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e LIBRARY

OF THE

UNITED STATES
DEPARTMENT OF AGRICULTURE.

Class _

Book£ (L ^

• „t Anrlcuftural Library
USDA, National Agrtcuiw

imoi Baltimore BiW
■\Q6h i -•« ' O070&-2351

U. S. DEPARTMENT OF AGRICULTURE,

Vt

BUREAU OF ENTOMOLOGY— BULLETIN No. 94.

L. O. HOWARD, Entomologist and Chief of Bureau.

INSECTS INJURIOUS TO FORESTS AND
FOREST PRODUCTS.

Kibfs'y, u. s n

I, DAMAGE TO CHESTNUT TELEPHONE AND TELEGRAPH POLES"
BY WOOD-BORING INSECTS.

B.y THOMAS E. SNYDER, M. F., Agent and Expert.

II. BIOLOGY OF THE TERMITES OF THE EASTERN

UNITED STATES, WITH PREVENTIVE

AND REMEDIAL MEASURES.

By THOMAS E« SNYDER, M. E., Entomological Assistant
Forest Insect Investigations.

WASHINGTON:

GOVERNMENT PRINTING OFEIOE.

1916.

B UBEAU OF EXTOMOLOGY.

L. O. Howard, Entomologist and Chief of B ureau .

C. L. Marlatt, Entomologist and Assistant Chief of Bureau.

E. B. 0 'Leaky, Chief Clerk and Executive Assistant.

F. H. Chittenden, in charge of truck crop and stored product insect investigations.

A. D. Hopeixs, in charge of forest insect investigations.

W. D. Hunter, in charge of southern field crop insect investigations.

: — , in charge of cereal and forage insect investigations.

A. L. Quaintance, in charge of deciduous fruit insect investigations.

E. F. Phillips, in charge of bee culture.

A. F. Burgess, in charge of gipsy moth and brown-tail moth investigations.

Rolla P. Currie, in charge of editorial work.

Mabel Colcord, in charge of library.

Forest Insect Investigations.

A. D. Hopkins, Forest Entomologist in charge.

H. E. Burke (in charge of Pacific Slope Station at. Placerville. Cal.), Josef Brun-
ner (in charge of Northern Rocky Mountain Station at Missoula, Mont.), T. E.
Snyder, W. D. Edxionston (in charge of Southern Rocky Mountain Station at
Colorado Springs, Colo.), F. C. Craighead, J. M. Miller ;in charge of seed insect
station at Ashland, Oreg.), and A. B. Chaaiplain, assistants in forest entomology .

L. C. Griffith, assistant in shade tree insects.

S. A. Rohwer, specialist on forest Hymenoptera (in charge of Eastern Station at East-
Falls Church, Va.).

A. G. Boylng, specialist.

C. T. Greene, specialist on forest Diptera.

W. S. Fisher, specialist on forest Coleoptera.

Carl Heinrich, specialist on forest Lepidoptera.

Jacob Kotinsky, entomological assistaiit.

William Middleton. scientific assistant.

35 2

PREFACE.

Bulletin 94, entitled " Insects Injurious to Forests and Forest
Products/' consists of two parts and an index.

Part I, "Damage to Chestnut Telephone and Telegraph Poles by
Wood-Boring Insects," by Thomas E. Snyder, comprises the results of
a special study of a serious damage to the base of standing chestnut
telephone and telegraph poles by the wood-boring larva of a beetle
designated by the author as the chestnut telephone-pole borer
(Parandra brunnea Fab.).

Part II, " Biology of the Termites of the Eastern United States,
with Preventive and Remedial Measures," by Thomas E. Snyder,
is based mainly on investigations and experiments conducted during
the past three years by Mr. Snyder in connection with his work in the
Branch of Forest Insect Investigations. It also includes unpublished
notes by Messrs. H. G. Hubbard and F. L. Odenbach. Termites are
among the most destructive insects to both crude and finished forest
products in North America, among which may be listed construction
timbers in bridges and wharves, telephone and telegraph poles, hop
poles, mine props, fence posts, lumber piled on the ground, railroad
ties, and the woodwork of buildings. The sudden crumbling of
bridges and wharves, the caving in of mines, and the settling of floors
in buildings, are sometimes directly due to the concealed work of
these insects. The use of untreated wood-pulp products, sucl^ as the
various composition-board substitutes for lath, etc., is restricted in
the Tropics and southern United States because of the ravages of
termites. In the cities of Washington, Baltimore, St. Louis, Cleve-
land, New York, and Boston, and throughout the eastern and south-
ern United States, damage by termites to the woodwork of buildings
is occasionally serious.

Methods of prevention and control against injuries to finished and
utilized forest products, etc., are based on the results of experiments
conducted by this branch of the bureau.

A. D. Hopkins,

Forest Entomologist.

CONTENTS

I. Damage to chestnut telephone and telegraph poles by wood-boring in- Page.

sects Thomas E. Snyder. . 1

Object of paper 1

Historical data '1 1

The chestnut telephone-pole borer (Parandra brunnea Fab.) 3

Character of the insect 3

Distribution 4

Character of the injury 4

Importance of the problem 5

Extent of damage and loss 6

Favorable and unfavorable conditions for destructive work 6

Associated wood-boring insects 7

Prevention of the injury 8

Publications on wood preservation and statistics on poles 11

II. Biology of the termites of the eastern United States, with preventive and

remedial measures Thomas E. Snyder. . 13

Introduction ". 13

Classification 14

Historical 16

Biological experiments 20

The termitarium 20

Communal organization 22

Situation of the nests 22

Number of individuals in colonies 25

The different castes — polymorphism 27

The sense organs 31

The functions of the castes 32

The life cycle 33

The metamorphosis — caste differentiation 33

Progressive development of nymphs 36

Seasonal variations in the colony 43

Eggs 43

Nymphs of reproductive forms 43

Neoteinic reproductive forms 44

Workers 44

Soldiers 45

Location of the colony in winter 45

Duration of development and life 46

Cannibalism 46

Situation of the different forms in the nest 47

The swarm, or so-called nuptial flight 48

The establishment of the new colonies 49

Copulation and the rate of egg laying 50

VI INSECTS INJURIOUS TO FOREST PRODUCTS.

II. Biology of the termites of the eastern United States, with preventive and

remedial measures — Continued. Page.

The royal pair and other reproductive forms 53

Occurrence in the United States 53

Historical 54

Description of the reproductive forms 65

Dates of the swarming of Leucotermes 68

Association .with ants 70

Termitophilous insects 71

Parasites 72

Summary and conclusions based on the results of the experiments 72

The damage to forest products 75

Preventives, remedies, and "immune" woods 76

Method of obtaining photographs for the illustrations 82

Bibliography 83

Index 87

ILLUSTRATIONS.

PLATES.

Page.

Plate I. Fig. 1. — The chestnut telephone-pole borer (Parandra brunnea):

Male and female beetles. Fig. 2. — The chestnut telephone-pole

borer: Young larvae, dorsal and lateral views. Fig. 3. — Damage

to an untreated chestnut telegraph pole near surface of ground by

the chestnut telephone-pole borer 2

II. Work of the chestnut telephone-pole borer. Fig. 1. — Gallery of the
chestnut telephone-pole borer showing pupal chamber with the
entrance plugged with excelsior-like wood fibers; work near base
of pole below ground. Fig. 2. — Mines of the chestnut telephone-
pole borer near surface of ground 4

III. All castes of Leucotermes virginicus in heartwood of a maple tree

infested by Parandra brunnea 22

IV. Book destroyed by termites 22

V. Pine barn sill cut into ribbons by termites 22

VI. Living, stag-headed chestnut tree infested for entire length by ter-
mites; the insects entered the tree at the base through a lightning

scar and honeycombed the heartwood 22

VII. Views of sheds constructed by Leucotermes virginicus to cover up
galleries exposed on cross sections of logs sawn from an infested

maple tree 26

VIII. Lateral views of fully developed nymphs of the first and second
forms, and lateral and dorsal views of neoteinic kings of Leuco-
termes flavipes 30

IX. The evolution of the soldiers of Leucotermes virginicus and L. flavipes
from the large-headed worker-like larvae, showing the changes at

the molts 34

X. Quiescent stages during the final molt of nymphs of the first form of
Leucotermes flavipes and the active molted nymph with wings

unfolding, lateral and dorsal views 38

XI. Quiescent stages during the final molt of nymphs of the second form
of Leucotermes flavipes and the pigmented neoteinic king, lateral

and dorsal views „ . 38

XII. Fig. 1. — "Royal cells," in which true queens of Leucotermes flavipes
were found. Fig. 2. — Royal chamber excavated in solid chestnut
wood, in which 40 neoteinic reproductive forms of Leucotermes

flavipes were found 46

XIII. Comparison of true and neoteinic or supplementary queens of Leuco-
termes flavipes 58

XIV. Neoteinic reproductive forms of Leucotermes virginicus 58

XV. Dorsal, lateral, and ventral views of neoteinic nymphal or supple-
mentary queens of Leucotermes flavipes 62

XVI. Comparison of abdomens of young, fertilized, true queen and young

and old mature kings of the same type of Leucotermes flavipes 62

XVII. Fig. 1. — Telegraph pole, with base charred, mined by termites.

Fig. 2. — Pine flooring honeycombed by termites 74

VII

Vm INSECTS IXJURIOTIS TO FOREST PRODUCTS.

TEXT FIGURES.

Page.
Fig. 1. The chestnut telephone-pole borer (Parandra brunnea): Full-grown

larva 3

2. The chestnut telephone-pole borer: Female beetle, head and pro-

notum of male beetle 4

3. The chestnut telephone-pole borer: Pupa 5

4. View of a portion of the large number of treated experimental stakes

under test as to the relative effectiveness of various preventives
against termite attack 19

5. A'iews of unfinished and finished termitarium. showing structure and

interior with termite-infested wood ' 21

6. Wood of outer layers at base of insect-killed southern yellow pine

honeycombed by termites 24

7. Suspended tubes constructed by termites of earth and excreted wood. 26

8. Broken-off pine sapling from basal end of which tubes in figure 7

were suspended toward stump 27

9. Nymphs and soldiers of Leucotermes virginicus; young nymphs, and

grayish-black bands on abdomen of worker of flavipes 29

10. Molting larvae of Leucotermes flavipes in the quiescent stage 34

11. Molting nymph of the first form of Termopsis angusticollis in the

quiescent stage 39

12. View of a swarm of Leucotermes flavipes showing a portion of the enor-

mous number of winged, colonizing sexed adults that constitute a
swarm, emerging from an infested stump .48

13. Dorsal, ventral, and lateral views of a fertilized, mature, true queen

of Leucotermes flavipes, showing the position of the spiracles 67

14. Red-oak block impregnated with paraffin wax honeycombed by ter-

mites after a five months' test 78

ERRATUM

Page 10, footnote a, for Prior us read Brionus.

IX

U. S. D. A., B. E. Bui. 94, Part I. F. I. I., December 31, 1910.

INSECTS INJURIOUS TO FORESTS AND FOREST PRODUCTS.

DAMAGE TO CHESTNUT TELEPHONE AND TELEGRAPH
POLES BY WOOD-BORING INSECTS.

By Thomas E. Snyder, M. F.,
Agent and Expert.

OBJECT OF PAPER.

It has recently been determined through special investigations
conducted principally by the writer that serious damage is being
done to the bases of standing chestnut telephone and telegraph poles
in certain localities by the grub or larva of a wood-boring beetle,
here called the chestnut telephone-pole borer.a The character and
extent of the damage under different conditions of site in several
localities have been determined, and poles treated with various pre-
servative substances have been inspected to compare the efficiency
of both chemicals and methods of treatment. These investigations
have resulted in the determination of practical methods of preventing
injury to poles by wood-boring insects.

HISTORICAL DATA.

The first information of serious damage to standing chestnut poles
by wood-boring insects was conveyed in a letter, dated December
15, 1906, from E. O. Leighley, a correspondent of this Bureau,
reporting damage to telephone poles in Baltimore, Md., by borers.
Mr. A. B. Gahan, assistant entomologist of the Maryland Agricul-
tural Experiment Station, College Park, Md., who investigated the
injury to the poles, stated that it was the work of a borer and was
located just beneath the surface of the ground. Mr. Gahan brought
specimens of the work and the insect to this office. The borers were
identified as cerambycid larvae, and later were determined to be the
chestnut telephone-pole borer (Parandra brunnea Fab.).

On December 16, 1906, Mr. H. E..HoDkins, division superintendent
of a telephone company, stated that the poles in West Virginia were

"Parandra brunnea Fab.; Order Coleoptera, Family Spondylidee,

64271°— Bull. 94, pt. 1— 10

Z INSECTS INJURIOUS TO FORESTS.

badly injured by borers and that these borers were abundant. On
March 8, 1907, he collected larvae from chestnut telephone poles at
Pennsboro, W. Va. These were determined to be the larvae of the
chestnut telephone-pole borer.

The writer on October 3, 1909, inspected some chestnut telegraph
poles which had been standing for about twelve years on New York
avenue, in Washington, D. C. The poles had been taken down under
orders from the city authorities, which necessitated the placing of
wires in conduits under ground, and they had been lying in piles for
about a month before they were inspected. The chestnut telephone-
pole borer had been working in the base of the poles, and white ants,
or termites, were associated with them. Twelve out of the 103 poles
examined had been damaged, some more seriously than others.

On October 15, 1909, Mr. H. E. Hopkins sent a reply to a request
by Dr. A. D. Hopkins for further information regarding insect dam-
age to poles in West Virginia. He stated that in one line built twelve
years ago (40 miles long, 36 chestnut poles to the mile, poles 20 to
40 feet long and 5 to 12 inches in diameter at the top) approximately
600 poles had been rotted off at the. top of the ground, and inspection
showed that 95 per cent of the damage was directly or indirectly
due to insects. Other lines in this division were reported to be in
about the same condition. It was later determined that most of the
insect damage was the work of the chestnut telephone-pole borer.

Dr. A. D. Hopkins states in a recent comprehensive bulletin a that
" construction timbers in bridges and like structures, railroad ties,
telephone and telegraph poles, mine props, fence posts, etc., are
sometimes seriously injured by wood-boring larvae, termites, black
ants, carpenter bees, and powder-post beetles, and sometimes reduced
in efficiency from 10 to 100 per cent." Thus, while it has been known
that almost all classes of forest products that are set in the ground
are seriously injured by wood-boring insects, the problem of insect
damage to standing poles, posts, and other timbers has never been
made the subject of a special investigation.

In May, 1910, this study was assigned to the writer, and, in addi-
tion to a study of the insects involved, investigations in cooperation
with telephone and telegraph companies have been conducted in the
District of Columbia, Maryland, Virginia, Pennsylvania, New Jersey,
and New York. Through the courtesy of the Western Union Tele-
graph Company several telegraph lines were inspected in July and
August, 1910, in Virginia, where the poles were being reset or replaced.
Here the butts of over 200 poles set under different conditions of
site were thoroughly examined for insect damage, and sometimes the

a Insect Depredations in North American Forests. <Bul. 58, Part V, Bureau of
Entomology, U. S. Department of Agriculture, p. 67, 1909.

Bui. 94, Part I, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate I.

Fig. 1.— The Chestnut Telephone-pole Borer (Parandra brun-
nea): Male and Female Beetles. Fig. 2.— The Chestnut
Telephone-pole Borer: Young Larv/e, Dorsal and Lateral
Views. Fig. 1, Slightly Enlarged; Fig. 2, Twice Natural
Size. (Original.)

Fig. 3— Damage to an Untreated Chestnut Telegraph Pole
near Surface of Ground by the Chestnut Telephone-pole
Borer. (Original.)

DAMAGE TO CHESTNUT POLES BY INSECTS. 6

entire pole was split open. In one line 10 to 12 years old (approxi-
mately 30 chestnut poles per mile, 25 feet long, about 6 inches diam-
eter at the top, 10 inches at the base, and apparently of second quality) ,
between Petersburg and Crewe, Va. — the poles had already been reset
once, east of Wilson, Va. — serious damage by the chestnut telephone-
pole borer rendered from 15 to 20 per cent of the poles unserviceable.
After the present second resetting it was esti-
mated that the poles can not last more than
four or five years longer. West of Wilson the
poles were naturally in much worse condition,
and many were broken off and only held up
by the wires on the sounder poles. In another
line examined, between Portsmouth and Boy-
kins, Va. (poles 30 feet long and apparently of
second quality) , serious damage by this borer
averaged about 10 or 15 per cent, and between
Boykins, Va., and Weldon, N. C, according to
a linesman, 50 per cent of the poles are badly
decayed near the surface of the ground.
Much of this damage, however, is due to fun-
gous heart rot. According to a statement
by the foreman of a resetting crew, between
Asheville, N. C, and Spartanburg, S. C, hundreds of chestnut poles
were badly decayed in the 67 miles of line reset, and were only held
up by the wires. The line was 15 years old. There was serious
damage by " wood lice" (termites) and also by "white wood worms.' '

THE CHESTNUT TELEPHONE-POLE BORER.

(Parandra brunnea Fab.)
CHARACTER OF THE INSECT.

Fig. 1. — The chestnut telephone-
pole borer (Parandrabrunneay.
Full-grown larva. (About
twice natural size. (Original.)

The chestnut telephone-pole borer is a creamy white, elongate,
stout, cylindrical, so-called " round-headed" grub or "wood worm"
(fig. 1), which hatches from an egg deposited by an elongate, flattened,
shiny, mahogany brown, winged beetle from two-fifths to four-fifths of
an inch in length. (Plate I, fig. 1 ; text fig. 2.) The eggs are probably
deposited from August to October in shallow natural depressions or
crevices on the exterior of the pole near the surface of the ground;
often the young larvae enter the heartwood through knots. The
young borers (Plate I, fig. 2) hatching therefrom eat out broad
shallow galleries running longitudinally in the sapwood, then enter
the heartwood, the mines being gradually enlarged as the larvae
develop. As they proceed, the larvae closely pack the fine excreted
boring dust behind them. This debris, which is characteristic of

4 INSECTS INJURIOUS TO FORESTS.

their work, is reddish to dunnish yellow in color and has a claylike
consistency. The mines eventually end in a broad chamber, the
entrance to which is plugged up by the excelsior-like fibers of wood
chiseled out by the strong mandibles of the larva. Here the resting
stage (fig. 3), or pupa, is formed, and in this chamber the perfect adult
spends considerable time before emerging. Often all stages from very
young larvae only about one-fourth inch long to full-grown larvae over

1 inch long, pupae, and
adults in all stages to
maturity are present in
the same pole. Adults
have been found flying
from July to Septem-
ber. As yet the sea-
sonal history of this
borer has not been com-
pletely worked out.

DISTRIBUTION.

Fig. 2.— The chestnut telephone-pole borer: Female beetle, three tniS insect IS Very

and one-half times natural size. Head and pronotum of male widely distributed
beetle. (Original). • t> r\ j. '

ranging from Ontario,
Canada, to Texas, eastward to the Atlantic coast, and westward to
Arizona and southern California. It is common throughout the
natural range of the chestnut — and in this connection it should be
observed that most of the chestnut poles are purchased from local
timber-land owners.

CHARACTER OF THE INJURY.

The injury to the poles consists in large mines in the wood near the
line of contact of the pole with the ground, necessitating the frequent
resetting or even the replacement of the damaged poles. These
irregular galleries of the grub (Plate II, fig. 1) run both horizontally
and longitudinally throughout the heartwood, and are sometimes 7
inches long, but vary with the individuals, which show great differ-
ences in size. The borers usually work in the outer layers of the
wood at the base of the pole for a distance of from 2 to 3 feet below,
and sometimes from 1 to 2 feet above the line of contact of the pole
with the surface of the ground. The greatest damage is to that area
just below and just above the surface of the ground (Plate I, fig. 3);
here the conditions of air and moisture are most favorable. Often
the entire butt up to a distance of from 4 to 6 feet and higher,
according to the depth of setting, is mined. The numerous galleries,
often very close together, completely honeycomb the wood in a zone

Bui. 94, Part I, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate II.

Fig. 1. Fig. 2.

Work of the Chestnut Telephone-pole Borer.

Fig. 1.— Gallery of the chestnut telephone-pole borer, showing pupal chamber with the en-
trance plugged with excelsior-like wood fibers; work near base of pole, below ground.
Fig. 2.— Mines of the chestnut telephone-pole borer near surface of ground. Natural size.
(Original.)

DAMAGE TO CHESTNUT POLES BY INSECTS.

3 to 4 inches in from the exterior of the poles; this so weakens the
poles that they break off close to the surface of the ground. The
basal 2 feet is usually sound. Even if the damage is not serious
enough to cause the poles to break off under strain, they are likely to
go down during any storm, and thus put the wire service out of com-
mission; such damaged poles are a serious menace along the right of
way of railroads. The beetle will attack poles that are perfectly
sound, but evidently prefers to work where the wood shows signs of
incipient decay; it will not work in wood that is "sobby" (wet rot),
or in very "doty" (punky) wood. It has not yet been determined
just how soon the borers usually enter the poles after they have been
set in the ground. However, poles that had been standing only four
or five years contained larva? and adults of this borer
in the heartwood, and poles that had been set in the
ground for only two years contained young larvae in the
outer layers of the wood.

Poles that appear sound on the exterior may have the
entire basal interior riddled, and the work of the borers
is not noticed until the poles break off. If merely iso-
lated poles are injured, the poles that are broken off are
held up by the wires and can be detected by the fact
that they lean over, but if several adjacent poles are
affected, especially where there is any unusual strain,
that portion of the line is very likely to go down. The
presence of the borers in injurious numbers can be de-
termined only by removing the earth from about the
base of the pole; the exit holes of the borer are found
near the line of contact with the soil. Often large,
coarse borings of wood fiber project from the exit holes.
Sometimes old dead parent adults are found on the
exterior of the poles underground. During August the
young adults may be found in shallow depressions on the exterior of
poles below the ground surface.

Fig. 3. — The
chestnut tele-
phone-pole bor-
e r : Pupa.
Slightly more
than twice
natural size.
(Original.)

IMPORTANCE OF THE PROBLEM.

The subject of the relation of insects to the rapid decay of chestnut
poles has not been thoroughly investigated in the past, but now that
the supply is becoming scarcer it is especially important to know
what are the various primary causes of the deterioration of these
poles, hitherto described under the vague term " decay." Although
the chestnut telephone-pole borer has not hitherto been considered an
insect of any economic importance, and has been described in ento-
mological literature as only living under bark, principally of pine, or

6 INSECTS INJURIOUS TO FORESTS.

in the decomposing wood of various species of deciduous and conif-
erous trees, the evidence is abundant that breeding in the bases of
chestnut poles is not a newly acquired habit. It has also been
determined that this beetle damages many species of living forest,
fruit, and shade trees that have been previously injured by fire or
other causes, and often leads to the destruction of trees that would
otherwise recover from such wounds, and while not normally a
primary enemy to trees, may thus become of more than secondary
importance.

The damage by the chestnut telephone-pole borer is especially
serious in consideration of the fact that in many parts of its range the
chestnut is threatened with extinction as a tree species on account of
the very severe ravages of the combined attack of an insect a and a
fungous disease. Further unnecessary drain upon the supply of
chestnut timber should be avoided by protecting that already in use
and thus prolonging its length of service.

EXTENT OF DAMAGE AND LOSS.

As more than one-fourth of the 3,500,000 round poles exceeding 20
feet in length used annually by telephone, telegraph, and other
electric companies are chestnut (Kellogg, 1909),6 and as this borer
has seriously damaged as high as 10 to 40 per cent, varying with
conditions of site, of the chestnut poles which have been set in the
ground for from ten to twelve years in lines in North Carolina, Virginia,
West Virginia, Maryland, and the District of Columbia, it is evident
that this insect is an important factor in decreasing the normal
length of service of the poles. c In lines from twelve to fifteen years
old the damage is much greater, and at the end of this number of
years of service any line in which poles of this species are set has to
be practically renewed. According to a statement in Forest Service
Bulletin 78 (Sherfesee, 1909), "approximately 4 per cent, or 5,908
feet board measure of the 147,720 feet board measure of standing
poles annually requiring replacement in the United States, is destroyed
by insects." If only chestnut poles be considered, at least 10 per
cent of the poles reset or replaced are injured by insects.

FAVORABLE AND UNFAVORABLE CONDITIONS FOR DESTRUCTIVE WORK.

The damage is apparently greatest and the borers are most abun-
dant where the poles are set in high or level dry ground under good
conditions of drainage. Such sites are the crests of railroad cuts
through low hills, slopes of " fills," and in cultivated or other fields.
Where the poles are in wet sites there is usually but little injury by

a Agrilus bilineatus Web.

& See list of publications, page 11.

c The average life of a chestnut pole is eight to ten years (Sherfesee and Weiss, 1909).

DAMAGE TO CHESTNUT POLES BY INSECTS. 7

woocl-boring insects except to that portion near the surface of the
ground. Conditions of drainage are more important than different
soil combinations, and the condition of the soil is more important
than its composition; i. e., where the soil is hard packed there is
apparently less damage than where it is loose. The quality and con-
dition of the poles before setting is a very important factor to con-
sider before arriving at any conclusions as to the relative longevity of
poles under various conditions of site. Green (unseasoned) or im-
perfectly seasoned poles are less durable than those thoroughly
seasoned. Poles that are defective a before setting, as they very
often are (i. e., showing evidence of incipient decay) , and poles that
have the heartwood mined by the chestnut timber worm, b the work
of which is very abundant, will, of course, decay much more
rapidly than poles that are in an absolutely sound condition.
The galleries of the chestnut timber worm afford an entrance to
the spores of wood-destroying fungi, and thus greatly accelerate
decay. White mycelium compactly filled these galleries throughout
many standing poles, thus clearly proving that these mines aid
greatly in enabling the fungous heart rot more rapidly and
completely to penetrate the entire heartwood of the poles. If the
injury by both wood-boring beetles and wood-destroying fungi
(between which there is a varying interrelation) be considered, then
in several lines from ten to twelve years old in North Carolina,
Virginia, and West Virginia at least 50 per cent of the poles are either
rendered unserviceable or their length of service is much shortened.

ASSOCIATED WOOD-BORING INSECTS.

It is not to be concluded that this wood-boring beetle is the only
insect that injures standing chestnut poles. Indeed, the most
common injury is by the "wood lice" or white ants.c In lines from
ten to twelve 3? ears old these insects have seriously damaged as high
as 15 per cent of the poles, and their work is often present, at least
superficially, in as high as 75 per cent of the poles under all conditions
of site. However, the damage is usually to the outer layers of the
wood, where it is moist or there is incipient decay, and is more
superficial and localized than that of the chestnut telephone-pole
borer. Nevertheless, white ants often completely honeycomb the
sound heartwood of poles, especially at the base. They work both
in sound wood, "doty" (dry rot) wood, and "sobby" (wet rot) wood.
Sometimes a large channel runs up through the core of the heart

a Often this evidence is the old galleries of the destructive two-lined chestnut
borer (Agrilus bilineatus Web.), showing that the tree must have been dead before
it was cut for a pole, and hence is more likely to be defective throughout the interior*,
in other instances heart rot is clearly present.

b Lymexylon sericeum Harr.

c Identified by Mr. Theodore Pergande of this Bureau as Termes flavipes Kollar.

8 INSECTS INJURIOUS TO FORESTS.

and the sides are plastered with clay, forming a hollow tube with
several longitudinal galleries. Their work often extends from 2 to 4
feet above the surface of the ground. They leave the outer shell
of the wood intact and work up through the longitudinal weathering
checks, covering the exterior of the pole with earth to exclude the
light. White ants will damage poles that have been set in the ground
only two years. Evidently they enter the pole from below the surface
of the ground. The habits and characteristics of these peculiar and
interesting insects have been thoroughly discussed in Circular No. 50
of this Bureau by Mr. C. L. Marlatt.

A giant round-headed borer a is sometimes found in the poles,
usually in association with the chestnut telephone-pole borer. In
poles where the wood is sound this borer apparently works as a rule
only in the outer layers of the wood, the galleries running longitudi-
nally through the heart below the surface of the ground. In poles
where there is decay it will completely honeycomb the he art wood
near the surface of the ground.

In several poles where the wood was "doty" a large Scarabseid b
which has before been found in decayed oak railroad ties was present
and caused the poles to break off sooner than they otherwise would.
The irregular galleries of the grub completely honeycomb the decayed
heartwood near or just below the surface.

A flat-headed borer c and wireworms d were found in galleries locally
in the more or less decayed heartwood of several poles. A large black
carpenter ante does some damage to sound poles set in dry ground
through woodland. This ant often widens the longitudinal weather-
ing checks and thus accelerates decay. A small black ant7 was very
numerous in many poles, but its work is usually confined to the
outer layers of the wood. The work is often throughout udoty"
poles. Injury by this ant is not primary, but it also widens weather-
ing checks, enlarges other defects, and induces more rapid decay.

PREVENTION OF THE INJURY.

Doctor Hopkins makes the following statement in a recent bulletin :0

Insect damage to poles, posts, and similar products can be prevented to a greater
or less extent by the preservative treatments which have been tested and recom-
mended by the Forest Service for the prevention of decay. These should be applied

a Prionus sp.

& Identified by Mr. E. A. Schwarz, of this Bureau, as Polymrechus brevipes Lee.

c Identified by Mr. H. E. Burke, of this Bureau, as Buprestis rufipes Oliv.

d Species of the family Elateridse. The large larvae of Alaus sp. were especially
injurious.

* Identified by Mr. Theodore Pergande as Camponotus pennsylvanicus Mayr.

/ Identified by Mr. Theodore Pergande as Cremastogaster lineolata Say.

0 Insect Depredations in North American Forests. <Bul. 58, Part V, Bur. Ent.,
XL S. Dept, Agr., p. 84, 1909.

DAMAGE TO CHESTNUT POLES BY INSECTS. 9

before the material is utilized for the purposes intended, or, if it be attacked after
it has been utilized, further damage can be checked to a certain extent by the use of
the same substances.

