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Bulletin 407
1921

Agric . - Foresti > . Adam Library

AUGUST, 1921

CORNELL UNIVERSITY ^^ . -^
AGRICULTURAL EXPERIMENT STA

THE PAINTED HICKORY BORER

E. H. DUSHAM

ITHACA, NEW YORK
PUBLISHED BY THE UNIVERSITY

*c

CONSENTS

Classification, history, and synonymy sr-i,-J.

Distribution .•*•* .'...- ^*i^Pi8i

Food plants 'Z&'&H—- • - l82

Economic importance ... -. . ^j&r- 184

Descriptions >"!* 186

The adult 186

The egg • • 187

The larva 187

The pupa 188

Life history and habits 188

The adult 188

Appearance of the adults 188

General activities 188

Feeding habits 189

Mating habits 190

Oviposition 190

The egg 191

Time of hatching 191

The larva 192

Activities of the newly hatched larva 192

Larval burrows 192

The pupal cell 193

The prepupal stage 194

The pupa 194

Pupation 194

Activities of the pupa 194

Transformation to adults 195

Seasonal history 195

Natural enemies 195

Recommendations for control and prevention of injury 196

Bibliography 198

173

'96025

THE PAINTED HICKORY BORER

Cyllene caryae Gahan

Order, Coleoptera Family, Cerambycidae

E. H. DUSHAM

The hickory is one of the most important of our American hardwoods,
not only on account of its fruit, but especially because of the valuable
qualities of its wood. " Tough as hickory " is not only a familiar expres-
sion but an apt phrase as well, for its wood is heavy, hard, strong, and
exceedingly tough. It is on account of these qualities that, more than
any other wood, hickory is used in the manufacture of such commodities
as are subject to excessive strain. Consequently, more than one-half of
the hickory cut today is tised in the construction of vehicles or their parts,
and at least one-third is made into handles for articles in which strength
and toughness are the main considerations. In addition to these, agri-
cultural implements consume three per cent of the entire cut, while the
remaining five per cent is used in the manufacture of athletic goods, porch
furniture, and various other commodities. The factory uses of hickory as
given by Kellogg (19 14)*, may be summarized in percentage as follows:

Vehicles 61

Handles 31

Agricultural implements 3

Sporting and athletic goods i

Other uses 4

•

Like most other trees, the hickory is subject to various insect injuries
which reduce the amount of timber suitable for manufacturing purposes.
It harbors a large host of different pests, some attacking the foliage,
fruit, and wood of living trees; others the wood of recently dead and
felled trees, sawlogs, handles, poles, and other unseasoned products from
which the bark has not been removed; while still others inflict serious
injury on the sapwood, even after seasoning, causing what is commonly
known as the " powder post " injury.

Prominent among the insects of the second category is the painted
hickory borer, Cyllene caryae Gahan.

CLASSIFICATION, HISTORY, AND SYNONYMY

The painted hickory borer belongs to the order Coleoptera, suborder
Phytophaga, family Cerambycidae, and subfamily Cerambycinae. The

1 Dates in parentheses refer to Bibliography, pages 198 to 203.

175

BULLETIN 407

family 'Cerambycidae includes medium- to large-sized beetles, of graceful
form, and, in most cases, with very long antennae, whence the common
name Longicorns,1 or long-horned beetles. For the most part their larvae
are wood borers, some of the most important enemies of our forest and
shade trees being found among their number.

The genus Cyllene, to which this species belongs, together with several
other groups now ranking as distinct genera, was formerly included in
the old genus Clytus. Because of definite, well-marked characters,
Newman (1840) described it as a distinct genus as follows:

Facies Clyti: caput parvum, fer& pronum, antennae maris corpore longiores, feminae
valdS breyiores, ii-articulatae; articulis externis manifestd crassioribus, 3-6 apice
spini brevi armatis: prothorax latus, dorso convexus, lateribus rotundatis dente postico
armatis: elytra convexa, apice i-spinosa: pedes simplices.

Cyll. spinifera. Fusca, obscura, lanuginosa: prothorax maculis duabus glabris orna-
tus: antennarum articuli basi cinerei: elytra fusca, maculis fasciisve lanuginosis cinereis
ornata. (Corp. long. .7 unc. lat. .225 unc.)

Inhabits South America. Specimens taken by Mr. Darwin are in the cabinet of
the Entomological Society. I find no description of this insect in Serville's " Nouvelle
Classification", but it is by no means uncommon in collections: it bears as close a
resemblance to Clytus, as Phacodes to Callidium.

This genus includes about thirty-five species, all of them inhabitants
of the new world. Their distribution is shown in table i.

TABLE i. THE DISTRIBUTION OF THE SPECIES OF CYLLENE

Place

Num-
ber of
species

Place

Num-
ber of
species

Place

Num-
ber of
species

Brazil

17

Argentina ....

-t

Patagonia

i

Mexico . .

K

United States. . .

ii

Bolivia

i

Chile *

2

Venezuela

-i

Nicaragua

i

Costa Rica

I

Honduras

i

Panama .

2

Hawaiian Islands

I

Cuba

i

Guatemala. . . .

I

Paraguay

2

Canada

2

Much confusion has existed regarding the identity of the species in
question. The earliest writers confounded it with the locust borer
(Cyllene robiniae Forster), it being then the common supposition that the
latter species had two broods annually, the first emerging in spring and
attacking the hickory, and the second emerging in late summer and
attacking the locust. Not only were the insects confused, but, until
quite recently, the hickory species itself was incorrectly named, being
known as Cyllene pictus Drury, a synonym of Cyllene robiniae Forster.
Consequently Gaylord (1843), Harris (1862), and other writers mentioned
this species as being found on goldenrod in late summer, and attacking
the locust, while Fitch (1859) and Rath von (1860) stated that the locust

THE PAINTED HICKORY BORER 177

borer (Clytus robiniae) was a common borer in the trunks and limbs of
black walnut, and was also quite destructive to hickory hop poles and
firewood.

Kirkpatrick (1856) seems to have been the first to realize that the
locust and hickory borers were two distinct species. He stated that two
species of Clytus were found in Ohio, one the locust borer and the other
the hickory borer. Furthermore, he said that Cyltus pictus was found
only in hickory wood, where it generally bored parallel with the grain,
causing powder post. He gave brief data concerning the time of pupation
and the emergence of the adult. However, he says that he never found
the adult depositing eggs in wood cut more than a year, even when it
had no other wood in which to deposit the eggs.

