptinidae, 116, 126, 136
rustic borer, 202
sapwood timberworm, 126
Scobicia bidentata, 1386
Stephanopachys spp., 136
tanbark borer, 197
Tomolips quercicola, 224
Trichodesma gibbosa, 133
Trypopitys sericeus, 133
wharf borer, 125
white-marked spider beetle,
137
Xyletinus peltatus, 1382
Xylobiops basilaris, 134, 502
termites:
eastern subterranean, 49
Formosan subterranean, 50
Incisitermes snyderi, 46
Kalotermes approximatus, 45,
53
Neotermes castaneus, 54
jouteli, 46, 53
Prorhinotermes simplex, 46,
54
southern dry-wood, 53
tropical rough-headed powder-
post, 54
(several more spp. also)
marine borers:
Bankia, 10, 11
Chelura, 3
Limnoria lignorum, 3
Martesia, 11
shipworms, 2, 10
Sphaeroma spp., 3
Teredo, 10
wood lice, 3

Yellow cucumber tree:
Archips magnolianus, 378
Yellow-poplar:
borers, bark and phloem—
Euzophera ostricolorella, 359
flat-headed apple tree borer, 165,
478
oak-bark scaler, 199

613

Yellow-poplar—Continued
borers, wood—

Ambrosiodmus tachygraphus,
269

Buprestis rufipes, 165

Chalcophorella campestris, 168

Columbian timber beetle, 265

flat-headed apple tree borer, 165,
478

Orchesia castanea, 125

pear blight beetle, 269

sapwood timberworm, 126

Xyleborus saxeseni, 269

leaf feeders—

614

beetles:
flat-headed apple tree borer,
165, 478
Odontopus calceatus, 221
mite, Tetranychus magnoliae, 7
moths:
Epimecis virginiara, 339
Paralobesia liriodendrana, 378
Phyllocnistis liriodendrella,
408

Yellow-poplar—Continued
leaf feeders—Continued

moths—Continued
polyphemus moth, 299
promethea moth, 298

sucking insects—

Abgrallaspis townsendi, 108

Chionaspis salicisnigrae, 109

European fruit lecanium, 105,
A75

Macrosiphum liriodendri, 84

periodical cicada, 74

tuliptree scale, 103

Yew:
leaf and root feeders—

Asiatic garden beetle, 149, 485
black vine weevil, 220
strawberry root weevil, 221

sucking insects—

Comstock mealybug, 100
cottony maple leaf scale, 101
Dysmicoccus cuspidatae, 99
Fiorinia externa, 111
Fletcher scale, 106

grape mealybug, 100

:
|

INSECT INDEX

This is an alphabetical listing of the insect species, both scien-
tific and common names. Bold face numbers indicate page where

the insect is discussed.

Abbottana clemataria, 346
Abgrallaspis—
ithacae, 108
townsendi, 108
Acanthocinus—
nodosus, 196
obsoletus, 196
pusillus, 196
Acantholyda—
angulata, 435
apicalis, 435
circumcincta, 435
erythrocephala, 23, 433
luteomaculata, 435
maculiventris, 435
pini, 435
zappei, 434
Acarina, 6, 35
Aceria phloceoptes, 8
Acholla multispinosa, 65
Acleris—
chalybeana, 385
logiana, 386
tripunctana, 386
varianda, 385
Acmaeodera pulchella, 160
Acmopolynema bifasciatipennis bi-
fasciatipennis, 472
Acordulecera, 436
Acossus—
centernesis, 388
populi, 388
Acrididae, 32, 33, 57
Acrobasis—
betulella, 354
caryae, 354
caryivorella, 353
demotella, 354
feltella, 354
indiginella, 354
juglandis, 353
rubrifasciella, 354
septentrionella, 354
Acronicta—
americana, 308
distans, 308
funeralis, 308
innotata, 308
interrupta, 308

Acronicta—Continued
leporina vulpina, 308
lepusculina, 308
lithospila, 308
modica, 308
morula, 308
oblinita, 308
retardata, 308

Actenodes acornis, 169

Actias luna, 298

Actrix nyssaecolella, 359

Adalia bipunctata, 102, 122

Adelges—
abietis, 22, 86
cooleyi, 87
lariciatus, 89
piceae, 22, 88
strobilobius, 22, 89

Adoxophyes furcatana, 375

Aedes, 416

Aegeria—
apiformis, 23, 398
tibialis, 398

Aegeriidae, 33, 34, 35, 38, 397

Aenea ostryaealla, 390

Agathidium oniscoides, 119

Agathis—
pini, 366
pumilus, 367, 405, 469, 473

A gonopterix—
argillacea, 395
nigrinotella, 395
pteleae, 395
robiniella, 395

Agonum spp., 117

Agrillaxia flavimana, 170

Agrilus—120
acutipennis, 163
anxius, 161
arcuatus torquatus, 163
betulae, 163
bilineatus, 160
celti, 163
cephalicus, 163
difficilis, 163
egenus, 163
fuscipennis, 163
horni, 162
juglandis, 163

615

Agrilus—Continued

lecontei, 163

liragus. 162

otivsus, 163
Agromyzidae, 33, 35, 36, 38, 426
Airora cylindrica, 120
Alaus—

myops, 153

oculatus, 153
Alebra, 69
Aleurochiton forbesu, 92
Aleyrodidae, 36, 92
Alleculidae, 116
Allotropa—

ashmeadi, 484

burrelli, 484

convexifrons, 100, 484
Alsophila pometaria, 334
Altica—

ambiens subspecies alni, 141

carinata, 142

subplicata, 142
Amblycerus robiniae, 137
Ambrosiodmus—

lecontei, 269

linderae, 269

tachygraphus, 269
Amorbia humerosana, 374
Amphibolips—

confluenta, 481

fulginosa, 481
Amphicerus bicaudatus, 136
Amphidasis cognataria

(see Biston cognataria)
Amphimallon majalis, 22, 150
Ampulicidae, 495
Anabrus simplex, 59
Anacampsis—

mnocuella, 393

rhoifructella, 393
Anacamptodes—

ephyraria, 338

larvaria, 338

pergacilis, 338
Anastatus disparis, 323, 475
Anatis 15-punctata, 123
Anchylopera plantanana, 371
Andricus cornigerus, 398
Aneflormorpha subpubescens, 194
Aneflus protensus, 199
Anisandrus—

minor, 234

obesus, 234

pyri, 269
Anisomorpha—

buprestoides, 57

ferruginea, 57
Anisota—

rubicunda, 301

senatoria, 300

stigma, 300

virginiensis, 301
Anobiidae, 31, 126, 131
Anobium punctatum, 22, 132
Anomala—

luciola, 151

oblivia, 150

orientalis, 22

616

Anomoea laticlavia, 145
Anomogyna eliminata, 309
Anopheles, 416
Anoplodera—
biforis, 197
canadensis, 196
minnesotana, 197
mutabilis, 197
nitens, 196
proxima, 197
vagans, 197
vittata, 197
Anoplonyx—
canadensis, 465
luteipes, 465
Anoplura, 40
Anormenis septentrionalis, 68
Antheraea polyphemus, 299
Anthocoridae, 65
Anthocoris—
borealis,
(see A. musculus)
musculus, 65
Anthomyiidae, 427
Anthophila pariana, 23, 396
Anthrax sinuosa, 444
Anthribidae, 116, 209
Anthrophorinae, 497
Antispila nyssaefoliella, 403
Ant lions, 43
Ants—
Allegheny mound, 491
Argentine, 492
black carpenter, 484, 486
Florida carpenter, 489
imported fire, 492
red carpenter, 489
Texas leaf-cutting, 490
velvet, 485
Anurogryllus muticus, 60
Apanteles—
congregatus, 296
lacteicolor, 469
melanoscelus, 323, 469
solitarius, 324, 470
Apantesis radians, 307
A patelodes—
angelica, 334
torrefacta, 333
Aphelia alleniana, 386
Aphelinus proelia, 107
Aphelopus theliae, 496
Aphidecta obliterata, 123
Aphididae, 31, 35, 36, 37, 77, 86
Aphidoletes thompsoni, 421
Aphids—19, 438, 64. 67, 77, 121, 424,
472
balsam twig’, 83
balsam woolly, 12, 24, 88, 90, 1238,
124
beech blight, 84
black pecan, 84
boxelder, 83
Cooley spruce gall, 87, 91
crapemyrtle, 84
eastern spruce gall, 86, 89
elm coxecomb-gall, 85
elm leaf, 84

Aphids—Continued
giant bark, 82
hickory gall, 92
Norway-maple, 84
pine bark, 89
poplar vagabond, 85
spruce gall, 43
white-pine, 83
woolly alder, 85
woolly apple, 84
woolly elm, 84
woolly elm bark, 84

Aphis craccivora, 84

Aphrophora—
parallela, 71
saratogensis, 72

Aphrophorinae, 71

Apidae, 39, 497

Apinae, 497

Apocrita, 432, 468

Apoidea, 497

apple bucculatrix, 409

Aptesis basizonia, 470

Arachnida, class 2, 5

Araneida, 5

Archips—
argyrospilus, 377
cerasivoranus, 376
fervidanus, 377
georgianus, 378
griseus, 378
infumatanus, 378
magnolianus, 378
negundanus, 378
purpuranus, 378
rileyanus, 378
rosanus, 23, 376
semiferanus, 378

Archodontes melanopus, 206

Arctiidae, 304

Arge—
clavicornis, 437
coccinea, 437
pectoralis, 437
scapularis, 437

Argidae, 437

Argypon flaveolatum, 337

Argyresthia-——
auveoargentella, 403
conjugella, 403
freyella, 402
laricella, 403
thuiella, 403

Argyrotaenia—
alisellana, 385
juglandana, 384
mariana, 385
occultana, 385
pinatubana, 384, 387
quadrifasciana, 385
quercifoliana, 384
velutinana, 384

Arhopalus rusticus obsoletus, 198

Arilus cristatus, 64
Arrhenodes minutus, 208
Arthropoda, 2

Asemum striatum, 198
Asilidae, 422

Asilus sericeus, 422
Aspen blotch miner, 407
Asphondylia—

azalea, 418

ilicicola, 418
Aspidiotus—

abietis, 112

ancylus,

(see Diaspidiotus ancylus)

hederae, 112

ithacae,

(see Absgrallaspis ithacae)
juglans-regiae,

(see Quadraspidiotus juglans-

regiae)
perniciosus,

(see Quadraspidiotus pernicious )

Asterocampa—
celtis, 294
clyton, 294
Asterolecaniidae, 35, 100
Asterolecaniinae, 101
Asterolecanium—
arabidis, 101
bambusae, 101
luteolum, 101
miliaris miliaris, 101
miliaris robustum, 101
minus, 101
pustulans, 101
puteanum, 101
quercicola, 101
variolosum, 22, 101
Atimia confusa, 198
Atta texana, 490
Attelabus—
analis, 223
bipustulatus, 223
nigripes, 223
rhois, 223
Atteva aurea, 403
Aulacidae, 483
Aulacus—
burquei, 483
digitalis, 483
lovet, 483
pallipes, 483
Automeris 10, 299

Bagworm, 21, 411, 472, 475, 477, 478

Baliosus—

nervosus,

(see B. ruber.)