It is often of prime importance to prevent injury from wood-boring insects, for the
reason that such injuries contribute to more rapid decay. Therefore anything that
will prevent insect injury, either before or after the utilization of such products, will
contribute to the prevention of premature deterioration and decay.

Through the courtesy of the American Telephone and Telegraph
Company and the Forest Service, about 40 chestnut poles set in a
test line near Dover, N. J., were inspected by the writer on July 15,
1910, in company with engineers of the telephone company and Mr.
H. F. Weiss, Assistant Director, Forest Products Laboratory, Forest
Service, to determine the relative merits of various methods of pre-
venting damage by wood-boring insects to the bases of poles. In
this line, which is eight years old, variously treated poles alternated
with untreated poles in order that each chemical preservative and
method of treatment might be given an absolutely fair test under
the same conditions of site. The poles were 30 feet long, 7 inches
in diameter at the top, and 33 inches in circumference 6 feet from
the base. In this inspection the earth was removed (to a depth of
about 1 foot) from the base of the pole, and then the pole was chopped
into to determine the rate of decay. This method of inspection for
insect damage is not very satisfactory. The various methods experi-
mented with in this test line were brush treatments with a patented
carbolineum preservative and spirittine, charring the butt, setting
the pole in sand, and setting it in small broken stone. It was found
that, although these methods may temporarily check the inroads of
wood-boring insects, they will not keep the insects out of the poles.
The most serious damage to the poles in this line was by white ants.
Other insect damage was by a large black carpenter anta and the
larvae of a round-headed borer. b

An inspection was made, between September 6 and 14, 1910, of
the bases of over 400 chestnut poles set in a similar test line near
Warren, Pa., and Falconer, N. Y. These poles were treated by the
creosote " open-tank" method of impregnation, and brush treatments
of creosote, wood creosote, creolin, two different carbolineum pre-
servatives, and tar; they had been set in the ground for a period of
five years. All these treatments, except the brush treatments with
creolin and tar, were efficient in preventing the attacks of wood-
boring insects, at least for a five-year period, in this northern climate.
There was but little damage by insects to the poles in this test line.
The most common injury to the untreated poles was by the large
black carpenter ants which widen the longitudinal weathering checks,
and hence induce more rapid decay. The work of the chestnut tele-

a Camponotus pennsylvanicus Mayr.
b Prionus sp.

10 INSECTS INJURIOUS TO FORESTS.

phone-pole borer was found in several poles, and this beetle was evi-
dently just beginning to attack these poles. There was some damage
by a round-headed borer. a No white ants or termites were present,
and this is evidently too far north for these destructive borers. A
report by inspectors of the American Telephone and Telegraph Com-
pany and the Forest Service on the remainder of the poles in this
test line (between Jamestown and Buffalo, N. Y.) not personally
inspected by the writer, showed that these conclusions can be applied
to all the poles in the line with the exception that there was super-
ficial injury by small black ants to two poles treated by brush treat-
ments of carbolineum avenarius and to two treated with wood creo-
sote; also, as the inspection progressed, injury by the chestnut tele-
phone-pole borer became more abundant and serious, and the borers
seemed to be established in the poles. The poles treated by the creo-
sote " open-tank" method of impregnation and by brush treatments
with creosote and with l'S. P. F." carbolineum remained uninjured.

Methods of treating poles superficially by brushing with various
preservatives have proved to be temporarily efficient in keeping
wood-boring insects out, if the work is thoroughly done and not only
the butt, but also the base, is treated. If the pole is not thoroughly
brushed, insects enter through the untreated or imperfectly treated
portions, especially through weathering checks and knots. Where
the base is left untreated, insects, especially white ants or termites,
enter the pole from below ground and, avoiding the treated portions,
come right up through the pole.

The few poles of southern yellow pine in a line near Bartley, N. J.,
inspected on July 15, 1910, which had been impregnated with creo-
sote by the Bethell cylinder-pressure process, 12 pounds of oil to the
cubic foot, and had been set in the ground since February, 1903,
were apparently absolutely free from signs of decay or damage by
wood-boring insects. In another line, running between Norfolk, Va.,
and Washington, D. C, the few poles (12 years old, of squared —
with the sapwood cut away — southern yellow pine) inspected on
August 10, 1910, near Portsmouth, Va., which had been impregnated
with creosote by the Bethell cylinder-pressure process, were also
apparently absolutely sound.

Thus, it is evident that impregnating the poles with creosote by
some standard process (either the open-tank or the cylinder-pressure
processes) will keep wood-boring insects out and preserve the poles
for a much longer period than they would last untreated. In the
open-tank method only the area most subject to the attacks of wood-
boring insects and deterioration in general (i. e., the basal 8 feet) is
treated, while by the cylinder-pressure processes the entire pole is
impregnated. Alternating less susceptible juniper (red cedar)6 poles

aPriorus sp. & Juniperus virginiana.

DAMAGE TO CHESTNUT POLES BY INSECTS. H

or pine poles thoroughly impregnated by some standard process in
the line with the chestnut poles would be a safeguard in holding up
an old line where the damage is found to be serious on resetting.

A list of some available publications on wood preservation is
appended.

12 INSECTS INJURIOUS TO FOEESTS.

PUBLICATIONS ON WOOD PRESERVATION AND STATISTICS ON

POLES.

1903. Von Schrenk, H. — Seasoning of timber. <Bul. 41, Forest Service, U. S.
Dept. Agr.

1906. Grinnell, H. — Prolonging the life of telephone poles. <Yearbook U. S.

Dept. Agr. for 1905, Extract No. 395.

1907. Crawford, C. G. — The open-tank method for the treatment of timber.

<Cir. 101, Forest Service, U. S. Dept. Agr.
1907. Crawford, C. G. — Brush and tank pole treatments. <Cir. 104, Forest Serv-
ice, U. S. Dept. Agr.

1907. Grinnell, H. — Seasoning of telephone and telegraph poles. <Cir. 103, Forest

Service, U. S. Dept. Agr.

1908. Sherfesee, W. F.— A primer of wood preservation. <Cir. 139, Forest Service,

U. S. Dept, Agr.

1908. Weiss, H. F. — Progress in chestnut pole preservation. <Cir. 147, Forest

Service, U. S. Dept, Agr.

1909. Sherfesee, W. F. — Wood preservation in the United States. <Bul. 78, Forest

Service, U. S. Dept. Agr., pp. 24, 25, Table I.
1909. Sherfesee and Weiss. — Wood preservation. <Rep. Natl. Conserv. Com.,
vol. 2, p. 663.

1909. Kellogg, R. S.— The timber supply of the United States. <Cir. 166, Forest

Service, U. S. Dept. Agr., pp. 20-21.

1910. Willis, C. P. — The preservative treatment of farm timbers. < Farmers' Bui.

387, U. S. Dept, Agr.

O

U. S. D. A., B. E. Bui. 94, Part II. F. 1. 1., February 17, 191.*

INSECTS INJURIOUS TO FORESTS AND FOREST PRODUCTS.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED
STATES, WITH PREVENTIVE AND REMEDIAL MEAS-
URES.

By Thomas E. Snyder, M. F.,

Entomological Assistant.
INTRODUCTION.

The following notes on the biology of the common termites, a or
" white ants/' of the eastern United States were, for the most part,
made while conducting investigations to determine the character
and extent of damage by termites and other forest insects to various
classes of crude and finished forest products and to devise methods
of preventing the injury. During 1910 and 1911 special investiga-
tions were conducted by the writer as to the character and extent
of damage to telephone and telegraph poles and mine props by wood-
boring insects. This contribution is thus based, largely, on these
investigations, as well as on additional experiments conducted dur-
ing the past three years by the writer in the branch of Forest Insect
Investigations and on some of the unpublished notes of the late
H. G. Hubbard and those of Rev. F. L. Odenbach.

" White ants" are among the most destructive insects of North
America to both crude and finished forest products, among which may
be listed construction timbers in bridges, wells, and silos, timbers
of wharves, telephone and telegraph poles, bean and hop poles, mine
props, wooden cable conduits, fence posts, lumber piled on the
ground, railroad ties, and the foundations and woodwork of buildings,
etc. The sudden crumbling of bridges and wharves, caving in of
mines, and settling of floors in buildings are sometimes directly due
to the hidden borings of termites. The use of untreated wood-pulp
products such as the various " composition boards" used as sub-
stitutes for lath, etc., is restricted in the Tropics and portions of the
southern United States because of the ravages of termites. In the
cities of Washington, Baltimore, St. Louis, Cleveland, and New
York, and throughout the Southern States damage by termites to

« Order Platyptera, Packard (1886), suborder Isoptera, Brulle (1832), family Meso-
termitidx, Holmgren, genus Leucotermes Silvestri.

13

14 INSECTS INJURIOUS TO FOEESTS.

the woodwork of buildings is occasionally serious. As far north as
Boston, Mass., damage of this sort occurs, and in Michigan cases are
reported in which furniture in buildings has sunk through the floors,
mined by " white ants."

The preventives and remedies against injury to forest products
and nursery stock herein given are based on the results of experi-
ments conducted by the Bureau of Entomology. The species to be
considered in this paper are Leucotermes jlavipes Kollar and Leuco-
termes virginicus Banks. The former is widely distributed a over
the United States, but the recorded distribution of the latter is more
Hmited.6

CLASSIFICATION.

Termites are naturally to be classed among that most interesting
group of social insects comprising the ants, bees, wasps, etc., since
they live in colonies which are made up of various highly specialized
forms or castes. Each of these forms has a distinct role in the proc-
esses of the social organization, as there is a well-defined division of
labor. In the systematic classification of insects, however, termites
are widely separated from the other social insects. These latter rep-
resent the highest and most specialized development, whereas termites
represent the lowest and are among the oldest of insects. Further-
more, in many points in their life habits termites are widely different
from the other social insects.

The tropical genus Termes, from which the family and generic
names of termites are derived, is a Linnaean creation c and appears
for the first time in 1758, in the tenth edition of Systema Naturae,
where it was placed among the Apt era, between the genera Podura
and Pediculus. Since then termites have been classed d among the
orders Neuroptera, Corrodentia, and Pseudoneuroptera, although
Brulle, in 1832 (Exped. Sc, Moree, t. 3 (ZooL), p. 66, Paris, 1832), had
founded the distinct new order Isoptera.e Packard/ in 1863, stated
that ' ' seven out of the eight well-established families of the Neuroptera
sustain a synthetic relation with each of the six other suborders.7'
Hagen7^in 1868, stated in regard to the error that he made in describ-

a Marlatt, C. L. The White Ant (Termes Jlavipes Koll.). U. S. Dept. Agr.. Bur.
Ent., Circ. 50, pp. 8, figs. 4, June 30, 1902. See p. 4.

b Banks, N. A new species of Termes. Ent. News., v. 18, no. 9, p. 392-393, No-
vember, 1907.

c Desneux, J. Isoptera, Fam. Termiticke, pp. 52, figs. 10, pis. 2. (Wytsman, P.,
Genera Insectorum, fasc. 25, Bruxelles, 1904, p. 1-3.)

d Feytaud, J. Contribution a Fetude du termite lueifuge. Arch. Anat. Micros.,
t. 13, fasc. 4, p. 481-606, figs. 34, 30 juin, 1912.

« Desneux, J. Loc. cit.

/ Packard, A. S. On synthetic types in insects. Boston Jour. Nat. Hist., v. 7,
no. 4, p. 590-603, fig. 4, June, 1863.

<?Hagen, H. A. Proc. Boston Soc. Nat. Hist., v. 12, p. 139, October 21, 1868,
Boston, 1869.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 15

ing a a damaged earwig (Forficula) from Japan as a wingless termite,
"that the three families, Termitina, Blattina, and Forficulina, are
coordinated and very nearly allied." Packard,6 in 1883, placed
termites in a new order, the Platyptera, including the Termitidse,
Embiidse, Psocidse, and Perlidse, and three years later he dismem-
bered c the Pseudoneuroptera into the Platyptera, Odonata, and
Plecoptera. Knower/ in 1896, stated that in the development of
the embryo of termites there is a resemblance to that of the Orthop-
tera. Desneux, in 1904/ stated that termites are derived phylo-
genetically from the "Blattides," an idea accredited to Handlirsch
in 1903, although Hagen, in 1855, had already formulated this
theory based on purely biological considerations. Comstock/ in
1895, also placed termites in the order Isoptera. Enderlein,^ in
1903, placed termites with the Embiidse in the order Corrodentia,
suborder Isoptera. In this order he also placed the Psocidse and
Mallophaga. Handlirsch h contests the affinity of the Termitidse with
the Embiidse; he further states* that termites are derived from all
deposits from the lower Tertiary on, but that all older fossils formerly
mistaken for termites by Hagen, Scudder, and Heer do not belong
to this order. Banks J (1909) considered termites to be in the
order Platyptera, suborder Isoptera, with two other suborders, the
Mallophaga and Corrodentia. Holmgren states^ that he believes
both groups, the "Termiten" and "Blattiden," are offshoots of a
more primitive group, the "Protoblattoiden." The oldest "Blat-
toiden" occur in the first part of the middle, upper Carboniferous
(Pottsville, North America), the oldest " Protoblattoiden," in the last

0 Hagen, H. A. On a wingless white ant from Japan. Proc. Boston Soc. Nat.
Hist., v. 11, p. 399-400, ilhis., February 26, 1868.

& Packard, A. S. Order 3, Pseudoneuroptera, IT. S. Dept. Agr., U. S. Ent. Com.,
3d Rept., p. 290-293, 1883.

c Packard, A. S. A new arrangement of the orders of insects. Amer. Nat., v. 20,
no. 9, p. 808, September, 1886.

d Knower, H. McE. The development of a termite — Eutermes (Rippertiif) A pre-
liminary abstract. Johns Hopkins Univ. Circ, v. 15, no. 126, p. 86-87, June, 1896.

e Desneux, J. Loc. cit.

/ Comstock, J. H. Manual for the Study of Insects. Ithaca, N. Y., 1895, p. 95-97,
fig. 104-106.

9 Enderlein, G. Uber die Morphologie, Gruppierung und Systematische Stellung
der Corrodentien. Zool. Anzeig., vol. 26, p. 423-437, figs. 4.

^Handlirsch, A. Zur Systematik der Hexapoden. Zool. Anzeig., vol. 27 (1904),
p. 733-769.

i Handlirsch, A. Die fossilen Insekten und die Phylogenie der rezenten Formen,
pt. 8, p. 1240, Leipzig, 1908.

3 Banks, N. Directions for Collecting and Preserving Insects, pp. 135, figs. 188,
Washington, 1909. U. S. Nat. Mus. Bui. 67. Platyptera, p. 6-7.

k Holmgren, N. Termitenstudien 1. Analomische Untersuchungen. K. Svenska
Vetensk. Akad. Handl., Bd. 44, No. 3, pp. 215, Taf. 1-3, Uppsala & Stockholm, 1909.
Die Verwandtschaftsbeziehungen der Termiten, p. 208-213.

16 INSECTS IXJUKIOUS TO FORESTS.

part of the coal measures (Allegheny, North America), but the
youngest "Protoblattoiden" occur in the lower Permian formation
of Europe. Holmgren considers the termites to be in a distinct
order, the Isoptera.

The order Isoptera, according to Holmgren (1911),a is divided into
three families, the Protermitidse, the Mesotermitidse, and the Meta-
termitidse. All of these families are represented in North America,
and while this paper is restricted to a discussion of species of the
genus Leucotermes Silvestri, subfamily Leucotermitina? Holmgren,
family Mesotermitidse Holmgren, a species in the genus Termopsis
Heer, subfamily Termopsinee Holmgren, family Protermitidas Holm-
gren, is briefly mentioned. Thus it will be seen that the species
under observation occupy a middle position between the highest
and the lowest genera in the systematic classification of termites.

HISTORICAL.

According to Desneux, Smeathman's marvelous descriptions in
Some Account of the Termites Which Are Found in Africa and Other
Hot Climates (London, 1781) are the real basis of scientific re-
searches on the biology of termites.5 Hagen c gives a resume of the
researches on termites up to 1860. Drummond's Tropical Africa
(London, 1889), in chapter 6, gives an interesting account of " white
ants." Froggatt, in Australian Ternritidae/ gives a resume of the
studies of various workers on termites. Saville-Kent, in Chapter
III of the Naturalist in Australia (London, 1897), describes the
habits of termites in the Tropics; Smeathman, Drummond, and
Saville-Kent in their popular accounts of termites stimulated interest
in the habits of these insects and led to scientific researches. Grassi
and Sandias,6 in their classical work, have outlined the results of
the more important biological researches. Sharp / gives an excellent

« Holmgren, N. Termitenstudien 2. Systematik der Termiten. K. Svenska
Vetensk. Akad. Handl.. Bd. 46, No.6, pp. 86, Taf. 1-6, Uppsala & Stockholm, 1911.
Ordnung Isoptera, p. 10-11.

& Smeathman, H. Some account of the termites, which are found in Africa and
other hot climates. In Philos. Trans. London, v. 71. p. 139-192, 3 pis., 1781. (Smeath-
man's observations were afterwards confirmed by Savage in 1850 and the late G. D.
Haviland.)

c Hagen, H. A. Monographie der termiten. Linnaea Entomologica, v. 10, 1855,
p. 1-144, 270-325; v. 12, 1858, p. 1-342, pi. 3; v. 14, 1860. p. 73-128.

d Froggatt, W. W. Australian Termitidae, Part I. Proc. Linn. Soc. N. S. Wales,
v. 10, ser. 2, p. 415-438, July 31, 1895. Distribution, p. 416^26.

e Grassi, B., and Sandias, A. The constitution and development of the society of
termites; observations on their habits; with appendices on the parasitic protozoa of
Termitidse and on the Embiidse translated by F. H. Blandford. Quart. Jour.
Micros. Sci. [London], v. 39, pt, 3, n. s., p. 245-322, fold. pi. 16-20, November, 1896,
and v. 40, pt, 1, p. 1-75, April, 1897.

/Sharp, D. Cambridge Nat. Hist,, vol. 5, Insects, Pt. 1. chap. 16. p. 356-390, Lon-
don, 1901.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 17

summary of the various writings on termites up to 1901. Escherich a
gives a resume of the more important work up to 1909, and also an ex-
tensive bibliography. Holmgren's treatise 6 deals with anatomy, sys-
tematic classification, development, and biology, and gives a summary
of the effect of the social life, as shown in the progressive and retro-
gressive development of termites. Bugnion c> d has recently made
observations that lead him to state that (in Eutermes lacustris
Bugnion and Termes spp.) the differentiation of the three castes
occurs in the embryo. Feytaud e has published an account of the
life history of Leucotermes lucifugus Rossi, a species closely related
to our common species of the eastern United States. His observa-
tions confirm those of Heath on the same species in this country.
He also made studies of the internal anatomy. The more important
contributions to our knowledge of termites are the results of re-
searches by Bobe-Moreau, Bugnion, Czervinski, Desneux, Doflein,
Drummond, Escherich, Feytaud, Froggatt, Grassi and Sandias,
Hagen, Haldemann, Haviland, Holmgren, Jehring, Latreille, Leidy,
Lespes, Fritz Mtiller, Perez, Perris, Petch, Rosen, Savage, Silvestri,
Sjostedt, Smeathman, Wasmann, and many others. In the United
States, Banks, Buckley, Hagen, Heath, Howard, Hubbard, Joutel,
Knower, Marlatt, Porter, Schaeffer, Schwarz, Scudder, Stokes, Strick-
land, and Wheeler have contributed to the knowledge of the habits
of our native species. The following paragraph gives a brief resume
of the more important researches on the biology of the common
species of Leucotermes in the United States.

H. A. Hagen contributed several articles on the habits of flavipes,
and has described forms f> 0 found by the late Mr. H. G. Hubbard, of
the Division of Entomology. Hubbard, many of whose unpublished
notes are herein included, was the first to find royal individuals, both

a Escherich, K. Die Termiten . . . erne biologische Studie, p. 2-7, Leipzig,
1909.

b Holmgren, N. Termitenstudien 3. Systematik der Termiten. Die Familie
Metatermitidae. K. Svenska Vetensk. Akad, Handl., Bd. 48, No. 4, 166 p., Taf.
1-4, Uppsala & Stockholm, 1912.

Blick auf dem mutmasslichen, stammesgeschichtlichen Entwicklungsverlauf der
Termiten, p. 129-153.

Holmgren, N. Termitenstudien 2. Systematik der Termiten. K. Svenska
Vetensk. Akad. HandL, Bd. 46, No. 6, pp. 86, Taf. 1-6, Uppsala & Stockholm, 1911.
Ordnung Isoptera, p. 10-11.

c Bugnion, E. La differenciation des castes chez les termites [Nevr.]. Bui. Soc.
Ent. France, 1913, no. 8, p. 213-218, April 23, 1913.

d Bugnion, E. Les termites de Ceylan. Le Globe: Memoires, Soc. Geog. Geneva,
t. 52, p. 24-58, 1913.

e Loc. cit.

/Hagen, H. A. The probable danger from white ants. Amer. Nat., v. 10, No. 7,
p. 401-410, July, 1876.

9 Riley, C. V. Social insects from psychical and evolutional points of view. Proc.
PAL Soc. Wash., v. 9, p. 1-74, figs. 12, April, 1894.

18 INSECTS INJURIOUS TO FOEESTS.

true and neoteinic, in the United States, and his material forms an
important auxiliary to later investigations. Much of Hubbard's col-
lecting in Florida and Arizona was done in company with Mr. E. A.
Schwarz, of the Bureau of Entomology, who has since published a some
of his observations on the habits of termites in southwestern Texas.
The Rev.F.L.Odenbach,S. J., of Cleveland, Ohio, has studied the hab-
its of our native species of termites since 1893, and has contributed
many manuscript notes. Mr. L. H. Joutel, of New York, has studied the
habits of our common species6 and has contributed some unpublished
notes. Dr. H. McE. Knower,clate of Johns Hopkins University, in 1894
published new and important contributions on the embryology of ter-
mites (Eutermes) . His observations are not in accord with those of
Bugnion, since he determined that the nasutus develops from a
worker-like larva nearly as large as a young nasutus, having 13
joints to the antenna and worker-like head and jaws. This worker-
like larva had a small head gland with no ''corne front ale" on the
outside of the head, although sections show essentially the same
structure in the gland as that of the nasutus. Mr. C. Schaeffer, of
Brooklyn, N. Y., was the first to record d the presence of a fertilized
true queen of flavipes in a colony. The researches e of Dr. Harold
Heath, of Stanford University, CaL, on the habits of California ter-
mites confirm the statements of Perez and Perris that some colon-
izing individuals of Leucotermes lucifugus succeed in establishing new
colonies. Mr. Nathan Banks, of the Bureau of Entomology, has
contributed several articles on our native termites, and has described
a new species of Leucotermes (yirginieus) T' from the eastern United
States. Mr. C. L. Marlatt, assistant chief of the Bureau of Ento-
mology, has described the distribution, life history, and destructive-
ness of flavipes in the United States and figured the castes.^7 Mr.
E. H. Strickland,^ Carnegie scholar in economic entomology at Bos-

a Schwarz, E. A. Termitidse observed in southwestern Texas in 1895. Proc. Ent.
Soc. Wash., v. 4, No. 1, p. 38-42, Nov. 5, 1896.

& Joutel, L. H. Some notes on the ravages of the white ant ( Termes flavipes). Jour.
N. Y. Ent, Soc., v. 1, No. 2, p. 89-90, June, 1893.

c Knower, H. McE. Origin of the "Nasutus" (soldier) of Eutermes. Johns Hop-
kins Univ. Circ, v. 13, No. Ill, p. 58-59, April, 1894.

d Schaeffer, C. Jour. N. Y. Ent. Soc, v. 10, No. 4, p. 251, December, 1902.

c Heath, Harold. The habits of California termites. Biol. Bui., v. 4, No. 2, p.
47-63, January, 1903.

/Banks, N. A new species of termes. Ent. News, v. 18, No. 9, p. 392-393,
November, 1907.

9 Marlatt, C. L. The White Ant {Termes flavipes, Koll.). II. S. Dept. Agr., Bur.
Ent., Circ, No. 50, rev. ed., pp. 8, figs. 4, June 27, 1908.

ft Strickland, E. H. A quiescent stage in the development of Termes flavipes
Kollar. Jour. N. Y. Ent. Soc, v. 19, No. 4, p. 256-259, December, 1911.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 19

ton, Mass., has described and illustrated a quiescent stage in the
development of the nymph of the first form of Leucotermes jlavipes,
which stage was discovered by Prof. W. M. Wheeler, of Bussey Insti-
tute.

While investigating damage to the bases of telegraph poles by
wood-boring insects a large fertilized true queen of Leucotermes
flavipes was found by the writer on August 12, 1910. a The finding of
this interesting form, usually considered to be rare in colonies of our
native termites, served as an incentive to further biological study on
the habits of our common species. On August 11, 1911, molting

Fig. 4.— View at Falls Church, Va., showing a portion of the treated experimental stakes under test,
as to the relative effectiveness of preventives against termite attack. (Original.)

larvae in the quiescent stage were observed by the writer for the first
time, in a colony in Illinois. Special investigations were begun at
Falls Church, Va., in March, 1912, to determine (1) the habits of our
common termites, (2) the effectiveness of various methods and chem-
ical wood preservatives in preventing attack by our native termites
(fig. 4), and (3) the ll immunity " (?)& or relative resistance of native
and tropical species of wood.c

a Snyder, T. E. Record of the finding of a true queen of Termes flavipes Kol.
Proc. Ent. Soc. Wash., v. 14, No. 2, p. 107-108, June 19, 1912.

& It is doubtful if any species of native wood of economic importance is absolutely
immune to termite attack.

c Impregnation of wood to resist insect attack. Amer. Lumberman, Nov. 15, 1913

20 INSECTS INJURIOUS TO FORESTS.

BIOLOGICAL EXPERIMENTS.
THE TERMITARIUM.

Early in March, 1912, work was begun on a large outdoor termi-
tarium, at Falls Church, Va. The projects outlined for study in
connection with the termitarium were (1) to observe the progressive
development of the termite from the larva to the adult, especially
in the case of the colonizing form; (2, to watch the insects when they
swarmed, to determine whether this swarming is a nuptial flight,
i. e., whether or not the sexes leave the nests in separate swarms
and at what time copulation occurs; (3) to observe how the new col-
ony is established, (4) in what proportion the pairs survive, and (5)
the conditions in the parent colony after the swarm is over.

The termitarium was merely a large cage in which the termites
could swarm under conditions as nearly similar to those in nature
as possible. The cage consisted of a bottom of galvanized iron in
the form of a rectilinear box sunk in the ground to the depth of a
foot, a framework of wood to support the wire netting, and a wooden
roof covered with tarred paper. The bottom of the cage consisted
of a galvanized-iron box 10 by 6 feet and 1 foot deep, of 27-gauge
galvanized sheet iron with an inch flange turned inwards. (Fig. 5, a.)
This box was raised off the ground on a wooden floor of seven-eighths
inch material, 4 inches wide, which was laid on 2 by 4 inch studding.
There was an air space of 1 inch between the sides of the box and the
earth, as well as a similar space between the sheet-iron bottom and
the earth. The top of the sheet-iron box was nailed to a framework
of 2 by 4 inch studding, which was supported on cedar posts, the top
of the box being several inches above the surface of the ground.
Loose earth was placed in the box to about half its depth, over which
was spread a shallow layer of black earth (leaf mold) from the forest
floor, for the purpose of retaining moisture. The wooden framework
consisted of 2 by 4 inch studding, which was covered on the inside
with galvanized- wire netting (14 squares mesh per inch) and rein-
forced at the side and ends by boards 4 inches wide by seven-eighths
inch thick. The cage is about 7 feet in height, and the roof, slanting
away from a shed against which it is built, acquires sufficient pitch
to shed rainfall. (Fig. 5, b.)

Two chestnut logs which had been cut and allowed to season
about one month before the experiment was begun were placed in
the cage. One was placed on end in the dirt, the top of the log
approximating the height of a stump, while the other, a tangential
section, was partially buried in the ground. The bark was loosened,
but was left intact on both logs. The logs were both sound, but
were kept moist. A number of small decaying branches and strips of
decaying chestnut boards were also laid flat upon the earth in the

BIOLOGY OF THE TEEMITES OF THE EASTERN UNITED STATES. 21

•r.a

c3 T3

3 "

22 INSECTS INJURIOUS TO FOEESTS.

termitarium, after a careful examination to determine that no
termites were already present. The earth in the cage was kept
sufficiently moist in the endeavor, in so far as possible, to approxi-
mate natural conditions. (Fig. 5, c.)

The termitarium was ready for occupancy on April 8, and a chest-
nut log infested with a colony of Leucotermes jiavipes was introduced.
This log had the bark on and was partially buried. On April 9 a
decaying oak stump containing a colony of the same species was
also placed in the cage and partially buried in the earth. Several
termite colonies in logs and stumps in the forest were kept under
observation at the same time, and seasoned logs and slabs with
loosened bark were placed near by under conditions similar to those
in the cage. The following notes are based on observations of
colonies of termites in the termitarium, colonies in small tin boxes,
and colonies in nature.

COMMUNAL ORGANIZATION.

SITUATION OF THE NESTS.

Termites in the eastern United States usually make their nests in
decaying stumps or in logs and even small pieces of wood on the for-
est floor, although they also inhabit dead standing trees as well as
injured living trees. They never form mounds as in the Tropics. These
soft-bodied insects always conceal themselves within wood or in
earth as means of protection against sunlight and their enemies, in
consequence of which much of the damage they do is hidden. Ter-
mites of the genus Leucotermes are essentially wood destroyers and
infest and seriously injure a great variety of crude and finished
forest products which are in contact with the ground. The longi-
tudinal excavations usually follow the grain of the wood,a and in the
more sound wood their work is confined to the outer layers, where
there is abundant moisture and incipient decay. A protective outer
shell of wood is always left intact, since all except the winged, sexed,
colonizing forms shun the light and are blind. Small, transverse,
round tunnels which nearly pierce the outer shell are to be found
when their tunnels closely approach the exterior. Sometimes the
thickness of this protective shell is less than one-half millimeter.
These may be soundings to see how nearly the surface is being ap-
proached, or merely unfinished excavations for the exit of the sexed
adults, or possibly they may be feeding burrows. Termites often
take advantage of the burrows of other wood-boring insects, enlarg-
ing and adapting them to their purposes; by these means they are

a Termite work can be readily distinguished from that of carpenter ants, whose ex-
cavations do not follow the grain. Sometimes in decayed wood, however, termites
construct long, deep, but narrow, transverse galleries across the grain, forming ledges
or shelves,

Bui. 94, Part II, Buieau of Entomology, U. S. Dept of Agriculture.

