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U. S. DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 69,
L. ©. HOWARD, Entomologist and Chief of Bureau.

THE CHINCH BUG.

BY

F. M. WEBSTER,

In Charge of Cereal and Forage-Plant Insect Investigations.

IssuED JUNE 21, 1907.

WASHINGTON:
GOVERNMENT PRINTING OFFICE.
19.07.

BUREAU OF ENTOMOLOGY.

L. O. HowArp, Hntomologist and Chief of Bureau.
GC. L. Mariatrr, Hntomologist and Acting Chief in absence of Chief.
R. 8S. Crirron, Chief Clerk.

¥. H. Currrennen, in charge of breeding experiments.

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

W. D. Hunver, in charge of cotton boll weevil investigations.

A. L. QUAINTANCE, in charge of deciduous-fruit insect investigations.

EE. F. PuHinures, in charge of apiculture.

D. M. Roaers, in charge of gipsy and brown-tail moth work.
A. W. Morrity, engaged in white fly investigations.

E.S8. G. Trrus, in charge of gipsy moth laboratory.

C. J. GILLISs, engaged in silk investigations.

R. P. Currin, assistant in charge of editorial work.

MABEL Cotcorp, librarian.

CEREAL AND FORAGE-PLANT INSECT INVESTIGATIONS.

i. M. WEBSTER, in charge.
G. I. Reeves, W. J. Puitiies, C. N. AINSLIE, special field agents.

to, DE PAR MENT - OF AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 69.

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

fee CINCH BUG.

BY

F. M. WEBSTER,

In Charge of Cereal and Forage-Plant Insect Investigations.

IssuED JUNE 21, 1907.

WASHINGTON:
GOVERNMENT PRINTING OFFICE,

Pe OIOET

LETTER OF TRANSMITTAL.

U. S. DEPARTMENT OF AGRICULTURE,
Bureau or Entomoroey,
Washington, D. C., March 5, 1907.

Str: [have the honor to transmit for publication the accompanying
manuscript entitled ‘“ The Chinch Bug,” by F. M. Webster, in charge
of the cereal and forage-plant insect investigations of this Bureau.
This is a thorough revision by Mr. Webster of his earlier account of
this destructive pest published in 1898 as Bulletin No. 15, new series,
of this office, and includes additional data based on observations made
during the past eight or nine years. I recommend that it be pub-
lished as Bulletin No. 69 of the Bureau of Entomology.

Respectfully, L226.
. O. Howarp,

FE, .tomologist and Chief of Bureau.
Hon. James WILson,
Secretary of Agriculture.

CONTENTS:

SUES OO ETON. Ss i ea mE a ee ee
ETE ts Tween ae wed eee RC le eee ee AE ye ie ia
SUL SMMet ANCL cMIMM IMITATIONS =.= 2. 2 es ee ee ee
CORRS eG ae ee ee ne eee een Ree

EKgg period and number of eggs deposited by each female
Descriptions of the different stages of development_______-_________-______
Wevelopment- andchabits) of thes youngs 22-3 2e2 2 ee ee
INTMDeT AO .ceNnera Ons aanm Welly: a ese SS ee
Destructiveness largely due to gregarious habits________________________
EO CVC ame | LeUT I iS eee eee noe eee cr ne none eee Oe ee ee ee oe
ImMSecisnthaanresmistaken tor cChinch DuUgs= ss Se eee
WOSSESECHUISeC aD Yy-sChil Che DUS S422 ae Se ee i ee ee
SCRAPE | GUS GS a ra ee ee ee

Influence of precipitation on the chinch bug-

intivence of temperature on the chinch bug. = 2-3 = eee
ENED Us UMTS a STN TM TS se ee ee ae a ee Se St

PASI Cipla Ce eee ee Se ee bee ee

Fungous enemies of the chinch bug determined
Field and laboratory experiments in Indiana___________________
First field applications of fungous enemies of the chinch bug_____
he works Of Proressor Snow, ime Wwansas= =) = ee
Other insects attacked by Sporotrichum globuliferum__ ~~~ _—
First artificial cultivations of Sporotrichum globuliferum
ReEsmisormneldsapplicanions in) Ohio22==2 =.= = ees
Meteorological influences favoring development of fungous ene-
MNES TO fee we Hi Chey Ul eae eee eee Sot Pe Se eee
ABPActerialvenemy-of the <chinchy bugs= == = = = ee es ee
The practical utility of fungous and bacterial enemies in fighting the
Ce TNL CG aL een ee ene ae ea Boe, Sa

OE VEN") GY ORS AWE ICES CO GC FDC |e er CLD, SAE SS ey aR OE Se eye ee

Other bird enemies of the chinch bug

LEVON ERO sg ae ety ee eee ee ee ene Soe ee ose

Invertebrate enemies of the-chinch bugs2.2=—. = 22 se
Remedialeandspreventive: Measures. -= 522 22

Destruction of chinch bugs while in hibernation_________-____________

Sowing decoy plats of attractive grains or grasses in early spring____

Difficulty of reaching chinch bugs in meadows_____________________

Watchtfulness necessary during protracted periods of drought

Utility of kerosene in fighting chinch bugs__________________________

Utility of deeply plowed furrows supplemented by the use of kerosene

emulsion

2) |
pas

on
oo

Or Or
He He

Or Om Ow
Oo ©

6 CONTENTS.

Remedial and preventive measures—Continued.
The ridge and: coal=tar method---_+*-- Se eee
Other barrier method s__ <2 =~. 2 2 a ee eee
Necessity for preventing chinch bugs from becoming established in
fields of wheat and ‘crass. 2 =) * eee
Summary of remedial and preventive measures_______-_-__-_-___-__
Probable orizin and diffusion of therchincehy pis
Indications of a probable distant origin and later diffusion__________
Unique appearancerandseresariois pivot ee eee
Occurrence of the long and short winged forms and their distributicn_
Relation of the inland and seacoast short-winged forms _________-____
Probable courserorcdithw stoma see ae eee ee eee ee
Habits of the European species, Blissus doriw Ferr__________________
Previous ideas on the diffusion of the chinech bug___________________
Reasonsitor the present theorysot dif usion= 9
MV CY ie = Shee ce A pe I ges a Or ee a

Page.
66

68

69
72
72
74
74
ri)
78
78
$3
Si
88
91

LLLUSTERAT IONS.

Fig. 1.—Map of North America showing areas infested by the chineh bug_
P= lmmavirerstices On thes chimeht lesa ee
3.—Blissus leucopterus: adult of long-winged form____---~--------
4.—Blissus leucopterus: adults of short-winged form__-~~--~~---~-_-

5.—Corn plant infested with chinch bugs____---------------__------
NESTS TVG RUSE CULIL Sma eee ee ee
pI OS TIVO GLIUCH. Cy a ee ee
SE = COnIMeEland “PUllGAit a= ee ee

9.—Brachyrhynchus granulatus: larva, pupa, and adults__-------~__
10.—Map showing areas in the United States over which the chinch
bug occurs in greatest abundance and may at any time become

CL OGSTECTC liye ee en es Se es 2 ee eee ee
11.—Map showing distribution of the chinch bug in Ohio in 189625. ==
12.—Map showing distribution of the chinch bug in Ohio in Soi
13.—Map showing distribution of the chinch bug in Ohio in NSO sas
14.—Map showing distribution of the chinch bug in Ohio in 1895 and
amount of precipitation over the State during May, 1895__--__-
LED Sa SUQLO SLS =e ee ee ee

WA SSVI COUCWUD 2S
17.—Map showing probable course of diffusion of the chinch bug over

IN@aue) AW aie oe ee ee ee
1 = Plissiiseaoric. lmmeatine: Stacege-=. ==!) Sees ee

Oe iS SUSU ONG AC UGS ee ee

~]

vai bern Cr BUG.

Few insects, and certainly no other species of the natural order to
which this one belongs, have caused such enormous pecuniary losses as
has the chinch bug, Blissus leucopterus Say. No other insect native to
the Western Hemisphere has spread its devastating hordes over a
wider area of country with more fatal effects to the staple grains of
North America than has this one. But for the extreme susceptibility
of the very young to destruction by drenching rains and to the less
though not insignificant destructiveness during rainy seasons of the
parasitic fungus, Sporotrichum globuliferum Speg., on both the adults
and young, the practice of raising grain year after year on the same
areas, as followed in the United States, would become altogether un-
profitable. Some of this insect’s own habits, emphasizing as they do
the effects of meteorological conditions, are the most potent influences
that serve to hold it within bounds, by giving its tendency to exces-
sive increase a decidedly spasmodic character.

DISTRIBUTION.

The genus Blissus is widely distributed over the world, occurring in
South Africa, Abyssinia, Algeria, Sicilia, southern Europe, northward
at least to the sand dunes of central and northern Hungary, India,
Japan, southern Russia, and in the Western Hemisphere in Buenos
Aires, and from Panama and the Island of St. Vincent northward
to middle California on the Pacific coast and Cape Breton on the
Atlantic. When we come to understand that the Hemiptera of the
world are far from being well known, and the faunas of South
America and central Africa have as yet been hardly studied at all,
we may well presume that future studies of the hemipterous insects
of these countries may unite some of the different areas now known
to be inhabited by the several species of this genus.

At present in the Old World this genus may be said to occur in the
Ethiopian, Oriental, and Palearctic life regions; while in the New
World it ranges from the Neotropical region at Panama and St.
Vincent into the Nearctic over the borders of the Boreal subregion in ~
British America.

Our American species, Blissus leucopterus Say, the only one of the
genus at present known in the Western Hemisphere, has been recorded
from St. Vincent and Grenada, West Indies, by Uhler; Cuba, by Stal:

9

10 THE CHINCH BUG.

Volcan de Chiriqui, Bugaba, and San Feliz, Panama, by Champion;
San Geronimo, Paso Antonio, Panzos, Champerico, and Rio Naranjo,
Guatemala, by Champion; Lower Purissima, Lower California, by
Uhler; Alameda, Cal., by Koebele; and in the vicinity of San Fran-
cisco, Cal., by both Uhler and Koebele; Orizaba, Mexico, by H. H.
Smith; Tamaulipas, Mexico, by Uhler; Mesilla Park, N. Mex., by
Cockerell; Florida, by Schwarz and Dr. J. C. Neal; Sydney, Cape
Breton, by W. H. Harrington; Muskoka, Ontario, Canada, by E. P.
Van Duzee, and Winnipeg, Manitoba, where a single specimen was
collected by Dr. James Fletcher and given by him to Mr. Harring-
ton, to whom I am indebted for information regarding its occurrence.
Inland, in the United States, it may be said to be generally distributed
from Texas to Manitoba. It is also very probable that its occurrence
along the Pacific coast is much more extended than is at present
known, as it has not been searched for to any extent in that region.
(See map, fig. 1.)
HIBERNATION.

The chinch bug hibernates in the adult stage, and though there may
be occasional exceptions, especially in the South, it has yet to be
observed in very early spring in any other than the adult stage, at
least in any locality north of Mexico. The writer observed pup in
central Illinois apparently in hibernation in company with adults
on November 11, but there is no proof that these survived the sue-
ceeding winter. In Tensas Parish, La., adults were abroad in con-
siderable numbers during March, 1887, yet there was no indication
of any young having wintered over. The adults were pairing and
seemingly engaged in oviposition, precisely as is to be observed in
the Northern States during May and June. No young were observed,
as most certainly would have been the case had they occurred there,
for observations were made in fields of young corn, where, had the
young bugs been present even in very limited numbers, they would
certainly not have escaped the rigid searching under and about the
bases of the leaves of the young corn plants.

Doctor Howard ¢ quotes Prof. G. F. Atkinson, at that time of Chapel
Mill, N. C., as having observed half-grown chinch bugs on crab grass,
about the Ist of October. The same authority also quotes Doctor Riley
to the effect that many of the chinch bugs pair in the fall preparatory
to seeking winter quarters, and also cites the fact that Mr. James O.
Alwood observed them pairing in a field of uncut pearl millet, Octo-
ber 27, 1887, on the grounds of the Ohio Agricultural Experiment
Station, then at Columbus, Ohio. Dr. Cyrus Thomas,’ in speaking

#'The Chinch Bug, by L. O. Howard; Report of the Commissioner of Agricul-
ture for the year 1887, pp. 51-88.
» Bulletin No. 5, U. S. Entomological Commission, p. 13.

iy

HIBERNATION.

of the possibilities of an occasional third brood in southern Illinois

, states that there were some evidences of this, but not

and Kentucky

eas

Va

WS ;
S585

oo aK
%

VO
secre

erage
GPCI

ey

b Exes

(Author’s illustration.)

o
8.

160

oo vor Mnte

v7

OS

eovtr _or| wer

Roe

stribution of the chinch bu

orth America showing di

Map of N

Fig. 1.—

«

sufficient to justify him in asserting it as a fact or to satisfy him of

its correctness.

12 THE CHINCH BUG.

It therefore seems probable that no young are produced as a result
of the late pairing, at least until spring, and it has yet to be shown
that the late appearing larve do not mature before the hibernating
season sets in, or else die during the winter. When we come to con-
sider the extreme susceptibility of the newly hatched chinch bug to
wet weather, less perhaps in case of the short-winged form, it will be
apparent that as we approach the Tropics the wet and dry seasons
would tend to influence the breeding seasons, as those individuals that
hatched before the close of the rainy season would be, in a measure at
least, continually eliminated, while those that hatched so late as to be
caught in the commencement of the rainy season would also be to an
equal extent destroyed, and thus, by continually restricting the breed-
ing period to certain months, establish a fixed law that would be ad-
hered to even under the somewhat different conditions which occur
farther to the northward. Unfortunately the date or dates on which
the young were observed by Mr. Champion, on Volean de Chiriqui,
in Panama, are unknown to the writer, and it is impossible to say
whether or not they were found during or near the dry season.

In an article on the hibernation of the chinch bug, Mr. C. L. Mar-
latt * calls particular attention to the fact that in Kansas the chinch
bug in autumn seeks the dense stools of some of the wild grasses in
which to hibernate, and to such an extent did this occur that it was
suggested as probably the normal hibernating habit of the species.

Before entering into a discussion of this matter, it will be well to
present two communications received from the late Dr. J. C. Neal, at
that time of Stillwater, Okla. As Doctor Neal was located in a sec-
tion of the country where, in many cases, civilization had not in-
fluenced to such a marked degree the natural insect fauna, the author
applied to him to secure some exact information in regard to the
chinch bug under such conditions. The correspondence, however,
was terminated suddenly by Doctor Neal’s death. The two letters
here given are among the last he ever penned. They are of a some-
what general nature, and will be referred to later in this discussion.

OKLAHOMA AGRICULTURAL AND MECHANICAL COLLEGE,
Stillwater, Okla., October 31, 1895.

My Dear Sir: Yours of the 28th just received. Last year was the first wheat
year in most of the new additions to this Territory, and from all sections the
cry was for infection, as ‘the bugs are ruining us.” I received letters from
every county in the strip and in the western sections. The most damage was
done in the extreme southern range of counties, and near Okarche (see map,
fig. 10) the damage was excessive. I do not think there is a single acre in this
or Indian Territory that is not saturated, so to speak, with the chinch bug.
You may put this whole area down as within the infested boundary line.
My belief is that the increase of country roads, the decrease of March fires,
the shiftless habits of the vast majority of our farmers in allowing volunteer

4 Insect Life, Vol. VII, pp. 282-234, 1894.

HIBERNATION. 13

wheat and oats to grow and wheat lands to remain fallow, and the planting
of new and better grass crops than the tough blue-stem, are direct causes of
what I believe a decided increase of this insect in Oklahoma during the last
five years. It would be amusing, if it were not so pathetic, to read the many
letters I get, something in this wise: ‘I planted wheat on sod land; the chinch
bugs destroyed it so badly that in February I plowed it up and sowed oats;
this, too, went the same way; I then planted corn, and when it was a foot high
the little bugs came by the millions and destroyed that; I then planted the land
to Kafir corn, and that will be ruined if you can not help me.” What could I
do for such a man? Had the bugs laid out a programme for their daily sus-
tenance, no better commissary-general could have been obtained for them than
he was, and I had to write him that his plan was the worst one possible for
him, and the best for the bugs, and that the only suggestion I could make,
from the bugs’ standpoint and for their benefit, would be to plant wheat again
so that they could have something for the coming winter’s food. In his case
it was a series of fatal mistakes from ignorance of the habits of the bugs.

Another thing which I believe adds materially to the increase of these pests
is the complete destruction of the prairie chickens, the decimation of par-
tridges, and the thinning out of all kinds of smaller birds, such as the cow black-
birds, bank sparrows, martins, larks, and other prairie birds. This section
is full of reckless boys and men who kill everything that flies, good, bad, and
indifferent, “for fun.”

Some years ago I went out on the Cherokee Strip, miles away from human
habitation, and saw some of the small birds—larks and killdees—busily picking
in the young grass, in early spring, and upon examination found these places
swarming with chinch bugs sucking the juices of the blue-stem grass.

“Almost any time in the winter when the weather is warm one can find chinch
bugs, and I have witnessed two “ flights”’ of these insects and determined them.
I should be glad to answer any more specific questions at any time.

With regards, I remain,
J. C. NEAL.

The second letter is a short note in reply to the author’s question
regarding the grasses fed upon by the chinch bug, their hibernating
habits, and developments:

STILLWATER, OKLA., November 20, 1895.

DEAR PROFESSOR WEBSTER: In reply to your postal, I would say that I do not
know, but will at once make observations and report at my earliest chance.

My belief is that the bugs attack all the grass family except the Cenchrus,
and that only is exempt on account of its bitter taste, which effectually shields
it from insects, as far as I have seen, both in this section and in Florida.

I will take the matter in hand at as early a date as possible and write you
progress and results.

Very respectfully, J. C. NEAL.

It is reasonable to infer from these letters that the chinch bug win-
tered over about the stools of grass, and that the birds were observed
to attack them there in early spring, as the statement is made that
later, when the young corn was a foot high, the little bugs came by
the million. This condition of affairs may be considered in connection
with the statements of Dr. Asa Fitch,* regarding his cbservations

a Second Report on Noxious, Beneficial, and Other Insects of New York, p. 283.

14 THE: CHINCH BUG.

in Illinois in the autumn of 1854, when in passing over the northern
part of the State he found the ground in some places, in the midst
of extensive prairies, covered and swarming with chinch bugs,
reminding him, as he says, “ of the appearance presented on parting
the hair on a calf that has been poorly wintered, where the skin is
found literally alive with vermin.” Further along in his report
(p. 290) he states that “so late as the forepart of October I met
several of these insects in the pupa state, and some of these I do not
doubt would pass the winter in that state, and therefore would not
deposit their eggs until the following spring.” That he did not find
these pupe in New York is shown by his statement on page 287 of the
same report, to the effect that he had “ met with but three specimens
in New York, occurring on willows in the spring of 1847 and May
12, 1851.” As shown farther on in this bulletin, there is no proof
that these pup did not develop to adults before winter, or die before
spring, and the conditions indicated would almost presuppose that
hibernation would take place on the prairies where the insects were
observed by Doctors Fitch and Neal. From personal recollection
the writer knows that the section of Illinois to which Doctor Fitch
refers was, at the time mentioned, but thinly populated, and there
were still very extensive tracts of the original prairie grasses miles
distant from woodlands.

In an interesting note by Mr. E. A. Schwarz“ on the hibernation of
the chinch bug, given in discussing Mr. Marlatt’s paper, previously
mentioned, attention is called to the fact that the hibernation of the
chinch bug had been observed by him, in its maritime home, in the
vicinity of Fortress Monroe, Va., which locality he had been in the
habit of visiting for a number of years, during the first warm days
of spring. The maritime flora and fauna are here late to awake, and
most insects peculiar to the seacoast can still be found in their winter
quarters by the end of April. By pulling up any good-sized stool of
grass and beating it out on the smooth surface of the sand or over a
cloth a multitude of various insects are sure to be found, and among
them always plenty of chinch bugs. These stools of grass not only
serve as winter quarters, but in summer the chinch bugs crawl into
them during the daytime to protect themselves from the fierce rays of
the sun.

In the timothy meadows of northeastern Ohio the writer has wit-
nessed cases where the chinch bugs had commenced their operations
along one side, worked part way across the field, killing the timothy
as they advanced, and continued their depredations the following
year precisely where they suspended work the autumn before, the
long-winged individuals only migrating in the intervening time.

— +

@Insect Life, Vol. VII, pp. 420-422, 1895.

- HIBERNATION. 15

In southwestern Maine, where this short-winged form has _ oe-
curred in more or less destructive numbers for upward of forty years,
and where it affects timothy in the same manner as in Ohio, both long
and short winged individuals, the latter in the majority, hibernate
under dead leaves, brush heaps, and similar débris in and about the
fields where they have ravaged the timothy. They do not appear to
select only the drier portions of such fields, but are found also liter-
ally swarming about the clumps of rushes (Juncus) that grow in the
low spots. Some of these low places become submerged in winter by
rains and melting snows, and the hibernating bugs are washed out
and killed.« Possibly others not observed might have remained
among the living timothy, as it is further stated that many hibernat-
ing individuals were to be found among the leaves of clover border-
ing on spots of timothy that had been killed out by them during the
preceding summer.

That the short-winged or maritime form must hibernate in or in
very close proximity to the field it infests goes without saying, and
it would appear that but for the cultivation of timothy it would
have become diffused inland from the coast less rapidly, if at all.
It is doubtful if this inland diffusion began until the country became
settled by the white man and timothy began to be grown by him as
a forage crop—a situation that would be coexistent with a dimuni-
tion in the number and extent of prairie and forest fires.

West of the Allegheny Mountains we encounter this short-winged
maritime form only in western Pennsylvania, northern Ohio, south-
ern Michigan, extreme northern Indiana, and equally extreme north-
ern Illinois. The writer once found a single short-winged individual
in southern Ohio, and a single individual that may or may not belong
to this species has been recorded from New Mexico by Prof. T. D. A.
Cockerell.

Except as indicated in the preceding paragraph, over this whole
country the long-winged form is the only one known, and its habits
are almost as unlike those of the maritime form as they would be
were the latter a different insect. Timothy culture has never ex-
tended to the Gulf coast, and the extensive growing of the crop over
this whole western country is of recent date, coexistent with the
advent of the white man. Here, therefore, timothy is not attacked
by chinch bugs.

The inland or long-winged form inhabits largely a prairie country,
and it would appear that, as these prairies were annually burned
over during the hibernating season, the form that became the most
scattered prior to hibernation would be likely to stand the best chance
of surviving. It seems to the writer that the wings of the chinch

a Nineteenth Ann. Rept. Maine Agric. Exp. Sta., 1903, pp. 41-52.

16 THE CHINCH BUG.

bug might have been, in early days in the Mississippi Valley, kept
up to a high standard of development by the necessity of such an
escape from prairie fires and not by the presence of Sporotrichum
globuliferum, as suggested by Professor Saj6 in his paper, a trans-
lation of which is included herein under the heading, “ Habits of
the European species, Blissus doria Ferr.”

As mentioned farther on, the advance of civilization having revo-
lutionized the face of the country, there has come a corresponding
change in the hibernating habits of the chinch bug. This insect
must now seek shelter in the limited patches of timber that are left
in the sections that were once entirely wooded and in the matted grass
along fences and roadsides, but especially among the fallen leaves
and rubbish that usually accumulate along Osage orange hedges.
Brush piles, old haycocks, strawstacks, and, in Ohio, at any rate,
shocks of corn fodder left standing in the fields through the winter,
all harbor chinch bugs during the hibernating season.

The fact that the insect hibernates in matted bluegrass along road-
sides and fences has been called in question by Professor Forbes and
by Mr. Marlatt, the former in his first report as State entomologist
of Illinois (p. 37) and the latter in Insect Life (Vol. VII, p. 282),
but notwithstanding this, in some parts of Ohio, in Indiana, and
Uhnois they do hibernate in just such places and can be found there,
especially during the winter and early spring following a season of
abundance, but the investigator must know how to search for them.
The writer has found them late in the fall collected under rails, half
buried in soil and dead grass, and in northern Illinois, while search-
ing for other insects in early spring, he was sure to find them in vary-
ing numbers with small Carabide, Staphylinide, and other early
appearing insects, on the under side of boards laid down in grassy
places, though no amount of searching the grass itself would have
revealed their presence. .

In the timothy meadows of northeastern Ohio the percentage of
long-winged individuals is always much greater in fall than in June,
showing that some, at least, hibernate there and migrate to the cui-
tivated fields in spring. In Maine, in the case of the maritime form,
of 565 bugs collected in hibernation in October, 1902, only 60 had long
wings.” In Kansas, where Mr. Marlatt made his observations, there
was still too much prairie, and the species was doubtless still adhering
to its ancient habits of hibernation. In southern Ohio the author has
found it attacking the wheat in May, in small isolated spots over
the fields, while there was nothing in the least to imply an invasion
from outside, but the wheat had been sown in the fall among corn,
and later the cornstalks cut off and shocked, remaining in this condi-
tion until the following spring. This occurred so frequently that

a@19th Rept. Maine Agric. Exp. Sta., 1903, p. 48.

HIBERNATION. . We

there seemed no room to doubt that the attacks had been caused by
adults wintering over in the corn fodder, and that these left their
winter quarters in spring to feed and breed on the grain growing
nearest at hand.

Prof. Herbert Osborn,* in giving a summary of his observations
on the chinch bug in Iowa in 1894, states that “ In a great majority
of cases, 90 per cent or more, the infested fields were directly adja-
cent to hedges or thickets or belts of timber, and in 75 per cent Osage
orange hedges were the most available shelter. In about 13 per cent
of the cases the evidence showed hibernation in grass and weeds, and
in some of these cases there could scarcely be a doubt that the hiber-
nating bugs were protected by a heavy growth of grass or weeds and
that they moved from these directly into the adjacent grain fields.”
Prof. Lawrence Bruner had previously called attention to the fact
that the chinch bug hibernated in great numbers about Osage orange
hedges in Nebraska. Doctor Lugger, in Minnescta, gives the follow-
ing as offering shelter to the bugs during winter: “ Rubbish of all
kinds, but chiefly that of hedges, wind-breaks, and along the edges
of woods, as well as corn fodder, logs, and even loose bark and
stones.”

While drenching rains are beyond all possible doubt fatal to the
newly hatched young, the adult bugs seem to be almost proof against
either wet or cold weather. It is doubtless true that very many
individuals die in their winter quarters, and in fact the writer has
found these dead in considerable numbers in some instances during
early spring, but it seems at least doubtful if either cold or wet would
entirely account for this fatality. It would seem that somewhere
and at some period in the past this hibernation has been more for
protection from natural enemies than against the elements, though
of course there might have been other reasons not discernible under
a changed environment. The pupa hides away to molt, though it
does not appear that this course is followed in the earlier stages, and
the reasons for this are not at all clear. That the adult is able to
withstand combined cold and wet weather is amply proved by the
observations of several people. Dr. Hy. Shimer, in Illinois, found
that those which were in corn husks filled with ice, even the chinch
bugs themselves being inclosed in the crystallized element, were able
to run about when they were thawed out, apparently unaffected by a
temperature that had varied from 15° to 20° below zero Fahr. It
seemed that when exposed to the sweeping prairie winds at that
temperature, with no protecting cover, they perished. Mr. G. A.
Waters, in the Farmers’ Review for October 19, 1887, relates that a
bunch of fodder that had fallen into a ditch washed out near a corn

4Chinch Bug Observations in Iowa in 1894, Insect Life, Vol. VII, pp. 280-232.
26608—No. 69—07 m

2

18 THE CHINCH BUG.

shock by heavy rains became covered with water that stood over
it long enough for a sheet of ice to form. When the water had sub-
sided the corn was husked and a number of chinch bugs were found
among the ears, where they had been immersed for a week or more;
yet on being exposed to the warm sun they began to crawl about in
a lively manner.

The Maine Agricultural Experiment Station some years ago @ car-
ried out a series of experiments with the maritime form to determine
the effect- of freezing. Ten long-winged and 6 short-winged bugs
were frozen in an open box for fifteen hours. Upon thawing out 2
gave no signs of life. After being kept for nine hours at a tempera-
ture of 65° the 14 surviving bugs were refrozen for fifteen hours and
then thawed out, when 5 long-winged and 3 short-winged revived. .
After nine hours at a temperature of 65° they were frozen a third
time for fifteen hours, during which time the minimum temperature
sank to 16° below zero. When thawed out all revived, but during
the following nine hours at 65° temperature the 3 short-winged bugs
and 2 of the long-winged ones died. The remaining 3 long-winged
were then frozen a fourth time for fifteen hours, after which none
revived.

In summarizing the results of these experiments, 25 in number,
it was found that complete submersion in water, even for a considera-
ble period, is not necessarily fatal. Freezing during submersion in
water is almost surely fatal. Freezing while exposed to dry atmos-
phere is generally fatal. Freezing in a moisture-laden atmosphere
is only occasionally fatal. It will be observed, however, that not all
of these results would necessarily follow corresponding experiments
with the inland long-winged form. :

SPRING, SUMMER, AND AUTUMN MIGRATIONS.

If there is an ample supply of proper food close at hand the chinch
bug simply crawls from its hibernating place, but if it is in the timo-
thy meadows of northeastern Ohio it does nothing but continue its
ravages where it left off the autumn before, except some of the
long-winged form, which very evidently fly to the wheat and corn
fields. In wheat fields—unless the migration has been from an ad-
joining field, in which case the attack is made along the edge nearest
thereto—the females do not seem to forsake their gregarious habits
entirely, as they do not scatter out evenly over the entire field, but ap-
pear to locate in colonies, and when the young hatch and begin to attack
the growing grain their presence is first disclosed by small whitening
patches, which increase in dimensions as the young become older and
more numerous. In low-lying fields these whitening patches more

ne

¢ Nineteenth Rept. Maine Agric. Exp. Sta., 1903, p. 48,

SPRING, SUMMER, AND AUTUMN MIGRATIONS. 19

commonly appear on the back furrows or on any slight elevations
that occur in the field. But on higher and level ground the whitening
areas are observed scattered over the entire field, and constantly
widening until the whole field appears to ripen prematurely and
crinkle down. When the migration is accomplished by crawling,
the females seem to spread only enough to afford food for the young
until the latter are able to make their own way from place to place.
The young remain clustered on the plant about which they were
hatched until this has been drained of sap, when they make their way,
almost in a body, to a second plant, and in this way an attack will be
pushed forward day after day.

In the spring the chinch bug probably lingers about its winter quar-
ters until a favorable day for migration occurs. ‘Transfer a typical
Indian summer day to early May, and perhaps raise the tempera-
ture a few degrees, and you have a day during which chinch bugs
may be seen on the wing, crawling along on fences, or at rest on the
tops of fence posts as if taking observations, and in reality, as the
writer has come to believe, to catch the scent of wheat or corn fields.
It is on just such a day as this that Aphodius serval Say will be
observed posted in precisely the same way, opening and closing the
leaves of its antenne, evidently to catch the scent of the fresh drop-
pings of animals. The same movements characterize Aphodius inqui-
natus HUbst. during the Indian summer days of autumn. The writer
has also observed the plum curculio, Conotrachelus nenuphar Ubst.,
acting in precisely the same way in late autumn.

While discussing the subject of chinch-bug migrations, it may be
best to state here that there is a second flight of chinch bugs in sum-
mer after the majority have become fully developed, and not as soon
as the individual reaches the adult stage, as Professor Saj6 has found
to be the case with the European species, Blissus doriw Ferr. A
migration by flight takes place in the fall, usually during the period
of Indian summer. The magnitude of such migrations depends in
the spring on the number of individuals that have been in hiberna-
tion, and in the summer and fall entirely on the abundance of the
species during the current year. If there has been no great abun-
dance during the spring the summer flight will not be lkely to attract
attention. During .the ‘invasion of 1896 in Ohio an individual
alighted on the writer’s hand while he was riding on a street car in
the heart of the city of Columbus. A heavy storm of rain has much
influence in scattering the bugs in midsummer, and just preceding a
heavy rain the writer has noted the fully developed adults very
abundant on Indian corn plants, while immediately after the storm
there would be very few to be found. As these storms were not
always accompanied by high winds, it is probable that it is the rain-
fall that scatters the insects.

20 THE CHINCH BUG.

In timothy meadows where the original attack has begun along one
side and gradually extended inward, the line of separation between
the entirely dead grass and that uninjured is frequently not over a yard
in width, and within this narrow, irregular strip we may have the
dead and brown, the yellowing indicating more or less serious injury,
and the perfectly healthy green of unattacked plants. This many-
colored border may change but little in the space of a week or ten
days, except to advance very materially, leaving the grass completely
dead or dried up, while the clover plants are uninjured. This indi-
cates that the females, after leaving their places of hibernation, do not
spread out over any large area, but to a certain degree maintain their
gregarious habits. The author believes that these habits have been
shaped by some past environment in which the species has been placed
for a long period of time, as, for illustration, the inhabiting of
bunches or tufts of grass more or less isolated from each other.

To what extent pairing takes place in these places of hibernation
before the insects make their way to the cultivated crops is a matter
of considerable uncertainty. From his own observations the writer
is inclined to believe that only a very insignificant minority follow
this course.

In his “ Wanderings of Insects ” Prof. Karl Sajé has called atten-
tion to the influence of electrical storms in the dispersal of insects,
and it is quite possible that adult chinch bugs may be thus affected by
the heavy thunder that usually accompanies these storms, during
which they seem to disappear from corn plants on which they had
previously congregated.

OVIPOSITION.

According to most writers the eggs are deposited either about or
below the surface of the ground, among the roots of the grass or grain.
It is more than likely that the place varies with the conditions, as the
eggs are not infrequently found above ground about the bases of the
plants, and even upon the leaves, though we have never found them
there, but have often found them under the sheath of grasses. It
would seem, then, that the eggs require a cool, damp, but not a wet
location.

.

EGG PERIOD AND NUMBER OF EGGS DEPOSITED BY EACH FEMALE.

Doctor Shimer states that each female deposits 500 eggs, scattering
them over a period of from ten days to three weeks, and as the adult
develops in fifty-seven to sixty days after the eggs are deposited, or
about forty-two days after hatching, it will be seen that some of the
earliest hatched young are well along toward full development by the
time the last eggs are being deposited. According to Doctor Riley,
the eggs hatch, on the average, in two weeks.

DESCRIPTION OF DIFFERENT STAGES. 21

In a series of breeding-cage experiments Prof. W. G. Johnson found
thai each female deposited from 98 to 237 eggs, the egg period lasting
from eighteen to twenty-one days, and the period of oviposition cover-
ing from thirty-eight to forty-two days. Forbes also records in his
Fifth Report (p. 44) experiments showing that the period of incuba-
tion may cover from twelve to twenty-two days. (See Forbes’s 19th
Report, pp. 177-183.) It must be remembered, however, that Pro-
fessor Johnson had but six females employed in his experiments and
that these were necessarily under an artificial environment.

DESCRIPTIONS OF THE DIFFERENT STAGES OF DEVELOPMENT.

The following descriptions of the egg and various stages of the
young bugs are taken from Riley’s Seventh Missouri Report, while
that of the adult is from
the original by ThomasSay,
as published in his Ameri-
can Entomology (Vol. I, p.
329, Le Conte Ed.) : - f

The egg.—Average length 0.03
inch, elongate-oval, the diame-
ter scarcely 4 the length. The
top squarely docked and sur-
mounted with four small :

: E ; Fig. 2.—Blissus leucopterus: a, b, eggs; c, newly hatched
rounded tubercles near the cén- larva; d, its tarsus; e, larva after first molt; /, same after
ter. Color, when newly laid, second molt; g, pupa; the natural sizes indicated at
pale or whitish, and translu- sides; h, enlarged leg of perfect bug; 7, tarsus of same,

still more enlarged; 7, proboscis or beak, enlarged.
(From Riley.)

cent, acquiring with age an
amber color, and finally show-
ing the red parts of the embryo, and especially the eyes toward tubercled end.
The size increases somewhat after deposition, and will sometimes reach near 0.04
inch in length. (Fig. 2, a, b.)

Larval stages—The newly hatched larva is pale yellow, with simply an
orange stain on the middle of the three larger abdominal joints. The form
scarcely differs from that of the mature bug, being but slightly more elongate ;
but the tarsi have but two joints and the head is relatively broader and more
rounded, while the joints of body are subequal, the prothoracie joint being but
slightly longer than any of the rest. The red color soon pervades the whole
body, except the first two abdominal joints, which remain yellowish, and the
members, which remain pale.

After the first molt the red is quite bright vermilion, contrasting strongly
with the pale band across the middle of the body, the prothoracie joint is
relatively longer, and the metathoracic shorter. The head and prothorax are
dusky and coriaceous, and two broad marks on mesothorax, two smaller ones on
metathorax, two on the fourth and fifth abdominal sutures, and one at tip of
abdomen are generally visible, but sometimes obsolete; the third and fourth
joints of antennze are dusky, but the legs still pale. After the second molt the
head and thorax are quite dusky and the abdomen duller red, but the pale trans-
verse band is still distinct; the wing pads become apparent, the members are
- more dusky, there is a dark-red shade on the fourth and fifth abdominal joints,

22 THE CHINCH BUG.

and, ventrally, a distinet circular dusky spot, covering .the last three joints.
(ULE epee KOT ee sa),

The pupa.—tin the pupa all the coriaceous parts are brown-black,. the wing-
pads extend almost across the two pale abdominal joints which are now more
dingy, while the general color of the abdomen is dingy gray; the body above
is slightly pubescent, the members are colored as in the mature bug, the
three-jointed tarsus is foreshadowed, and the dark horny spots at tip of abdo-
men, both above and below, are larger. (Fig. 2, g.)

The adult.—Blackish, hemelytra white with a black spot.

Inhabits Virginia.

Body long, blackish, with numerous hairs. Antenne, rather short hairs;
second joint yellowish, longer than the third; ultimate joint rather longer
than the second, thickest; thorax tinged with cinereous before, with the basal
edge piceous; hemelytra white, with a blackish oval spot on the lateral middle ;
rostrum and feet honey-yellow; thighs a little dilated.

Length less than three-twentieths of an inch.

I took a single specimen on the Hastern Shore of Virginia.

The whiteness of the hemelytra, ine which is a blackish spot strongly con-
trasted, distinguishes this species readily.

To the foregoing description of the adult Dr. Asa Fitch, in his
second report on the Insects of New York, adds brief descriptions of
nine varieties, all, with but one exception,
being based upon slight variations in color,
some, perhaps, being due to immaturity, the
single exception being the short-winged in-
land form, of which variations from the
nearly wingless to fully winged are shown
in figures 3 and 4.

Leaving, then, out of consideration the
color varieties as arranged by Doctor Fitch,
we have a long-winged form (fig. 3) in
which individuals from the eastern portion
FIG. 3.—Blissusleucopterus: adult Of the country differ from those found in the

of long-winged form. Much West by being more hairy and robust, as

eee te pointed out by Mr. Van Duzee, and a short-
winged form (fig. 4), found along the seacoast, and in the North
Atlantic Coast region, extending inland as far as the country adjacent
to the Great Lakes.

DEVELOPMENT AND HABITS OF THE YOUNG.

The newly hatched young are very active, and the first to appear
may be observed with their progenitors about the bases of wheat, corn,
or grass plants, and later all stages are seen mingling together, having
little appearance of belonging to the same species, so greatly do they
vary in size and color in their several stages of development.