ruber, 141, 142
Bankia, 10, 11
Bapta—

semiclarata, 337

vestaliata, 337
Basicladus celibatus, 411
Basilarchia archippus,

(see Limenitis archippus)
Bassareus lituratus, 145
Bassus pumilus,

(see Agathis pumilus)
Battaristis vittella, 394
Bees—436, 483, 484, 485, 497

bumble, 497
carpenter, 497, 498

Bees—Continued
cuckoo, 497
digger, 497
honey, 497
leaf-cutting, 486, 497
mining, 497
Beetles—113
alder flea, 141
ambrosia, 17, 21, 120, 124, 264
Asiatic garden, 149, 485
bamboo powder-post, 135
bark—17, 20, 21, 29, 118, 119, 120,
227, 450, 502
birch, 263
eastern ash, 249
eastern juniper, 248
flat, 124
four-eyed spruce, 264
hickory, 237, 270
native elm, 250, 499
oak, 255
peach, 240
smaller European elm, 10, 19,
24, 29, 154, 235, 475, 499
belid, 209
black turpentine, 244
blister—120
ash-gray, 120
striped, 120
brentid, 208
carrion, 119
checkered, 154
click, 153
Columbian timber, 265
darkling, 125
death-watch, 131, 132
dermestid, 156
drug store, 131
eastern death-watch, 132
eastern hercules, 152
eastern larch, 248
eastern spruce, 247
engraver, 502
European snout, 219
eyed click, 153
false powder-post, 134
flat powder-post, 189
furniture, 132
giant stag, 145
goldsmith, 152
green June, 152
ground, 117
hide, 156
hister, 119
ivory-marked, 192
Japanese, 19, 151, 485, 486
lady—121
convergent, 123
twice-stabbed, 105, 110, 111, 123
two-spotted, 122
larder, 156
leaf—137
cottonwood, 138
elm, 138, 431, 474
European elm, 431
imported willow, 143, 477
larger elm, 141
long-horned, 170

‘618

Beetles—Continued
lyctus—
old world, 130
southern, 129
western, 130
May, 483
ostomatid, 120
passalid, 119
pear blight, 269
pitted ambrosia, 266
powder-post, 17, 21, 126, 128, 155
red-pine cone, 257
red turpentine, 246
rhinoceros, 152
sap, 121
seed, 137
snout, 208
softwood powder-post, 132
southern cypress, 248
southern pine, 240, 477, 502
spider—136
brown, 137
white-marked, 137
spruce timber, 268
stag, 145
striped ambrosia, 267
tiger, 118
timber, 125
tumbling flower, 126
western pine, 28
white-fringed, 224
white-pine cone, 256
Belidae, 209
Bessa harveyi, 458
Bethylidae, 484
Bethyloidea, 484
Bibarrambla allenella, 395
Bibio albipennis albipennis, 417
Bibionidae, 417
Biorhiza forticornis,
(see Xanthoteras forticone)
birch tube maker, 354
Biston cognataria, 346
Bitoma carinata, 124
black-bellied clerid, 155
Blastobasidae, 31, 395
Blattidae, 61
Blennocampa caryae,
(see Eriocampa juglandis)
Blepharipa scutellata, 323, 430, 478,
483

Bombyliidae, 423
Borers—21,
alder, 188
American plum, 359
apple twig, 136
ash, 400
ash and privet, 199
Australian-pine, 166
banded-ash, 184
banded-hickory, 178
black-horned juniper, 195
black-horned pine, 194
Bostrichidae, 126, 134
boxelder twig, 369
broad-necked root, 200
bronze birch, 29, 161, 478
bronze poplar, 162

Borers—Continued

buprestid, 475, 496

cedar tree, 198

cerambycid, 475

chestnut bark, 196

cottonwood, 179

cypress bark, 197

dogwood, 398

dogwood twig, 189

eastern larch, 199

elder, 193

elm, 188

elm bark, 197

fig tree, 199

flatheaded, 156

flat-headed apple tree, 165, 478

flatheaded bald cypress sapwood,
160

flatheaded fir, 169

flatheaded sycamore
167

gall-making maple, 201

hairy pine, 198

hard maple bud, 413

hardwood stump, 205

hemlock, 168, 483

hickory spiral, 163

jack-pine shoot, 368

Large flatheaded pine heartwood,
167

lesser cornstalk, 358

lilac, 400

linden, 187

living-beech, 183

living hickory, 183

locust, 17, 29, 64, 176

locust twig, 373

maple callus, 399

maple petiole, 463

mesquite, 178

mulberry, 193

mulberry bark, 193

oak branch, 183

oak sapling, 183

cak stem, 194

old house, 206

painted hickory, 178

pecan, 398

persimmon, 400

pin-hole, 264

pine bark, 196

pitch mass, 399

pole, 197

poplar, 185, 483

poplar-and-willow, 218

poplar-butt, 202

red-headed ash, 184

red oak, 180

rhododendron, 398

rhododendron stem, 190

ribbed pine, 206

roundheaded, 170

roundheaded apple tree, 187

rustic, 202

shot-hole, 238

small cedar bark, 198

spined bark, 192

sugar maple, 180

heartwood,

Borers—Continued
sumac stem, 190
tanbark, 197
thorn-limb, 188
turpentine, 164
two-lined chestnut, 160
wharf, 125
white oak, 182
white-pine shoot, 367
wood, 17, 21, 450

Bostrichidae, 33, 39; 116, 126,

134
Brachymeria—
compsilurae, 478
ovata, 478
Brachyrhinus—
ovatus, 221
sulcatus, 220
Brachys—
aeruginosa, 169
ovata, 169
tesselatus, 169
Braconidae, 468, 469
branconids, 469, 476
Brentidae, 34, 38, 116
Brephos infans, 334
bristletails, 41
Brochymena—
carolinensis, 62
quadripustulata, 62
Brown prionid, 205
Bruchidae, 31, 117, 137
Bucculatrix—
ainsliella, 410
canadensisella, 409
coronatella, 410
luteella, 410
packardella, 410
pomifoliella, 409
quinquenotella, 410
recognita, 410
Budworms—
black-headed, 385, 472
jack pine, 381
spruce, 25, 378, 472, 477
Bugs—
assassin, 64
boxelder, 66
coreid, 66
damsel, 65
doodle, 43
flower, 65
lace—62, 65
azalea, 64
basswood, 64
elm, 63
hackberry, 63
hawthorn, 64
oak, 63
rhododendron, 64
Sycamore, 63
lygaeid, 67
mealy—438, 67, 99, 424, 484
Comstock, 100
grape, 100
long-tailed, 100
striped, 100
palmetto pill, 223

127,

619

Bugs—Continued
plant, 65
royal palm, 67
spined soldier, 62
spittle—
alder, 73
dogwood, 74
meadow, 74
pecan, 71
pine, 71
Saratoga, 72, 472
stink, 62
tarnished plant, 65
true, 61
wheel, 64
Buprestidae, 33, 34, 35, 36, 38, 39,
115, 156, 467
Buprestis—
apricans, 164
lineata, 165
maculipennis, 165
rufipes, 165
salisburyensis, 165
striata, 165
Butterflies—
brush-footed, 292
comma, 292
monarch, 293
mourning-cloak, 293
swallowtail, 291
Byrrhidae, 114, 119
Byrrhus, 119

Calaphis—

betulaecolens, 84

betulella, 84
Calcaritermes nearcticus, 47, 54
California five-spined ips, 28
Caliroa—

cerasi, 23, 457

lineata, 457

(near) quercuscoccineae, 457
Callidium—

antennatum hesperum, 194

schotti, 195

texanum, 195

violaceum, 195
Calligrapha—

multipunctata bigsbyana, 140

scalaris, 140
Calliphoridae, 428
Callirhy tis—

cornigera, 480

floridana, 479

futilis, 480

gemmaria, 480

operator, 480

palustris

(see Dryocosmus quercuspalus-
tris )

punctata, 479
Callisto geminatella, 408
Callosamia promethea, 298
Calocalpe undulata, 337
Calosoma—

calidum, 117. 118

frigidum, 117

620

Calosoma—Continued
scrutator, 117, 344
sycophanta, 117, 118, 323
willeoxt, 117

Cameraria—
cincinnatiella,

(see Lithocolletis cincinnatiella)
hamadryadella,
(see Lithocolletis hamadrya-

Camnula pellucida, 58

Campaea perlata, 346

Camponotus—
abdominalis floridanus, 489
caryae discolor, 490
castaneus, 490
ferrugineus, 489
Mississippiensis, 490
nearcticus, 489
pennsylvanicus, 486
pylartes fraxinicola, 490
rasilis, 486
tortuganus, 490

Campoplex frustranae, 365

Campsomeris annulata, 489

Canarsia ulmiarrosorella, 359

Cankerworms—128,
fall, 21, 334, 389, 472, 4838
spring, 339

Carabidae, 114, 117

Carabus auratus, 323

Carcelia—
lagoa, 350
laxifrons, 430

Carpenterworms—
aspen, 388
little, 389
pecan, 387

Carphoborus bifurcus, 239

Carulaspis carueli, 111

Carynota stupida, 71

Carybruchus gleditsiae, 137

Caryomyia—
holotricha, 420
sanguinolenta, 420
tubicola, 420

Casebearers—
birch, 405
cigar, 406
elm, 404
larch, 24, 405, 469, 473, 478
pecan cigar, 404
pecan leaf, 353
pecan nut, 354

Caterpillars—
eastern tent, 328, 476, 477, 483
forest tent, 19, 330, 429, 472, 475
maple seed, 369
prairie tent, 330
puss, 349
red-humped, 316
saddleback, 347
Sonoran tent, 330
Sspiny-elm, 293
tent, 62, 327
ugly-nest, 376
unicorn, 317

Caterpillars—Continued
variable oak leaf, 315
walnut, 312, 472
western tent, 330
yellow-necked, 311
zebra, 309

Catocala, 308

Catogenus rufus, 124

Caulocampus acericaulis, 463

Cecidomyia—
catalpae, 419
cerasiserotinae, 419
cupressi, 419
foliora, 419
ocellaris, 419
pellex, 419
piniinopis, 418
poculum, 419
reeksi, 419
resinicola, 419
verrucicola, 419

cecidomyids, 476

Cecidomyiidae, 30, 33, 36, 417, 484

centipedes, 4
Centuroides vittatus, 9
Cephalcia—
fascipennis, 435
fulviceps, 435
marginata, 435
Cephaloidae, 116
Cephenomyia—
phobifer, 431
pratti, 431
Cephidae, 467
Cephoidea, 467

Cerambycidae, 31, 33, 34, 35, 36, 38,

39, 115, 116, 170, 484
Ceramica picta, 309
Ceraphronidae, 483
Ceratomia—

amyntor, 295

catalpae, 295

undulosa, 295
Ceratopogonidae, 416
Cerceris fumipennis, 496
Cercopidae, 34, 36, 71
cercopids, 496
Ceresa bubalus,