Plate III.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate IV.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate V.

1

IfM'TO if! 1 f

BIB

'I'

ft j illlf !

tftj WL

11

fT'iliilllff)'1 ! 1 MmI "

ullllff)?.!-1-'} s'\t[1

Ji

■ IT' ^H jt j_ J ;,'

$lii ill

wllil NU

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ttPV 11

Hflfi

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Biiii'lSH !■

<

Damage by Termites.

Pine barn sill cut into ribbons by Leucotermes sp. at Mayfield, Kans. (Original.

Bui, 94, Part II, Bureau of Entomology, U S. Dept. of Agriculture.

Plate VI.

Damage by Termites.

Living stag-headed chestnut tree, 50 feet in length, Falls Church, Va. a, Complete length of
tree showing stag top and lightning scar on side; b, view of interior showing heartwood
completely honeycombed ; c, view of north side of tree showing width of scar, and honey-
combed interior. (Original.)

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 23

able to penetrate more rapidly to the heartwood and honeycomb the
interior.

Termites quickly disintegrate the wood of dead trees and stumps,
which soon becomes converted to humus, the rapidity with which
this is done depending on the relative resistance of the species of
wood. This beneficial role in nature, however, is offset by the enor-
mous destruction they accomplish in rapidly rendering insect, fire,
and disease killed timber unmerchantable and by the damage they
inflict to the roots and trunks of injured living trees. Termites will
infest the heartwood of living trees injured at the base by fire, dis-
ease, or other insects (PI. Ill), and sometimes in such trees they
excavate upward, throughout the dead heartwood, longitudinal tun-
nels, irregular in diameter, the sides of which are lined with earth
mixed with excrement. These insects also infest the roots of living
trees, finding ingress through abandoned burrows of the large,
roundheaded (Prionid) borers. Sometimes they girdle young trees —
forest-tree nursery stock, for example — -eventually cutting the trees
off near the ground, examination disclosing that the stems were
honeycombed. This is not necessarily due to the presence of dead
wood near by, since termites will tunnel for long distances under-
ground. While usually confining their work to moist or decaying
timber or to vegetable material of any sort, and to books (PL IV)
and papers that are somewhat moist, termites will attack seasoned,
dry wood, provided there is access to moisture elsewhere; i. e., they
use moist frass and earth in extending the burrows, thus creating
more favorable conditions. In the Southern States termites will
infest the bark and outer layers of the wood of the base of yellow
pines killed by barkbeetles before the foliage has all fallen; trees
that have been killed in the spring and show reddish-brown needles
and much fallen foliage being infested by the middle of August.
Trees killed in the spring will also have the outer layers of wood of
the base honeycombed by the following December. (Fig. 6.) The
larger-celled, thin-walled spring wood is eaten away first, leaving the
smaller-celled, harder summer wood uneaten. (11. V.)

Where the heartwood is decayed in a standing living tree termites
will work for a considerable distance above the ground, completely
honeycombing the heartwood. In a living chestnut tree at Falls
Church, Va., with the decayed heartwood exposed in a long scar,
termites had infested the heartwood to a height of from 45 to 50 feet
above the ground. The outer shell of living sapwood was intact.
(PL VI.)

Termites are quite effective in clearing fields of old snags and
stumps, but this benefit is offset by the damage they do to posts and
buildings.

62896°— Bull. 94, pt 2—15 2

24

INSECTS INJURIOUS TO FORESTS.

Termites (Leucotermes spp.) also inhabit subterranean
Drummond has compared tropical species of termites to earthworms
and declared that they are equally as beneficial to man in aerating
the soil. After swarming, many of the sexed adults excavate shallow
cells in the earth under small pieces of decaying wood, and later enter
the wood. The royal cell is constructed in decaying wood or in the

earth slightly below the surface of
the ground. Termites usually in-
fest wood by entering from the
ground underneath, rather than
directly on the exposed surface, the
latter being usually the habit at the
time of the swarm. (Termopsis
angusticollis Walk, usually infests
wood by gaining ingress through
wounds and abrasions.) None of
the sound traplogs with loose bark
in the termitarium or in the forest
was infested except at the point of
contact with the ground, but ter-
mites in pairs have been found un-
der loose split bark on decaying
logs where more moisture was pres-
ent. Workers and soldiers are fre-
quently to be found in the spring
in small pieces of decaying wood
lying on the ground, and termites
probably extend old colonies or es-
tablish new colonies by means of
subterranean tunnels. During the
winter the members of the colony
are to be found in a labyrinth of
undergound passages.

These excavations are of varying
size and shape, and extend in all di-
rections. Some of the tunnels are
partitioned off into separate cham-
bers, while others are unpartitioned
runways. In the main runways the very young are absent. The
partitions consist either of uneaten portions of the wood or small coni-
cal piles of moist earth mixed with frass (excreted, finely digested
wood) of clay-like consistency. Sometimes in broad, shallow chan-
nels a small irregular mass nearly blocks the channel. The sides of
the channels are smooth, and the uneaten masses of wood which
serve as barricades appear as little islands and are distinct because of

i , I illiu

\m

lift ifSiS

1

:

hi

m

: $ j ! i | - .. -Li

*» . .>?

|P#

Hii? 1 V

m

HBIif

['. - - . .._ L,

Fig. 6. — Work of termites in insect-killed southern
yellow pine. Tree killed in the spring; wood at
base honeycombed by following December.
Spartanburg, S. C. (Original.)

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 25

the rough appearance due to the pores and cell walls of the wood.
The walls of these channels are spotted with little piles of finely
digested, excreted wood, giving the wood a characteristic mottled
appearance.

Large cavities encountered by termites when working in the wood
of logs, poles, or trees with decayed heartwood or hollow core are
usually filled up with moist earth mixed with frass, the whole having
a clay-like consistency and a conglomerate appearance due to the
irregular deposits of excreted, finely digested wood. The "doty"
hollow cores in the bases of infested poles are filled up in this manner.

In all their operations the termites carefully wall up and conceal
themselves. Usually there is but little evidence on the exterior of
infested wood to indicate the presence of termites, and they may not
be detected until the interior is completely honeycombed. Sexed
adults swarming from infested buildings are always a warning of their
presence. Again, the outer surface is covered over with longitudinal
viaducts of small diameter constructed of earth (PL VII, figs. 1 and 2),
and whereas the interior channel is smooth, the exterior has a rough
granular appearance. These viaducts are resorted to in order to
extend the colony or to reach some object, such as decayed wood, or
when their pathway crosses some impenetrable substance, the object,
of course, being always protection from the light — all except the
winged sexed adults shun the light — and their numerous enemies.
Viaducts, or " sheds," can be seen running up to a considerable height
above the ground in the longitudinal weathering checks on poles and
posts, as well as between the crevices in the bark of infested dead
and living trees, and uncovered viaducts are found under the loose
bark on dead trees. Viaducts in the interior of hollow-cored poles or
trees may be of large diameter and may consist of several irregular
longitudinal interior channels.

Termites sometimes resort to another type of viaduct, which for
descriptive purposes may be called suspended tubes. On May 26,
1912, at Elkins, W. Va., Dr. A. D. Hopkins found termite tubes of earth,
5 to 6 inches long, hanging from the end of a Virginia scrub pine
sapling which had fallen, leaving the broken base 2 feet from the
ground. The termites had evidently infested the base of the sapling
through the ground before it fell and were trying to make connection
with the ground by means of these suspended tubes. (Figs. 7 and 8.)

These covered viaducts or sheds, the uncovered viaducts, and the
tubes — all constructed by termites of earth and excreted wood — are
fragile.

NUMBER OF INDIVIDUALS IN COLONIES.

Young colonies of Leucotermes jiavipes — that is, colonies but
recently established, in decaying wood or in the ground under decay-
ing wood, by sexed couples that have swarmed — are small, and since
the rate of egg-laying by the young queens is remarkably slow, the

26

INSECTS INJUEIOUS TO FOKESTS.

increase in numbers is also correspondingly slow. Observations of
such incipient colonies in the spring of 1912 and 1913, after the swarm,
the time of which varies with the season, from the middle of June to
early July, indicate that from 6 to 12 eggs are normally to be found
with pairs of jiavipes. While the brood first hatched is relatively
small, contrary to the habits in the other social insects coition is
repeated and the young colony gradually increases in numbers.

On April 25, 1912, at Falls Church, Va., a small colony or sub-
colony of Leucotermes Jiavipes, the branch of a larger colony, was
found under a small chestnut slab sunken in the ground. The day
was warm and bright and many members of the colony were congre-

Fig. 7.— Suspended tubes, constructed by termites of earth and excreted wood.
(Original.)

gated under the slab above ground. An attempt was made to cap-
ture the entire colony in order that the relative proportions of the
various castes might be ascertained. Although nymphs of the first
form constituted the most abundant caste, some few transforming
nymphs and a few sexed adults were present, as well as a few workers
and soldiers. Workers constituted the second most abundant caste.
Nymphs of the second form appeared more active than the
nymphs of the first form. The following possibly is not a fair state-
ment, either of the number of individuals in the colony or the rela-
tive proportions of the different castes, since (1) the nymphs were
probably above ground to take advantage of the warm afternoon
sun which would aid their development — a common procedure; (2)
only a few workers and soldiers were necessary to attend them and
no doubt many more were still in the subterranean passages; (3)

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate VII.

3 if-

lo

-I ?

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 27

many escaped and the workers would escape more easily than the
nymphs; (4) probably, unconsciously, more effort was made to cap-
ture nymphs and soldiers than workers. The following figures,
however, will probably approximate the relative abundance of castes.
Nymphs of the first form, 279; nymphs of the second form, 86;
nymphs of the first form in the quiescent stage, 31 ; individuals in the
stage following, that is, with wings unfolded and held away from the
body, 3; immature sexed adults without pigmentation, 17; immature
sexed adults with gray pigmentation, 26; nearly mature adults, 5;
nymphs of the second form molting, 4; workers, 93; soldiers, 24;
the total being 568. Nymphs of the second form were only one-
fourth as numerous as nymphs of the primary form, and, including
all stages to sexed
adults, gave a count
of 90 to 361, respec-
tively.

On March 22, 1913,
at Black Mountain,
N. C, nymphs of the
second form (Jiavipes)
taken in a small de-
caying oak branch on
the ground greatly
outnumbered nymphs
of the first form, al-
though usually the lat-
ter is by far the more
numerous form.

Due to the wander-
ing habits of species of
this genus, it is difficult to estimate the size and extent of an old,
well-established colony, which may branch out over several acres of
grounds However, the number of individuals in well-established,
permanent colonies probably runs up into the ten thousands, since
from 5,000 to 10,000 (estimated) eggs, scatteringly or in clusters the
size of a pea, were found in a large colony- of Leucotermes virginicus
near Chain Bridge, Va., on June 19, 1913. This colony, which was
in a large decaying black oak log, consisted of a large number of
workers and soldiers, and numerous larvae.

THE DIFFERENT CASTES— POLYMORPHISM.

In a termite colony there are several different forms, or castes, of
mature individuals, as well as those of different castes in the various
stages of development. The castes are the workers, the soldiers, the

Fig. 8.— Broken-off pine sapling from basal end of which tubes in fig-
ure 7 were suspended by termites toward stump. (Original.)

a The spreading out of a colony is largely due to increase in numbers and consequent
need of fresh supply of food; that is, decaying wood.

28 IXSECTS INJURIOUS TO FORESTS.

colonizing winged adults of both sexes, the supplementary or neo-
teinic° reproductive forms (often in large numbers), both "ergatoids"
and nymphal ' meotenes, ' ' and the single true royal pair. Besides these
mature forms there are freshly hatched, undifferentiated larva?, differ-
entiated larva?, and nymphs of the first and second forms. Of course,
all these forms are not present in a colony at the same time, as there
is seasonal variation.

Workers are developed from larva? that will not mature the sexual
organs, but, unlike the bees, are of both sexes. They are dirty white
in color, are large-headed and soft-bodied, with 10 segments to the
abdomen, and at maturity are approximately 5 millimeters in length
mjiavipes and 3.5 millimeters in virginicus. The antenna? consist of
15 to 17 segments exclusive of the base mjiavipes and 13 to 15 in
virginicus. The workers constitute the most injurious wood-de-
stroying form and have well-developed mandibles. The left man-
dible has five pointed teeth, the fifth tooth with a broad base, and the
inner margin having parallel carina?. The right mandible has two
pointed teeth, the third and fourth teeth being broad, and the fourth
with parallel carinse. The mandibles evidently have both tearing
and rasping functions. The labrum is rounded. The worker ter-
mites possess a large intestinal paunch and the contents enable the
outline of this paunch to be seen through the tissue of the abdomen.
Workers are blind.

Soldier termites (fig. 9, a) are more highly specialized workers,
being also developed from large-headed workerlike larva? that will
not mature the sexual organs, and the caste is represented by both
sexes. While they are soft-bodied, the head, which is pigmented
yellowish-brown, is chitinized and is more oblong and elongate than
in the worker, tapering slightly toward the apex, or being slightly
broader at the base. The mandibles, which are enormously devel-
oped, are long, slender, saberlike, with no marginal teeth, chitinized,
and of a yellowish-brown color. The body is of a dirty white or pale
yellowish color. The labrum of the soldier is more narrow than
that in the worker, elongate, and subelliptical. The '"meiiton,"
which is chitinized, is convex, elongate, and more slender in com-
parison with the worker. Mature soldiers are from 6 to 7 millimeters
in length mjiavipes and 4.5 to 5 millimeters in length hi virginicus.
The antenna has from 14 to 17 segments mjiavipes and 15 segments
in virginicus. The soldiers, as well as the sexed adults, of these two

aGrassi, B~, and Sandias, A. Op. cit., p. 249: "The term neoteinia has been intro-
duced by Camerano (Bui. Soc. Ent. Ital. [v. 17], 1885, pp. 89-94) to denote the per-
sistence during adult life of part or all of the characteristics normally peculiar to the
immature, growing, or larval stages. - * * Xeoteinia, or the persistence of larval
characteristics, does not necessarily imply that anticipation of sexual maturity which
is usually connoted with the use of the term pedogenesis, which, moreover, is
strictly applied to agamic reproductions."

BIOLOGY OF THE TEEMITES OF THE EASTERN UNITED STATES. 29

species can be differentiated. The soldiers, like the workers, are
wingless and blind. Individuals of both castes complete their devel-
opment in less than a year.

The nymphs of the reproductive form develop from larvae that will
mature the sexual organs. The term "nymph" is applicable,

V, ^

<^3 w

2 a

03 £i
~ C3

e a

t-J

Si

Ca Sh .Sf

e 1 «

8 « s

s a "§

according to Lespes and Hagen, to individuals "with the wing-
rudiments easily distinguishable to the naked eye."a Individuals
the wing-rudiments of which are present, but only distinguishable

are termed larvae. Nymphs are white

when under magnification,

a Grassi, B., and Sandias, A. Op. cit., p. 264.

SO INSECTS INJURIOUS TO FORESTS.

and soft-bodied, and when fully developed and ready to molt for the
last time are from 7 to 7.5 millimeters in length in flavipes and 4.5
to 5 millimeters in virginicus. In flavipes the antenna consists of
from 16 to 17 and in virginicus from 14 to 15 segments. The man-
dibles are practically the same as in the worker, and are probably
very necessary in effecting an exit from the old colony. R. D. Grant
states0 that in a Missouri Pacific Railroad Co. engine house the
rafters were injured and the cement of the brick walls built 14 years
previously was perforated. Mr. C. L. Marlatt has specimens of
plaster which was laid on metal lathing in a building at Charlotte,
N. C, which had been mined in order to allow the winged adults to
escape from heavy wooden beams which had been honeycombed.
Also, in the establishment of the new colony the young royal couple
have the excavating to do.

There are two forms of nymphs (PI. VIII, figs. 3, a, b), namely, the
primary form, with elongate wing pads, that develops into the winged
sexed adult, and the "second form" (Lespes), with short wing pads —
mere buds — which represents an arrested early stage of the nymph
of the primary form, or even a larva. Nymphs of the secondary
form are slightly more elongate, and develop the sexual organs
without progressing further, instead of completing their normal
development to the winged, pigmented, sexed adults that swarm.
These nymphs, after becoming sexually mature and attaining a straw-
colored pigmentation — normally after the nymphs of the primary
form have developed to sexed adults — become supplementary royal
individuals, kings, and queens, but never ( ?) leave the parent colony.
They do not. possess functional eyes.

The sexed individuals, when ready to swarm, are castaneous-black
or light brown in color, have two pairs of long, filmy wings, and are
so chitinized that they can bear full sunlight. They possess both
functional compound eyes and simple eyes or ocelli. The body — ex-
cluding the wings, which are slightly longer than the entire insect
and project some distance beyond the abdomen when "in situ " — is
slightly less in length than in the case of the nymphs. The entire
body of the sexed individual is from 9 to 10 millimeters in length in
flavipes and from 7.5 to 8 millimeters in length in virginicus. There
are from 16 to 18 segments to the antenna in. flavipes and from 15 to
16 antennal segments in virginicus. The sexes are as easily distin-
guishable as in the case of fully developed nymphs. "(1) The sev-
enth [abdominal sternite] (the apparent sixth) is strongly developed
and semicircular, with the rounded edge posterior in the female and
very short in the male. (2) The eighth sternite, which is reduced
to two lateral lobes in the female, is small and entire in the male.

a Grant, R. D. Jour. Proc. Acad. Sci. St. Louis, v. 3, p. cctxix, November 19, 1877.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate VIM.

Fig. l.—a, Lateral view of fully developed
nymph of primary form; b, lateral view of
neoteinic king— compound eyes without
pigmentation.

Fig/2. — Dorsal view of same: a, Showing pubes-
cence and tapering abdomen of neoteinic
.king; b, nymph of the primary form.

Fig. 3.— a, Fully developed nymph of the first form with elongate wing pads before final
molt; b, nymph of the second form with short wing pads before final molt; c, neoteinic
king after final nymphal molt. (Original. )

Stages of Leucotermes flavipes.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 81

(3) The ninth sternite nearly resembles the eighth/ ,a The genital
" appendices" in the male are attached to "what is apparently the
eighth, but is really the ninth sternite, the first being fused with the
metasternum."6 There are two segmented appendages, or cerci, at-
tached to the abdomen. After swarming, the sexed adults become
royal individuals.

Larvae, or young, are undifferentiated individuals which, after
further development, attaining chitinization and pigmentation, change
to differentiated individuals. The larvae, like all the other stages of
termites, are active. When young or freshly molted the individuals
are more transparent and white and the segmentation of the body at
this stage is more sharply defined. "The younger the individual, or
the more recent its ecdysis, the thinner is the chitin." c The term
larva is applied (1) to any individual which has not yet attained full
size, mature chitinization, and pigmentation; (2) in the case of the
sexed individuals, to those with the wing-rudiments not easily dis-
tinguishable to the naked eye; and (3) in case of the soldier, to the
large-headed, undifferentiated, workerlike forms.

The eggs are white, slightly reniform, and those laid by true queens
(queens that have swarmed) are approximately 1 millimeter in length
in jiavipes. The eggs are usually found in clusters or scattered
singly in the galleries.

THE SENSE ORGANS.

Termites are essentially subterranean in habit and in consequence
all castes of Leucotermes are blind except the colonizing individuals.
The soldiers have compound eyes, but without pigmentation; in
some neoteinic royal individuals the pigmentation of the compound
eye is not visible to the naked eye (PI. VIII, fig. 2, a), but in most
cases there is a slight pigmentation of variable intensity. All castes
except the colonizing individuals shun the light.

Although blind, termites are known to possess other sense organs.^
The antennas are important tactile sense organs, and often individuals
may be seen feeling their way by means of these appendages. The
antennae of the colonizing individuals are pitted, and from these
pits, which appear as white depressions on the pigmented antennal
segments, prominent hairs arise. A. C. Stokes, in an article entitled
"The sense organs on the legs of Termes Jiavipes Koll.," e describes

aGrassi, B., and Sandias, A. Op. cit., p. 306.

&Ibid., p. 271.

c Ibid., p. 256.

d Miiller, Fritz. Beitrage zur kenntniss der Termiten. Jenaische Ztschr, Bd. 9
(n. F. Bd. 2), p. 241-264, pi. 10-13, Mai 8, 1875. See p. 254, pi. 12, figs. 32, 34.

e Stokes, Alfred C. The sense-organs on the legs of our white ants, Termes Jiavipes
Koll. Science, v. 22, no. 563, p. 273-276, illus., November 17, 1893.

32 INSECTS INJURIOUS TO FORESTS.

and figures "sensory hairs"; "sensory pits"; "tibial spurs with
prominent basal apertures, across which extends a delicate membrane
(auditory organs?)"; "pilose depressions"; "surface markings";
etc. Grassi ° mentions tactile, "very long, fine, readily vibratile
hairs" on the body, and states that the cerci also appear to be "es-
sentially tactile." It is believed that there is a relation between the
convulsive movements frequently observed, that is, the sudden
jerking of the whole body, and these sense organs, and that indi-
viduals are thus enabled to communicate, or at least give danger
or distress signals. The convulsive movements made by the workers
and soldiers, when the royal pair are disturbed in the cell, are very
violent and indicate great agitation.

There is a characteristic musty or acrid odor which can be easily
detected in colonies of Leucotermes, and individuals frequently
can be seen to follow directly in the path taken by others, but as
termites usually travel in well-worn channels this may be due to
tactile sense alone.

THE FUNCTIONS OF THE CASTES.

The social economy of termites is somewhat similar to that of other
social insects, although in many respects totally different. Undif-
ferentiated young hatch from the eggs, are active, and in turn trans-
form to the differentiated individuals of the castes after a series of
quiescent stages and molts.

As the name implies, the workers are those individuals that make
the excavations, extend the colony, and care for and protect the
royal couples and young.

The soldiers, more highly specialized workers, are of less importance
functionally than the workers — just as the anther transformed to the
petal in the common pond lily, Castalia sp., is less important function-
ally than the other anthers — yet both serve a purpose. Just before the
time of swarming, the members of colonies become restless, and as
the sexed adults emerge numerous workers and soldiers congregate on
the outskirts of the colony near the exit holes with heads toward the
exterior. The duty of the soldiers is apparently entirely protective,
but they do not appear to be very effective, at least when the colony
is opened and they are exposed to the attack of ants, etc.

The pigmented, winged, sexed adults, developed from nymphs
of the first form, are the colonizing individuals which swarm in
enormous numbers in the spring and found the new colonies, and
when finally established at the head of a new colony, which they
have reared, they become the royal couple. Unlike other social
insects, the male continues to live, even after the fertilization of the
female, and the king and queen inhabit the royal cell together, there

a Grassi, B., and Sandias, A. Op. cit., p. 267-269.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 33

being repeated coition. Termites are sometimes polygamous, at
least in incipient colonies. The young queens care for the young and
carry the newly hatched larvae in their mouths to safer places in the
nest when the colonies are disturbed. (Nymphs of the first form also
have been seen performing this duty in old colonies.) The workers
attend the larvae in old colonies.

In the life cycle of termites, however, there is so much variation in
the development of the castes that it leads to a rather complex life
history. Young are kept in a retarded, undifferentiated state and can
speedily be turned into substitute reproductive forms of both sexes.
These neoteinic royal individuals are used (1) as substitutes for the
true royalty that have swarmed and (2) in splitting up the old colony
into new and independent colonies. Fully developed nymphs of the
second form are to be found in colonies in the spring. They never
complete their normal development, which consists of the acquisition
of wings and the mature pigmentation, but assume the characteristic
pale straw color after the final molt. The sexual organs are developed
after the acquisition of pigmentation. They become neoteinic repro-
ductive forms, but always ( ?) remain in the parent colony.

Each caste has a distinct function, there being a well-defined
division of labor, which, however, is not strictly adhered to. Ter-
mites properly represent the phenomenon of polymorphism, i. e.,
u species in which one or both of the sexes appear under two or more
distinct forms,a" both sexes being equally polymorphic.

THE LIFE CYCLE.

THE METAMORPHOSIS CASTE DIFFERENTIATION.

The metamorphosis of termites, while formerly considered to be a
contrastingly simple type of insect development, is in reality very
complex; indeed, there may be said to be different types of develop-
ment for the castes. In the development of the worker, as in the
order Thysanura, there is no external change or metamorphosis, the
freshly hatched worker larva being active and of the same form as
the adult worker. In the development of the soldier, however,
marked changes in form occur, the mature soldier, with pigmented
head and saberlike mandibles, being developed from a large-headed,
white, workerlike larva (PL IX). The winged, pigmented, sexed
adult is developed from a small-headed, white larva, there being a
still more radical change. However, what may popularly be termed
the "antlike" form can be distinguished in all stages in the devel-
opment of all the castes, hence termites can hardly be classed with
insects with et complete metamorphosis ' as the true ants, where there

a Wheeler, W. M. Concerning the polymorphism of ants. Bui. Amer. Mus. Nat.
Hist., v. 23, p. 50-93, 6 pis., January, 1907. See p. 50.

34 INSECTS INJURIOUS TO FORESTS.

are the three successive stages of larva, pupa, and adult, although
termites pass through pupalike "quiescent" stages in the molting of
larvas and nymphs, with temporary periods of inactivity. Further-
more, great variation is possible in the metamorphosis, as is seen in the
development of "neoteinic" or lt supplementary" reproductive forms,
as the "neotenes" and "ergatoids." Therefore, in general, termites
may be classed with insects having " incomplete metamorphosis,"
as the locusts and roaches. In this type of development the form of
the body is always essentially the same as that of the adults. The
development of the larva and nymph to the winged sexed adult ter-
mite greatly resembles the development of the locusts and roaches.
In the genus Leucotermes there is no apparent metamorphosis in
the case of individuals of the worker caste. The freshly hatched
worker larva is active, there being a very simple development. The

Fig. 10.— Leucotermes flavipes: Quiescent stage of molting larvse. Enlarged 10 times.

(Original.)

change from larva to adult is a gradual growth, the white, large-
headed larva passing through a series of quiescent stages and molts.
The adult worker is of a dirty grayish-white color. In the soldier
caste and the reproductive forms the metamorphosis is more com-
plicated, and marked changes in form occur during the development.
Young molting larvae in the " quiescent stage" a were first observed
by the writer on August 11, 1911, near Jerseyville, 111., in a large,
wide, longitudinal channel in the decaying heartwood of the butt of
a white cedar telegraph pole.5 Previous to the molt the larva falls
over on its side and passes through a quiescent stage, the head being
bent down to lie on the ventral side of the body along which the
antennas and legs, also, lie extended in a backward direction. This
gives the larva the appearance of being doubled up (fig. 10). After
the skin has been shed the larva resumes its normal activity.

a Strickland, E. H. Loc. cit.

& Snyder, T. E. Changes during quiescent stages in the metamorphosis of ter-
mites. Science, n. s., vol. 38, No. 979, pp. 487-488, October 3, 1913.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate IX.

f r

'*!## it*1

d

F*»ltiit5

Leucotermes virginicus and L. flavipes.

Molting soldier nymphs: a, L. virginicus, head showing mandibles and iabrum of soldier nymph
just molted from workerlike larval form; b, L. flavipes, same at later molt; c, L. flavipes, same
atlatermolt, d, L. flavipes, mature soldier; d', ventral view snowing convex, slender, chitinized,
pigmented "menton." Photograph from specimens in balsam slides. Enlarged 16 times.
(Original.)

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 35

The antennae increase in the number of segments by the subequal
division of the third segment a independent of the molts in L. luci-
fugus Rossi.

In the metamorphosis of the soldier, as has been shown by Grassi a
{Calotermes flavicollis Fabricius and Leucotermes lucifugus Rossi),
Knower b (Eutermes sp.), and Heath6 (Termopsis angusticollis Wal-
ker), there is a radical change from a large -headed larva to the
soldier caste. This change takes place during c a quiescent stage.
The worker, like the soldier, also develops from a large-headed larva ;
that is, in the colony there are two types of larvae, the large-headed
and small-headed, the former normally developing to workers and
soldiers, while the latter become the reproductive forms. According
to Grassi there are four molts in the development of the asexual forms
of Leucotermes lucifugus, whereas the sexed forms pass through five
molts. Holmgren states d that in Leucotermes the soldiers are
polymorphic and originate e from at least five different larval stages,
while the polymorphic workers originate from three different larval
forms. The undifferentiated larvae present possibilities of develop-
ment in all directions, but the development of Leucotermes shows
that the breeding possibilities, while great, are by no means so great as
in Calotermes. This would indicate progressive development toward
stability, as is also indicated by the absence or rarity of neotenes and
ergatoids in the more highly developed termites (Termes and Euter-

a Grassi, B., and Sandias, A. Op. cit., p. 270.

1* Op. cit.

c Snyder, T. E. Loc. cit.

^ Holmgren, N. Termitenstudien 3. Systematik der Termiten — Die Familie
Metatermitidae. K. Svenska Vetensk. Akad. Handl., Bd. 48, No. 4, pp. 166, 4 pis.,
Uppsala and Stockholm, 1912. Blick auf dem mutmasslichen, stammesgeschicht-
lichen Entwicklungsverlauf der Termiten, p. 129-153.