As a rule the bugs confine themselves to the lower portion of the
plants attacked, but may later push their way upward, especially if the
lower portion becomes tough and woody, finally covering the plants in

DEVELOPMENT AND HABITS OF THE YOUNG. DS

patches, as seen in figure 5, where they are shown on a stalk of young
corn. Mr. E. A. Schwarz relates a curious exception to this habit in
Florida upon sand oats, /niola paniculata, where the entire develop-
ment of the insect is undergone upon the highest part of this tall
plant and not close to the bottom. Mr. Schwarz has given as a proba-
ble reason for this the fact that strong winds are continually blowing
the fine, sharp sand through among the lower parts of the plants,
rendering it nearly or quite impossible for the bugs to remain in that
situation, thus forcing them to seek their sustenance farther up the
plants. While figure 5 gives a good representation of the appearance
of a corn plant when the chinch bugs are present in excessive numbers.
yet the writer has invariably found that these bugs much prefer a
stallx that has been blown down by the wind or partly broken off by
the plow and left lying nearly flat upon the ground.

In timothy meadows the very young are to be found only by pull-
ing away the soil from about the bulbous roots and drawing down

Fic. 4.—Blissus leucopterus: adults of short-winged form. Much enlarged (original).

the dead sheaths that usually envelop them. An observer may even
pull up a tuft of grass entire, and yet, unless he examines in this way
closely, may overlook them, so snugly are they thus ensconced among
the roots. If driven to forsake a tuft of grass the young bugs move to
another and crawl downward, and are soon to be found as snugly
settled as before. It is only when they are older and well advanced
toward maturity that they work to any extent above ground, and
even then only in cases where they are present in great numbers.
Singularly enough, where infested meadows are plowed up and
planted with corn the females seem to ignore the young corn plants
and select the occasional stray clumps of timothy that cultivation has
failed to destroy and deposit their eggs about these, so that later the
young may be swarming about these last, while hardly one is to be
found about the young corn. This is precisely the opposite of what
is observed farther west.

24. THE CHINCH BUG.

Although lying externally on their food plants, and notwithstand-
ing the fact that the young may attack the bases or even the roots of
some of these, the species is essentially an external feeder, and appears
while thus engaged almost totally indifferent to possible attacks of
natural enemies. When not feeding, however, there is at times a
tendency to hide away under the sheaths of young corn or beneath
clods of earth or bunches of coarse stable manure, where this has
been recently appled and left more or less exposed on the surface of
the ground. The writer has noted this in cases where neither an
uncomfortable temperature nor wet weather necessitated protection.

As has been shown in the description of the larval stages, there are
four molts between the egg and the adult state. Just how the molt-
ing larve act we have never been able to determine; neither have we
witnessed pupation, but a fully developed pupa that is ready to molt
is easily distinguished by its larger size and more tightly fitting skin,
which is almost shining white on the median ventral surface of the
abdomen. It now hides itself away, seemingly preferring to get
under the sheaths of grasses or grains; but if these are not con-
venient it will crawl under loose clods, or even into crevices in the
ground. While thus hidden away the pupa skin splits along the
back and the fully developed adult makes its way out, leaving the
empty skin behind. These last are very frequently mistaken for
dead chinch bugs, and, when moldy, the farmer is very likely to sup-
pose that they are bugs which have been killed by the fungus Sporo-
trichum globuliferum, if this has been applied in the fields.

On first emerging from the pupa the adult is generally of a dull
pink color, except the wings, which are white, exclusive of the veins;
these being of the same pinkish hue as the body. In a short time
these colors change to the normal ones of the species, but during the
breeding season these newly developed adults may be observed crawl-
ing about with the young of all stages as well as the maturely colored
adults.

If this development has been taking place in a wheat field and the
grain is harvested at this time, or if from any other cause the food
supply becomes suddenly exhausted, all sizes of larvee with pups and
adults will start off on foot to hunt for a fresh supply. Though
many individuals may now have become fully developed, and, so far
as can be determined, possess wings entirely fitted for active service,
nevertheless they will crawl along a dusty road or across freshly
plowed fields in company with their less fortunate fellows, seemingly
never for a moment supposing that they can span the intervening
space by flight. The writer is totally unable to account for this phe-
nomenon in the species at this time, the disinclination to use the wings
being so wholly unlike the habits of B. doriw, as shown by the careful
and painstaking observations of Professor Sajé in Hungary. Again,

NUMBER OF GENERATIONS ANNUALLY. 25

the seeming desire on the part of the pupz to secrete themselves while
transforming to adults does not at all coincide with the idea of a
supposed immunity from attacks of natural enemies. Surely our
species of Blissus has not always lived where natural enemies were as
few as they are with us at the present time. Even where we have
both the long-winged and short-winged forms occurring together in
timothy meadows there is no such haste exhibited on the part of the
former to escape from the companionship of the latter, as observed
by Professor Saj6. We know, however, that our species certainly
does enjoy a considerable immunity from natural enemies, though its
conspicuous colors in both the larval and adult stages contrast very
strongly with those of its usual food plants and its presence is still
further advertised by its strangely persistent gregarious habits. We
have come to suppose the species to be, in part at least, protected
from attack by its vile odor, and so, indeed, it may be in the United
States, but the writer fully believes that somewhere in its southern
habitat it will be found to have one or more enemies, like the ant,
Eciton hamatum Fab., of Central America, for illustration. Our
native ants, however, will seldom attack even the young.

NUMBER OF GENERATIONS ANNUALLY.

Over the most of its area of habitation in North America, at least,
the chinch bug is two brooded, though in northeastern Ohio the
writer has totally failed to detect the second brood, or, in fact, to
perceive any indications that a second brood occurs; but this will be
referred to later. As previously shown, there is not sufficient proof
at hand to warrant the statement that there is, even in the far South,
a partial third brood. It is probable that the number of broods of
this species annually has been primarily decided in its home in the
tropical regions by the wet and dry seasons occurring there, and that
we have in the North these same broods occurring at slightly differ-
ent periods under the influence of a change from wet and dry to hot
and cold seasons.

Belt, in his Naturalist in Nicaragua, has the following to say with
regard to the seasons on the northeastern side of that country: ‘“ The
rains set in in May and continue with occasional intermissions until
the following January, when the dry season of a little more than three
months begins” (p. 103). ‘The heaviest rains fall in July and
August, and at those times the brooks are greatly swollen.” ‘ In
September, October, and November there are breaks of fine weather,
sometimes lasting for a fortnight, but December is generally a very
wet month, the rains extending far into January, so that it is not
until February that the roads begin to dry up” (p. 104). It seems
that possibly we have here the key to the secret of the number of

26 _ THE CHINCH BUG.

broods annually of the chinch bug. That this insect may be able to
adapt itself still further to changed latitude and environments and
become single brooded is not at all impossible. As illustrating the
ease with which insects, at least some of them, can change their
habits to correspond with their environment, we have in South Aus-
tralia the following facts regarding the codling moth, Carpocapsa
pomonella I, of which, though being still double brooded, “ the
winter caterpillars hatch into moths irregularly from the beginning
of October until the middle of November and deposit their eggs
accordingly, giving rise to a succession of young caterpillars until
the beginning of December. About the third week in December the
first moths of the second brood begin to appear and deposit eggs, and
members of this second generation of moths continue hatching and
egg laying until the end of February.”

The author’s notes on the chinch bug in northeastern Ohio are
as follows: Very young larvee, with what appeared to be their pro-
genitors, were observed at Jefferson, Ashtabula County, within 11
miles of the shores of Lake Erie, June 16, 1893, there being no
advanced larve among them. On August 27, 1896, a few miles
south, at West Andover, in the same county, only adults were
observed in two days’ search, though some of these showed by their
color that they had but recently passed the pupal stage. In this
latter locality, May 7, 1897, the sexes were pairing, but no young
"were present so far as could be observed, while to the south and west
of this locality, June 8 and 9, precisely the same conditions obtained
as to the bugs, no young appearing at this time. Quite copious rains
might have destroyed the young, but within 15 miles of these locali-
ties, on July 14 of this year, larvee were found after first molt and
stages intervening between these and the adults. Near Youngstown,
on October 8, 1897, only adults were present, pairing was not in prog-
ress, and the insect was not pairing in Ashtabula County on August
27, 1896. June 9, 1898, only two very young larve could be found
at Salem, about 15 miles southwest of Youngstown.

Up to October 17, 1898, no young of a second brood had been
observed, though careful search had been made from time to time in
the fields and meadows of northeastern Ohio, and a large number of
adults which developed in July and August, and since kept in con-
finement, had not only not reproduced, but had shown no disposition
whatever to pair. On the other hand, in southwestern Ohio, in the
vicinity of Cincinnati, on September 24, where the species occurred
in abundance, fully 75 per cent were pups, the remainder being made
up of larve, some of them quite young, and adults in about equal

a

4George Quinn, in Journal of Agriculture and Industry, South Australia, Vol.
ito akiey

NUMBER OF GENERATIONS ANNUALLY. 27

proportions, some of the latter showing by their immature colors
that they had but just passed the pupal stage.

Hatching is not fully in progress in the Northeast before the 25th
of June, only an occasional individual having passed the first molt
before the 10th of July. In the hght of the information that has
been gained by these observations, the occurrence of a second brood
of young in northeastern Ohio is doubtful.

The late Dr. J. A. Lintner, in his studies of the outbreak of this
insect in New York State in 1882 and 1883, seems to have relied
much on the published habits of the species farther west—as, indeed,
the writer has himself done until recently—and made no exact
studies of the species at that time; and in his annual report, where
the outbreak is discussed, no absolute proof of the existence of a sec-
ond brood in New York is presented. The occurrence of a second
brood of young in northern Illinois, as indicated by Doctor Fitch,
has always been considered as settled, and in a more northern lati-
tude than northern Ohio, so that there must be some other influences
besides latitude to account for the phenomenon. That the species
has occupied this territory for many years is indicated by the obser-
vations of Mr. E. P. Van Duzee, of Buffalo, N. Y., who wrote that
the insect was as abundant twenty-three years ago as at the present
time, so that whatever effect on the,insect the recent occupation of
the country might have had, that effect has passed away and a con-
dition of what we might call equilibrium now exists here.

On July 7, 1889, in the extreme northern part of Indiana, the
writer found an abundance of young which had not yet molted for
the first time. Dr. A. S. Packard records adults as pairing at Salem,
Mass., June 17, 1871, as quoted by Doctor Lintner, while the latter
gentleman” records the young as occurring in Lawrence County,
N. Y., about June 5, 1883.

Hardly have the latest hatched young of the first brood developed
to the adult before the young of the second brood begin to appear.
In southern Ohio this is about the first week in August. Generally
these young do little injury, because the wheat has long since been
harvested and the corn is usually too far advanced and tough to
offer a desirable source of food supply, except in cases where fields
have been planted very late, and here the writer has known them to
work considerable injury, especially in seasons of severe drought that
prevented the rapid growth of the plants. Fall attacks on wheat are
rare, and the injury is never of a serious nature, as it is usually the
case that by the time the young wheat is large enough to invite attack
the chinch bugs are searching for winter quarters.

«Second Report State Entomologist of New York, pp. 148-164, 1885.
b Loe. cit., pp. 158, 159, 164.

28 THE CHINCH BUG.

In the timothy meadows of northern and northeastern Ohio, how-
ever, the principal injury is done during August and September, and
in favorable weather on into October. Now, if we allow sixty days
for development from the egg, it would be September before the ap-
pearance of the adults of the brood to which these various young
belonged. If all eggs were deposited immediately, it would be
November before the adults of the second brood would begin to
occur, a condition of affairs that has never been observed. <As pre-
vously shown in this bulletin, the first brood is fully developed in
northeastern Ohio by the first of September, but there certainly is
no indication that a second brood of young is developed during Sep-
tember and October. It would seem, then, that from northern Ohio
through New York, New England, and probably to Nova Scotia
the adults from the first brood of larve winter over, and that there is
here but one annual brood.

DESTRUCTIVENESS LARGELY DUE TO GREGARIOUS HABITS.

Attention has been directed previously to the gregarious habits
of the chinch bug, and we only refer to the phenomenon again be-
‘ause it is to this that its destructiveness is largely due. It is not
the excessive numbers, but the persistency with which they will
congregate en masse on limited areas, that renders their attacks
so fruitful of injury. With an ample supply of food the young
develop and leisurely diffuse themselves over the adjacent fields,
and there are neither swarming flights nor migrations. In 1884,
in northern Indiana, a small field of wheat was severely attacked
by chinch bugs. At harvest there was every prospect of a migration
from the field of wheat to an adjacent one of corn, and the bugs were
present in sufficient numbers to have worked serious injury to the
latter: but the wheat had grown up thinly on the ground, and there
had sprung up among the grain a great deal of meadow foxtail grass,
Txophorus (Setaria) glaucus, and panic grass, Panicum crus-galli,
and to these grasses the bugs transferred their attention, finishing
their development thereon, and later, so far as could be determined,
they scattered by flight out over the adjacent fields, working no fur-
ther injury. Pedestrian migrations may continue for a fourth of a
mile or even more, but on reaching a suitable food supply the tend-
ency of the bugs is to congregate upon their food plants until these
are literally covered with individuals: varying in color from the
black and white of the adults to the bright vermilion of the more
advanced larvae. (See fig. 5.) Whatever tendency there is ex-
hibited toward a wider diffusion is confined to the adults, the others
remaining and leaving in a body only when the plant on which
they have congregated has been drained of its juices and has begun

FOOD PLANTS. 29

to wither, when they simply crawl to the nearest plants and again
congregate upon these as before. In case the migration has been
to a field of corn, if this is badly overgrown with either of the two
grasses previously named, the bugs will collect upon the latter, and
unless the corn plants are very small they will not as a rule attack
them until the grass has been killed. Some farmers have gone so
far as to claim that a benefit is derived from a certain abundance
of chinch bugs, the statement being made that the bugs will lll
out these grasses to an extent that nothing else will. It is clear
that the acquisition of wings is not the signal for the adults to
abandon the companionship of the larve and pup, yet they do
gradually disappear from among them. It is possible that the
disposition to pair does not exist until the individual has reached
a certain age beyond seeming maturity, and that it is not until the
passion for mating has overcome their gregarious inclination that
they are disposed to migrate. Or it may be that the phenomenon
may be explained on the supposition that when the pairing season
approaches the males scatter out in order to find females with which
they are not akin, thus following out natural selection and prevent-
ing a continual interbreeding. Over the northern United States,
at least, the injury in cultivated fields is done almost entirely by the
young bugs, but in the timothy meadows the damage is due as much,
if not more, to the depredations of the adults.

FCOD PLANTS.

As to food plants, there can be no doubt that these consisted origi-
nally of the native grasses. This is amply proved by the observa-
tions of Fitch and Le Baron, in Ulinois; Dr. J. C: Neal, in Florida
and Oklahoma; Marlatt, in Kansas; Schwarz, in Florida; and by
those of Mr. Henry G. Hubbard in the midst of the Colorado desert
in California. Regarding this last statement, Mr. E. A. Schwarz
wrote as follows:

You may be interested to learn that chinch bugs were collected this year
(1897) on March 28 by Mr. H. G. Hubbard, at Salton, in the midst of the
Colorado desert of California, This locality is considerably below the ocean level,
and represents an ancient extension of the Gulf of California. Even at the
present time the Salton Basin is occasionally flooded, the water entering through
New River, which runs from the mouth of the Colorado River into the Salton
Basin. The specimens were taken on a species of coarse grass which is
incrusted with a saline deposit.

No wonder that the chinch bug is accused of being a seashore
species !

Of cultivated grasses, or such as occur in cultivated fields, probably
Txophorus glaucus and Panicum crus-galli are the favorites, though
millet and Hungarian grass are apparently nearly as attractive. As

30 THE CHINCH BUG.

early as 1845, in Illinois, Dr. William Le Baron, afterwards State
entomologist, gave the food plants of the chinch bug as follows:

iG. 5.—Corn plant two feet tall infested with chinch bugs. (Author’s illustration.)

% * ok

“all kinds of grain, corn, and herd’s-grass” (timothy).¢
But to this day in Illinois, as shown by the observations of Professor

@ Prairie Farmer, December, 1845.

INSECTS MISTAKEN FOR CHINCH BUGS. ol

Forbes and the writer, the species will attack timothy only in cases
where it is compelled to do so by reason of a lack of other food. In
addition to the preceding, Doctor Howard gives broom corn, sorghum,
chicken corn, Bermuda grass (Capriola dactylon), bluegrass (Poa
pratensis), crab grass (Syntherisma sanguinalis), and bottle grass
(Ivophorus viridis), and also states that in the rice fields near
Savannah, Ga., in August, 1881, he observed the winged adults upon
the heads. Prof. H. A. Morgan wrote that in 1897 it had become.a
serious enemy to “ Providence ” rice in Louisiana, where for two years
it had seriously injured corn, and the writer was informed through
other sources that 1t proved injurious to corn again in 1898. Adults
have often been found collected in the silk of belated ears of corn in the
fields in September, when all other parts of the plant had either become
too old and tough to afford nourishment or else had been killed by the
frosts of autumn. Prof. Lawrence Bruner has recorded the insect as
feeding upon so-called wild buckwheat (Polygonum dumetorum or P.
convolvulus).© The writer has never seen
chinch bugs attack bluegrass (Poa pra-
tensis), and has seldom witnessed them in-
juring oats, but on September 27, 1904, he
observed larvee, pupx, and adults, the last
all fully winged, attacking Arrhenatherum
(oat grass) on the experiment farm of the
University of Tennessee, at Knoxville.
Over the western country the major por- F%9- &—Nysius angustatus: b, pupa;
: s M ¢c, mature bug. (From Riley.)
tion of the damage done is to fields of

wheat, barley, rye, and corn, the outbreak generally originating in
wheat or barley fields and the bugs migrating at harvest to the corn-
fields. (See fig. 5.) In the eastern part of the country, where the
timothy meadows are the most seriously infested, this is not the case,
and here the migrations are as likely to be to the timothy meadows as
to the fields of corn, where both are equally within reach. Besides,
everything indicates that a very large proportion of the adults may
hibernate in these meadows, even making their,way thereto in the
autumn.

INSECTS THAT ARE MISTAKEN FOR CHINCH BUGS.

Messrs. Osborn and Mally” have given a list of twelve species of
Hemiptera which have been mistaken with more or less frequency for
the chinch bug, the list being as follows: ;

Nysius angustatus Uhl., the false chinch bug (fig. 6), is probably

the most frequently mistaken for the true chinch bug, as it often

@ Report Commissioner of Agriculture, 1887, pp. 57-58.
6 Bul. No. 32, Iowa Agr. College Exp. Sta., pp. 363-885.

32 THE CHINCH BUG.

breeds in considerable numbers under purslane, amaranth, ete., and
more than any other insect resembles the chinch bug. It is, however,
of a light-gray color, which will always distinguish it from its more
destructive fellow.

Ischnodemus falicus Say, or the long chinch bug, as it is sometimes
called, is much larger and longer than the true chinch bug.

Ischnorhynchus didymus Zett. is more robust, of
a light-tawny color, with prominent, glassy wings.

Peliopelta abbreviata Uhl. is, next to the false
chinch bug, probably the most often mistaken for
the true insect, and especially is this true in local-
ities where the brachypterous form of Blissus
leucopterus abounds, viz, in timothy meadows. Its
broader head and body, however, quickly enables
one to distinguish it.

Geocoris fuliginosus Say, G. borealis Dallas, G.
Fie. 7.—Piesma cm byllatus Say, and G. limbatus Stal, according to
erea. (From Riley.)

Osborn and Mally, have all been confused with the
chinch bug in Iowa. These are all broader and flatter than the true
chinch bug, the head being nearly as wide as the thorax.

Ligyrocoris sylvestris ia is larger than the true chinch bug, and
its wings are quite dark instead of white.

Prapesonotus nebulosus Fall. is a trifle larger and its body is not
so black as in the chinch bug.

Cymodema tabida Spin. is longer than the true chinch bug, of a
light brown color, and the ends of the wings are glassy.

Triphleps insidiosus Say, or the insidious flower bug (fig. 15), as
it is more commonly called, is another bogus chinch bug, though an
enemy of the true pest, as previously stated.

Piesma cinerea Say, the ash-gray leaf bug (fig. 7), is often mis-

taken for the true chinech bug, though its form
differs greatly from that of the latter. It is often
quite abundant, but not in grain fields or rt
meadows.

Corimelena pulicaria Germ., the flealike negro
bug (fig. 8), has been confused with the chinch *
bug; though it does not in the least resemble the Pais tee Bienes
latter, either in form or color, and its confusion is
probably to be accounted for by the fact of its being occasionally
found in wheat fields in considerable numbers.

Brachyrhynchus granulatus Say (fig. 9) has been mistaken for
the chinch bug in Ohio, and in a way that was somewhat amusing.
Farmers in southern Ohio, during the winter of 1896-97, were burn-
ing over the woodlands with a view to destroying the hibernating
insects, when there came several discouraging reports to the’ effect
that such a course would be ineffective, as the bugs were wintering

LOSSES CAUSED BY CHINCH BUGS. 33

in the tops of trees, especially where the tops were dead, under the
bark and often from 50 to 75 feet from the ground. This was a
piece of astounding information, to the writer at least, and it was
only after securing specimens that he was able to solve the mystery.
This insect, in all stages of development except the egg, hibernates
under loose bark. It is broader and much flatter than the true chinch
bug, but the wings are white and the body black.

The object in calling attention to these bogus chinch bugs is to
prevent their confusion with the true Blissus leucopterus, as in some
cases people finding them and supposing them to be the true pest
are likely to become panic stricken and often destroy property un-

‘necessarily, so notorious has the name * chinch bug” become in the

_United States.
LOSSES CAUSED BY CHINCH BUGS.

It would appear that this pest first made its presence known by
its ravages in the wheat fields of the North
Carolina farmers; for we are told | ~ Pee that “in
1785 the fields in this State were so overrun
with them as to threaten a
total destruction | | of the grain.
And at length the crops
were so de- :
stroyed in some ie : {
districts that eal fe Sage Ne |
farmers were oo ae Seu Oe
obliged to By oaeendi | " SW f°.
abandon the es | a
sowing of wheat. | | ng |
It was four or ~ mises :
five years that

© P Fic. 9.—Brachyrhynchus granulatus: a, early nymph; 0, adult; ec, late
they continued nymph. All enlarged (original).
so numerous at this time.” ¢

In the year 1809, as stated by Mr. J. W. Jefferys,’ the chinch bug
again became destructive in North Carolina to such an extent that
in Orange County farmers were obliged to suspend the sowing of
wheat for two years. In 1839 °¢ the pest again became destructive in
the Carolinas and in Virginia, where the bugs migrated from the
wheat fields at harvest to the corn, and in 1840 there was a similar
outbreak, and both wheat and corn were seriously injured. In all
of these cases, however, there is no recorded estimate of the actual
financial losses resulting from the attacks of the chinch bug. Accord-

@ Webster on Pestilence, Vol. I, p. 279. Not seen. Quoted from Fitch.
6 Albany Cultivator, first series, Vol. VI, p. 201,
¢ The Cultivator, Vol. VI, p. 103.

26608—No. 69—07 m——3_*

34 THE CHINCH BUG.

ing to Le Baron, during the years from 1845 to 1850 the insect
ravaged over Illinois and portions of Indiana and Wisconsin, and
in 1854 and 1855 it again worked serious injury in northern Illinois.
The writer’s earliest recollection of the chinch bug and its ravages
in the grain fields of the settlers on the prairies dates from this
last outbreak. Mr. B. D. Walsh estimated the loss to the farmers
of Illinois in 1850 at $4,000,000, or $4.70 to every man, woman, and
child living in the State. The earlier outbreaks, though the occa-
sion of smaller money loss, were even more disastrous; for the
destruction of the grain crops in those pioneer days not only took
away all-cash profits, but also deprived the early settlers of their
very living, and in some cases. reduced them to starvation.

In 1863, 1864, and 1865 the insect was again destructive in I]linois
and other Western States, its ravages being especially severe in 1864,
when we have another attempt at computation of the financial loss.
Dr. Henry Shimer, of Mount Carroll, Ul., who had carefully studied
the chinch bug, estimated that “ three-fourths of the wheat and one-
half of the corn crop were destroyed by the pest throughout many
extensive districts, comprising almost the entire Northwest.” In
criticising the doctor regarding another point, Messrs. Walsh and
Riley, in The American Entomologist (Vol. I, p. 197, 1869), admit
that the estimate was “a reasonable one,” and, taking it as a basis,
with the actual cash price per bushel, computed the loss at about
30,000,000 bushels of wheat and 128,000,000 bushels of corn, with a
total value of both amounting to over $73,000,000. Of course all
computations of this sort are necessarily only approximately cor-
rect, but there is more likelihood of an under than an over estimate
in this case.

There was a serious outbreak of the chinch bug in the West again,
in the year 1868, and again in 1871, but in 1874 the ravages were both
widespread and enormous. Doctor LeBaron computed the loss in
1871 in seven States, viz, Iowa, Missouri, Illinois, Kansas, Nebraska,
Wisconsin, and Indiana, at $30,000,000.. Doctor Riley computed the
loss in Missouri alone in the year 1874 at $19,000,000, and added the
statement that for the area covered by Doctor LeBaron’s estimates in
1871 the loss in 1874 might safely be put down as double, or upward
of $60,000,000.2. Dr. Cyrus Thomas, however, estimates the loss to
the whole country for the same year at upward of $100,000,000.°

The next serious outbreak of the chinch bug of which we have the
losses resulting therefrom computed, occurred in 1887, and covered
more or less territory in the States of Kentucky, Ohio, Indiana,

a Second Report State Entomologist of Illinois, p. 144.
b Seventh Report State Entomologist of Missouri, pp. 24-25

lod

¢ Bulletin No. 5, U. S. Entomological Commission, p. 7.

LOSSES CAUSED BY CHINCH BUGS. 35

Illinois, Wisconsin, Minnesota, Iowa, Missouri, and Kansas. In this
case the damage was estimated by the United States statistician, Mr.
J. R. Dodge, at $60,000,000, the heaviest losses occurring in Hlinois,
Iowa, Missouri, and Kansas. This gives us as the estimated loss
in the thirty-eight years, 1850 to 1887, both inclusive, the enormous
sum of $267,000,000.

There was a serious outbreak in Kansas, Iowa, Minnesota, and I]li-
nois, having its beginning probably as early as 1892, but reaching its
maximum severity, as in Ohio, in 1896. The loss in Ohio during the
vears 1894, 1895, 1896, and 1897 could not have fallen far short of
#2,000,000. The farmers of this State in many cases were entirely

Fic. 10.—Map showing areas in the United States over which the chinch bug occurs in
greatest abundance and may at any time become destructive. (Original.)

unfamiliar with the chinch bug and its ravages, and therefore were
unable to account for the damage that it worked in their fields until
some time after. This was especially true of the timothy meadows in
the northeastern part of the State; so that there were probably many
fields, both of grass and of grain, that suffered seriously, and, in fact,
im some cases were ruined by the chinch bug without the owners being
aware of the cause. For this reason, while the computed loss appears
large, it seems to me to be entirely reasonable. Of the losses ocea-
sioned in other States during the years above indicated no definite
computations are available, but they were severe, and must have
amounted to millions of dollars. If we could have careful estimates
of the loss during the last fifteen years, it would in all probability

@ Report of U. S. Commissioner of Agriculture, 1887, p. 56.

36 THE CHINCH BUG.

swell the amount to considerably in excess of $330,000,000 for the
period from 1850 to 1906. Within the last ten years the insect has
become more injurious in Oklahoma, western Kansas, and northern
Texas, localities not included in these estimates, and although the
spring rains serve to destroy the young bugs, outbreaks in northern
Texas and Oklahoma are not rare in fields.of wheat, corn, and barley.
If the indirect losses were to be added, the amount would indeed be
enormous. During the outbreak in Ohio at least two farmers became
discouraged, and, thinking that the loss of their crops by the attack
of chinch bugs would result in their financial ruin, in their despond-
ency they sought relief in suicide.

When we take into consideration that the financial losses as above
estimated have not fallen upon the entire nation, but almost without
exception upon the nine States previously named (see fig. 10), it will
be seen that this diminutive insect constitutes a formidable enemy to
the agriculturist of these States. In fact, small as it is, this pest has
cost the people of these nine States a sum of money which, a few
years ago, would have defrayed the entire expense of the National
Government for a whole year. Fire excepted, there is probably no
other factor that has caused such an enormous financial loss within
the same period over the same area of country.

NATURAL CHECKS.

All adverse natural influences affecting the chinch bug will be
treated under this head, with the exception of animal and vegetable
foes, which are considered here as natural enemies.

INFLUENCE OF PRECIPITATION ON THE CHINCH BUG.

There is probably no more potent factor in restraining the increase
in numbers of this species than is to be found in meteorological influ-
ences consequent upon rain. The fact has long been known that the
years of greatest abundance of the chinch bug were preceded by a
series of years during which there had been a deficiency in the rainfall
over the area of country devastated by this species. In fact, it has in
a general way come to be understood that dry seasons are favorable
and wet seasons unfavorable for the development of the chinch bug,
though the details of the phenomenon have never been very carefully
and elaborately worked out. The entomological and meteorological
records of the past have, however, clearly shown that the amount of
the annual rainfall is not a safe guide in this problem. Chinch bugs
have occurred in excessive numbers during years of heavy
precipitation.

The term “ wet season,” so frequently used in this connection, is an
indefinite one, but if the term “ season ” be restricted to the period of
time intervening between the vernal and autumnal equinoxes we shall

INFLUENCE OF TEMPERATURE ON CHINCH BUG. 37

have more definite grounds upon which to base our studies of meteoro-
logical influences. Thus applied, the terms “ wet” and “ dry ” sea-
sons would include within them the two breeding periods of the
chinch bug, at least largely so, north of latitude 30° N. But the his-
tory of this species has shown that there may be an excess of rainfall
during this critical period and that still a sufficient number of insects
may develop to work serious injury over considerable areas of coun-

aim!

Vl

GUERNSEY
=
6

,
,
y

m
: \

H1g. 11.—Map showing distribution of chinch bug in Ohio in 1896. (Author’s illustration. )

try. This is due to two, and perhaps more, causes. In the first
place, an unusually heavy rainfall at long intervals, while bringing
up the total for a given period, may have but little effect in reducing
the number of chinch bugs, while a much less amount of precipitation
coming at short intervals and in the midst of the hatching season
would cause a far greater mortality among the young. And, in the
second place, the precipitation may come at the beginning or even

38 THE CHINCH BUG.

before the commencement of this breeding season or just at the close
thereof, thus enabling the major portion of the young to reach a
period in their development wherein they are little, if at all, sus-
ceptible to the effects of drenching rains. This was clearly illus-
trated in southern Ohio during the spring of 1896, and again in 1897.
Throughout southern Ohio, in 1896, between latitude 38° 30’ and 39°
40’, as the reports of the United States Weather Bureau show, there

OEFIANCE

PAULDING

ate
TUSCARAWAS
eit. ere

OELAWARE

LICKING
4 FRANKLIN
MADISON

PICKAWaY

Fig. 12.—Map showing distribution of chtnech bug in Ohio in 1907. (Author's illustration. )

had been but very little rain up to May 11, and no general
rain until May 25. The effect upon the young bugs, judging from
the destruction which they caused, would seem to have been to
destroy only the latest to hatch, leaving the earlier developing
young sufficiently advanced to withstand the effects of the later and
heavier rains. The accompanying map (fig. 11) will show the areas
over which chinch bugs were reported marked thus =—, while the

INFLUENCE OF PRECIPITATION ON CHINCH BUG. 39

area seriously ravaged is indicated thus #, showing that the rain
came too late in such a section to ward off an outbreak of the pest.
According to the Weather Bureau reports also, the distribution of
rain in May, 1897, differed materially from that of the same month
of 1896, in that in 1897 the major portion of the rain fell prior to
the 15th, the remainder of the month being rather dry, the only

general precipitation occurring on the 23d and 24th, with a much

ASMTABULA

AULOING

Ss
SS

COSHOCTON HARRISON

j
y)

Fic. 13.—Map showing distribution of chinch bug in Ohio in 1894. (Author’s illustration.)

lighter rain on the 28th. But here again the amount was insufficient
to’ ward off serious injury, as is indicated by map (fig. 12), the same
symbols being used here as before. In this case it was probably the
latter portion of the brood that survived, as a personal inspection
of the country early in the month failed to reveal the presence of
young bugs, though they were certainly present in abundance at a
corresponding period of the preceding year.

40 THE CHINCH BUG.

That the amount and frequency of rain during the month of May
has very much to do with the ravages of chinch bugs where sufficient
numbers have wintered over to produce the requisite number of
young, is further shown by the fact that in 1894 the only locality
where serious ravages were committed was in Wyandot County, as
shown on map (fig. 18), and this was one of the few areas in Ohio

where the precipitation during that month was less than 8 inches.

i

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ic. 14.—Map showing distribution of cninch pug in Ohio in 1895, and amount of pre-
cipitation over the State during May of the same year. (Author’s illustration.)

Except over a circular area covering less than one-half of the county
the amount of precipitation was 3 to 5 inches, and this area includes
that ravaged by the chinch bugs during the following month.

Still more striking, however, is the relation between the two
phenomena during the following year. The last of this series of
maps (fig. 14) shows the area over which chinch bugs were reported

INFLUENCE OF PRECIPITATION ON CHINCH BUG. 4]

and the area where their injuries were the most severe; also, by
horizontal lines, the areas over which the amount of precipitation
was the least. From this it will be observed that in all of the seri-
ously affected area, and in nearly all of the area over which the
pest was reported at all, the precipitation during the month of May,
1895, was from 1 to 2 inches,.the extension of the point westward
into Shelby County being especially interesting. It may be said
with regard to the occurrences outside of this area of light precipita-
tion that the exact localities were probably not indicated, as the
information was secured from farmers, and their locations as indi-
cated on the map were their post-office addresses, which might have
been several miles away in any direction, and the isolated points of
attack were often based upon one or two reports. If exact localities
could have been obtained, and the precise area of precipitation indi-
cated, the connection between the two phenomena would have been
shown more correctly, and would probably have revealed an even
greater uniformity than is now apparent. It must be understood,
however, that in these calculations northeastern Ohio is excluded,
and the writer believes that what is true of the rest of the State
will be found to be equally correct as regarding territory occupying
the same latitude westward to the limit of this area of distribution.
While it is probable that the effect of precipitation during August
would have a similar influence on the second brood of young, and,
consequently, upon the number of adults which would go into winter
quarters, yet a careful study of the two factors shows that meteoro-
logical conditions in August have a far less influence upon the fol-
lowing brood than do those of May.

Owing to causes which are as yet unknown to the writer the same
laws do not apply to the northeastern part of Ohio and to what we
have termed the west-bound tide of migration. Here, and as against
the more or less short-winged form of chinch bug, meteorological con-
ditions appear to exert a far less potent influence. What is true of
meteorological conditions during May elsewhere in Ohio, seems to
be partly true of June in the northeastern portion of the State,
though there is not the evidence of the effect of precipitation here
that we have elsewhere. Doctor Lintner, in his Second Report, while
discussing the outbreak of the chinch bug in New York during 1882-83,
calls attention to the fact that both in 1881 and 1882 there was an
excess of precipitation. On page 158 of his report Doctor Lintner
says that spring, summer, and autumn were exceptionally wet. In
spring heavy and continued rains flooded meadows which, later,
showed the effect of chinch-bug attack. Even at haying time while
the bugs were young and, according to all accounts, easily killed
by heavy rains, they persisted in multiplying and living despite
the fact that rains were so freauent and severe that only a portion

492 THE CHINCH BUG.

of the hay could be gathered in a proper condition. This was the
state of affairs on July 5 when the hay was cut, and on October 10
Doctor Lintner stated that owing to continued rains grass was still
lying in the fields and could not be gathered, while fields of oats
remained unharvested. In all of the reports given of this outbreak
it was stated that the damage was first observed in August or Sep-
tember, and it is believed that this will hold good as applied to
northeastern Ohio.

As has been stated, the females oviposit as a rule at or just below
the surface of the ground, and the young make their way upward
in order to secure food. In case of cultivated grains this mode of
procedure is absolutely imperative, as the bases of the plants are at
that time too tough and woody to offer sufficient food. But in the
case of timothy the conditions are entirely different, as the bulb of
this plant, situated just below the surface of the ground and con-
venient to the place of oviposition, furnishes an ample supply of
food without making it necessary for the young to crawl upward
in order to secure it. Then, too, the surface of the ground in cul-
tivated fields is nearly or quite free of dead leaves and stems, there
being little but the vertical-growing plants to afford protection from
the weather. In timothy meadows the surface of the ground is
usually covered to the depth of an inch or more with dead and decay-
ing stubble and leaves, and the top of the ground itself is often more
or less loose and meilow in the immediate proximity to the bulbs of
the plants. It would appear that we might here have a partial solu-
tion of the problem of the vital effects of precipitation:on the young
bugs. Besides, for aught we know, the progeny of this quite short-
winged form may be better able to withstand naturally the effect of
drenching rains than that of the east-bound long-winged form. We
must recollect that in the one case the progenitors have worked their
way over hot, arid plains as well as cool, damp prairies, while in the
other case the tide of migration lay between the more elevated lands
and the sandy beaches of the seashore where there was always a more
or less near proximity to the ocean, until the tide of migration left the
seashore and drifted westward over New York and onward into
northeastern Ohio. (See map, fig. 17.)

This influence of precipitation on the young chinch bugs while in
the act of hatching, and that of temperature upon the adults in winter,
some illustrations of which have been included under the subject of
hibernation, are the only cases where meteorological conditions appear
to have a direct effect on this species. As previously shown, the tem-
perature effects are, largely at least, unfavorable for such adults as
may happen to be more or less unprotected during the hibernating sea-
son. Upon this point it might be well to suggest that this protection,

INFLUENCE OF PRECIPITATION ON CHINCH BUG. 43

which may be composed of leaves and dried grass, may be burned away
in early winter and thus leave the insects without the expected protec-
tive covering, or this covering may be still further augmented by a
mantle of snow, which, remaining for a more or less protracted period
of time, counteracts the influences of temperature, and the latter then
becomes a factor of secondary importance in the problem of life among
chinch bugs. It is very doubtful if temperature is as vital in its ef-
fects as are the indirect influences of precipitation during the breeding
season.

It has long been understood that the two species of entomogenous
fungi, Sporotrichum globuliferum Speg. and Entomophthora aphidis
Hoifm., both of which attack the chinch bug, require for their rapid
development an atmosphere heavily charged with moisture, and that
without this neither of these becomes sufficiently abundant to cause
any serious mortality among the insect host, but this matter will
receive attention in the discussion of these parasitic foes farther on.

INFLUENCE OF TEMPERATURE ON THE CHINCH BUG.

The writer would call attention here to a possible influence of
temperature upon what he has termed the west-bound tide of migra-
tion. When the time arrives for the hibernating adults to leave
their winter quarters and disperse over the fields prior to oviposition,
if the weather should prove too severe they have but to remain in
these quarters a while longer until more favorable weather. Thus,
along the northern Atlantic coast the season is generally much later
near the shore than it is a few miles inland, and Mr. Schwarz ¢
has called attention to the influence which this phenomenon exerts
upon the chinch bug. Now, this retardation amounts probably to
about a month in spring, which would have a tendency to delay ovi-
position, especially among the short-winged females. If this were
continued through a long period of time, consequent upon the slow
movement of this tide of migration northward along the coast, it
would hardly be surprising to find that this retarded activity in
spring had become so characteristic as to be retained after this tide
had swept to the westward, and resulted in the species being thus
single brooded in the East, while it is double brooded in the east-
bound tide of migration in the West. This effect of a long habita-
tion along the shores of the northern Atlantic would be to some extent
encouraged by the prolonged northern winter and the correspond-
ingly shorter period during which the species could breed, and thus
instead of the effects of the old environment becoming obliterated
they might be continued, or, as in case of the fore-shortening of the
wings, still further intensified. If the effect of this prolonged period

a Insect Life, Vol. VII, p. 422.