(see Stictocephala bubalus)
Ceruchus piceus, 145
Cerura—

borealis, 318

cinerea, 318

multiscripta, 318

occidentalis, 318
Chaetexorista javana, 348, 431
Chafers,

European, 150

pine, 150

rose, 150
Chaitophorus—

populicola, 84

steveusis, 84
Chalastogastra,

(see Symphata)
Chalcididae, 31, 36, 478
Chalcidoidea, 471

Chalcophora—
georgiana, 168
liberta, 167
virginiensis, 167, 168

Chalcophorella campestria, 168

Chamaemyiidae, 425

changa, 60

Charadra deridens, 309

Chariessa pilosa, 154, 155

Chelifer cancroides, 10

Chelura terebrans, 3

chermid, pine leaf, 91

Chermidae, 35, 36

Chermidae, 36

chigger, 8

Chilocorus—
bivulnerus, 102, 105
stigma, 123

Chilopoda, 2, 4

Chionaspis—
americana, 108
corni, 109
furfura, 108
lintneri, 109
salicisnigrae, 109

Chironomidae, 416

Chlaenius erythropus, 117

Chlamisus platani, 145

Chlorophorus annularis, 22, 207

Chloropidae, 425

Choristoneura—
biennis, 381
conflictana, 382, 383
fractivittana, 384
fumiferana, 378
houstonana, 384
occidentalis, 381
pinus, 381
pinus maritima, 383
rosaceand, 383, 384

Chramesus—
chapuisti, 239
hicoriae, 239

Chrysididae, 484

Chrysidoidea, 484

Chrysis shanghaiensis, 484

Chrysobothris—
adelpha, 167
azurea, 167
blanchardi, 167
dentipes, 167
femorata, 165
floricola, 165
harrisi, 167
neopusilla, 167
orono, 166
pusilla, 167
scabripennis, 167
sexsignata, 167
texana, 167
tranquebarica, 166
trinervia, 167
veridiceps, 167

Chrysocharis laricinellae,

(see Kratochviliana laricinellae)

Chrysoclista linneella, 23, 390

621

Chrysomela—

crotchi, 22, 123

interrupta, 22, 138

knabi, 138

scripta, 138
Chrysomelidae, 32, 117, 137, 477
Chrysomphalus—

dictyospermi, 112

obscurus,

(see Melanaspis obscura)

tenebricosus,

(see Melanaspis tenebricosa)
Chrysopa oculata, 43
Chrysopidae, 43
Crysops, 422
cicada killer, 496
Cicadas—

dog-day, 74

periodical, 74
Cicadellidae, 36, 68
cicadellids, 496
Cicadidae, 34, 36, 74
Cicindela, 118
Cicindelidae, 114, 118
Cimberis, 223
Cimbex americana, 438
Cimbicidae, 437
Cinara—

abietis, 83

atlantica, 83

canadensis, 83

carolina, 83

curvipes, 80

juniperivora, 83

pini, 83

sabinae, 83

strobi, 83

taedae, 83

tujafilina, 83

watsoni, 83
Cincinnus melsheimeri, 347
Cincticornia pilulae, 420
Cingilia catenaria, 345
Cinyra gracilipes, 170
Cisidae, 115
Citheronia—

regalis, 303

sepulchralis, 303
Citheroniidae, 300
Clastoptera—

achatina, 74

obtusa, 74

proteus, 74

undulata, 73
Cleis picta, 123
Cleptes, 484
Cleptidae, 484
Cleridae, 115, 154
Clistogastra,

(see Apocrita)
Clostocerus tricinctus, 1438
Closterocerus cinctipennis, 441
Cnidocampa flavescens, 23, 347
Coccidae, 33, 36, 37, 101
Coccinella—

novemnotata, 123
transversoguttata, 123

622

Coccinellidae, 115, 121
Coccoidea, 92
Coccophagus insidiator, 473
Cochliomyia—
hominivorax, 428
macellaria, 429
cockroaches, 54, 61, 495
Coeloides scolytivorus, 249
Coelostethus notatus, 133
Colaspis pini, 142
Coleophora—
alniella, 404
atromarginata, 404
caryaefoliella, 404
fletcherella,

(see C. serratella)
fuscedinella, 23, 405
laricella, 23, 404
limosipennella, 23, 404
querciella, 404
salmanii,

(see C. fuscedinella)
serratella, 406
tilaefoliella, 404
ulmifoliella, 404

Coleophoridae, 404

Coleoptera, 18, 19, 22, 28, 31, 32, 33,
34, 35, 36, 38, 39, 113, 423, 469,

471, 475, 477, 478, 484
Coleotechnites—

apicitripunctella, 390

dorsovittella, 391

juniperella, 391

piceaella, 391

thujaella, 391

variella, 391
Collembola, 41
Colocasia—

flavicornis, 309

propinquilinea, 309
Colopha—

ulmicola, 85

ulmisacculi, 85
Colydiidae, 114, 116, 124
Colydium lineola, 124
Compsilura concinnata, 311,

324, 330, 430, 478
coneworm, spruce, 355
Conocephalus brevipennis, 59
Conophthorus—

coniperda, 256

resinosae, 257

taedae, 258

virginianae, 258
Conopia—

acernt,

(see Sylvora acerni)

acerrubri, 402

bolteri, 402

proxima, 402
Conopidae, 424
Conostigmus virginicus, 483
Conotrachelus—

affinis, 227

anaglypticus, 227

aratus, 227

carinifer, 227

Conotrachelus—Continued
caroliniensis, 433
crataegi, 227
hicoriae, 227
juglandis, 227
NaSO, 227
nenuphar, 227
posticatus, 227
retentus, 227

Contarinia—
canadensis, 418
juniperina, 418
negundifolia, 418
virginianiae, 418

Coptodisca splendoriferella, 404

Coptotermes formosanus, 22, 45, 46,
50

Coreidae, 66

Corthylus—
columbianus, 265
punctatissimus, 266

Corticotomus cylindricus, 120

| Corydalidae, 42

Corydalus cornutus, 42

Corythucha—
aesculi, 64
arcuata, 63
associata, 64
bellula, 64
betulae,

(see C. pallipes)
canadensis,

(see C. mollicula)
celtidis, 63
ciliata, 63
cydoniae, 64
cyrta,

(see C. pallipes)
elegans, 64
juglandis, 64
mollicula, 63
pallipes, 63
pergandei, 64
pruni, 64
pyriformis,

(see C. pruni)
salicis,

(see C. mollicula)
ulmi, 63

Cosmoptergidae, 390

Cossidae, 33, 38, 386

Cossonids, 120, 124

Cossoninae 223

Cossonus—
corticola, 223
COSSONUS, 223
impressifrons, 223
impressus, 223
platalea, 223

Cossula magnifica, 387

Cotalpa lanigera, 152

Cotinis nitida, 152

Cregya oculata, 156

Crematogaster—

| ashmeadi, 492

cerasi, 491

clara, 492

Crematogaster—Continued
laeviuscula, 492
lineolata, 492

Cremifania nigrocellulata, 425

Cressonia juglandis, 297

crickets—59, 424
mole—60

European, 60

southern, 61

Mormon, 59
short-tailed, 60
snow tree, 60
tree, 472, 475

Croesia—
albicomana, 385
semipurpurana,

(see C. albicomana)
Croesus latitarsus, 462
Crossocerus ambiguus, 496
Crustacea, Class 2, 3
Cryphalus,

(see Taenioglyptes)
Cryptococcus fagi, 22, 97, 411
Cryptorhynchus lapathi, 22, 218
Cryptotermes—

brevis, 45, 47, 54
cavifrons, 47, 54

Cryptothelea gloverei, 413

Cryptulocleptus dislocatus, 253

Crypturgus—
alutaceus, 239
atomus, 239
borealis, 239

Crytepistomus castaneus, 22, 221

Ctenicera—
nitidula, 154
triundulata, 154

Ctenocepalides—
canis, 41
felis, 41

Ctenuchidae, 304

Cucujidae, 114, 115, 124

Cucujus clavipes, 124

Cuerna costalis, 70

Culex, 416

Culicidae, 415

Cupes concolor, 118

Cupesidae, 114, 118

Curculio—
auriger,

(see C. say)

caryae, 226
caryatrypes, 226
humeralis, 226
longidens, 226
neocarylus, 226
orthorhynchus, 226
pardalis, 226
proboscideus,

(see C. caryatrypes)

sayi, 226
sulcatulus, 226

Curculionidae, 31, 32, 33, 34, 35, 36,
38, 117, 209

cutworms, 309

Cymatodera bicolor, 155

Cynipidae, 31, 478, 479

623

cynipids, 476
Cynipoidea, 36, 478
Cyrtolobus discoidalis, 71
Cyzenis albicans, 337

daddy-long-legs, 10
Dahlbominus fuscipennis, 441, 442,

446, 448, 473
Dasineura—

balsamicola, 418

communis, 418

gleditchiae, 418

pseudacaciae, 418
Dasychira—

atomaria, 320

basiflava, 320

cinnamomea, 320

meridionalis, 320

plagiata, 320

tephra, 320

vagans, 320
Dasymutilla occidentalis occidentalis,

485

Datana—

angusi, 311

contracta, 312

drexeli, 312

integerrima, 312

major, 313

ministra, 311

perspicua, 312
Deilinea—

erythemaria, 337

variolaria, 338
Dendrobiella—

aspera, 136

sericans, 136
Dendroctonus—

frontalis, 240

obesus, 247

piceaperda,

(see D. obesus)

simplex, 248

terebrans, 244

valens, 246
Dendrotettix—

australis, 58

quercus, 58
Depressaria betulella, 395
Dermacentor variabilis, 9
Dermestes—

lardarius, 156

maculatus, 156
Dermestidae, 115, 156
Derocrepis aesculi, 145
Derodontidae, 124
Desmia funeralis, 350
Desmocerus palliatus, 193
Deuteronomos magnarius, 346
Diachlorus ferrugatus, 422
Dialeurodes—

chittendeni, 92

citri, 92
Diapheromera—

blatchleyi, 57

femorata, 55, 57

velti, 57

624

Diapriidae, 483
Diaspididae, 106
Diaspidiotus—
aesculi, 108
ancylus, 108
liquidambaris, 108
Diaspis carueli,

(see Carulaspis carueli)
Dibrachys cavus, 477
Dicaelus purpuratus, 117
Dicentria lignicolor, 316
Dicerca—

divaricata, 168

lurida, 168

obscura, 168

punctulata, 168

tenebrica, 168

tenebrosa, 168
Dichelonyx—

albicollis, 149

elongata, 149

subvittata, 149
Dichomeris—

ligulellus, 393

marginelius, 23, 393, 394
Dimmockia incongrus, 474
Dinoderus minutus, 135
Diodyrhynchus, 223
Dioryctria—

abietella, 23, 355

abietivorella,

(see D. abietella)

amatella, 356
cambticola, 358
clarioralis, 354
disclusa, 358, 475
pygmaeela, 358
reniculella, 355
zimmermani, 354, 356, 357
Diplopoda, 2, 4
Diplotaxis—
liberata, 149
sordida, 149
Diprion—
frutetorum, 23, 451, 473
hercyniae, 23, 452, 473
similis, 23, 450
Diprionidae, 439
Diptera, 18, 19, 23, 30, 31, 33, 35, 36,
838, 415, 471, 475; 476; 4772-478;
483, 484
Dircacea quadrimaculata, 125
Dolichopodidae, 423
Dolichopus vittatus, 423
Doraschema—
alternatum, 194
wildii, 193
Dorcus parallelus, 145
Drepanaphis—
acerifoliae, 84
carolinensis, 84
nigricans, 84
sabrinae, 84
Drepanosiphum platanoides, 84
Drino bohemica, 431
Drosophilidae, 425
Dryinidae, 496