« The recent discovery by McClung, Stevens, and Wilson (Wilson, E. B., Heredity
and miscroscopical research: Science n. s., v. 37, No. 961, p. 814-826, May 30, 1913)
of the association of an odd number of chromosomes, in the divisions of the spermato-
cytes— that is, cells formed by the division of the "spermospore," the male germinal
cell — with the determination of sex may also be applicable to caste differentiation in
termites. Wheeler (Wheeler, W. M., The polymorphism of ants, with an account
of some singular abnormalities due to parasitism: Bui. Amer. Mus. Nat. Hist., v. 23,
p. 1-93, pis. 6, Jan., 1907), however, states, with reference to ants, that nourish-
ment, temperature, and other environmental factors merely furnish the conditions
for the attainment of characters predetermined by heredity, that is, with Weismann
he believes that the characters that enable us to differentiate the castes must be repre-
sented in the egg, but with Emery he believes the adult characters to be represented
in the germ as dynamic potencies or tensions rather than morphological or chemical
determinants. Holmgren states (op. cit., p. 140) that in termites, as the result of
the method of feeding, three potential germ plasms are released in at least three direc-
tions * * * and that there must be a germ plasm correlation which finds its
expression in the caste correlation.

36

INSECTS INJUBIOUS TO FORESTS.

mes), wherein, however, there may be several true queens0 in the
one colony and where polygamy exists. In insects supposed to
represent the most primitive, or lowest and least developed, types
this is a rather complex metamorphosis. Here also there is much
variation in the life cycle and no strict rule is followed. (See chart.)
Apparently the individual development is entirely subservient to the
needs of the colony. This ability of adaptation of individual to
circumstance leads to a complex economy.

A representation1 of some of the successive stages in the development of the various forms
or castes in the life cycle of Leucotermes flavipes Kollar as found in colonies in the eastern
United States. All these reproductive forms are not present in the same colony at the
same time.

(A)

(1) Young, undifferentiated larvae.

(B)(2) Large-headed larvae that (3) Larvae that wi
will not mature sexual or- organs,
gans.

mature sexual (4) Reserves for emergency repro-
ductive forms.

(C) (5) Larvae of
soldiers.

(6) Lar v ae of
workers.

(D) (7) Soldiers. (8) Workers.

(9) Nymphs of
the first form
with long
wing pads.

(12) W inged,
p ig men ted,
sexed adults.

(10) Nymphs
of second form
(Lespes) with
short wing
pads.

(11) Reserves for emergency
productive forms.

(13) The true (14) Nymphal (15) Nymphal (16) Larval

royal pair, neoteinic sub- emergency emergency

with wing stitute repro- substitute substitute

stubs. ductive forms. reproductive reproductive

forms. forms.

1 See correlated forms of Leucotermes lucifugus Rossi. Grassi, B., Ein weiterer Beitrag zur Kenntnis des
Termitenreiches. Zool. Anz, Bd. 12, No. 311, pp. 355-361, July 8, 1889. "Ubersicht der im Termitenstaate
vorkommenden Formen- Termes lucifugus, p. 360); id., Ein weiterer Beitrag zur kenntniss der Termiten-
reiches. Ent. Nachr., Jahrg. 15, No. 14, pp. 213-219, July, 1889; Holmgren,N ., Termitenstudien 3. Syste-
matic der Termiten-Die Familie Metatermitida?. K. Svenska Vetensk. Akad. Handl., Bd. 48, No. 4,
p. 148, Scheme B, Uppsala & Stockholm, 1912 (table showing parallel evolution and at what molts changes
or development occur).

PROGRESSIVE DEVELOPMENT OF THE NYMPHS.

Colonies of Leucotermes spp. in the northeastern United States are
dormant during the winter, the insects retiring to the more remote

a. John. O. Notes on some termites from Ceylon. Spolia Zeylanica, v. 9, pt. 34,
p. 102-116, 1913.

Riley, C. V. Termite economy. Proc. Biol. Soc, Wash., v. 9, p. 71-74, April,
1894.

Escherich, K. Termitenleben auf Ceylon, Jena, 1911, p. 45-46.

BIOLOGY OF THE TERMITES OE THE EASTERN UNITED STATES. 37

galleries in the wood or to the subterranean passages of the nests,
At Falls Church, Va., in 1912 it was not until March 11, in 1913.
February 20, and in 1914, March .23 that signs of activity were
observed in colonies of flavipes. Nymphs of the first form with
"short wings" (Fritz Muller) or "wing pads" (Hagen) were present
on March 11, 1912. At this time the wing pads were well developed,
being about two and one-half times the length of the segment from
which they originated, and had a yellowish tinge. The antennae
consisted of from 16 to 17 segments, excluding the base, and the line
of demarcation between the basal segments was less distinct. On
March 29, 1912, the ocelli were visible in the nymphs; antennae,
head, and thorax were acquiring a tinge of yellowish-brown, and the
compound eyes were becoming pigmented — a reddish-brown. On
April 18, 1912, and April 8, 1913, at Falls Church, Va., nymphs of
the first form when fully developed and ready for the final molt
could be readily distinguished by the opaqueness of the elongated
wing pads, the filmy, yellowish-brown, loosening skin, which be-
comes separated from the body, particularly posteriorly, and the
reddish-brown pigmentation to the compound eyes. (PI. VIII,
fig. 1, a.) The nymphs gradually increase in size and pass through a
series of molts and quiescent stages until the final molt, when the
wings are unfolded. Packard, who figures ° the stages in the growth
of the wing in flavipes, states that the wings are simply expansions.
During the latter part of April, 1912, nymphs with short wing pads,
or those of the second form (Lespes) (PL VIII, fig. 3, b), were found in
colonies. Their antennae had from 1 6 to 17 segments. These nymphs
appear to be more active than nymphs of the first form and have
but slight pigmentation of the compound eye.

During the final molt — which occurred from April 18 to 27, 1912,
April 8 to 17, 1913, and April 22 to May 2, 1914, at Falls Church, Va.,
in case of the nymphs of the first form (flavipes) — nymphs of both the
first and second forms pass through a " quiescent stage " b (Pis. X and
XI), which closely approaches the pupal stage of insects with com-
plete metamorphosis. This quiescent stage apparently serves the
same purpose as the pupal stage, since the most marked changes,
both external and internal, take place during this molt ; it is, however,
of short duration. On April 18, 1912, the first nymphs (flavipes) in
this stage were observed in the outer layers c of a decaying stump . All
of the nymphs in this colony were fully developed, but very few had

o Packard, A. S. A textbook of entomology. New York, 1903, p. 140.

& Strickland, E. H. Loc. cit.

c Larvae or nymphs in the quiescent state are usually to be found isolated in small
but deep, transverse conical nitches or shelves in the nest where they are not liable to
disturbance by the movements of the other members of the colony. Possibly they
seek out such secluded places, usually near the outlying galleries, beforehand, or may
be carried there while helpless, by the workers. Clusters of eggs are also found in
similar nitches.

38 IKSECTS INJURIOUS TO FORESTS.

yet begun to molt. A large number were placed in a covered tin box
3^ by 5 by 1J inches, with disintegrated wood and moist earth in the
bottom. This was to be a check box, and the conditions here were
more natural than in the small corked vials, in which over 100 other
nymphs had been kept in the dark, one nymph to each vial. The
development of each nymph was watched and the time necessary for
the various changes from nymph to adult was noted. On June 8
fully developed nymphs of virginicus, with opaque wing pads, were
observed to molt in a similar manner. Nymphs in all stages described
for flavipes were observed molting until June 1 1 .

As E. H. Strickland has already figured and described these
changes in the nymphs of the first form (flavipes), his accurate
description is here quoted in detail, with a few comments. Until
this description appeared, the quiescent stage in the development
from nymph to adult for this species was undescribed. His obser-
vations were made from nymphs collected in the neighborhood of
Boston, Mass., May 7, 1910.

The mature nymph becomes very sluggish and finally all movement ceases; it
then falls over on its side and the head is bent down till it lies on the ventral side
of the body, along which also the antennae and legs are extended in a backward
direction, * * * while the wing pads are bent downward till they lie laterally
along the sides of the body. * * * It will be at once noticed that while in tnis
position the nymph is to all appearances a quiescent "pupa libera." There does
not appear to be an ecdysis immediately prior to this quiescent period, however,
so I would hesitate to describe it as a true pupal state though it undoubtedly has
the same physiological function.

This quiescent stage lasted in the few specimens observed for a peri 3d varying
from four to about nine hours.0 The duration in time seems to be controlled to a
large extent by the amount of moisture in the earth surroundiug the pupa for when
specimens were placed in perfectly dry earth they were unable to pass beyond this
stage of development, & while the greater the amount of moisture, the shorter the
period. During this stage the last nymphal skin splits across the head and along
the dorsum, and is slowly worked downward and backward till a large portion of it
hangs freely from the apex of the abdomen on the ventral side. The legs are the
last part c of the body to be freed from this skin, which then becomes detached as
a much crumpled mass. As soon as the wings are liberated they begin to move
away from the body at their base. This is apparently due to the tracheae in the
basal portion of the wing becoming inflated. The inflation, however, does not
extend beyond the suture along which the wing is subsequently broken off, and
the distal portion remains tightly folded. * * *

The ecdysis described above is the last in the development of the imago, for the
insect now disclosed is the sexually complete d adult; it does not, however, become

a Varying from about 3§ to 12 hours at Falls Church, Va,— T. E. S.

& A considerable amount of moisture apparently is essential to normal develop-
ment. Specimens placed in small, individual, corked vials molted normally, while
others placed in vials the mouths of which were lightly plugged with cotton developed
abnormally, with distortions, or not at all.- — T. E. S.

c Sometimes the antennae are the last part to be freed of the cast skin in case of
nymphs of both the first and second forms. — T. E. S.

d At this stage the sexual organs are not yet fully functionally matured.— T. E. S.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate X.

Leucotermes flavipes.

a and b, Lateral views of the quiescent stage of nymphs of first form; c, dorsal view of active
molted nymph showing how wings are held away from the body. Photographs from ether-
ized specimens, a', Lateral view of quiescent stage of nymphs of the first form (skin cast);
b', dorsal view of same; e', active molted nymph of first form with wings unfolding. From
alcoholic specimens. (Original.)

Bui. 94, Pa-tl

O £
o -

U — ,

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 39

active as soon as it emerges but remains for about a quarter of an hour in the same
position as that in which the ecdysis occurred. During this time, however, the
head is slowly drawn upward to its normal position and the insect finally struggles
to its feet. Its movements are at first very awkward and uncertain but after a few
minutes it is actively running about. As before mentioned, the greater portion of
the wings remains closely folded together so that at first sight they appear as abnor-
mally placed wing-pads. A close examination with a hand lens shows them to con-
sist of the very compactly folded wing. * * *

After these young adults have been running about for an hour or so the main portion
of the wing begins to expand; the basal portion becomes expanded before the apical
part begins to unfold, but the infla-
tion gradually works toward the apex
till the typical fully winged though
pigmentless adult is produced. The
wings continue to be held away from
the body till this process is complete,
after which they are folded from the
base in an overlapping position over
the abdomen. The ensuing pigmen-
tation of the body is gradual a and
does not appear to be affected by the
presence or absence of light; the en-
tire body turns black through shades
of yellow and brown till in about
twenty-four hours the sexually com-
plete imago is ready for swarming.

It will be seen that the whole pe-
riod intervening between the normal
nymphal stage and the typical pig-
mentless adult staged occupies only
some nine to ten hours, and this ap-
parently accounts for its not having
been recorded before, even though it
appears to be perfectly normal, c for
it has occurred in different localities
in two successive years and all nymphs
taken passed through these stages be-
fore completing their development.

An illustration of the thorax of Leu-
cotermes flavipes with unexpanded
wings was given by Packard in his
Text-book of Entomology, but he
here described it as a late nymphal
wing pad, otherwise there seem to be no references to either of the stages herein fig-
ured and described.

Fig. 11.— Tcrmopsis angusticollis: Quiescent stage of
nymph of the first form. Enlarged 7 times. (Original.)

a The borders of the chitinized parts first take on pigmentation, passing through
shades of gray to castaneous to the castaneous-black of the mature adult, the abdomen
being the last to take on pigmentation; there are often abnormalities in development —
T. E. S.

b The whole period intervening between the fully developed nymph and the
mature pigmented adult is about one day and one-half to two days for individuals, and
about 1 week to 10 days for the colonies.— T. E. S.

c These stages are absolutely normal and necessary to the progressive development
of nymphs of species of both the genera Leucotermes (I. Jlavipes and virginicus) and
Termopsis (angusticollis) (fig. 11). — T. E. S.
62896a-^Bull. 94, pt 2—15 — -3.

40 INSECTS INJURIOUS TO FORESTS.

During this final molt the females of nymphs of both the first and
second forms normally lose the genital appendices;0 that is. the
genital appendices are present in nymphs of both sexes before this
molt, but afterwards only in mature winged males and supplementary
kings, developed from nymphs of the second form; these appendices
can be readily seen on the cast skins. In egg laying complementary
queens of Termopsis angusticoUis (with no indication of wing pads),
genital appendices were present, though absent hi true queens.

On April 25, 1912, at Falls Church, Ya., molting nymphs (flavipes)
of the second form were found. The progressive development of
over 100 nymphs was noted, and apparently the molting is similar to
that of nymphs of the first form. Before the quiescent stage is
reached, the nymphs assume a straw-colored hue. Various stages of
molting nymphs were observed through April 28. After the molt,
the head and prothoracic segments darken, especially on the borders.
The abdominal segments also darken. Sometimes there are gravish-
black, longitudinal, pigmented markings on the head. Some of the
nymphs have very short wing pads, mere buds, while in others the wing
pads are more elongate, the pair on the metathorax reaching down to
and slightly overlapping the (apparent) second abdominal tergite.
This difference can be seen in supplementary or neoteinic reproductive
forms and illustrates the fact that the growth of nymphs of the first
form is arrested in various stages of development, as does (?) the
presence of genital appendices in complementary queens of Termopsis.

Sometimes, due to unfavorable moisture conditions, there is an
abnormal development of nymphs of the first form. Individuals
may be observed with partial pigmentation to the chitinized parts,
acquired before the quiescent stage or before the wings are unfolded;
that is, the wings may be in various stages of development, from the
opaque-colored, elongate whig pads to partially unfolded, or unfolded
but still crumpled, wings. There is great individual variation in
the manner of molting. Adults with mature body pigmentation but
with distorted, poorly developed wings, or even with opaque wing
pads, have been observed emerging from the parent colony at swarm-
ing time. A swarm6 of flavi pes occurred at Falls Church, Ya., on
May 8, 1912, and another on April 25, 1913, while in the case of
virginicus the swarm occurred on June 1, 1913.

This quiescent stage, or uEuhestadium," has been described by
Hagen.c He states (in regard to the final molt of nymphs of the
first form) that the skin bursts on the prothorax: and in order to

a Grassi and Sandias. Op. cit., p. 306.

& It is from 7 to 10 days after the last sexed adults in colonies have acquired wings
and mature pigmentation that the swarm occurs.

c Hagan, H. A. Monographie der Termiten. Linnaea Entomologica, v. 12,
p. 337-338, 1858. - . :

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 41

get out of the old skin, the insect doubles up. The insect lies on
the ground during the molt. Grassi, while he did not describe this
stage, knew that during certain ecdyses in the development of nymphs
and soldiers important changes took place. Odenbach, January 13
to 24, 1896, observed in an artificial nest indoors (manuscript notes)
molting larvae oi flavipes in the quiescent stage, as if dead. He states
that the molting process lasts three and one-half hours, that workers
assisted, and that the skin is eaten. His observations are prac-
tically the same as those of Strickland. Holmgren describes this
stage in the larva of Rhinotermes taurus Desneux, Escherich a figures
larvae of Termes obscuripesWeLsmsum, and Bugnion5 figures a soldier
of Termes liorni Wasmann in this quiescent stage. Holmgren was
the first to state that a quiescent stage occurs in connection with
each molt, and to note the internal as well as external changes that
occur during these molts.

The writer has observed quiescent stages of undifferentiated (?)
larvae, larvae of nymphs of the first form, nymphs of the first and
second forms, and larvae of workers and soldiers of Leucotermes
flavipes and L. virginicus and soldiers and nymphs of the first form
of Termopsis angusticollis .c Differentiated nymphs of the first
form of L. virginicus only 2.5 mm. in length have been observed.
Bugnion states d that since he has found nasuti larvae of Eutermes
lacustris Bugnion 1.3 mm. in length he believes that the differentia-
tion is effected in the embryo for the three castes. The young
nasutus with the distinct ucorne frontale" is figured. This is not
at all in accordance with Knower's statements and drawings of the
development of the nasutus of Eutermes (rippertii ?) which developed
from a worker-like larva, and with Grassi's description in Calotermes
flavicollis Fabricius and Leucotermes lucifugus and the writer's de-
scription in L. virginicus of the development of soldiers from worker-
like larvae. Bugnion further states e that in the higher termites
the differentiation of caste reaches perfect expression.

Observations by the writer of molting soldier larvae of Leucotermes
spp. show that the differentiation takes place during a " quiescent
stage '-' rather late in the life cycle.

From the first to the middle of August, 1913, freshly molted pig-
mentless soldier nymphs of flavipes in the stage preceding maturity
were noticeable in colonies in Virginia. From the middle of June

a Escherich, K. Op. cit., p. 43.

& Bugnion, E. Le Termes Horni Wasm. de Ceylan. Rev. Suisse Zool., t. 21,
no. 10, p. 299-330, pi. 11-13, juin, 1913. See p. 305-309.

c Snyder, T. E. Loc. cit.

d Bugnion, E. Les termites de Ceylan. Le Globe: Memoires Soc. Geog. Geneva,
t. 52, p. 24-58, 1913.

« Bugnion, E. La differenciation des castes chez les Termites [Nevr.]. Bui. Soc.
Ent. de France, 1913, no. 8, p. 213-218, April 23, 19.13. See p. 217.

42 INSECTS INJURIOUS TO FORESTS.

to the first part of July, 1914, molting large-headed soldier larvae
and nymphs of Leucotermes Jlavipes were found in colonies in Vir-
ginia. After the first part of July freshly molted, pigmentless
nymphs of soldiers were common in colonies. On August 17, 1913,
molting soldier larvae were found in the quiescent stage in a colony
of virginicus at Chain Bridge, Va. During the quiescent stage differ-
entiation took place. Larvae to all external appearances undiffer-
entiated or of the worker type (as shown by the head, mandibles
with marginal teeth, and labrum of the still adhering larval skin),
the individuals (virginicus) being over 3 millimeters in length, in
the quiescent condition, develop at this molt to pigmentless nymphs
of soldiers (PL IX, a), with more elongate, soldier-like head, and
mandibles without marginal teeth. In this stage the head, mandibles,
labrum, and "menton" (Bugnion) have not attained the shape or
length of those of the mature soldier, there being at least one later
molt to maturity. After the first radical change, the head is not
pigmented (the only pigmentation being at the inner margin at the
tips and base of the mandibles, and at the tips of the maxillae) , not
elongate, rounded, tapering toward the base (broad at apex), the
mandibles shorter and broadening at base. The labrum is elongate,
subelliptical, tapering at apex and slightly at base, wider than in
the mature form; the "menton" is convex, tapering toward base,
wider than in the mature form. The antennae have 14 segments.

After the next molt the nymph is as yet shorter than at maturity,
being from 4.5 to 5.5 millimeters in length in Jlavipes, and the head is
more elongate but still broader at the apex, with the mandibles,
labrum, and u menton" more elongate and slender. The antennas
have from 14 to 15 segments. At this stage the mouthparts and
borders of the antennal sockets are slightly pigmented. (PI. IX, b.)
After another molt the full size of the mature nymph is attained
and there is pigmentation of the chitinized parts (PI. IX, c), there
being three molts from the large-headed, worker-like larva to the
mature pigmented soldier. (PL IX, d.)

It will be noted that there is a gradual elongation of the parts, as
the mandibles, labrum, and "menton/ ' and that these parts become
more slender and loose in width as the mature form is reached. a In
this connection it might be of interest to state that in the neoteinic
individuals (neoteinic reproductive forms with short wing pads) the
head, thoracic segments, and abdominal tergites and sternites are
both longer and broader than in the reproductive forms that develop
from nymphs of the first form; that is, the structure at this younger
retarded early stage is more gross.

a Snyder, T. E. Changes during quiescent stages in the metamorphosis of termites.
Proc. Ent. Soc. Wash., v. 15, no. 4, p. 162-165, pi. 6-7, Dec., 1913.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 43

Freshly molted, immature, pigmentless nymphs of soldiers of
Termopsis angusticollis , obtained from material collected by B. T.
Harvey at Ashland, Oreg., August 28, 1913, show the same stages of
development. In conclusion, therefore, it may be stated that in case
of Leucotermes spp. and Termopsis angusticollis the differentiation
occurs during a molt and quiescent stage rather late in the life cycle
of the insect, the larvae having belonged to all external appearances
to the undifferentiated group.

SEASONAL VARIATIONS IN THE COLONY.

There are wide differences in the composition of colonies of Leuco-
termes at various seasons of the year.

EGGS.

In well-established colonies of flavipes, eggs and newly hatched or
young larvae have been found by Hubbard, in Florida, in April, May,
and June; by the writer, in Virginia, in April, May, June, July, August,
September, and October; in Illinois, in August; and by Odenbach, in
Ohio, in September. In incipient colonies, the first eggs are laid, in
Virginia, by the middle of June or July in case of flavipes and in July
or August in the case of virginicus. In well-established colonies of
virginicus, in Virginia, eggs have been found in late May and early
June. In cases of artificial nests or colonies of flavipes, kept under
observation indoors, and in cases of infested buildings, eggs have
been found by Odenbach and the writer in January, February,
March, April, May, July, August , September, November, and De-
cember. Odenbach's nests were from South Brooklyn, Ohio.

In case of both flavipes and virginicus, in long-established colonies
the period of maximum egg production is from the middle of May to
early September in Virginia; that is, during the warm months.

NYMPHS OP REPRODUCTIVE PORMS.

Nymphs of the first form of flavipes with easily discernible wing
pads were found in the quiescent stage of the molt on August 5, 1913,
at Veitch, Va. By the middle of September they had well-devel-
oped wing pads — two and one-half times the segment from which
they originated — and antennae with from 16 to 17 segments. The
nymphs were from 5.5 to 6.5 millimeters in length and the compound
eyes had acquired the primary reddish pigmentation. There is
apparently an unaccountable annual variation, since in 1913 and
1914 nymphs were very abundant in colonies in the fall, while in
1912 they were comparatively scarce. In March, up through the
time of molting, nymphs of the first form are present in the out-
lying channels of colonies in great numbers.

44 IXSECTS INJURIOUS TO FORESTS.

In incipient colonies nymphs are not produced during the first year
that the colony is established; but are developed every year in old,
well-established colonies. Nor are nymphs of the first form produced
the first year in "orphaned'' colonies. Nymphs of the second form
are commonly to be found during the latter part of April to early
May in Virginia (March in Korth Carolina), and they occur in vary-
ing numbers associated with nymphs of the first form. Xymphs of
both forms are well developed by the middle of September of the year
in which sexual maturity is attained, in jiaripes.

Colonizing individuals of Jiaripes appear from the early part of
April to May in Virginia and Maryland (earlier in infested bundings).
In the case of virginicus they do not appear till a month later, or the
end of May or early June in Virginia and Maryland. After having
attained the mature pigmentation they soon swarm.

The royal individuals oijiavipes — the pahs of winged sexed adults
that have swarmed — are to be found together in the royal cell. In
incipient colonies of termites, unlike the other social insects, the male
assists in the establishment of the colony and continues to cohabit
with the queen, there being repeated coition.

XEOTEINIC REPRODUCTIVE FORMS.

Xeoteinic reproductive forms are normally( l)a developed from
nymphs of the second form after the swarm: the males continue to
cohabit with the females. Immature neoteinic reproductive forms
are to be found at the time the winged sexed adults are attaining
mature pigmentation — the end of April in Virginia. Maturity is
probably attained shortly after the swarm, namely. May to June,
although mature neoteinic reproductive forms have been found before
the swarm.

In case of virginicus, neoteinic reproductive forms, produced from
nymphs of the second form, are matured, fertilized, and egg laying
by July to August. In case of flavipes, these reproductive forms
are matured by May to June. There is a seasonal variation.

WORKERS.

Workers are always present hi colonies except those just estab-
lished by colonizing sexed adults: they are permanently present in
the colony and constitute the most numerous caste.

a These forms can be produced at any time necessary. On April 9. 1912, a decaying
stump infested with jiaripes was removed from the forest and placed in the termi-
tarium; this was at a time when the nymphs of the first form were nearly mature.
On November 18 the termites had entered the ground and two neoteinic ergatoid(?)
forms were found about 3 to 4 inches below the surface in chambers in the earth. Xo
nymphs of the first form were present. They could be distinguished from the workers
by the larger size, distended abdomen, straw-colored pigmentation, and sharper seg-
mentation of the abdomen. Rudimentary wing pads were present. The antennae
had 15 to 16 segments. Workers solicited drops of liquid from the female by stroking
the abdomen with their palpi.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 45

SOLDIERS.

Soldiers are also always present in colonies, except those just estab-
lished by colonizing sexed adults, usually a few being present in
incipient colonies; they are always relatively much less numerous
than the workers. From the first to the middle of August, 1913,
freshly molted, pigmentless soldiers of jiavipes were common in colo-
nies in Virginia, where they were found as late as the middle of
October. On August 17, 1913, near Chain Bridge, Va., molting
larvae and nymphs of soldiers of virginicus were found.

LOCATION OF THE COLONY IN WINTER.

By the middle of November to December, depending upon the
season, termites retreat to the subterranean passages of the colony,
the earth under infested logs being riddled by a labyrinth of galleries.
In case of very large logs, termites may remain in the more impene-
trable inner galleries in the heartwood. In Virginia they remain in this
retreat until the last of February or first or last part of March, depend-
ing on weather conditions.

Indeed, the center of activity in termite colonies changes with
the seasons, due to the varying needs of the colonies as to condi-
tions of warmth and moisture, which are essential to life and devel-
opment. In spring when there is abundant moisture, open, wooded
southern exposures are favorable, and the outlying galleries of
colonies are teeming with developing nymphs, whereas during the
heat of summer conditions would be too dry. Consequently in
summer termites bury themselves more deeply in the wood or earth
in less exposed galleries, in moist, shady sites, and in winter usually
enter the ground to escape the cold. a In autumn, developing larvae
of the castes, and nymphs of soldiers and sexed adults, are to be
found in the outlying galleries where the warmth will enable more
rapid development. Therefore colonies apparently depleted at cer-
tain seasons, at others will be infested. Again, the Leucotermes
colony readily migrates, and the site is liable to abandonment if
conditions become unfavorable. A single colony may extend to
and inhabit several adjacent stumps or trees, and it is often impos-
sible to define the limits of a colony or the line of demarcation be-
tween different colonies in a region where termite colonies are abund-
ant and there are many decaying stumps or logs; hence what is
apparently a separate colony may be only a branch connected with
the main colony by subterranean passages. If colonies are cut into
and the reproductive forms removed, the colony quite frequently
abandons the nest. The reproductive forms are capable of move-
ment, and it may be that old colonies branch out by means of neo-

o As the higher altitudes are attained, termite colonies in the earth under stones
are more common; that is, in the North Carolina mountains and in canons in Arizona.

46 INSECTS INJURIOUS TO FORESTS.

teinic reproductive forms which eventually establish new colonies.
However, this is only a theory, but otherwise what becomes of the
large number of nymphs of the second form in colonies in the spring ?
Surely they are not needed in the parent colony any more than the
winged sexed adults, and it may be that they are impelled to leave
the colony by the same irresistible force that induces the swarm.
However, it is probable that workers and soldiers accompany them
from the parent colony and that by means of subterranean passages
they establish the new nest. Indeed, these forms may be the nu-
cleus of the small bands of foraging workers and soldiers infesting
decaying branches mentioned frequently in literature. The alterna-
tive is, of course, that such bands become isolated from the parent
colony and rear the reproductive forms. It seems that both meth-
ods may be possible and necessary.

DURATION OF DEVELOPMENT AND LIFE.

The eggs of jlavipes hatch in about two weeks after they are laid.
Workers developed from eggs laid on July 15, 1912, were 4.5 mm. in
length by the following December, with 13 segments to the antennae.
Both workers and soldiers complete their development within one
year.

Definite data on the duration of life of any individuals of Leuco-
termes, not excluding even the royal pair, are lacking. However, the
males or true kings of jlavipes continue to live with the true queens
after copulation, the royal individuals probably living at least Rre
years ;a neoteinic queens live at least one year and probably as long as
true queens. Smeathman conjectures that a queen of Termes belli-
cosus Smeathman when 3 inches long is about 2 years old.

CANNIBALISM.

There are many instances to show that termites are cannibalistic in
their habits; all dead or injured individuals are eaten; also, according
to Odenbach, larvas that have difficulty in molting.

It is not at all rare to find, especially in cases of workers, a narrow*,
grayish band, with black, scalloped, turned-up edges usually on the
dorsum of the abdomen, but also sometimes present on the ventral
surface and as a plate on the legs. This band is sometimes present
on the abdomen of soldiers, and it also occurs on the thorax and
head of workers. Possibly these black bands are healed-over wounds
where the insects have bitten one another (fig. 9, c) , or they may be due
to a bacterial or fungous disease, or to both wound and disease.

These bands occur on workers and soldiers of both jlavipes and
virginicus.

a Heath, H. The longevity of members of the different castes of Termopsis angus-
ticollis. Biol. Bui., v. 13, no. 3, p. 161-164, August, 1907.

Jul. 94. Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate XII.

Leucotermes flavipes.

Fig. 1.— a, Abandoned burrow of Lymexylon sericeum in solid wood of chestnut telegraph pole,
in which a fertilized true queen was found; b, cell excavated in decaying wood by young
royal couple.

Fig. 2.— Royal cell in solid chestnut in which 40 neoteinic royal individuals, for the most part
queens, were found. (Original.)

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 47

On August 12, 1914, several workers with these bands on the body
were taken from a colony of flavipes at Falls Church, Va., and placed
in a small tin box with decayed wood and earth. Normal soldiers
were also placed in the box. On October 2, 1914, the workers with
the black bands were still alive and apparently in the same condition;
the soldiers had no bands.