Horn (1863), referring to the hickory borer under the name Arhopalus
pictus Drury, described the character of the larval galleries in hickory
wood.

The next reference to the hickory borer was made by Walsh (1864).
Discussing the spread of the locust borer westward, he mentioned that as
much as six years previous to the arrival of that insect he had split an
adult male Clytus pictus out of a stick of hickory in Rock Island, Illinois,
and that in the course of the next two or three years, he had found two
more specimens in the same neighborhood. He believed this was proof
that the locust and hickory species were distinct, for the hickory species
had evidently been in existence in Illinois all the time, but had fed on
hickory and walnut quite unnoticed until the arrival of the locust borer.
He also added that Professor Sheldon, of Davenport, Iowa, had told him
that for many years back he had repeatedly discovered Clytus pictus in
hickory wood, and that, so far as he was aware, the locusts of Davenport
had not been attacked by this insect.

Later, Walsh (1864) definitely showed, by anatomical characters, that
the locust and the hickory species were distinct. In studying the adults
of both species, he found that there were quite remarkable differences
in the males, but none in the females. These differences in the males
he tabulated as follows:

Hickory feeding "b Locust feeding "b

1. Antennae, when relaxed and laid i. Antennae, when relaxed and laid
close and straight along the back, reaching close and straight along the back even in
beyond the tip of the elytra by the whole the specimen which has the longest ones,
length of the terminal joint (n). not attaining the tip of the elytra by a

space equal in length to the two terminal
joints (10 and n).

2. Antennae from % more robust to 2. Antennae much less robust, except
twice as robust, especially towards the the few last joints, and less tapered from
base. base to tip.

178 BULLETIN 407

Hickory feeding 'b Locust feeding 'b

3. Terminal or nth joint of antennae 3. Terminal or nth joint of antennae
full ? longer than the penultimate, and scarcely | or \ longer than the penultimate,
composed of two portions connected by the division into two portions barely
an indistinct connate suture foreshadowing discoverable, and the terminal portion
a 1 2th joint, (as in Purpuricenus tf and not suddenly slenderer from base to tip.
in Tragidion annulatum^ Lee.,) which

suture is more distinct on the inferior
surface. The basal portion of the nth
joint as long as joint 10, the terminal
portion, which is suddenly slenderer from
base to tip, more than £ as long as joint 10.

4. Elytra widened at base and tapered 4. Elytra much less tapered and shaped
towards their tip, so that the two together exactly as in the 9 of both the two races,
just before the extreme tip equal the basal i.e., with the lateral edges subparallel.
width of one of them.

5. The 2nd or W-shaped band on the 5. The W-shaped band on the elytra
elytra in two of the Philadelphia specimens colored yellow, exactly like the other
and the Illinois specimen whitish, in the bands, in all my 15 specimens.

other Philadelphia specimen centrally

whitish, but decidedly varied with yellow . : >

on the two exterior arms of the W.

6. Legs proportionally %-% longer and 6. Legs proportionally no longer or
stouter than in 9 . stouter than in 9 .

The 9 antennae are exactly alike, being in both races a little more than £ as long
as the body, with the terminal joint equal in length to the penultimate or perhaps
very slightly longer, and no perceptible difference in the robustness of the whole antenna.
The general appearance of the two 9 9 and of the cf of the locust-feeding race is
very similar, but, owing to the shape of the elytra, the cf of the hickory-feeding race
has a different and Leptura-like habit. So closely indeed does the cf of the locust-
feeding race resemble the 9 of both races that until a recent period I had always sup-
posed that all my specimens of that race — some 30 or 40 in number — were 9 9 , and
that the unique cf which I possessed of the hickory-feeding race, was the normal cf of
the species. In all the 9 9 of both races the W-shaped band on the elytra is as
yellow as the other bands. Whether there is any distinction in the larva state is
unknown, as the larva of the locust-feeding form has never yet been critically
examined. * *

* * * It is a doubtful and disputed question in Entomological Archaeology, whether
Drury's name pictus or Forster's name robiniae has the priority, as Drury was the
first to describe the insect, and Forster the first to name it. We may therefore,
with even-handed justice, appropriate the name robiniae to the locust-feeding race
with short and slender cf antennae and leg, which appears in September, and the
name of pictus to the hickory-feeding race with long and robust cf antennae and legs,
which appears in May and June.

Both Forster's and Drury's descriptions, however, show that the species
they refer to are identical, and are the ones which inhabit the locust. The
names Cyllene robiniae and Cyllene pictus are, therefore, synonymous,
and Walsh's application of this name to the hickory species was accordingly
unjustified.

Riley (1876) gave a brief description of the larva and imago of the
hickory borer, together with a few notes on its life history, and men-
tioned the characteristics distinguishing it from Cyllene robiniae.

THE PAINTED HICKORY BORER 179

Horn (1880) differentiated the hickory-boring species from the locust-
boring species as follows:

C. pictus Drury

The prosternum is longer than wide, truncate at tip and not prolonged. The second
joint of hind tarsus glabrous at middle. The elytra are obliquely truncate at tip but
not prolonged. The antennae of the male are always longer than the body resembling
in this respect antennatus.

Lives in Hickory wood and appears early in Spring, and is abundant wherever that
tree occurs and is often very destructive.

C. robiniae Forst.

Closely resembles pictus in form and coloration but differs in the antennal characters
of the male and the structure of the hind tarsi. The prosternum is as broad as long
being wider than in pictus and also truncate at tip. The legs are also shorter than in
that species, the hind thighs not reaching the tip of the abdomen in the male.

Infests Locust wood (Robinia pseud-acacia), and appears in the Autumn. I have
seen a variety of this species with the W-shaped band entirely obliterated.