44 THE CHINCH BUG.

of hibernation has been to reduce the number of broods, then it will
have to be considered as a natural check, in that to a certain extent it
prevents excessive abundance by reducing the number of offspring.
This would also account for the rather surprising immunity that has
heretofore been enjoyed by the northeastern portion of the country
from the ravages of this destructive species.

NATURAL ENEMIES.

It is possible that there are some reasons which might appear to
justify the placing of fungous enemies of the chinch bug among the
natural checks, as they no doubt do exert a more or less powerful
influence in that direction, but it seems more convenient to include
them among natural enemies, especially as one at least has come to be
applied artificially to overcome the insect. The fact that the abun-
dance and consequent influence of these fungous enemies is almost
entirely dependent upon meteorological conditions is sufficient to
place them in a secondary position, even though they may under
favorable meteorological conditions act as natural checks. Al, doubt-
less, have other host insects, and the two most important have been
known to break out again and again spontaneously and destroy myr-
iads of chinch bugs when the latter were present in excessive numbers.
But this has taken place only in connection with the necessary pre-
cipitation; hence these fungi become natural enemies only under
certain favorable weather conditions; and though their season of
most potent effect is during the time when the chinch bug is develop-
ing from the egg to the adult, yet as shown by observation they may
exert powerful and fatal effects among the adults, where these last
have congregated together in masses.

PARASITIC FUNGI.

The two species of entomogenous fungi to which reference has just
been made are L'ntomophthora aphidis Hoffman * and Sporotrichum
globuliferum Speg,’ both having probably been associated in destroy-
ing the chinch bug spontaneously in the fields, and doubtless were
distributed to correspondents by Professor Snow and others to be
artificially established in fields where there was an overabundance of
chinch bugs. For this reason it is impossible to separate the effects
of the two in the earlier literature, even the first observations of Dr.
Henry Shimer“ probably applying to their joint effect.

« Hoffman, in Fresenius’s “ Entomophthorew,” p, 208, figs. 59-67,
»b Spegazzini, ‘Fungi Argentini,” II, p. 42.
¢ Proc. Acad. Nat. Sci. Phila., May, 1867.

PARASITIC FUNGI. 45

Doctor Shimer, however, was the first to call attention to the wide-
spread and fatal effects of fungous diseases among chinch bugs,
and while his explanations therefor seem now crude and illogical,
his observations were made with such care and accuracy that we have
not yet had occasion to materially revise them. Under date of July
16, 1865, he makes this observation: “* * * [found many dying
on the low creek bottom land from the effects of some disease, while
they are yet in the larval state—a remarkable and rare phenomenon
for insects thus in such a wholesale manner to be dying without at-
taining their maturity, and no insect enemy or other efficient cause
to be observed capable of producing this important result.” Again,
under date of July 22: “ On low grounds the chinch bugs are dead
from the disease above alluded to, and the same disease is spreading
to the hills and high prairies.”

Under this date also he speaks of the very wet weather, and states
that in a barley field the chinch bugs began to die at about the same
time that they did on the low creek bottom, and that they rapidly met
the same fate, so that few of them lived to find their way to a neigh-
boring cornfield, while under date of August 8 he states that of those
that migrated to the cornfields “ very few are to be found remaining
alive; but the ground around the base of the cornhills is almost liter-
ally covered with their mouldering, decomposing dead bodies. They
are dead everywhere, not lying on the ground alone, but sticking to
the blades and stalks of corn in great numbers, in all stages of de-
velopment, larva, pupa, and imago.”

“This disease among the chinch bugs was associated with the long-
continued wet, cloudy, cool weather that prevailed during a greater
‘portion of the period of their development. * * *”

These are precisely the conditions under which these fungi have
been observed to prove the most fatal to the chinch bug during recent
years, where their introduction among the host insects was accom-
plished by artificial means. Although Doctor Shimer probably never
anticipated the artificial cultivation of his “ disease,” and the results
which have since been obtained from its artificial dissemination in the
fields, yet his careful and painstaking studies must ever be associated
with the application of fungous diseases in the destruction of insects
in America. It is certainly to be regretted that such practical ento-
mologists as Mr. B: D. Walsh and Dr. C. V. Riley should have
expressed themselves so discouragingly regarding Doctor Shimer’s
observations and conclusions, Doctor Riley, as late as 1870, even
going so far as to ridicule the theory of disease being in any way
responsible for the death of the chinch bugs observed by Doctor
Shimer.*

@Second Report State Hntomologist of Missouri, pp. 24-25, 1870.

46 THE CHINCH BUG.

It was not until 1879 that an entomologist came to the rescue of
Doctor Shimer’s theory of disease among chinch bugs. Dr. Cyrus
Thomas, in Bulletin No. 5 of the United States Entomological Com-
mission, 1879, page 24, stated that while Doctor Shimer’s plague
among chinch bugs was somewhat extraordinary, yet it was in accord-
anee with facts that he had himself ascertained in reference to other
insects, and in proof he cited a similar wholesale destruction of flies in
southwestern Virginia and eastern Tennessee in the year 1849, and
also a similar epidemic among grasshoppers in western Minnesota,
Dakota, and northern Iowa in 1872. This position of Doctor Thomas
in support of Doctor Shimer may be regarded as a second step in our
advance in a knowledge of the influence of meteorological conditions
on the chinch bug. It paved the way for further research in this
direction.

FUNGOUS ENEMIES OF THE CHINCH BUG DETERMINED.

While the subject of epidemic and contagious diseases of insects
was discussed to a greater or less extent among scientific men, there
was a decided lack of actual experimentation, and none at all with
‘the fungous parasites of the chinch bug until 1882 and 1883, when
Prof. S. A. Forbes began what ultimately proved to be a long series
of studies of the chinch bug and its natural enemies. At this time,
1882, Professor Forbes was more especially interested in the bacterial
diseases of the chinch bug, and though he found, at Jacksonville, I1.,
many specimens of dead chinch bugs embedded in a dense mat of
white fungous threads, which sometimes almost hid the body and
reminded him of the fatal disease previously reported by Doctor
Shimer, yet except to secure from Prof. T. J. Burrill a determination
of this fungus as belonging to the Entomophthora no progress was:
made in the study of this particular phase of the chinch-bug problem.?

In July, 1887, Professor Forbes found attacking the chinch bug in
Clinton County, IL, a second fungus, which he determined as be-
longing to the genus Botrytis, but this conclusion has since been
revised and the species is now known as Sporotrichum globuliferum
Speg. This discovery of a second species of entomogenous fungi and
its separation from the Entomophthora comprises what may be justly
termed a third step in the advancement of our knowledge of this prob-
lem. Professor Forbes, however, seems to have still been too deeply
interested in his bacterial studies to pay any special attention to the
other phases of his problem, further than to record the occurrence of
his new Botrytis in various localities in Illinois, and in one instance
on a beetle, Parandra brunnea (observed by Mr. John Marten, at
Champaign), and, similarly, to note the occurrence of the still spe-
cifically undetermined Entomophthora.?

a Twelfth Report of the State Entomologist of Illinois, pp. 47-51, 1882.
> Sixteenth Report of the State Entomologist of Illinois, pp. 46-49, 1888.

PARASITIC FUNGI. 47

The scene of action now changes from Ilhnois to Kansas, and to
Prof. F. H. Snow belongs the credit of first applying the knowledge

that had been gained up to that time (1889) by confining supposed
healthy chinch bugs with others affected by either one or the other of
the fungi, or possibly both Entomophthora and Sporotrichum, and
using the bugs thus infected for the propagation, in the field, of the
disease from which they had died.

As early as 1887-88 Professor Snow expressed, in the Sixth Bien-
nial Report of the Kansas State Board of Agriculture, the opinion
that “in the warfare of man against his insect foes a most valuable
ally will be found in the bacterial and fungoid diseases which may be
artificially introduced when nature fails to come to our aid,” an
opinion at that time largely based upon the investigations of Pro-
fessor Forbes and his own observations of the chinch bug in Kansas,
thus paving the way for the experiments of 1889. Professor Snow
had now obtained a specific determination of the fungous disease as
(Empusa) Entomophthora aphidis Hoffman, although there is some
ground for the suspicion that Sporotrichum globuliferum was also
present.

Entomophthora aphidis was already known to affect Hemiptera in
Germany and the United States. Dr. Roland Thaxter @ states that, as
early as 1886, his attention had been called to the attacks of this fun-
gus on aphides in the greenhouses at Cambridge, Mass., where it acted
as a decided check, and later, in 1887, Dr. L. O. Howard had called his
attention to great quantities of aphides dying with the same disease
on clover near the Agricultural Department buildings in Washington,

DEC.

FIELD AND LABORATORY EXPERIMENTS IN INDIANA.

On July 20, 1889, the writer, at that time a special agent of the
Division of Entomology of the United States Department of Agri-
culture, stationed at Lafayette, Ind., received, through the kindness
of Professor Snow, enough material with which to make some experi-
ments, the chinch bug being at that time very abundant at Lafayette,
and an exceptionally good opportunity thus being offered for ex-
perimentation. The results of these experiments were published in
detail in Bulletin 22 (old series), United States Department of Agri-
culture, Division of Entomology (pp. 55-63), but as this was the
first series of experiments carried out with a view of testing with
exactness the precise effects of varying degrees of temperature and
atmospheric moisture on the growth of the Entomophthora, and care-
fully following out the progress of the disease under varying meteor-

4 Memoirs Boston Society of Natural History, Vol. IV, p. 176.

THE CHINCH BUG.

48

ological conditions, the matter 1s here republished in full, the bulletin
in which it was originally included being now out of print.

These diseased bugs were placed under glass with living ones from the fields,
the latter being provided with food and kept thus confined for fifty-three hours,
when the major portion of them were placed on several hills of corn seriously
infested by bugs, the remainder with the dried remains received from Professor
Snow being scattered about over a small area of young wheat sown for experi-
ment and also swarming with young chinch bugs. The hills of corn on which
the bugs had been placed were isolated from others, equally badly infested, by
narrow frames of boards placed on the ground and the upper edges covered with
tar. This last precaution was taken in order to prevent communication with
other hills, intended as checks on those used directly in the experiment. The
area of young wheat over which infested bugs had been placed was not inclosed,
but its limits carefully marked. Five days after, July 27, a single bug was
found on one of the isolated hills of corn which had very evidently died from the
effects of Hntomophthora, and by the 80th enough others were found to show
that the fungus had fully established itself and the barriers about the isolated
hills were removed. On August 2 dead bugs covered with Entomophthora were
found in considerable numbers about hills of corn 25 feet from where the orig-
inal colonies had been placed and also throughout and even 55 feet beyond the
area of young wheat over which dead and affected bugs had been distributed.
Daily observations were now made, but the progress of the disease seemed to
come to a standstill. From the 5th of August up to the 9th it was almost im-
possible to get sufficient material outside to enable me to carry on laboratory
experiments. August 15 the spread of Entomophthora appeared to have taken
on new life, and diseased bugs were becoming much more numerous. August 15
found diseased bugs 172 feet from any place where they had been previously
observed. August 20 diseased bugs were very abundant over all of the area
where disease had been distributed, and two days later examples were found
a quarter of a mile from the starting point of the disease. Immediately after
this, however, another halt was observed, both in the intensity of attack and
rapidity with which it spread, due either to the dry weather or to the fact that
the bugs had now all reached the adult stage and had become diffused over the
country, no longer congregating together. From either one or the other, or
both of these causes, I lost track of the Entomophthora and was not able to again
find it in the fields. It seems proper to state here that chinch bugs were not
at any time excessively abundant. The greatest numbers were in the exact
localities where the disease was first distributed, the congregating at these
places being brought about by the close proximity to’a large number of small
experimental plats of wheat, and when this was haryested the bugs collected en
masse on the corn and young wheat. In connection with these facts, it is also
interesting to note that from July 15 to August 31 there were ten days on which
rain fell. The dates of these rains and the amount of precipitation is given

below:

ome | Precipi- Precipi-

Date. tation. Date. tation.

Inches. Inches.
SY: WAT ace 2 Ske Oe eee eee (OO) |i UID Peace jot o tb oonssemeeserineasfsscos 0.78
LQ 28 hsrehienon, cae eee een 1, 25 BOL h assess ecis Jae oe ee oe eee . 50
DO so Bic AeA oteja< iia ee Oe yA) aN PANT Oey O mee ee eeepc ein einl<iniaiaicio telaie = laleeeeeee 3.36
A EE Ee gee ee oS . 04 IS. Soa See eee sn eens .15
DG eae dets Hoe nies Ue 2a Cee ee 218 VA sdaplegese thm dewsskt weincmssee meee . 02

* PARASITIC FUNGI. 49

With a view of learning whether or not there was any difference as regards
susceptibility to the attack of Entomophthora between bugs in different stages
of development, a series of experiments was begun, as follows:

Young plants of Setaria glauca were transplanted to a box, and upon each
plant was placed a dead bug covered with the fungus, and also healthy larve;
larvee just on the point of pupation ; pupze just prior to reaching the adult stage,
and fully developed adults, each stage being placed on separate plants, and each
covered with a small inverted glass vial designated by lettering. As checks,
another series was prepared, like the first in every particular. The soil in
‘the box was kept well moistened, and the plants remained fresh. This experi-
ment was made on August 2, about the time when the attack outside began to
diminish in intensity. The following are the results of examinations on the
dates indicated, the original experiments being indicated by capitals and the
checks by small letters, thus—A—a, adult; B—b, young larvee; C-c, older larve;
D-d, pupe.

Date. | Ak | a. | B. b. Cc. | ee D. | d.

| | | |
Aur. 5 Healthy --| Healthy ..| Healthy ..; Healthy ..| 1 dead....| Healthy .| 1 dead...) 1 dead.
Au 6} 1 dead ..../ 1 dead....| Healthy ..| Healthy ..| 1dead....| Healthy .| 3 dead...) 1 dead.
AMP ey | ALudeat—. io Glad on jordead:... | dead=._-l\3 dead!..-.| 1 dead =5_)- os... on2- 5 dead.
Aug. 16 All dead..} All Ce ge All dead. .| All dead..| Alldead..| All dead.| All dead.| All dead.

On the same day this experiment was begun a second was also commenced.
like the first in every particular, except that the healthy bugs used in experi-
mentation were exposed to fungus-infected individuals for only five he irs and
then placed under their respective glasses. As a result, on August 15, thirteen
days after, none had died, thus strongly indicating that the Entomophthora did
not exist generally in the fields, and that it could not be communicated during
a period of five hours’ exposure.

On August 7 a large number of healthy bugs were placed under glass, with a
number which had recently died from Entomophthora, the moisture in the vessel
being absorbed by calcium chloride. A check experiment was also commenced,
where the material and the conditions were the same, except the humidity of the
atmosphere, care being taken to have the latter as nearly saturated with mois-
ture as possible. August 10 the original experiment was divided and a portion
of the healthy bugs removed and placed in a damp environment, the remainder
being kept under the original dry conditions. The results on August 22 were
as follows: In the original experiment, where the healthy bugs had been con-
tinually in dry quarters, not a single bug had died from Entomophthora. Not
only this, but none of those which had been removed after three days and placed
in dry quarters had died, showing that the disease was not contracted and did
not develop in healthy bugs, though kept exposed in a dry atmosphere for fifteen
days, nor could it be originated by placing in a damp atmosphere for twelve days
bugs which had been exposed to contagion for three days in dry quarters. The
results with the check experiment were quite different. Within five days after
being confined with the Entomophthora the healthy bugs began to die from
effects of the disease, and in three days more every one had died from the same
cause, their bodies being covered with spores.

Still another experiment was tried, which consisted in confining a large num-
ber of healthy bugs with others diseased in a damp environment, and when the
fungus had destroyed a portion the remainder were divided and a part removed
to dry quarters. The result was that while those left in damp confinement con-

26608—No. 69—07 m——4

50 THE CHINCH BUG.

tinued to die, none of those inclosed in dry environment were destroyed. As the
fungus had by this time become distributed over the experiment farm so that I
could not tell with certainty whether material from the fields was in a perfectly
healthy condition or not, no further experiments were made in this direction.
From the foregoing it will be observed that the essential element in all of
these experiments was an abundance of moisture, without which the HEnto-
mophthora could neither become established nor flourish after it had gained
a footing. Again, the extent to which the disease will prove contagious will
depend upon the number of bugs. Without great numbers massed together

comparatively few would contract the disease. To sum up the matter, there is °

little hope for relief to the farmer from the influence of Entomophthora, except
when chinch bugs are abundant and massed together in great numbers, and
during a period of wet weather. I have succeeded in getting the fungus estah-
lished at two widely located points in Indiana, and do not consider it at all
difficult to introduce in localities where chinch bugs are abundant, provided
the weather is favorable. But if it is ever utilized by the farmer, which seems
to me to be at present 2 matter of considerable doubt, it will only be after
the pest has become very abundant, during the time between the first larval
and adult stages and in a wet time. After the Entomophthora has been intro-
duced into a certain field it will become diffused only in proportion as the
bugs travel about and healthy bugs come in contact with spores from. those
which have died from the disease. This will not be very great until the pupal
stage is reached.

The larve of chinch bugs seem to in some way understand that while molting
they will be well-nigh helpless, and hence hide themselves away in vast num-
bers in secluded places. Under such conditions the spores thrown from dis-
eased bugs would reach a larger number of their fellows. I have found adults
but recently molted affected by the Entomophthora. After the bugs acquire
wings and scatter themselves over the country, the liability to contagion will be
again reduced, unless in case of very severe invasions, where, from force of
numbers, congregating on or about food plants becomes a necessity. Hence
the introduction of the fungus among lary will at first proceed but slowly,
and only in extreme cases and under favorable conditions can it be expected
to proceed much more rapidly among adult bugs. In short, the only way that
this fungoid disease seems capable of being employed in agriculture is by
the establishment of some central propagating station to which farmers can
apply and receive an abundant supply of infected bugs on short notice. By
this means they could take advantage of a rainy period of a week or ten days,
and, if they can contrive by sowing plats of millet and Hungarian to mass
the bugs in certain localities about their fields, they might accomplish some-
thing toward warding off an invasion. But.the possibility of overcoming an
invasion after it is fully under way, as is almost sure to be the case during a
dry season, it must be confessed is not very encouraging. My failure after
repeated experiments to produce this Entomophthora in the vicinity of Lafay-
ette without the importation of germs is decidedly against the theory that
might be advanced that the northeastern portion of the State was kept free of
destructive invasions by reason of this disease brought about by wet weather.
There is as yet no reason to believe that the disease has ever existed in that
section of the State.

The fungus entering into these experiments was determined as an
Entomophthora by Dr. J. C. Arthur, and the probability is that it
was LZ’. aphidis, though it is possible that Sporotrichum was also
present and remained unobserved.

i PARASITIC FUNGI. 51
FIRST FIELD APPLICATIONS OF FUNGOUS ENEMIES OF THE CHINCH BUG.

As has been stated, the credit for first confining healthy chinch bugs
with those diseased and utilizing the individuals thus infected by
transporting them to sections of the country supposedly free from the
disease in order to create new areas of infection, belongs to Prof.
F. H. Snow. During October, 1888, the year prior to that during
which Professor Snow began his experiments, Prof. Otto Lugger,
of Minnesota, collected a quantity of diseased chinch bugs at the
experiment station at St. Anthony Park and distributed them to
eighteen different localities in the southern part of the State where
the pest was known to occur in destructive abundance. The diseased
material was sent out in tin boxes by mail, and the contents of the
boxes, on arrival at their destination, were simply thrown in any
field where there was an abundance of chinch bugs. Later in the
season the condition of affairs where these distributions had been
made was such that “ careful search in the majority of places failed
to produce a single living specimen, while the traces of the disease
were found everywhere.” With a spirit of caution and exactness
in every way most commendable on the part of Professor Lugger,
he says: “The disease spread so rapidly that even corn growing
near wheat fields crowded with chinch bugs was entirely protected,
and no bugs had entered it in all the places visited by myself. But
the writer is by no means satisfied that the disease was really intro-
duced in this manner. Is it not possible that the disease was already
there, unknown to anyone, and that the writer had simply reintro-
duced its germs? The reason for this belief is based upon the fact
that too large an area was infested by the disease—too large to be
readily accounted for by the short time in which the atmospheric
conditions were apparently in its favor.”

In this case Professor Lugger states that both Entomophthora and
Sporotrichum were present and the latter was sent by him to Pro-
fessor Forbes, so there is the same confusion of the two fungi in this
case that existed in the writer’s experiments-in Indiana, except that in
the one case it was certain that Entomophthora was present, while in
the other it was the Sporotrichum.

THE WORK OF PROFESSOR SNOW IN KANSAS.

Although Professor Snow had the experience and observations of
Shimer, Forbes, and Lugger to aid him in his first efforts to apply
the knowledge gained by these gentlemen, yet it must be said that it
has been largely due to his untiring energy and perseverance that the
use of these fungi has reached the present state of importance. It
will hardly be saying too much if we state that his persistent un-

a University of Minnesota Experiment Sta., Bul. 4, Oct., 1888, pp. 40-41.

52 THE CHINCH BUG.

daunted labors, in the face of much skepticism and opposition, has
won for him the admiration of his fellow-workers, even among those
who were long in extreme doubt as to the success of his labor. He has
done more than any other one person to call attention to the possi-
bilities of practical benefits to be derived by farmers themselves; has
done more to advertise the merits of these fungous diseases among
the masses than anyone else, and, in fact, has made the “ chinch-bug
fungus ” almost a household word over the entire United States.

It is therefore all the more to be lamented that he should have ac-
cepted and published in his several reports the unsubstantiated state-
ments of farmers whose testimony on a matter of this nature is, as
every entomologist knows, absolutely worthless unless accompanied
by specimens. His own personal experience in this direction and in
several States had long ago led the writer to disregard all reports
relating to the efficiency or inefficiency of these fungous diseases
among chinch bugs, when such came from the ordinary farmer with-
out being accompanied by specimens for examination. The cast
pupal skins of the chinch bug pass with nonentomologists very well
for dead bugs, and if the former have been attacked by the ordinary
white molds the deception, except to the eye of an expert, is com-
plete.

There is probably not an entomologist who has distributed these
fungous diseases among farmers who has not found just such con-
ditions as did Professor Lugger in Minnesota, where it was impos-
sible to determine whether these diseases had been introduced arti-
ficially or whether they were already present and had been over-
looked. In the writer’s experience, while receiving chinch bugs from
different parts of Ohio to be infected with the disease, consignments
have come to him with the insects dying and others dead and covered
with Sporotrichum, showing that this was already present and that
the very utmost that we could expect to accomplish would be to aid in
locally spreading the contagion. Besides this, the writer has sent
material to farmers sufficient to start the fungus in their fields.
knowing perfectly well that it would be a considerable time before
actual benefits could by any possibility be expected to materialize,
and within a week received the astonishing information that the fun-
gus was so perfectly successful that the bugs all disappeared within a
few days after the application of the disease. There is little doubt
that the distribution of upward of 7,000 boxes of these fungi to
the farmers of Kansas has accomplished a vast amount of good, but
beyond this it is impossible to go. Of Professor Snow’s laboratory
work or the labors of himself and assistants in the fields no criticisms
can be made, and there will be occasion to quote from these in future
pages of this bulletin.

‘ PARASITIC FUNGI. 53

Sporotrichum globuliferum, or at any rate the fungus which is now

passing under that name, was first found by Professor Forbes to

“infest the chinch bug in Illinois in 1887, and its destructive effects
observed in the fields in the autumn of 1888.

Since the last-mentioned date the writer distributed upward of
3,000 packages of this fungus to the farmers of Ohio during the out-
break of the chinch bug in the State in 1895, 1896, and 1897,and knows
from personal observation and study that it is under certain favor-
able conditions a deadly foe of this species, that its use under these
conditions is practicable, and that if its application can be made simul-
taneously with the commencement of the breeding season it will prove
effectual. This statement is made for the reason that as late as 1895
Dr. M. C. Cook, in his popular work on entomogenous fungi, “ Vege-
table Wasps and Plant Worms” (p. 120), states that “no species of
this genus is known to have occurred on living matter, as they are
saprophytes pure and simple, and then, probably, only as the stroma
or conidia of some fungus of higher organization, possibly the Sphe-
riacel.” This statement was made in discussing S. denswm, but on
the following page (121), after dealing with S. globuliferum, he ap-
pends the following: “ The remarks made under the previous species
are applicable to this, which is not entitled to rank as a parasite, but
rather as an accidental development upon one out of many forms of
decaying animal matter.”

OTHER INSECTS ATTACKED BY SPOROTRICHUM GLOBULIFERUM.

Spegazzini“ described the species from Argentina as occurring on
the dead bodies of beetles, notably Monocrepidius and Vaupactus
wanthographus Germ. Besides Parandra brunnea Fab., Professor
Forbes has recorded this fungus on Lachnosterna and a number of
other Coleoptera, and also on lepidopterous larvee, as well as on the
young of other insects, while the writer has infected, artificially,
Epicauta pennsylvanica De G. and witnessed an instance of accidental
infection of Megilla maculata De G., but failed*to infect the harle-
quin cabbage bug ((/urgantia histrionica Hahn) even when these
were placed among dead and dying chinch bugs in the breeding cages.
In both cases these beetles were almost entirely covered by the fungus
after having to all appearances died from its effects. With respect
to this matter one point is clear, either the determination of this
fungus is incorrect or else Doctor Cook has made a very serious mis-
statement, which ought to be corrected. It is but just to state, how-
ever, that Professor Forbes, in his eighth report (p. 23), calls atten-
tion to the fact that it is closely allied to Botrytis, and would be
placed by some botanists under that genus now.

a Spegazzini, Fungi Argentini, ii, p. 42.

54 THE CHINCH BUG.

FIRST ARTIFICIAL CULTIVATIONS OF SPOROTRICHUM GLOBULIFERUM.

In April, 1891, Dr. Roland Thaxter succeeded in cultivating S.
globuliferum artificially on agar-agar, and a month later Professor
Forbes made similar cultures on the mixture of corn meal and beef
broth, this last being an exceedingly valuable discovery, as it revolu-
tionized our method of distributing the fungus by securing chinch
bugs to be kept for a time with those diseased and then sent out to be
scattered over the fields—a cumbersome method which was never satis-
factory. The writer’s own work in Ohio was based on material ob-
tained from Professor Forbes, and the first year he distributed in-
fected chinch bugs, but after that he used the artificial base of beef
broth and corn meal, finding the latter far more satisfactory to
handle, and, so far as could be determined, equally effective.

RESULTS OF FIELD APPLICATIONS IN OHIO.

In regard to the writer’s own experience, it is unnecessary to go
into details, except to state that, under the most favorable laboratory
conditions, he was able to kill apparently perfectly healthy chinch
bugs within three days after bringing them in contact with the Spo-
rotrichum. In the fields, during the season of 1895, though upward
of 750 packages of diseased bugs were sent out to farmers, and some
astonishing reports of results received therefrom, yet his own obser-
rations led him to believe that in many cases these were rather more
imaginary than real. Over the areas where local showers occurred
during the season of development of the first brood of young the
effect was much more satisfactory. But in many cases the request
for help came late, and soon after the fungus was applied the bugs
scattered out over the fields, disappearing to the eyes of the ordinary
farmer, who, of course, attributed all to the effect of the Spo-
rotrichum. In 1896, however, meteorological conditions changed,
and at last the writer had the good fortune to secure the very oppor-
tunity for which he had been waiting for years. All through April
and up to the 10th of May in southern Ohio there was little rain, and
even during the remainder of the latter month the hght rains hardly
sufficed to break the drought, so that there was a perfect breeding
season for the chinch bug during the forepart of the breeding period.
The result was that over some sections (see fig. 7) there were myriads
of young bugs. ‘Then the rains came on, and there were presented
the two essential requisites for success with the fungus, viz, chinch
bugs and wet weather.

Soon the demands for supplies of Sporotrichum began to pour in,
and 1,200 packages were distributed within a few weeks, instructions
being given to place the contents of the boxes where the chinch bugs

PARASITIC FUNGI. JO

were massed in greatest abundance, giving preference to the lower
and damper localities in the fields.

After the distribution had been finished, the sections where the out-
break of chinch bugs had been the most severe and where the larger
portion of the Sporotrichum had been distributed were visited. There
was certainly no mistaking the effect of the fungus. Going to the
place in a field (generally a wheat field) where the fungus had been
introduced, the track of the chinch bugs as they moved in any direc-
{ion was in many cases almost lterally paved with the dead bugs
more or less enveloped in their winding sheets of white. Along
ravines, dead furrows, or other depressions, the ground would be
nearly white, the dead diminishing in numbers as the higher grounds
were reached, though these were by no means free from corpses. In
one instance the bugs had left a field of wheat at harvest, the Spo-
rotrichum having been applied there before the movement began,
and entered an adjoining cornfield. The way was marked with
white, not only the surface of the ground, but on stirring up the
mellow soil of the edge of the cornfield it was found to be literally
full of dead chinch bugs to the depth of 2 or 3 inches, the white
fungus-covered bodies strongly contrasting with the black color of
the rich loam. Not only this, but under the sheaths of the leaves
and on the leaves themselves hundreds of dead were to be found on
the outer rows of corn, on the grass and weeds, and, indeed, almost
everywhere. Millions of chinch bugs were certainly destroyed in this
one field.

In other fields, where the number of bugs had been less, the dead
were less numerous, and then they were more apt to be scattered over
the leaves of the corn, as in such cases a diseased bug seems to be ani-
mated with a desire to crawl upward on any object which presents
itself, just as a larva of the clover-leaf weevil, Phytonomus punctatus
Fab., when attacked by E'ntomophthora spherosperma (Fres.) will
climb to the tip of a vertical blade of grass and coil itself around it,
and, holding it in the grasp of death, remain in that position so
strongly attached that the winds and rains fail to dislodge it until it
has become disintegrated. In other localities, where no Sporotrichum
had been distributed, the ravages had certainly been greater and the
writer failed to find any indication of the presence of the fungus.
So far as his observation extended, unless there were a sufficient
number of chinch bugs massed to become injurious, the fungus had
but little effect upon them. In other words, the massing appeared
to be an essential requisite. Whether this was sufficient of itself, or
whether the effect of massing was to reduce the vitality of the indi-
vidual bug, and thus render it more susceptible to the spores of the
fungus, it is impossible for the writer to decide; but he has long

56 THE CHINCH BUG.

suspected that the latter was the true solution of the problem. We
know that most domestic animals or fowls thrive best and are
the most vigorous when kept in small flocks, while among humans
the maximum of health and minimum of disease is obtained where
the individuals are scattered over a moderate area per capita and the
atmosphere is dry and pure; low, damp, and ill-ventilated quarters,
when overcrowded, being especially fatal, particularly to the young.
The individual in perfect health and vigor may in one sense be said
to be above and out of reach of disease, and before the two can be
brought together there must be some interacting element that will
bring the individual down to a point where he can be reached by the
disease; that is, the disease can rise only to a certain plane and the
victim needs to be first attacked by some element not necessarily
fatal in itself, but sufficiently depressing to bring the individual
down to where he can be grasped by the disease.

2

METEOROLOGICAL INFLUENCES FAVORING DEVELOPMENT OF FUNGOUS ENEMIES OF
THE CHINCH BUG.

When human beings are overcrowded and some disease is intro-
duced among them, everyone knows the effect of a low, damp locality
under a high temperature and with both air and water more or less
stagnant. Even the once healthy and vigorous are more or less
reduced and enervated by their environment, and thus brought within
the influence of the deadly disease. Again, if an individual is
stricken and forsakes his miasmatic surroundings for those more
salubrious, the disease may still overcome him, but seldom spreads
to others, except such as come in actual contact with either himself or
his belongings, while if not too much reduced before changing his
habitation the chances are much more favorable for his recovery.

It seems to the writer that in this matter of metorological conditions
and their relation to the effect of entomogenous fungi on the chinch
bug we are really dealing with the same problem in a different field.
The young chinch bug which has not yet come into possession of its
full measure of strength, and the spent females, which have lost theirs,
fall easiest as the prey to these fungi, while the fully developed bugs,
endowed with health and vigor, appear to be to some extent immune
to the attacks of these enemies, and if not massed in large bodies they
seem still more likely to escape destruction. In the timothy meadows
of northeastern Ohio an occasional dead adult has been found in late
autumn, but the fungus had certainly not claimed many victims,
though both the long and the short winged forms were present in con-
siderable abundance, clustered about the roots of grass. With Forbes
the writer believes that after becoming fully matured the chinch bug
‘is, largely at least, beyond the reach of Sporotrichum. What is the

A BACTERIAL ENEMY OF THE CHINCH BUG. 57

element that serves to enervate and reduce the older larvae and pupzx,
as well as many recently developed adults among them? Is there
nothing that, not of itself fatal, so acts upon the system of the bugs
that they are brought into a condition of susceptibility—a sort of
“ 9o-between,” so to speak, but which demands atmospheric moisture
before it will rise to an aggressive state?

A BACTERIAL ENEMY OF THE CHINCH BUG.

Forbes finds that the bacterium Bacillus insectorum Burrill is nor-
mal to the chinch bug and occurs always in the intestinal coeca, and
the writer has often wondered if this were not the very reducing ele-
ment. In a paper contributed to the “American Practitioner,” Sep-
tember, 1891, he describes the effect of this bacteria on the cceeca as
completely destroying the secreting epithelium, the cells of which
break down and disappear, leaving the delicate tubes filled with a
vast mass of microbes with some small intermixture of droplets of
fat and a little nondescript débris, the result of cellular decomposi-
tion. Now, it certainly seems to the writer that we may here have the
very enervating element necessary and which, in order to become sufli-
ciently aggressive to perform its functions perfectly, requires the
very conditions afforded by frequent showers, without which it is
comparatively helpless. We know very well that human beings are
far more susceptible to disease when weakened by fatigue, dissipa-
tion, or other forms of exhaustion, and under such conditions suc-
cumb to disease when they would otherwise enjoy immunity there-
from. We will not, however, follow this further, but submit it as a
problem well worthy of careful consideration and study. In his own
experiments with Sporotrichum globuliferum the writer has found
that under the most favorable conditions the fungus will attack even
the youngest larve, while Forbes states that it will also attack the
eggs, but in the fields it seemed generally most prevalent among the
more advanced larvee, pupe, and newly developed adults, though
much depends upon meteorological conditions and the abundance of
chinch bugs, as well as the time during the breeding season when
the fungus is doing its work. That is to say, there is a time at the
beginning of the breeding season when there are only adults and
young larve; later there will be larve of various ages, and, toward
the last, few if any of these, but all will be either pupe or adults.
For some reason it seemed more difficult to get the Sporotrichum to
work satisfactorily when the chinch bugs were beginning to breed
than later on, the last of June and the early part of July. These
facts are mentioned here to show that, judging by their effects, these
fungi hold a secondary place.

58 THE CHINCH BUG.

THE PRACTICAL UTILITY OF FUNGOUS: AND BACTERIAL ENEMIES IN
FIGHTING THE CHINCH BUG.

Regarding the practicability of utilizing these entomogenous fungi
in agriculture, there seems no reason to revise a statement made ten
years ago, viz, that this can be done only in cases of excessive abun-
dance and during wet weather, the basis for infection being provided
by some central propagating station from which farmers can receive
promptly an abundant supply. The writer believes that for himself
he could manage to get considerable benefit from their use in destroy-
ing chinch bugs, provided he were located within the area of the fre-
quent occurrence. This could be done only by watching the seasons
carefully, and in case there should occur two years in succession
wherein the breeding periods were covered by drought, then every
preventive measure known should be adopted, notably the burning
of leaves, dead grass, and other rubbish during winter or early spring,
followed up by sowing small plats of early millet, Hungarian grass,
or, better yet perhaps, spring wheat, in low damp places in the fields,
with a view of attracting the females or in fact massing the bugs,
and then freely applying the fungi in their midst. Whether the
average farmer, with his somewhat crude ideas of entomology, can do
this successfully or not is very uncertain. It is almost impossible to
determine even a few weeks in advance whether a season is to be fa-
vorable or unfavorable to the development of the chinch bug, which
would of itself cause occasional false alarm, and the precautionary
measures rendered entirely unnecessary by a few timely and drench-
ing rains just at the critical time. Before we can expect to be emi-
nently successful in this matter, not only the farmer but also the
entomologist and meteorologist have much to learn.

THE BOBWHITE OR QUAIL.

Chinch bugs have few important enemies among the birds of the
northern United States. To what extent the coast birds feed upon
them it is impossible to say, but inland the common quail or bobwhite
(Colinus virginianus) is the only species that can be said to devour
any considerable number. As this is one of our most highly prized
game birds, it is slaughtered annually in tremendous numbers.

The following list will show the degree of protection offered the
quail by legislative enactment in the States where the chinch bug is
the most destructive. The close season for quail in the several States,
during which killing is prohibited by law, is as follows: ¢

Maine, all the year.

New York, December 1 to November 1, except in some counties where it is

from December 1 to October 16. In Rensselaer County it is from December 1
to October 1 and in Richmond County all the year until 1908.

aFarmers’ Bulletin No. 265, pp. 18-25, 1906.

THE FROG. 59

Pennsylvania, December 1 to November 1.
Ohio, December 5 to November 15.
Indiana, January 1 to November 10.
Illinois, December 20 to November 10.
Minnesota, December 1 to October 1.
Iowa, December 15 to November 1.
Missouri, January 1 to November 1.
Nebraska, December 1 to November 15.
Kansas, December 15 to November 15, with some exceptions where the close

season extends to March 11, 1908.¢
Oklahoma, February 1 to October 15.
Texas, February 1 to November 1.

The breeding season from latitude 38° northward to Canada begins
in May, and continues through July and occasionally into September
or even October. A young bird just from the nest was taken in
Wayne County, Ohio, September 5, 1887.2 There are probably two,
and southward three, broods each season, and, while rather prolific, the
quails are kept well reduced in numbers, at times to the verge of exter-
mination over considerable sections of country. They are hunted
incessantly and slaughtered without consideration, except for gain.
Some also are killed by flying against electric wires, while entire
coveys are sometimes smothered or frozen under the snow. As a
result, the helpfulness of the quail against chinch bugs is greatly
diminished.

OTHER BIRD ENEMIES OF THE CHINCH BUG.

Among the other bird enemies of the chinch bug are the prairie
chicken, redwing blackbird, catbird, brown thrush, meadowlark,
house wren, tree swallow, and horned lark, but there is little doubt
that in seasons when the chinch bug is excessively abundant the com-
paratively few eaten by all of these birds is insufficient to reduce the
numbers of the pest to any extent.

THE FROG.

Dr. Cyrus Thomas quotes Professor Ross and others as stating that
the common frog is an enemy of the chinch bug. While this is prob-
ably true, it is nevertheless well known that comparatively few frogs
frequent grain fields as a rule, and thus the benefit derived from their
attacks is of too little importance to merit further notice.