Chanvallon, according to Hagen,a recommends placing arsenic in
termite nests, and since the insects are cannibals and the dead are
eaten, a large number can be killed in this manner.

SITUATION OF THE DIFFERENT FORMS IN THE NEST.

The reproductive forms are not necessarily to be found in a ll royal"
cell situated in the more remote parts of the nests, as in tropical
species, but are usually in the more sound or solid wood (PI. XII,
figs. 1, a and 2). In colonies recently established by colonizing indi-
viduals the eggs and young are present in a definite royal cell, where
they receive the care of the queen. In well-established colonies no
forms are permanently present with the reproductive forms, and there
apparently is no well-defined royal cell.

The royal cell, excavated in decayed wood by the sexed adults
that have swarmed, is a broad, oval chamber, the entrance to which
is but slightly larger in diameter than the abdomen of the queen at a
period 14 months after swarming (PI. XII, fig. 1, 5).

Most of the 40 neoteinic reproductive forms found at Falls Church,
Va., May 27r 1912, were congregated in a single chamber in the
solid wood of a chestnut slab. This chamber was a broad but shallow
longitudinal cell in the solid, sound wood. The entrances to this
chamber were but slightly larger in diameter than the abdomens of
the fertilized queens. Other neoteinic individuals were found in
shallow, broad, oval cells in the wood and in earth under the slab
(PI. XII, fig. 2).

The nymphs are usually present in the more remote passages of the
nest, except during the spring, when they are in the outlying channels,6
where the warmth of the sun will hasten their development.

a Hagen, H. A. Monographie der Termiten. Linnsea Entomologica, Bd. 10,
1855, p. 35.

b Developing larvae, nymphs, or immature adults are normally to be found, tem-
porarily at least, in that part of the nest where there are the most favorable conditions
of heat and moisture for their rapid development — changing with the seasons. In
the spring and autumn these forms occur under the bark on decaying stumps and
under decaying wood or bark sunken in the ground in open sunny sites, always being
in the outlying galleries where the warmth will enable more rapid development.

There are apparently no permanent sites used as "nurseries," as is the case in
tropical species of termites. However, young larvae are seldom found in the main
unpartitioned runways, but rather in partitioned galleries, where they will not be
disturbed by the activities of the other members of the colony. Often they are in
broad but shallow unpartitioned galleries in the more sound wood of the interior
heartwood of logs, etc.

48

INSECTS INJURIOUS TO FORESTS.

Unpartitioned channels or runways usually contain only workers
and soldiers; the partitioned channels contain larva? and nymphs
besides. The young, freshly hatched, developing larva? are often
found in broad but shallow moist galleries in the interior of the
heartwood. Extensions of an old colony, consisting of subterranean
passages leading to small pieces of decaying wood, contain only
workers and soldiers.

As previously stated, during the winter all the castes seek the more
remote or subterranean passages of the nest.

THE SWARM OR SO-CALLED NUPTIAL FLIGHT.

Under normal conditions in the Southern States.0 in early April
and May, in the case of the more common species, flavipes, and in
June in the case of virginicus, the colonizing individuals emerge in

en ormous numbers from
small holes in the wood
of stumps, logs, poles,
fence posts, foundation
timbers in buildings,
and the roots of trees
and from the ground.
It is 7 to 10 days after
the last sexed adults
have acquired wings
and mature pigmenta-
tion that the swarm
occurs. The winged in-
sects usually crawl upon
some elevation before
taking flight: before
swarming they teem
over the tops of infested stumps (fig. 12) and festoon brush lying on
the ground in order to get a start. If a sexed adult loses its wings
while at a height above ground (as on the top of a stump) it jerks
itself up in the air in endeavoring to get down. Numerous workers
and soldiers are congregated in the outer layers of the wood near the
exit holes at the time of emergence. These colonizing individuals
differ from the other soft-bodied castes in that they are larger, of a
castaneous color, and are highly specialized and developed for the
purpose of swarming and starting new colonies. In addition, they
are sexed and have eyes and wings. While it is true that they are
weak fliers and are preyed upon by many insectivorous animals —
birds, lizards, insects, etc. — yet some escape to found new colonies.

a The first swarm which contains the greatest number of individuals does not
occur outdoors until the ripening of the pollen of the flowers of dogwood ( Cornus
ftorida), which is also influenced by seasonal and geographical variations.

Fig. 12.— Leucotermes flavipes: View of a swarm of sexed adults
emerging from a stump at Palls Church, Va.; a portion of the
enormous numbers constituting a swarm. (Original.)

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 49

After the adults have flown a short distance in an irregular
" wobbly" manner, they fall to the ground, and by catching the
tips of the wings against some object and turning sideways they
pry them off at a suture or line of weakness near the base, leaving
stubs. This triangular, basal portion of the wing, or stub, is thick-
ened and more chitinized than the wing and is also pubescent up to
the suture, a possible aid in breaking off the wing after flight.

The male follows the female tirelessly and persistently, with head
close to her abdomen, and touches her abdomen with the antennae.
Often the male and female run in a circle of small diameter, and
sometimes the pursued turns pursuer, apparently attracted by some
secretion at the posterior end of the body.a Sometimes as many
as three individuals may be seen running off together. This is
apparently due to sexual attraction, an amatory procedure pre-
liminary to pairing, which accomplishes the purpose of bringing
the sexes together. This continues for several days after the flight.
The sexes are attracted to each other at a period several days before
swarming, as is evidenced by the fact that when a colony is broken
into there is a short flight, followed by loss of the wings, after which
the male follows the female in the same manner as after normal
swarming.

Neither of the terms " swarm" or " nuptial flight" is appropri-
ate in referring to the emergence of the colonizing sexed adult
termites, since the insects after a short flight separate into pairs,
or the males and females may even "pair" (but do not "mate"
sexually till later) with individuals of other colonies, and never con-
gregate again in the same colony, but form many new colonies. In
the case of bees, on the contrary, after the swarm subsides the insects
all together form one new colony. Furthermore, copulation does
not take place at the time of the swarm, which is not a "nuptial
flight."

THE ESTABLISHMENT OF NEW COLONIES.

Many investigators have considered that the foundation of new
colonies by winged sexed adults was impossible, and was not the
purpose of the swarm, but Ferris (1876), Perez (1894), and Heath
(1903) disproved this.

For several days after swarming the now wingless sexed adults
can commonly be found together under small pieces of decaying
wood, lying on the ground, ultimately disappearing, either to excavate
shallow cells in the ground, in decaying wood (PI. XII, fig. 1, b), or to
take possession of an old abandoned insect burrow. The entrance
to this now royal cell is but slightly wider than the abdomen. It

a Heath, H. The habits of California termites. Biol. Bui., v. 4, no. 2, p. 47-63,
figs. 3, January, 1903. See p. 54.

50 IXSECTS IXJUEIOrS TO FORESTS.

is here that copulation probably takes place. Doubtless many
colonizing pans, even after escaping then numerous enemies at
the time of the annual swarm, fail to become established in a new
colony owing to unfavorable moisture conditions. Sometimes sev-
eral pans are found together in the same cell in a piece of wood, or
perhaps one male and two females, or vice versa, but these repro-
ductive forms, unlike the neoteinic forms, are not normally polvg-
amous. The young royal couple, after finding suitable shelter, for-
age for themselves and the abdomens of both sexes increase sliehtlv
in size, becoming swollen. This is probably due to feeding a and
development of the sexual organs.5 Heath states that "everything
is apparently sacrificed to lightness of body" at the time of the
swarm. This results in wider dispersal.2 Copulation probably
does not take place till about a week after the swarm, when the
couples are established together in the royal cell, as sexed adults
of Jlavipes that swarmed on May S. 1912. were in royal cells on
May 15. the male no longer f oho wing the female about.

While it is not essential that the colonizing pairs, the young kings
and queens, be adopted by foraging workers and soldiers, it is pos-
sible that this sometimes occurs. It is significant that small branch
colonies of workers and soldiers are to be commonly found under
decaying pieces of wood and in the ground after the swarm.

COPFXaTIOX AXD THE RATE OF EGG LAYTXG.

Actual copulation was not observed during these investigations,
but observations indicate that copulation does not take place till
after the male and queen are established in the royal cell, and copu-
lation at the time of swamiing outside the nest is not very probable,
as the genitals of the males are in a very imperfect stage of devel-
opment.0 Other observers have noted the process in the case of
Jlavipes.

Haviland states. d "'In Ttrmes malayanus I have reason to think
that the king fertilizes the eggs after they are laid: indeed, copu-
lation in the case of kings and fully grown queens of most species of
the genus Termes is apparently impossible." [ ?]

a Heath. H. Loc. cir.

b M tiller. Fritz. Beitrage zur kenntniss der Termiten. Jenaische Ztschr. f. rued..
Bd. 7. Heft 3. p. 333-35S. fig:. 11. pis. 19-20. Marz 7. 1S73. See p. 337-351.
I. Die Geschlechtstheile der Soldaten von Calotermes.

II. Die TVohnungen unserer Termiten. p. 341-358.

c Hagen. H. A. Some remarks upon white ants. Proc. Boston Soc. Xat. Hist.,
v. 20. p. 121-124. December 4. 1878.

d Haviland. G. D. Observations on termites, with descriptions of new species.
[Read 3d June. 1897.] Jour. Linn. Soc. ivLondon •. Zool.. v. 26. p. 355-442. pi. 22-25.
April 1. 1898.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 51

Grassi states:® "On 'April 17, 1891, about 11 a. m., I detected the
king and queen in coitu in a glass containing a small Calotermes
nest * * *. They stood end to end in a straight line with the
tips of their abdomen applied to each other * * *. It is there-
fore certain that the connection takes place in the nest, and is re-
peated at intervals." Sandias states5 that he observed what
appears to be a similar process between substitute forms still far
from maturity, being only about a fortnight old.

The Rev. F. L. Odenbach, of St. Ignatius College, Cleveland, Ohio,
has noted (MSS.) the copulation of neoteinic royalty (Leucotermes
flavipes), namely, the mating of supplementary queens, time and
again in artificial nests. On March 11 and 29, 1898, he observed
many sexed adults pairing, coitu lasting about three minutes. He
further states that the same queen has connection repeatedly with
different males. On December 27, 1899, in describing the pairing
of neoteinic royalty with short wing pads, he states: "The introduc-
tion is a lively play with feet and feelers, heads looking in opposite
ways, the bodies curved together so as to make a circle, then the
male slips along the body of the female until the organs meet; then
they stand in one line, heads looking in opposite directions. The
body is moved backward and forward, hinging on the legs and
finally to both sides, as if they wished to telescope the abdomens.
Time of connection, about one minute."

Heath states c * * * "almost a fortnight after swarming, I
have on several occasions seen the royal pair of Termopsis in coitu.
With their bodies closely appressed end to end in a straight line they
remain from one to ten minutes in contact."

Egg laying in the case- of flavipes begins about the middle of June
or July, varying with the season, or about one and one-half months
after the swarming. While the eggs hatch in about 10 days after
they are laid, larvae of varying sizes are often present, since they do
not all hatch uniformly. Most of the first broods develop workers
and a few soldiers, as the workers constitute the caste most necessary
to the conduct of the young colony. At this time the queen and the
male still occupy the royal cell together and the queen, with abdomen
only slightly distended, cares for and carries about the eggs and later
the young larvae in her mouth, when the colony is disturbed. The
royal cell is kept clean and the sides are now smoothed.

Recently hatched larvse are fed on prepared food and do not eat
wood until later in their development. On January 8 and 15, 1896,
Odenbach observed workers to draw a white substance of the con-
sistency of butter from the anus of neoteinic queens and devour it.

a Grassi, B., and Sandias, A. Op cit., p. 285-286.

b Ibid., p. 386.

c Heath, Harold. Op. cit., p. 52.

52 INSECTS INJURIOUS TO FORESTS.

On November 18, 1912, workers were observed to solicit liquid from
the anus of a larval or ergatoid (?) queen, at Falls Church, Ya. On
August 5, 1913, a fertilized, fully developed queen, collected at
Yeitch, Ya., about 14 millimeters in length, ejected a clear white
liquid from the anus, when disturbed.

The rate of egg laying of the young and active queen is not very
rapid, as clusters of eggs in varying numbers from about 6 to 12
were observed in several cells with single pairs. The new colony at
first is very small, and even after the rearing of the first brood of
workers and soldiers the increase in numbers is not rapid. In July,
1912, at Falls Church, Ya., about 12 small white eggs in a cluster
were observed in a royal cell with young royal individuals of jlavipes.
At least three were observed, probably 2 males and 1 female. These
had been captured after the swarm on May 8, in the earth under a
small piece of decaying wood. On July 29 the newly hatched larvae
were observed, and on October 30 seven workers and one soldier sur-
rounded a single royal pair. Fragments of the chitinized parts of
another adult were found near the royal cell. The abdomen of an
egg-laying female under observation, 13 months after swarming,
was oblong and somewhat distended, the segments of the abdomen
being slightly separated and showing white between. On October
30, when the female in the royal cell was disturbed, she continually
moved the end of the abdomen, curving it ventrally under the body.
This alternate rising and falling of the abdomen has been described
by Smeathman as a constant "peristaltic" movement, in the case
of tropical species. No eggs had been laid since the first were de-
posited in July, and it is believed that this so-called "peristaltic"
movement in case of Leucotermes is merely the result of alarm, and
has no direct bearing on egg laying. During this time the male still
occupied the cell with the female and both were active. Eventually
the abdomen of the female becomes immensely distended through
the development of the ovaries, but in the case of certain species of
Leucotermes the queen still retains the power of locomotion.

It will thus be seen that development under the foregoing con-
ditions is at best a slow process and not at all comparable to that
which takes place in tropical species, where growth of the queen and
the rate of egg laying is correspondingly rapid.

On February 21, 1913, nine or more additional eggs were observed in
a cluster near the royal cell of the above-mentioned pair. This cell
was in a small decayed branch, placed on moist earth, and isolated
in a tin box. On February 24 the first freshly hatched larva was
observed. The abdomen of the queen at this time was not markedly
distended. On May 16 freshly hatched larva? were again present in
this colony. On August 15 six eggs, as well as newly hatched young,
were present in the royal cell. The male still cohabited with the

BIOLOGY OF THE TEEMITES OF THE EASTERN UNITED STATES. 53

queen, and the abdomen of the queen was not as yet markedly-
distended.

While the recently hatched young are active, they are dependent
on the care of the parents or upon the workers for food.

Wheeler a states, "In incipient ant colonies, the queen mother
takes no food, often for as long a period as eight or nine months, and
during all this time is compelled to feed her first brood of larvae
exclusively on the excretions of her salivary glands. This diet,
which is purely qualitative, though very limited in quantity, pro-
duces only workers and these of an extremely small size (micrer-
gates)." In incipient termite colonies (Leucotermes and Termop-
sis b) the young royal couple share the royal cell, excavated in
decaying wood, at which time the abdomens of both the sexed adults
increase slightly in size, and they take food — that is, wood. The first
larvae develop to workers and a few soldiers, both forms being smaller
than normal individuals or those in well-established colonies. No
nymphs of sexed adults are produced during the first year. The
rate of egg laying of a fully developed true queen is much more rapid.
On August 5, 1913, at 5 p. m., a true queen, about 14 millimeters
long, which had been taken in the root of a dead chestnut tree above
ground, was isolated with the king in a cell in wood. By 9 a. m. on
August 6, more than 12 eggs had been laid. When captured in the
tree there were several hundred eggs as well as numerous recently
hatched larvae near by.

The antennae of some true royal pairs that have swarmed are
apparently entire at a period of seven months after swarming; how-
ever, the segments were not actually counted. In other pairs the
antennae of both sexes are mutilated.

THE ROYAL PAIR AND OTHER REPRODUCTIVE FORMS.
OCCURRENCE IN THE UNITED STATES.

Feytaudc gives a historical account of the frequency of occur-
rence of reproductive forms of L. lucifugus in Europe and figures the
reproductive forms. Between April and September in the eastern
United States the several types of reproductive forms of our common
species of termites are to be found in colonies in decaying wood
above ground; that is, the pigmented, true royal pair with wing
stubs, developed from the sexually mature adults; the supple-
mentary neoteinic forms, with pale straw-colored pigmentation and
short wing pads, developed from nymphs of the second form; and
the ergatoids and neoteinic larval forms, with straw-colored pigmen-
tation and no wing pads or rudiments, developed from mouldable
larvae. It is believed that since these forms are mobile and that in

« Op. cit., p. 68. & Heath, H. Op. cit., p. 57. c Loc. cit.

54 INSECTS INJUEIOUS TO FORESTS,

old, well-established colonies there is apparently no definite perma-
nent royal cell, these forms inhabit the subterranean passages in
wood or in the earth below the frost line during the winter. During
the warm months, probably to facilitate the processes of reproduc-
tion and development of the young, they inhabit the passages in
decaying wood above ground. The occurrence of true royal pairs
is not rare,a but supplementary or neoteinic reproductive forms are
apparently more common in colonies.

HISTORICAL.

The following is a historical record, in chronological sequence, of
the occurrence of reproductive forms in colonies of Leucotermes
jlavipes and L. virginicus in the United States, together with notes on
the conditions under which they were found.

The first queens of Jlavipes taken h in the United States were found
by the late H. G. Hubbard in a colony in Florida and were of the
neoteinic type, or supplementary form, with short wing pads. Hub-
bard also found the first neoteinic kings, although he makes no men-
tion of them and may not have recognized the fact that he had found
both sexes of neoteinic reproductive forms. Some of the neoteinic
queens that Hubbard collected are deposited in the Hagen collection
at Cambridge, Mass., but most of the specimens are in the collection
of the United States National Museum. The number of specimens now
present in the vials at the museum is probably not as great as when
originally collected, since Hubbard gave certain of the royal individuals
to Hagen. Hubbard's note, dated " Enterprise, Fla., May 19, 1875,"
recording the finding of the first reproductive forms in the United
States, reads: " Termes jlavipes (determined by Hagen), females with
their eggs from small, rotten log in road near lake shore; females not
in separate cells, several together." [This vial also contained two
supplementary kings and nymphs of the first form.] Another note
dated April 4, 1882, Crescent City, Fla., reads: "Termes jlavipes,
nymphs, queens, and eggs from galleries in pine log; Trichopsenius
and Anacyptus were found in this nest." [A neoteinic king was also
present in the vial.] A note dated "May 11, 1883, Crescent City,
Fla., in pitch pine," is in a vial containing 11 neoteinic queens, 5 neo-
teinic kings, and 631 eggs with the embryos in various stages of de-
velopment. [Nymphs of the first form were also present in this vial.]

a It was formerly thought that true queens did not exist in colonies in the United
States and Europe (proper). According to E. A. Schwarz (Termitidae observed in
southwestern Texas in 1895. Proc. Ent. Soc. Wash., v. 4, no. 1, p. 38-42, Nov.
5, 1896), there are but few permanent nests, headed by true royalty, of Leucotermes,
due to the wandering habit of the genus; that is, the frequent moving of colonies
would necessitate such rarity.

& Hagen, H. A. The probable danger from white ants. Amer. Nat., v. 10, p. 401-
410, July, 1876. See p. 405.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 55

Mr. Louis H. Joutel,a of New York City, found a number of ferti-
lized, egg-laying neoteinic queens occupying the same cell, 9 in one
colony and 14 in another. The number varies with the colony, as
has since been ascertained, and many neoteinic royal individuals may
be present in a small colony, where they would be more needful. Mr.
Joutel spent several years in the study of termites, and in correspond-
ence with the writer states that on two separate occasions he has
found true queens — the first shortly after the finding, in 1893, of the
above-mentioned neoteinic queens. This would be the earliest record
of the finding of a true queen of flavipes in this country. He further
states that on a later occasion he found two true queens at Peekskill,
N. Y., on the same day, July 14, under about the same conditions.
The queens were located in a a* * * dead hickory stump, about
12 inches diameter, and were in the upper part of the tunnels among
the workers. There was nothing to suggest a queen cell and no eggs
to be found, although I looked over them carefully. They were found
in stumps about 20 to 30 feet apart." In commenting on a statement
made by the writer b that a true queen was inactive in a burrow
when discovered, Mr. Joutel states: "The three [true queens] that I
found were very active, and while they did not move quite as fast as
the larvae [workers], it was due only to their size." Mr. Joutel is quite
right, as I have since found out, and the instance cited was probably
due to the queen being caught in a burrow too narrow for her dis-
tended abdomen, in trying to escape, rather than being confined in a
cell the entrance to which was narrower than the size of her abdomen.
While Mr. Joutel was the first actually to find a true queen, Mr. C.
Schaeffer,a of Brooklyn, N. Y., was the first to record the finding of a
fertilized, true queen of flavipes, at Moshulu, N. Y., July 16, 1902.
He has kindly loaned the specimen to me for study. This queen is
approximately 8 millimeters in length, the abdominal tergites and
sternites not being widely separated, is markedly pubescent, and the
antennae are mutilated.

In describing the true queens which he found, Mr. Joutel states
that they resembled the queen that Mr. Schaeffer found. He fur-
ther states, "* * * they were all apparently broader than the
one you figure in relation to its length. The heavy chitinized parts
looked like little dots on the surface and did not take up as much
space in relation to the rest of the surface as those parts do in the one
you figure. [True, in living specimens.] The first one had its
antennas complete; of the other two, one had two segments missing
on one antenna and the other had three segments wanting —

a Loc. cit.

& Snyder, T. E. Record of the finding of a true queen of Termes flavipes Kol. Proc.
Ent. Soc, Wash., v. 14, no. 2, p. 107-108, June 19, 1912.
62896°— Bull. 94, pt 2—15 1

56 IXSECTS INJURIOUS TO FORESTS.

these happened to be on the same side." He also says, "I have re-
peatedly taken a pair at swarming time and bred the eggs from them,
but have never got them to colony size/7 and that "* * * in
stating that I found only three queens, I had reference to the fully
developed ones. On Staten Island, in company with Mr. W. T.
Davis, I came across a small log that had eight separate cells with a
king and queen in each. A few had three and one four (individuals).
They had eggs but no young. Some had perfect antennas and others
had segments missing." He states: "One small colony that I kept
alive several years that had only about two or three dozen workers
swarmed each year, a few individuals only, but I broke up and exam-
ined each fragment of wood and sifted the ground several times but
never found a trace of either kind of queen; the whole colony was con-
tained in a gallon jar. Other colonies that had thousands of workers,
which I kept alive about 22 months from time of collecting, never
swarmed or had young, but finally died of old age, I presume. They
got smaller and seemed to have more fat in their makeup." Mr.
Joutel also says, "Light, I found, was not objected to by termites,
unless it was too strong, as long as they felt that they were covered,
that is, they worked under cover, not necessarily in the dark." This
is true, but even in metal termitariums with sliding, thick, red glass
covers, termites, while at first they actively wandered about on the
surface of the earth, and were apparently unaffected, soon sought
cover under decaying pieces of wood or in the earth.

The Kev. F. L. Odenbach, S. J., has, since 1895, made observations
on the habits of Leucotermes flavipes in artificial colonies. He has
roughly sketched in his notes two types of neoteinic queens that he
has found. The neoteinic kings he describes as being compressed
laterally, and therefore seeming to have a ridged back. One neoteinic
queen, with short wing pads, that he found pairing was 10 millime-
ters in length and had a markedly distended abdomen. Another
gravid queen that he figures is apparently adapted from a nymph of
the first form, since long, well-developed wing pads are present. The
abdomen is distended and the abdominal tergites are separated. He
states :

These reproductive forms were from a very large nest I took up in South Brooklyn,
near Cleveland, Ohio, in the fall of 1895.

I placed the termites in a large glass globe. In September, 1897. I discovered a
large mass of eggs from which were developed all the different nymph forms known to
me, true winged males and females also in large numbers. These latter chased about
the nest in such wild disorder for a few days that the workers fell upon them and de-
stroyed them to the very last one. It reminded me of the slaughter of drones in a
beehive. At the time it seemed to me that the wild orgies. [?] which as a rule occur
outside of the nest in midair, disconcerted the workers and soldiers who did the next
best thing to restore order and quiet in their household.

From the above large nest I caused a colony to migrate into a Lubbock nest, and
in this nest I found the different neoteinic reproductive forms. First, the nymph

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 57

with the long wing pads. She laid eggs, and I repeatedly induced her to do so by
the same means by which I first caused ant workers to lay eggs (1885).

If they seem numb with cold, I place my hand on the glass plate and this induces
the activity.

This queen was quite different in shape and color from those I will mention below,
being larger and of a lighter color. She was slow in her movements and did not change
her location very often. She was tended by the workers, which could hardly be said
of the others, since they were too restless.

The others, reproductive individuals, were nymphs of different kinds, with different
shaped wing pads, but none with as long ones as those of the individual mentioned
above. They also laid eggs, but during one of their wild rushes around the nest were
given then quietus. I now had only the above nymph and the one with no visible
wing pads left. This latter I thought to be a true queen (?), since the treatment of
her was nothing like that accorded to the nymph with long wing pads. She laid eggs,
but seemed to be disregarded by the workers.

On March 26, 1895, at Haw Creek, Fla., Hubbard found an imago
(a male) with the wings gone, in a colony. The antennae have seg-
ments missing. His note reads, uIn old, rotten pine log in swamp
hummock; soldiers, larvae, and imago with wings gone."

King a also states that he has found a black form (male) with wing
stumps in a colony and that it is a swift runner. "Again, there is
associated with Termes jlavipes a clear black form, variable in size,
some with wing stumps, and others, so far as I can see now, without
being cleared, appear to have none. [?] I have only met with five of
these forms so far; one measured 6 mm. in length, another 5 mm., and
two of these measure 4 mm." He states that the fifth form was sent,
not long since, to Dr. L. O. Howard, with notes, who referred it to
Mr. E. A. Schwarz, the latter stating that it was a fully developed
male with wing stumps.

Mr. King further says :

a * * * When I first observed its appearance with Termes
jlavipes, and in the nest with it, I supposed it to be a species of a
staphylinid beetle, so swift were its movements that they made them
quite deceptive. They are very swift runners and hard to capture.
Further observations will be necessary to determine whether these
are new species or not. It is my impression, however, that they are
of a different type."

He does not state at what period of the year he found this form.

These forms found by Hubbard and King were possibly royal indi-
viduals, and in the case of the specimens collected by Mr. King they
were possibly of both species, jlavipes and virginicus.

H. G. Hubbard found a true queen of Leucotermes, probably luci-
jugus (a species very similar to jlavipes 6). As in virginicus the ocelli
are nearer to the compound eyes than in jlavipes. L. virginicus

a King, G. B. Termes flavipes Kollar and its association with ants. Ent. News,
v. 8, No. 8, p. 193-196, October, 1897. See p. 194.

bHagen, H. A. Monographie der Termiten. Linnsea Entomologica, v. 12, 1858, p.
174-180 and 182-185.

58 INSECTS INJURIOUS TO FORESTS.

Banks has as yet been found only in Virginia, West Virginia (Hop-
kins), Maryland, North Carolina, and Illinois (Snyder). Leucotermes
lucifugus Rossi occurs in the United States and "is found in
Texas, Kansas, Colorado, and southern California, and perhaps else-
where." a The species lucifugus of Mediterranean Europe, according
to E. A. Schwarz,6 is probably native to America (Mexico). Accord-
ing to Dr. Knower, flavipes has been introduced into Japan and is
firmly established. This species has also been introduced into Europe
and has been destructive in the vicinity of Vienna.

This true queen, found by Hubbard, is slightly over 13 millimeters
in length, has the abdominal tergites and sternites more projecting,
and has not as greatly distended an abdomen as the true queen found
in Virginia ; that is, the chitinized parts are less fused than in the older
queen. (PL XIII, b.)

The note in Hubbard's field diary recording the discovery of the
first fertilized true queen reads as follows:

June 20th, 1898, Santa Rita Mountains, Madera Canon (Southern Arizona). We
ascended a ravine filled with majestic sycamore trees under which the ground was wet
with numerous springs, but entirely tramped by cattle and devoid of smaller vegeta-
tion. * * * I found under a stone in a little dry mound in a wet springy spot
on the mountain side, a colony of true Termes, among which, in a cell cavity just
beneath the stone was a single matured gravid female, or queen, which certainly had
been winged; took eggs, larvae, workers, and soldiers with the queen. This is the
first instance known of a true queen in the genus Termes. There were no supple-
mentary or nymphal queens in this colony and no male was found. I explored the
entire colony, which was not a large one.

Prof. Harold Heath, of Stanford University, CaL, has described the
life cycle of Leucotermes lucifugus in California,0 which is very similar
to that of our common species of the East. He figures a true "primary"
queen, as only 8 months old, however, that has a markedly distended
abdomen and the abdominal tergites separated — probably an error,
due to transposed descriptions under the figures, according to obser-
vations by Feytaud d and the writer.

A true queen, with wing stubs (flavipes), found by the writer was
in an abandoned burrow of Lymexylon sericeum Harr. (PI. XIII, b)
in the decaying butt of a chestnut telegraph pole near Portsmouth,
Va., on August 12, 1910. This queen was inactive, since the burrow
was no wider than her abdomen, and apparently she was unattended.
The abdomen was greatly distended and oblong in shape, the ab-
dominal tergites and sternites being widely separated. In the bright
sunlight the abdomen appeared to have a yellowish green tinge.
The length of the queen was approximately 14 millimeters. The
antennas were mutilated.

a Howard, L. O. The Insect Book. p. 356. New York, 1901.

b Loc. cit. See p. 39.

c Heath, Harold. Loc. cit.

<* Feytaud, J. Op. cit., p. 567, fig. 21.

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE XIII.

Fig. l.—a, Supplementary queen with straw-colored pigmentation to chit-
inized parts: b, true queen with castaneous pigmentation to chitinized
parts. (Original.)

Fig. 2. — Supplementary and true queen, showing deeper
pigmentation of chitinized parts of the true queen.
(Original.)

Leucotermes flavipes.

Jul. 94, Part II, Bureau of Entomology, U. S. Dept of Agricultur

Plate XIV.

Leucotermes virginicus and L. flavipes.