Moffat (1882) in speaking of the confusion that existed in literature
between the hickory and locust borers, differentiated the two species as
follows :

In pictus the body is uniformly more slender and tapers more behind than robinia.
In pictus the antennae is [sic] decidedly longer, that of the females reaching to the
end of the body, and that of the males beyond. The third or W-band on the wing
covers is noticeably more delicate than in robinia, and quite white in contrast to the
yellow of the other's markings, a characteristic I have never seen in any robinia I ever
met with (and I see them here in hundreds every fall feeding on the Goldenrod), and
one which would of itself make it quite easy to pick out my pictus, male or female,
from amongst any quantity of robinia they might be mixed with.

That same year, Horn (1882), fearing that there might be some difficulty
in properly distinguishing the species by the characters mentioned by
Moffat (1882), again referred to the more definite characters for disting-
uishing the two species as given by him in 1880, and added, as another
distinguishing character, that in pictus the W-shaped band rarely joins
the transverse band, while in robiniae this band nearly always joins the
transverse band at the suture.

Riley and Howard (1892) mentioned specimens of the hickory borer
having been sent to them with the complaint that they were abundant in
dwelling houses. The insects had emerged from hickory wood that was
used for fuel.

Bruner (1893) stated that hickories in Nebraska were subject to attack
by this beetle. He gave a brief description of the adult, also mentioning
the time of its emergence, and suggested control measures.

Hopkins (1895) mentioned finding a hickory log containing a great
number of the larvae and pupae of the hickory borer, and having numerous
holes in the bark, indicative of the emergence of the adults. However,
no adults were found in the wood altho larvae and pupae were abundant.

Webster and Mally (1897) reported the receipt of a section of an osage-

180 BULLETIN 407

orange post, one and one-half feet long and four inches in diameter, together
with a complaint that a worm was destroying many fence posts of this
kind of timber. The insect causing the damage was the hickory borer,
and, in all, twenty-seven adults emerged from this one piece of wood.

Hopkins (1898), referring to the developing of broods for three years
in succession in hickory by the Cerambycidae, mentioned the hickory
borer as one of them. He supposed that the late arrivals were the result
of retarded development, but an examination of the wood showed that
new wood was being continually infested.

Webster and Mally (1898) discussed this insect at length. They
described the adult, and gave brief notes on the life history. They talked
of the destructive work of the insect on osage-orange fence posts and
offered suggestions for the prevention of injury.

The next discussion of this species was furnished by Felt (1905) who
reviewed the history of the insect up to that time, gave notes on the life
history, described the adult and larva, and listed the natural enemies
attacking it.

Until 1908 the insect had been called Clytus or Cyllene pictus, which,
as has already been stated, was a synonym of Cyllene robiniae. Realizing
this mistake, Gahan (1908 a) described the hickory borer as a new species,
giving it the name of Cyllene caryae. He further stated that it could be
distinguished from Cyllene robiniae as follows:

From C. robiniae, Forst., which they greatly resemble in markings, C. caryae and
its varieties may be distinguished by the thicker, longer, and usually darker-coloured
antennae, by the two widely separated yellowish or whitish spots on each of the meta-
thoracic episterna, the limitation of the sexual puncturation of the male pronotum to
the anterior part, and in having the intercoxal process of the prosternum nearly parallel-
sided.

In C. robiniae the antennae are generally reddish brown in colour, somewhat slender,
as a rule distinctly shorter (never longer) than the body in the male; the yellow pubes-
cence forms an almost continuous band on each of the metathoracic episterna, when
it is broken up into two spots the interval between the spots is always narrow; the
intercoxal process of the prosternum widens out posteriorly, its sides being curved
instead of almost parallel. The sexual puncturation of the pronotum of the male is
much more extensive than in C. caryae ; it covers the greater part of each side, forms an
anterior transverse band, and two bands running backwards on the disk, diverging a
little behind and dilating each into an oval or rounded spot between the middle and
the base; it is similar in character to that occurring in the North- American species
C. decora, Oliv., and the Brazilian C. mellyi, Chevr.

Daecke (1910) mentioned a character which would help to distinguish
the two species at a glance. He stated that he was not aware that the
upper side of the abdomen of beetles had been used as a means of differ-
entiating closely allied species, and accordingly exhibited two specimens
of Cyllene pictus and Cyllene robiniae with the wings spread. The upper
side of the abdomen of pictus showed reddish, with the anal segment
black, while that of robiniae was all black. Both species had two rows

THE PAINTED HICKORY BORER 181

of golden yellow lunules along the abdomen, caused by yellow pile; but
the anal segment of pictus showed one median, dorsal, narrow and oblong
spot of gold, while the corresponding spot in robiniae was obtusely tri-
angular.

Another somewhat extensive discussion of this insect was set forth by
Webb (1911). A list of the trees attacked was given and also, the range
of the insect. Brief notes were furnished concerning the life history of
the species and recommendations were made for preventing the injury
which it causes.

The synonymy of this species is as follows:

Arhopalus pictus Drury

1863 Horn, G. H. Proc. Erit. Soc. Philadelphia, vol. i, p. 30.

Clytus pictus Drury

1864 Walsh, B. D. Proc. Ent. Soc. Philadelphia, vol. 3, p. 420-422.

1882 Moffat, J. A. Can. ent., vol. 14, p. 200.

1883 Packard, A. S. Guide to the study of insects, p. 497.

Cyllene pictits Horn (nee. Drury)

1880 Horn, G. H. Trans. Amer. Ent. Soc., vol. 8, p. 135-136.
1882 Horn, G. H. Can. ent., vol. 14, p. 240.
1887 Leng, C. B. Entomologica Americana, vol. 2, p. 196.
1897 Wickham, H. F. Can. ent., vol. 29, p. 149.