Dryocoetes—

affaber, 263

autographus, 263

betulae, 263

caryl, 264

granicollis, 264

piceae, 264
Dryocosmus quercuspalustris, 482
Dryophthorus americanus, 223
Dynastes tityus, 152

cuspidatae, 99

obesus, 99

Eacles—

imperialis, 303

imperialis pini, 304
Eburia quadrigeminata, 192
Ecdytolopha insiticiana, 373
Ectodemia—

heinrichi, 413

populella, 413
Ectropis crepuscularia, 339
Elachertus cocoeciae, 474
Elaphidion—

incertum, 193

mucronatum, 192
Elaphidionoides villosus, 192
Elasmidae, 474
Elasmopalpus lignosellus, 358
Elasmuche lateralis, 62
Elateridae, 31, 114, 153
Elatophilus inimica, 65
elder leaftier, 351
elm calligrapha, 140
Empididae, 423
Empoasca, 69
Enaphalodes—

antomerius, 182

cortiphagus, 181

rufulus, 180
Enchenopa binotata, 71
Encyclops coerulea, 199
Encyrtidae, 474
engraver—

hackberry, 239

pine, 262

southern pine, 260
Ennomos subsignarius, 343
Enoclerus—

lecontei, 155

nigripes, 155

quadriguttatus, 249
Eotetranychus—

hicoriae, 7

matthyssei, 7

populi, 7

querci, 7

weldoni, 7
Epargyreus clarus, 294
Evhemeroptera, 44
E'picauta—

cinerea, 121

fabricii, 120

pestifera, 120

subglabra, 121

torsa, 121

vittata, 120

Epicnaptera americana, 333
Epilampsis laricinellae, 405
Epimecis virginiaria, 339
Epinotia—

aceriella, 371

lindana, 374

nanana, 23, 371

similana, 374

solandriana, 371
Epipaschidae, 351
Episimus argutanus, 360
Epizeuxis aemula, 309
Erannis tiliaria, 25, 341
Erinna—

abdominalis, 421

lugen, 421
Eriocampa juglandis, 464
Eriococcidae, 97
Eriocraniidae, 414
Eriophyes—

caulis, 7

fraxiniflora, 7

parapopuli, 7

pyri, 7
Eriophyidae, 7
Eriosoma—

americanum, 84

crataegi, 84

lanigerum, 84

rileyi, 84
Ernobius—

granulatus, 133

mollis, 133

tenuicornis, 133
Erynniopsis rondani, 431
Erythroneura, 69
Essigella pini, 83
Euceraphis—

betalae, 84

lineata, 84

mucidas, 84
Eucharitidae, 475
Eucosma—

cocanda, 368

gloriola, 367

monitorana, 368

sonomana, 368

tocullionana, 368
Eucrada humeralis, 133
Eufidonia notataria, 338
Eugnamptus, 223
Eulachnus—

agilis, 83

rileyi, 83
Eulimneria rufifemur, 362
Eulophidae, 472
Eulype hastata, 337
Eupelmella vesicularis, 475
Eupelmidae, 475
Eupelmus—

allynvi, 475

cyaniceps amicus, 475

cyaniceps cyaniceps, 475

pini, 475
Eupethecia—

filmata, 337

luteata, 337

625

Eupethecia—Continued Flies—Continued

palpatata, 337 bot, 431
transcanadata, 337 cherry fruit, 424, 483
Eupteromalus nidulans, 325 chloropid, 425
Eurytetranychus buxi, 7 damsel, 45
Eurytomidae, 477 dance, 423
Eurytoma— deer, 421
magdalidis, 477 dobson, 42
pint, 477 dragon, 45
pissodis, 477 flesh, 429
verticillata, 477 flower, 423
tylodermatis, 477 fruit, 424
Eustrophus tomentosus, 125 greenbottle, 429
Eutetrapha tridentata, 188 harvest, 74
Eutettix, 69 horse, 421, 483
Eutrombicula alfreddugesi, 8 house, 428
Euura, 465 ichneumon, 421
Euxoa scandens, 309 leaf miner, 426
Euzophera— long-legged, 423
magnolialis, 359 louse, 428
ostricolorella, 359 March, 417
semifuneralis, 359 May, 44
Evaniidae, 482 robber, 422
Evora hemidesma, 360 sand, 416
Exartema— serewworm, 27, 428
permundanum, 374 -snipe, 422
quadrifidum, 374 soldier, 421
Exenterus— stable, 428
abruptorius, 448, 471 stone, 44
canadensis, 442, 444 striped horse, 422
Exorista larvarum, 323, 430 tachina, 429
Exoteleia— tachinid, 429, 478
chilleotti, 392 thick-headed, 424
dodecella, 392 tsetse, 428
nepheos, 392 vinegar, 425
pintfoliella, 391 walnut husk, 424
white, 438, 474
Fascista cercerisella, 394 flower thrips, 42
Fentonia marthesia, 316 Formica exsectoides, 491

Fenusa— Formicidae, 31, 32, 39, 486

dohrnui, 28, 456

pusilla, 23, 454

ulmi, 23, 455
Feralia jocosa, 309
Ferrisia virgata, 100

Frankliniella tritici, 42
fruitworm, green, 309
Fulgoridae, 32
Fumaria casta, 411

Figitidae, 478 Galerita—
filament bearer, 342 bicolor, 118
filbertworm, 373 janus, 118
Fiorimia— gall midges—
externa, 111 balsam, 418
theae, 112 boxelder, 418

flatid planthoppers, 68
Flatidae, 68

willow beaked, 420
galls—

flatids, 496 acorn pip, 480
fleas, 40 gouty oak, 480, 481
cat, 41 hackberry-nipple, 76
dog, 41 horned oak, 479
human, 41 large oak-apple, 481, 482
oriental rat, 41 noxious oak, 481
Flies—415 oak fig, 482
bee, 423 oak flake, 481
big-headed, 423 oak potato, 480
black, 416 ribbed bud, 480

black blow, 429

black cherry fruit, 424

black horse, 421
blow, 428

626

succulent oak, 482
Gargaphia tiliae, 64
Gasteruption, 483
Gaurax apicalis, 426

Gelechiidae, 390
geometers—
chain-spotted, 345
notched-wing, 346
Geometridae, 334
Geopinus incrassatus, 118
giant hornet, 494
Gibbobruchus mimus, 137
girdlers—
birch and beech, 201
twig, 191, 502
Gluphisia septentrionis, 317
Glycobius speciosus, 180
Glyphipterygidae, 396
Glyptoscelis—
barbata, 145
pubescens, 145
Gnathotrichus—
aciculatus, 269
materiarius, 268
gnats—
fungus, 417
Gnophothrips fuscus, 42
Gobaishia ulmifusa, 85
Goes—
dibilis, 183
pulcher, 183
pulverulentus, 183
tessellatus, 183
tigrinus, 182
Goniozus—
electus, 485
longinervis, 485
Gossyparia spuria, 22, 98
Gracillaria—
azaleella, 409
bimaculatella, 409
negundella, 408
packardella, 409
pulchella, 409
purpuriella, 409
quercinigrella, 409
sassafrasella, 409
syringella, 23, 409
Gracillariidae, 406
grape leaf folder, 350
Graphium marcellus, 292
Graphocephala versuta, 70
Graphognathus, 224
Grasshoppers—
clear-winged, 58
differential, 58
eastern lubber, 58
lesser migratory, 58
long-horned, 59
red-legged, 58
short-horned, 57
two-striped, 58
Griselda radicana, 374
grubs—
common cattle, 431
northern cattle, 431
white, 146, 485
Gryllidae, 34, 59
Gryllotalpa gryllotalpa, 22, 60
Gryllotalpidae, 31
Gryllus, 60

Gyponana, 69
Gypsonoma haimbachiana, 369
Gyroptidae, 40

Habrolepis dalmani, 475
hackberry stargall, 76
Halisidota—

caryae, 304

harrisii, 306

maculata, 305

tessellaris, 305
Haltichella—

rhyacioniae, 478

xanticles, 478
Haploa—

clymene, 307

lecontei, 307
Harpalus spp., 118
Hedia chionosema, 374
Heliozelidae, 403
hellgrammites, 43
Hemerobiidae, 43
Hemerocampa—

definita, 319

leucostigma, 318
Hemichroa crocea, 23, 463
Hemicoelus—

carinatus, 132

gibbicollis, 132

umbrosus, 132
Hemicrepidus bilobatus, 154
Hemileuca—

lucina, 300

maia, 299

nevadensis, 300

Hemiptera, 18, 28, 36, 61, 68, 475

Hepialidae, 35, 414
Hesperiidae, 294

Heterarthrus nemoratus, 23, 453
Heterobostrychus aequalis, 22, 136

Heterocampa—
bilineata, 316
biundata, 316
guttivitta, 316
manteo, 315
umbrata, 316

Hexapoda, 2

Hexarthrum ulkei, 224

hickory horned devil, 303

hickory shuckworm, 372, 472, 477,

485
Hippelates, 426
Hippoboscidae, 428
Hippodamia convergens, 123
Histeridae, 114, 119
Holcocera lepidophaga, 396
Holostrophus bifasciatus, 125
Homadula anisocentra, 23, 396
Homaledra sabalella, 390
Homalodisca—

coagulata, 70

insolita, 70

Homoptera, 22, 31, 32, 33, 34, 35, 36,
37, 42, 67, 471, 472, 474, 475, 496

Hoplocampa—
halcyon, 464
lacteipennis, 465

Hoplocampa—Continued
montanicola, 465
oskina, 464
pallipes, 465

hopper, three-cornered alfalfa, 71

Hormaphis—
hamamelidis, 85
spinosus, 85

horned passalus, 119

hornets—493
bald-faced, 494
giant, 494

horntails—465, 471
black and red, 466
blue, 466
white-horned, 466
yellow-horned, 466

Hubbellia marginifera, 59

Hyalophora cecropia, 298

Hylastes—
exilis, 252
porculus, 252
salebrosus, 252
tenuis, 252

Hylecoetus lugubris, 126

Hylemya platura, 427

Hylobius—
aliradicis, 217
congener, 217
pales, 213
pinicola, 216
radicis, 215
rhizophagus, 216
warreni, 216

Hylocurus—
bicornis, 252
biorbis, 252
harnedi, 252
langstoni, 253
rudis, 252
spadix, 252

Hylotrupes bajulus, 22, 206

Hylurgopinus rufipes, 250

Hylurgops pinifex, 252

Hymenoptera, 18, 19, 28, 28, 31, 32,
33, 34, 36, 39, 423, 424, 431, 475,
476, 477

Hyparpax—
aurora, 317
perophoroides, 317

Hyperaeschra stragula, 313

Hyperaspis—
congressis, 105, 123
signata, 105

Hyperetis amicaria, 342

Hyphantria—
cunea, 306
textor, 306

Hypoderma—
bovis, 431
lineatum, 431

Hypothenemus—
chapuisi, 254
dissimilis, 254
eruditus, 254
georgiae, 254
interstitialis, 254