, L. virginicus, neoteinic larval reproductive form; b, L. flavipes, neoteinic nymphal repro-
ductive form (queen); cl, c2,L. flavipes, neoteinic larval queens, dorsal and ventral views
(note absence of wing pads). From etherized specimens. Enlarged 10 times. (Original.)

BIOLOGY 'OF THE TERMITES OF THE EASTERN UNITED STATES. 59

Mr. W. B. Parker, of the Bureau of Entomology, has found two
large fertilized true queens of Leucotermes lucifugus in redwood hop-
poles in California. These queens were found in May, one being
approximately 8 millimeters in length, with 11 segments to the
antennae, the other approximately 6 millimeters in length, with
antennae broken. Like the queen found by Mr. SchaefTer, the ab-
domens were not fully distended.

At Falls Church, Va., during the seasons of 1912, 1913, and 1914,
numerous nymphs of the second form, in stages before, during, and
after molting, were found by the writer; also young and mature
neoteinic reproductive forms and young and mature true royal indi-
viduals.

On April 23, 1912, at Falls Church, the first colony of flavipes was
found with a nymph of the second form present. The abdomen
was noticeably broader, and this nymph was active and associated
with nymphs of the first form. The nymphs of the first form were
fully developed and in various stages to the sexed adults with imma-
ture pigmentation. Freshly hatched young larvae were present. The
day was sunny and warm.

On the same day a colony of flavipes was found in a decaying tulip
tree stump, established well in the heartwood. Sexed adults with
mature castaneous-black pigmentation were present and flew when
the colony was disturbed. They were not present in large numbers,
and all were in the outer layers of the wood. No fully developed
nymphs in the stage just before the final molt were present, which is
remarkable, as they were apparently present in all other near-by
colonies. Very young nymphs and half -grown nymphs, as well as
larvae just hatched and young to medium-sized larvae, workers, sol-
diers, and two neoteinic males (mature kings?) were present. They
were active and in the outer layers of the wood. The abdomens
of these males tapered markedly, being narrower at the end of the
abdomen. They had a yellowish brown pigmentation, and in one
king the compound eves were without a trace of pigmentation.
(PI. VIII, fig. 2, a.)

A small colony of flavipes was found on April 25, 1912, under a
small chestnut slab lying on the ground, exposed but sunken com-
pactly into the moist earth. The day was sunny and warm. Fully
developed nymphs of the first form were the most abundant stage,
although some few transforming nymphs and a few sexed adults
with dark pigmentation, as well as a few soldiers and workers, were
present. The next most abundant caste to nymphs of the first form
was nymphs of the second form, of which 90 were present and 4
were molting. They were active, much more so than nymphs of the
first form. Most of these forms were either in shallow tunnels in the
earth or on the earth, but a few were found in the decaying wood.

60 INSECTS INJURIOUS TO FORESTS.

On the same day a single nymph of the second form was taken in a
small colony under a strip of chestnut bark compactly pressed into
the ground; this nymph was associated with nymphs of the first form.
In two other colonies single nymphs of the second form were found
associated with sexed adults under the bark on decaying chestnut
stumps. Normally, but relatively few nymphs of the second form,
as compared to nymphs of the first form, are present in colonies.

In another colony about one dozen nymphs of the second form,
one of which was molting, were found associated with adults with
pigmentation in various stages of development to maturity. A few
of the latter with mature pigmentation had only one pair of wings
unfolded, whereas others were present with the wings at the tips
still compactly folded.

In a near-by colony, nymphs of the second form that had com-
pleted the final molt were present with sexed adults, but unlike the
foregoing they are darker, some being pigmented. Very young
larvae were present in the outer layers of the wood. The acquisi-
tion of pigmentation in neoteinic royalty apparently is a sign of
maturity and old age.

On May 27, 1912, 40 neoteinic forms, for the most part fertilized
queens, together with females in which the abdomen was not greatly
distended, were found in the more solid wood of a decaying chestnut
slab on the ground. (PI. XII, fig. 2.) Smaller forms, males, with the
abdomen not oblong or distended were present in the colony. Most
of these forms were congregated in the longitudinal royal chamber,
in which young were also present. According to Grassi, a in case of
lucifugus, "The colony must therefore rear fresh (complementary)
kings every year [?], which become mature in August and Septem-
ber, fertilize the queens, and die." In colonies in Virginia neoteinic
reproductive forms of jiavipes are being developed from nymphs of
the second form during April and May and are matured by May to
June. In case of virginicus these reproductive forms are matured
by July to August. There is a seasonal variation. There is no
conclusive evidence that the kings do not five as long as the queens.
They are not always present with the queens in colonies, but being
so much more active are more likely to elude capture.

Two sexed adults, a royal couple, were captured by C. T. Greene
at Falls Church, on November 12, in the frass in an old burrow of
Prionoxystus robinise Peck, in the base of a living chestnut tree.
Three royal individuals were present in the frass, and were located
from 2 to 3 inches deep in the burrow. Four to six (at most) young-
were present, and, according to Mr. Greene, apparently all were
workers.

a Grassi, B., and Sandias, A., op. cit., p. 298.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 61

On November 18, 1912, two larval (ergatoid?) reproductive forms
with a yellowish pigmentation and rudimentary wing pads were
found in passages in the ground under a decayed stump at Falls
Church.

Nymphs of the second form greatly outnumbered nymphs of the
first form in a colony of Jlavipes taken in a decaying oak limb on the
ground on March 22, 1913, at Black Mountain, N. C.

On April 17, 1913, at Falls Church, molting nymphs of the second
form of Jlavipes were found together with molting nymphs of the
first form, some of the former of which had already molted and
attained the characteristic yellowish pigmentation.

Molting nymphs of the first and second forms of virginicus were
found on April 30, 1913, at Falls Church, under decaying slabs of
wood on the ground.

Mr. H. S. Barber, of the Bureau of Entomology, on May 30, 1913,
on the Virginia shore of the Potomac River, opposite Plummers
Island, Md., found a neoteinic reproductive form of Jlavipes. This
was a queen exceeding 6 millimeters in length, with wing buds and
17 segments to the antennae. This queen was associated with
workers, soldiers, and young in decaying pine wood on the ground.
(PL XIV, b.)

On May 26, 1913, nine neoteinic queens of Jlavipes with distended
abdomens and one king were found near Charteroak, Pa., in the
interior of a decaying maple stump. Freshly hatched young and
young larvae were numerous. The queens were not in separate cells,
but were active and associated with workers and soldiers. (PI. XV.)

Mr. H. G. Barber, under the title "Another Queen of the White
Ant Found," states :a

While on a collecting excursion last Fourth of July with Mr. Charles E. Sleight, at
Lake Hopatcong, N. J., I found a small colony of white ants beneath a small log-
where they had made some tunnels along the ground beneath the log. Among the
individuals was captured a fully developed queen, which was preserved and presented
to the local collection of insects, at the American Museum of Natural History.

A large fertilized true queen and a king of Jlavipes were found on
August 5, at Veitch, Va., in a large colony associated with workers,
soldiers, and young and several hundred unhatched eggs in the
sound outer layers of wood, about 1 inch in from the exterior, near
the base of a dead standing chestnut tree. This tree had died at
least two years previous, but this may not approximate the age of
the colony, since termites often infest the bases of living trees by
obtaining entrance through old abandoned insect burrows. The
king was hidden beneath the queen, which was about 14 millimeters
in length. The antennae of the queen in this instance were mutilated

a Barber, H. G. Another queen of the white ant found. Jour. N. Y. Ent. Soc, v.
22, no. 1, p. 73, March, 1914.

62 INSECTS IXTUPJOrS TO FORESTS.

and the workers and soldiers were very solicitous. It is evident that
the queen is fed on food prepared by the workers, since her abdomen
became markedly shrunken when she was isolated.

On August 7. 1913. another true royal couple of this species was
found near Chain Bridge. Va.. in the interior of a decayed yellow
poplar (tulip tree log lying on the ground. The colony was small
and the abdomen of the queen was not fully distended, beins: more
hat than oblong. > PI. XVI. b, king. I

On August 15. 1913. at Falls Church, a young neoteinic repro-
ductive forml female of virginicus was found in an experimental
stake of decaying yellow pine which had been set in the ground a
year previous. The queen ( 1 1 was of a dirty yellow color with four
grayish-black longitudinal markings on the epicraniuni. and grayish-
black markings between the coxa? and rues o thoracic andmetathoracic
tergites. The length was 4.5 millimeters. The segments to the
antenna? were mutilated, one antenna having 11 segments. (PL
XIV. a. There was no pigmentation to the compound eyes.

On September 17. 1913. near Chain Bridge. Va.. numerous nymphs
of the second form of flavipes of mature size were found associated
with nymphs of the first form in the sound heartwood of a decaying
black locust stump. The nymphs of the first form had well-de-
veloped wing pads.

On November 3. 1913. at Falls Church, nymphs of the second form
oi flavipes were found associated with those of the first form in the
sound wood of a decayed oak stump near the ground. Young were
present in this colony.

In 1914. on April 17. the first fully developed nymphs of the second
form of Leucotermes flavipes were found in colonies at Falls Church,
Va.

A large true queen oiflavipes was found in a decayed oak stump
about 5 feet high on July 17. 1911. at Falls Church. The tree had
been dead for at least three to four years, and the stump was about 14
inches in diameter and still had the bark on. This queen, which was
oblong but somewhat quadrate — 4 nullirueters in width — was 14j
milhmeters in length (measured while alive a . and in color had a
slight tinge of greenish-yellow or opaqueness. The antenna? had seg-
ments missing. The colony was very large. Numerous eggs were
present in the galleries in the decayed wood, and the queen was
found in an elliptical cell about 2 inches in the wood from the exte-
rior: the sides of the cell were cleanly eaten out. The royal cell was
situated about 1^ feet above the surface of the ground. The male
was not found.

The abdomens of queens become further elongated during the killing and fixing
process, and hence in photographs of preserved specimens there is an apparent loss
in width.

Jul. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate XV.

*'-'••• .#

Leucotermes flavipes.

a, Dorsal, b, ventral, and c, c', lateral views of abdomen of fertilized nymphal neoteinic
queens, a and c', Enlarged 7 times; b and c, enlarged 10 times. (Original.)

94, Part II, Bureau of Entomology, U. S. Dept. of Agricultut

Plate XVI.

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BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 63

In a near-by dead oak tree about 1 foot in diameter, which had
been dead probably for two years, another true queen of this species
was found on the same day. This queen was oblong and approxi-
mately 12^ millimeters in length (measured while living). The royal
cell was in the decayed wood about one-half inch from the exterior
at the base of the tree near the roots. The colony was large in num-
bers. Hundreds of eggs in clusters were in shallow galleries under the
bark. The male was not found. Sometimes eggs in large clusters
were found in deep, transverse, conical ledges or notches, attended
by workers.

When the royal cell is cut into and the queen removed, the large
number of attending soldiers and workers become very much ex-
cited, as evidenced by constant convulsive movements or sudden
jerking of the whole body.

In this same woodland, on July 20, four neoteinic queens of
flavipes were found in the more solid wood of a low (the wood above
ground being nearly disintegrated) decayed oak stump, near the
surface of the ground. One queen was of the normal neoteinic type,
developed from a nymph of the second form, being 9 millimeters in
length, with wing pads present. The compound eyes were pig-
mented, and the antennae consisted of 16 segments. The other three
neoteinic queens were of a type not previously found in colonies by
the writer. These queens, of a straw-colored pigmentation, were
7, 6, and 5 millimeters in length (measured while alive), respectively.
In shape the abdomens are oblong quadrate, like those of the true
queens. The antennae comprise 15 segments, which have a tinge of
grayish-brown pigmentation to the dorsal surface. No wing-pads or
rudimentary buds are present. The compound eyes are without
pigmentation. The head, thoracic segments, and tergal and sternal
"nota" (chitinized tergal and sternal areas) are not as broad as in
neoteinic reproductive forms developed from nymphs of the second
form. In these larval queens, as in true queens, the segments of the
abdomen are less projecting than in normal neoteinic queens, and the
"nota" are less semicircular. The mouth parts and legs are also less
gross in structure. Indeed, these queens more nearly approach the
true queens as to these points. (PL XIV, c.) They were prob-
ably developed from larvae of the sexed forms. The outline of rows
of numerous eggs in various stages of development could be seen
through the body tissue under a high-power Zeiss binocular. The
body tissue of normal neoteinic queens is coarser and thicker. This
is the first time that reproductive forms of apparently different types
have been found in the same colony of this species. The colony of
termites in which these queens were found was small in numbers.

All the queens found by the writer in July, 1914, at Falls Church
were captured within an area of ground less than an acre in extent.

64 IXSECTS INJURIOUS TO FORESTS.

It is believed that the spread or distribution of a colony may be
largely dependent on the supply of decaying wood near by; that is,
if there is a large amount near by (as colonies in large dead trees) the
colony will not branch out over a large area.

On August 5, 1914, at Falls Church, a true royal pair of flavipes
was found in a cell in the more solid wood of a decaying oak chopping
block, that is, a section of a log that had been put to this use in the
woods. The cell was in a knotty area of solid wood about H feet
from the ground but in the outer layers. The king was hiding
beneath the queen, and is 6 millimeters in length. The abdomen is
distended and the antennae mutilated. The queen was of large size
(probably 10 to 12 millimeters in length) but was crushed in cutting
into the royal cell. The colony was large, and galleries extended from
the ground up through this block and another similar block placed on
top of it. The termites had filled in the crevices between the two
blocks with clay, and larva?, pupa?, and adults of Homovalgus sqiia-
miger Beauvois, a scarabseid beetle, were present in the termite
galleries in the clay or in pupal cells.

In the same woods several young royal pans of flavipes were found
established in incipient colonies in the outer layers of a decaying
chestnut slab partially sunken in the ground. Each pair was in a
shallow cell excavated in the wood and was surrounded by a few
young larva? (a half dozen to a dozen) . A few unhatched eggs were
present in some of the cells. In one of the cells three adults were
present instead of simply one pair. This is quite often the case in
incipient colonies.

At Yeitch, Va., on August 12, eight neoteinic reproductive forms
of flavipes were found in a decaying yellow pine stump. They were
in the more solid wood about 1 foot from the ground, but in the
outer layers. Five were females and three were males or kings.
The females were all about 7 millimeters in length, with abdomens
distended with eggs, but with the segments not markedly separated
except near the end of the abdomen, where the latter was lumpy and
distorted, due to distension. The males, with narrow, slender abdo-
mens, were all about 6J millimeters in length, but had, instead of the
straw-colored pigmentation of the females, a darker pigmentation
streaked with grayish-black markings on the head and borders of
the tergites and sternites and between the coxa? and the mesothoracic
and metathoracic tergites. The antenna? of these reproductive
forms consisted of 16 to 17 segments. These forms were developed
from nymphs of the second form. The colony was not very large,
but numerous unhatched eggs and young larva? were present.

At Lake Toxaway, N. C, on August 27, 1914, an ergatoid queen
of Leucotermes virginicus was found under a large flat rock in a shallow
cell in the earth. Workers, soldiers, and young were aggregated

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 65

under the stone, but the colony was not large. This queen had no
trace of wing pads or pigmentation to the compound eyes, if present.
The abdomen is oblong-ovate, being distended; its length is 5 millime-
ters. The antennae are mutilated. Workers and soldiers surrounding
the queen ran up and touched her with the antennae and then evi-
denced excitement and alarm by the convulsive jerking of the body
backward and forward.

This locality is a wooded, rocky hillside at an elevation of about
3,400 feet above sea level. There had been surface fires through the
forest. It was noted that termite colonies were unusually abundant
in the earth under stones in this locality, which is apparently the case
as the higher elevations are reached.

On September 15, 1914, near Chain Bridge, Va., well-developed
nymphs of both the first and second forms of jlavipes were found in a
colony in the more solid wood of a decaying chestnut stump. There
is an annual seasonal variation in the degree of development the
nymphs of the first form have attained by late fall; sometimes the
wing pads are long and the nymphs apparently nearly mature; in
other years in March these nymphs will still have comparatively short
wing pads. There is sometimes also a variation in the different
colonies.

DESCRIPTION OP THE REPRODUCTIVE PORMS.

As has been previously stated, the abdomens of both the young
queen and male increase slightly in size and become distended after
swarming. The abdomen of queens (flavipes) that had laid eggs
during July, in January (9 months after swarming) were oblong and
somewhat distended, the segments of the abdomen being slightly
separated and showing white tissue between. The queens are dark
castaneous in color and the males more blackish. The legs, tarsi,
and tibiae are markedly light yellowish in color in both sexes. After
the swarming the abdomens of the males become only slightly dis-
tended.

The gradual distension of the abdomen of the queen, brought
about by ovarian development, necessitates the separation of the ab-
dominal tergites and sternites, and the connecting tissue between the
abdominal tergites, pleurites, and sternites becomes remarkably dis-
tended, or more probably, according to Hagen,a as stated by Riley,
there is actually further growth after the insect has reached the
imaginal stage. It may be noticed from the illustration of the true
queen found by the writer on August 12, 1911 (PL XIII, b), that
there are two brownish scars located on the pleural tissue, on the right
side of the abdomen. These were not noticed until the queen was

a Riley, C. V. Termites, or white ants. Proc. Biol. Soc. Wash., v. 9, pp. 31-36,
Apr., 1894.

66 INSECTS INJURIOUS TO FORESTS.

removed from the cell, and were probably due to injury received in
shipment, as the queen was not taken from the royal cell, but a small
block of wood was cut out of the pole, the whole having been placed
in a vial of alcohol. The queen was partly out of the cell and the
scars were probably due to abrasions by small fragments of jagged,
projecting wood. This is stated in detail, because it might be
thought that the woimds were "battle scars."'

Attached to the mesothorax and metathorax in true queens are
the stubs of wings, lost at the time of swarming. The head, thorax,
and scutellar area ("nota") of the abdominal segments of true queens
are more heavily chitinized and more deeply pigmented with casta-
neous brown than in the neoteinic queens, developed from nymphs
of the second form, which are straw-colored (PI. XIII, a); conse-
quently, the nonfunctional eyes and ocelli are not so prominent in
neoteinic queens, which never develop wings and which always0 (?)
remain in the parent colony. The head, thoracic tergites, and ab-
dominal tergites and sternites are both longer and broader than in
the true queens, which same differences are apparent in the nymphs.
(PI. VIII.) In true queens the mesothoracic and metathoracic ter-
gites have a distinctively irregular shape. The chitinized ,;nota" of
the abdominal tergites and sternites more markedly approach the
semicircular in shape and are much more projecting in neoteinic
queens developed from nymphs of the second form. However, in
younger (smaller) true queens (flavipes and lucifugus) the tergites
and sternites are slightly more projecting than in older queens.
Furthermore, the legs are more slender and the mouth parts slightly
smaller (less gross) in true queens. The mesothorax and metathorax
and pigmented, chitinized "nota" have a distinctive shape in the
neoteinic larval queens. lPI. XIY, c.)

In matured true queens (of both flavipes and lucifugus) ribbons of
parallel ovules of various sizes and stages of development are visible
(under high-power Zeiss binocular) through the tissue of the abdo-
men, where there are no deposits of fat. Sections through the body
show an enormous ovary development, with ribbons of ovules in
progressive stages of development. In a lateral or dorsal view of the
abdomen of the true queen, small, round spiracles can be seen, set
in at the base of the lateral slope of the tergites. (Fig. 13. a and c.)
The spiracles are approximately similarly placed on queens of spe-
cies in the genera Calotermes Hagen, Termopsis Heer, and Eutermes
Fr. Muller. In some tropical species, as in Termes beUicosus, the
spiracles are located in the pleural tissue of queens with enormously

a It may be possible that snbcolonies or offshoots of large old colonies are estab-
lished by these mobile queens and workers and soldiers by means of subterranean
passages to decaying wood.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 67

distended abdomens, an indication of displacement by actual post-
adult growth ( ?). In tropical species of Eutermes the enormous de-
velopment of the ovaries in old queens crowds the digestive and ex-
cretory organs to the ventral surface of the abdomen.

Situated on the vertex of the epicranium is a small depression
which appears as a rather prominent white spot, under the binocular
microscope, in workers, nymphs, and neoteinic royal individuals of
flavipes. This depression, slightly larger in diameter than an ocellus,
is also present in the soldiers and in colonizing individuals. This

mentioned by Grassi,& as a "* * *

[existing in lucifugus] " (only ?) in the

is the "retrocerebral gland"
[gland of unknown function
nymph of the first form,
the perfect insect, and
the soldier. It elimi-
nates a transparent se-
cretion which can be
squirted out for some
distance."]

While the abdomens
of neoteinic queens ap-
parently never become
as elongate as in true
queens, they become as
much distended, but
have not the oblong or
quadrate shape, being
more oval, or wider
near the end of the ab-
domen, which tapers
markedly. At the end
of the abdomen of true
queens the chitinized
parts are more compressed or fused than in the neoteinic forms.

Fertilized, neoteinic queens, developed from nymphs of the second
form, range in length from 9 to 12 millimeters. The males are of the
same length as the nymphs from which they develop ; their abdomens
are compressed laterally and taper toward the end, and " therefore
seem to have a more narrow ridged back" (Odenbach).

Neoteinic or supplementary royal individuals are obtained from an
arrested early stage in the development of the nymphs of the first

a Homologous to the .small head gland of worker-like larvae of Eutermes, which lar-
vae develop to nasuti, with a nose-like process (?).
b Grassi, B., and Sandias, A. Op. cit., p. 317.

a b e

Fig. 13.— Leucotermes flavipes: Fertilized, true queen; dorsal (a), ven-
tral (&), and lateral (c) views of abdomen. Drawn from specimen
preserved for three years in alcohol. Note position of spiracles.
(Original.)

68 IXSECTS INJURIOUS TO FORESTS.

form, or even larvae, as young are kept in an undifferentiated state,
which can be speedily turned into reproductive forms that serve as
substitutes. Fritz Miiller0 compares the modes of diffusion and re-
production to those of plants which continue the species by means of
cleistogamic as well as perfect flowers, the neoteinic forms correspond-
ing to the cleistogamic flowers which are supplementary, emergency
forms for use in case the perfect flowers (or winged colonizing forms)
should fail. The winged forms would furnish a possible, but not
probable, escape from interbreeding, since, usually, males and females
from the same colony pair. Neoteinic supplementary forms are pro-
duced not only to counterbalance the loss of true royalty but also
for the purpose of extending the colony.

DATES OF THE SWARMING OF LETJCOTERMES.

Colonizing individuals of Leucotermes flavipes usually swarm in
the forenoon during the first part of April and May in the southern part
of the eastern United States. Farther north the swarm occurs later,
usually the last of May or early June. Hagen 6 mentions an excep-
tionally large swarm which occurred in Massachusetts. The sexed
adults normally emerge earlier in infested buildings. According to
E. A. Schwarz, sexed adults swarmed from infested beams in the
floor in the basement of the old United States National Museum on
March 15, 1883, the second year after the opening of the museum.
The following year they swarmed during the latter part of March.

On April 16, 1910, sexed adults, possibly from the same colony,
swarmed from crevices between the bricks in the sidewalk opposite
the old National Museum.

On March 30, 1908, sexed adults came up through cracks in the
floor in a building at Philadelphia, Pa.

Mr. Schwarz states, in an article entitled ''Termitidas Observed in
Southwestern Texas in 1895" : c

Terraes flavipes Kol. —Common throughout southwestern Texas and very destructive
to woodwork in houses. An immense swarm of winged individuals [lucifugus?]
issued from several houses at San Diego [Tex.], on October 25. I was informed that
in early spring another flight takes place in buildings infested by termites [flavipes?].
"What appeai-s to be the same species is also common in sticks and branches lying on
the ground in the chaparral, but I failed to get the winged form from such situations.

a Miiller, Fritz. Beitrage zur Kenntniss der Termiten. III. Die Nymphen mit
kurzen Flugelscheiden (Hagen), "nymphes de la deuxieme forme" (Lespes). Em
Sultan in seinem Harem. Jenaische Ztschr., Bd. 7, Heft 4, p. 451^63, figs. 3. Novem-
ber 18, 1873.

& Hagen, H. Some remarks upon white ants. Proc. Boston Soc. Xat. Hist., vol.
20, p. 121-124, December 4, 1878.

c Proc. Ent, Soc. Wash., vol. 4, no. 1, p. 38-42, November 5. 1S96. See p. 38.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 69

W. D. Hunter states a:

Termes flavipes K. is not uncommon in Texas, where swarms occur ordinarily
during the early part of the season. In 1911, however, the insect did not come into
notice until about the middle of October [lucifugus?]. At that time much more than
usual numbers were to be seen throughout the State.

Often several swarms emerge from the same colony in the same
year. At MeDonogh, Md., in April, 1913, according to a correspond-
ent, sexed adults swarmed from the woodwork of a house, the beams
being honeycombed. A very large swarm issued on April 6, another
on April 13, a third on April 18, and a fourth swarm issued April 25,
comprising in all four distinct swarms. In size, however, the first
swarm, from my observations, is usually the largest. (Fig. 12.)

Leucotermes virginicus swarms during the forenoon in the vicinity
of Washington, D. C, usually one month later than flavipes, or in
early June. However, on August 11, 1913, at Falls Church, Mr.
William Middleton observed a large swarm to emerge from a small
chestnut corner stake which was set in the ground. The colony,
which was not large, swarmed at 12.30 p. m.

Leucotermes lucifugus, according to Heath,6 swarms "* * *
at different times between the months of October and April * * *."
"It usually takes place about 11 a. m."

During the early part of May, 1909, sexed adults of Leucotermes
sp. (probably flavipes) were observed by the writer emerging in great
numbers from the ground in the fenced yard of an old house located
in a pine grove near Woodville, Tyler County, Tex. The swarm
occurred in the forenoon of a warm, sunny day.

The following notes were made by members of the branch of
Southern Field Crop Insect Investigations, in charge of Mr. W. D.
Hunter:

On September 12, 1910, a light shower fell, and before the rain had entirely stopped
the air was full of flying termites; this lasted one hour. On December 16, 1910, there
was a swarm [lucifugus] at Dallas, Tex. After a rain at Dallas, October 13, 1911, there
was a swarm; another rain brought them out on October 16, 1911. On October 18,
1912, there were swarms of termites at Dallas, following a two-day's rain.

It will be seen from the foregoing that reliance, in the determination
of species, can be placed on the dates of swarming, since the species
flavipes never swarms in the fall in the eastern United States.
Apparently rainfall has no influence on the time of swarming, as is
the case in Texas.

a Hunter, W. D. Some notes on insect abundance in Texas in 1911. Proc. Ent.
Soc. Wash., vol. 14, no. 2, p. 62-66, June 19, 1912. See p. 63.
b Op. cit., p. 52.

70 IXSECTS INJURIOUS TO FORESTS.

ASSOCIATION WITH ANTS.

Grassi a and Escherich b have given interesting accounts of the rela-
tions between terrnites and ants. S. A. Forbes c states ih&t jlavipes
has been found associated with Formica scJiaufussi Mayr. H. C.
McCookd states that •jlavipes occurring under stones in the neighbor-
hood of the Alleghenies were seized and carried off by the mound build-
ing ants (Formica exsectoides Forel) when disturbed. J. C. Branner e
refers to the common ants as being enemies of the termites. King /
has noted the association of termites with ants. The following brief
notes have been made while investigating damage by termites to trees
and forest products:

Termites and ants are commonly to be found inhabiting the same
log or stump, yet ants are the enemies most to be feared by termites,
as they will capture and carry away the members of a disorganized
colony. Ordinarily the relations between termites and ants seem
to be neighborly and peaceful. If the termite colony is opened
up and disorganized, the ants at once take advantage of the oppor-
tunity and carry away the termites, which offer but little resistance.
Ants of several species may be attracted to such a helpless colony
from a distance. The soft-bodied soldiers are apparently not very
effective in such emergencies, although in the narrow channels of
the colony, where the powerful head with open mandibles is the only
front presented to the marauding ants, they afford some protection
to the colony.

Two species of carpenter ants (Camponotus pennsylvanicus Mayr
and Cremastogaster lineolata Say^) are the ants which more com-
monly have been found associated with termites in the eastern
United States. The latter species, due to its small size and rapidity
of movement, is a most formidable enemy.

Ants greatly diminish the number of the colonizing individuals at
the time of the swarm, carrying them away as they are running
about on the ground. Soldiers and workers guard the breaches from
which the sexed adults have emerged, possibly to keep the greedy
ants from following their prey to the parent nest.

a Grassi. B.. and Sandias. A. Op. cit,. p. 282-283.

5 Escherich. Iv. Die Termiten oder weissen Anieisen. Erne biologische Studie,
p. 122-126, Leipzig, 1909.

c Forbes. S. A. Nineteenth Eeport of the State Entomologist on the Noxious and
Beneficial Insects of the State of Illinois, p. 198, Springfield, 111.. 1S95. The white
ant in Illinois (Termes jlavipes, Kollar), p. 190-204.

d McCook, H. C. Note on mound-making ants. Proc. Acad. Xat. Sci. Phila. for
1879, p. 154-156, August 12, 1879.

e Branner, J. C. Geologic work of ants in tropical America. Bui. Geol. Soc.
Amer., v. 21, p. 449-496, rigs. 11. pi. 35, August 20. 1910. See p. 478-479.

/ Loc. cit.

9 Ident'_ 3d by Mr. Theodore Pergande. of the Bureau of Entomology.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 71
TERMITOPHILOUS INSECTS.

The presence of termitophilous insects or "guests" in colonies of
Leucotermes Jiavipes in the United States has been recorded by
several writers on termites. Mr. E. A. Schwarza has published an
extensive list of Coleoptera associated with Jiavipes. Inquilines, or
guests, are found only in permanent colonies and, as brought out
by Mr. Schwarz, b might be of importance in establishing the original
habitat of a species; that is, if a termite species had peculiar inqui-
lines (guests that do not occur among other species of termites) in one
country and none in another, it would indicate that the termite
species was native to the country where the inquilines occur in its
colonies. Mr. H. G. Hubbard found the staphylinid beetles Tri-
chopsenius and Anacyptus in a colony with supplementary royalty,
April 20, 1882, near Crescent City, Fla. He also found peculiar
wingless psocids which resemble young termites in a colony of the
latter, near Haw Creek, Fla., March 26, 1895. The note in his field
diary reads:

Termitophilous psocid found with termites in large log of pine, swampy hummock
of Prairie Farm. Several specimens in alcohol, together with worker of termite.
* * * The resemblance to a young termite is perfect, especially in mature speci-
mens. * * * The psocid is, however, much more active than the termite and very
difficult to capture. Immature specimens were not rare. * * * The immature
specimens inhabit the galleries of the termites, but are not so apt to be found among
the termites as in their immediate vicinity.