Cyllene picta Drury

1884 Harrington, W. H. Rept. Ent. Soc. Ontario, 1883, p. 46.

DISTRIBUTION

The distribution of the hickory borer is in all pfobability coextensive
with that of its food plants, the hickories. Kirkpatrick (1856) recorded
it from northern Ohio as breeding in hickory. Walsh (1864) stated that
about six years previously he had split this insect from hickory at Rock
Island, Illinois, while Professor Sheldon, of Davenport, Iowa, informed
him that he had found it in hickory at that place for many years back.
Walsh (1866) reported the insect rare in Illinois, having taken only three
specimens during a seven-years collection. He stated, however, that it
was common near Philadelphia, Pennsylvania, in walnut and hickory.
Larvae of this species were sent to Walsh from Kansas in 1867, for identi-
fication. Rogers (1880) reported that it was one of the common species
in Ontario. Smith (1890) reported the insect as being generally dis
tributed, tho rare in New Jersey. In 1892, C. C. Black sent several
specimens from Connecticut to Riley, stating that they had emerged from
hickory firewood. Bruner (1893) mentioned it as being very destructive

182 BULLETIN 407

in Nebraska. The same year, Hopkins (1893) reported it from West
Virginia, where it was boring in mulberry and hickory. Johnson (1898)
mentioned its time of appearance at Philadelphia, Pennsylvania, while
Dr. Hamilton reared it in the southwestern part of that state. Webster
and Mally (1897) recorded it as being injurious to fence posts in Ohio.
That same year, Lintner (1897) received specimens from Brooklyn, New
York, where they had emerged from firewood. Wickham (1897) listed
it from Ontario and Quebec. Lugger (1899) recorded it among the beetles
injurious to fruit-producing plants in Minnesota. Smith (1900) stated
that it was found thruout the state of New Jersey. Moffat (1902)
received specimens of this insect from London, Ontario. Felt (1905)
reported it as common in New York State where, it was found frequently
on hickory. Gahan (1908 a) recorded a variety of this species from Texas,
and another variety from Villa Lerdo, Durango, Mexico. Douglass
(1912) stated that it was not a very common species in Indiana. Webb
(1911) believed that its range was coextensive with that of the hickory.
The writer has taken this insect at State College, Pennsylvania, at Welland,
Ontario, and at Ithaca, New York. In all three localities, it was abundant
during May and June.

According to the above data the insect has been reported from Ontario,
Quebec, Connecticut, New York, Ohio, New Jersey, Pennsylvania, West
Virginia, Minnesota, Illinois, Iowa, Kansas, Nebraska, Indiana, Texas,
and Durango, Mexico.

FOOD PLANTS

The hickory borer is injurious mainly to the hickories, especially the
shagbark variety (Carya ovata [Mill] K. Koch). In 1862, T. B. Ashton
bred it from black walnut (Juglans niger L.) in Pennsylvania, where he
said the larvae were not uncommonly found in that tree. Riley (1880)
stated that it also . attacked pecan (Carya illinoensis [Wang] K. Koch)
and butternut (Juglans cinerea L.). Hopkins (1893) recorded it as
attacking the dead branches and small trees of mulberry (Morns sp.)
in West Virginia. Beutenmuller (1896) added honey locust (Gleditsia
triacantkos L.) to its list of food plants. LWebster_and Mally (1897)
said that it had developed a fondness for osage orange (Madura pomifera
[Raf.] Schneider) hardly second to its partiality for hickory. Moffat
(1902) received from A. Morde, London, Ontario, specimens of the insect
which had been reared on a stick of bitternut hickory (Carya cordiformis
[Wang] K. Koch). Fisher and Kirk (1912) stated that they had reared this
insect from wild grape (Vitis sp.). Blatchley (1910) said that it occurred
on the elm (Ulmus sp.) and that the larvae bored in the wood of that
tree. In July, 1917, Henry Dietrich, of Ithaca, New York, reported

THE PAINTED HICKORY BORER

<

183

PLATE I

THE PAINTED HICKORY BORER

i, Male and female. 2, Eggs (slightly enlarged). 3, Eggs (much enlarged). 4,
Larva (natural size). 5, Work of larva in hickory.

1 84 BULLETIN 407

finding a female with her ovipositor inserted in a branch of black birch
(Betula lenta L.). The same year J. N. Knull, of Harrisburg, reported
that he had bred the insect on hackberry (Celtis sp.). At State College,
Pennsylvania, the writer has observed this insect running over the trunks
of dead pitch pines (Pinus rigida Mill), and has seen the females inserting
their ovipositors into crevices in the bark. An examination of such
places, however, failed to show that any eggs had been deposited. The
writer has taken this insect in shagbark hickory, osage orange, and honey
locust. Attempts were made to rear it in elm, but the females would not
oviposit in it, even tho no other material was supplied.

In brief, then, the hickory borer has been reported as breeding in the
following kinds of wood: shagbark hickory, black walnut, pecan, butternut,
mulberry, honey locust, osage orange, bitternut hickory, elm, wild
grape, and hackberry.

ECONOMIC IMPORTANCE

The hickory borer is capable of causing considerable damage. While
it is never present in very great numbers, and does not attack healthy
trees, its work on recently killed, standing trees and on felled timber, as
well as on unseasoned products from which the bark has not been removed,
makes it of considerable economic importance on account of the value
of the wood destroyed.

It attacks recently killed, standing timber and felled material from
which the bark has not been removed. However, the writer has never
found it boring in such material after it had been dead for more than one
year, altho repeated attempts were made to have females oviposit in
such material. It enters not only the trunks of recently dead and felled
material but also the branches, even those as small as one and one-half
inches in diameter. In fact, it seems to prefer the smaller trees and
branches to the large trunks, for the galleries are always more numerous
in such material. In all cases the most severe injury is due to the pupal
cells which penetrate the larger trunks and branches to a depth of from
one to two inches, while in smaller branches they extend to the heart-
wood. •

Hickory is particularly injured by this borer, for the sapwood, which
until recently has been considered the only part of the tree suitable for
the manufacture of vehicles, agricultural implements, and various other
commodities, is so riddled by the larval galleries that the manufacturer's
profit is considerably reduced, resulting in a higher price for the finished
product. Years ago, when the hop industry consumed large numbers of
hickory poles, the wood for this purpose was often seriously damaged,
especially if it was left standing in the woods during the danger season.

THE PAINTED HICKORY BORER

185

BULLETIN 407

PLATE II

THE PAINTED HICKORY BORER

6, Larva (enlarged). 7, Holes to exterior made by larva before pupation. 8, Pupa
(natural size). 9, Pupae (enlarged). 10, Pupa in pupal cell. u, Pupal cells showing
plugs of splinters and sawdust.

1 86 BULLETIN 407

But now that elm has supplanted hickory for this purpose, the economic
loss has been considerably reduced. Material used in the manufacture
of porch furniture is frequently so injured by these larvae that it is very
easily broken.