628

Hypothenemus—Continued
obscurus, 254
quercus, 254
rotundicollis, 254
salicis, 254
Hypoxylon, 162
Hypulus concolor, 125
Hyssopus, 474

Ibalia—
anceps, 478
ensiger, 478
maculipennis, 478
scalpellator, 478
Ibaliidae, 478
Icerya, 95
Ichneumonidae, 468, 469, 470
Ichneumonoidea, 468
ichneumons, 470, 476
Ichthyura—
albosigma, 311
apicalis, 311
brucei, 311
inclusa, 310
strigosa, 311
Idiocerus, 69

Incisitermes—
schwarzi, 46, 53
snyderi, 46

Incurvariidae, 413

Ips—21, 450

avulsus, 260
borealis, 262
calligraphus, 260, 502
grandicollis, 260, 502
latidens, 262
perroti, 262
perturbatus, 262
pini, 262, 502
Iridomyrmex—
humilis, 492
pruinosus, 492
Isoptera, 22, 31, 39, 45
Ithycerus noveboracensis, 209
Itoplectis—
4-cingulatus, 471
conquisitor, 412, 471
viduata, 471
Ixodes scapularis, 9

Janetiella coloradensis, 420
Janus abbreviatus, 467
Japanagromyza viridula, 427

Kalotermes approximatus, 46, 53
katydids—59
angular-winged, 59
broad-winged, 59
fork-tailed bush, 59
Kermes—
galliformis, 97
kingit, 97
pettiti, 97
pubescens, 97, 475
trinotatus, 97
Kermococcidae, 97

Keys—
aphids, 78
beetle larvae, 113
Buprestidae adults, 157
Buprestidae larvae, 159
cerambycid larvae, 171
injuries by Scolytidae and Platy-
podidae, 229
orders and families based on types
of injury, 30
lepidopterous larvae, 275
powder-post beetle larvae, 126
scale insects, 93
termites, 46
Kleidocerys resedae geminatus, 67
Knulliana cincta, 178
Kratochviliana laricinellae, 454, 473

Lacewings—

brown, 43

golden-eye, 43

green, 43
Lachnus, 82
Lacon—

avita, 154

discoidea, 154
Lacosma chiridota, 347
Lacosomidae, 347
Laemobothridae, 40
Laetilla coccidivora, 102, 105, 360
Lagoa crispata, 350
Lagriidae, 116
Lambdina—

athasaria athasaria, 345

athasaria pellucidaria, 345

endopiaria, 345

fiscellaria, 344
Lamellicornia, 114
Lapara—

bombycoides, 297

coniferarum, 297
Laphria—

flavicollis, 422

index, 422

thoracica, 422
large aspen tortrix, 383
Laricobius erichsonti, 89, 124
Lasiocampidae, 327
Lasioderma, 127, 327
Laspeyresia—

anaranjada, 372

caryana, 372

erotella, 373

ingens, 373

toreuta, 372

youngana, 371
Latrodectus mactans, 5
leafhopper, 43, 68, 496

white-banded elm, 69, 500
leaf miners, 406

arborvitae, 403, 478

aspen, 408

basswood, 141

birch, 454

boxwood, 418

elm, 455

leaf miners—Continued

European alder, 456

gregarious oak, 406

holly, 426

lilac, 409

locust, 143, 472, 478

sassafras, 409

solitary oak, 406

tupelo, 403

unspotted, 408
leaf rollers—

basswood, 350

fruit tree, 377

hickory, 384

locust, 358

oblique-banded, 383

red-banded, 384
lecanium—

European fruit, 105, 475

oak, 106

taxus, 106
Lecanium—

caryae, 106

corni, 105

fletcheri, 106

nigrofasciatum, 105

quercifex, 106

quercitronis, 106
Lecanodiaspis tessalatus, 112
Leperisinus—

aculeatus, 249
Lepidoptera, 19, 23, 24, 31, 32, 33,

34, 35, 36, 38, 274, 415, 423, 430,

469, 471, 474, 475, 478, 484
Lepidosaphes—

ulmi, 109

yanagicola, 109
Leptocoris trivittatus, 66
Leptogaster flavipes, 422
Leptoglossus—

corculus, 66

occidentalis, 66
Leschenaultia fulvipes, 300
Lespesia—

ciliata, 298

frenchii, 299
Leucopis obscura, 425
Leucopomyia pulvinarie, 102
Lexis bicolor, 307
Lice—

book, 42

chewing, 238

jumping plant, 76

plant, 72

sucking, 40

wood, 3
Lichenophanes—

armiger, 136

bicornis, 136
Limacodidae, 347
Limenitis—

archippus, 293

arthemis, 294

astyanax, 294
Limnoria lignorum, 3
Liothrips umbripennis, 42
Lipoptena, 428

629

Litaneutria minor, 55
Lithocolletis—
aceriella, 407
bethunella, 407
cincinnatiella, 406
corylisella, 408
crataegella, 407
hamadryadella, 406
hamameliella, 407
lucetiella, 407
ostensackenella, 407
robiniella, 407
salicifoliella, 407
tremuloidiella, 407
trinotella, 407
Lithophane—
antennata, 309
latinicerea, 309
locust, post-oak, 58
Lonchaea—
corticis, 425
polita, 425
Lonchaeidae, 425
Longistigma caryae, 82
looper—
false hemlock, 344
green spruce, 338
hemlock, 344, 478, 483
linden, 341
saddle back, 339
Lophodonta—
angulosa, 313
ferruginea, 313
louse—
body, 40
crab, 40
wood, 3
Loxosceles reclusa, 5
Lucanidae, 145
Lucanus elaphus, 145
Lycia ursaria, 342
Lyctidae, 39, 126, 128
Lyctus—21, 127
brunneus, 130
cavicollis, 130
linearis, 130
opaculus, 130
planicollis, 129
Lygaeidae, 67
Lygus lineolaris, 65
Lymantor decipiens, 254
Lymantriidae, 318
Lymexylidae, 38, 116
Lymexylonidae, 125
Lymire edwardsii, 304
Lynchia americana, 428
Lyonetiidae, 409
Lytta muttalli, 121

Machimia tentoriferella, 395
Macremphytus, 465
Macrobasis—

fabricii, 120

unicolor, 120

Macrodactylus subspinosus, 150

Macropsis, 69
Macrosiphum liriodendri, 84

630

Macroxyela, 433
Madiza glabra, 426
Magdalis—

armicollis, 224

austera, 225

austera substiga, 225

barbicornis, 225

barbita, 224

hispoides, 225

olyra, 225

pandura, 225

perforatas, 224

salicis, 225
maggot—

apple, 424

seed corn, 427
maggots, 415, 427
Magicicada—

cassini, 75

septendecim, 74
Malacosoma—

americanum, 328

californicum lutescens, 330

californicum pluviale, 330
_disstria, 339

tigris, 330
Maladera castanea, 23, 149
Mallophaga, 40
mantid—

Carolina, 55

Chinese, 55

European, 55

narow-winged, 55
Mantidae, 54
Mantis religiosa, 55
maple phenacoccus, 99
maple leaf cutter, 413
Margarodidae, 34, 35, 37, 95
Marmara fasciella, 408
Martesia, 11
Matsucoccus—

alabamae, 96

gallicola, 96

matsumurae, 96

resinosae, 22, 95
Mayetiola rigidae, 420
Medetera, 423
Megachilidae, 33, 497
Megacyllene—

antennata, 178

caryae, 178

robiniae, 176
Megalodontoidea, 432
Megalopyge opercularis, 349
Megalopygidae, 348
Megaphasma denticrus, 57
Megarhyssa—

atrata atrata, 465

lunator, 471

macrurus macrurus, 465
Megastigmus—

amelanchieris, 476

laricis, 476

specularis, 476
Megaxyela, 433
Melanagromyza—

schineri, 427

tiliae, 427

Melanaspis—

obscura, 108

tenebricosa, 108
Melandrya striata, 125
Melandryidae, 115, 116, 124
Melanolestes picipes, 65
Melanolophia canadaria, 338
Melanophila——

acuminata, 169

aeneola, 169

drummondi, 169

fulvoguttata, 168, 169

notata, 169
Melanoplus—

bivittatus, 58

bruneri, 58

differentialis, 58

femurrubrum, 58

punctulatus, 58

sanguinipes, 58
Melissopus latiferreanus, 373
Melittomma sericeum, 126
Meloidae, 32, 120
Melophagus ovinus, 428
Melsheimer’s sack bearer, 347
Membracidae, 34, 36, 70
membracids, 496
Menoponidae, 40
Mesitiopterus kahlii, 57, 484
Mesoleius tenthredinis, 458, 471
Messa populifoliella, 457
Metapalma spectabile, 475
Metcalfa pruinosa, 68
Meteorus versicolor, 469
Micracis—

meridianus, 253

suturalis, 253

swainei, 253
Micracisella—

nanula, 253

opacicollis, 253

Microbregma emarginatum emargi-

natum, 133
Microcentum—
retinerve, 59
rhombifolium, 59
Microwesia misella, 123
Micrutalis calva, 71
midges—416
gall, 417
gouty pitch, 418
juniper, 418
red-pine needle, 418
millipedes, 4
Mindarus abietinus, 83
Minthea rugicollis, 23, 131
Miridae, 65
Misogada unicolor, 318
mites—5, 18, 43
itch, 8
maple bladder-gall, 7
pear leaf blister, 7
pine bud, 8
southern red, 6
spider—6
carmine, 7
four-spotted, 7
Schoene, 7

mites—Continued
spider—Continued
spruce, 6
Mnemonica auricyonia, 414
Mollusca, Phylum, 2, 10
Monarthropalpus buxi, 23, 418
Monarthrum—
fasciatum, 267
mali, 267
Monellia—
caryella, 84
costalis, 84
microsetosa, 84
Monelliopsis nigropunctata, 84
Monocesta coryli, 141
Monochamus—
carolinensis, 205
maculosus, 205
marmorator, 205
notatus, 205
scutellatus, 203
titillator, 203
Monoctenus melliceps, 439
Monodontomerus—
aerus, 323, 476
dentipes, 451, 476
indiscretus, 477
japonicus, 477
minor, 477
montivagus, 477
Monophylla terminata, 155
Moodna ostrinella, 359
Mordellidae, 116, 126
Mordwilkoja
vagabunda, 85
mosquitoes, 415
moths—
acorn, 396
American dagger, 308
apple fruit, 403
bagworm, 410

brown-tail, 24, 325, 480, 431, 469,

472, 476
buck, 299
carpenterworm, 386
casebearer, 404
cecropla, 298, 477
cherry scallop shell, 337
clearwing, 397
cottonwood dagger, 308
crinkled flannel, 350
cynthia, 297
dark tussock, 320
definite-marked tussock, 319