@In Rawlins County, under restrictions, November i to January 1.
vA preliminary List of the Birds of Wayne County, Ohio, by Harry C. Ober-
holser. Bul. Ohio Agric. Exp. Sta., Tech. Ser., Vol. 1, No. 4, p. 270.

60 THE CHINCH BUG.

INVERTEBRATE ENEMIES OF THE CHINCH BUG.

Of the invertebrate enemies the same may be said as of the frog.
The writer has occasionally found a chinch bug containing a species
of Mermis, “ hair snake.” Also occasionally ants may be seen drag-
ging these bugs away, while lady-beetles have sometimes been found
to devour them, as recorded by Walsh and Forbes. Perhaps the
worst insect enemies of the chinch bug are to be
found among its comparatively near relatives,
the insidious flower bug, 7'7riphleps insidiosus
Say (Anthocoris pseudo-chinche of Fitch’s See-
ond Report) (fig. 15), and Milyas cinctus Fab.
(fig. 16), the latter being reported by Doctor
Thomas as the most efficient of the insect ene-
mies of this pest, while Doctor Riley found that
the former also attacked it. Professor Forbes
ascertained by examinations of the contents of
Fic. 15,—Triphleps insidio- the stomach of a ground beetle, Agonoderus

sus Fab. (From Riley.) allipes Fab., that one-fifth of the total food of
this species was composed of chinch bugs. Doctors Shimer and
Walsh both claim that lace-wing flies (CArysopa) destroy chinch
bugs, and they are doubtless correct. The writer has also very often
found dead chinch bugs entangled in spider webs, though whether
killed for food or by accident it has been impossible to determine.
It will be seen, however, that the combined influence of all of the
natural enemies of the chinch bug, parasitic fungi excepted, is far
too weak to offer any material protection to :
the agriculturist against this pernicious
enemy of his crops, with nothing to promise
an improved condition of affairs in this
direction in the future. There may some-
times appear hymenopterous parasites of
the eggs, but we have as yet no proof of the
existence of such in this country, and only
suspect the possibility of such a phenome- |
non because other allied species have similar _—_‘F"¢- 16.—Milyas cinetus Fab.
: ; é (From Riley.)

enemies, which destroy their eggs. In
short, the immunity of the chinch bug from attacks of other organ-
isms is so striking that it has attracted the attention of all entomolo-
gists who have made a study of the species, and all accept this as indi-
cating that it is an exotic, not originally belonging to our insect fauna.

REMEDIAL AND PREVENTIVE MEASURES.

The list will include all remedial and preventive measures that have
been found to possess the merit of reasonable efficiency and practica-
bility. These may not all prove applicable in all localities or under

REMEDIAL AND PREVENTIVE MEASURES. 61

every variety of circumstance, and the farmer will often have to fit
his protective measure to meet weather conditions, location of field
and its surroundings, as well as the thousand and one other variations
of a similar nature.

DESTRUCTION OF CHINCH BUGS WHILE IN HIBERNATION.

The first effort that may be made with a view to warding off an at-
tack of chinch bugs is to destroy them in their winter quarters. This
ean be accomplished by burning all dried grass, leaves, or other rub-
bish during winter or early spring. Forbes (First Report, p. 37) and
Marlatt (Insect Life, VII, p. 232) have cast some doubt upon the
statements to the effect that the chinch bug hibernates to any great
extent among dried grass, leaves, and rubbish, but the evidence is so
overwhelmingly in favor of the assertions of nearly every entomologist
who has studied the insect in its hibernation to the effect that it does
select such places in which to pass the winter that there is hardly any
use of raising the question at all. A good illustration of the fact that
large numbers of chinch bugs may be in hiding in such places and
escape detection is shown by the fact that a quantity of dried leaves
from about a vineyard located on a narrow neck of land about a
quarter of a mile from the Bay of Sandusky on the one side and about
14 miles from the shore of Lake Erie on the opposite side were col-
lected late in April and brought to our insectary and placed in a
breeding cage. At the time of collecting the leaves only an occasional
chinch bug was to be observed, but under the warm atmosphere of the
insectary they began to stir themselves, and soon demonstrated that
there had been a large number ensconced unseen among the dried and
curled dead grape leaves. So it is with the matted grass along road-
sides and fences, especially the Virginia worm rail-fence.

While it is not possible to find the hibernated chinch bugs by
searching, yet if pieces of boards are laid down on the grass in early
spring the bugs will collect on the under side and may be found
there, or they may be discovered by the method of collecting known
to entomologists as sifting. The burning of all such grass will de-
stroy thousands of bugs in their winter quarters; but sometimes the
matted bluegrass remains green in winter, or the weather is not
sufficiently dry to enable the farmer to burn over such places. In
such cases a flock of sheep, if given the freedom of the fields during
winter and spring, will eat off all living vegetation and trample the
ground with their small feet, so that not only is all covering for the
bugs removed, but also the bugs are trampled to death. The ease
with which the narrow strip of grass land along a post and wire
fence can be kept free of matted grass and leaves, as compared
with that along a hedge or rail fence, indicates that there may be

62 THE CHINCH BUG.

an entomological factor connected with the modern fence that has
been overlooked, giving it, in this respect, an advantage over the
more ancient form. Shocks of fodder corn left in the fields over
winter certainly afford protection for many chinch bugs, as also
will coarse stable manure spread on the fields before the chinch bugs
have selected their place of hibernation in the fall. In short, the first
protective measure to be carried out is a general cleaning up in winter
or early spring either by burning or pasturing or both.

SOWING DECOY PLATS OF ATTRACTIVE GRAINS OR GRASSES IN EARLY SPRING.

Judging from the manner in which the wintered-over adults are
attracted to hills of young corn, wheat fields, or plats of panic grass
and foxtail, it has always seemed to the writer practicable to take ad-
vantage of this habit and sow small patches of millet, Hungarian
grass, spring wheat, or even corn, early in the spring and thus bait
the adults as they come forth from their places of hibernation. Their
instincts will prompt them to seek out the places likely to afford the
most desirable food supply for their progeny, and if an artificial
supply can be offered them that will be more attractive than that
furnished by nature, the bugs will certainly not overlook the fact,
but will take advantage of it to congregate and deposit their eggs
there, whereupon eggs, young, and adults can, a little later, be sum-
marily dealt with by plowing both bugs and their food under and
harrowing and rolling the ground to keep the former from crawling
to the surface and escaping. The writer has thoroughly tested this
method in a case where the bugs, young and old, had taken posses-
sion of a plat of neglected ground overrun with panic grass (Panicum
crus-galli), which was mown and promptly removed and the ground
plowed, harrowed, and rolled before the bugs could escape, thus
burying them beneath several inches of soil out of which they were
unable to make their way, and as a consequence they were almost
totally annihilated, hardly 1 per cent making their escape to an
adjoining cornfield.

DIFFICULTY OF REACHING CHINCH BUGS IN MEADOWS.

There is, however, some doubt in regard to the practicability of
applying these measures in meadows. Meadow lands can be burned
over with perfect safety to either the grass or clover, if done while the
ground is frozen, but there is danger of injury if burned over in
spring, and it is somewhat doubtful if the hibernating chinch bugs
would be killed unless the surface of the ground was heated to a
degree that the grass and clover plants would hardly be able to
withstand.

REMEDIAL AND PREVENTIVE MEASURES. 63

Infested areas of meadow land could be plowed, it is true; but the
work would have to be done very carefully, else the grass and stubble
would be left to protrude above ground along each furrow and consti-
tute so many ladders by which the chinch bugs could easily crawl out
and make their escape. Where the ground will admit of subsoiling, or
where a “ jointer ” plow can be used, this latter difficulty can be easily
overcome. Usually, however, the chinch bug works too irregularly in
a field to permit of plowing under infested areas without disfiguring
it too much for practical purposes, especially in the case of meadows,
unless it be where the bugs have migrated en masse from an adjoining
field, when a narrow strip along the border can often be sacrificed to
good advantage. In many instances the heroic use of the plow in
turning under a few outer rows of corn would have saved as many
acres from destruction. In the majority of cases it is the fault of the
farmer himself that these measures are not effective, as he will sel-
dom take the trouble to burn the dead leaves, grass, and trash about
his premises at the proper time, and when there occurs an invasion
of chinch bugs, instead of resorting to heroic and energetic measures
to conquer them on-a small area he usually hesitates and delays in
order to determine whether or not the attack is to be a serious one,
and by the time he has decided which it is to be, the matter has gone
too far, and the chinch bugs have taken possession of his field. This
is especially true in the West, where the bugs breed exclusively in the
fields of wheat and remain unobserved until harvest, when they sud-
denly and without warning precipitate themselves upon the growing
corn in adjacent fields. In fighting the chinch bug promptness of
action is about as necessary as it is in fighting fire.

WATCHFULNESS NECESSARY DURING PROTRACTED PERIODS OF DROUGHT.

It has always appeared to the writer as though a little watchful-
ness on the part of farmers during periods of drought might enable
them to determine whether or not chinch bugs were present in any
considerable numbers in their fields, in time to interpose a strip of
millet between the wheat and corn, to be utilized later as previously
indicated. Instances have come under observation where, the wheat
fields being overgrown with panic grass and meadow foxtail, the bugs
transferred their attention to these as soon as the wheat was harvested,
and a prompt plowing of the ground would have placed the depre-
dators beyond the possibility of doing any serious injury. If the
weather at the time is hot and dry, a mower may be run over the
stubble fields or along the borders of them, cutting off grass, weeds,
and stubble, as the case may be, leaving them to dry in the hot sun,
when, in a few hours, they will burn sufficiently to roast all bugs

64 THE CHINCH BUG.

among them, and, while not destroying every individual, this will
reduce their numbers to such an extent that they will be unable to
work any serious injury.

In case the weather at the time should, on the contrary, be wet and
rainy, so that it is impossible to mow and burn, the prompt distribu-
tion of the fungus Sporotrichum will prove of immense value; for in
this case the more the bugs are massed over a limited area, the more
fatal will be the effects of the fungus, and especially will this prove
true if the land is low and inclined to be damp. This statement will
also hold good with reference to meadow lands during the breeding
season, though later the adults do not appear to succumb to the
effects of the fungus nearly as readily, and the writer has found the
fungus present in spring among masses of hibernating individuals,
with little indication of its contagious nature, only an occasional
individual being attacked.

UTILITY OF KEROSENE IN FIGHTING CHINCH BUGS.

In fighting the chinch bug there is at present no more useful sub-
stance than kerosene, either in the form of an emulsion or undiluted.
From its penetrating nature, prompt action, and fatal effects on the
chinch bug, even when applied as an emulsion, it becomes an inex-
pensive insecticide, while it has the further advantage of being an
article of universal use in every farmhouse, and is therefore always at
hand for immediate use. The emulsion has the further advantage of
being capable of sufficient reduction in strength to prove fatal to insect
hfe and yet not injure the vegetation upon which such may be depre-
dating. Diluted and ready for use, the emulsion is prepared as fol-
lows: Dissolve one-half pound of hard soap in 1 gallon of water,
preferably rain water, heated to the boiling point over-a brisk fire, and
pour this suds while still hot into 2 gallons of kerosene. Churn or
otherwise agitate this mixture for a few minutes until it becomes of a
cream-like consistency and, on cooling, will form a jelly-like mass
which adheres to the surface of glass without oiliness. For each gallon
of this emulsion use 15 gallons of water, mixing thoroughly. If
appled to growing corn it will be best to use the emulsion either dur-
ing the morning or evening, say before 8 a. m. or after 5 p. m., as at
those times it will be less likely to affect the plants than if applied
in the heat of the day.

Where an invasion of the chinch bug is in progress from a field of
wheat to an adjoining field of corn, as an illustration, the marginal
rows of corn can frequently be saved, even after the bugs have massed
upon the plants, by spraying or sprinkling them freely with kerosene
emulsion, being careful not to get much of it directly into the crown
of the plants and using a sufficient quantity so that the emulsion will

REMEDIAL AND PREVENTIVE MEASURES. 65

run down the outside and reach such bugs as are about the base of
the plants. This treatment will kill the bugs clustered upon the corn,
and in case of those on the way to the field, while it will not keep
them out, it will cause a halt in the invasion, and thus give the
farmer an opportunity to put other measures in operation, one of
which will include the use of kerosene in another manner. If a deep
furrow is plowed along the edge of the field, running the land-side
of the plow toward the field to be protected, the furrow will form a
temporary barrier to the incoming hordes.

UTILITY OF DEEPLY PLOWED FURROWS SUPPLEMENTED BY THE USE OF
KEROSENE EMULSION.

In dry weather the sides of this furrow can be made so steep and the
soil so finely pulverized that when the chinch bugs attempt to crawl
up out of the furrow they will continually roll back to the bottom,
where they can be sprinkled with either kerosene alone or with the
much less expensive emulsion, and killed. In case of showery weather,
which prevents the sides of the furrow from remaining loose and dry,
the bottom. can be cleared out with a shovel, making it more smooth
and the sides more perpendicular, thus rendering it so much easier to
follow along the bottom than to attempt to climb the sides. If holes
are dug across the bottom at distances of, say, 30 or 40 feet, the bugs
will fall into them and can be still more easily disposed of by the use
of kerosene. That both of these measures are thoroughly practicable
the writer has ample personal experience in evidence, and knows that
under most conditions that are likely to obtain, prompt and efficient
application is all that is necessary. During a few days this work
will demand the closest watchingand application, but fields of grain
can be protected thoroughly and effectually if these measures are
faithfully carried out, and the expense of time and money will be
found to be less than in almost any other plan that has been up to
this time discovered. In his own experience, in no case has a field
attacked by a migrating army of chinch bugs come under the writer’s
observation, but that might have been saved from very serious injury
by the prompt use of either of these measures, though under some
conditions the farmer might find it advantageous to apply some of
the other methods of protection here given.

THE SURFACE AND COAL-TAR METHOD.

The objections made by farmers to the use of most forms of these
barriers is that the finest pulverized soil soon becomes incrusted by
even the slightest rainfall and the bugs then pass over it without
difficulty, while barriers of boards are expensive.

26608—No, 69—07 m——5

66 THE CHINCH BUG.

It is feasible to eliminate both by simply smoothing off a path
along the margin of an infested field where such a one adjoins the one
to be protected. This can be done with a sharp hoe, and as the
margins of wheat fields usually become compacted it is but little trou-
ble to thus clear off a path a foot or more in width, smooth as a floor,
with the surface almost as hard. - In the midst of this path post holes
are sunk as in the bottoms of furrows, and a train of coal tar is run
between them, being so arranged that it will reach the post hole near
the inner side opposite the field from which the bugs are migrating.
In this way as the bugs reach the train of coal tar they will follow
along until they reach the post hole, while those meeting with the
post holes will usually divide and, following around it, join with the
flow of bugs moving along the train of coal tar. The result is that
they become congested in the acute angle where the coal-tar train is
intercepted by the post holes. Those in the apex of this angle can
not turn back, and thus are continually pushed into the post holes by
those behind. As the bugs, varying from the red larve to the
almost black pupe, mass along the line of coal tar they have much the
appearance of a reddish-brown stream running into the holes. From
these holes there is no escape and here the bugs can be readily killed
by sprinkling with kerosene. The shghtest train of coal tar is suffi-
cient to obstruct the passage of the bugs, and light rains will not
affect its efficiency. In dry weather these trains of coal tar soon be-
come covered over with dust and must be renewed; but in showery
weather there is no dust, and if the coal tar is renewed daily or, at
most, twice each day it will accomplish its work and nothing further
will be needed than to kill the bugs that have collected in the post
holes. This measure is inexpensive and can be promptly put into
operation if the coal tar is at hand. The writer has been able in this
way to effectively protect a field of corn surrounded on two sides by
a wheat field literally overrun with chinch bugs at harvest and during
a time when light showers were occurring, frequently several times
each day.

THE RIDGE AND COAL-TAR METHOD.

Differing quite materially from the preceding are the various com-
binations of coal tar and ridges of earth, smoothed and packed along
the apex, or, instead of the ridge of earth, 6-inch boards, such as
are ordinarily used for fencing, placed on edge and the upper edge
coated with tar. Forbes has reported excellent results from the ap-
plication of a line of coal tar put directly upon the bare ground where
the surface. has been rendered compact by a recent fall of rain. Even
in this series of protective measures kerosene can be used to great
advantage. In the experiment recorded by Professor Forbes the coal
tar was put upon the ground between a wheat field and a cornfield

REMEDIAL AND PREVENTIVE MEASURES. 67

from an ordinary garden sprinkling pot from which the sprinkler
had been removed and the orifice of the spout reduced in size with a
plug of wood until the tar came out in a stream about the size of the
little finger and made a line on the surface of the ground about three-
fourths of an inch in width. Post holes were sunk along the line
from 10 to 20 feet apart on the side next to the wheat field, thus prac-
tically completing the barrier, and the chinch bugs being unable to
cross the line of tar accumulated in the post holes in vast numbers,
where they were killed; and those bugs that had already entered the
cornfield before the barrier was constructed were prevented from
spreading farther by tar lines between the rows of corn, the infested
corn itself being cleared of bugs by the application of kerosene emul-
sion. The same writer states“ that several farmers in Vermilion
County, Ill., prepared for the coal-tar line by hitching a team to a
heavy plank and running this, weighted down with three or four
men, over the ground once or twice until a smooth, hard surface had
thus been made to receive the tar. If the barrier was to be made in
sod, a furrow was plowed and the bottom of this made smooth by
dragging the plank along the bottom. In both cases post holes were
sunk along the tar lines, and in these were placed cans or jars into
which the bugs fell in myriads and were destroyed.

On one farm of 250 acres a coal-tar line 90 rods in length was re-
newed once each day and killed about 8 gallons of chinch bugs. In
the case of another farmer there were 300 rods of tar lines with post
holes, cans, etc., which resulted in destroying about 10 bushels of
chinch bugs. <A 6-gallon jarful was destroyed in less than half a day
at one point on the line. In this last instance the lines of tar were
renewed three times a day, but even then less than a barrel of tar
was used. Still another farmer, with 120 rods of tar line, used about
a third of a barrel of tar and did not lose a hill of corn; he caught
chinch bugs by the bushel. In some of the cases cited the tar line
was run in a zigzag course, the post holes being situated at the angles,
and in others leader tar lines were run obliquely to the main tar line, .
one end terminating at the traphole, but both of these plans were
afterwards regarded as unnecessary, a single straight line being en-
tirely sufficient and less expensive. ‘The numerous cases where these
methods were put into execution with entire success and at small
expense is the best possible proof of their practical utility. If
a farmer is situated near town, where refuse tin cans are dumped in
any locality where they can be got out of the way, he can select the
larger of these, set them in the post holes and partly fill them with
kerosene and water. The water being heavier than the kerosene will
sink to the bottom, leaving a stratum of kerosene on the surface.

_@ Twentieth Report of State Entomologist of Illinois, p. 39, 1898.

68 THE CHINCH BUG.

The chinch bugs falling into this will be forced down by the weight
of those coming after, and thus all will be passed through the kero-
sene into the water below. This will obviate the necessity of fre-
quently emptying the cans or treating their contents. It may also
be stated that where the post holes are quite deep and enlarged at
the bottom the bugs falling into them will perish without further
attention.

OTHER BARRIER METHODS.

Professor Snow, working in Kansas, followed a somewhat different
method and one that, under certain conditions, might be found supe-
rior to that used by Professor Forbes, or the furrow and kerosene
method applied by the writer in Ohio. This modification consists in
throwing up a double furrow, known among farmers as “ back fur-
rowing,” and thus forming a ridge, the top of which is smoothed
and packed with a drag hawing a concave bottom of the form of the
ridge to be made. If the bottom of this drag is covered with zinc, it
will be found to keep bright and polished and by this means make
a smoother ridge. The substances used were coal tar as it came
from the gas works and crude petroleum as taken from the oil wells.
The former is the more easily obtained, except in certain localities,
and will probably be found the more practical, as it stands on the
surface better and is not so readily washed away by rains. Both of
these substances are, however, offensive to the bugs, and they will
seldom attempt to cross them or even come close enough to touch
them, but on approaching will turn and run along the ridge in the
evident hope of finding a gap through which they can pass. Post
holes were dug on the outside of the line, but close up to it, so that
the bugs in passing along beside the tar line would crowd each other
into them. Professor Snow suggests that it will be better to con-
struct this barrier several weeks prior to its being needed, as then the
tar line has but to be run along the ridge and the post holes dug, when
the whole system is complete and the chinch bugs can be thus shut
out from the first.

With these barriers of either ridge or furrow and the use of coal
tar or crude petroleum, supplemented by kerosene emulsion, a very
large percentage of the injury from chinch bugs may be prevented,
and, in fact, with a reasonable degree of watchfulness and prompt ac-
tion, all injury from migrating hordes may be prevented. The use of
tarred boards set on edge or slightly reclining might, under some
circumstances, take the place of the ridge or furrow, but these cases
will be exceptional, and the use of kerosene emulsion will probably

a Fifth annual Report of the Director of the Experimental Station of the Uni-
versity of Kansas, for the year 1895 (1896), pp. 45-47,

REMEDIAL AND PREVENTIVE MEASURES. 69

be found equally practicable here, as also will the post holes for col-
lecting the chinch bugs. This method is merely cited in order to call
attention to its possible use where the others are found impracticable.
The plowing of furrows has been in vogue since the first writings of
Le Baron and the second report of Doctor Fitch, and may be utilized
in other ways than those previously mentioned. A heavy log
dragged back and forth in this furrow will pulverize the soil in dry
weather, and Forbes has recorded the fact that where this has a
temperature of 110° to 116° F. it is fatal to the young bugs that fall
into the furrow, even if they are not killed by the log. As 120° is
not uncommon in an exposed furrow on a hot summer day, it will be
observed that there may be cases where this method will be found
very serviceable, and especially is this likely to prove true in a sandy
soil with a southern exposure. In sections of the country where
irrigation is practiced these furrows may be flooded and in this way
rendered still more effective without the expenditure of either time
or money to keep them in constant repair. Doctor Riley long ago
laid considerable stress on this measure, believing it of much value,
especially in the arid regions of the far West. The same writer ad-
vised the flooding of infested fields, wherever it could be done, for ¢
day or so occasionally during the month of May. It is hardly prob-
able, however, that this will often be found feasible except in rice
fields, where it 1s sometimes practiced.

NECESSITY FOR PREVENTING CHINCH BUGS FROM BECOMING ESTABLISHED
IN FIELDS OF WHEAT AND GRASS.

In the foregoing it will be observed that prevention of migration
has been the chief end in view either by destroying the chinch bugs
in their hibernating quarters, and thus preventing the spring migra-
tion to the breeding places, or by various traps and obstructions to
prevent them from migrating from such places to others not already
infested. The great problem remaining to be solved is to prevent their
breeding in wheat fields at all. As has been shown, it is absolutely
impossible, with our present inability to forecast the weather months
in advance, to be able to foretell whether or not an outbreak of chinch
bugs is likely to take place. There may be an abundance of bugs in
the fall—enough to cause an outbreak over a wide section of coun-
try—and these may winter over in sufficient numbers to cause some
injury in spring, yet a few timely, drenching rains will outbalance all
of these factors, and our wisest prognostications fail of proving true.
It is this very factor of uncertainty that renders unlikely the success-
ful carrying out, over any large area of country, of any protective
measures where, as in this case, the benefit to be derived will only be
realized nearly a year afterwards, if at all. The average farmer,

70 THE CHINCH BUG.

when smarting under a heavy loss, will often take such long-range
precautions as to sow belts of flax, hemp, clover, or buckwheat around
his wheat field once; but if the chinch bugs do not appear, and he sees
the useless investment of time, labor, and seed, he will be likely to con-
clude next year to take the risk and do nothing. For the present, then,
we have no method whereby we can prevent the chinch bugs from
taking up their abode in wheat fields or timothy meadows and
raising their enormous families there, except to destroy the adults
in their winter quarters.

The writer once tried to destroy the young in a wheat field by
spraying with kerosene emulsion the small areas of whitening grain
that indicated where the pests were massed in greatest abundance.
The result was unsatisfactory, and it is very doubtful if it is possible
to apply this measure with any degree of success, and we are forced
to the conclusion that, for the present at least, we shall be obliged
to rely upon the measures previously given. It therefore becomes
of the utmost importance to clean up the roadsides, and along fences
and patches of woodland, as well as any other places likely to afford
protection for the hibernating chinch bugs. There are of course
obstacles in the way of carrying out this plan generally over any
large area of country, and especially in sections where the rail fence
predominates. But as the country gets older it will be found that
it is not chinch bugs alone that seek these places in which to pass
the winter, but myriads of the other insect foes of the farmer as
well, and that careful attention to the condition of roadsides, lanes,
hedgerows, and waste places about the farms, during the season
when insects seek out these places wherein to pass the winter, will
pay well for the time expended in that direction. It may come
about that some phase of the street-cleaning reform may invade the
country, and it is certain that if such were to occur it would, in time,
save the country enough to go far toward reducing the expense of
securing good roads. In fact, the term “ good roads” ought to
include the proper care of the roadsides, as well as the grading and
macadamizing of the roadbed itself.

There are at present so-called weed laws in many States, and,
though more or less of a dead letter in most cases, these laws are
steps in the proper direction. The time when insect pests will be
looked upon in the eye of the law as so many public nuisances, and
the harboring of them a corresponding crime, may be a long way off,
but as it gradually draws nearer to us we shall come to learn that,
after all, it is the rational view to take and will go far toward solv-
ing not only the chinch bug problem, but many others of a similar
nature. So far as the chinch bug is concerned, when we burn over
the waste lands and accumulated rubbish about our farms in autumn
or winter, we are simply applying the same check that the dusky

REMEDIAL AND PREVENTIVE MEASURES. 71

savage did when he lighted the prairie fires, though unwittingly and
for an entirely different purpose. In the timothy meadows of the
northeastern portion of the country, where, for lack of wings fitted
for locomotion, the chinch bug does not so largely migrate to the
waste lands in autumn, the problem is somewhat different, and it will
require some careful experiments to determine the exact effects of
burning over the meadow lands in winter, both on the hibernating
chinch bugs and on the grass roots. There can be little doubt, how-
ever, that a rapid rotation of crops, so as not to allow the short-
winged form to become thoroughly established in a meadow, and the
burning over of waste places, thus destroying such rubbish and
débris as will serve to offer hibernating places for the long-winged
form, will go far toward settling the chinch bug problem in grass
lands.

As previously stated, the chief drawback in putting preventive
measures in force is in the difficulty of foretelling an invasion. In
northeastern Ohio in 1897 hundreds of acres of timothy meadow
were destroyed after the hay crop had been removed, but so late that
the farmers did not suspect the true condition of their meadows until
the spring of 1898, when the young grass failed to put forth and an
examination revealed the fact that the roots had been killed, the
abundance of chinch bugs pointing unerringly to the cause of the
trouble, though in many cases a heavy crop of hay had been removed
the previous year where now the ground was entirely bare. While
in the case just cited a previous knowledge of the presence of chinch
bugs in these meadows might not have enabled the owners to have
saved them in the fall of 1897, yet the fall plowing of the land, possi-
bly early enough to have sown the ground to fall wheat, would have
buried the majority of the bugs so deeply in the soil as to have killed
vast numbers of them and thus prevented their migrating to other
lands in the spring of 1898. A rotation of crops that would have in-
cluded grass for not to exceed two successive years, followed by wheat,
would have amounted to precisely the same remedial measure as the
one suggested.

A ease in northeastern Ohio has come to the writer’s notice where
an infested timothy meadow was plowed late in the fall of 1897.
Late in April of 1898 this ground was cultivated, rolled, and har-
rowed several times and most carefully and completely prepared for
corn, which was planted, but with the result that a portion of the field
was attacked and destroyed by chinch bugs, largely of the brachypter-
ous or short-winged form. An examination about June 10 revealed
the bugs in considerable numbers about the still remaining plants, but
scattered over the field were more or less numerous clumps of timothy,
in some cases apparently having been killed by the chinch bugs, while
in others these were literally swarming about the dying but still

2 THE CHINCH BUG.

green clumps of grass, thus showing that the former had either not
been buried by the plowing and cultivation of the ground, or else the
grass had not been thoroughly covered, and thus ladders had been left
whereby they were enabled to climb to the surface.

SUMMARY OF REMEDIAL AND PREVENTIVE MEASURES.

In summing up the matter of remedial and preventive measures
for the control of the chinch bug, it may be stated that the insects can
be destroyed in their places of hibernation by the use of fire. They
can, under favorable meteorological conditions, be destroyed in the
fields, if present in sufficient abundance during the breeding season,
by the use of the fungus Sporotrichum globuliferum, if promptly and
carefully applied. They can be destroyed while in the act of migrat-
ing from one field to another by tarred barriers or deep furrows sup-
plemented by post holes, and by being buried under the surface of the
ground with the plow and harrow; or the latter method can be ap-
pled after the bugs have been massed upon plats of some kind of
vegetation for which the bugs are known to have a special fondness,
which decoys should be so arranged as to either attract the females
and induce them to oviposit therein, or they should be arranged with
the idea of intercepting an invasion from wheat fields into cornfields.
When these decoys have been turned under with a plow and the sur-
face immediately smoothed and packed by harrow and roller, the bugs
will be destroyed. While in the cornfields they can be destroyed on
the plants by the application of kerosene emulsion. Without vigi-
lance and prompt action, however, only indifferent results are to be
expected from any of these measures.

PROBABLE ORIGIN AND DIFFUSION OF THE CHINCH BUG.

For the farmer engaged in attempts to check the ravages of the
insect in his fields the question of origin, or how it came to reach
him, will at the time have little interest for him. It will suffice that
it is present in overwhelming numbers, and what he will most desire
will be to learn how to rid his premises of its most unwelcome
presence in the most summary manner possible.

If, however, the farmer happens to be a thoughtful and observing
man he will sometimes wonder how it is that, except in Virginia and
the Carolinas, a person need not be very aged in order to remember
a time when the chinch bug was an unknown factor in his profession,
with a possible value far too small to merit consideration. If he
happens to reside in northeastern Ohio or in some portions of New
York, and has spent some time in I]linois, Iowa, Kansas, or Minne-
sota, he will probably marvel at the striking difference in appearance

PROBABLE ORIGIN AND DIFFUSION. 12

between many of the chinch bugs of his own locality and those found
in any of the last-mentioned States, and will probably be able to
satisfy himself of their identity only by the similarity of their vile
odor. Again, he will probably be equally at a loss to understand
why it is that his own timothy meadows are overrun by these pestifer-
ous insects and destroyed, while in other localities, perhaps less than
100 miles away, similar meadows are left untouched, the injury
there being confined to the wheat and corn fields.

If wondering leads to questioning, as it often does among the
constantly increasing number of educated and up-to-date farmers,
it will not satisfy him to receive an evasive or obscure reply to his
query as to why such differences exist, for if he can not get a clear
explanation he will want ideas, theories, or possibilities. He wants
the best explanation possible to give until some one finds out a better
one, realizing that had mankind been perfectly satisfied with the
knowledge that a stroke of lightning would split a tree or destroy
human life, and had stubbornly refused to listen to possibilities or
to anything but facts, we would not now be able to understand and
utilize electricity in the many ways that we do at the present time.
Such men understand perfectly that the solution of most problems
in natural science must of necessity commence with theories which
must be patiently tested and adopted or rejected as the results
demand, while the scientific man knows that the solution of one
problem often opens up the way for the solution of another, the
last not infrequently having an entirely different application from
the first.

The science of applied entomology is growing rapidly and becoming
both broader and deeper, and it is not enough simply to tell the hus-
bandman what an insect is and how to kill it. He must have some-
thing along with that information to set his own mind to thinking, to
work out problems or improve upon the solutions already given him,
otherwise it is much like giving money to a professional beggar. If
we can not give facts based upon demonstrations, then give the best
explanation possible, even though it be a theory which is only ex-
pected to stand until some one does better. It is for the thoughtful,
progressive farmer, as well as the student of geographical distribu-
tion, that this possible solution of the problem of the chinch bug has
been prepared, and while the full practical value of the ideas ad-
vanced has yet to be demonstrated, this of itself can not be urged as
sufficient grounds for not sending it forth for study and consideration.

Thanks to the careful observations of Professor Sajé, on the Euro-
pean species of chinch bug, Blissus doriw, it is now for the first time
possible to compare the habits of this species with our own.

74 THE CHINCH BUG.
INDICATIONS OF A PROBABLE DISTANT ORIGIN AND LATER DIFFUSION.

In the United States our chinch bug, Blissus leucopterus, has a
number of peculiar characteristics, which, while having an economic
interest, also point to a probable previous condition differing some-
what from the present, and not in all cases tending toward its present
numerical strength. On the other hand, we find that it is now fol-
lowing some probably ancient habits which do not appear to be of
any special benefit, but rather the reverse.

In the first place, over its area of greatest destruction, it appears to
prefer level tracts of country where the damp conditions consequent
upon frequent rainfalls remain the longest, and in the second place,
the period of spring oviposition is for the most part included within
that during which the spring rains of the United States usually
occur—that is to say, throughout the great grain belt, east of the
Rocky Mountains, April and May are not normally months of severe
drought, and it is during these two months that the larger portion
of the eggs are deposited. As in the reverse of this, however, the
period of fall oviposition, August and September, is far more likely
io be favored by a lack of precipitation. These conditions do not
always obtain, and it is because of the fluctuations that the insect
is able to reach its maximum in point of numbers.

Another factor which plays quite an important part in reducing the
number of adults maturing during unfavorable seasons may be found
in the almost universally gregarious habits of the young, thereby ren-
dering the ravages of fungous diseases the more universal and fatal.
In all of these peculiar characteristics as well as in some anatomical
features, it would seem as if we had a series of guide posts, so to
speak, which indicate more or less clearly the ancient home of the
species, and at least throw some light on its origin and diffusion.

JTNIQUE APPEARANCE AND GREGARIOUS HABIT.

Mr. Kk. A. Schwarz“ some time ago called attention to “ the unique
appearance of the full-grown chinch bug, with its white wings and
chalky-white pubescence,” which, he declared, “ forcibly indicates
that the insect is either a psammophilous or a maritime species,” and
expressed the opinion that its geographical distribution fully bears
out the theory that it belongs to the latter class. The same author
states that the species has the habit of clustering about the roots of
tufts of grass along the Atlantic coast, from Florida to Atlantic City,
N. J., and Mr. W. H. Harrington ® observed it to have the same habit
along the seashore at Sydney, Cape Breton, in 1884. The late Dr.

@Insect Life, Vol. VII, p. 420.
bCan. Ent., Vol. XXVI, p. 218.

Lad

PROBABLE ORIGIN AND DIFFUSION. rips

J.C. Neal, while at Stillwater, Okla., wrote me that he had observed
the species to have the same habit in that Territory, miles from any
human habitation. Dr. Asa Fitch* found them swarming amidst
extensive prairies in Illinois, in 1854, while more recently Mr. C. L.
Marlatt has witnessed the same phenomenon in Kansas.’ In short,
this gregarious habit seems to be most tenaciously adhered to wher-
ever these insects are found in any numbers. When migrating from
one field to another, after crossing a roadway or plowed field they
will at once flock together on a few plants along the margin of the,
to them, new field instead of scattering about, two or three to a plant.
It may also be added that Mr. Koebele found the species in large
numbers along the seashore not far from San Francisco, Cal., in the
first, second, and third stages of development, on a species of grass
growing along the coast.

It has not, so far as is known to the writer, been observed in similar
places along the shores of the Great Lakes, though search has been
made for it there, but it occurs in destructive abundance in timothy
meadows inland in northern and northeastern Ohio 25 to 75 miles dis-
tant, most generally clustering about the roots of grass—which, by the
way, is about the only vegetation attacked—as the species is described
as doing along the seacoasts. It may also be stated that it seems to
hibernate there precisely as observed by Mr. Marlatt in Kansas;
Doctor Neal in Oklahoma; Mr. Schwarz in Virgina in the vicinity
of Fortress Monroe, and as the earlier observations of Doctor Fitch
in Illinois would imply. Thus we find this habit of clustering upon
the plants attacked to be a constant one, and where the natural grass
vegetation has not been displaced by farm crops, thus leaving the
ground more or less bare during winter, the chinch bugs continue to
hibernate there. With these two characteristic habits generally fol-
lowed over the great area inhabited by the species in North America,
we may add a third possible factor in the problem of origin and
diffusion of the species which, though an anatomical dimorphism,
may be discussed as likely to throw considerable light upon the prob-
able ancient habitat of the insect.

OCCURRENCE OF THE LONG AND SHORT WINGED FORMS AND THEIR
DISTRIBUTION.

The occurrence of both the long and short winged forms, inter-.
mixed along our seacoasts and in the northeastern section of the
country, but not elsewhere, shows very plainly that this dimorphism
is not due to the temperature of any particular locality, but must
be regarded as having been brought about by disuse of the wings

« Second Report, Insects of New York, p. 283.
b Insect Life, Vol. VII, pp. 2382-254.

76 THE CHINCH BUG.

for a considerable period of time, thus indicating a seashore habit
on the one side, while the total lack of the short-winged form else-
where indicated otherwise. ; .

In a paper presented before the Entomological Society of Wash-
ington,* “On the insects found on Uniola paniculata in southeastern
Florida,” by Mr. EK. A. Schwarz, the author stated that Blissus leu-
copterus occurred in large numbers on the upper part of the plant,
the imagos and larger young among the ears and the smaller indi-
viduals between the upper blades. Mr. Schwarz attributes this habit
to the tough woody nature of the storm-beaten plant nearer the
ground, thereby driving the insects to the more tender though more
exposed portion of the plant. In connection with this statement the
writer tells us that the imsect occurs in that southern latitude only
in the short-winged form, and that in the examination of thousands
of specimens from that region he had never found a single long-
winged specimen. Under date of May 4, 1896, Mr. W. H. Harring-
ton wrote of this species as follows: “ In September, 1890, I found
it at Aulac, almost on the border between New Brunswick and
Nova Scotia. It seemed not uncommon and occurred under stones,
about the roots of grass, in a pasture adjoining the marsh where I
found Diabrotica longicornis, the pasture being on the upland
skirting the marsh. Both the long and short winged condition
occurred, asin Cape Breton.” Dr. A. S. Packard communicated
to Dr. J. A. Lintner the following extract from his diary: “ June 17,
1871, at Salem, Mass., chinch bugs with wing covers extending over
the basal third of the abdomen, seen in copula, end to end.¢ In the
serious outbreak of this insect in the timothy meadows of northern
New York, in 1882 and 1883, about 20 per cent of the bugs were of
this short-winged form.” ¢

Although Dr. Asa Fitch, as early as 1855, refers to this form along
with nine others, he does not give the source from which he obtained
specimens, but just previous to this he says (p. 287) that he had met
with but three specimens from his own State, and these were found
on willow in the spring of 1847.° Had any of these been of the short-
winged form he would have been very hkely to have mentioned the
fact. Mr. EK. P. Van Duzee? states that he had known of the occur-
rence of the species in western New York as early as 1874, and had
also found it at Ridgeway and Muskoka, Ontario. Ordinarily the
short-winged form predominates, but in hot, dry summers the chinch

4Proc. Ent. Soe. Washington, Vol. I, p. 104. Read Nov. 3, 1887.
» Canadian Entomologist, Vol. XIV, p. 218.