The damage inflicted by the. hickory borer on black walnut, a wood
used to a considerable extent at the present time in the manufacture of
furniture, and especially, of gun-stocks, is bound to result in an increase
in the cost of this material.

The effect of the insect's work on osage orange is almost as injurious as
on hickory. This wood possesses much natural durability, and posts
and poles made of it remain sound for many years, even under conditions
favorable to decay. This natural durability is greatly decreased by the
hickory borer's work. Fence posts of this wood, lying in piles, are often
actually so honeycombed that they are fit only for firewood.

Invariably, severe damage results when the cut material is left in the
woods during the danger period. Not only do the burrows in the wood
decrease its value, but they also afford a means of access to fungus
diseases and moisture, which soon render the wood useless except for
firewood, and its value for even this purpose is much reduced.

DESCRIPTIONS

The adult
Gahan (1908 a) described the adult insect as follows:

Dark brown above, marked with pale yellow or yellowish- white pubescent bands —
two on the head, four on the pro thorax, and seven on the elytra; the first thoracic band
placed at the anterior border and very narrow; the first and second elytral bands almost
directly transverse, the third W-shaped, the fourth and fifth angulated and interrupted,
the sixth consisting of a rather strongly arcuated band on each elytron, the seventh
forming a border to the apex; body beneath banded with yellowish pubescence; meta-
thoracic episterna marked each with two yellow spots, the interval between which is
of a dark brown colour and as wide as or wider than either of the spots; legs reddish;
antennae dark brown, sometimes more or less reddish in parts.

d* Antennae rather thick, extending past the apex of the elytra; third to sixth joints
subdentate posteriorly at the apex; pronotum with a small punctate area on each side
rather close to the anterior border; sides of pro thorax very finely and densely punctate.

9 Antennae extending to the middle of the elytra, not quite so thick as in the male.
Length 10-22, breadth 3-7 mm.

The females are easily differentiated from the males by the shorter legs
and antennae, and more robust, compactly built body.

Gahan recorded two varieties of this species: one from Texas, with
pubescent bands an ashy white instead of yellow, and with the pronotum
of the male marked anteriorly with a small punctate area on each side
of the mid-line in addition to the lateral punctate area; and the other
from Villa Lerdo, Durango, Mexico, with the pubescent bands ashy

THE PAINTED HICKORY BORER 187

white or pale yellow, and the pronotum of the male marked anteriorly
with a median sub-rotundate area, which is joined, by means of a trans-
verse punctate band, with each of the punctate areas.

The egg

The egg is elongate-oval in shape, milky white in color, and measures
2 millimeters in length by .85 millimeters at its broadest point. The
chorion is very delicate and smooth, lacking ridges or other characteristic
markings. There is a considerable difference in the curvatures of the
dorsal and ventral surfaces, the ventral surface being noticeably more
convex. However, the shape may vary considerably, since the chorion
permits them to be easily modified to fit the crevices in which they are
deposited.

The larva

Packard (r88i) described the larva as follows:

Body thick; mouth-parts black; head reddish behind the antennae. Prothoracic
segment (first behind the head) large and broad, being one-half as long as broad; flat
and broad above, the upper surface being lower than that of the succeeding segment;
the anterior edge thickened, being slightly corneous; a mesial deeply impressed line,
especially on the hinder two-thirds, where it becomes a broad, deep angular furrow,
dividing the tergum into two quadrant-shaped halves; the outer edge of the segment
rises above the flattened tergal portion, which is sparsely covered with hairs; the latter
thicker along the sides of the body. The body contracts in width behind the 4th
abdominal segment; the upper side of each of the first six abdominal segments (corre-
sponding to those segments in the beetle) is raised into blister-like swellings, especially
on the 5th and 6th segments, which are much narrower than the four preceding segments.
These dorsal swellings are smooth and free from fine hairs. Abdominal segments 7-9
convex above, not swollen, and the abdomen is narrowest between the 5th and 6th
segments. A pair of large spiracles on the mesothoracic segment, and a pair on each
of the first eight abdominal segments.

Antennae 3- jointed; the two basal joints being of the same length; the basal one
being one- third stouter than the 2nd; the third joint filiform, and one-half as long as
the 2nd joint, and ending in two or three hairs. The thin membranous labrum is
divided into two parts, the basal solid, the terminal portion forming a moveable flap,
overlapping and reaching nearly to the end of the mandibles when closed; the basal
portion is shorter than broad, being broadly trapezoidal and smooth; the outer division
is broader than long, the edges being rounded, so that it is almost broadly ovate (trans-
versely) and smooth, covered with long hairs. It is pale membranous, of a testaceous
hue. Mandibles black, very thick and stout, with obtuse, rounded edges; they are
almost as long as the base is broad. Maxillae membranous, flattened; maxillary palpi
2-jointed. Labium membranous, with a transverse chitinous band near the insertion
of the 2-jointed palpi; both joints short; second one-half as thick as the first; edge hairy,
the hairs reaching to the ends of the palpi. Length of body 0.50 inch; breadth of
prothoracic segment, 4.2 mm.; breadth of head, 3.2 mm.

There is considerable variation in the length of the full-grown larva,
the range being from J to if inches. The larva also shortens con-
siderably just before changing to the pupa. In all the larvae examined
by the writer, the maxillary palpi were three- join ted, instead of two-
jointed as described by Packard (1881).

i88 BULLETIN 407

The pupa

The pupa is a typical, naked, coleopterous, Cerambycid pupa, brownish
in color, and measuring from 9 to 22 millimeters in length. The femur
and tibia of each leg are folded against each other, and lie transversely
on the ventral surface of the body with the tarsi projecting backward,
along the center. The wing pads are folded backward, passing under the
prothoracic and mesothoracic legs, with the tips covering the inner parts
of the mesothoracic legs on the ventral side. The antennae project for-
ward dorsad to the first two pairs of legs and then curl inward and are
held close against the body. There are numerous pointed granulations
on the dorsal side of the prothorax, and similar sharp spines on the
abdominal segments. On the penultimate segment, these spines are
somewhat larger, and are recurved anteriorly at the tip. There are seven
in a row near the posterior margin, two others more anteriorly situated,
and three more in front of these. The last segment has four similar
projections in a row.