European pine shoot, 12, 21, 24,
361, 470, 471, 472, 474, 475, 477,

478

eye-spotted bud, 367
flannel, 348
geometrid, 334
giant silkworm, 297

FY DSYs ice Sal Oe ZA Ae Zo mod,
Go le23.03 205 450 sre oilers GO),

472, 474, 475, 476, 483
hag, 347
hickory tussock, 304
hornet, 398
imperial, 303

moths—Continued
io, 299
jack-pine shoot,
lappet, 333
leaf roller, 374
leopard, 386
luna, 298
Nantucket pine tip, 363, 472, 475,
477, 478, 485
Nevada buck, 300
notodontid, 309
nun, 430
olethreutid, 360
oriental, 347, 349, 431, 484
pale tussock, 305, 306
pepper and salt, 346
pine bud, 392
pine tube, 384
pine tussock, 320
pitch-twig, 365
polyphemus, 299
promethea, 298
regal, 303
royal, 300
rusty tussock, 318
satin, 24, 324, 430, 470, 472, 475,
483
saturnid, 478
shoot, 474
slug caterpillar, 347
smeared dagger, 308
spear-marked black, 337
sphinx, 294
spotted tussock, 305
spruce bud, 369
spruce seed, 371
sycamore tussock, 306
tent caterpillar, 327
tiger, 304
tip, 474
tussock, 318
underwing, 308
wax, 390
western-pine tip, 365, 478, 485
white-marked tussock, 318, 476,
478, 483
winter, 336
Zimmerman pine, 356
Musca domestica, 428
Muscidae, 31, 428
Muscina spp. 428
Mutillidae, 485
Mycetophilidae, 417
Mycodiplosis alternata, 421
Myeloborus—
fivazi, 259
ramiperda, 259
Mymaridae, 472
Myrmeleontidae, 43
Myzocallis—
alhambra, 84
alnifoliae, 84
bella, 84
caryaefoliae, 84
discolor, 84
kahawaluokalani, 84
melanocera, 84

632

Myzocallis—Continued
tiliae, 84
ulmifolvi, 84

Nabidae, 65
Nabis sordidus, 65
Nacerdes melanura, 23, 125
Nadata gibbosa, 313
Nalepella tsugifoliae, 8
needle miner—

pine, 391

spruce, 360
Neichnea laticornis, 156
Nematidium filiforme, 124
Nematocampa limbata, 342
Nematus—

abbotti, 461

fulvicrus, 461

hyalinus, 461

limbatus, 461

ostryae, 461

pinguidorsum, 461

populi, 461

-pinguidorsum, 461

tibialis, 461

ventralis, 461
Nemosoma cylindricus,

(see Corticotomus cylindricus)
Neoclytus—

acuminatus, 184

caprea, 184

fulguratus, 184

jouteli, 184

mucronatus, 184, 186

muricatulus, 184

muricatulus muricatulus, 184

scutellaris, 184
Neodiprion—

abbott, 447

abietis, 446.

americanum,

(see N. t. taedae)

compar, 450

dubiosus, 447

excitans, 19, 449

hetricki, 450

lecontei, 439

maurus, 450

merkeli, 450

nanulus nanulus, 445

nigroscutum, 450

pinetum, 446

pinirigidae, 450, 473

pratti banksianae, 442

pratti paradoxicus, 443, 473

pratti pratti, 441

rugifrons, 447

sertifer, 23, 447

swainei, 444

taedae linearis, 443, 445

taedae taedae, 443

virginianus, 447

warreni, 450
Neolecanium cornuparvum, 103
Neolygus invitus, 65
Neotychodes trilineatus, 199

Neotermes—

castaneus, 54

jouteli, 46, 53
Nephopteryx—

subcaesiella, 358

subfuscella, 358

virgatella, 358
Nepticula sericopeza, 23, 413
Nepticulidae, 34, 413
Neptyia—

canosaria, 344

semiclusaria, 344
Nerice bidentata, 314
Neuroptera, 18, 42, 476
Neuroterus—

fioccosus, 481

noxiosus, 481

quercusbatatus, 480
Neurotoma—

crataegi, 435

fasciata, 435

inconspicua, 435
Nicobium castaneum, 134
Nicrophorus, 119
nine-spotted ladybird, 123
Nitidulidae, 115, 121
Noctuidae, 308
Norape ovina, 350
no-see-ums, 416
Notodonta simplaria, 318
Notodontidae, 309
Notolophus antiqua,

(see Orgyria antiqua)
Nuculaspis californica, 111
Nygmia phaeorrhoea, 23, 325
Nymphalidae, 292
Nymphalis—

antiopa, 293

j-album, 292

oak-bark scaler, 199
oak-bark scarrer, 181
Oberea—

ferruginea, 190

myops, 190

ocellata, 190

pallida, 190

ruficollis, 190

schaumi, 190

tripunctata, 189

ulmicola, 190
Obolodiplosis robinae, 420
Obrussa ochrefasciella, 413
Odonata, 45
Odontopus calceatus, 221
Odontota dorsalis, 143
Oecanthus—

exclamationis, 60

fultoni, 60

latipennis, 60

pini, 60
Oecophoridae, 395
Oodemeridae, 39, 116, 125
Oeme rigida, 200
Oestridae, 431
Oiketicus abbotii, 411

Olene plagiata,

(see Dasychira plagiata)
Olethreutidae, 31, 38, 36, 360
Olfersia fumipennis, 428
Oligonychus—

aceris, 6

bicolor, 6

boudreauxi, 6

cunliffer, 6

ilicis, 6

letchworthi, 6

milleri, 6

newcomeri, 6

propetes, 6

ununquis, 6
Oligotrophus—

betulae, 238, 420

papyrifera, 419
Omalophora salicis, 74
Oncideres—

cingulata cingulata, 191

cingulata texanus, 192

pustulatus, 192
Oncometopia orbona, 70
Ooctonus aphrophora, 472

Ooencyrtus kuwanai, 323, 474, 475

Operophtera—

bruceata, 336

brumata, 336
Oracella acuta, 100
Orchesia castanea, 125
Orgilus obscurator, 362, 470
Orgyria antiqua, 23, 318
Orius insidiosus, 65
Ornithoica, 428
Ornithomyia, 428
Orthopleura damicornis, 156

Orthoptera,, 195723) 7316 43253350 34,

54, 475, 476, 477, 478
Orthosia hibisci, 309
Orthosoma brunneum, 205
Orthotomicus—

caelatus, 263

VICINUS, 263
Orussidae, 432, 467
Oscinella coxendix, 426
Osmoderma—

eremicola, 153

scabra, 153
Ostomatidae, 115, 120
Othniidae, 116

Pachybrachys—
carbonarius, 145
othonus, 145
peccans, 145
tridens, 145

Pachylobius picivorus, 217

Pachypsylla—
celtidisastericus, 76
celtidisgemma, 76
celtidisinteneris, 77
celtidismamma, 76
celtidisvesicula, 76
venusta, 77

633

Packardia geminata, 348
Paleacrita vernata, 339
palmerworm, 393
Palthis angulalis, 309
Pamphiliidae, 433
Pandemis—

lamprosana, 375

limitata, 375
Panthea—

acronytoides, 309

furcilla, 309
Pantographa limata, 350, 352
Paonias—

excaecatus, 297

myops, 297
Papaipema furcata, 309
Papilio—

cresphontes, 292

glaucus, 291

troilus, 291
Papilionidae, 291
Paraclemensia acerifoliella, 413
Parallelodiplosis florida, 420
Paralobesia liriodendrana, 373
Parandra brunnea, 197
Paranthrene—

dollii castanea, 402

dollii dollivi, 402

palm, 401

simulans simulans, 402

tabaniformis, 402

tricincta, 402
Parasetigena agilis, 328, 430
Parectopa robiniella, 408
Parharmonia pint,

(see Vespamina pini)
Paria—

quadrinotata, 145

sexnotata, 145
Passalidae, 119
Pealius azaleae, 22, 92
pear-slug, 457
Pediculoides ventricosus, 392
Pediculus humanus, 40
Pelecinidae, 483
Pelecinus polyturator, 483
Pelidnota punctata, 153
Pemphigus, 79
Pentatomidae, 62
Penthimia, 69
Pergidae, 436
Periclista, 468
Perilampidae, 476
Perilampus hyalinus, 476
Periphyllus—

americanus, 84

lyropictus, 84

negundinis, 83
Perisierola punctaticeps, 485
Pero—

honestarius, 344

morrisonarius, 344
Peronea variana,

(see Acleris variana)
persimmon psylla, 77
Petalium—

bistriatum bistriatum, 133

seriatum, 133

634

Petrova—

albicapitana, 366

comstockiana, 365

houseri, 367

virginiana, 365
Phaenicia sericata, 429
Phaeoura quernaria, 342
Phalangida, 10
Phalonia rutilana, 23, 374
Phaloniidae, 374
Phampiliidae, 433
Phanerotoma sp. 372
Phanomeris phyllotomae, 454, 470
Phasgonophora sulcata, 478
Phasmatidae, 33, 55
Phenacaspis pinifoliae, 109
Phenacoccus—

acericola, 22, 99

dearnessi, 99

serratus, 99
Pheosia rimosa, 313 |
Phigalia titea, 25, 339 |
Philaenus spumarius, 74 |
Philopteridae, 40
Philosama cynthia,

(see Samia cynthia)
Phloeosinus—

canadensis, 249

dentatus, 248

pini, 249

taxodti, 248

texanus, 249
Phloeotribus—

dentifrons, 240

frontalis, 240

liminaris, 240 |
Phlogistosternus dislocatus, 156
Phlyctaenia—

coronata, 351

tertialis,

(see P. coronata)

Phobetron pithecium, 347
Phobocampe disparis, 323
Phormia regina, 429
Phthirus pubis, 40
Phthorophloeus,

(see Phloeotribus)
Phycitidae, 31, 32, 34, 38
Phyllaphis, 80
Phyllidae, 36
Phyllobius—

intrusus, 218

oblongus, 23, 219
Phyllocnistis—

liquidambarisella, 408

liriodendrella, 408

magnoliella, 408

populiella, 408
Phyllophaga—

crenulata, 148

drakei, 146

forsteri, 148

implicta, 148

luctuosa, 147

prunina, 148

prununculina, 148

rugosa, 148

tristis, 148

Phyllotoma nemorata,

(see Heterarthrus nemoratus)
phylloxera—

pecan, 92

pecan leaf, 92
Phylloxera—

caryaecaulis, 92

devastatrix, 92

notabilis, 92

nyssae, 92

rileyi, 92
Phylloxeridae, 33, 36, 37, 86
Phymatodes—

dimidiatus, 197

testaceus, 197

varius, 197
Physocnemum—

andreae, 197

brevilneum, 197

violareipenne, 197
Physokermes piceae, 106
Physostegania pustularia, 338
Phytobia—

amelanchieris, 426

clara, 427

posticata, 427

pruinosa, 426

pruni, 426

setosa, 426
Phytomyza ilicicola, 426
Phytoptus pini, 8
pigeon tremex, 466
Pikonema—

alaskensis, 459

dimmockti, 460
Pimpla turionellae, 362, 471
pine colaspis, 142
Pineus—

coloradensis, 91

floccus, 91

pinifoliae, 91

similis, 91

strobi, 22, 89
Pipunculidae, 423
Pissodes—

affinis, 213

approximatus, 211

dubius, 213

fisket, 213

nemorensis, 212

rotundatus, 213

strobi, 209
pitch nodule maker, 366
Pitedia uhleri, 62
Pityoborus comatus, 258
Pityogenes—

hopkinsi, 258

meridianus, 258

plagiatus, 258
Pityokteines sparsus, 263
Pityophthorus—

annectens, 255

balsameus, 255

biovalis, 255

cariniceps, 255

cascoensis, 231

consimilis, 231

crinalis, 255

Pityophthorus—Continued
dentifrons, 255
liqudambarus, 255, 277
natalis, 255
nudus, 231
opaculus, 255
patchi, 255
puberlus, 255
pulchellus, 255
pulicarius, 255
pullus, 231
rhois, 255
scriptor, 255
shepardi, 255
tonsus, 231