¢ Lintner’s Second Report, State Entomologist of New York, p. 164.
@d Second Report, State Entomologist of New York, p. 156.

e Second Report on Noxious Insects of New York, p. 291.

f Canadian Entomologist, Vol. XVII, pp. 209-210, 1886.

PROBABLE ORIGIN AND DIFFUSION. LC

:
bugs mostly acquire fully developed wings. He had never found the
species 1n grain fields of any sort, but always in grass lands, generally
in timothy or clover, but sometimes in wild grasses. Of eleven speci-
mens collected from under the bark of an old log by Mr. J. Pettit, of
Grimsby, Ontario, in 1866, and sent to Mr. B. D. Walsh for determi-
nation, all were of the short-winged form.” It was these specimens
that doubtless led Doctor Riley” to call attention to the fact that
in Europe there are many genera of half-winged bugs which occur
in two distinct or “ dimorphous ” forms with no intermediate grades
between the two, viz, a short-winged or sometimes a completely
wingless type and a long-winged type. Frequently the two occur
tpgether and copulate promiscuously, while sometimes the long-
winged type occurs in particular seasons, especially in very hot sea-
sons, While more rarely the short-winged type occurs in a different lo-
cality from the long-winged type, and usually in that case in a more
northern locality. In northeastern Ohio the species occurs during
some years in great abundance and very largely at least on timothy.
Here the short-winged form is very largely in the majority, and in
the spring of 1897, of 1,900 specimens collected indiscriminately,
only about 400 were of the long-winged type.

In northern Indiana, where the insect occurs but rarely, this short-
winged type does not predominate; but aside from these two locali-
ties, with an acquaintance with this species running over forty years,
chiefly in Indiana and Illinois, the writer has never met with the
short-winged type among millions of adults. If this short-winged
type occurs elsewhere to the westward, except along the Pacific
coast, where both forms have been collected by Koebele and others,
it has not been found by entomologists, even to the northward as
far as Minnesota, Winnipeg, and Manitoba, while to the eastward
of this Mr. Van Duzee collected the brachypterous form on Muskoka
River, Ontario, near the lake of that name.° | On comparing speci-
mens from New York with a large series from Kansas, the former
were found to be quite uniformly more robust, with longer hairs
on the/pronotum.@

It would seem that here we have evidence of two distinct tides of
migration, the one sweeping north and eastward, while the other has
mainly been to the north and westward, meeting the former in north-
eastern Ohio and northern Indiana, and possibly somewhere farther
to the north in British America. The two, besides differing in the
Jength of the wings, are sufficiently unlike in appearance to attract
the attention of students of Hemiptera.

«Practical Entomologist, Vol. II, p. 2%.

6 Second Report on the Insects of Missouri, p. 22, 1870,
éCan. Ent., Vol. XXI, p: 3, 1889.

@ Loe. cit., Vol. XVIII, p. 209,

whe) THE CHINCH BUG.

RELATION OF THE INLAND AND SEACOAST SHORT-WINGED FORMS.

It is possible that the short-winged form of chinch bug found in
Ohio is precisely the same form as that found along the seacoasts,
but it seems to the writer that the inland form originating from this
maritime short-winged element, instead of acquiring wings of normal
length as it drifted away from the coast, has really moved in the
other direction, and the wings have become still further aborted.

It will be observed by the illustrations given of both the inland and
maritime short-winged forms (see figs. 8 and 4) that in some of the
former the wings have become so aborted as to become almost invisi-
ble, while in the latter, though the wings are very much shortened,
they are nevertheless very clearly to be observed. It would seenf,
then, that we might reasonably presume that the species was orig-
inally long-winged, but, living along the seashore, the winged indi-
viduals have either flown each year inland or else been blown into the
sea to such an extent that a short-winged form has thus been evolved
which was unable to migrate and not easily blown into the sea. In
pushing inland while the country was still inhabited by the aborig-
inees another source of destruction would confront these insects in
the annual recurrence of fires whereby vast areas of country were
burned over in autumn, winter, or early spring, and these must have
destroved very many of the hibernating insects, while such indi-
viduals as migrated to sections not so burned over would escape
destruction.

PROBABLE COURSE OF DIFFUSION.

Let us suppose that the species originally worked its way north-
ward from South America, or even Panama, along the lowlands be-
tween the more mountainous interior and the Gulf of Mexico until
it reached Texas with its vast areas of level country extending not
only across the State itself, but northward into British America, and,
generally speaking, with the exception of the Ozark Mountains in
Missouri and Arkansas, eastward to the Appalachian system extend-
ing from Cape Gaspé, Quebec, Canada, to northern Alabama. This
area is more or less covered with a grass flora that affords ample food
for these insects, and it would seem that there was here offered every
incentive to migration broadly to the northward and eastward, and
at the same time there would be the Gulf coast along which those
individuals which either could not or did not migrate inland could
make their way as had their progenitors along the coast in Mexico.
(See fig. 17.)

Now, it would appear as though the short-winged individuals, if
there were any such, would remain along the coast, while the long-
winged individuals would, at least more or less of them, migrate in-
land, and at least some of these, but far more of those unable to fly,

PROBABLE ORIGIN AND DIFFUSION. 79

SO THE CHINCH BUG.

would be annually destroyed by the prairie fires, thus eliminating
whatever tendency there might be to perpetuate the brachypterous
forms, and develop a fully winged more or less nomadic race which,
as it slowly advanced inland, lost all vestige of its brachypterous
ancestry, if such had existed.“

On the other hand, we might expect the shore-inhabiting in-
dividuals to continue in their progress along the coast, the winged in-
dividuals continually migrating inland, leaving a mixture of the two
forms to push forward to the east coast of Florida—where as late as
1906 it attacked grass on lawns about Palm Beach—and northward
along the Atlantic to Cape Breton. As soon as this migration had
passed the southern terminus of the Allegheny Mountains the inland
spread would, very largely at least, be restricted to the area lying be-
tween the eastern slope of these mountains and the coast, thus leav-
ing the whole area to the west to be occupied by the northward tide
of migration instead of that from the east. East of the Mississippi
River and: south of the Ohio River the country is more heavily tim-
bered and the prairies are lacking, so that forest fires would here
take the place of prairie fires; but in the Southern States the woods
are composed more largely of pine, and Doctor Lugger, in Minnesota,
found that the chinch bug did not invade the region on which only
pine and other Conifer grew, but that the more southern counties
of his State, which are more or less wooded with deciduous trees, were
invaded. He also called attention to the fact that before the country
was settled by the whites these timbered lands were burned over fre-
quently, probably annually, but now the wooded areas are confined to
small tracts interspersed among the farms, and as these are not an-
nually burned over they afford suitable shelters for the chinch bug
during winter, and the grain fields of the farmer afford ample food
during the summer, while on the prairies which are burned over such
is not the case.?

Along the eastern coast the chinch bug has never been especially
destructive to the wheat crop north of North Carolina, where, accord-
ing to Doctor Fitch,° the earliest depredations occurred in 1783, while
Webster @ states that it threatened total destruction to the grain in
1785; but since that time the ravages have not been nearly as severe as
farther west in the Mississippi River Valley. In 1899, 1900, 1901,

aProf. H. A. Morgan, then entomologist of the State Experiment Station of
Louisiana, writing under date of May 30, 1898, states that he has never found
the brachypterous form of chinch bug in that State, and the writer did not
observe a single individual of these among the many macropterous specimens
taken by himself in that State.

do Virst Annual Report of the Entomologist of the State Experiment Station of
the University of Minnesota, 1895, p. 26.

-e Second Report on Noxious, Beneficial, and other Insects of New York, p- 278.

@d Webster on Pestilence, Vol. I, p. 279.

PROBABLE ORIGIN AND DIFFUSION. Si

and 1902 this maritime form destroyed the timothy in the vicinity of
Reidsville, N. C. This is on the southern border of+imothy culture
along the Atlantic coast, and some years ago an attempt was made to
grow timothy in that section. The grass did very well until the above-
mentioned attack occurred, and by 1905 there was but little remain-
ing. Strangely, too, nowhere along the Atlantic coast do we find the
short-winged individuals far inland until we reach New York and the
New England States, and what is equally perplexing they do not there
attack grain, but grass, whereas to the southward, except near the sea-
coast, it is the grain fields that are devastated by the long-winged
form. In other words, throughout New England, New York, north-
eastern Ohio, northern Indiana, and the Dominion of Canada we have
both the long and short winged individuals occurring together, but
depredating almost or quite exclusively upon timothy (Phleuwm pra-
tense).

In Bulletin 17, old series, Division of Entomology, U. S. Depart-
ment of Agriculture, Dr. L. O. Howard, the author, stated that in
1886 a timothy meadow located near Wakeman, Huron County, Ohio,
was considerably injured by chinch bugs. Since that time the species
has never been reported from that section of the State, and the writer
has found that depredations of that particular character are only
committed by the more or less brachypterous race. This has been
supposed to be largely confined to the northeastern portion of the
State, though there seems to be no good reason why it should not
appear in northwestern Ohio also. Owing to these facts this single
occurrence in meadows, recorded by Doctor Howard, formerly puz-
zled the writer greatly.

During the fall of 1898 there came reports of very serious destruc-
tion of meadows in Huron and Lorain counties, which lie contiguous
to each other, the cause being attributed to the dry, hot weather. But
an examination of the meteorological records for that section revealed
the fact that there had been no weather condition sufficiently severe
to affect timothy meadows in that way. A survey of the affected
meadows during early spring of 1899 revealed the presence of great
numbers of brachypterous chinch bugs hibernating in these meadows,
and the problem was solved. The species had doubtless been doing
more or less injury since 1886, entirely unknown to the farmer or any-
one else, thus showing the extent to which its secluded life in
meadows protects it from observation. This section of the State
since 1886 has been more largely devoted to dairying, and the
meadows are not as rapidly rotated with other crops as when the
cereals were grown more extensively.

«Extract from correspondence of Prof. Franklin Sherman, jr., State ento-
mologist.

26608—No. 69—07 6

82 THE CHINCH BUG.

In Ohio, which appears to be the frontier of destructive abundance,
the line separating the habitat of the combined forms and that of the
macropterous form, exclusively, indistinctly marks the line of separa-
tion between the most serious depredations and almost total immunity
of attack on timothy meadows by chinch bugs. To the west and
south of this a short-winged adult chinch bug is rarely seen, timothy
meadows are seldom attacked, and then only where fields of small
grain or corn are not in easy reach; as, for illustration, where the in-
sect happens to breed in a wheat field surrounded by timothy, and,

when the grain is harvested, there is no other recourse left it but to-

attack the grass. In the opposite direction from our line, however,
the conditions are quite the reverse. Here, while fields of wheat are
eccasionally badly injured, thousands of acres of timothy meadow
have been entirely killed out from its attack.

The area of destructive infestation of timothy meadows seems to
extend on the east in Ohio from Lake Erie to the Ohio River at the
northernmost point of West Virginia, and on the west, in the vicinity
of Sandusky, it extends only 25 or 30 miles from the lake shore. In
limited numbers the area of distribution extends westward, probably
narrowing gradually, around the lower end of Lake Michigan into
northern Illinois, where it seems to be on the increase, though still
far from common. <As will be shown further on, this form is not
likely to become destructive where timothy is grown in rotation with
other farm crops.

So far as it 1s possible to determine, there are a considerable number
of winged adults produced in this area every year—perhaps from 30
to 50 per cent some seasons—and these breed in the grain fields; but
at wheat harvest, instead of migrating to the corn, as is done else-
where, they go by preference to the timothy meadows. In western
New York, where both the long and short winged forms occur, Mr.
Van Duzee wrote that he had never found an individual of either
form in grain fields, but that they both literally swarm in timothy
during some years. Doctor Lintner told the writer that in the serious
outbreak of this pest in the meadows of New York in 1882 and 1883
about 20 per cent were of the short-winged form. Doctor Perkins has
recorded an attack of the chinch bug in a timothy meadow in northern
Vermont. Whether or not the short-winged form was the depredator
in this last-named locality the writer is unable to say, but, generally
speaking, the short-winged form is unknown at any considerable
distance from the coast, except in Maine, New York, Ohio, Ontario,
and northern Indiana, and but rarely does it occur in either form in
the two latter localities.

Just why this short-winged form should occur in such abundance in
the three States named is a matter that the writer is at present unable
fully to explain; but it does seem that this difference in food habits as

HABITS OF THE EUROPEAN BLISSUS DORIA. 83

between the two forms and the limited distribution of the short-
winged form inland might open the way to a solution of the mystery.
The writer believes that the insect is primarily a tropical macrop-
terous species, and that it has followed the coast from South America
along the Gulf and Atlantic northward to Cape Breton, and along
the Pacific coast to San Francisco and possibly beyond; also that it
spread from northern Mexico and Texas northward as far as Winni-
peg, subsisting upon the native grasses, and in the meantime spread-
ing also to the eastward to northern Indiana and Ohio.

} On the other hand, from the Atlantic coast there has originated a
tide of diffusion the trend of which has been westward, the bugs
here partaking more of the nature of their seashore ancestors, more or
less of them being of the short-winged form, which their less nomadic
habit has served to further emphasize. This tide of diffusion has en-
countered what the western tide did not, at least until much later,
namely, the timothy meadows of the Caucasian agriculturist, and,
adapting itself to this food
plant, has held closely to it,
thus avoiding the necessity
of seasonal migration. In
northeastern Ohio and pos-
sibly in northern Indiana
and northern Illinois the |
western tide of diffusion
has met the eastbound tide
and is perhaps amalgamat-
ing with it. (See map,
fig. 1%, iustrating sup- Fig. 18.—Blissus doria: a, first nymph; ec, second; b, third;
posed direction of diffusion d, fourth. (From illustrations prepared in the Bureau
of chinch bug.) ren gees

Although not at all conclusive evidence, it might be added that the

single specimen taken at Winnipeg by Doctor Fletcher was of the
macropterous form, while the single example taken by Mr. Van Duzee
at Muskoka, Canada, was of the brachypterous form; and this, with
the fact that the specimens from the island of Grenada were of the

_former and the Florida coast specimens of the latter exclusively,
shows that latitude and climate have no effect.

>

HABITS OF THE EUROPEAN SPECIES, BLISSUS DOR FERR.

Prompted apparently by a review of one of the writer’s papers
read before the eighth annual meeting of the Association of Economic
Entomologists at Buffalo in 1896, Prof. Karl Sajo, formerly of the
Kg. Ung. Staatliche Entomologische Versuchsstation, at Budapest,
published a short paper on “Unser Blissus doriw,” * which is so full of

4 Tilustr. Wachenschrift ftir Entomologie, Vol. II, pp. 449-451, July 18, 1897.

84 THE CHINCH BUG.

interest that the writer has reproduced it here, together with figures
of the larval, pupal, and adult stages of the insect (figs. 18 and 19).
Professor Saj6 writes as follows:

In the article on the eighth annual meeting of the Association of Economic
Entomologists (No. 26, pp. 401-403, Illustr. Wochenschrift fiir Entomologie) the
very instructive observations of Mr. Webster on the “chinch bug” (Blissus
leucopterus) in the State of Ohio were discussed.

In view of this communication I will give more in detail that which I have
observed concerning our European species of this genus, namely Blissus dorie
Ferr.

Like the North American larger species, the smaller European one appears in
two forms, namely, the wingless and the winged. The first describer of this
species, Ferrari, in Genoa, recognized only the wingless form, which with its
aborted wings looks very much like Hemipteron-nymphs, and probably by all
entomologists who previously saw it was not considered as a sexually developed
adult, but only the immature form
of some already known species. I
discovered the winged form seven-
teen years ago (1880) in the steppes
sand desert, called “ Nyires” of
the Kis-Szent-Miklos, and de-
scribed the same.¢@

I at that time made known the
characters of the immature forms,
which can not be confused with
the individuals which have reached
complete sexual development, in
that the immature individuals are

vermilion red while adult indi-

Fig. 19.—Blissus doriw: Wingless form at left; winged viduals are dark brown. It is
form at right. (From illustration prepared in the
Bureau of Entomology.)

interesting that the relationship
between the winged (macropte-
rous) and the wingless (brachypterous) individuals of the American and Huro-
pean species is very different. For while in America those individuals which
reach maturity are almost always winged, with us in Europe they are in gen-
eral only short-winged, and individuals capable of flight are not observed ;
and the fully developed macropterous individuals were not thus far, according
to my knowledge, found in any other place than in the central Hungarian sand
dunes already named, and here they occurred only on a single little portion which
only measured a few paces in diameter. It was a “ Dunenhugel” (sandy hill)
covered with high, scattered poplars, whose fallen, dried foliage sparsely cov-
ered the ground.

Here lived the colonies of Blissus doriw on the bases of the bushy, growing
grass, almost under the surface of the ground, and well concealed. The habits
of the European species are also in the main similar to those of its American
relative, since the latter also lives only on grasses, and during its development
also lives very close to the surface of the ground.

It is extremely remarkable that, even though B. doriw is very widely distrib-

aK, Saji: “Die bisher unbekannte makroptere Form yon Blissus dorie Ferr.”

99

Entomolog. Nachrichten, 1880, p. 255.

HABITS OF THE EUROPEAN BLISSUS DORIA. 85

uted here, and is met with not only on the “ Flugsande” (sand drifts), but also
in the hilly regions (e. g., on the southern exposure of the hill which stands
between Duka and Szod, in the midst of bluffs or rolling hills), the winged
specimens were to be found only on the very small “ Blissus Island” under the
poplars. But here also they were found but rarely, and only then when the
transformation from the pupa to adult stage was in full force. When there
were ho more pup to be found, then also the search for long-winged individuals
was in vain.

This appearance I explain in this way: That the winged examples, as soon as
they were able to fly, quickly flew away and disappeared in order that they
might serve as progenitors for new colonies.

But the place of discovery has since been transformed into an immense yvine-
yard by the Government, whereby grass, poplars, and also B. dori@ had to disap-
pear from thence. For four years I have, though seeking with the greatest
diligence, been unable to get track of the winged specimens anywhere in this
region, even though I know of a number of colonies of this species upon my own
premises. While formerly I captured a few specimens each year and gave them
partly to museums and partly to entomologists, I scarcely hope to attain such
interesting finds in the future.

The difference just mentioned between those individuals capable of flight and
those not capable of flight in our species and also in the transatlantic species
ean hardly be accidental, but may be sought for in the influences of environment.

Next there crowds to the front the fact that in North America B. lewcopterus is
continually subjected to the attack of its deadly fungus parasite to a high degree,
and its colonies die out as soon as rainy, moist atmosphere prevails. Conse-
quently, the Blissus species living there must always hunt new habitats and be
wandering continually to far distant localities. For this wings are of course
necessary, and only by means of these is the species enabled to sustain itself at
such a high grade of importance that it can, now here, now there, become a yeri-
table plague to agriculture.

With our European species it seems, on the contrary, in regard to many
points to be otherwise; for, while her habits in the main are similar to those of
her sister across the sea, yet there are found many important differences in
their environment.

Blissus doriw never congregates in such close masses as we read of in the
American reports. It forms only insect islands, and even individual families
seem to scatter out to some distance. In the steppes, moreover, the growth of
grass is not matted, but stands in isolated bunches on the partially bare ground
the bunches being not infrequently separated by several paces.

Our species will not go into cultivated fields. I have never found even a
single specimen among forage plants that have been sown, and already this
condition is one of the reasons why the European species does not cluster
together in such uninterrupted masses.

If, then, this is true the attacks of entomogenous fungi will hardly be able to
create such havoc in B. doria as it does among B. leucopterus in America.

I have also during eighteen years never observed a wholesale dying off in the
localities of occurrence known to me. The fungus 8S. globuliferum has perhaps
never attacked it, and even though the European form were susceptible to
similar pestilences, yet it is always hardly to be doubted that the fungus in the
European homes of B. doriew would not find favorable circumstances in that
here during the period of development of this species in normal years great
drought prevails. Rains lasting for a number of days, with continued moist
and warm atmosphere, belong, with us, among the rarities, especially during

86 THE CHINCH BUG.

the summer, and it is the young stages that are especially sensitive to the fun-
gus attack, as has proven to be the case in America.

Among insects there may possibly be found Blissus enemies, even though the
extremely penetrating odor of this bug, which is identical with that of the one
living in beds in houses, may serve as a protection.

Taking all of this together, we observe that our European species is in less
danger than the American, and that it is not subjected to catastrophes of total
destruction, so far as has yet been observable in the stationary localities of occur-
rence in the open field, for I have never yet observed a sudden disappearance
from the localities known to me. It is not necessary, therefore, for it to be
continually hunting up new fields in which to thrive, and there was no appar-
ent reason which in the struggle for existence would have given preponderance
to the long-winged form; and so in time, in the generation of our species,
which originally, perhaps, was full winged, the winged form became less and
less numerous, until to-day we see almost entirely brachypterous individuals
in the adult stage, exactly the same as in the bedbug, Acanthia lectularia, with
this difference, that among the swarming masses of the latter nowadays not
a single example with fully developed wings can be found, fortunately for us.

It is evident that the long-winged tendency in B. doriw is disappearing, and
the time may come when one will be unable to find any long-winged specimens.
The designated dangers, on the contrary, against which the chinch bug must
fight in North America require very strong migratory powers, and, conse-
quently, well-developed wings, through which this especially significant differ-
ence between B. doriv and B. leucopterus has been brought about.

As to the question whether or not our species shall be considered injurious, I
can answer that it in no wise belongs to the entirely indifferent insects, but, on
the contrary, contributes to the complete drying up of the rather sparse grasses
of our steppe meadows during the summer. But since it has not thus far
housed in the cultivated fields, it can not be placed upon the black list of seri-
ous depredators. Whether, moreover, in the future, when in consequence of
the continued destruction of its herding meadows, its original food plants dis-
appear more and more, B. dori@ may become, like so many other insect species,
a depredator through necessity can only be conjectured. We have in this re-
gard already recorded entirely too many remarkable transformations in the
menu of other species to disregard entirely the possibility of a similar transfor-
mation in the life habits of our B. dori.

IT wish also at this time to state, for the benefit of our many readers who may
not be familiar with it, that in the dimorphic bugs, especially those in which
the macropterous and brachypterous forms are found simultaneonsly, the
former possess a much stronger and broader thorax than the latter. As a
result of this difference in their physical structure, one is, when comparing
them for the first time, easily inclined to designate them as two distinct species.

In addition to this, there is in Blissus the strikingly beautiful coloration of
the long-winged specimens, whose clayus and corium are light ocher-yellow, and
the unusually large membrane, which is about twice as large as corium and
clayus together and of an entirely milk-white color, making the long-winged
individuals very prepossessing. The individuals with rudimentary wings, on
the contrary, are of an obscure chocolate-brown. The larvee are, as has already
been stated, of a bright vermilion-red color, marked with black.

With the foregoing, relative to the habits of an allied species of
Blissus, it seems to the writer that we can the better understand how,

a@Translated from the German by Mr. C. W. Mally.

PROBABLE ORIGIN AND DIFFUSION. S7

under one set of conditions, all traces of a short-winged form might
entirely disappear, while with another set of conditions this tendency
might not only be perpetuated, but greatly emphasized. The two
species, B. Jeucopterus and B. doriw, are fully illustrated in all stages
of development, as well as both macropterous and brachypterous
forms. (See figs. on pp. 21, 22, 23, 83, 84.) For specimens of the
latter species, B. doriw, we are indebted to Professor Sajé.

PREVIOUS IDEAS ON THE DIFFUSION OF THE CHINCH BUG.

Formerly it was supposed that the chinch bug was a native of the
Atlantic coast States, and that it made its way westward with the
advance of civilization and the consequent progress of wheat growing.
This theory was based upon the fact that the original description was
drawn up from a specimen from the eastern shore of Virginia, col-
lected by Mr. Say himself,‘ and, as before stated, the earliest destruc-
tion on record caused by this insect occurred in North Carolina, and
it also committed great depredations in Virginia in 1839. Up to this
time it had been Fans that it was a southern oes confined to
the country south of latitude 40° north. But about this time chinch
bugs appeared in I]linois, at Nauvoo, simultaneously with the settle-
ment of the Mormons at that place, and as many supposed that this
sect brought the bugs to the couhtry with them, they were toe
termed * Mormon lice.”

In his second report, page 284, Doctor Fitch states that Mr. Wil-
liam Patten, of Sandwich, Dekalb County, I1., informed him that the
chineh bug first appeared in that locality in 1850. Mr. Patten, the
father of Prof. Simon Patten, of the University of Pennsylvania,
and the writer’s father, settled in the immediate vicinity of Sandwich,
TIL, in 1852. This was ten years after the Pottawattamie chief, Shab-
bona, and his tribe had migrated to Kansas or Nebraska, the writer
does not remember which, but he does recall that 1t was about this time
that the prairie fires ceased to occur over any wide areas, as the prairies
were no longer fired annually by the Indians. The whole country was
fast being occupied, and he well remembers that the settlers would
decide upon a certain date on which they would set fire to the wild
grass—in late autumn—so that all could be prepared. It may also be
stated that there were very few timothy meadows at that time, as the
wild grass afforded an abundance of hay, and not until years after
did cultivated grasses come into general use. The writer also knows
from personal experience and observation that with the decrease in
prairie fires there came an increasing abundance of chinch bugs,
which attacked the wheat fields of the farmer. Up to about 1862
these fields were largely of spring wheat, but about that time there

@The complete writiugs of Thomas Say, edited by Le Conte, Vol. I, p.

SS THE CHINCH BUG.

was a rapid decline in the growing of this grain in northern Illinois.
It seems possible that spring wheat might be more liable to attack
from chinch bugs than fall wheat, as the former is, at the time when
chinch bugs seek out their breeding grounds, more tender and inviting
than the latter. Mr. Walter Young, writing from Galesville, Wis.,
states that his spring wheat was totally destroyed in 1897, though
there had been none sown for ten years previous on the premises, and
while the chinch bugs did not ordinarily do much injury, just as soon
as spring wheat was sown they returned, as it were, and destroyed it.

If spring wheat is so attractive to chinch bugs in spring as this
would indicate, might it not be used for baits instead of millet, as is
advised further on, in order to draw off the females in spring when
seeking localities for oviposition ?

This was in a country where there was comparatively little timber,
the only forests, if such they could be called, being along the streams
of water. The writer is confident that the chinch bug did not sud-
denly make its appearance in that section, but that with the increase
of grain growing and the decrease of prairie fires its effects began to
be more and more marked. Since then Prof. S. A. Forbes has secured
information of the occurrence of these insects in sufficient numbers
to attract attention as early as 1823 in southern Uhnois, and within
°5 miles of New Harmony, Ind., where Thomas Say resided and did
the most of his entomological work.

REASONS FOR THE PRESENT THEORY OF DIFFUSION.

It seems to the writer that in all of this we have good grounds
for supposing that the chinch bug occupied the most of the country
prior to its occupancy by the white man, and that its first depreda-
tions were caused by its own advance coming in contact with the
advance of civilization; and the simultaneous cessation of forest and
prairie fires, with the displacement of the native grasses by large
areas of wheat, so combined that the points of contact were in Ih-
nois in the West and Virginia and North Carolina in the East. Not
until within the last twenty-four years has the chinch bug been
known to work serious and widespread injury east of the Allegheny
Mountains, north of Virginia; and west of these mountains they have
done scarcely any damage north and east of a line drawn from
Chicago southeast to Cincinnati. Thousands of farmers in Ohio
never saw a chinch bug until within the last thirteen years, and there
are thousands more in northwestern Ohio, southern Michigan, and
northern Indiana that, even yet, would not be able to recognize one
were they to see it among their growing grain, or even 1f in abund-
ance. But in considering this matter the fact must be borne in
mind that timothy meadows are not burned over annually as were the

PROBABLE ORIGIN AND DIFFUSION. 89

forests and prairies, and the stubble does not die with the harvesting
of the crop as in wheat, and therefore annual migrations are not
necessary for the bugs in order to preserve life. In a_ timothy
meadow the species may live on and reproduce year after year with-
out ever being obliged to abandon the field. It was the wheat fields
of the West that the eastbound macropterous tide of migration found
confronting it in Ilinois, and the smaller fields of grain and timothy
meadows that the combined macropterous and brachypterous forms,
more or less maritime and northbound, came in contact with along
the Atlantic coast, while at the present time the two tides of migra-
tion have met in northeastern Ohio and northern Indiana.

In figure 17 is illustrated the theoretical directions and courses
taken by each of these tides of migrations from the tropical regions,
and in figure 1 the areas over which the species is now known to
occur in Central and North America are indicated.

The writer believes that this same course of migration has been
pursued, at least in the West, by the several species of Diabrotica,
and especially D. longicornis Say, and to a less extent by another
species of Hemiptera, Murgantia histrionica Hahn and possibly also
by Dynastes tityus L., while the two latter with others are now
working northward along the Atlantic coast. Besides, the westward
tide of migration has been followed in all probability by Pontia rape
L., Phytonomus punctatus Fab., Hylastinus obscurus Marshm., and
Crioceris asparagi 1, all of which have first become destructively
abundant west of the Allegheny Mountains in extreme northeastern
Ohio. The last four species, having been introduced from Europe,
have undoubtedly migrated westward.

With an almost total lack of natural enemies in the United States,
and with nearly all of its closest allies belonging in Mexico and the
West Indies, it would seem as though we were in possession of addi-
tional evidence of the chinch bug’s tropical origin. Besides this the
name “chinch bug” is of Spanish origin, and this language has
never been in common use in North America except in Florida and
the country along the Mexican border.

The species certainly prefers the low country to the higher, and is
seldom found in any numbers at an altitude of over 2,000 feet. Gen-
erally its habitat is 1,000 or lower. The altitude where it was
found breeding on Volcan de Chiriqui, in Panama, is 6,000 feet; and
of its habitations in Guatemala, San Geronimo, is 3,000 feet; Panzos,
2,000 feet ; Champerico, sea level, and Rio Naranjo, about 2,000 feet,
while in Colorado it occurs sparingly near Fort Collins at an eleva-
tion of 5,500 to 6,000 feet, while Professor Cockerell did not find it at
all in the same State at elevations of 7,000 to 8,000 feet. On Mount
Washington, in New Hampshire, it has been found only once, and this

90 THE CHINCH BUG.

time by Doctor Packard, on the summit, which has an elevation of
6,500 feet. Rx

In his own experience, running over something like forty-five
years, the writer has never witnessed serious injury by chinch bugs
to crops on hilly land. It may be stated, however, that all of his
studies of the insect have been carried on in a level country, Ohio
being the most uneven and hilly, but even here all of the outbreaks
observed were on level areas. In Minnesota, however, Doctor Lugger
found that those grain fields which were most seriously injured
were located near the edges of woods or on slopes. In some pub-
lished observations of Professor Osborn, in Iowa, kindly placed
at the writer’s disposal by Doctor Howard, we find that in 1894 about
90 per cent of the infested fields examined by Professor Osborn were
on high ground and about 80 per cent of the fields were hilly and
ridges, In most cases the damage being first apparent upon the
higher portions of the fields. The exceptions were where the chinch
bug had evidently hibernated in wild grass and weeds occurring in
the lower places, and these had been very dry for the twelve months
preceding the damage of that year. Besides, both the Iowa and
Minnesota areas are below 1,000 feet elevation.

The area over which the chinch bug is more especially abundant
and destructive comprises such a variety of soils and geological
formations that a study of these factors at once shows that neither
has any material influence in the distribution of the species, at least
in the United States. In its northernmost habitat it would not be
at all surprising that it should prefer a sandy, rather than a clay,
soil, the former being looser and warmer on or near the surface.
(See fig. 10.)

In conclusion, then, on this point it may be stated that if Blissus
leucopterus originated in the Western Hemisphere it was probably
near the Tropics, and it is not impossible that its generic ancestors
may have been carried from Europe or Africa by either the north
equatorial or the main equatorial Atlantic currents, landing them
on the northern shores of South America or on some closely located
islands, from which the species has spread coastwise around the
Caribbean Sea and the Gulf of Mexico, as previously indicated. In
this connection it 1s interesting to note that specimens from Grenada,
collected on the Mount Joy and Caliveny estates by Mr. H. H. Smith
in June and September, show that the species here attains a large
size and is more variable, both in size and markings, than is com-
monly found to be the case in the eastern United States.’

@ See paper by the writer on Origin and Diffusion of Blissus leucopterus and
Murgantia histrionica, in Journal of Cincinnati Society of Natural History, Vol.
XVIII, February, 1896. .

+ Uhler on Hemiptera-Heteroptera from St. Vincent and Grenada. Proc. Zool.
Soe., London, 1894.

INDEX.

Page.
Agonoderws, pallepes. enemy ofochineh) bug. 2] -s = ee ee 5
PMMA TM TOOGs Plamice Oly NG SUIS UO USUMUUS == 5 se Se Te Ese et ees 32
ANLEMOGOMS«)SCULdO-Ghiniche — Ul nupnleps (nSidioswus- 22 =2 = ke 60
AMIS (UNE SAIDSY Loy CO) cats aH) eH wy gt Ea a ee ee ee ee ee G
Aphides, Entomophthora aphidis a fungus enemy______________________- 47
AN NOGLUSIUVOMULUCUS mile Tabi On the DG eae = ee ee Se se 19
RE UC UeeTin Teel aN Set eA ie ee ee ee ae ES a eee SL 1¢
Enh enAnentin: sOOde DL Amit Of. Glimehs DU aoe ee Boe Se 8 ee oe 31
Bacillus insectorum, bacterial enemy of chinch bug_____-_______________ DT
Bacterial <oneniness Of sehen, (Oe ae se ee ee Sil, Bs:
Barley, food plant of chinch bug______ oe a et Pca NE Sa oe eh 2 oil
Barrens susSesa Sana st gem CMe Ue = =a io) ee > te 65-69, 72
iBiackpirdsredwine, cheniy of Chinchybugsa2 = eee 2 ee ee ee 59.
BI SSUSHOISiDUTOMe Oh AeNUS=: == 2a esa ee Semi Se pees Sees £
COA DUAR GE SoL GEN OU OTS aR ee RS PS is eee eae Nec 6) oe Be Ee 7 83-ST
migratory Habit contrasted with that of chinch bug________ 19
no disinclination to use wings, as with chinech bug________ 24
leucopterus. (Nee also Chinch bug.)
lO GHiy AnECOnd Satire et ee ae es Bese = 9-10
Bluegrass. (See Poa pratensis.)
Bobwhite. (See Colinus virginianus. )
Botrytis, Sporotrichum globuliferum closely allied______________________ ae
wrong determination of Sporotrichum globuliferum ~~~ ~~~ — 46
Brachyrhynchus granulatus, mistaken for chinch bug____-~____-_________ 32-33
Buckwheat, wild. (See Polygonum dumetorum or P. convolvulus.)
BurnineovertomMestroyrchincehsbugs= 25. 228s 2s ee ee 61-62, 72
Cabbage bug, harlequin. (See Murgantia histrionica.)
Copmoladaciylon=toodsplantio® chinghibuge ==). 92222 e eee oie
Carpocapsa pomonella, change of habits in South Australia______-____-____ 26
Canomadwenemy-of Chiimnchmitos= a= as. ee ee LE ol ge) bese 2 Ea 59
Genchrussonly crass not attacked by chinch bite - = 22225225. 2a 3
Chinch bug. (See also Blissus leucopterus.)
achilieed CSErip Ont ee ee a Oe SR Senne a ee OP:
AP Wedikall Ge he UN Casa ee eee es Se a oe ees 74-75
POAC CET ANGRY TINT CS epee EES Taeey gl pS ee ENC
behavior when attacked by Sporotrichum globuliferwm_—_—_— DD
HeneiiimaAllecedsstherehr@ni= 2 let ee ee Ol eet ei ee Ass Dy
UT MR @TIOTIT OS pees et tee eer eee see Sle es ee 58-5)
GESCRIP MONS Ole SUC SE a ee Fe a a ee Ss apee 21-22
destructiveness largely due to gregarious habits_____________ 28-29
GE ElOD NIG eee eer eee eae ner Pee eee ee 22-25
AiO MEV IOUS IOGHSAS o enh ges oS oa RS fe 87
WLOM ADI SACOUES Cis hae a it ae 2! ety ee 78-83
LeASOUSsrOLEPresen i uneOn y= =. n= 2 ery See Se 88-90:

92 INDEX.

Page.
Chinehibues GiSeriPucion, Leo Srey Nn Cea a es 10
Vertical soa = Se sek ee ee eee 89-90
HNO AOvea ooh hal yearn \yreruMoeee Ss 18
freezing: Pee eee ee Se ee ee ee 17-18
ego) eseriptoms2 === mee ee ee ek ee eee Zi
Period: 22> 232 2 wee ee Soe eee | eee ee 20-21
eres, number depositede.2 322-5 2e Te ee eee es 20-21
false. (See Nysius angustatus. )
food” plants: {> -8t. 02 Ses caw Se et eed oe eee 29-381
FOrM tromiGrenada laren diver tally ems eae ee 90
FOL AMEN my oats as See ee ME I I ee ee 59
fungous’ enemies ——— = ew he ee ee 44-58
POL AC LLG AUD avail eye eet ee eee ara ee eee 58
Fenerations rannvalily = St ee ea ea 25-28
erecaniouss Nabites=== =.= ee, Mere te De te eh AS go T4-T5
habits account largely for destructiveness________ 28-29
habits of PyOUNS: Bee © eee A Be ee ee er 22-25
iMeSr Ma GON 2a eS Sl ay pa ee ade gd a eee 10-18
OMSL OKAY oME NO 3648
LEM PEVAUUTG ; se ee es ee a 4844
insects amistakensthereiors= = 2 Sass Bae ee ee eee 31-383
invertebratevenemies 222. ee 2 ee Ase she Ee 60
larvallstages descriptions. =). = eee oe et eee 21-22
long. (See /schnodemus falicus.)
long-winged form and short-winged form___________________ 22,
occurrence and dis-
CEUPUbIOMy 2 (5-17
LOSSES SSeS Sane ee ee ie ee ee 33-36
meteorological influences favoring fungous enemies__________ 56-57
LOM REAENHO IM CONay 1EOKONE vim Seteeelal (OVE iwool_ 24
migrations in spring, Summer, and autumn__~__________ 18-20
NAMeLOL Span Shvonigin ss see ee eee 89
mature ECW CK Se S= ke SR ee ee ed ee ee ee 36-44
TNE TINGS cee es nae 7 ee I ee ee eee ee 44-60
Ook Hray Enoel Chiartweom, jo —— 72-90
OV MpOsiGlO mse ee ee ee 20-21
pupa; Gescriptiont os os hee ee Tee ee ee eae eee 22
remedial and preventive measures___-=_- == 60-72
short-winged, relation of inland and seacoast forms __________ 78
WHrysopas enemies; of ce bya Chay ye ee ee ee 60
Clover-leaf weevil. (See Phytonomus punctatus.)
Coalstar, use ine barriers cams ty china Chay fo oes eee ee 65-68
Codling moth. (See Carpocapsa pomonella.)
Colinus, Vinginianus Enemy, Ofre Min Chae eee ee en 58-59
CO POMMOD AOS (CMT OGH®, aaNeserouaes leo 19
Conimeleng pulicaria, mistaken! forsehimnchy pices ee ee 32
Grorinas Voyeroraroaty sxoxorel javleways (oye ClayuaKelat |oybyeee ee 3
Chicken; Loos plaiatey Of: 1c tine TT ee ee ara 31
foods plant.of chinehe bug = Se eee ee eee ee 30
Cow blackbirds, decrease held responsible for increase of chinch bugs____ 15
Crab grass, occurrence of half-grown chinch bugs in fall_=-_--__________ 10

OMOGCETIS, GSP AT AG1,. COULSE: Ot: CL SO Tee ee ee 89

INDEX. 93

Page
Ciinodena Tiorda- mistaken tom chinch Dues 2222252 ee ee 3D
Decoy plats of grains and grasses, in combating chinch bug_________-_--~- 62, (2
DraproticasCOULSe! Ole GitbUSION =. one. t Seeie es i es ee See 89
LOTUGLGO Te1Ut Seer er ey es aoe pe Da Rae 2 Se Fs A ote Soe 76
COUESCLORCIMEUSTOMe a2 an Be eee Sees 89
PO TMOTO RH MEST ee MUCUS SQL Su CQO RUC te ee ee ees eee a ee ee 84
CEL AUNONEY a nl OKs ene OE ee ee A ee ee 22, 15-17
Dispersal: of&snsects, eftected) by Stonms == = 5 = ee 19, 20
DIStribmhioneeocrapnicalwoL, echinche pues = es oe eS 9-10
MET Galle nO lm GHUNC a Urea es ee ee et ee oe Se ee J 89-90
Drought necessitates watchfulness against chinech bug_-____--____-_____~_ 63-64
DUMGASUGSMUULY RUS COMLSCL OL ACitsUISl@Mea = = as 2a ee eee 2 89
HGOneNAaMmanen.«chemynoh ChinGhypuce ses soe eee se ee 25
LPS COIs: — ELOMODNUOLUNU DUS aa ee ee AT
Entomophthora aphidis, fungous enemy of aphides_____________--___-___- AT
Ghinchip ase se ee ie 44, 47
spherosperma, fungous enemy of Phytonomus punctatus— 55
Epicauta pennsylvanica, attacked by Sporotrichum globuliferwm__——~--~~ 53
Hires. prairie, relation-to numbers of chinch bugs2==2=2- 2") 2s_S ss 87-89
LG UuIMestOMAeSELO y VGlin Chie eae ae Se See a Fe ee oe 6
Flower bug, insidious. (See T'riphleps insidiosus.)

mreeZinome-ect vommaclhiiin Clips se— = See 2s a eae 17-18
HNO omen e MyaO ls Chin Chg Dore se =e tee et ee ee ee sls Se SS ee 59
MUM OUSTeneMml ES Oichin Covi a= sas ee eee ee oe te 44-58
GeterminaLiOns wes ee ea SS ee 46-47
field and laboratory experiments in Indiana_ 47-50
Lest el Gea DlGalTtON sae oes wee pill

meteorological influences favoring deyelop-
TUCSNOY Ee eee Spe SAO See ee oe eee 56-57
DERACEICH IE thi yee ees eS Se ee ee 58
work of Professor Snow in Kansas____--~—_ 51-53
Geoconsnoredis- mistaken tom chinchyDue =) === ee eee 32
HLL S Mis talcenetoim Chinchepug= === =) === =n == os ae 1 B2
hivgOsUsS Mistaken Lom chinchw buses. == ==. s=== = Bey
hmbanis mistakenstorzchinchs pugs == 22s aes 2s ee 32
Craineallaiaindsroodlplantsiotichinchybug.==2— =) a= 2 2 Sse ee 30

Grass, Bermuda. (See Capriola dactylon.)
blue. (See Poa pratensis.)
bottle. (See Irophorus viridis. )
crab. (See Syntherisma sanguinalis. )
meadow foxtail. (See /rophorus glaucus. )
oat. (See Arrhenatherum. )
panic. (See Panicum crus-galli.)
Grasses, all but Cenchrus attacked by chinch bug_—________-__-____---- 13
“Hair snake.” (See Mermis.)
Herd’s grass. (See Timothy.)