LIFE HISTORY AND HABITS

The adult
Appearance of the adults

The cold and rainy spring of 1917 without doubt considerably retarded
the emergence of the adults, for the material kept in the insectary and the
records of the insects emerging from hickory used for firewood during the
winter show that warmth has a considerable influence on the time of
emergence. Observations were made on material kept at the insectary,
where normal conditions were simulated as much as possible, and on a
pile of hickory trunks and branches in the field which had been infested
the preceding spring. The first adults emerged June 6, in the insectary,
when three males were observed. The writer's records show that at
State College, Pennsylvania, in 1915, the first adults were observed on
May 12. As the weather became warmer, the number emerging gradually
increased until June 27. From then until July 5, the number of adults
appearing gradually decreased, no more being found after that date. In
the field, the males were the first to appear, several days elapsing between
their first aooearance and that of the females.

General activities

The hickory borer is a sun-loving insect. On bright, sunny days during
May and June they may be found on the trunks and branches of recently
killed trees or on felled timber, running rapidly back and forth. Rarely
are they found on such material when it is well shaded. When disturbed,
they either drop suddenly to the ground and hide among the herbage,

THE PAINTED HICKORY BORER 189

or else fly swiftly away, for their powers of flight are well developed. On
bright, warm days they work from about ten o'clock in the morning until
about half past five in the afternoon, with a maximum activity between
three and four o'clock. At night, and during cool, cloudy, or rainy days,
they hide under flakes of the bark, and are quite inactive. At such times
the pieces of bark covering them can be removed without causing the
beetles either to take flight or to drop to the ground, tho they sometimes
run about slowly until they again find shelter under some other piece
of bark.

During the period of activity the males are always more numerous
than the females. The males also appear earlier in the day than the females
and are much more active than the females, coursing up and down the
trunks and branches, chasing away ants or other insects which may alight,
and frequently fighting with each other. The females, on the other hand,
move about rather slowly, possibly due to their shorter legs and greater
bulk. At intervals they stop to examine places on the bark or to try
crevices with their ovipositors. Both sexes, when handled, make the
characteristic squeaking sound so common to beetles of this family.

Feeding habits

Like many of the Cerambycids, the adults of this species are pollen
feeders. For some time the writer was puzzled as to what the food plants
of the beetles could be, for while males and females, confined in cages,
copulated freely, yet oviposition did not occur, and both sexes died in
three or four days when no food was supplied. Finally, the alimentary
canal of a female, taken while ovipositing in the field, was dissected and
was found full of pollen. An examination of all the flowers in bloom at
that time was then made, with the result that the beetles were found
actively feeding on the pollen of the flowers of hawthorn (Crataegus sp.).
From then on, no difficulty was experienced in breeding the insects in
captivity. With a supply of these blossoms in the cages, copulation took
place and eggs were deposited as usual. This would seem to indicate
that food is necessary before oviposition takes place. This also agrees with
the observations of Garman (1916) on Cyllene robiniae, a closely related
species. In feeding on the pollen, these beetles clamber over the flowers,
and at a distance are often mistaken for wasps which are also common
on flowers at that time, and which they much resemble in coloration.
After feeding, they usually clean off their antennae and tarsi by drawing
them thru the mouth parts. While Crataegus was the only flower on
which the writer found the beetles feeding, yet Mr. V..R. Haber, of Ithaca,
New York, mentioned having found them feeding on the pollen of alder
catkins in Ohio.

i QO BULLETIN 407

Mating habits

Mating occurs shortly after emergence, as adults which emerged June
6 were found copulating the following day. The males are very aggressive
in seeking the females, often fighting vigorously among themselves for
possession of the female, biting each other about the head and appendages.
On June 13, a small-sized male attempting to copulate with a female,
and being roughly pushed aside by a larger male, seized his larger opponent
by the hind leg and held on for several seconds until vicious bites on the
head and legs caused the smaller male to loosen his hold. In mating,
the front legs of the male are clasped around the female's abdomen, the
middle and hind legs trailing on the ground. In this position, the female
drags him about with her over the trunks and branches, stopping only
during the act of copulation. Copulation occurs frequently and lasts
but a few seconds. On June 13, mating occurred eight times in fifteen
minutes. When copulation is completed, the penis and ovipositor pull
away with considerable difficulty, both being considerably extended from
the abdomen in the process. Several males may mate with a single
female.

Opposition

Oviposition takes place very soon after copulation, as females which
were observed mating on June 13 deposited eggs the following day. The
eggs are always placed in crevices or under scales of the bark. Smooth
branches are rejected unless there are rough spots or crevices present
where eggs may be laid. To facilitate observations, the writer obtained
branches with very smooth bark, and cut slits in them at various intervals
which provided favorable places for the deposition of eggs. These were
placed in cages which each contained a single fertilized female, and obser-
vations were made. Before ovipositing, the female walked around,
exploring the slits with antennae and ovipositor. Having found an
apparently suitable place, the female inserted her ovipositor into it, moving
it about in a manner suggesting the movements of a house-fly's mouth
parts in exploring a bit of food. If the crevice was unsuitable for depositing
an egg, the ovipositor was withdrawn and another crevice explored. This
process was continued until a suitable place was found and then the eggs
were deposited.

The number of eggs laid in a single crevice varies. In the insectary,
where artificial scars were made in very smooth bark, the number of
eggs placed in a single crevice varied from one to fourteen. Under normal
conditions, in the field, the greatest number of eggs found in a single
crevice was five.

THE PAINTED HICKORY BORER 191

Observations were made at the insectary on the total number of eggs
laid by a single female. Of the twelve specimens observed, the maximum
number of eggs deposited by a single female was fifty-six, the minimum
being thirty-eight. While large cages were used and conditions as natural
as possible were provided, yet dissection of the abdomens of these females
at death showed that not all the eggs had been deposited, quite a few
remaining in the ovaries in different stages of development. Whether
the entire number of eggs is deposited under natural conditions in the
field was not determined. Most of the eggs are deposited on the first
day. For example, the female which deposited a total of fifty-six eggs
laid forty-six the first day, eight the next, and two on the third day.

After copulation, the males become inactive and die in about a week's
time. The females, as a rule, die within three to five days after oviposition.