King c records three inquilinous staphylinid beetles as associated
with Jiavipes, PJiilotermes pilosus Kraatz, Hornalota sp., and Tachy-
porus jocosus Say.

All stages of small scarabseids, Homovalgus squamiger Beauvois and
Valgus canaliculatus Fabricius, are commonly found associated with
termites in decaying wood in Maryland, Virginia, West Virginia, and
North Carolina, and are probably truly inquilinous. From the mid-
dle of July, 1914, prepupal larvas and pupaB of Homovalgus squamiger
were commonly found in decaying wood infested by termites and in
the galleries of termites in Virginia. Adults of this beetle begin to
mature about the middle of August. The larvae construct oval pupal
cells in the decayed wood or make them of earth; the interior is
smooth and glossy. This beetle is probably a true inquiline.

a Schwarz, E. A. Termitophilous Coleoptera found in North America. Proc.
Ent. Soc. Wash., vol. 1, no. 3, p. 160-161, March 30, 1889.

Schwarz. E. A. Additions to the lists of North American termitophilous and
myrmecophilous Coleoptera. Proc. Ent. Soc. Wash., vol. 3, no. 2, p. 73-78, January
8, 1895.

b Schwarz, E. A. Termitidas observed in southwestern Texas in 1895. Proc. Ent.
Soc. Wash., vol. 4, no. 1, p. 38-42, November 5, 1896.

c Op. cit., p. 196.

62896°— Bull. 94, pt 2—15 5

72 INSECTS INJURIOUS TO FOEESTS.

Beetles of the family Pselaphidse are frequently found in decaying
wood near termite nests, and some are known to be their guests.
Adults of TmesipJiorus carinatus Say a were found in decaying wood
in which colonies of Jlavipes were present at Falls Church on March
18, 1912. Mr. E. A. Schwarz has included this species in his list of
myrmecophilous beetles as " often found among ants of various spe-
cies." b Adults of Batrisus virginise Casey a were found in decaying
wood infested with virginicus on the same day.

Adults of the staphylinid, PTiilotermes pennsylvanicus Kraatz a were
collected with Jlavipes near Kane, 111., August 11, 1911, in the butt
of a decaying white cedar telegraph pole. The species is a true
inquiline, and the beetles are very active. On August 16, 1913, near
Chain Bridge, Va., an adult of PTiilotermes sp. (possibly Juchsii
Kraatz) (determined by Mr. H. S. Barber of the Bureau of Entomology)
was found in a colony of virginicus. Blatchley c records PTiilotermes
pilosus Kraatz and P. fuclisii Kraatz in the nests of Jlavipes in
Indiana.

PARASITES.

Termites are infested externally with mites and internally with
protozoan parasites, but no internal or external feeding insect para-
sites have been recorded by Leidy/ Grassi,6 or Porter/ Grassi
states that the presence of these protozoa in the intestine retards sex-
ual development, as evidenced in the case of workers and soldiers.
He further states that they are normally absent in the reproductive
forms and newly hatched larvae.

SUMMARY AND CONCLUSIONS BASED ON THE RESULTS OF
THE EXPERIMENTS.

The following conclusions are based on observations of colonies in
the termitarium, colonies in small tin boxes, and other colonies in
the forest at Falls Church.

Colonizing individuals of both sexes swarm together from colonies
of Leucotermes Jlavipes and L. virginicus from about 11 a. m. to 1
p. m., the length of time occupied by the adults in emerging being
about one hour. No evidence of the separate swarming of the sexes

a Identified by Mr. E. A. Schwarz.

& Schwarz, E. A. Myrmecophilous Coleoptera, found in temperate North America,
Proc. Ent. Soc. Wash., v. 1, No. 4, p. 237-247, May 15, 1890.

c Blatchley, W. S. The Coleoptera or beetles of Indiana, p. 343-344, Indianapolis,
1910.

d Leidy, J. On intestinal parasites of Termes Jlavipes. Proc. Acad. Nat. Sci.
Phila. [v. 29] for 1877, p. 146-149, June 26, 1877.

Leidy, J. The parasites of the termites. Jour. Acad. Nat. Sci. Phila., ser. 2, v. 8,
p. 425-447, pis. 51-52, February, 1881.

e Grassi, B., and Sandias, A. Op. cit., p. 11-13.

/Porter, J. F. Trichonympha and other parasites of Termes Jlavipes. Bui. Mus.
Comp. Zool., v. 31, no. 3, p. 45-68, pis. 6, October, 1897.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 73

or seasonal dimorphism has been observed. However, all the indi-
viduals do not necessarily emerge from the same colony at the same
time, since there may be several swarms from the same nest. On
May 14, 1912, at Falls Church sexed adults of flavipes emerged,
although not in great numbers, from colonies, whereas there was
evidence, by the discarded wings on the ground, of an earlier swarm.
Most of the colonizing individuals had swarmed from other colonies
on May 8; yet some individuals, not in the enormous numbers of
the first swarm, were observed swarming from the same colonies on
May 14. While there may be as many as four swarms, the first is
usually the largest. This may be explained by the fact that there
is an uneven development of individuals. Indeed, a few retarded,
winged, sexed adults (virginicus) may remain in the colony till July
24 (near Kane, 111.) and early August (District of Columbia) or be
found, as individuals, flying. A large swarm of virginicus has
emerged from a colony as late as August 11, 1913, at Falls Church.

The winged insects usually crawl up to some elevated place before
taking flight. There is an enormous mortality of the colonizing
individuals, and insectivorous animals destroy them in great num-
bers. The swarm is not a " nuptial flight."

The so-called " amatory passages" possess a sexual significance,
and there is a mutual attraction between the sexes several days
before the normal period of swarming. This can be observed if the
colony is disturbed and the colonizing forms emerge prematurely, and
is evidenced even before the loss of the wings. This attraction,
probably due to some secretion, continues till after the royal pair is
established in the royal cell and copulation has taken place. Copula-
tion probably occurs a week after the swarm of flavipes; that is, on
May 15 adults that had swarmed on May 8 were in the royal cell and
apparently no longer following each other about, head close to abdo-
men, as previously. There is evidence that individuals of neither
sex are sexually mature at the time of swarming, and that there is
further development before copulation, as can be noted in the increase
in size of the abdomen of both sexes.

The colonizing individuals are not all irretrievably lost.a The
establishment of new colonies by these forms is a normal process.
Although there is an enormous mortality at the time of the swarm,
and a still further diminution in numbers due to inability to become
established under favorable conditions, yet some pairs do become
established. These sexed adults are not necessarily monogamous.

a Perris, E. Nouvelles promenades entomologiques, Termes lucifugus. Ann. Soe.
Ent. France, ser. 5, t. 6, p. 201-202, 1876.

Perez, J. Sur la formation de colonies nouvelles chez le termite lucifuge ( Termes
lucifugus). Compt. Rend. Acad.,Sci. (Paris), t, 119, No. 19, p. 804-806, Nov. 5, 1894.

Heath, Harold. Loc. cit.

74 INSECTS INJURIOUS TO FORESTS.

True royal pairs are independent of workers in the foundation of
new colonies; that is, it is not necessary that they be found and
established in royal cells by foraging workers and soldiers, although
it is possible that this occurs, which would then constitute an inde-
pendent colony. New colonies established by sexed adults that
swarmed were found both in the termitarium (where conditions were
similar to those in nature) and in the forest. The king and queen
are equally important; they continue to cohabit and coition is
repeated. The first brood reared by young true queens that have
swarmed consists of workers and soldiers of smaller size than the
normal form.

Neoteinic royal individuals are to be found commonly in colonies
of jiavipes in the eastern United States; they are "ergatoids" or
"neotenes" and are developed by retarding the normal development
of nymphs of the first form at an early stage, and from young larva?;
sometimes as many as 40 or more, consisting of many queens and a
few males, may be present in the same colony. The males are
polygamous. It is believed that these forms are provided (1) when
overcrowded old colonies are split up and new, independent colonies
are to be established, or (2) through the actual loss of the true royalty
or by the accidental separation of some members of the colony from
the royalty. This method of the formation of new colonies is sure,
and probably much more rapid in the case of establishment by the
true colonizing forms, as the royalty would receive the care of the
workers, would not have to forage for themselves, and their only
function would be reproduction. The number of eggs laid and the
rate of increase would necessarily be much more rapid, due to the
following facts: (1) The abdomens of mature, fertilized, supple-
mentary queens are nearly as fully distended as in the case of true
queens; (2) the large number of supplementary royalty possible to
be present in a single colony and consequent increase in the number
of eggs laid, and (3) the proper care and nourishment the royalty
would receive.

It is often noted that the antennas of the reproductive forms are
mutilated, that is, have a greater or less number of segments missing.
However, this is not always the case in young reproductive forms.
The loss of the segments in individuals in long-established colonies
might be due to the treatment the latter receive from the workers, or
from being dragged about by them, but the antenna? are sometimes
mutilated in individuals in incipient colonies.

There is a series of molts and " quiescent stages" in the develop-
ment of the larva? of the castes; caste differentiation occurs during
such a stage, which corresponds somewhat to the pupal stage in
insects with complete metamorphoses.

In conclusion, the more important facts may be summarized as
follows: There is great variation in the life cycle of Leucotermes;

Bui. 94, Part II, Bureau of Entomology, U. S. Dept. of Agriculture.

Plate XVII.

Fig. 1. — Chestnut telephone pole, with base
charred but basal area untreated. This
pole has been standing near Dover, N. J.,
for eight years. ( Photograph loaned by
United States Forest Service.)

r1 I U

Fig. 2. — Pine flooring honeycombed by ter-
mites at New Iberia, La. (Original.)

Damage by Termites.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 75

these insects are adapted to meet emergencies successfully and over-
come obstacles without the disorganization of the colony. New
colonies may be established (1) by the sexed colonizing adults that
invariably swarm and leave the parent colony; (2) by neoteinic royal
individuals, produced from nymphs of the second form which never
( ?) leave the parent colony, or from young larvae, as in colonies
orphaned after the nymphs of the first form have nearly completed
their development; (3) by neoteinic reproductive forms supplied to
orphaned colonies, which may be derived from nymphs of the first or
second forms, or larvae. Nests headed by true royalty are not rare,
but many difficulties are to be surmounted in their establishment by
sexed adults ; such recently established colonies are small in number.
Nests headed by neoteinic reproductive forms are more commonly
to be found, as this is a more sure and more rapid method of estab-
lishment. Colonies established by neoteinic reproductive forms
necessarily increase in size more rapidly due to the numerous egg-
laying queens and the care and food they receive from the workers.
Subcolonies or temporary colonies are frequently found with only
workers and soldiers present; these subcolonies, which furnish
increased facilities for habitation and food supplies, are possibly
offshoots from the parent colony or nest and are established by
means of subterranean passages, which are extended for long dis-
tances by foraging workers and soldiers.

THE DAMAGE TO FOREST PRODUCTS.

Termites seriously injure construction timbers in bridges, a wharves,
and like structures; telephone and telegraph poles b (PL XVII, fig. 1),
hop poles,c mine props, d fence posts and rails or boarding; lumber
piled on the ground, railroad ties set in the ground (not where there
is stone or slag ballast or heavy traffic), woodwork (PI. V; PL XVII,
fig. 2) in new and old buildings/ and especially seriously damage the
wooden boxing or "conduits" of insulated cables placed in the ground
(to the detriment of the insulation) ; tent pins and ridge poles, wooden

a Hagen, H. A. The probable danger from white ants. Amer. Nat., v. 10, no. 7,
p. 401^410, July, 1876.

& Snyder, T. E. Insects injurious to forests and forest products. Damage to chest-
nut telephone and telegraph poles by wood-boring insects. U. S. Dept. Agr., Bur.
Ent., Bui. 94, pt. 1, pp. 12, figs. 3, pis. 2, December 31, 1910. See p. 9-10.

c Parker, W. B. California redwood attacked by Termes lucifugus Rossi. Jour.
Econ. Ent., v. 4, no. 5, p. 422-423, October, 1911.

d Snyder, T. E. Insect damage to mine props and methods of preventing the
injury. U. S. Dept. Agr., Bur. Ent., Circ. 156, pp. 4, July 13, 1912. See p. 2-3.

^Marlatt, C. L. The white ant. (Termes flavipes Koll.). U. S. Dept. Agr., Bur.
Ent., Circ. 50, rev. ed., pp. 8, figs. 4, January 27, 1908.

Hopkins, A. D. Insect injuries to forest products. U. S. Dept. Agr. Yearbook
1904, p. 381-398, 1905. White ants, or termites, p. 389-390.

76 INSECTS INJUBIOUS TO FORESTS.

beehives and tree boxes; wooden electrotype blocks, and books (PL
IV) and documents stored in damp, dark places, etc. ; timber in con-
tact with the ground being especially liable to serious damage. Often
the damaged material has to be removed and replaced, or rebuilt.
The wood of no species of native tree of commercial importance is
"immune" to attack, although some are relatively more resistant
than others. Such damage has occurred as far north as Boston and
the shores of the Great Lakes, but greatly increases as the Tropics are
approached. In the Southern States termites are especially destruc-
tive to wooden underpinning, beams, and flooring (PI. XVII, fig. 2),
and all other material of wood accessible in buildings. They enter
buildings by means of tunnels through the ground, by way of wooden
joists, or by means of covered paths (minute "sheds" constructed of
earth and excrement of the superficial consistency of sand), leading to
the woodwork over the surface foundations of stone or other material
which they can not penetrate. This enables them to avoid the light.
Thus termites silently, secretly, and ceaselessly work their insidious
damage, instinctively never perforating the exposed surface of timber,
except to enable the sexed adults to swarm. Sometimes the emerg-
ence of these winged forms is the first indication of their presence,
but at other times joists and floors collapse without warning.

PREVENTIVES, REMEDIES, AND ''IMMUNE" WOODS.

Forest products in contact with the ground should be impregnated
with coal-tar creosote, which is a permanent preventive against
attack by our native termites. Coal-tar creosote has many prop-
erties which would recommend its use in this respect, for it is also a
fungicide, and, being insoluble in water, will not leach out in wet
locations. These requirements furnish objections to many chem-
icals that otherwise are very effective insecticides. The various
methods of superficially treating timber, as by charring, by brushing,
or by dipping with various chemical preservatives, among which are
creosotes, carbolineums, etc., have proven to be temporarily effec-
tive iii preventing attack,0 if the work is thoroughly done. If not
thoroughly done, termites enter through the untreated or imper-
fectly treated portions, especially through weathering checks and
knots. Where the bases of poles, mine props, etc., are left untreated
termites enter the timber from below, and, avoiding the treated
portions, come up through the interior. Charred timber is effective
against termite attack for a period less than a year, although it is not
seriously damaged at the end of one year. It will readily be
seen that neither brushing nor spraying the exterior after place-

a Snyder, T. E. Insects injurious to forests and forest products. Damage to chest-
nut telephone and telegraph poles by wood-boring insects. IT. S. Dept. Agr., Bur.
Ent., Bui. 94, pt. 1. pp. 12, figs. 3, pis. 2, Dec. 31, 1910. See p. 9-10.

BIOLOGY OF THE TEEMITES OF THE EASTEEN UNITED STATES. 77

ment, as is sometimes practiced, is effective in keeping out termites,
since the portion that sets in the ground could not be treated, and it
is usually at this point that termite attack occurs.

Before treating timber with chemical preservatives, especially
where the brush method is employed, it is essential that the timber
be thoroughly seasoned, otherwise penetration by the preservative
will be retarded.

A treatment with "blue oil" is recorded as apparently effective in
protecting wood against the attacks of "white ants," or termites,
besides acting as a preservative (fungicide) generally. "Blue oil"a
is the residue left in the distillation of mineral oils after the isolation
of kerosene (petroleum) and paraffin; (a) the oil to be a shale product;
(b) its specific gravity (at 60° F.) to be 0.873 to 0.883; (c) its flashing
temperature to be not lower than 275° F. (close test).

Many patented wood preservatives, advertised as effective against
wood borers, often merely contain simple preservatives, as for in-
stance, linseed oil, to which a slight odor of oil of citronella has been
imparted, or contain simple poisons. For timber to be set in the
ground, brush coatings with linseed oil are not effective against
termites.

An English firm manufactures a saccharine solution which prob-
ably contains a salt as arsenic 6; this patented treatment is supposed
to be efficient against wood-boring insects, especially termites. The
wood is seasoned by immersing in the saccharine solution at 120
to 140° F. This process is being tested.

Impregnation with chlorinated naphthalene may prove effective
against termites, as a preservative for woodwork, in interior finish,
where a requirement is that the preservative should not "sweat"
out, or stain the wood. Treated wood blocks buried in the ground
with termite-infested logs were not attacked after a test of nearly
six months. c Impregnation with paraffin wax was not effective
(fig. 14). If the wood is not in contact with the ground, impregna-
tion treatments with bichlorid of mercury and zinc chlorid are effec-
tive. The mercury and zinc in this form are both soluble in water.

H. W. Bates, in a paper entitled "On the prevention of destruc-
tion of timbers by termites," Transactions of the Entomological
Society of London, 1864, Vol. I, p. 185, cites preventive measures.
M. J. Berkeley, in The Technologist, (London), 1865, Vol. V, p. 453,
gives remedies based on the report by the committee of inquiry into

"The protection of timber against white ants. Trans. Roy. Scot, Arbor. Soc, v. 23,
pt, 2, p. 227-228, July, 191 0.

Dixon, W. B. Protection from "white ants" and other pests. Nature, v. 85,
no. 2148, p. 271, December 29, 1910.

& Chemical abstracts, v. 7, no. 2, p. 408, January 20, 1913.

c Impregnation of wood to resist insect attack. Amer. Lumberman, no. 2009, p. 32,
November 15, 1913.

78

INSECTS INJURIOUS TO FORESTS.

the ravages by white ants at St. Helena. Froggatt gives a preventive
measures applicable in New South Wales.

Fig. 14. — Red oak block, impregnated with paraffin wax, honeycombed by termites after five months '
test. This was buried in the ground with termite-infested logs. (Original.)

° Froggatt, W. W. "White ants, with some account of their habits and depredations.
Misc. Pub. no. 155, Dept. Agr., Sydney, N. S. Wales, Sydney, 1897. See p. 6-8,
account of depredations and methods of prevention.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 79

The wood-pulp products and various patented " composition
boards" used as substitutes for lath, etc., might be made termite proof
by adding during their manufacture such poisons as white arsenic,
antimony, bichlorid of mercury, zinc chlorid, etc; tests are being con-
ducted.

In general, serious damage by termites to the wood of fire or
insect killed, standing, merchantable timber can be prevented if the
timber is utilized within from one to two years from the time that it
was killed, depending on the species of wood and the locality — one
year for pine (less in the Southern States) and two years for chestnut.

Forest tree nursery stock should be planted in ground that has
been plowed deep late in the fall of the year in a region where injury
by termites is common.

Marlatt a states regarding white-ant infestation in buildings that
setting foundation beams or joists in concrete is only a partial
protection, since in the settling of the house the concrete will crack
and afford entrance to the insects. Some protection is afforded by
removing decaying stumps or posts, etc., adjacent to buildings, by
drenching infested timbers with kerosene, and by removing infested
joists in cellars and drenching the ground where they were set with
kerosene (or carbon bisulphid). Where the injury is confined to
exposed woodwork in buildings, hydrocyanic-acid gas fumigation b
is to be recommended, the infested beams and joists beneath being
exposed, if possible, by opening up the floors.

Certain species of wood appear to be naturally highly resistant to
termite attack. Species of wood that, so far as determined by test,c
have been resistant to attack by our native termites are all tropical
species and woods too expensive for ordinary use, including teak
(Tectonia grandis) from Siam and Burma, greenheart (Nectandra
rodisei) d from South America and the West Indies, "peroba" (several
species of Aspidosperma) from South America, and mahogany
(Swietenia mahoghani) from tropical America. Hagen6 states that,
according to Kirby, " Indian oak" or teak (Tectonia grandis) and
"ironwood " (Sideroxylon) of India are immune to attack by termites.
This immunity ( ?) or relative resistance of ironwood is not due to
hardness, since Asiatic termites attack the hardest wood, Lignum-
vitse, but to the presence in the wood of substances (as oils or alka-

« Op. cit.

& Howard, L. O., and Popenoe, C. H. Hydrocyanic-acid gas against household
insects. TJ. S. Dept. Agr., Bur. Ent., Circ. 163, p. 8, November 29, 1912.

c Impregnation of wood to resist insect attack. Amer. Lumberman, no. 2009, p. 32,
November 15, 1913.

^This wood has been superficially attacked by termites after 12 months' test; the
wood was eaten to the depth of one-eighth inch.

« Hagen, H. A. Monographie der Termiten. Linnsea Entomologica, Bd. 10, p.
44-45, 1855.

80 INSECTS INJURIOUS TO FORESTS.

loids) repellent or distasteful to termites.0 The presence of tyloses
or of gums may be factors in determining h the durability and
resistance of hardwood species. Hagen further states that, while teak
is not destroyed by termites, he believes that even teak will be
attacked when it has become old or been long exposed to the air.
Hagen states that it is useless to consider tannin as a preservative,
since, according to Williamson, termites will destroy leather. Some
species of woods native to the Philippine Islands are apparently
immune to termite attack.

Capt. Ahern, Chief of the Philippine Forestry Bureau, quotes
Foreman, p. 390, as stating0 that the "anay," or native termite, "eats
through most woods (there are some rare exceptions, such as 'molave/
'ipil/ 'yacal/ etc.)." Capt. Ahern states that —

The following woods are not subject to attack by anay: "Dinglas" [Eugenia brae-
teata Roxb., var. roxburghii Duthie, family Myrtacese], "ipil" [Afzelia bijuga Gray,
family Leguniinosse], "molave" [Vitex littoralis Dene, family Verbenaceae], and
"yacal" [Hopea plagata Vidal, family Dipterocarpese].

"Tindalo" [Afzelia rhomboidea Vid.] is attacked by "anay" when there is no other
wood in the vicinity.

"Baticulin" [Litsea obtusata B. and H., F. Vill, fam. Laurineag] is attacked by
"anay, " but is not damaged or destroyed, except such parts as are buried underground.

TEST WITH THE WHITE ANT.

Mr. D.N. McChesney, master mechanic at the depot quartermaster shops in Manila,
found last February that his trunk (made of an American spruce) had been invaded
by white ants and was almost entirely destroyed; the clothes contained in the trunk
were also eaten. He placed the trunk on the ground and near it pieces of the following
woods:

Result of 30 days' contact with ants.
American woods:

Oregon pine Entered and eaten; a mere matter of time

for complete destruction.

Bull pine Eaten more readily than Oregon pine.

Spruce Do.

Western hemlock Not touched.

California redwood [Sequoia semper- Ants tried, but discontinued after a slight
virens]. effort.

California white cedar. .*. Do.

Native woods:

Molave Ate a little of it; deepest hole about one-
fourth inch.
Narra [Pterocarpus indicus Wild., fam- Ate a little of it; deepest hole about one-
ily Leguniinosse]. half inch.

Painted wood Ants worked under paint and ate the

wood readily.

° The quality of hardness in wood, while not rendering a species of wood immune
to termite attack, is an important factor in determining the relative resistance of
species of woods. Hardness is a factor in the grading of mahogany lumber.

b Gerry, M. "Tyloses: Their occurrence and practical significance in some Ameri-
can woods. " Jour. Agric. Research, vol. 1, no. 6, p. 462-464, March 25, 1914.

c Ahern, G. P. Important Philippine woods. 112 pp., col. pis. Forestry Bur.,
Manila, P. I., January 2, 1901. See p. 91.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 81

Hemlock and redwood a are badly honeycombed by our native
species of termites on the Pacific coast, and California white cedar
is honeycombed by the termites of the Pacific coast.

European cypress (Cupressus sempervirens Linn.) is reported b
damaged by termites at Rochefort, France.

Cedrus deodar, from India, and Cedrus atlantica, of the mountains
of northern Africa, are reported to be immune ( ?) to termite attack.

Red "deal" is less liable to attack than white "deal." c

The following three species of wood remained untouched by ter-
mites for three year's in the District of Pretoria, Transvaal: d " Lead-
wood" (Combretum prophyrolepsis) , " black ironwood " (Olealauri-
folia) , ' ' vaalbosch " (Brachylsena discolor) .

In case of certain species of pines, with an extremely resinous heart-
wood, as the "fatwood" of the longleaf pine (Pinus palustris) of the
Southern States, while termites honeycomb the sapwood, the heart-
wood apparently is resistant. Odenbach states that turpentine is
very repellent to termites in artificial nests. In southern Rhodesia e
the wood of the "mopani" tree (Copaifera mopani) withstands ter-
mite attack for years, and is therefore very suitable for straining
posts for fences, though unfortunately not a timber that can be cut
and squared.

Tests of the relative resistance of various native and exotic woods
have been begun, but as yet no definite conclusions can be drawn.
The heartwood of the following native species of wood is relatively
more resistant to attack by our native termites: Black locust; black
walnut f (cases on record where heavy beams supporting flooring in a
building in Baltimore, Md., were completely honeycombed); eastern
white cedar (Chamsecyparis thyoides); eastern red cedar or juniper;
bald cypress of the Southern States; western red cedar (Thuja plicata)
of Washington, Oregon, and California; incense cedar (Libocedrus
decurrens) of Oregon and California; and Monterey cypress (Cupressus
macrocarpa) of California. All these species of woods, however, are
attacked by termites.

a Parker, W. B. Loc. cit.

b Hagen, H. A. Monographie der Termiten. Linnaea Entomologica, Bd. 10,
p. 1-144, 1855. See p. 133.

c French, C. Handbook of the destructive insects of Victoria, pt. 2, p. 141, Mel-
bourne, 1893.

d Howard, C. W., and Thomsen, F. Notes on termites. Transvaal Agr. Jour., v. 6,
No. 21, p. 85-93, illus., October, 1907; v. 7, No. 27, p. 512-520, April, 1909; v. 8, No.
29, p. 86-87, October, 1909.

e Jack, R. W. Termites or "white ants." Rhodesia Agr. Jour., v. 10, No. 3,
p. 393-407, pi., February, 1913.

/ Impregnation of wood to resist insect attack. Amer. Lumberman, No. 2009^
p. 32, November 15, 1913.

82 INSECTS IXJUEIOUS TO FORESTS.

Hagen a briefs the records of various travelers as to the immune (?)
or termite-resistant wood species of the countries of the world.
Froggatt b states that in Australia red pine is more resistant than clear
pine; that " jarrah" is said to be resistant, though not immune; and
that desert cypress when sawn up appears to be resistant, but in the
form of logs is not immune. The Rev. Joseph Assmuth, S. J.,c of
Bombay, India, states that "deal" and "pukka" teak are injured
by termites: he gives photographs of the damaged specimens of wood.
In answer to a letter of inquiry he states:

The "'pukka" teak is what is called here in India "Burmese teak," Tectonia
grandis L. "Pukka" means genuine; it is used here in opposition to the less reliable
timber of '"Malabar teak." though both come from the same species of wood. The
difference of both lies. I am told, in the seasoning of the timber, or rather in the differ-
ent mode of felling the trees. In Malabar they cut off a ring of bark from near the base
of the tree, so that the tree dries up while standing still erect, and is then felled. This
seems to cause a gradual withdrawal of the oils contained in the wood, which makes
the wood more liable to the attack of white ants. In Burmah the tree is felled as it
stands and allowed to dry up lying on the ground. Thus the peculiar oils remain in
the wood and are preserved in it, and consequently this timber is less palatable to
the white ants, and shunned by them until in course of time the oils evaporate also.
Then the white ants go for it too. Such, at any rate, is my theory. I can't explain
otherwise why the one sort is at once attached by white ants, whereas the other remains
for a longer period, at least, immune. The case therefore is briefly this: Malabar teak
(here also called "jungle teak") is attacked by the white ants from the beginning.
Burmah or "pukka" teak remains safe for a certain period, sometimes longer, some-
times shorter, but it is not absolutely safe either.

"Dealwood" is the common European wood used for boxes and the like; it is usu-
ally timber of Abies. Picea. or Pinus. It is the wood most readily attacked by ter-
mites of different species.

Kanehira/ tabulates the results of experiments with a large
number of species of woods as to their relative resistance to termite
attack in Formosa, Japan. Reasons are advanced as to the probable
causes which render the wood "termite proof" ( ?). The conclusions
he draws can not be accepted without further details as to how the
tests were conducted and after a longer period of experimentation.
Certain of the woods he lists as "immune" are known to be attacked
by termites.

METHODS OF OBTAINING PHOTOGRAPHS FOR THE ILLUSTRATIONS.

The photographs of the insects reproduced in this paper were made
after a method devised by Mr. H. S. Barber, of the Bureau of Ento-
mology. The specimens were placed either between horizontal

a Loc. cit.

b Froggatt, W. W. WHteants. (Termitidse.) Misc. Pub. 874. Dept. Agr. X. S.
Wales, Sydney, 1905, p. 43-44.

c Assmuth, J. Wood-destroying white ants of the Bombay presidency. Jour.
Bombay Xat. Hist. Soc, v. 22, no. 2, p. 372-384, 4 pis., September 30, 1913.

d Kanehira. On some timbers which resist the attack of termites. Indian For-
ester, v. 40, do. 1, p. 23-41, January, 1914.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 83

glass slides submerged in alcohol or immersed in water in a chamber
made by gluing glass slides to thin slips of rectangular cover glass
with heated balsam. In this water-tight compartment the speci-
mens can be placed in vertical or horizontal positions. Ether is
used to clear out the cell, which is filled with boiled water, thus
avoiding the formation of air bubbles. If the specimens are placed
head downward, movements of the antennas, due to vibration, can
be avoided. Cells of various sizes are used, or one large cell can be
utilized by placing thin slips of cover glass back of the specimens to
hold them firmly in position. By clamping the cell to the lens
holder of a dissecting microscope placed horizontally, focusing can
be conveniently accomplished by adjustments of the screw.