EM eRianoneore- chinGhy bugs = ===. ae ee ee Pe Se ee eee 10-18
Hungarian grass, food plant of chinch LS Ss a vp ee ees ee 29
HAUS Gi USROUSGIEUS. =COUtSCLOLs CithUSION == === n= 22 2 et 2 ee _ 89
AMSeCis dispersal: eiecteds byeStOnMs == — 2 eas ee ee eee ee 19520
Tschnodemus fakeus, mistaken for chinch bug=_----_------_--=+----_--_ 32

Ischnorhynchus didymus, mistaken for chinch bug———-----~-------_.-__ 32

94 INDEX.

Page
TLOPLOTUS OLAUCUS, LOOM PVAtAOL «CLUES yr ee eee a 29
VUTIGIS) LOOGNPLAME, Of Eli GHG yee eee 31
JUNCUS, place onhibernatiomrOf, Chin eh sb te se ee ee ee eee 15
iXerosene emulsion, use against chinch buge—--_-2-_ 2 eee 64-65, 72
use againstichinch: bug so = Sat eee se ee eee 64-65
+ Ralidées,’ iobserved feedin= on chinch DusS222 222 eee 128
Lace-wing flies. (See Chrysopa.) ;
Lachnosterna, attacked by Sporotrichum globuliferum___________________ 53
ady-beetles, enemies of chinch buge.22e ss 2 ee eee ee 60
Barks norned: enemy or chincgh hugs === ee ee 59
UALS 2 ODSCLVEO) TECMIMSs Orne CLIN hs VS eee als}
Leaf-bug, ash-gray. (See Piesma cinerea.)
Tigyrocoris sylvestris, mistaken! for chinch bulges s= = = see 32
TOSSES! frome Chui CH py eee Le a LE = 30-36
Martins, decrease held responsible for increase of chinch bugs__________ 1B
INR OHA we Ke, GCravsronye Oye (MVE \oKR =a s 7 ES ew 59
Meadows, difficulty of combating chinch bug therein_______~________- -___ 62-63
Megilla maculata attacked by Sporotrichuwm globuliferuwm___—~-—~----_~ 53
Mernmis! parasite: ot <chinch (bugte 2522 sae eee 60
Meterological influences favoring fungous enemies of chinch bug________ 56-57
Migrations Of chinch= pues =.2) 22 4 eee eee eee 18-20
pedestrian.—-s:-8 S| 2 eee Ae ee ee eens 24-28
Nillets toods plantaokeachineh: bug2=-2— = a eee 29
Miigas einciusenemyeaor Chineh pus. =a a ee ee 60
Monocrepidius, attacked by NSporotrichum globuliferum_—_~~-~-________ = 53
“Mormon lice? early local mame for: CHCl Ul ase ee 87
VEO ONAN MUSUPLONAGE, COULSE sO tea Cts UST OMe eee eee ee ee 89
not attacked by Sporotrichwn globuliferwmn_____— 53
Naupactus xanthographus, attacked by Sporotrichum globuliferwn___———— 53
Negro bug, flea-like. (See Corimelaena pulicaria.)
Nysius angustatus, mistaken for chinch Hine es bees Soe el ee 31-382
Oats; food plantiot chinch J\oug2s 22-3 ee eS ee ee ee ou
Ovipositionv of <chinchebu ges. a es ee ee ee eee 20-21
Panicum crus-galti food, plant of chinch bug == 29
Parandra brunnea, attacked by Sporotrichum globuliferum___~——-__----_- 53
‘Partridges,” decimation heid responsible for increase of chinch bugs__- als
Pasturines remedy stor chinchipl ca] 2s e ee eee 62
Peliopelta abbreviata, mistaken for chinch) pws 2 o== = ee eee 32
Phleum pratense. (See Timothy.)
Phytonomus punctatus, behavior when attacked by Hntomophthora
SPROLOS DELI Se SE dee 2 ee eee 55
Course Vor (diitusion 222 Se ee ee eee 89

Piesma cinerea; mistaken. tor chinch bugs. 2 ee eee 7 TE ge
Plowine, LULLOWS, barriers ta San SiG wc katie la ty ee ee oe

(ep)
ive)
=] =] 0
bo bo bo

EO «CES TUOY: aC LLC l ny oo eo ee ee ce ee 63,
Plum curculio. (See Conotrachelus nenuphar.)
POO. Dr atensiss LOOd plants Ofc GHC HS Ul es ee ee ee eee 31
Polygonum dumetorum or P. convolvulus, food plant of chinch bug____- 51
Pontia rape, course of diitusione====— = 89
Prairie-chicken— enemys orchinch )ug= === ee 13, 59
Precipitation; anfluence on chinch pues 22 22322 eee 3648

Preventive measures against chinch bug, difficulty in practice___--__- ats 69-71

me

INDEX. 95

Page
Rurslane: foods plant Of MYSCUS) (ONOUSEMTUS == ee) 32
Quail. (See Colinus virginianus. )
Rain. (See Precipitation. )
Rice eehOOU em lamGr One Gham Clik Wea 22 Se ee 2 ee er ee Sil
Rushes. (See Juncus. )
ExeKe meLOOGE |) Lclliter Oe GMI Cl OU Seen See ee be ee Se ee ee 3
Salton Basin, occurrence of ehinch bug—_-__--—-—__~__ ER ea ne 20 29
Sand oats. (See Uniola paniculata.)
Setaria glauca. (See Ixrophorus glaucus.)
POLS MI ehOOCE Plante Ole GhimMChnDUS= = a= 2 oe Pee ees 31
“ Sparrows, bank,’ decrease held responsible for increase of chinch bugs__ 60
SMG erewepsOCCwLEeU Cen ChinGhpUSSe= == === = se SS eee 60
Sporotrichum globuliferum, first artificial cultivations_____________-_____ Ht
INSCCISeaveael eC ee te es Se De
DALLA SiiChOMm- ehinche pues ==. ==. es 44, 46, 47, 53
results of field applications in Ohiog__________ 54-56
WISerACaINstechinch pte ses oe (HE Tw
NLORMS mini MCencesin dispersal iO iNSCCtS=— ==" eae ee 19, 2¢
Sl Omer (ROG men enya @ lari Chile ates ee Sere ee 5Y
Syntherisma sanguinalis, food plant of chinch bug______________________ 3
Remmeratwreyininence on: Chinch buge= === = seas se a 43-446
irish roOwn -ehemyeOhremineh WUe ss. 2 es eee Ee a ee 59
himMoniyatoodeplantcOl Chimehs ges == Se ee ee es ZanoQ re
Trap crops (See Decoy plats.)
MA HesOnvOLUSAMGULLOSIES) IMiStalcen form ehinGhe pea == 22.5.) ee ee oe
MMOLE DSMINSIAOSWs, enemy: Or ehineh) PWG. 2- 52 = 22a ee ee 60,
MUS talseniemOles CHIMGH Cy OU =s ae eee ee a 32
Uniola pan iculata, abnormal habit of chinch bug thereon________________ Zawto
Mine tie OOO planar sOfechimcht Iyer ee ee ee eee 3
NMnen hose wenennvaOtes Clim Cli. DU Geese es ee ee ee 59

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U. S. DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY— BULLETIN No. 70. ,

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

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REPORT OF THE MEETING

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INSPECTORS OF APIARIES,
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Bes DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 70.

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

REPORT OF THE MEETING:

OF

INSPECTORS OF APIARIES,

SAN ANTONIO, TEX., NOVEMBER 12, 1906,

ISSUED JUNE 17, 1907.

WASHINGTON:
‘GOVERNMENT PRINTING OFFICE.
TOOL.

BUREAU OF ENTOMOLOGY.

L. O. Howarp, Entomologist and Chief of Bureau.
C. L. MArRLATT, Entomologist and Acting Chief in absence of Chief.
R. S. Cuiirton, Chief Clerk.

F. H. CHITTENDEN, in charge of breeding experiments.

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

W. D. Hunter, in charge of cotton boll weevil investigations.

KF. M. WesstTer, in charge of cereal and forage-plant insect investigations.
A. L. QUAINTANCE, in Charge of deciduous-fruit insect investigations.
D. M. Rocers, in charge of gipsy and brown-tail moth work.

A. W. Morrity, engaged in white fly investigations.

EK. 8. G. Titus, in charge of gipsy moth laboratory.

C. J. GILuiss, engaged in silk investigations.

R. P. Curriz, assistant in charge of editorial work.

MaBeL Coucorp, librarian. ©

APICULTURAL INVESTIGATIONS.

EK. F. PHiuips, in charge,
G. F. WHITE, exrpert in bacteriology.
J. M. RANKIN, in charge of apicultural station, Chico, Cal.
I’. G. Fox, assistant in apiary.
B. N. Gates, collaborator, Worcester, Mass.
JESSIE HE. Marks, apicultural clerk.
9

“a

LETTER OF TRANSMITTAL.

U.S. DerartMent or AGRICULTURE,
Burrau or Enromowoey,
Washington, D. C., March 18, 1907.

Sir: I have the honor to transmit the manuscript of the proceed-
ings of a meeting of inspectors of apiaries held in San Antonio. Tex.,
November 12, 1906. The meeting together and conference of the per-
sons interested in the eradication of the diseases which are such a
drawback to apiculture can not fail to bring out many points of
importance. In such a meeting the subject is presented in a way
which is not possible in articles written for journals devoted to bee
keeping or for publication in other forms. There is no organization
of inspectors and no funds are available by which these proceedings
may be published, and since this meeting was to a large extent the
result of the efforts of members of the Bureau of Entomology, and
since these men took such an active part in the meeting, it would seem
fitting that the proceedings be issued as a publication of the Depart-
ment of Agriculture. I therefore recommend that this manuscript
be published as Bulletin No. 70 of this Bureau.

Respectfully,
L. O. Howarp,
Entomologist and Chief of Bureau.
Hon. James Wiison,
Secretary of Agriculture.

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The meeting of inspectors of apiaries was held on the Monday fol-
lowing the close of the National Bee Keepers’ Association convention,
November 12, 1906, at San Antonio, Tex., as a result of a call issued
by Mr. N. E. France, inspector of apiaries for Wisconsin, Mr. W. Z.
Hutchinson, inspector of apiaries for Michigan, and the writer.

The object of this meeting was to get together for consultation the
men interested in the eradication and control of bee diseases. The
closer cooperation of these men in their work can result only in good
to apiculture and is greatly to be desired. Inspectors are chosen
from among the practical bee men of the county or State, and the
majority of them are familiar with their work on entering the service.
They also accumulate a vast amount of information concerning dis-
eases, most of which never reaches the bee journals or gains publicity
in any way. A meeting of these men brings out many points which
would otherwise remain unknown.

The meeting at San Antonio was most interesting and valuable, and
since much.that was said there has never been published it seems
desirable to issue the proceedings of the meeting in the form of a bul-
letin to add to the knowledge of the bee-keeping public on bee diseases.

At the close of the meeting the writer was asked by the inspectors
present to prepare a list of questions based on the laws now in force
for the control of bee diseases. This lst was prepared at once and a
copy sent to each of the inspectors whose name and address could be
obtained. At the same time the questions were taken up for detailed
examination, and various persons were consulted on points of impor-
tance which arise. This work is not yet complete, for it has assumed
proportions which were entirely unexpected at the beginning. It was
originally intended that this discussion of the laws should be inserted
as an appendix to the present report, but this would only delay the
present publication. This aspect of the subject may therefore be sub-
mitted for future publication as soon as it can be prepared. It is
very important that the best possible wording be used in a law to con-
trol bee diseases.’ The bee keepers of several States which do not have
such laws are at present interested in this subject.

6 PREFACE.

In the preparation of the manuscript for publication it was neces-
sary to rewrite the articles by Doctor White and the present writer
because the manner of presentation of these subjects in a meeting is
not suited for publication. It was also necessary to edit the discus-
sions, for the stenographic report was inaccurate in numerous places.
It is believed, however, that even if the exact words of each person
are not recorded, the meaning is the same as was intended to be con-
veyed. In certain places it has seemed desirable to enlarge somewhat
on certain things which were said by the writer. This bulletin there-
fore can not be called a verbatim report, but it represents nevertheless
the proceedings of the meeting.

It is hoped that similar meetings may be held in the future.

EE. Pairs;
In charge of Apiculture.

=yUy}

CONE NEES:

7

PAPERS.
Page.
The Bacteriology of Bee Diseases________-_____ _---~_______ G. F. White__ 10
The Present Status of the Investigation of Bee Diseases___H. F. Phillips__ 22
Apiary Inspection in New York State__.________________¢ has. Stewart__ 5)
’ American Foul Brood on the Pacific Coast_____________ Fred A. Parker__ a7
The History of Bee-Disease Inspection in Wisconsin ______. N. EH. France__ 73
TOPICS OF DISCUSSION.
Syl LOMIUS Toke WECRMISEa Segre = Soe: ty Nahe ea en a et ey mea ne 18
CEO ar ENT LG eae ae LSS Ge Ute Tes ee ee ae ra Of ee eo ee 19
tia mIBOMU Ol CIS CASO Ss 528 oF eSe ye de ee a 20
BERNICE CG GLO Free AE OTM Gone ae ee a ane Set gle Arm eee dah iy ag ee ear ue ae 2
VET REVE TT Voy Me COM ge bs eA ye ey eee ek pe eB Ne a IER pea Ca cele 1 ary pata aad ec ee ATU Sei ee ae 20
IBRSKERM ee is ye Nel (0 N OVELE) OVS eee is manta ala epee ht eee ey Be AGREE eee p15) Ll 21
Sal BURT Leal ON ELOLONG Las ee anh Se ies red BAIS LN Senet Cael Sev RR ona Sea og Ae Et 21
me CHoneGaLried lO OWECLSe a see ya ee a ye aes Se oe SES 23
Bas eAUUpT fey coh Osta STO COTES SS pes etc ere DP Eh eS ae Wake See a i ee Ps
PADI GA GlONSKONPW Ce CI SGAISES ae = a peas Se ee eT le ee J ee ee Dey
Existence of American foul brood and European foul brood in the same
EO) | TN ae cere Beery oe ie ot ls Seen ae ee eee Gh g Ue ee ee De
Geographical distribution of American foul brood______________________ 53
MMe cEeoO ha ChimatenG Ms valbnlen Cease sa eis ses SE eco eo ie ea eyes. 3 54
ASSOCIA MONGOfANSPeClOrs) Ol play en = ee Le ee eS 54
eRe AM en burOTAW CC) CISCASCGs oss nen Le ee eg ae 61
Hireciwot requeenino-OnsdiSeases = — sss se a ee oe 6
VIS eatin @ Tate eer ee cea SS are ne ore Sie ae ee end NO 69
PR cian terial ete ean Mahe gta ena SEAMED: Meee irk seis Bato? 70
ESPEN Ses Ole WOMMeMites 925 > aa ern as Le ie eka SS ae ia
Boilineshoneyatromadusea sed colomiesese ic 2a sae eee 72
INET CT 2 ee tain opened app eee Be Nay 2 men ee ae eee ghee NEI ig eS th OU
LEBUST RA TION:
Page.
PuaTE I. Portion of comb from colony infected with American foul brood
Frontispiece.

REPORT OF THE MEETING OF INSPECTORS OF APIARIES, SAN
ANTONIO, TEX., NOVEMBER 12, 1906.

The meeting was called to order in Market Hall at 9.30 a. m. by

Dr. E. F. Phillips, of the Bureau of Entomology, Department of
Agriculture, Washington, D. C., who addressed the members as fol-
lows:
_ Lapies anp GENTLEMEN: Last summer it was my pleasure and
privilege to visit a considerable number of the men at work on bee-
disease inspection throughout the United States. After talking with
them and going with them on inspection trips, it became evident that
there is a great deal yet to be done in regard to the making of better
methods of inspection in work against bee diseases, entirely apart
Trom the scientific aspect of the subject. .

On the 3d of last August Mr. N. E. France, inspector of apiaries
for Wisconsin, who is attending this meeting, and Mr. W. Z. Hutchin-
con, the inspector of apiaries for Michigan, and myself met in Mil-
waukee, Wis., to talk over certain plans for making bee-disease in-
spection more effective. A circular letter, addressed to the persons
now acting as inspectors in the various counties and States of the
United States, was drawn up, suggesting that they meet in some place
this fall to take up the work of bringing about more uniformity in
the methods employed. The meeting of to-day is, the result of that
circular letter.

There are several things in work against bee diseases that are not
at all clear to the bee-disease inspectors and others interested in the
subject, and we wish to take some of them up for discussion to-day.
We have with us Dr. G. F. White, of the Bureau of Animal Industry,
U.S. Department of Agriculture, Washington, D. C., who has made
» most thorough scientific study of the cause of bee diseases, and we
shall first ask him to give his demonstrations and the result of his
work. After he finishes I shall myself attempt to summarize the
investigations of bee diseases which have previously been made by
different men. It is perhaps well to take up the scientific aspect of
this work first in order to make this clear, and later we shall take up
_the methods of inspection and treatment and the form of desirable
Jaws controlling bee diseases.

9

10 MEETING OF INSPECTORS OF APIARIES.

Ti would perhaps be well for the inspectors of apiaries in the
United States to be organized in some way so that there might be
greater uniformity in the work and more harmonious cooperation.
In Butfalo several years ago such an association was organized. At
that time Mr. France, Mr. Hutchinson, and several others met to-
gether and organized an association of bee inspectors of the United
States and Canada, but they had only an organization meeting and
have never met since. Such an organization is desirable, but per-
haps not all of those that would care to take part in the association
are present.

Doctor White will now give us a demonstration on the brood dis-
eases of bees, a subject on which he has made exhaustive studies.

THE BACTERIOLOGY OF BEE DISEASES.¢

By Goh} Wirt eho
Of the U. S. Department of Agriculture.

The object of this paper is to discuss briefly the science of bacteri-
ology as it is used in the study of bee diseases and to give a summary
of the results of my work on these diseased conditions.

In our discussion of bacteriology, or the science which deals with
bacteria, it may be well to consider the subject under the following
headings: (1) The nature of bacteria; (2) their distribution; (3) the
methods for studying them, and (4) the results of their activity.

THE NATURE OF BACTERIA.

Bacteria are considered by some scientists to be a form of life
lower than either animals or plants, but by the majority of authors
they are looked upon as plants, and we shall so consider them at this
time without going into a detailed discussion of the arguments in
favor of such a view. Bacteria, often referred to as germs, microbes,
or parasites, ave, then, very small plants, so small indeed that they
must be magnified 600 diameters or more before they can be seen.

a At the meeting of the inspectors of apiaries Doctor White gave a demon-
stration of the work which he has done on the brood diseases of bees, showing,
in illustration of his address, slides and cultures of the various bacteria under
consideration. In view of this fact the stenographic report of his address is not
clear on all points, since the demonstrations are lacking. It has, therefore,
seemed best for Doctor White to write the article here published, giving a
popular discussion of this phase of the work which would ‘be intelligible without
the demonstration. The substance of his remarks is all included in this paper
except the part pertaining to the work of other bacteriologists, which is dwelt
on at some length in the article herein published on ‘‘ The Present Status of
3ee Disease Investigation.”” This method of handling the subject in a published
report will make the subject much clearer to those who did not attend the
ineeting at San Antonio.—H. F. P. :

THE BACTERIOLOGY OF BEE DISEASES. 11

These plants, then, constitute an invisible flora, which we can see only
by the use of a microscope of very high magnifying power. The
morphology, or the structure, including form and size, is principally
of two types, rod-shaped or cylindrical and round or spherical. The
size of bacteria varies. Those which are rod-shaped usually measure
from 1 to 3-6 microns in length and from one-half to 1 micron in
diameter. A micron is the unit of measure for very small objects
and is equivalent to g5}55 of an inch. For example, if a single bac-
terium of the rod form measures 2 microns, it would take 12,500
placed end to end to measure | inch in Jength. The spherical bacteria
or cocci have about the same diameter as the rod-shaped ones.

Bacteria grow or multiply after a manner called fission; that 1s,
after increasing in size they become constricted in the middle, which
constriction finally severs the rod completely, and we then have two
bacteria where there was but one before. Undgr favorable conditions
for growth, each bacterium divides by fission every twenty minutes,
or, in other words, gives rise to three generations in one hour. Such
being their marvelous rate of increase, a little calculation will demon-
strate that countless millions may be formed in a short time under
favorable conditions, which are proper temperature, moisture, food
in correct proportions, and the absence of much light. The tempera-
ture most favorable for the growth of a species of bacteria which is
able to produce a diseased condition in animals is approximately the
temperature of the animal which is affected by such species. Mois-
ture is universally necessary. The food must not be too concentrated.
Light inhibits the growth of bacteria. Direct sunlight is bacteri-
eidal; that is, it kills bacteria.

Many species produce spores when the conditions are not favorable
for the multipheation of bacteria. These are small bodies formed in
the bacteria (probably never more than one in a single bacterium)
which are somewhat comparable to the grain in wheat and corn.
These spores constitute a resting stage and usually also a very resist-
ant stage, for high temperature and strong disinfectant solutions are
necessary to kill them. It is these spores which probably make the
control of the bee diseases more difficult. When the spores again gain
access to a suitable “soil,” for example, the body of an animal, they
germinate and a new growth takes place as before.

Many species of bacteria have the power to move when they are in
a liquid medium, while others do not. This ability to move is due to
long, slender processes, which we call flagella, extending from the
body of the bacterium.

THE DISTRIBUTION OF BACTERIA.

Bacteria are very widely distributed. Everyone is familiar with
the very wide distribution of the higher members of the plant king-

i MEETING OF INSPECTORS OF APIARIES.

dom, as the trees, shrubs, flowers, and grasses, and we may for con-
venience refer to these plants as the visible flora. There is also an
invisible flora, made up of the plants we can not see except with the
aid of the miscrope. This flora includes the very minute plants:
referred to as bacteria, and also the yeasts and some fungi. The dis-
tinct species of plants which belong to the invisible flora outnumber
by far those which are visible to the naked eye. These microscopic
plants are found upon the surface of the animal body and along the
digestive tract; they are found in the soil, in the food we eat, and in
the water and milk we drink, but are not found within the normal
tissues of animals and higher plants.

HOW BACTERIA ARE STUDIED.

The morphology or structure of bacteria is studied with the aid of
a microscope of high*magnification. Since the number of distinct
species of bacteria is so extremely large, and since the shapes assumed
by them are so few, it is obvious that many different kinds must look
alike under the microscope. This is a point of considerable value in
connection with bee-disease work, since in some cases attempts have
been made by the use of the microscope alone to determine what
species of bacteria was causing certain diseased conditions. With
our present knowledge it is not possible to make a positive diagnosis
of these diseases with the microscope alone. With the microscope we
are able to determine usually only the genus to which any bacterium
belongs. If we are trying to identify Bacillus alvei, for example, we
are able with the microscope alone to say only that it is a Bacillus,
since it is seen to be a straight rod. Some other means is necessary to
determine the species (a/ve/) to which it belongs. For this purpose
we use artificial media or “ soils” in which pure cultures of the bac-
teria are inoculated or planted.

The media in common use are bouillon and sugar-free bouillon,
gelatin, agar, and sugar-free bouillon to which has been added small
amounts of various sugars known in chemistry as glucose, lactose,
saccharose, maltose, and levulose. In addition to these media, use is
made of potato, milk, and milk to which litmus has been added, so
that the reaction—whether acid, alkaline, or neutral—may be noted.
The bouillon is prepared from beef juice to which some peptone and
salt are added. Sugar-free bouillon is similar, except that the muscle
sugar has been removed. Gelatin is made from pure sheets of gel-
atin somewhat similar to that used in cooking, to which bouillon is
added. The bouillon affords the food for the bacteria or other small
plants, while the gelatin keeps the medium solid at ordinary tempera-
tures. Agar-agar (or simply agar) is the dried stem of a certain sea-
weed which liquefies on heating; to this is added bouillon, as.in the

THE BACTERIOLOGY OF BEE DISEASES. 30)

vase of gelatin, which solution congeals, as does gelatin, on cooling.
The milk used is cow’s milk with the butter-fat removed. Before
using, all these media are sterilized by heat to kill all bacteria or fungi
which might be present.

Having prepared these soils in this way, before the inoculation of
them the bacteria must be obtained in pure culture. By pure culture
is meant the growth of one species only in a medium. Such a cul-
ture is obtained by diluting a small quantity of the material, e.g,
decayed larve, containing the bacteria with a relatively large amount
of liquefied agar, and then pouring it into a shallow sterile glass
box (Petri dish). In this way we get only a few bacteria scat-
tered throughout a thin layer of the medium. Each bacterium then
begins to grow, and after a few hours it has produced a large num-
ber, which, being massed together, we are able to see with the naked
eye. This mass of bacteria, having been produced from one individ-
ual, constitutes a colony, and such a colony can contain but one spe-
cies, therefore we speak of it as pure. Pure cultures are then made
by inoculations from such a colony. The next step is to identfy this
species which we now have isolated from all other species. To do
this we inoculate a few or all of the differential media mentioned
above. After inoculating and growing the bacteria in these different
media or soils at about the body temperature for a day or longer we
observe the effect upon the various media produced by the growth of
the bacteria and the appearance of the growth in or upon these media.
All these phenomena and appearances we speak of as cultural char-
acters. Having obtained in this way the cultural characters of a spe-
cies of bacteria, we are able to classify it by comparing these cultural
characters with the cultural characters of known species.

To illustrate this, let us take for example Bacillus coli communis,
found normally in the intestine of man and many animals, including
the intestine of the adult bee, Bacillus alvei, found in European
foul brood, and Bacillus larva, found in American foul brood.
Bacillus coli communis by its growth in bouillon causes the latter to
become heavily clouded; Bacillus alvei makes it feebly clouded;
while Bacillus larvae does not grow at all in this soil and the bouillon
remains clear. In gelatin Bacillus coli communis grows very well
and does not liquefy the medium by its growth; Bacillus alvei grows
very slowly and only feebly and liquefies the gelatin; while Bacillus
larve does not grow at all in this medium. When Bacillus coli com-
munis is planted on potato it produces a brownish growth; Bacillus
alvei on this medium produces a lemon-yellow growth, and Bacil-
lus larve fails to show any growth. When Bacillus coli communis
is planted in milk there follows a rapid souring of the milk and a
firm coagulation of the casein; Bacillus alvei produces a soft coagu-

14 MEETING OF INSPECTORS OF APIARIES.

lum which is followed by a slow digestion or liquefaction of the
easein; Bacillus larve does not grow in milk. In litmus milk,
Bacillus coli communis produces a large amount of acid, which is
indicated by the change of the litmus to the red color; Bacillus alvei
produces no marked change in reaction, and Bacillus larve does
not grow in this medium. In the bouillons to which the sugars,
glucose, lactose, saccharose, ete., have been added, there is produced
by the growth of Bacillus coli communis, gas, and a large amount of
acid; Bacillus alvei does not produce gas and only a small amount
of acid by its growth in the media containing sugars, while Bacillus
larve does not grow when planted in these “soils.” (I shall
speak later of a medium upon which Bacillus larve will grow.)

It is by these differences which we observe in the growth upon the
various media and the effect produced upon the different media by
the growth of the bacteria that we are able to determine one species
of bacteria from another.

THE RESULTS OF THE ACTIVITY GF BACTERIA.

In the consideration of this question it is convenient to divide the
bacterial flora into two groups—nonpathogenic, or those which do not
produce disease, and pathogenic, or those which do produce disease.
Some of the nonpathogenic bacteria are economically very important
as scavengers. The bodies of dead animals and plants are largely
brought to decay by them. The flavors of butter, cheese, and wines
are thought to be improved by the growth of bacteria or other micro-
organisms—the fungi and the yeasts. Others of these micro-organ-
isms ruin the food, causing the souring of milk, the spoiling of fruit,
etc. Many diseases in man and animals are known to be caused by
bacteria, as tuberculosis, diphtheria, glanders, and anthrax.

I wish now to speak briefly of how bacteriology has been used in
the study of bee diseases, and to summarize the results which have
been obtained. For a more detailed account you are referred to a
bulletin issued by the Bureau of Entomology of the United States
Department of Agriculture—Technical Series, No. 14, * The Bacteria
of the Apiary, with Special Reference to Bee Diseases,” issued
November 6, 1906.

From what has been said one would naturally infer that in every
apiary, whether diseased or not, there are en the hives, combs, and
bees a large number of bacteria that are perfectly harmless. If one
is trying to find in a diseased apiary the species of bacteria which is
the probable cause of the trouble, what is the method of procedure ?
Suppose there were two herds of cattle on adjoining farms and the
cattle on one farm were dying while those on the other remained well.
If it were suspected that some plant which the cattle were ating wasthe

THE BACTERIOLOGY OF BEE DISEASES. 15

‘ause of death, naturally the plant would be selected which was
found on the farm where the animals were sick and which was not
found on the farm where the animals. remained well. This is
exactly the kind of reasoning used when we are looking for the
bacteria which are causing the diseases among bees. This neces-
sitates, as you see, the study of all the bacteria which are present in
any apiary, whether diseased or not, as well as those in diseased
aplaries.

At the time we began the work on bee diseases, in June, 1902, the
disorders which were causing the greatest trouble were known to
bee keepers as black brood, foul brood, pickle brood, and paralysis.
After the study of a large number of samples of brood affected by
disease which was being called black brood and the finding of
Bacillus alvei in all of them, it is very clear that this disease is
the same as that investigated by Cheyne in 1885 and called by him
“foul brood;” he first described Bacillus alvei. “ Black brood” was
a name given by Dr. William R. Howard, of Fort Worth, Tex., to a
disease which he thought existed in New York State, and he described
as its cause Bacillus milii. After a careful search in New York State
for a disease containing Bacillus milii we were unable to find it, and
there seems to be. no ground for the description of a new disease.
What has been called black brood by Doctor Howard is obviously the
type of foul brood which we now distinguish as European foul brood.

In the decaying larve and dried scales found in the cells in the
disease which was receiving the name of foul brood there were seen
by the use of the microscope a very large number of the spores of bac-
teria, and in the larve in the early stage of the disease there were
observed bacteria in the rod form. When these spores were planted
upon the media or soils which have been explained earlier in this
paper, they would not grow. It became necessary, then, to devise a
soil in which the growth could be obtained. After a number of un-
successful attempts, a medium or soil was made from healthy bee
larve in which the spores would germinate and the bacteria would
grow. By astudy of this species, which was found in the dead larve
of this disease and which was not found in the healthy apiary, it was
evident that it was not Bacillus alvei, and, since Bacillus alvei is not
present at all, we know that this disease is not the foul brood which
Cheyne had reported in his work in 1885. Since it is not this type of
foul brood, what could it be? By carefully reviewing all the work
which had been done by others, the conclusion was inevitable that this
diseased condition had not been described properly from a bacterial
standpoint as a disease separate and distinct. from the foul brood of
Cheyne, but that the mistake had been made for a long time of calling
two different and distinct diseases which affected the brood of bees
by one name. This condition was reported to the New York State

16 MEETING OF INSPECTORS OF APIARIES.
6

department of agriculture in a report to that department made in
January, 1908, and another in January, 1904. In the latter report
this condition, for want of definite information, was referred to as
“ X brood” and the bacillus es Bacillus X. At the suggestion of
Dr. E. F. Phillips, of the Bureau of Entomology, United States
Department of Agriculture, it was thought best for very good rea-
sons to retain the name foul brood in the name of each disease and
add a qualifying word to designate the difference between the two
diseases. ‘ European ” is added to foul brood to designate the disease
which Cheyne studied in England (Europe) in 1885, and “American ”
is added to the foul brood which was first studied in the United States
(America). We distinguish, then, European foul brood and Ameri-
‘an foul brood. Both of these diseases of the brood of bees seem to
be found in Europe as well as America. It must therefore be remem-
bered that these names do not put any stigma on either country,
Europe or America, but, on the contrary, Europe is thereby given
the credit of having first studied the European foul brood and
America for having first studied American foul brood.

In a study of the so-called * pickle brood ” we are unable to sug-
gest from a bacteriological standpoint any cause for the disease. A
study has been made of the bacteria found upon the healthy adult bee
and those found in the intestine and also the bacteria found upon and
within the adult bees suffering with palsy or paralysis, but so far no
suggestion can be made from a bacteriological standpoint as to the
cause of this disorder of the adult bee.

To conclude, I shall read, with your permission, the summary of
the work reported in the bulletin referred to above.‘

SUMMARY TO PART I.

The results of the study of the bacteria found normally in the apiary may be
briefly Summarized as follows: f

(1) The temperature of the hive approximates that of warm-blooded animals.

(2) Upon adult bees and upon the comb there occurs quite constantly a species
of bacterin which we refer to in this paper as Bacillus A, and which, it is
helieved, is the organism that some workers have confused with Bacillus alvei
which is universally present in European foul brood.

(3) There cceurs very constantly in the pollen and intestine of adult bees a
species here referred to as Bacillus B.

(4) From the combs Bacterium cyaneus, Saccharomyces roseus, and a Micro-
coccus referred to here as Micrococcus C, have been isolated and studied.

(5) Toney from a healthy hive is, as a rule, sterile.

(6) The normal larvire are, as a rule, sterile.

(7) There is an anaérobe found quite constantly in the intestine of the
healthy honey bee. It is referred to in this paper as Bacterium D.

(8) From the intestine there have been isolated and studied the following
micro-organisms: Bacillus cloace, Bacillus coli communis, Bacillus cholera suis,

a'Technical Series, No. 14, Bur, Ent., U. 8. Dept. Agric,

THE BACTERIOLOGY OF BEE DISEASES. 17

Bacillus subgastricus, Bacterium mycoides, Pseudomonas fluorescens liquefaciens,
and two referred to as Bacillus EH, and Saccharomyces F'. Others less frequently
present have been isolated, but not studied.

(9) In two samples of brood with unknown disease there was found a species
of yeast plant here referred to as Saccharomyces G.

SUMMARY TO PART II.

Following is a brief summary of the results of the present investigation of bee
diseases :

(1) There are a number of diseased conditions which affect the apiary.

(2) The disease which seems to cause the most rapid loss to the apiarist is
European foul brood, in which is found Bacillus alvei—first isolated, studied,
and named by Cheshire and Cheyne in 1885.

(3) The distribution of Bacillus alvei in the infected hive is as follows:

(a) The greatest number of infecting germs are found in the bodies of dead
larve.

(bo) The pollen stored in the cells of the foui-brood combs contains many of
these infecting organisms.

(c) The honey stored in brood combs infected with this disease has been
found to contain a few bacilli of this species.

(d@) The surface of combs, frames, and hives may be contaminated.

¢e) The wings, head, legs, thorax, abdomen, and intestinal contents of adult
bees were found to be contaminated with Bacillus alvei.

(f) Bacillus alvei may appear in cultures made from the ovary of queens
from European foul-brood colonies, but the presence of this species suggests
contamination from the body of the queen while the cultures are being made
and has no special significance.

(4) The disease which seems to be most widespread in the United States we
have called American foul brood, and the organism which has been found con-
stantly present in the disease we have called Bacillus larve. This disorder
was thought by many in this country and other countries as well to be the foul
brood described by Cheshire and Cheyne, but such is not the case.

(5) From the nature of American foul brood it is thought that the organism
has a similar distribution to that of Bacillus alvei.

(6) It appears that Huropean foul brood was erroneously called *“‘ New York
bee disease ’ or “ black brood” by Dr. William R. Howard in 1900.

(7) There is a diseased condition affecting the brood of bees which is being
called by the bee keepers “ pickle brood.” No conclusion can be drawn from
the investigation so far as to the cause of the disease.

(8) Aspergillus pollinis, ascribed by Dr. William R. Howard as the cause of
pickle brood, has not been found in this investigation and is not believed by the
author to have any etiological relation to the so-called ‘“‘pickle brood.”

(9) Palsy or paralysis is a diseased condition of the adult bees. No con-
clusion can yet be drawn as to its cause.