The egg
Time of hatching

The eggs hatch in from six to ten days. Observations were made on
a batch of one hundred and fourteen eggs deposited by three females on
separate branches, on June 19. These were collected and placed where
conditions would be as natural as possible. The results of the experiment
appear in table 2.

TABLE 2. THE TIME OF HATCHING OF 114 EGGS, DEPOSITED ON JUNE 19

Date of hatching

Number
of eggs
hatched

Tune 2S

18

J 26..::: .. • ::

24.

27

•i-i

28

oo

•7Q

2O

From the above data it will be seen that the first eggs hatched six days
after they were laid, while the last to hatch required ten days. The
majority of them, however, hatched eight days after they were laid.
Observations made on a batch of eggs kept in the laboratory, where con-
ditions were more dry and warm, showed that the eggs hatch in from
five to seven days.

A few days before hatching, the fully formed larvae may be seen thru
the chorion, actively twisting about. Just prior to hatching, the egg
enlarges slightly at the anterior end, due to the size of the larval prothorax.
The chorion is undoubtedly broken by the muscular contractions of the

192 BULLETIN 407

larva, for suddenly it bursts at the anterior end, and as the larval head
is pushed thru, the opening is gradually enlarged until a slit extends more
then one-half the length of the egg. No egg burster of any kind was
found on the larva.

The larva

Activities of the newly hatched larva

The newly hatched larva has the appearance of most Cerambycid
larvae. It measures 1.8 millimeter in length, while the prothorax, which
is the broadest part of the larva, measures .9 millimeters in width. The
mouth parts are black, while the head, behind the mouth parts, is brownish
in color. The different segments are sparsely covered with delicate,
long hair.

Immediately after hatching, the tiny larvae migrate in under the scales
of the bark, where the eggs have been deposited, as far as the size of the
prothorax will allow, and then, using the scale of bark as a brace, begin
to burrow in towards the wood. It takes about twelve hours to burrow
deep enough so that they are completely concealed and during this time
fine, powdery sawdust is pushed out behind them. Under natural
conditions, where the eggs are laid in clusters of only four or five, there
is but little mortality among the young larvae, each individual being
able to establish a burrow by itself. But when a large number of eggs are
laid in a single crevice, as was done by the specimens kept at the insectary,
there is not room enough for all of them to get started. Accordingly,
two or more penetrating into a single burrow, one is punctured by the
mouth-parts of the other and is killed. Out of fifty-two larvae hatching
in a single piece of hickory, only twelve were able to survive and establish
burrows. Moreover, when the bark is smooth and no crevices or scales
are present to serve as braces for them in beginning their burrows, they
are unable to penetrate the bark and soon die. In like manner, the larvae
are unable to burrow into material from which the bark has been
removed.

Larval burrows

Having penetrated the bark, the young larvae drive their long, linear
burrows with the grain of the wood, grooving the inner bark as well as
the sapwood. Each burrow usually begins as a small gallery -^ inch
wide and of about the same depth. Gradually increasing in width and
depth, it may extend for a distance of from 8 to 12 inches or even more,
after which it usually bends on itself to form a long, narrow, irregular U,
whose arms may be either in contact with each other or slightly separated ;
on the other hand, the gallery may consist of several loops which are either

THE PAINTED HICKORY BORER 193

distinct or else cross each other in places. Prom beginning to end, however,
all the galleries increase in width and depth, until, just before the pupal
cell is formed, they measure from J to J inch wide and J inch in depth.
In small branches, in which a large number of larvae are at work, the
sapwood may be entirely honeycombed to a depth of an inch or more,
the different burrows being indistinguishable from each other. The
nature of the wood seems also to have a decided influence on their shape,
those in wood free from knots being more or less regular and linear, while
those in knotty material may be very irregular. On felled material, the
galleries may extend in either direction with the grain of the wood, but
in standing timber the majority of them first extend downward.

The burrows are firmly packed with the coarse, sawdust-like borings
chewed off by the larvae, whose gnawings can be heard for a considerable
distance and resemble the noise made by the so-called " sawyers " in pine.
The writer was unable to determine whether all of the gnawed material
passed thru the body of the larvae. That dissections showed the alimen-
tary canal full of sawdust-like particles, and that the material filling the
burrows was so tightly packed and stuck together that it took considerable
work to dislodge it, are facts tending to confirm this view. Furthermore,
it is not easy to see how the larvae could store the loose sawdust so
compactly, were it not previously moistened by passage thru the alimen-
tary canal. Here and there along the course of the burrows, air holes
are eaten thru the bark to the exterior, the number of these for a single
burrow varying from one to four.

The pupal cell

The larva becomes full grown in from ten to twelve weeks. At the
end of that time it gnaws a large, oval-shaped hole thru the bark to the
exterior, and begins the construction of the pupal cell. This extends
into the solid wood to a depth of from i to 2\ inches, and then parallel
with the grain of the wood for about 2 inches. In making the pupal
cell, the larva apparently does not eat the sawdust, for as soon as an
appreciable amount is chewed off, the larva, using its head as a scraper,
pushes it to the exterior thru the hole previously made in the bark. After
pushing out one load, the larva backs down into the burrow again and soon
reappears with another supply. This process is continued until the entire
pupal cell is completed. Evidence of the borer's attack on timber is
therefore seen at this time in the large, oval holes in the bark and in
the great amount of sawdust pushed out, which accumulates in small
heaps beneath infested material. The making of large holes thru the bark
before pupating seems rather unusual, as many of the Cerambycid larvae

ip4 BULLETIN 407

do not make them. These were evidently the holes to which Hopkins
(1897) referred when he stated that the numerous holes in the bark indi-
cated that the adults had emerged, altho nothing but fully matured
pupae were found in the wood.

When the pupal cell is hollowed out, the larva, by means of its mandibles,
splits off some splinters very much resembling short pieces of excelsior,
\ to | inch long, from the sides of the burrow. With these it closes up
the pupal cell a short distance below the point where it begins to run
parallel to the grain of the wood. After these splinters have been packed
firmly in place, so as to form a compact plug, they are reinforced on the
inside with a plug of sawdust.

In fallen timber, the pupal cells may extend in either direction, but
in standing timber all those examined by the writer extended downward.