Plagiodera versicolora, 23, 143

Plagiometriona clavata, 145

Plagodis—
phlogosaria kuentzingaria, 342
serinaria, 342

planthoppers, 68

Platybregmus canadensis, 134

Platycerus quercus, 145

Platycotis vittata, 71

Platygasteridae, 484

Platygaster pini, 418

Platypodidae, 34, 117, 272

Platypus—
compositus, 273
flavicornis, 272
quadridentatus, 273

Platytetranychus—
multidiquituli, 7
thujae, 7

Plecoptera, 44

Plectrodera scalator, 179

Plegaderus spp., 119

Pleroneura brunneicornis, 433

plum curculio, 227

Podapion gallicola, 218

Podisus maculiventris, 62

Podosesia—
syringiae fraxini, 400
syringiae syringiae, 400

Poecilonota—
cyanipes, 169
thrueura, 170

Polistes, 495

Polycaon stoutii, 136

Polydrusus impressifrons, 23, 220

Polygonia—
comma, 292
interrogationis, 292

Polygraphus rufipennis. 264

Polymoechus brevipes, 153

Polynema striaticorne, 472

Polyphylla—
hammondi, 149
occidentalis, 149
variolosa, 149

Pomphopoea—
aenea, 120
polita, 121
say, 120

Pompilidae, 493, 495

Ponana, 69

Popilius disjunctus, 119

Popillia japonica, 23, 151

635

poplar tent maker, 310
Porthetria dispar, 23, 320
praying mantids, 54
Priocera castanea, 156
Prionoxystus—

macmurtrei, 389

robiniae, 388
Prionus—

imbricornis, 200

laticollis, 200

pocularis, 200
Pristaulacus—

bilobatus, 483

rufitarsis, 483
Pristiphora—

chlorea, 459

erichsonu, 23, 458, 460

geniculata, 28, 459, 460

siskiyouensis, 459

sycophanta, 459
Prochoerodes transversata, 346
Prociphilus—

bumeliae, 83

corrugatans, 85

fraxinifoli, 85

imbricator, 84

tessellatus, 85
Proctotrupoidea, 482
Prodiplosis morrisi, 420
Profenusa—

canadensis, 457

lucifex, 457

mainensis, 456

thomsoni, 456
Prolimacodes—

badia, 348

scapha, 348
Prorhinotermes simplex, 46, 54
Prosapia bicincta, 73
Prospaltella perniciosa, 107
Proteoteras—

aesculana, 369

willingana, 369
Protobarmia porcelaria, 346
Protocalliphora, 429
Protozoa, 2
Pselactus spadix, 224
Psen, 496
Pseudaonidia duplex, 111
Pseudaphycus malinus, 100
Pseudaulacaspis pentagona, 112
Pseudobogeris, 431
Pseudobrachium mandibulare, 484
Pseudocneorhinus bifasciatus, 23, 219
Pseudococcidae, 99
Pseudococcus—

adonidum, 100

comstocki, 22, 100

maritimus, 100
Pseudolucanus capreolus, 145
Pseudophilippia quaintancii, 106
Pseudopityophthorus—

asperulus, 256

minutissimum, 255

Pruinosus, 256

pubescens, 256
Pseudoscorpionidae, 10
pseudoscorpions, 10

636

Pseudotachinomya webberi, 342
Pseudothysanoes—
lecontei, 253
rigidus, 253
Psilocorsis—
faginella, 395
fletcherella, 395
obsoletella, 395
quercicella, 395
reflexella, 395
Psilus politus, 483
psocids, 42
Psocoptera, 42
Psorophora, 416
Psychidae, 410
Psylla—
annulata, 77
buxi, 77
carpinicola, 77
floccosa, 77
galeaformis, 77
trimaculata, 77
Psyllidae, 35, 76
psyllids—424
blistergall, 76
boxwood, 77
budgall, 76
petiolegall, 77
Pterocomma—
populifoliae, 84
salicis, 84
smithae, 84
Pteromalidae, 477
Pteronidea ventralis,

(see Nematus ventralis)
Pterophylla camellifolia, 59
Ptilinus—127

pectinicornis, 133
pruinosus, 133
rufiicornis, 133
Ptinidae, 116, 126, 136
Ptinus—
clavipes, 137
fur, 137
Ptosima gibbicollis, 169
Pulex irritans, 40
Pullus impexus,

(see Scymnus impexus) |

Pulvinaria— |
acericola, 102
floccifera, 103
innumerabilis, 101

punkies, 416

Pyraustidae, 350

Pyrochroidae, 115

Pyrrhalta—
cavicollis, 140
decora decora, 140
luteola, 23, 138
tuberculata, 140

Quadraspidiotus—
forbesi, 107
gigas, 107
juglansregiae, 107
perniciosus, 22, 106 |

question-mark, 292
quince curculio, 227

Ramosia rhododendri,

(see Synanthedon rhododendri)
Raphia frater, 309
Recurvaria—

apicitripunctella,
(see Coleotechnites apicitripunc-
tella)
piceaella,
(see Coleotechnites piceaella)
thujaella,

(see Coleotechnites thujaella)
Reduviidae, 64
Reticulitermes—

arenicola, 47

flavipes, 47, 48, 49, 50

hageni, 47, 49

tibialis, 46, 49

virginicus, 47, 50
Rhabdophaga salicis, 23
Rhagionidae, 422
Rhagoletis—

cingulata, 424

completa, 424

fausta, 424

pomonella, 424
Rhinotermitidae, 47
Rhipiceridae, 115
Rhopalidae, 66
Rhyacionia—

adana, 362

buoliana, 23, 361

frustrana bushnelli, 365

frustrana frustrana, 363

frustrana neomexicana, 365

rigidana, 363, 477, 478

sonia, 365

subtropica, 365
Rhydinofoenus, 483
Rhynchaenus—

pallicornis, 223

rufipes, 222
Rhynchophorus cruentatus, 223
Rhyncophora, 208
Rhysodidae, 114
Ricinidae, 40
Romalea microptera, 58
Romaleum rufulum,

(see Enaphalodes rufulus)

saddled prominent, 316, 472
Salebrica subcaesiella,
(see Nephopteryx subcaesiella)
Salpingidae, 115
Samia—
cynthia, 23, 297
cecropia,
(see Hyalophora cecropia)
sandflies, 416
Sannina uroceriformis, 400
Saperda—
calcarata, 162, 185, 186
candida, 187
concolor, 188
cretata, 188
discoidea, 187

Saperda—Continued
fayi, 188
imitans, 188
lateralis, 188
moesta, 188
mutica, 188
obliqua, 188
tridentata,
(see Hutetrapha tridentata)
vestita, 185
Sapygidae, 486
Sarcophaga—
aldrichi, 429
houghi, 429
Sarcophagidae, 429
sarcophagids, 476
Sarcoptes scabei, 8
Sarcoptidae, 8
Saturniidae, 297
sawflies—117, 436
argid, 437
Arkansas pine, 443, 445
balsam-fir, 446, 473
birch, 437
birch leaf-mining, 453, 470, 473
black-headed ash, 464
brown-headed ash, 463
cimbicid, 437
conifer, 439
dusky birch, 462
elm, 438, 472
European pine, 12, 19, 24, 447, 470,
471, 473
European spruce, 18, 19, 24, 431,
452, 470, 471, 4738, 477
green-headed spruce, 460
introduced pine, 28, 450, 476
jack-pine, 19, 442
larch, 458, 471
loblolly pine, 443
mountain-ash, 459
nesting-pine, 434
pin oak, 457
plum web-spinning, 435
red-headed pine, 19, 4381, 439, 473
red-pine, 445, 473
stem, 467
striped alder, 463
Swaine jack-pine, 24, 444
Virginia pine, 19, 24, 441, 473
web-spinning, 433
white-pine, 446
willow, 461
willow shoot, 467
yellow-headed spruce, 459, 472
sawyers—
balsam fir, 205
Northeastern, 205
southern pine, 203
spotted pine, 205
white-spotted, 203
scales—67, 121, 424
armored, 106
bamboo, 101
beech, 97, 501
black pine-leaf, 111
brown elm, 105
camphor, 111

637

scales—Continued

cottony maple, 101

diaspine, 43

elm scurfy, 108

euonymus, 111, 112

English ivy, 101

European elm, 98, 473

Fletcher, 106

gloomy, 108

golden oak, 101, 474

hemlock, 108

holly, 101

juniper, 111

magnolia, 103, 104

maple leaf, 102

margarodid, 95

obscure, 108

oleander, 101

oystershell, 109, 110, 475

persimmon, 112

pine needle, 109

pine tortoise, 103, 123

pine twig gall, 65, 96

pit, 100

Putnam, 108

red-pine, 65, 95, 123

San Jose, 106, 475

scurfy, 108

soft, 101

spruce bud, 106

terrapin, 105

tuliptree, 103

walnut, 107

white peach, 112

willow scurfy, 109

woolly pine, 106

yellow oak, 101
Scaphoideus luteolus, 69, 70
Scapteriscus—

abbreviatus, 60

acletus, 61

vicinus, 60
Scarabaeidae, 31, 32, 145
scarabaeids, 485
scarabs, 145
Scarites subterraneus, 118
Scelionidae, 483
Schizolachnus—

lanosus, 83

piniradiatae, 83
Schizonotus sieboldi, 144, 477
Schizura—

concinna, 316

ipomaeae, 317

leptinoides, 317

unicornis, 317
Sciaphila duplex, 374
Scleroderma, 484
Scobicia, bidentata, 136
Scolioidea, 485
Scolidae, 485
Scolopendridae, 4

Scolytidae, 31, 33, 34, 35, 38, 39, 117,

227, 265
Scolytus—
fagi, 238
mali, 237
multistriatus, 23, 235

638

Seolytus—Continued
muticus, 239
piceae, 239
quadrispinosus, 235
rugulosus, 238
sulcatus,

(see S. malt)
scorpion, striped, 9
Scorpionidae, 9
scorpions, false, 10
Scudderia—

curvicauda, 59
furcata, 59

Scymnus—
impexus, 123
lacustris, 105, 123

Seirarctia echo, 307

Semanotus—
ligneus, 198
litigiosus, 198

Semiothisa—
bisignata, 338
granitata, 338
_ocellinata, 338
sexmaculata, 338

Serica—
intermixta, 149
peregrine, 485
sericae, 149
tristis, 149
vespertina, 149

Serropalpus barbatus,

sheep ked, 428

shield bearers, 403

shipworms, 2, 10

Sibine stimulae, 347

Silpha, 119

Silphidae, 114, 119

silver-spotted skipper, 294

Simulidae, 416

Sinea spinipes, 64

Sinodendron rugosum, 145

Siphonaptera, 40

Sirex—
abbottu, 466
cyaneus, 466
edwardsi, 466
juvencus, 23, 466
nigricornis, 466