The best results are obtained by etherizing recently killed specimens
that have not been placed in alcohol. A 72 millimeter focal length
lens brings out the characters to the best advantage, if the enlarge-
ment is 6 to 8 diameters.

Many of the photographs were taken by Mr. H. S. Barber and some
by Mr. H. B. Kirk at the eastern field station. Most of them, how-
ever, were made by the photographic laboratory of the United
States Department of Agriculture.

BIBLIOGRAPHY.^

This bibliography consists mainly of publications on our native
termites or closely allied species; some little-known publications are
included. Hagen, up to 1860, Froggatt, Holmgren, Escherich, and
Feytaud give extensive bibliographies.

Andrews, E. A., and Middleton, A. R. Rhythmic activity in termite communities.

Johns Hopkins Univ. Circ, n. s., 1911, no. 2 (whole no. 232), p. 26-34, illus.,

February, 1911.
Atkinson, G. F. Some Carolina insects. 1st Rpt. So. Car. Exp. Sta. for 1888,

p. 19-56, 1889.
Banks, N. Two new termites. Ent. News, v. 17, no. 9, p. 336-337, illus., Novem-
ber, 1906. Cryptotermes cavifrons, from Florida, and Termopsis laticeps, from

Arizona, described.
Bidie, G. White ants — termites eroding glass. Nature, v. 26, no. 675, p. 549, October

5, 1882. t

Buckley, S. B. Description of two new species of termites from Texas. Proc. Ent.

Soc. Phila., v. 1, p. 212-215, May, 1862 (1863). Hamitermes (Termes) tubiformans,

from Texas, p. 212, and Eutermes cinereus, from Texas, described.
Casey, T. L. A new genus of termitophilous Staphylinidse. Ann. N. Y. Acad.

Sci., v. 4, p. 384-387, March, 1889.
Coryell, J. R. The termite pest of the old world. Sci. Amer., v. 59, no. 10, p. 151,

September 8, 1888, illus.
Derry, D. E. Damage done to skulls and bones by termites. Nature, v. 86, no.

2164, p. 245-246, April 20, 1911.
Dixon, W. A. Protection from "white ants" and other pests. Nature, v. 85, no.

2148, p. 271, December 29, 1910.

a The references in the bibliography, as well as those in the footnotes, have been
verified by the librarian of the Bureau of Entomology, Miss Mabel Colcord.

84 INSECTS INJURIOUS TO FORESTS.

Fitch, A. Fourth Report on the Noxious and Other Insects of the State of New York,

1858, p. [8] or Trans. Termes frontalis. N. Y. State Agr. Soc, v. 17 (1857), p.

694, 1858.
Forbes, S. A. The white ant in Illinois. (Termes flavipes Kollar.) 19th Rpt. State

Entomologist . . . of Illinois for 1893 and 1894, p. 190-204, 1895. Lists economic

publications on termites.
Grant, R. D. [Ravages of Termes flavipes.] Trans. Acad. Sci. St. Louis, v. 3,

Jour, of Proc, p. cclxix, November 19, 1877.
Hagen, H. A. Report on the Pseudo-Neuroptera and Neuroptera collected by

Lieut. W. L. Carpenter in 1873 in Colorado. Ann. Rpt. U. S. Geol. Survey

Ter. for 1873, by F. V. Hayden, p. 571-606, 1874.
White ants destroying living trees and changing foliage in Cambridge, Mass.

Canad. Ent., v. 17, no. 7, p. 134-136, July, 1885.
The female of Eutermes rippertii. Psyche, v. 5, no. 157-159, May-July, 1889

p. 203-208.
Haviland, G. D. Observations on termites; with descriptions of new species. Jour.

Linn. Soc. [London], Zool., v. 26, no. 169, p. 358-442, pis. 22-25, April 1, 1898.
Observations on termites or white ants. Ann. Rpt. Smithsn. Inst, for year

ending June 30, 1901, p. 667-678, pi. I-IV, 1902.
Hubbard, H. G. Notes on the tree nests of termites in Jamaica. Proc. Boston Soc.

Nat. Hist., v. 19, p. 267-274, December 26, 1877, printed March, 1878.

Insects affecting the orange, Washington, 1885. Chap. IX, p. 121-125, White

ants or "wood-lice."
Kellogg, V. L. Are the Mallophaga degenerate psocids? Psyche, v. 9, no. 313,

p. 339-343, May, 1902.
Knower, H. McE. The development of a termite — Eutermes (Rippertii?) . A pre-
liminary abstract. Johns Hopkins Univ. Circ. , v. 15, no. 126, p. 86-87, June, 1896.
The embryology of a termite, Eutermes (Rippertii?). Jour. Morphol. [Boston],

v. 16, no. 3, p. 505-568, pi. 29-32, August, 1900.
A comparative study of the development of the generative tract in termites.

Johns Hopkins Hospital Bui., v. 12, no. 121-122-123, p. 135-136, 2 figs., Balti-
more, April-May- June, 1901.
Lintner, J. A. Entomology. Proc. Albany Inst., v. 2, p. 48-50, 1878.
Osborn, H. On the occurrence of the white ant (Termes flavipes) in Iowa. Proc.

Iowa Acad. Sci., v. 5, for 1897, p. 231, 1898.
Osten-Sacken, C. R. Extract from a letter by Baron R. Osten-Sacken, on the

specimens of Termes found by him in California. Proc. Boston Soc. Nat. Hist.,

v. 19, p. 72-73, for January 3, 1877, June, 1877.
Packard, A. S. Third Report U. S. Entomological Commission, Washington, 1883,

p. 326-329.

On the systematic position of the Mallophaga. Proc. Amer. Philos. Soc,

v. 24, p. 264-272, 13 figs., September 2, 1887.
A Textbook of Entomology, New York, 1903.
Parker, W. B. California redwood attacked by Termes lucifugus Rossi. Jour.

Econ. Ent., v. 4, no. 5, p. 422-423, October, 1911.
Riley, C. V., editor. Amer. Ent., v. 3 (n. s., v. 1), p. 15, January, 1880. Termi-

tophilous insects collected by E. A. Schwarz recorded.
Riley, C. V., and Howard, L. O. Termites swarming in houses. U. S. Dept. Agr.,

Div. Ent., Insect Life, v. 6, no. 1, p. 35, November, 1893.
Scammell, E. H. White ants. Knowledge, n. s., v. 4, p. 10-12, January, 1907.

Protection of wood by a newly devised process.
Schwarz, E. A. Staphylinidae inquilinous in the galleries of Termes flavipes. Amer.

Ent., v. 3 (n. s., v. 1), p. 15, January, 1880.

Termitophilous Coleoptera found in North America. Proc. Ent. Soc. Wash.,

v. 1, no. 3, p. 160-161, 1889.

BIOLOGY OF THE TERMITES OF THE EASTERN UNITED STATES. 85

Scudder, S. H. Remarks upon the American white ant. Proc. Boston Soc. Nat.
Hist., v. 7, p. 287-288, 1860.

Further injury to living plants by white ants. Canad. Ent., v. 19, no. 11, p.
217-218, November, 1887.

More damage by white ants in New England. Psyche, v. 6, no. 177, p. 15-16,
January, 1891.

The fossil white ants of Colorado. Proc. Amer. Acad. Arts and Sci., v. 19
(n. s., v. 11), p. 133-145, 1884.
Smith, E. F. White ants as cultivators of fungi. Amer. Nat., v. 30, p. 319-321,

April, 1896.
Snyder, T. E. Record of the finding of a true queen of Termes flavipes Kol. Proc.
Ent. Soc. Wash., v. 14, no. 2, p. 107-108, pi. 3, June 19, 1912.

Changes during quiescent stages in the metamorphosis of termites. Proc. Ent.
Soc. Wash., v. 15, no. 4, p. 161-165, December, 1913.
Stokes, A. C. The sense-organs on the legs of our white ants. Termes flavipes Koll.

Science, v. 22, no. 563, p. 273-276, November 17, 1893.
Warren, E. Some statistical observations on termites, mainly based on the work of
the late Mr. G. D. Haviland. Biometrika, v. 6, no. 4, p. 329-347, illus., March,
1909.
W^heeler, W. M. The embryology of Blatta germanica and Doryphora decemlineata.
Jour. Morphol., Boston, v. 3, no. 2, p. 291-386, pi. 15-21, September, 1889.

The phylogeny of the termites. Biol. Bui., v. 8, no. 1, p. 29-37, December,
1904.

The fungus-growing ants of North America. Bui. Amer. Mus. Nat. Hist., v. 23>
p. 669-807, pi. 49-53, September, 1907. Including Pt. IV, 1, The fungus-growing
termites, p. 775-786.

RELATION OF TERMITES TO THE ORIGIN OP HOG-WALLOWS AND PRAIRIE MOUNDS.

Barnes, G. W. The hillocks or mound-formations of San Diego, California. Amer.
Nat., v. 13, no. 9, p. 565-571, September, 1879.

Branner, J. C. Decomposition of rocks in Brazil — Burrowing animals — Ants. Bui.
Geol. Soc. Amer., v. 7, p. 295-300, February 4, 1896.

Ants as geologic agents in the Tropics, Jour. Geol., v. S, p. 151-153, February-
March, 1900.

Natural mounds or hog-wallows. Science, n. s., v. 21, no. 535, p. 514-516,
March 31, 1905.

Geologic work of ants in tropical America. Bui. Geol. Soc. Amer., v. 21,
p. 449-496, fig. 11, pi. 35, August 20, 1910.

Forshey and Copes. Mounds. Proc. New Orleans Acad. Sci., v. 1, no. 1, p. 18-20,
March 1, 1854.

Hilgard, E. W. Physico-geographical and agricultural features of the State of Cali-
fornia. U. S. Dept. Int., Census Off. 10th Census U. S., v. 6. Report on cotton
production in the United States, Part II, p. 663-741. Hog-wallows, p. 676-677,
693.

Le Conte, J. Prairie mounds. Proc. Cal. Acad. Sci., v. 5, p. 219-220, January, 1874.
Hog-wallows or prairie mounds. Nature, v. 15, p. 530-531, April 19, 1877.

McCook, H. C. Note on mound-making ant's. Proc. Acad. Nat. Sci. Phila., 1879,
p. 154-156, 1880.

Turner, H. W. Hog-wallow mounds. U. S. Geol. Survey, Ann. Rept. 17, 1895-96,
Pt. I, p. 681-683, pi. 33, 1896.

Veatch, A. C. Formation of natural mounds — Ant-hill theory. U. S. Geol. Survey,
Prof. Paper no. 46, p. 55-56 and 58-59, 1906.

Wallace, A. R. Glacial drift in California. Nature, v. 15, p. 274-275, January 25,
1877.

ADDITION AL COPIES

OF THIS PUBLICATIOX !MAY BE PBOCTBZ^ FBOM

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INDEX.

Page.

Afzelia bijuga, wood resistant to termite attack 80

Afzelia rhomboidea, wood partially resistant to termite attack 80

Agrilus bilineatus in chestnut, destructive work 6

Alaus sp., damage to chestnut telephone and telegraph poles 8

Anacyptus, ''guest," or inquiline, of Leucotermes flavipes 71

' ' Anay, ' ' name for native termite of Philippines . 80

Ant colonies, feeding of larvae by queen 53

Antimony against termites 79

Ants —

carpenter, work distinguished from that of termites 22

white. (See Termites.)

Aptera 14

Arsenic—

saccharr .e solution against termites 77

white, against termites 79

Aspidosperma spp., wood resistant to termite attack. 79

"Baticulin." (See Litsea obtusata.)

Batrisus virginise, association with Leucotermes virginicus 72

Beehives, wooden, destruction by termites 76

Bichlorid of mercury against termites 77, 79

Bisulphid of carbon against termites 79

Blatta germanica, bibliographic reference 85

' ' Blattiden " 15

" Blattides " 15

Blattina 15

"Blattoiden" 15

"Blue oil " against termites 77

Books, destruction by termites 76

Brachylsena discolor, wood resistant to termite attack 81

Bridge timbers, destruction by termites 13

Brush treatments of —

chestnut poles against insects 9-10

forest products against termites * 76

Bupresiis rufipes, damage to chestnut telephone and telegraph poles 8

Cable conduits, wooden, destruction by termites 13, 75

Cage for rearing and studying termites 20-22

Calotermes, copulation 51

Calotermes flavicollis —

nymphs, development 41

soldier, metamorphosis 35

Camponotus pennsylvanicus —

association with termites 70

damage to chestnut telephone and telegraph poles 8, 9

Cannibalism among termites 46-47

Carbolineums against termites 9, 10, 76

55514—16 2 . 87

88 INSECTS INJURIOUS TO FOREST PRODUCTS.

Cedar— Page.

California white, wood eaten by termites 81

California white, wood not eaten by termites in experiment 80

Eastern red. (-See Juniper.)

Eastern white. (See Cham-secy pans thyoides.)

incense. (See Libocedrus decurrens.)

red. (See Juniper us virginiana.)

Western red. (See Thuja plicata.)

Cedrus atlantica, wood reported "immune" to termite attack 81

Cedrus deodar, wood reported "immune " to termite attack 81

Chamxcyparis thyoides, heartwood comparatively resistant to termite attack 81

Charring forest products against termites 9; 76

** Charring the butt " treatment of chestnut poles against insects 9

Chestnut borer, two lined. (See Agrilus bilineatus.)
Chestnut telephone and telegraph poles —

damage by wood-boring insects 1-12

prevention of injury by insects 8-11

Chestnut telephone-pole borer. (See Parandra brunnea.)

Chestnut timber, time for utilization to prevent damage by termites 79

Chestnut timber worm. (See Lymexylon serieeum.)

Chlorinated naphthalene against termites 77

Combretum prophyrolepsis, wood resistant to termite attack 81

i ' Compositi on b oards ' ' —

destruction by termites 13

treatment to render them termite proof 79

Copaifera mopani, wood resistant to termite attack 81

Cornus florida, ripening of pollen synchronous with first swarm of termites 48

Corrodentia 14

Cremastogaster lineolata —

association with termites 70

damage to chestnut telephone and telegraph poles 8

Creolin treatment of chestnut poles against insects 9

Creosote —

coal-tar, a fungicide 76

coal-tar, impregnation of forest products against termites 76

treatments of chestnut poles against insects 9-11

Creosotes against termites '. 76

Cryptotermes cavifrons, bibliographic reference , 83

Cupressus macrocarpa, heartwood comparatively resistant to termite attack 81

Cupressus sempervirens , wood eaten by termites 81

Cylinder-pressure process treatments of chestnut poles against insects 10-11

Cypress —

bald, heartwood comparatively resistant to termite attack 81

desert, of Australia, wood comparatively resistant to termite attack 82

European. (See Cupressus sempervirens .)
Monterey. (See Cupressus macrocarpa.)

^Beal, red, " wood more resistant to termite attack than "white deal " 81

*' Deal, white, " wood less resistant to termite attack than "red deal " 81

*'DeaIwood," wood attacked readily by termites 82

^'Dinglas." (See Eugenia bracteata var. roxburghii.)

Pipping treatments of forest products against termites 76

Documents, destruction by termites 76

Poryphora decemlineata, bibliographic reference 85

Harwig. (See Forficula.)

INDEX. 89

Page.

Elateridse, damage to chestnut telephone and telegraph poles 8

Electrotype blocks, wooden, destruction by termites 76

Embiidse 15

"Ergatoids" —

definition of term 28

development 34

Eugenia bracteata var. roxburghii, wood resistant to termite attack 80

Eutermes cinereus, bibliographic reference 83

Eutermes laeustris —

differentiation of three castes in embryo 17

nasuti, development 41

Eutermes rippertii, bibliographic reference . 84

Eutermes (Rippertiif), bibliographic references 15, 84

Eutermes sp. , soldier, metamorphosis 35

Eutermes spp., displacement of digestive and excretory organs in distended

queens 67

Fence posts, rails, and boarding, destruction by termites 13, 75

Floors, destruction by termites 14

Forest products, crude and finished, destruction by termites. 13, 75-76

Forficula . 15

Forficulina .' 15

Formica exsectoides, association with Leucotermes fiavipes 70

Formica schaufussi, association with Leucotermes fiavipes 70

Foundations, wooden, of buildings, destruction by termites. 13, 76

Greenheart. (See Nectandra rodisei.)

Hamitermes ( Termes) tubiformans, bibliographic reference 83

Hemlock, western, wood not eaten by termites in experiment 80

Hemlock, wood eaten by termites 81

Hog- wallows and prairie mounds, relation of termites thereto, bibliography 85

Homalota sp., " guest," or inquiline, of Leucotermes fiavipes 71

Homovalgus squamiger —

"guest," or inquiline, of termites 71

in galleries of Leucotermes fiavipes 64

Hopea plagata, wood resistant to termite attack. 80

Hydrocyanic-acid gas against termites 79

"Indian oak." (See Tectonia grandis and Teak.)

Insects damaging chestnut telephone and telegraph poles, prevention of injury . . 1-12

"Ipil." (See Afzelia bijuga.)

"Tronwood, black." (See Olea laurifolia.)

"Ironwood." (See Sideroxylon.)

Isoptera 13, 14, 15, 16

" Jarrah," wood comparatively resistant to termite attack 82

Juniper, heartwood comparatively resistant to termite attack 81

Juniperus virginiana poles less susceptible to insect attack than chestnut poles. 10-11

Kerosene against termites 79

Larva, forms to which term is applicable 29

"Leadwood." (See Combretum prophyrolepsis.)

Leucotermes '.". 13-16

Leucotermes fiavipes (see also Termites of Eastern United States and Termes
fiavipes) —

adults, colonizing winged, description 30-31

association with ants 70

cannibalism 46-47

90 INSECTS INJURIOUS TO FOREST PRODUCTS.

Leucotermes fiavipes, etc. — Continued. Page.

colonies, new, establishment 49-53

copulation 50, 51

' description of forms, or castes 28-31

distribution 14

egg, description ' 31

egg laying, rate 51, 52, 53

eggs, seasonal occurrence in colony 43

former studies thereon 17-19

"guests, " or inquilines . 71-72

installation in termitarium 22

neoteinic reproductive forms, seasonal occurrence in colony 44

neoteinic royalty, mating 51

number of individuals in colony 25-27

nymph, primary form, description .' 30

nymph, secondary form, description '. 30

nymphs, development 37-42

nymphs of reproductive forms, seasonal occurrence in colony 43-44

parasites : 72

reproductive forms, description 65-68

reproductive forms, occurrence, historical 54-65

seasonal variation in colony 43-45

sense organs 31-32

soldier, description 28-29

soldiers, seasonal occurrence in colony , 45

successive stages in development of the various forms or castes 36

summary and conclusions based on results of experiments 72-75

swarming dates 68-69

swarm, or so-called nuptial flight . . . . . 48

worker, description 28

Leucotermes lucifugus (see also Termes lucifugus) —

antennal segments, increase 35

colony formation, bibliographic reference 73

former studies thereon 18

forms, bibliographic reference 36

molts, number .,r 35

nymphs, development 41

reproductive forms, description 65-68

reproductive forms, occurrence 53-54, 57-58, 59, 60

soldier, metamorphosis 35

studies on life history and internal anatomy 17

swarming dates 68, 69

Leucotermes virginicus (see also Termites of Eastern United States) —

adults, colonizing winged, description 30-31

cannibalism 46

description of forms, or castes 28-31

distribution 14

eggs, seasonal variations in colony 43

"guests, " or inquilines 72

mention 18

neoteinic reproductive forms, seasonal occurrence in colony 44

number of individuals in colonies 27

nympn, primary form, description 30

nymph, secondary form, description 30

INDEX. 91

Leucotermes virginicus, etc. — Continued. • page

nymphs, development 38-42

reproductive forms, occurrence 54, 57-58, 60, 60-62, 64-65

seasonal variations in colony 43-45

soldier, description 28-29

soldiers, seasonal occurrence in colony 45

summary and conclusions based on results of experiments 72-75

swarming dates ' 69

swarm, or so-called nuptial flight 48

worker, description 28

Leucotermitinae 16

Libocedrus decurrens, heartwood comparatively resistant to termite attack 81

Lignumvitse, wood not resistant to termite attack 79

Linseed oil against termites 77

Lit ea obtusata, wood partially resistant to termite attack , . 80

Locust, black, heartwood comparatively resistant to termite attack 81

Lumber piled on ground, destruction by termites 13-75

Lymexylon sericeum —

in chestnut, destructive work 7

true queen of Leucotermes flavipes in abandoned burrow 58

Mahogany. (See Swietenia mahoghani.)

Mallophaga 15

Mesotermitidse 13, 16

Metatermitidae 16

Mine props, destruction by termites 13, 75

"Molave." (See Vitex littoralis .)
"Mopani tree." (See Copaifera mopani.)

Naphthalene, chlorinated, against termites 77

Narra. (See Pterocarpus indicus.)

Nasuti 41

Nectandra rodisei, wood resistant to termite attack 79

" Neoteinic " reproductive forms of termites, development 34

"Neotenes" —

definition of term 28

development 34

Neuroptera 14

Nursery stock, forest tree, prevention of damage by termites 79

Nymph, forms to which term is applicable 29

Odonata 15

Olea laurifolia, wood resistant to termite attack 81

"Open-tank " treatments of chestnut poles against insects 9-11

Orthoptera 15

Paraffin wax against termites 77

Parandra brunnea —

character of insect 3-4

damage and loss to chestnut poles, extent 6

damage to chestnut telephone and telegraph poles, general account. . . . 1-3, 3-7

distribution. 4

importance as injuring chestnut poles 5-6

injury, character 4-5

Pediculus 14

Perlidse 3.5

"Peroba." (See Aspidosperma spp.)

Philotermes fuchsii, association with Leucotermes flavipes ' 72

92 IXSECTS INJURIOUS TO FOEEST PRODUCTS.

Page.

Philotermcs pennsylvariicus. "guest," or inquiline, of Leucotermes flavipes 72

PMlotermes pilosus, ••guest/' or inquiline. of Leucotermes flavipes 71-72

PMlotermes sp. (possibly/wcft-sn), association with Leucotermes virginicus 72

Photographs of termites for illustrations, methods of obtaining them 82-83

Pine-
bull, ■wood eaten by termites 80

clear, of Australia, wood less resistant to termite attack than that of red

pine 82

longleaf. (See Pinus palustris.)

Oregon, wood eaten by termites 80

red. of Australia, wood comparatively resistant to termite attack 82

timber, time for utilization to prevent damage by termites 79

Pinus palustris, heartwood resistant to termite attack 81

Platyptera 13. 15

Plecoptera 15

Podura 14

Poles-
bean and hop, destruction by termites 13. 75

chestnut telephone and telegraph, damage by wood-boring insects 1-12

chestnut telephone and telegraph, prevention of injury by insects 8-11

statistics, list of publications thereon 12

telephone and telegraph, destruction by termites 13, 75

Polym&chus brevipes, damage to chestnut telephone and telegraph poles 8

Prairie mounds and hog wallows, relation of termites thereto, bibliography 85

Prionoxystus robinix, royal couple of Leucotermes flavipes in old burrow '60

Prionus sp.. damage to chestnut telephone and telegraph poles 8. 9. 10

Protermitida? 16

'•Protoblattoiden*' 15. 16

Pseudoneuroptera 14. 15

Psocidae 15

Psocids. wingless, "guests." or inquilines, of Leucotermes flavipes — 71

Pterocarpus indicus. wood partially resistant to termite attack SO

Railroad ties, destruction by termites 13. 75

Rearing termites, cage therefor 20-22

Redwood-
California. (See Sequoia sempervirens.)

wood eaten by termites SI

Rhinotermes taurus. resting stage of larva 41

Saccharine solution and arsenic against termites 77

Sand setting for chestnut telephone and telegraph poles against insects 9

Sequoia sempervirens, wood not eaten by termites in experiment 80

Sideroxylon. wood resistant to termite attack 79

Silo timbers, destruction by termites 13

Snyder, Thomas E., paper —

''Biology of the Termites of the Eastern United States, with Preventive

and Remedial Measures -; 13-85

"Damage to Chestnut Telephone and Telegraph Poles by Wood-boring

Insects" - 1-12

Spirittine treatments of chestnut poles against insects 9

Spruce, wood eaten by termites - SO

£tone, small broken, setting for chestnut telephone and telegraph poles against

insects 9

1 ' Supplementary " reproductive forms of termites, development 34

INDEX. 93

Page.

Swietenia mahogha?ii, wood resistant to termite attack 79

Tachyporus jocosus, "guest, ' ' or inquiline, of Leucotermes flavipes 71

Tar treatment of chestnut poles against insects 9

Teak. (See Tectonia grandis.)

"Teak, Burmese." (See Tectonia grandis .)

"Teak, Jungle." (See "Teak, Malabar.")

"Teak, Malabar, " wood comparatively resistant to termite attack 82

"Teak, Pukka, ' ' wood comparatively resistant to termite attack 82

Tectonia grandis (see also Teak) —

wood resistant to termite attack 78, 80

Telephone-pole borer, chestnut. (See Parandra brunnea.)

Tent pins, destruction by termites 75

Tent ridge poles, destruction by termites 75

Termes 14

Termes bellicosus —

longevity 46

position of spiracles in distended queens 66-67

Termes flavipes (see also Leucotermes flavipes) —

bibliographic references 14, 84, 85

damage to chestnut telephone and telegraph poles 7-8

Termes frontalis, bibliographic reference 84

Termes horni, resting stage of soldier. . .• V 41

Termes lucifugus (see also Leucotermes lucifugus) —

bibliographic reference 84

Termes malayanus, copulation 50

Termes obscuripes, resting stage of larva 41

Termes spp., differentiation of three castes in embryo 17

Termitarium for study of termites of Eastern United States 20-22

' ' Termiten " 15

Termites, damage to chestnut telephone and telegraph poles 2,3, 7-8

Termites of Eastern United States (see also Leucotermes flavipes and L. virginicus) —

adults, colonizing winged, description , . 30-31

adults, winged sexed, function 32-33

association with ants 70

bibliography 83-85

biological experiments 20-22

biology, with preventive and remedial measures 13-85

cannibalism 46-47

caste differentiation 33-36

castes 27-33

castes, function 31-32

classification „ 14-16

colonies, new, establishment 49-53

colony, location in winter 45-46

colony, seasonal variations 43-45

communal organization 22-27

copulation > 50-53

damage to forest products 75-76

descriptions of forms, or castes 27-31

development, progressive, of nymphs 36-43

duration of development and life 46

egg-laying rate '. 50-53

eggs, description 31

94 INSECTS INJURIOUS TO FOREST PRODUCTS.

Termites of Eastern United States, etc. — Continued. Page.

eggs, seasonal occurrence in colony 43

forms, or castes, situation in nest 47-48

functions of castes 31-32

"guests, " or inquilines 71-72

historical 16-19

"immune" (i. e., resistant) woods 76-82

larvae, description 31

larvae, situation in nest 47, 48

life cycle 33^6

metamorphosis 33-36

neoteinic reproductive forms, seasonal occurrence in colony 44

neoteinic reproductive forms, situation in nest 47

neoteinic royal individuals, function 33

nests, situation 22-25

number of individuals in colonies 25-27

nymphs of reproductive forms, seasonal occurrence in colony 43-44

nymphs, primary form, description 29-30

nymphs, progressive development 36-43

nymphs, secondary form, description 30

nymphs, situation in nest 47

parasites 72

photographs, methods of obtaining them for illustrations 82-83

polymorphism 27-33

preventives of injury 76-82

relation of hog wallows and prairie mounds thereto, bibliography 85

remedies 76-82

reproductive forms, description 65-68

reproductive forms, occurrence 53-65

reproductive forms , situation in nest 47

resistant woods 76-82

"royal cell," situation in nest 47

royal pair and other reproductive forms 53-68

soldiers, seasonal occurrence in colony 45

seasonal variations in colony 43-45

sense organs 31-32

situation of different forms in nest 47^48

soldiers, description 28-29

soldiers, function 32

soldiers, situation in nest 48

summary and conclusion based on results of experiments 72-75

swarming dates 68-69

swarm, or so-called nuptial flight 48-49

winter location of colony 45-46

workers, description 28

workers, function 32-33

workers, situation in nest 48

Termitidae 15

Termitina 15

Termitophilus insects 71-72

Termopsinae 16

Termopsis 16

INDEX. 95

Termopsis angusticollis — Page.

duration of development and life 46

nymphs, development 39-41, 43

soldier, metamorphosis 35

Termopsis laticeps, bibliographic reference 83

Thuja plicata, heartwood comparatively resistant to termite attack 81

Timber, time for utilization to prevent damage by termites 79

' ' Tindalo. ' ' (See Afzelia rhomboidea.)

Tmesiphorus carinatus, "guest," or inquiline, of Leucotermes flavipes 72

Tree boxes, wooden, destruction by termites 76

Trichopsenius, "guest," or inquiline, of Leucotermes flavipes 71

Turpentine, repellent to termites 81

Tyloses, significance of presence in woods, bibliographic reference 80

"Vaalbosch." (See Braehylsena discolor.)

Valgus canaliculars, "guest," or inquiline, of termites 71

Vitex littoralis, wood resistant to termite attack 80

Walnut, black, heartwood comparatively resistant to termite attack 81

Well timbers, destruction by termites 13

Wharf timbers, destruction by termites '. 13

White ants. (See Termites.)

Wood creosote treatments of chestnut poles against insects 9-10

' < Wood lice . " (See Termites . )

Wood, painted, eaten by termites 80

Wood preservation, list of publications thereon 12

Woodwork of buildings, destruction by termites 13, 14, 75, 76

"Yacal." (See Hopea plagata.)

Zinc chloride against termites 77, 79

o

%y and Plant Quarantia§