The prepupal stage

When the pupal cell is completed, the larva ceases activity and passes
into the prepupal stage, during which it is found at the further end of the
cell, always facing the plug of splinters and sawdust. However, if the
plug is damaged or destroyed, more sawdust will be gnawed to repair
the damage. During this stage both the length and bulk of the larva
slightly decreases. The duration of the prepupal stage varies with the
temperature and humidity. In the laboratory, where it was warm and
dry, this period lasted, on an average of fifteen days, while in the insectary,
under normal conditions, it was much longer, lasting from twenty-three
to sixty-three days. During the latter part of the prepupal stage, the
color of the larva varies from a yellowish white to a brownish color, and
the antennae and various other appendages become faintly discernible
thru the transparent cuticula.

The pupa
Pupation

In the laboratory, pupation began as early as August 23, while in the
field the first pupae were found on September n. Pupation continues
from this time until November 12, by which time all the larvae have
changed to pupae, the majority of them transforming during the month
of October. The last larval skin may remain attached to the end of the
pupa, or else lie free in the pupal cell.

Activities of the pupa

Thruout this stage the pupa remains at the further end of the pupal
cell, facing the plug of splinters and sawdust. It remains quiescent
unless disturbed, at which time the abdomen is moved back and forth.

THE PAINTED HICKORY BORER 195

If placed at the plugged end of the cell, it quickly moves itself back to
the opposite end by means of the spines on the thorax and abdomen.

Transformation to adults

In the laboratory the pupa began to show the characteristic markings
of the adult by the middle of October. However, they did not transform
to adults until the following February. In the field, the winter is passed
in the pupal stage. All of the pupae under observation at the insectary
were killed by the extremely cold weather during the winter of 1917,
so that records on the transformation to the adult stage were not obtained.
Likewise it was found that pupae in infested material in the field had
also been killed by the unusually cold winter of that year. This agrees
with the observations of Craighead (1918) and other workers on forest
insects, who also reported a great mortality among wood-boring insects,
due to the severe cold.

However, pupae obtained from infested material, late in the winter
of 1919, transformed to adults during that spring and emerged. The exact
time of transformation from pupae to adults was not determined.

SEASONAL HISTORY

The adult insects emerge during May and June, and sometimes, during
the first few days of July, depending on the season. They feed on the
pollen of Crataegus and probably on that of other flowers. The elongate-
oval, whitish eggs are deposited in crevices or under flakes of the bark.
These hatch in from six to ten days, and the young larvae, burrowing
thru the bark, then construct an irregular, U-shaped burrow, which grooves
both the bark and the solid wood, and is tightly packed with the sawdust-
like borings. The larvae become mature in from ten to twelve weeks,
at the end of which time they construct the pupal cell, first opening a
hole to the exterior thru which the borings are to be expelled. When
the pupal cell is hollowed out, the larva retreats into it, plugs up the
entrance with splinters and sawdust, and enters into a prepupal stage
lasting from twenty-five to sixty-three days. The transformation to
pupae begins about the middle of September, and by November 12, all
are in the pupal stage. In Ithaca, the winter is therefore spent as pupae.
The transformation to the adult beetles takes place in the spring.

NATURAL ENEMIES

The hickory borer has very few natural enemies. Riley (1874)
mentioned finding the larva of an Elaterid (Hemirhipus fascicularis
Fabricius) following the burrows of the hickory borer in hickory and

196 BULLETIN 407

pecan trees probably with the purpose of feeding on the larva of the borer.
Packard (1881) figured a Nitidulid larva which he stated probably preyed
on this and other hickory insects. Chittenden (1893) reared Bracon
erythrogaster Brulle and Doryctes radiatus Cr. from hickory wood infested
almost exclusively with the hickory borer. The writer has found Bracon
erythrogaster Brulle in considerable numbers ovipositing in the galleries
of this insect. Infested material has also yielded quite a few adults of
B. erythrogaster. This parasite seems to be quite effective in checking
the borer, as a large number of specimens split out of wood had been
killed by these parasites. The borers are killed during the larval stage,
and when parasitized, turn blackish in color and are much wrinkled
and flattened, due to the fact that their contents have been eaten out
by the parasite.

Very often that part of the pupal cell outside 'of the plug of splinters
and sawdust, is utilized by a small Eumenid wasp which there constructs
its nest of clay and rears its brood. These clay nests block the entrance
effectively and doubtless prevent parasites from gaining access to the
pupal cells during this vulnerable stage of the insect's life.

RECOMMENDATIONS FOR CONTROL AND THE PREVENTION OF INJURY

Since the hickory borer attacks only recently dead and felled material,
all attempts should be centered on methods of preventing injury as
much as possible. As previously stated, this insect is a sun-loving species,
rarely being found in well shaded places. To test this out, the writer
piled branches of hickory in the woods where there was abundant shade,
and in relatively close proximity to infested material from which the
adults were emerging. None of this material was attacked by the hickory
borer, and were it not for the fact that other species caused much damage
to the wood in such a location, the piling of hickory in shady places would
be an effective means of preventing injury.

Standing dead timber should be felled as soon as possible, preferably
within a few weeks or months after it has died. Such timber should be
utilized at once, or, if this is not possible, the logs should be barked or
else . put in water. Where either of these methods are employed injury
will be prevented, for the larvae cannot breed in the barked logs, and
water will kill whatever stages of the insect may be present, or will
equally well prevent oviposition.

Unbarked logs, posts, poles, and such material should never be left
lying in the woods during the danger season, which extends from May i
to August i . They should either be utilized at once, or barked, or placed
in water.

THE PAINTED HICKORY BORER 197

Infested material which cannot be used by the manufacturers should
be utilized as soon as possible for firewood, preferably before the insects
have had time to emerge. In this way large numbers of them may be
destroyed.

All material which cannot be used, such as slashings, should be burned
as soon as possible, for it is in such places that the insect finds excellent
breeding places. When such useless mater al has been left in the woods
until it has become infested, it should be burned before the beetles have
had an opportunity to emerge the following spring.

The destruction of Crataegus in the vicinity of large tracts of hickory
would also be beneficial, in that it would destroy the food supply of the
adult beetles at the time of emergence. The elimination of such trees
would be beneficial in any case, as they are the native hosts of many
of our most injurious insect pests.

198 BULLETIN 407

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