Siricidae, 465

Siricoidea, 465

Sisyrosea textula, 348

skeletonizers—
apple-and-thorn, 396
birch, 409
maple trumpet, 371
oak, 410, 411, 478
palm leaf, 390

skippers, 294

Smerinthus—
geminatus,

(see S. jamaicensis)

jamaicensis, 297

Smodicum cucujiforme, 189

Solenopsis saevissima richteri,
492

spanworms—

Bruce, 336

—

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23,

spanworms—Continued
elm, 343, 484

Sparganothis—
acerivoranda, 375, 386
dilutocostana, 375
fulgidipenna, 375
pettitana, 375
sulfureana, 375
tristriata, 375

Spathimeigenia aurifrons, 441

Sphaeroma, 3
Sphecidae, 39, 496
Sphecius speciosus, 496
Sphecoidea, 495
Sphingidae, 294
Sphinu—
chersis, 296
drupiferarum, 297
kalmiae, 296
luscitiosa, 296
Sphinx—
catalpa, 295, 483
elm, 295
great ash, 296
walnut, 297
spiders—5, 18, 495
black widow, 5
brown recluse, 5
Spilochalcis—
albifrons, 478
flavopicta, 478
mariae, 478
Spilococcus juniperi, 99
Spilonota ocellana, 23, 367
Spirobolus marginatus, 4
Spissistilus festinus, 71
spittlebugs, 424
springtails, 41
spruce scolytus, 239
Stagmomantis—
carolina, 55
floridensis, 55
Staphylinidae, 114

Stenocorus inquisitor lineatus, 206

Stenodontes dasytomus, 205
Stenoscelis brevis, 223
Stephanidae, 468
Stephanitis—
pyrioides, 64
rhododendri, 22, 64
Stephanoderes,
(see Hypothenemus)
Stephanopachys—
cribratus, 136
densus, 136
hispidulus, 136
TUJOSUS, 136
substriatus, 136
Sterictiphora prunivora, 437
Sthenopis—
argenteomaculatus, 413
thule, 413
Stictia carolina, 496
Stictocephala—
bubalus, 70
militaris, 71
Stilpnotia salicis, 23, 324
Stiretus anchorago, 62

Stomoxys calcitrans, 428
Stratiomyidae, 421
Strongylium, 125

Sturmia inconspicua, 431

sumac datana, 312
swallowtail—
spice-bush, 291
tiger, 291
zebra, 292
Sylvora—
acerni, 399
acerni buscki, 399
Symmerista—
albicosta,

(see S. canicosta)
albifrons, 314
canicosta, 314
leucitys, 314

Symphata, 432
Symphoromyia, 422
Synanthedon—
acerrubri, 399
castanea, 399
geliformis, 399
pictipes, 399
pyri, 399
rhododendri, 398
sapygaeformis, 399
scitula, 398
viburni, 399
Synchlora aerata, 336
Syneta ferruginea, 145
Syrphidae, 423
Systena marginalis, 144

Tabanidae, 421
Tabanus—
abactor, 422
atratus, 421
costalis, 422
lineola, 422
nigrovittatus, 422
quinquevittatus, 422
sulcifrons, 422
Tachinidae, 429
tachinids, 476
Tachydromia, 423
Tachys spp., 117
Taeniogly ptes—
fraseri, 259
rubentis, 259
rufiicollis, 259
Taniva albolineana, 360

Tarsostenus univittatus, 128, 155

T elamona—
decorata, 71
reclivata, 71

Telea polyphemus,

(see Antheraea polyphemus)

T elenomus—
alsophilae, 344, 483
bifidus, 483
californicus, 483
catalpae, 483
clisiocampanae, 484
dalmani, 483
geometriae, 483
hemerocampae, 483

639

Temelucha interruptor, 362, 471
Temnochila virescens, 120
Tenebrionidae, 35, 38, 116, 125
T enebroides—
bimaculatus, 120
corticalis, 120
dubius, 120
T enodera—
angustipennis, 55
aridifolia sinensis, 55, 56
Tenthredinidae, 32, 38, 34, 36, 453
Tenthredinoidea, 436
Tephritidae, 30, 424
Teredo, 10
termites—45
arid-land subterranean, 49
eastern subterranean, 48
Formosan subterranean, 50
nonsubterranean, 52
southern dry-wood, 53
subterranean, 21, 47

tropical rough-headed powder-

post, 54
Tethida cordigera, 464
Tetracis lorata, 346
Tetraleurodes mori, 92
Tetralopha—

asperatella, 62, 351, 352

melanogrammos, 353

militella, 353

robustella, 351

sp. near asperatella, 351
Tetranychidae, 6, 31, 32
Tetranychus—

canadensis, 7

cinnabarinus, 7

ellipticus, 7

homorus, 7

magnoliae, 7

schoenei, 7
Tetrastichus—

brevistigma, 140, 474

holbeini, 474

rugglesi, 474

turionum, 362, 474
Tetropium cinnamopterum, 199
Tetrya bipunctata, 62
Tettigoniidae, 59
Thamnosphecia—

rubrofascia, 399

sigmoidea, 399

various species,

(see Synanthedon)
Thanasimus dubius, 154
Thaumastocoridae, 67
Thecodiplosis piniresinosae, 418
Thelia bimaculata, 496
thrips, 41, 43
Thylacites incanus, 23, 220
Thysanidae, 474
Thysanoes—

berschemiae, 253

fimbricornis, 253

lobdelli, 253
Thysanoptera, 41

Thyridopteryx ephemeraeformis, 411

Thysanura, 41
Tibicen canicularis, 75

640

ticks—

American dog, 9

black-legged, 9
tile-horned prionus, 200
timberworms—

chestnut, 126

oak, 208

sapwood, 126
Tingidae, 36, 62
Tiphia—

asericae, 485

inornata, 485

poppilliavora, 152, 485

vernalis, 152, 485
Tiphiidae, 485
Tipulidae, 484
Tolmerus notatus, 422
T olype—

laricis, 333

velleda, 333
Tomolips quercicola, 224
Tomostethus multicinctus, 463
Tortricidae, 374
Tortricidia flexuosa, 348
Torymidae, 476
Torymus rugglesi, 476
Toumeyella—

liriodendri, 103

numismatica, 103

parvicornis, 105

pini, 105

sp. near numismaticum, 105
Townsendiellomyia nidicola, 430
Toxotrypana curvicauda, 425
Trachykele lecontei, 169
Tragosoma desparius, 198
treehoppers—70

buffalo, 70

two-marked, 71
Tremex columba, 466
Trichiocampus—

gregarius, 461

irregularis, 461

viminalis, 23, 461
Trichiosoma triangulum, 438
Trichiotinus, 153
Trichodestidae, 40
Trichodesma gibbosa, 133
Trichogramma minutum, 324, 472
Trichogrammatidae, 472
Trichopria tabanivora, 483
Trigonura elegans, 478
Tricorynus, 134
Trioza—

diospyri, 77

magnoliae, 77

tripunctata, 77
Trisetaceus cupressi, 8
Trogoxylon—

aequale, 131

caseyi, 131

parallelopidedum, 131
Trombiculidae, 8
Tropaca luna,

(see Actias luna)
Tropidosteptes amoenus, 65
Trypodendron—

betulae, 268

Try podendron—Continued weevils—Continued

bivittatum, 268 red elm bark, 224, 478
lineatum, 267 small chestnut, 226
retusum, 268 strawberry root, 221
scabricollis, 268 white-pine, 209, 423, 426, 475, 477
Trypopitys sericeus, 133 willow flea, 222
twig pruner, 192 whiteflies—92
Tylonotus bimaculatus, 199 azalea, 92
Tymnes tricolor, 145 citrus, 92
Typhlocyba, 69 mulberry, 92
rhododendron, 92
Unaspis euonymi, 111, 112 Winthemia datanae, 311
Urocerus— wireworms, 153
albicornis, 466 YURI Ne
cressoni, 466 butternut woolly, 464
flavicornis, 466 carpenter, 388
green-striped maple, 301
inch, 334

Valentinia glandulella, 396

Vanduzea arquata, 71 orange-humped maple, 314

Vasates— orange-striped oak, 300, 472
aceris-crummena, 7 pink-striped oak, 301
quadripedes, 7, 8 red-humped oak, 314

Vespa crabro germana, 494 secondary screw, 429

Vespamima pini, 399 screw, 27, 428

Vespidae, 39, 493 spiny oak, 300

Vespoidea, 493

Vespula—
arenaria arenaria, 494, 495
maculata, 494

viceroy, 293

measuring, 334

Xanthonia decimnotata, 145
Xanthoteras forticorne, 482
Xenochalepus dorsalis,

(see Odontota dorsalis)

walkingsticks—55 Xenopsylla cheopsis, 41
giant, 57 Xenotemna pallorana, 386
two-striped, 57 Xestobium rufovillosum, 23, 132
the walkingstick, 55, 484 Xiphydria—

Walshiidae, 390 abdominalis, 467

warbles, 431 hicoriae, 467

wasps—436, 483, 484, 485 maculata, 467
gall, 479 mellipes, 467
potter, 493 tibialis, 467
tiphiid, 485 Xiphydriidae, 467

webworms— Xyela—
ailanthus, 403 alpigena, 432
fall, 64, 306, 472, 476, 483 bakeri, 432
juniper, 393 minor, 432
mimosa, 396 obscura, 432
oak, 377 styrax, 432
pine, 65, 351 Xyelidae, 432
pine false, 433 Xylastodoris luteolus, 67

weevils— X yleborus—
apple flea, 223 affinis, 270
arborvitae, 218 cavipennis, 271
Asiatic oak, 221 celsus, 270
black elm bark, 224, 478 dispar, 271
black vine, 220 ferrugineus, 270
broad-nosed bark, 223 howardi, 271
deodar, 212 lecontei, 271
fungus, 209 obesus, 271
Japanese, 219 obliquus, 271
large chestnut, 226 pecanis, 271
New York, 209 pint, 271
northern pine, 211 rubricollis, 271
pales, 213 saxeseni, 269
pecan, 226 say1, 271
pine gall, 218 tachygraphus, 271
pine root collar, 213 volvulus, 271
pitch-eating, 217 xylographus, 270

641

Xyletinus— X ylotrechus—Continued

lugubris, 132 quadrimaculatus, 201

peltatus, 132 sagittatus, 202
Xylobiops basilaris, 134, 502 undulatus, 203
Xyloblaptus quadrispinosus, 136
Xylococculus betulae, 96 yellow jackets, 493
X ylocopa— Yponomeutidae, 402

micans, 498

virginica texana, 498 Zabrachia polita, 421

virginica virginica, 498 Zanolidae, 333
Xylocopinae, 497 Zeiraphera—
Xylophagidae, 421 diniana, 370
Xyloryctes jamaicensis, 152 fortunana, 370
Xylosandrus— ratzeburgiana, 23, 369

germanus, 271 Zelleria haimbachi, 403

zimmermani, 271 Zelus exsanguis, 65
Xyloterinus politus, 264, 267 Zengophora scutellaris, 23, 144
Xylotrechus— Zenillia—

aceris, 201 protuberans, 332

annosus, 203 vulgaris, 342

colonus, 202 Zenodochium coccivorella, 396

obliteratus, 202 Zeuzera pyrina, 23, 386
642

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