(10) Formaldehyde gas as ordinarily used in the apiaries is insufficient to
insure complete disinfection.

CONCLUSIONS.

In a paragraph the author wishes, if possible, to present the status of the bee
diseases in this country. It should be remembered, firstly, that ‘ black brood ”’
can now be dropped from our vocabulary, and probably does not exist ; secondly,
that the term ‘‘ foul brood”’ was being applied to two distinct diseases. One of
these diseases we now refer to as European foul brood, because it first received

30547—No. 70—07 m——2

18 MEETING OF INSPECTORS OF APIARIES.

a scientific study from a European investigator. We refer to the other disease
as American foul brood, because it was first studied scientificaily in America.
There is one more disorder in the brood of bees which has attracted considerable
attention—the so-called ‘ pickle brood.” There are, then, these three principal
diseases: European foul brood, American foul brood, and the so-called “ pickle
brood.”

Doctor Puiuies. We surely have all been glad to listen to Doctor
White in his most interesting account of his work. It will be well at
this time to ask him any questions concerning this work which may
have come to mind. Before opening this subject for discussion I
wish to say that after this discussion I shall take up in detail the
works which Doctor White has criticized. Consequently, if you have
no objection, we will hold over until later any discussion of these
papers.

Mr. C. P. Dapant. As I understand it, there exist these two bacilli
(Bacillus alvei and Bacillus larve) and also Bacillus mesentericus
vulgatus. Wave you samples of all three of the bacilli?

Doctor Wurre. Yes, sir; that (pointing to slides) is Bacillus lar-
ve, that (showing cultures) is the Bacillus alvei, and the next, Bac7-
lus mesentericus. There are a number of varieties of Bacillus mesen-
tericus, and vulgatus is one of them.

SYMPTOMS OF BEE DISEASES.

Mr. Dapant. Will you please give us a description of the two dis-
eases—that is, of the conditions arising when Bacillus larve and
Bacillus alvei are present in the combs?

Doctor Wutre. I should like to ask Doctor Phillips to answer that
question.

Doctor Puimups. I shall simply quote from Doctor White’s bulle-
tin. There was issued from the Bureau of Entomology some time
ago a small circular, Circular No. 79, entitled “ The Brood Diseases
of Bees,” and in this circular was included a description of the two
diseases which Doctor White has been studying. Doctor White was
kind enough to quote in his bulletin from Circular No. 79, and I shall
read the descriptions.

AMERICAN FOUL BROOD.

American foul brood (often called simply “foul brood’) is distributed
through all parts of the United States, and from the symptoms published in
European journals and texts one is led to believe that it is also the prevalent
brood disease in Europe. Although it is found in almost all sections of the
United States, there are many localities entirely free from disease of any kind.

The adult bees of an infected colony are usually rather inactive and do little
toward cleaning out infected material. When the larve are first affected, they
turn to a light chocolate color, and in the advanced stages of decay they become
darker, resembling roasted coffee in color. Usually the larvee are attacked at
about the time of capping, and most of the cells containing infected larvee are

GEOGRAPHICAL DISTRIBUTION 19

capped. As decay proceeds, these cappings become sunken and perforated, and,
as the healthy brood emerges, the comb shows the scattered cells containing
larvee which have died of disease, still capped. The most noticeable charac-
teristic of this infection is the fact that when a small stick is inserted in a
larva which has died of the disease, and slowly removed, the broken-down
tissues adhere to it and will often stretch out for several inches before break-
ing. When the larva dries, it forms a tightly adhering scale of very dark-brown
color, which can best be observed when the comb is held so that a bright light
strikes the lower side wall. Decaying larvee which have died of this disease
have a very characteristic odor, which resembles a poor quality of glue. This
disease seldom attacks drone or queen laryiee. It appears to be much more
virulent in the western part of the United States than in the East.

EUROPEAN FOUL BROOD.

European foul brood (often called “ black brood”) is not nearly as widespread
in the United States as is American foul brood, but in certain parts of the coun-
try it has caused enormous losses. It is steadily on the increase and is con-
stantly being reported from new localities. It is therefore desirable that bee
keepers be on the watch for it.

Adult bees in infected colonies are not very active, but do succeed in cleaning
out some of the dried scales. This disease attacks larvee earlier than does
American foul brood, and a comparatively small percentage of the diseased
brood is ever capped. The diseased laryvee which are capped over have sunken
and perforated cappings. The larvae when first attacked show a small yellow
spot on the body near the head and move uneasily in the cell. When death
occurs, they turn yellow, then brown, and finally almost black. Decaying larvze
which have died of this disease do not usually stretch out in a long thread when
a small stick is inserted and slowly removed. Occasionally there is a very
slight “ ropiness,” but this is never very marked. The thoroughly dried laryie
form irregular scales which are not strongly adherent to the lower side wall of
the cell. There is very little odor from decaying laryze which have died from
this disease, and when an odor is noticeable it is not the “ glue-pot ” odor of the
American foul brood, but more nearly resembles that of soured dead brood.
This disease attacks drone and queen larvee very soon after the colony is in-
fected. It is as a rule much more infectious than American foul brood and
spreads more rapidly. On the other hand, it sometimes happens that the disease
will disappear of its own accord, a thing which the author never knew to occur
ina genuine case of American foul brood. European foul brood is most destruc-
tive during the spring aud early summer, often almost disappearing in late
stimmer and autumn.

GEOGRAPHICAL DISTRIBUTION.

Mr. France (Wisconsin). When I was with Doctor Phillips and
Inspector Hutchinson in Michigan studying the difference between
American and European foul brood, it occurred to me that it was
possible to bring together at this time specimens of diseased brood
from different localities. In my own city (Platteville, Wis.) I found
samples of diseased comb and had reserved them for this meeting, but
unfortunately three of the four samples in my possession contained
moth larvee, and it was impossible to tell anything about the disease.
The only one that I still have is now in my grip.

Mr. Dapanr. Where were those samples from /

20 MEETING OF INSPECTORS OF APIARIES

Mr. France. From Michigan.

Doctor Puinuirs. Our first accurate knowledge of European foul
brood in the United States was the epidemic in New York State, and
most bee keepers still look on the disease as still being confined to that
State. However, European foul brood is now found in New York,
Vermont, Massachusetts, Connecticut, New Jersey, Pennsylvania,
Ohio, West Virginia, Michigan, Indiana, and Illinois. Reports have
been received at the Bureau of Entomology from all those States.
The disease is rapidly going west.

COMPARISON OF DISEASES.

Mr. Witu1am Arcuiery (Texas). Which of the two diseases is con-
sidered worse, the American or the European foul brood ?

Doctor Puiturs. That is a point which was simply suggested in
the descriptions just read. European foul brood spreads more rap-
idly than the other, but at the same time it will at times absolutely
disappear of its own accord, which is something that the American
foul brood seems not to do. We have these two factors over against
each other, and I should as soon try to eradicate one disease as the
other. As far as loss is concerned there seems to be very little differ-
ence,

Mr. J. Q. Smirn. (Tllinois). How many specimens have been sent to
the Bureau of Entomology from Illinois?

Doctor Puiiurrs. Two, I believe.

Mr. Smitri. I was in correspondence with some or the persons hav-
ing this disease among their bees, and I advised them to send samples
to you.

INFECTION IN HONEY.

Mr. R. A. Hotexamp (Missouri). Have the bacteria of both dis-
vases been found in honey from infected hives 4

Doctor Wurrr. We believe that the infecting agent may be present
in honey in each case, since the experience of the bee keeper has been
that the infection of a healthy colony has followed the feeding of
honey from hives affected by either disease.

INFECTION OF LARVAE.

Mr. A. H. Anprrson (Utah). How does the larva become infected ?

Doctor Pumps. The manner by which this is brought about is not
all entirely certain, but the facts would tend to show that it is through
feeding on infected material.

Doctor Wuire. It would appear that internally the contamination
is caused by being carried by worker bees. As they move over the
combs and clean out the cells they come in contact with contaminated
material. On any part of the body one can find Bacillus alvei; so I
shall leave it to you, as you are more conversant with bees.

INFECTION CARRIED TO FLOWERS. D1

Doctor Pumps. The bacteria in a diseased colony are present
everywhere. They are found all over the adult bees, on the queens, on
the outside of the comb, and every place else. They do not, however,
grow in honey; they quickly go from the rod condition to the spore
condition and remain in the latter conditien indefinitely when in
honey. According to the statement just made, it would seem that a
bee from an infected hive would always carry disease. The fact is,
however, that if the bees have been away from this infection for
some time they will not transmit the disease. Give them a new clean
hive with no food, so that all the honey is used up from the inside
of the body. The infection from the outside does not seem to spread
the disease if no brood is reared for a few days.

Mr. Suiri. I believe that the reason why the bee loses the infection
is because a certain time elapses before the comb is drawn out and
young larve are present which are large enough to become infected.
But I know this fact: If you shake bees from a diseased colony onto
combs that contain healthy larvee, you might as well leave the larve,
for disease at once appears. I have tried that.

BACTERIA IN QUEENS.

>

Mr. Dapanr. In either case have the bodies of queens been in-
spected ?

Doctor Wuirr. The bodies of queens have been inspected, and while
the internal organs contained these organisms, the ovaries seldom do,
and where Bacillus alvei is found in the ovary, or in our cultures
made from the ovary, they occur very seldom, and the probability is
that they get there through contamination in making the cultures
‘ather than from being found in the ovary itself. The ovaries are
very small and one must work with instruments that are sufficiently
large to handle. It is almost impossible to take cultures from the
ovary and not get contamination from the outside.

(74 = sa ”
BLACK BROOD.

Mr. H. H. Roor (Ohio). I thought I understood Doctor White to
say that the disease called black brood has not been found in New
York.

Doctor Puimuires. What Doctor White said was that there is no such
thing as black brood. The name black brood was a blunder.

INFECTION CARRIED TO FLOWERS.

Mr. Smrru. Then if, as you say, the contamination is always
present on the adult bees from diseased colonies, why is it not pos-
sible to carry it to the blossoms and leave it on the pollen, so that the
next bee visiting the same flower would carry germs to its hive?

Doctor Puruures. It is, of course, possible, but highly improbable.

22 MEETING OF INSPECTORS OF APIARIES.
VITALITY OF SPORES.

Mr. France. As to the duration of this bacillus in the spore form,
how long can it remain in honey and still have vitality to grow under
proper conditions ? |

Doctor Puruuies. I have never determined any limit. It is known
that very old honey from an infected hive will transmit disease to a
colony.

Mr. France. In my State (Wisconsin) we had an experience bear-
ing on this point where combs contained American foul brood. The
bees had died, leaving the combs containing dead dried-up larvee, and
the owner, anxious to start in bees again, put the hives away in the
granary, and four years afterwards hived bees on them and American |
foul brood started anew.

Mr. D. H. Cocesuari (N. Y.). When honey was shipped from |
Cuba several years ago and scattered all over the United States, if |
it was left where bees from this country could get to it, the disease
was certainly scattered broadeast.

|
;

PUBLICATIONS ON BEE DISEASES.

Doctor Prixiips. I desire at this time to announce the publication
of three pamphlets on bee diseases issued by the Bureau of Ento-
mology. Circular No. 79,“ The Brood Diseases of Bees,” was issued |
three or four weeks ago. I have just this morning received copies |
of a paper by Doctor White, Technical Series, No. 14, entitled, ‘* The |
Bacteria of the Apiary, with Special Reference to Bee Diseases.”
This was issued on November 6, and was received here this morning.
T have also a pamphlet here from the Bureau of Entomology con-
taining all the laws in force relative to bee-disease inspection. This
is a reprint from Bulletin No. 61, The Laws in Force Against
Injurious Insects and Foul Brood in the United States.” A recent
order of the Secretary of Agriculture has put a stop to the free
distribution of bulletins, but they can be purchased from the Superin-
tendent of Documents, Government Printing Office, Washington,
D.C. The price of Technical Series, No. i4, is 10 cents.

The following paper was then read by Doctor Phillips:

THE PRESENT STATUS OF THE INVESTIGATION OF BEE DISEASES.

By BE. EF. Puts, Ph D.,

Of the U. S. Departinent of Agriculture.

I wish to take up two or three phases of this work about which
Doctor White has been speaking, and te add some additional points.
In regard to the history of the investigation of bee diseases prior to |
1885, I can do no better than to quote the historical résumé contained

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 23

in a paper by Prof. F. C. Harrison, entitled “ Foul Brood of Bees,”
published as Bulletin 112 of the Ontario Agricultural College:

In all probability the first definite reference to foul brood is by Aristotle
(Historia Animalium, Book IX, ch. 27), who mentions an inertness which
seizes the bees and causes a bad smell in the hive. He also suggests that bees
are liable to become diseased when the flowers on which they work are
attacked by blight. Bee dysentery as well as foul brood causes a bad odor ;
but in the former disease the spotting and consequent smell are usually outside
the hive.

Columella (De Re Rustica, Book IX, ch. 18) mentions a bee pestilence and
an annual distemper which seizes the bees in spring. Pliny (Natural History,
Book XI, ch. 19, A. D. 79) writes of a disease of bees, but as he uses the same
term as Aristotle he has probably copied it from the latter author.

Schirach (Histoire des Abeilles, ch. III, p. 56, La Haye, 1771), in 1769, was
the first writer to name the disease “foul brood.” He says:

It is dangerous and a most destructive disorder to the bees, a genuine plague
when the complaint has reached a certain stage. The cause can be attributed
to two sources: (1) The putrid (or tainted) food with which the bees feed
the larvee for lack of haying better. (2) By the mistake of the queen bee in
misplacing the larvee in their cells, head upside down. In this position the
young bee, unable to get out of its prison, dies and rots away.

Further, Schirach clearly distinguishes between foul brood and chilled brood,
gnd mentions the fact that putrefaction follows the death of the brood from
frost, but in this case “it is only an accident and not a disease.”

The remedy Schirach recommended was to remove all diseased combs from
the infected hives and to keep the bees fasting for two days, after which they
are furnished with other cakes of wax and a suitable remedy given, “as a
little hot water mixed with honey, nutmeg, and saffron, or a syrup composed
of equal parts of sugar and wine seasoned with nutmeg.” Thus, as Cowan
(Journal of the Royal Agricultural Society, Vol. VI, Part IV, 1895) remarks:
“We had given us nearly one hundred and thirty years ago a method of cure
almost identical with what is by some claimed as new to-day.”

Tessier (L’Eneyclopedie Methodique, Abeille, p. 52, 1765) about the same
time as Schirach says that when the lary die in their cells it Causes an in-
fection in the hive which makes the bees sick. It is then necessary to drive
away or sometimes move the bees from the hive, and to take care to fumigate
the infected hive if it is going to be used again. It is necessary, in order to
avoid the same inconvenience, to take out the parts of the comb that may be
moulded by reason of the dampness. Duchet (Culture des Abeilles, p. 315,
Vevey, 1771), who wrote on bees in 1771, does not mention any disease that
can be certified as foul brood, but he describes bee dysentery.

Della Rocea (Traite Complet sur les Abeilles, Vol. III, p. 261, Paris, 1790),
vieaire-general of Syra, an island in the Levant, relates with much detail the
history of an epidemic of foul brood, which caused great destruction in the
island during the years 1777 to 1780. Della Rocca describes very minutely the
symptoms, destruction, and mistakes that were made in attempting to combat
the disease. He says:

The disease manifests its presence by defects in the combs filled with brood,
and which only contain a putrid mass; instead of the bee pup there is only
rottenness in the cells, which, however, being capped, always preserve a healthy
appearance. If these cells are broken open, a blackish liquid flows out, which
spreads the infection through the hive. This disease only manifests itself
in cells which contain a nearly mature larva or a capped one. The bees them-
selves remain in good health, and work with the same activity, but their
numbers decrease daily. This disease, however, was not so general in a hive

‘

94 MEETING OF INSPECTORS OF APIARIES.

but that a small portion escaped. Some new “bees emerged, but in too small
numbers to supply the daily losses.’ Thus a hive attacked by this scourge will
perish from scarcity of population. At first it was not noticed that this disease
was epidemic, and the hives emptied by death of the bees were filled with fresh
swarms, and these contracted the same disease and perished. Yet another
mistake was made. The remains of the hives that were lost were taken into the
streets of the town to expose them to the sun in order to extract all the wax,
and the bees from the neighborhood sucked up the honey, caught the disease,
and communicated it to other hives, and all, without exception, perished in a
short time. The epidemic having reached the island spread everywhere, and
the mortality among the bees was general, either from eating infected honey, or
from stopping up the infected combs, or from the bees nourishing their brood
on infected honey.

Della Rocea criticises Schirach’s statement regarding the misplacement of
the larvee by the queen as a cause of the disease, because “ everybody knows
that the queen has nothing else to do but deposit eggs. These are then cared
for and nourished by the bees; and when the larva is nearly ready to change
into the pupa, the bees close the cell, and every position which is given the larva
depends on their good pleasure and not on the queen’s.”’ Della Rocca himself
thinks that ‘“‘some pestilential blight had without doubt corrupted the quality
of the honey and the dust from the anthers,” and recommends “ burning every-
thing without pity, as there is no other resource when the disease is well
established, as the pest is without doubt the most terrible in the natural history
of bees.”

Neither Wildman (Treatise on the Management of Bees, London, 1796), Keys
(Ancient Beemasters Farewell, London, 1796, Woolridge), Needham (Brussels
Memoirs, Vol. II, 1780, Rhein), Reaumur (Memoirs pour Servir a l’Histoire
Naturelle des Insectes, T. V., p. 1784), and other authors about the same time
(latter end of the eighteenth century) mention this disease.

Bevan (The Honey Bee, London, 1827) names the disease * pestilence,” and
also quotes Schirach’s name, “foul brood,’ and says regarding it that the
“pestilence has been attributed to the residence of dead larvze in the cells,
from a careless deposition of ova by the queen. * * * It has also been
attributed to cold and bad nursing; that is, feeding with unwholesome food.”

Nothing further of note appears in bee literature till the year 1860, when
Doctor Leuckhart (Bienen Zeitung, Hichstadt, 1860, p, 282) writes that he was
formerly of the opinion that foul brood was caused by the same fungus
(Panhistophyton ovatum) which is noticed in a disease of the silkworm, but
now, after observation and experiment, is quite certain that the disease is
caused by neither vegetable nor animal parasite. He also notes that the term
“foul brood” is applied to a number of diseases affecting bees.

Molitor Muhlfeld (Bienen Zeitung, Eichstadt, 1868, p. 95) recognizes two
forms, one contagious and the other not contagious, and thinks that the only
cause of contagious foul brood is a fly (Jehneuwmon apium mellificarium) which
lays its eggs on the young larvee of the bee.

A discovery of note was Preuss’s (Bienen Zeitung, 1868, p. 225), in 1868. He
contradicts Muhlfeld’s statement about the fly, and states that foul-brood cells
can be detected by the sunken cap. With a microscope magnifying 600 diam-
eters he found small, dust-like bodies, with a diameter of =}, mm., belonging to
the genus Cryptococcus (WKutzig), and called the specific form alvearis, likened
it to the fermentation fungus (Cryptococcus fermentum), and thought that the
last germ gained access to the young bee and changed to Cryptococcus alvearis.
He notices that many experts lay the cause of the disease to fermenting food,
but the larve are easily contaminated by the fermentation fungus, which is
always present in the air. He also mentipns the enormous rapidity with which

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 25

the fungus multiplies, and gives an elaborate calculation of the number that
might be found in a cell containing a deceased larva.

As might have been expected, Preuss’s statement aroused considerable dis-
cussion at the meeting of German bee-keepers a short while after the publication
of his paper.

Vogel (Bienen Zeitung, Nos. 21 and 22) expressed doubt as to whether
C. alvearis was the real cause of foul brood or only a consequence of the disease,
but on the whole agreed with Preuss.

Wiegand (Bienen Zeitung, Nos. 21 and 22) agreed with Preuss’s theory, and
in giving his experience said that the disease was introduced into his apiary
through honey brought from a distance.

Pollman (Bienen Zeitung, Nos. 21 and 22) believed that the disease was intro-
duced by feeding honey from Havannam, where, when extracting the honey,
both brood and honeycomb were mashed up and the honey pressed out.

Doctor Leuckhart (Bienen Zeitung, Nos. 21 and 22) agreed with those who
thought the disease due to a fungus, but discredited the supposition that it was
the same as the fermentation fungus mentioned by Preuss, and rather thought
it was related to the silkworm fungus and that most of the brood diseases
ending in death were called “foul brood,” while they were really something
else. He believed that the fungus was present in the eggs of the queen when
laid.

Geilen (Bienen Zeitung, Nos. 21 and 22) believed that the disease came from
the putrefying remains of animal bodies upon which the bees alighted,

Muhlfeld (Bienen Zeitung, 1869, No. 3) again, in 1869, presented his former
views and also those of Preuss and gave directions for maintaining the health
of bees. He recommended the boiling of the honey and a use of carbolic acid
in the strength of 1: 100, or permanganate of potash 1: 300, as disinfectants.

Lambrecht (Bienen Zeitung, 1870, No. 2.) thought that foul brood was caused
by fermentation of the bee bread.

Hallier (Bienen Zeitung, 1870, No. 2) considered it no specific disease, but
thought it was probably produced by different fungi.

Cornallia (Bienen Zeitung, 1870, No. 5) proved contrary to the above and
found a fungus which he thought developed foul brood. He called it Crypto-
coccus alvearis and used carbolic acid, potassium permag, and lime water as
disinfectants.

Fisher (Bienen Zeitung, 1871, pp. 105-125) advanced a new foul-brood theory
in 1871, which somewhat follows the view of Liebig regarding the silkworm
disease and plant diseases. According to this theory, the predisposing cause
was insufficient nourishment, especially short stores for winter and spring.
Shortage of pollen supply was the next cause. Fisher tried to prove his views
by the practical experience of bee keepers and explained that the first result of
repeated and continued feeding was an increase in the production of bees; and
a consequent disproportion between brood and brood feeders arose, which should
be looked upon as another cause of foul brood. The disease, he said, might be
lessened or exterminated by applying means to reduce the production of brood, as
the removing of the queen and the area which the brood occupied. Foul brood
is probably the cause of a quantitative dearth of nourishment and a consequent
degeneration of the bees. The appearance of fungus growths was only a see-
ondary matter.

Schonfeld (Bienen Zeitung, 1874, pp. 201 and 261) infected several hives
with foul brood, and when it had fully developed he took a comb of the rotten
brood to the Physiological Institute at Breslau and had it submitted to a micro-
scopical examination by Doctors Cohn and Hidam (Bienen Zeitung, 1874, pp. 201

26 MEETING OF INSPECTURS OF APIARIES.

and 261). This examination showed that in every dead larva and in each foul
brood cell, whether the contents were yet white and fluid or brown, tenacious,
and ropy, there were to be found long oval bodies, which Preuss called ‘“ micro-
cocci.” Close to and among them, Cohn was the first to find, with the most
powerful of the five microscopes that were used, a countless number of slender
pale rods, joined together, and which he at once identified as bacteria of the
genus Bacillus. The length of a single rod was about 6 micromillimeters, but
many of them were two and three jointed, so that these foul brood bacteria
microscopically resembled the anthrax bacteria, though of course they were
different physiologically and in the manner in which they acted as ferments.

It is not surprising when we remember the state of bacteriological knowledge
in 1870, that Preuss should have mistaken micrococeci for the spores of a bacillus.

In 1885 the first investigation which merited close attention was
published in the Journal of the Royal Microscopical Society, entitled
* The Pathogenic History and History under Cultivation of a New
Bacillus (B. alvez), the Cause of a Disease of the Hive Bee Hitherto
Known as Foul Brood,” by Frank R. Cheshire, F. R. M.S., F. L. S.,
and W. Watson Cheyne, M. B., F. R. C. S. One point is here to be
especially noticed, there were two authors of this paper. The paper
was divided into Part I, Pathogenic History, by Mr. Cheshire, and
Part II, History Under Cultivation, by Mr. Cheyne, and with the
latter part Mr. Cheshire had nothing whatever to do. Bee keepers
are generally giving Mr. Cheshire the credit for this work, but it is
clear that Mr. Cheyne, the man who did the bacteriological work,
should be the one to get the credit. The description of the disease,
contained in Part I, is as follows:

The nature of foul brood as a germ disease.—If a comb be removed from
near the center of a healthy hive during the summer months its cells will
normally be filled with eggs, larvee, and pupze in every stage of development.
The eggs as left by the ovipositor of the queen or mother adhere commonly by
the end to the base of the cells they occupy, and favored by the high tempera-
ture constantly maintained within the hive, the germinal vesicle at about three
days matures into a larva ready for hatching. These eggs I have shown as lia-
ble to the disease even before they leave the body of the mother, but most careful
microscopic examination is needful to make this apparent (and of which I
shall speak presently more particularly). On the contrary, the larvse, which
are constantly fed by the workers, so change in appearance soon after in-
fection that a practiced eye at once detects the presence of the disease. Whilst
healthy their bodies are of a beautiful pearly whiteness, lying, at first floating,
in the abundant pabulum the nurses are ever ready to supply. As they grow
they curl themselves at the bottom of the cells until they become too strait
for their occupants, which now advance to the head to be in readiness for the
cocoon spinning, which follows upon the close of the eating stage. When the
disease strikes the laryze they move uneasily in their cells, and often then pre-
sent the dorsal surface to its mouth, * * * so that mere posture is no insuffi-
cient evidence of an unhealthy condition. The color changes to yellow, passing
on by degrees toward a pale brown, whilst the skin becomes flaccid and opaque ;
death soon occurs, when the body, now shrunken by evaporation, lies on the
lower side of the cell, increasing in depth of tone, until in a few days. nothing

sy pau i hall ee,

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 27

mere than a nearly black scale remains. Should the larvie, however, escape con-
tamination until nearly the period of pupahood, they are sealed over in the nor-
mal way by a cover made of pollen-grains and wax, * * * and which is
pervious to air. The cover furnishes a screen, op which part of the cocoon is
soon after spread, but the inhabitant of the cell is marked out for death, and
before very long the capping or sealing sinks and becomes concave, and in it
punctures of an irregular character appear, * * * and this is a nearly
conclusive sign of the diseased condition of the colony. The sense of smell is
also appealed to, as a peculiar, very offensive, and extremely characteristic
odor now escapes from the diseased combs. The bees, in addition, lose energy,
but become unusually active in ventilating their hive by standing at the door,
heads toward home, and flapping their wings persistently so that a strong out-
current, and as a necessary consequence, a correspondingly indraught are set
up. Should any attempt be made at removing a dead larva which has as-
sumed a deep brown tint, its body tenaciously adhering to the cell wall will
stretch out into long and thin strings like half-dried glue. The microscopist can
easily explain this. The thin chitinous aerating sacs and trachez do not un-
dergo decomposition at all easily, and these remaining, occasion the peculiarity
referred to. * * * The disease iS terribly infectious, and once started soon
spreads from cell to cel! and not unfrequently from stock to stock.

Mr. Cheshire was doubtless quite familiar with the disease of the
brood and this description of the symptoms, we may assume, was not
made from any one case, but from his entire experience. At that
time two diseases of the brood were not recognized. We are justi-
fied in concluding that both diseases existed in England at that time,
as they do now, and doubtless Mr. Cheshire had seen both without
realizing that he was dealing with two distinct maladies. In this
description he speaks of the disease as attacking brood at* various
times, for he says: “ Should the larva, however, escape contamination
until nearly the period of pupahood,” ete. His description before
that sentence applies as well to what we now call European foul
brood as to American foul brood, while the latter description, espe-
cially where he speaks of the odor and ropiness, is undoubtedly drawn
from experience with what we now call American foul brood. There
is, at any rate, ground for the supposition that Mr. Cheshire was.
dealing with two diseases.

In the case of Mr. Cheyne, however, the case was entirely different.
He was probably not familar with the brood disease from practical
experience. He also probably did all his work from one specimen, for
he says: “On August 11, 1884, Mr. Cheshire brought to me a piece
of comb containing larve affected with foul brood, with which I per-
formed the following experiments.” In describing this specimen he
says: “These larve were dead, of a yellowish color, and almost
liquid.” This description certainly applies as well to European foul
brood. Since the original description of Bacillus alvei is so important
in this work, it may be well to quote entire Mr. Cheyne’s part of this
paper. This is practically unavailable to bee keepers, since it is con-

28 MEETING OF INSPECTORS OF APIARIES.

tained in a journal to which few bee keepers can have access. It is
as follows:

On August 11th, 1884, Mr. Cheshire brought to me a piece of comb containing
larvee affected with foul brood, with which I performed the following experi-
ments: Selecting cells which were closed, but which Mr. Cheshire thought con-
tained diseased larvie, I brushed them over with a watery solution of bichloride
of mercury (1:1000) to destroy the organisms on the outside. With several
forceps, that had been heated and allowed to cool, the covering of the cell was
picked off so as to display the diseased larvee. These larvee were dead, of a
yellowish color, and almost liquid, and on examination afterwards their juices
were found to contain numerous moving bacilli. By means of a heated platinum
wire, tubes of meat infusion rendered solid by gelatin (10 per cent), or by
Japanese isinglass, were inoculated from several of these larvee and kept at a
suitable temperature. Development of bacilli, microscopically similar to those
seen in the juices of the larvee, occurred. The characteristics of this develop-
ment will be presently described. Further, in the tubes, kept at the body
temperature, there was not only a development of bacilli but also of spores.

These bacilli, as seen in the larval juices, measure about ;,'55 inch in length and
zos00 Inch in breadth. They are rounded or slightly tapering at their ends and
often have a clear space near one end. In the juices of the larvze during life
they apparently do not produce spores, although after death spores abound.

In the cultivation in the peptonized meat infusion, rendered solid by agar-agar,
the bacilli vary considerably in size, their average length being -,1,; inch, some be-
ing as small as ;>/99 inch and others as large as 5,55 inch. When they have at-
tained the latter size, division of the rod seems to begin. They are always somewhat
pointed at their ends. Their average breadth is ;5455 inch, varying from ;;1,5, to
25 00

The spores are largish oval bodies, averaging in length ;,455 inch (varying from
73100 tO yoz00 Inch), and in breadth 53455 inch (varying from 745, to 35455 inch).

In the agar-agar material the spores are generally arranged side by side in
long rows, and in old cultivations only a few bacilli can be seen, some forming
spores, some without any indication of spores. That these small bacilli can
produce such large spores seems at the first glance at a microscopical specimen
almost inconceivable, but I have been able to trace on the one hand the develop-
ment of the spores in the rods, and on the other the sprouting of the spores into
adult bacilli. This can be done in the following very simple manner:

Take a number of glass slides, each having a moderate-sized cell hollowed
out in its middle; clean it and pass through a Bunsen flame several times to
destroy any bacteria on its surface. With a brush apply a little vaseline around
the depression, and then place the slide under a glass shade to keep it from
the dust. Clean a number of cover glasses, purify them in the flame, and place
them on a pure glass plate beneath another shade. With a fine, pure pipette
put a small drop of sterilized cultivating fluid (meat infusion with peptone) on
the center of each of these cover glasses; then with a fine platinum wire inocu-
jate each of the drops with the spores, or with nonspore-bearing bacilli; rapidly
invert them over the cell, press down the cover glass so as to diffuse the
vaseline around its edge, and place the slides in an incubator kept at the
temperature of the body. These slides are removed at different intervals of
time, and as soon as each is taken out the cover glass is turned over and the
drop of fluid rapidly dried. The specimen can then be stained, mounted in
Canada balsam, and studied at leisure. This method seems to me to be much
more satisfactory than the observation of the organisms swimming about in

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 29

the drop of fluid, while the specimens can be kept permanently and compared
with one another.

In order to study the growth of the spores, I used a cultivation on the agar-
agar cultivating material which had been kept at the temperature of the body
for fourteen days, and which consisted almost entirely of spores, though a few
bacilli were present. As the result of several experiments, I have got a series
of preparations which have been taken at various times (15 minutes, 30 min-
utes, 40 minutes, 1 hour, 14 hours, 1 hour 50 minutes, 2 hours, 2 hours 20
minutes, 2 hours 50 minutes, 2 hours 55 minutes, 3 hours 20 minutes, + hours
20 minutes, 5 hours, 5 hours 35 minutes, 5 hours 40 minutes, and 7 hours 50
minutes), and the course of events is shown in Plate X.¢ The bacilli
stain with various anilin dyes—best, I think, with methyl-violet; but the
spores resemble the spores of other bacteria in not taking on the stain. The
cover glasses on which the organisms are dried are passed three times through
the gas flame and floated on the surface of a fairly strong watery solution of
methyl-violet for one or two hours. They are then washed in water, and
afterwards laid in weak acetic acid (1 per cent) till no more stain comes out.
They are again washed in water, allowed to dry at the ordinary temperature,
and mounted in Canada balsam. A spore-bearing cultivation shows the bacilli
stained violet and the spores unstained, with the exception of their outline,
which is of a faint violet color. In most cases no trace of the rod in which
the spore was formed can be seen. The first change which is observed on
cultivation is that in many cases the outline of the rod in which the spore was
formed becomes faintly visible. This can be seen in 15 minutes, and is, I
think, simply due to swelling by the fluid, as it is also evident to some extent
in the case of spores soaked in water for the same length of time. In from
half an hour to an hour it is evident that the bacilli which were present in the
original material are beginning to multiply, and a considerable number of rods
are now seen containing spores. It is evident that these spores are newly
formed, as extremely few bacilli containing spores were seen in the original
material, whereas in the preparations taken from a half hour to an hour a
considerable number are present. That some of the rods, instead of growing
by fission, at once proceed to form spores is probably to be explained in this
way. When the cultivation was removed from the incubator, some bacilli were
growing by fission, some were forming spores, and some had passed into a state
ready to form spores. The first go on growing by fission, the last complete
their spore-formation, which was arrested by removal from the warm tempera-
ture. That actively growing rods would not have formed spores so early
is evidenced by the facts observed in the second series of observations on the
formation of spores. The next thing that is observed is that several of the
spores take on the stain, and are as intensely violet as the adult bacilli. The
number of the spores which take on the stain in this way goes on increasing
as time passes, till in about four hours almost all the spores stain violet. In
three hours the first indication of sprouting of these spores becomes evident.
The stained part of the spore loses its oval shape, becomes elongated, and is
soon seen to burst through the spore-capsule at one part. It then presents the
appearance of a short rod, with a pale envelope embracing one end. This rod
gradually leaves the spore-capsule and then goes on multiplying as a full-
grown bacillus. In specimens taken from four to five hours all stages of
growth can be seen, and the remains of the ruptured spore-capsules are evident.

The bacilli appear to grow mainly by fission, but I have seen appearances

a@Not reproduced here.

30 MEETING OF INSPECTORS OF APIARIES.

which seem to me only explicable on the supposition that they also grow by
sending out buds from one end. <A bacillus may be seen with a small, some-
what conical stained peint attached to one end, though separated by a marked
division. This is certainly not the common mode of growth by fission, for there
the rod seems to divide into two pretty equal halves, while here we have but a
minute piece attached to one end.

The mode of formation of spores may be traced in a similar manner to that
described above in the case of the sprouting of the spores. It is, however, as a
rule, necessary to leave the organisms to grow for a much longer time than in
the former instance. I have not found development of spores as a rule before
twenty-three hours, but this depends very much, apparently, on the amount of
fluid that was present and the number of bacilli introduced at the time of
inoculation. The first thing noticeable is that the red begins to swell and
becomes spindle shaped. The swelling, which generally affects the middle of
the rod, may in some cases be most marked toward one end, increases in size,
and the center of the swelling gradually ceases to take on the stain. The eap-
sule of the spore is apparently also formed within the rod, and is not merely
the cuter part of the rod. In three or four hours the rod is seen to have almost or
completely disappeared, leaving the spore lying free or within the faint outline of
the original bacillus. It seems to me that the view that spore formation occurs
when the food is getting exhausted is correct, for the time at which this appear-
ance is found depends greatly on the drop placed on the cover glasses, and I
have found in one experiment that in one specimen, after twenty-three hours,
most of the rods were forming spores, while in another specimen where the drop
was much larger there was no trace of spore formation after twenty-eight
hours. I have here described the results of my earlier and rougher attempts
to study the formation of spores. I have, however, now improved the method
in the following way. As I have just now shown, the period at which spores
are first seen seems to depend mainly on the amount of fluid used and the
number of bacilli introduced, and as in the above method, both these factors
vary in each case, one can not get a regular series of preparations showing the
different stages at different times. In studying the sprouting of spores the
amount of fluid and the number of spores does not matter, for if sufficient nutri-
ment is present and a proper temperature is maintained the spores must sprout,
and probably they always take about the same length of time. The difficulty of
obtaining a series of specimens illustrating spore formation is easily obviated
in the following manner. Take a pure flask containing a small quantity of
sterilized infusion and inoculate it from a cultivation containing only bacilli.
Place it in the incubator for two or three hours so that the bacilli may increase
somewhat in number and diffuse themselves through the liquid. Thus the culti-
vating material contains bacilli pretty equally diffused through it, and if after
shaking the flask drops of equal size are taken, each will probably contain
about the same number of bacilli. The minutest quantity of fluid can easily be
obtained by means of a syringe having a fine screw on its piston and a large
nut revolving on this screw. The circumference of the nut being equally divided
into a number of small segments, the same quantity of fluid can always be
expelled from the syringe. By proceeding in this way equal-sized drops con-
taining an equal number of bacilli can be used and a regular series of specimens
obtained. I have found that using two-fifths of a minim containing one bacillus
and keeping the specimen at 36° C., the earliest appearance of spore formation
was evident in forty-one hours.

Leaving these matters, which are of great interest not only in regard to the
Bacillus alvei, but to all spore-bearing bacteria, and which I have therefore

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 81

dwelt on at length, we must pass on to the further consideration of this par-
ticular organism. The first point to be determined in investigating its relation
to foul brood was whether this was a new bacillus, unknown except in connec-
tion with this disease of bees, or whether it was a more or less well-known
form. To ascertain this point with regard to micro-organisms the microscope
is of little use; recourse must be had to the study of their life history, more
especially to their peculiarities of growth on different soils. Of all the ma-
terials employed as cultivating media, Koch’s gelatinized meat infusion is
the most useful for purposes of diagnosis. This is composed of an infusion
of meat containing 1 to 8 per cent of pepton, 10 per cent gelatin, made neutral
by carbonate of soda, and thoroughly sterilized. This material was first in-
troduced with the view of having a highly nutritive solid and at the same
time transparent medium on which to carry on pure cultivations, but it was
soon found that owing to the remarkably diverse ways in which different
micro-organisms grew in it, it could be used as a means of diagnosis of the
kind of organism, a means more certain than any other which we at present
possess. For purposes of diagnosis, as well as with the view of carrying on
pure cultivations, this material is used in three ways. While the material is
still fluid a small portion is poured into a number of pure tubes plugged with
cotton wool, sterilized, and allowed to solidify. <A fine platinum wire, heated
in a flame and allowed to cool, is dipped into the material containing the
bacterium in question, and then, after the removal of the cotton-wool plug, is
rapidly plunged down through the gelatin to the bottom of the tube and then
withdrawn. The plug is reinserted and the tube kept at a temperature suita-
ble for the development of most forms of bacteria, but not high enough to
melt the gelatin. If growth takes place at this temperature it occurs either
on the surface around the point of entrance of the needle, or along the needle
track, or in both places, and the appearance of the growth varies remarkably,
according to the different species of micro-organisins studied. The second way
is to liquefy and pour out a little of the gelatinized material on micrescopic
slides or on larger plates of glass which have been sterilized by heat. These
plates are placed in glass vessels containing moist blotting paper to prevent
drying of the gelatin and to protect them from the dust. After the gelatin
has solidified the purified platinum needle charged with the bacteria is drawn
rapidly over the surface of the gelatin. Bacteria are sown along the track,
grow there, and the whole can be placed under a microscope and the charac-
teristics of the growth studied with a low power. In the third mode a tube
of the gelatin mixture is inoculated with a very minute quantity of the bac-
teria. The tube is then placed in water at the body temperature to melt the
gelatin. When the material is melted, it is thoroughly shaken up to diffuse
the bacteria through it and, while still liquid, is poured out on sterilized glass
plates kept in a moist chamber, as in the former case. Solidification very
soon occurs, and the bacteria being caught at.various parts of the gelatin
grow there in the form of groups or colonies, which can be observed under the
low power of the microscope. I shall now describe the characteristics of the
Bacillus alvei when cultivated in these three modes.

(a) Test-tube cultivations.—lf an infected needle be plunged into a tube of
gelatinized meat infusion in the manner described above, growth occurs both
on the surface and along the needle track. On the surface the bacilli shoot out
in all directions from the point of entrance of the needle, forming a delicate
ramifying growth on the top of the gelatin; the characteristics of this growth
will be presently described under b. Along the track whitish irregular shaped
masses appear, which slowly increase in size and run together. In a few days

32 MEETING OF INSPECTORS OF APIARIES.

processes are seen to shoot out from these masses, which may extend through
the gelatin for long distances from the track, being thickened at yarious parts
and clubbed at the ends. These processes do not appear to join one another at
their ends. A very beautiful and characteristic appearance is got where very
few bacilli are introduced with the needle and where, therefore, at various parts
of the track, more especially at the lower part, individual bacilli or groups of
bacilli are planted at a considerable distance from each other. In a few days
minute round whitish specks become visible to the naked eye. These increase in
size till in about ten days shoots begin to appear. These radiate from the
central mass in all directions and become nodular at various parts, as described
above. When such a cultivation is old, the white branches disappear, and only
little whitish collections of bacilli are seen at various parts. On examining such
a tube with the pocket lens, however, numerous watery-looking tracts are seen
running through the gelatin from the central mass to the whitish collections.
The gelatin at the upper part of the track generally evaporates, to some extent
giving rise to the air-bubble appearance so characteristic of the cholear bacillus.
These are the appearances seen where the material contains gelatin in the pro-
portion of 10 per cent. Where less gelatin is present, the naked eye appearances,
while possessing the same characteristics, are somewhat different. The shoots
are much more numerous and appear much more rapidly, giving rise to a
haziness around the middle track, which with the pocket lens is seen to consist
of numerous delicate branches clubbed at the ends, as in the former case. I
think the amount of pepton present also makes a difference in the appearance,
though of this point I am not yet absolutely certain. The most characteristic
growth is, however, obtained when the material contains 3 per cent pepton as
well as 10 per cent gelatin, the shoots being then less numerous and much
coarser. And I can easily understand that this would be the case, for the bacilli
would have a large supply of nutriment in their immediate vicinity without the
necessity of having, so to speak, to spread out through the gelatin in search of
food as may be the case where no pepton, or only a small amount, is present.
This appearance is quite characteristic of this bacillus and is not seen in the
cultivation of any other organism that I know of. The bacilli of anthrax and
of mouse septicemia also spread out from the needle track, but the appearance
of their cultivation is quite different. In anthrax delicate threads, not clubbed,
shoot out from the track, soon anastomosing with other threads and forming a
delicate network throughout the gelatin. In mouse septiczemia the appearance
is that of a delicate cloudiness spreading through the gelatin. These foul-brood
bacilli, growing in this material, render it liquid after a time, the liquefaction
beginning at the surface and only spreading slowly downwards, but ultimately the
whole tube becomes liquid. After two or three weeks’ growth the appearance
presented by the tube is that of a layer of liquid at the upper part, and the
growth along the needle track with the other appearances described at the
lower part. The liquid portion is clear, except at the bottom of the liquid. where
there is a loose white flocculent deposit of bacilli, and on the surface there may
be a very thin scum. The liquid becomes yellowish in colour after a time and
gives off an odor of stale but not ammoniacal urine, or what may be better
described as a shrimpy smell. This yellowish colour and the peculiar odour
have been found by Mr. Cheshire to be distinctive of the diseased larve.

(b) If gelatin be poured out on a plate, allowed to solidify, and then stroked
with an infected needle, we learn the explanation of the appearances seen jn the
test-tube cultivations. The bacilli at first grow along the needle track, but very
soon they are seen to be collecting at parts forming pointed processes. From
the processes the bacilli grow out into the gelatin, often a single series of rods,
in Indian file, or two or three rods side by side. These processes are not quite

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 33

straight, but tend to curve, and at a little distance from the track they grow
round so as to form a circle. From this circle, which may be formed of single
bacilli, the process continues forming a fresh circle farther on. The bacilli in the
circle increase in number till ultimately it becomes completely filled up, and we
have a nodule consisting of bacilli in the course of the shoot. These shoots may
also join one another, forming a curved anastomosis, and the gelatin in the
immediate vicinity of the bacilli becoming liquid, a series of channels are
formed in the gelatin containing fluid in which the bacilli swim backwards and
iorwards. Later on parts of these channels become apparently deserted by the
baciili, so that the circles look to the naked eye as if they were detached from
the main track, but with a low power of the microscope the empty channels
can be traced.

It is impossible to give a proper idea of the appearance of the growth. The
forms assumed are the most beautiful shapes I have ever seen, but they are
yery numerous, always, however, retaining the tendency to ferm curves and
circles; thus we have the explanation of the appearances previously described
in the test-tube cultivations.

(c) The appearances of the colonies on plates on which the mixture of
bacilli and gelatinized infusion has been poured out is also very characteristic.
The earliest appearance of colonies is a small oval or round group of bacilli.
This group is not homogeneous in appearance under a low power of the micro-
scope, but lines indicating the bacilli are seen in it. It very soon becomes pear-
shaped, and from the sharp end of the pear processes begin to pass out into
the gelatin as before described.

These bacilli do not grow below 16° C. The best growth in gelatin is obtained
at a temperature of about 20° C. They grow most rapidly in cultivating
materials kept at the body temperature. Very few spores are formed at the
lower temperatures, but they appear rapidly and in large numbers at the body
temperature. I have several times observed bacilli containing spores swimming
about freely. The reaction of the medium is not of any great importance, but
a neutral medium is apparently the best. The bacilli swim freely in fluids with
a slow oscillating movement.

They grow rapidly at the body temperature in meat infusion with pepton
and rendered solid by agar-agar, but the appearance of their growth is not
nearly so characteristic as in gelatin. This, indeed, is the case with most
bacteria, so that agar-agar preparations, though very useful for carrying on
pure cultivations at the temperature of the body, are of little value for diag-
nostic purposes. They grow most rapidly on the surface of the agar-agar,
forming a whitish layer, but the shoots described above in the case of gelatin
do not occur, or only very imperfectly, in agar-agar. Here the bacilli arrange
themselves apparently side by side, and, producing spores in this position, we
have as a result, after a few days’ cultivation, long rows of _spores lying side
by side with here and there an adult bacillus.

On potatoes they grow slowly, forming a dryish yellow layer on the surface.
They grow very slowly indeed at the lower temperature. In order to get good
growth it is necessary to keep the potato at the body temperature.

In milk they grow well at the body temperature, and in a few days cease
coagulation of the milk, which also assumes a yellowish colour and gives off the
odour previously described. The coagulum is not firm, like that caused by the
Bacterium lactis, but is like a tremulous jelly, and may remain for a consider-
able time without the separation of any fluid, but ultimately it becomes liquid,
and after some months assumes the appearance of a dirty, brownish-yellow,
glairy fluid. It is very slightly, if indeed at all, acid.

30547—No. 70—07 M——3

34 MEETING OF INSPECTORS OF APIARIES.

They grow extremely slowly in coagulated blood serum, though kept at the
body temperature, and there form very long filaments with comparatively few
spores.

In meat infusion kept at the temperature of the body they grow readily,
causing muddiness, and after a few days a slight but not tenacious scum. The
same peculiar odor is also developed here, more especially if the infusion con-
tains a considerable amount of peptone. I do not think that there is any change
in the reaction of the fluid; I generally make the infusions faintly alkaline,
and after the growth of this organism in it it is faintly alkaline.

These characteristics show that this is a new bacillus, and one which, so far
as my knowledge and experience goes, is only found in foul brood. The constant
presence in large numbers of a characteristic organism in a disease and its
absence elsewhere must, according to our accumulating experience, afford a
strong presumption that the organism is the cause of the disease. In the case
of foul brood this matter has been completely proved by the following experi-
ments, the details of which will be found in Mr. Cheshire’s part of this paper.
With a cultivation in milk he sprayed a comb containing a healthy brood,
allowing the spray to act only on a particular part of the comb. This part
and no other became affected with foul brood. He has also succeeded in
infecting adult bees by feeding them with material containing these cultivated
bacilli.

IT have also had the opportunity of watching the effect of feeding flies with
material containing spores and bacilli. I was one day testing some milk in
which these bacilli were growing; a large blue-bottle fly settled on it and com-
menced to eat. I at once put a large glass funnel over the insect, leaving plenty
of air. When I came to the laboratory twenty-four hours later, the fly was in
the sitting posture on the table and was dead. Its juices were full of these
bacilli, as shown by microscopical examination and by cultivation.

Other animals which I have tested are more or less refractory to this bacillus.
I have kept cockroaches for days in a box in which was milk containing these
bacilli mixed up with sugar. I have also kept them in a box containing a piece
of paper which had been thoroughly smeared with the spores. None of them
died, but I can not be certain that in either case they ate any of the material,
for I never saw them even near it.

I inoculated two mice and one rabbit with a spore-bearing cultivation without
effect.

I injected half a syringeful of a spore-bearing eultivation into the dorsal.

subcutaneous tissue of each of two mice. One of these died in twenty-three
hours, the other seemed unaffected, but in the second case I doubt whether
the full quantity was introduced. In the case of the mouse which died, the
seat of injection and the neighbouring cellular tissue was found to be very
cedematous, but no microscopic changes were apparent in the internal organs.
Numerous bacilli were found in the cedematous fluid, as also a number of
spores which had not been sprouted, and there were also a few bacilli in the
blood taken from the heart. This was proved, of course, by cultivation as well
as by microscopical examination. On examining sections of the various organs
no morbid changes were found, and only very few bacilli were seen in the
blood vessels.

At the same time that I injected the mice I injected a syringeful of the same
cultivation subcutaneously into a guinea pig. This animal died six days later
with extensive necrosis of the muscular tissue and skin and cheesy-looking
patches distributed through it. There was no true pus. On making sections
of the necrosed tissue numerous bacilli, apparently Bacillus alvei, were seen,

.

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 35

but there were other bacteria and also micrococci, as of course would be the
case on account of the death of the skin. No micro-organisms were seen in the
internal organs. It thus remains questionable whether the necrosis was due
to the Bacillus alvei or not, more especially as I have since injected three
guinea pigs subcutaneously with spore-bearing cultivations, but without effect.
I must reserve the action of these bacilli on the higher animals for further
investigation, as well as several other points of interest in regard to this
organism to which I have not here alluded.

I venture to think that when all the evidence brought forward by Mr. Ches-
shire and myself is carefully weighed no doubt can be entertained that this
bacillus is new to science and is the cause of foul brocd. Many questions of
course still remain open, requiring further investigation into the life history
of the disease.

The next investigator to take up a bacteriological treatment of bee
diseases was Prof. J. J. Mackenzie, bacteriologist of the provincial
board of health of Ontario. The results of this work were published
in the Ontario Agricultural College Report for 1892, pages 267-273.
At the request of the Bee Keepers’ Union of Canada certain things
were taken up which had a very practical bearing on the question of
eradicating the prevalent disease.

Professor Mackenzie knew of but one disease, probably, and having
in hand the work of Cheshire and Cheyne, assumed that the disease
found in Canada is the same as that deseribed by Cheyne. This is a
natural mistake after the confusion in the diagnosis by Cheshire. — It
was not the object of this investigation to demonstrate what organism
produces the diseased condition, but, assuming that Bacillus alvei
causes the trouble, to determine what resistance to heat the organ-
ism has.

No adequate description, such as would allow us to make any com-
parisons with Bacillus alvei, is included in Professor Mackenzie’s
paper. We do know, however, or at least have every reason to believe,
that European foul brood was not found in Canada at that time and
is not prevalent there now. I have been informed personally by Mr.
William McEvoy, the veteran inspector of Ontario, that the disease
which we now designate as American foul brood is the prevailing
disease in Canada. It seems reasonable to suppose, therefore, that the
samples taken to Professor Mackenzie by Mr. Holtermann and others
did not contain any Bacillus alvei.

Professor Mackenzie does not indicate in his paper that he had any
difficulty in getting the organism with which he worked to grow on
ordinary media. Bacillus larve, which is present in American foul
brood, does not grow in such media, however, so there is but one con-
clusion to be reached, and that is that he was dealing with some non-
pathogenic form and not with Bacillus larvw. Since the bacillus
described by Doctor White as Bacillus A is found on combs, both
diseased and healthy, and somewhat resembles Bacillus alvei, it may
be that this is what Professor Mackenzie had.

’

36 MEETING OF INSPECTORS OF APIARIES.

This paper has probably come to the notice of but few bee keepers
in the United States, because the report of the agricultural college is
not widely distributed. To make it available for comparison, there-
fore, it is included here.

THE Fout Broop Bacitius (B. ALVEL) ; ITS VITALITY AND DEVELOPMENT.

[From Ontario Agricultural College Report for 1892, pp. 267—273.]

Mr. J. J. Mackenzie, B. A., bacteriologist of the provincial board of health of
Ontario, read the following paper :

GENTLEMEN: At the request of your secretary, Mr. Holtermann, I undertook
for your union some investigations on the subject of foul brood, the results of
which I propose giving you in this paper. Although it is almost a year now
since I undertook this work under the auspices of the Agricultural and Experi-
mental Union, it is by no means exhausted, and there are many points which
require to be further elucidated, which I have not had time as yet to touch on,
owing to the fact that investigations on foul brood had to be carried on simul-
taneously with my regular laboratory work. 'These points J hope to work at
next summer, and reserve the privilege of reporting again to your unicn on the
results of further investigation.

The subject of foul brood is an old one to apiarists and an intensely interest-
ing one to Canadian bee keepers, but in reading over the bee journals one can
not help being struck with the great want of unanimity amongst bee men as to
the disease, how it should be treated, how it is spread, and on many other points.
Some would have us believe that the disease arises de novo whenever insanitary
conditions prevail; others claim that there is a specific infection and where the
disease arises it must have originated from previously existing disease; some
claim that the honey is the only method of transmittal ; others that it is not, and
so on. On every point there seems to be plenty of arguments pro and con.

I have attempted in my work to take hold of some of these controverted
points from a bacteriological standpoint in order to aid in coming to some defi-
nite conclusion. Some of these points I should consider settled from the results
of previous investigation ; but as many bee men do not seem prepared to accept
this, my work will have value as confirming what has already been done.

Before an association which includes many practical bee keepers it would be
superfluous to enter upon a minute account of the clinical features of the dis-
ease. Most of you know them better than I do. I certainly would not be pre-
pared to “spot” foul brood in an apiary, although I certainly think I can under
the microscope. The infectious character of the disease has been generally
accepted for many years, but not until Cheshire and Watson Cheyne worked it
out scientifically was it definitely proved. They isolated a bacillus (Bacillus
alvei) which they found in the diseased brood and which they cultivated on
nutrient media for many generations, finally reinfecting perfectly healthy brood
from these pure cultures. This evidence to a bacteriologist is absolutely con-
clusive that Bacillus alvei is the specific cause of foul brood. Consequently,
when I began my investigations on some samples of diseased brood which were
sent me through Mr. Holtermann, I looked at once for Bacillus alvei. Micro-
scopically and by means of bacteriological methods I had no difficulty in isolating
2 bacillus which corresponds in all points to Bacillus alvei. It is a bacillus
similar to that of Cheshire’s in size, produces spores which are somewhat
thicker, giving the bacillus a clubbed appearance. On agar jelly it grows
rapidly so as to cover the whole surface. In gelatin its growth is very peculiar,
shooting out from the infected point in all directions. On potatoes it produces

~

+
PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 37

a yellow growth. All these characters show conclusively that it is identical
with Bacillus alvei. There seems no doubt, therefore, that the foul brood which
we have in Ontario is the same disease and produced by the same bacillus as in
other places.

Many prominent bee keepers, both here and in the States, however, maintain
that wherever unsanitary conditions are allowed to prevail, wherever chilled
brood is allowed to putrefy, or decapitated drones are left to decay in the
hive foul brood may arise de novo. This is not a new theory, either in bee
keeping or in medicine, but unfortunately it is a theory which is not supported
by the results of investigation. Diphtheria naturally will develop more readily
if unsanitary conditions are present, but it certainly will not develop if the
Bacillus diphtherie is absent. The same is true of other diseases, and conse-
quently when we come to consider such a decidedly infectious disease as foul
brood and learn the facts about it which such men as Cheshire have told us we
naturally come to the same conclusion. If I were to maintain that a Carnio-
lan queen might lay an egg which would develop into a humble bee, bee men
would be inclined to think that not only my bee knowledge, but also my scien-
tific knowledge, was at fault; but yet in all the bee journals I find many promi-
nent bee keepers maintaining that an ordinary microbe which produces putre-
faction may become metamorphosed into the specific cause of foul brood. It
it is easy enough, however, to combat such an opinion upon a priori grounds,
but not quite so easy to offer convincing proof.

In order to do this I thought it worth while to try some experiments. With
this end in view I obtained some comb containing chilled brood and endeavored
to isolate Bacillus alvei from it, but without success.

There were plenty of other bacteria, but none which presented the well-
marked morphological characters peculiar to Bacillus alvei. Again I had sent
to the laboratory a piece of perfectly healthy comb. I killed the brood by chill-
ing. Then I infected some of the cells from a pure culture of Bacillus alvei.
I allowed all the chilled brood to putrefy in a moist chamber for two weeks, at
the end of which time I obtained Bacillus alvei again from the cells which had
been artificially infected, but could find no traces of it in the other cells. I
left this comb in a moist chamber for several months and again examined, but
with the same results. In the cells in which Bacillus alvei had been placed it
was still to be found; in the others it was not present.

It seems to me that an experiment such as the above conclusively shows that
there is a distinct difference between foul brood and ordinary putrefaction.

In considering the subject of the vitality of Bacillus alvei the first question
which naturally arises is its power to resist heat. We know that bacilli which
produce spores and those which do not stand in entirely different positions in
this regard. The sporeless bacillus is destroyed at a much lower temperature
than one which contains spores. Consequently in considering the question of
the vitality of Bacillus alvei, which produces spores very quickly and easily, we
may confine our attention entirely to the vitality of the spore.

This is of special interest, as the question has been repeatedly raised whether
it is dangerous to use comb foundation made from foul-broody wax. Does the
temperature to which the wax is raised in the maaufacture of comb foundation
sufficiently destroy the vitality of the spere? Can the spore germinate and
infect the brood when once inclosed in the wax?

These questions have been raised by many careful thinkers among bee men,
and certainly deserve attention. The second point ought to be considered first,
since if surrounding a spore with a film of wax prevents its germination, we
need pay no further attention to the question of heat. The crucial test of this

388 MEETING OF INSPECTORS OF APIARIES.

would naturally be, supply a healthy colony with comb foundation known to
contain the spores and observe the result. This I had hoped to try with the
assistance of your-secretary, but other work came up which interfered with the
earrying out of this experiment and consequently it had to be postponed until
next year. However, I was able to perform one experiment which throws
some light on the subject. Mr. Holtermann, the secretary of your union, sent
me several pounds of very fine wax, such as is used for the manufacture of
comb foundation. I cultivated the Bacillus alvei upon agar jelly until I had
a large quantity of the bacilli containing spores; this was carefully scraped
off the jelly and dried first in the air and then over sulphuric acid. The result-
ing grayish mass was pulverized with a sterilized pestle and mortar, and finally
mixed thoroughly with the melted wax kept at a temperature sufficiently low
to prevent the immediate destruction of the spores by heat. By this means an
enormous humber of spores were introduced into the wax. After stirring the
wax for some time in order to insure a proper mixing it was allowed to cool.
This, as you all know, takes some time when dealing with a considerable quan-
tity. During the cooling I was careful not to disturb the wax.

After it had solidified I set out to discover if I could again obtain my
bacillus from the infected wax. If it would germinate in the nutrient media, it
certainly would in the bees, and that point was to a certain extent settled.
Now I obtained the following results:

From the upper layers of the infected wax I was unable to obtain cultures of
the Bacillus alvei, either by melting the wax in the nutrient jellies or by allow-
ing particles of the unmelted wax to fall on the surface of these jellies.

From the under layers, however, the results were different; particles of wax
placed on nutrient agar in an oven kept at 98° F. became surrounded in twenty-
four hours with a luxuriant growth of Bacillus alvei. When the wax was
melted into the agar or into beef tea, I also obtained the bacillus, consequently
it looks as if the mere fact of enveloping the spores with a film of wax was not
sufficient to prevent germination. I confess I can not understand how a spore
could germinate when surrounded with a film of wax. Spores in germinating
require moisture, and if a spore is completely embedded in wax, it can not
obtain sufficient moisture to germinate; I would rather believe, therefore,
that in this particular experiment the spores had not each an envelope of wax,
but that many of them were partially free from wax. Now, if this was the case
in my experiment, where I endeavored to make the incorporation of the spores
in the wax as thorough as possible, I certainly think it may frequently be
the case when foul-broody wax is used and no particular precaution taken.
That even when spores are thoroughly surrounded by wax they may not be
freed occasionally by the workers is a point which requires further elucidation
and upon which I intend to try some experiments next year.

In looking through the bee journals, however, I find it everywhere main-
tained by foundation makers that they never knew of a case of foul brood
originating from foul-broody wax; and I have yet to discover a well authenti-
‘ated case where this has occurred. What explanation can we offer of this
widespread opinion?

I explained to you above that I was unable to cultivate Bacillus alvei from
the upper layer of the infected wax. Your secretary also sent me a small
specimen of wax which he stated he knew to be from foul-broody comb. This
I examined repeatedly for foul brood, but was able to obtain it only once. I
think we must look to the physical conditions for an explanation of the free-
dom from infection through comb foundation. The difference in the specific
gravity of the bacteria and of melted wax is so great that throughout the

PRESENT STATUS OF INVESTIGATION OF BEE DISEASHES. 39

process of manufacture the bacteria tend to fall to the bottom. The first
refining of the wax must, of course, remove the greater quantity, and the vast
majority of the remainder will settle to the bottom during the process of
foundation manufacture. But that the simple process of mixing the infected
material with the melted wax is not sufficient to prevent germination I think is
shown by the results quoted above, where simple fragments of infected wax
when placed on agar jelly gave rise to a culture of Bacillus alvei.

This question I hope to touch on again after I have had an opportunity of
supplying healthy bees with foundation made from infected wax.

The other question is whether the temperature to which wax is raised dur-
ing foundation making is sufficiently high to destroy the spores of foul brood.
In order to decide this question there are several points to be noted. The
first is the character of the heat. We know that moist heat will destroy bac-
teria and their spores much more quickly than dry heat, and Mr. Corneil, of
Lindsay, has raised this point several times, claiming that the heat to which
the bacteria are exposed in melting wax is not moist heat but dry heat, conse-
quently we must heat to a high temperature and for a long time in order to de-
stroy the spores. The point is undoubtedly well taken, and can only be settled
by direct experiment. In order to determine the temperature at which the spores
are destroyed in melted wax, I used a method that was first described by
Koch. Sterilized silk threads were saturated with a beef-tea culture of Bacillus
alvei in which there were large numbers of spores. These threads were then
allowed to dry and in the dry state were preserved. These dried threads were
introduced into the melted wax and allowed to remain in it for a definite time
at a fixed temperature. At the end of that time the thread was introduced into
the melted agar or into beef tea heated to the melting point of wax, and thor-
oughly shaken, so as to separate the wax as much as possible from the threads ;
then the culture medium was rapidly cooled, and the tubes placed in the ordi-
nary cultivating oven kept at 98° EF. If I obtained a growth of bacilli, I con-
cluded that the threads had not been sufficiently heated in the wax; if I did
not, I concluded that they had been sufficiently heated. The following are
my results:

Atel 2 eh (OOLSCS)) :

For one-quarter of an hour: Growth.

For one-half an hour: Growth.

For one hour: Growth.

For one hour and a half: Growth.

For two hours: Growth.

For two hours and a half: No growth.
At 194° BY. (905)C.):

For one-half hour: Growth.

For one hour: Growth.

For two hours: Growth.

For three hours: No growth.

For four hours: No Growth.

On the other hand, a temperature of 122° F. (50° ©.) did not destroy the
spores in twenty-four hours.

I have repeated these experiments several times with the same results, so
that I would conclude that to destroy the foul brood in wax it is necessary to
heat to a temperature of at least 194° F. for at least three hours. Now the
question arises, does this take place during the process of manufacture of comb
foundation? In order to get as much data as possible on the subject I wrote
to Mr. Larrabee, of Michigan Agricultural College, as he had kindly offered me

40 MEETING OF INSPECTORS OF APIARIES.

any assistance in his power. He applied to two prominent’ foundation makers
for information. From their replies it is apparent that, for a short time at any
rate, during the refining and purifying of the wax it reaches a temperature
quite at or near 212° F. During sheeting, however, it apparently does not reach
a temperature much above the melting point, say 175° F. They both seemed to
agree that steam heat for too long a time injures the quality of the wax.

In the American Bee Journal, 1891, page 470, we find some statements on the
subject in a reply by two prominent foundation makers to an article by Mr.
Corneil upon the dangers of infected comb foundation. One of them, Mr.
Dadant, states that in refining it is heated for some time at 212° F. and is kept
liquid for twenty-four hours. The other, Mr. M. H. Hunt, states that it is kept
at the boiling point for six or seven hours. If these are the actual tempera-
tures reached during foundation making, I am inclined to think there is little
danger from foul brood in that direction.

I thought it possible that the whole question could be settled by introducing
a certain amount of some disinfectant—say beta naphthol—into the melted wax,
but my results have not been satisfactory. Apparently even the introduction of
1 per cent beta naphthol into wax did not hasten materially the destruction of
the spores. I was able to demonstrate the presence of living spores in wax
containing 1 per cent beta naphthol and heated for two hours to 194° F.

From all these facts, and taking into consideration also the physical fact of
the settling of the bacilli to the bottom, I should think that with reasonable care
in the preparation of comb foundation the dangers of infection from this source
would be slight. But that the spores may germinate after being mixed with the
wax, I think I have shown.

Why the spores of the Bacillus alvei are killed so quickly in the melted wax
I am not able to explain, but it may be due to the fact that the wax itself
when heated to such a temperature has an antiseptic value. That the spores
resist other antiseptics as strongly as do the spores of anthrax I have proved
by testing.

Cheshire and others recommend a solution of 2 per cent carbolic acid for
disinfecting the hive after removing infected comb, but on actual experiment
with the infected silk threads I have found that 2 per cent carbolic acid did
not kill the spores in six days. These results are similar to those obtained
by Koch for the spores of anthrax, and show that 2 per cent carbolic acid
can not be relied on to destroy the spores. However, the question of the
value of antiseptics I will take up more in detail later on in this paper.

I would like to say a word or two now on the methods of treating the
disease. There are practically two methods: first, the starvation method;
second, the method of medicated sirup. Mr. McEvoy’s method of treatment,
it seems to me, is practically a modification of the starvation method. The
first method is widely used both here and in the United States, whilst in
Pngland and Europe generally the second method is adhered to.

Considering the vitality of the spores of foul brood, it would seem at first
sight useless to try any process which did not recognize as its foundation
the destruction of the germ. I find, however, that many prominent bee
keepers who have had practical experience with the method of starvation,
or Mr. McEvoy’s method, accept it as successful. I have not had an oppor-
tunity to examine colonies which have been cured in this manner, and so can
not say that the bacilli have disappeared. I hope next summer to test this
question more fully. We may, however, examine into the rationale of the
method. In conversation with Mr. Corneil, of Lindsay, he made a suggestion
which may be quite familiar to you all, but which seems to me the only explana-

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 41

tion. That suggestion was that either starvation or comb building carried
the infected nurses past the period at which they act as nurses and gave them
a chance to rid their intestines of the germ. If this is combined with a
removal to absolutely clean hives, with new foundation, it may succeed, but
I must say that absolute cleanliness in this respect must be insisted on.
As I said above, I have not had any opportunity of investigating the results
of these methods practically, and so can not speak with certainty.

The fact of the presence of the bacilli in the workers and in the queen bears,
to a certain extent, upon this question. Cheshire and others make the state-
ment that the bacilli are found in the intestines of the workers and in the
ovaries of the queens. My own experience confirms this. I have found them
repeatedly in the workers, and in five queens from infected hives I succeeded
in obtaining the bacillus from the ovaries of three. That they are not always

- present in the ovaries of the queens from diseased colonies is certain; their
presence there is apparently accidental. For instance, in the case of one last
year’s queens in a hive rather badly diseased I was unable to find the bacillus,
whilst in a six weeks’ queen from a hive in which there were only a few dis-
eased ceils I succeeded in finding it.

Cheshire’s statement that he found a bacillus in an egg of an infected queen
seems to me to require confirmation. I have not been able to find the eggs in-
fected myself, but it is a question which would require very Jong and careful
investigation before one could be able to deny or confirm such a statement.

In the second method of treatment by medication I do not think that an
absolute destruction of the spores takes place, any more than in the starvation
-method. As I have shown above, 2 per cent carbolic acid was not sufficiently
strong to destroy the spores, consequently it is not likely that 0.2 per cent (1
part in 500) would be strong enough. I tried 0.2 per cent, but found it quite
unsuccessful. Its action then must have another explanation. To test this I
made up a sterilized beef broth containing 1 per 500 of carbolic acid, and in it
placed my infected silk threads. ft found that there was no indication of
growth. These threads were then taken out and placed in ordinary sterilized
beef broth, and I obtained a luxuriant growth, i. e., the 0.2 per cent carbolic
acid in the culture fluid, although it did not destroy the spores, prevented their
germination. That, then, is the explanation of the value of carbolated sirup
in the treatment of foul brood, it prevents the germination of the spores. ‘The
bee journals contain numerous examples of cases where carbolated sirup pro-
duced an improvement, but as scon as it was stopped there was a relapse. It
is evident that here again, as in the starvation process, there must be com-
bined an extremely thorough cleaning up, so that the best possible results
may be obtained from the treatment. Medicated sirup does not destroy the
spores, it simply prevents their development and gives the bees a chance to
rid themselves of the infection, and in that respect I certainly think resembles
the starvation process. Its advantage over that is that it can be carried on for
a longer time.

In the course of these experiments I tried another substance which has been
much used since Lortet’s work on the subject, viz, beta naphthol. I do not
think myself, from recent work on this substance, that beta naphthol should be
ranked very high as an antiseptic, mainly on account of its insolubility in
water. I found, however, that a beef broth containing 1 per 1,000 beta naph-
thol would not allow spores of Bacillus alvei to germinute, and consequently
had an equal value with 1 per 500 of carbolic acid. It has an advantage over
cearbolic acid on account of the disagreeable taste of the latter, and I think
would be more acceptable to the bees.

49 MEETING OF INSPECTORS OF APIARIES.

Salicylic acid in sirup has apparently the same effect, but I would not recom-
mend the addition of borax, as Behring has shown that borax lowers con-
siderably the antiseptic value of salicylic acid.

I tested also formic acid in the same way, but my results so far have not
been satisfactory, owing to the uncertain strength of my sample of formic acid.
I prefer to reserve a report upon it and other substances which I wish to try
until later.

Mercuric chloride I have not tested, as I do not think it wise to use it around
the hive. The idea of using a 1 per 1,000 solution to spray the diseased combs,
as suggested sometimes, is, I think, absurd, and would be a rather serious
operation for any living brood.

You will see that I consider all these methods of treatment do not in them-
selves necessarily presuppose the destruction of the spores, but depend upon the
fact that for a longer or shorter period the spores are prevented from germinat-
ing, and in this period they are eliminated from the infected bees. Whether the
vitality of the bees themselves has an effect upon the elimination or destruc-
tion of the spores is a point which would be extremely interesting, but one on
which at present we have no definite information. From the results of bac-
teriological work on other diseases we know that the animal body is engaged
in a constant warfare with the disease germs which may be introduced, and
this also may be the case in foul brood. Much more extended investigations,
however, would be necessary to prove this. It is much safer for apiarists to
accept the possibility of a recurrence of the disease after a course of treatment,
owing to the lodgment somewhere of some of the spores of Bacillus alvei, and
by care and cleanliness remove this possibility. To do this the hives and
frames in which a foul broody colony has lived must be sterilized, and this may
be done in various ways. For the sterilization of material by disinfectants
there was a tendency formerly among bacteriologists to run to such dis-
infectants as corrosive sublimate, carboliec acid, ete., but later work has shown
that there are a number of common chemicals which will act just as well, or
perhaps better. Corrosive sublimate has lost much of its reputation as a dis-
infectant within the last few years, and carbolic has been shown to be not
nearly so powerful as at first supposed.

For cleaning hives and frames which are suspected to contain the spores of
foul brood a hot 10 per cent solution of soft soap is perhaps as effectual as
any that can be recommended. <A good strong solution of washing soda, when
hot, is also very active, destroying the spores in a few minutes. Both these are
certainly better than 5 per cent carbolic for disinfecting the hives and frames,
as their cleansing properties are so much better than it, and Behring has
shown that 5 per cent carbolic requires at least three hours at bloogl heat to
destroy the spores of anthrax. In case the soap or the washing soda is used,
however, it must be used as hot as possible. Of course anything which is of no
value should be burned.

I trust that in this paper I have thrown a little light upon some of the facts
in connection with the disease of foul brood, but, as I stated in the begin-
ning, I reserve the privilege of submitting to you at a future meeting the re-
sults of next summer’s work.

Before closing I desire to express my thanks to your able secretary, Mr.
Holtermann, for the assistance which he has given me, and also to Mr. Corneil,
of Lindsay, for advice and for the use of volumes of all the principal bee
journals, which he has supplied me with; also to Mr. Larrabee, of Michigan
Agricultural College, in connection with the subject of comb foundation.

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 43

The next investigation to be considered is that by Prof. I. C. Harri-
son, professor of bacteriology of the Ontario Agricultural College.
Previous to this Dr. Wiliam R. Howard published a paper on the
subject, but this can be discussed better at a later time. In the paper
by Professor Harrison, previously mentioned, the author gives a
detailed: description of the bacillus with which he worked. The
description is as follows:

THE ORGANISM.

Bacillus alvei, Cheshire and W. Cheyne, 1885, from the larvee of bees suf-
fering from the disease known as foul brood, la loque (Fr.), and faul
brut (Ger.).

Morphological characteristics —In form the organism is a slender bacillus,
with ends slightly pointed and rounded. ‘In the larval juices it is about +55

000
of an inch in length and ,,!55 in breadth. On agar the bacilli vary considerably
in size, averaging ;;, inch, some-small as z5$59 inch, and others as large as

soy inch. When they have attained the latter size, division of the rod seems to

begin. They are always somewhat pointed at their ends. Their average
breadth is sgg55 inch, ranging from ;5355 to zshpq imch (Cheshire and W.

Cheyne). Klamann (Bienenwirtschaftliches Centralblatt, Hannover, 1888, pts.
18 and 19) states that a clear space often appears in bacilli with pointed ends.
From agar cultures twenty-four hours old, at 37° C., the bacilli average 4 y in
length and 1.0 in breadth. On gelatine cultnnes grown at 22° C., they are
somewhat shorter. They grow singly, but occasionally form chains of various
length.

Stains.—With the ordinary aniline stains the bacilli colour rather badly
(Hisenberg, Bakteriologische Diagnostik, Hamburg, 1891, p. 298, and Klamann,
Bienenwirtschaftliches Centralblatt, Hannover, 1888, pts. 18 and 19). The best
stains are methylene blue and methyl violet. The bacilli accept Gram’s stain,
but the spores are not colored by it. I find the most satisfactory stain in methyl
violet.

Capsule.—No capsule has been demonstrated by Welch’s method.

Flagella.—The bacilli are actively motile and possess a single flagellum at one
pole. The motility of the bacillus is quite pronounced in fresh cultures obtained
from bouillon, agar, and gelatine. The flagella stain by Pitfield’s, Loefflers’s, and
Van Ermegen’s method.

Spore iSO niphs oR RORES are formed by the bacillus, and are large oval bodies
averaging in length ;,4,, inch, and in breadth 5,455 of an inch. On agar the
spores are arranged in long rows, side by side, and are greater in diameter than
the cells from which they are derived. The earliest appearance of spore forma-
tion takes place in forty-one hours, at 36° C. (Cheyne), but in some cases it is
even sooner. The spores are formed in the center of the rod, and the formation
occurs as follows: The rod begins to swell and become spindle-shaped. Occa-
sionally the swelling is more marked at one end than in the center. The spindle-
shape increases in size, and the center of the swelling gradually ceases to take
the stain. The capsule of the spore is apparently formed within the rod and is
not merely the outer part of the rod. In three or four hours the rod is seen to
have almost or completely disappeared, although parts of the faint outline of
the ordinary bacillus may be noticed.

Germination of spores.—Under favorable conditions the beginning of the ger-
mination of the spores takes place in about three hours. The spore loses its oval

+4 MEETING OF INSPECTORS OF APIARIES.

shape, becomes elongated, and is soon seen to burst through the spore capsule.
It then presents the appearance of a short rod, with a pale envelope embracing
one end. The rod gradually leaves the spore capsule, and then goes on multi-
plying as a full-grown bacillus. According to Hisenberg (Bakteriologische
Diagnostik, Hamburg, 1891, p. 298) the spores are decolorized by the tubercle
bacilli stain, but preparations may be obtained by using the Ziehl-Neilsen stain
and alcohol for decolorization. The spores also stain by the method of Neisser.

Polymorphism.—Variations in size and shape may be brought about by growth
in acid media, or in media containing different sugars. These variations occur
also in the same culture, subjected to exactly similar conditions of growth.

Involution forms.—Abnormal forms are especially abundant when the bacillus
is grown on blood serum; peculiar Y-like forms and clubbed shapes are of
common occurrence, and relatively few spores are found.

BIOLOGICAL’ CHARACTERS.

Bowillon— ‘In meat infusion at the temperature of the body they grow
rapidly, causing muddiness and, after a few days, a slight but not tenacious
scum” (Cheshire and W. Cheyne). In bouillon, with a reaction of + 0.08
(Report of Convention of American Bacteriologists, Journal American Public
Health Association, Vol. XXIII, 1898), at 87° C., there is a slight turbidity in
fourteen hours, especially noticeable when the tube is shaken. In twenty-four
hours, the liquid is uniformly turbid, with a very fine sediment. In forty-
eight hours the turbidity increases and a pellicle commences to form. Reaction
of the culture at this time, +0.07. After ninety-six hours the broth is clear,
with a pellicle, white, rather massive, and somewhat tenacious. There is also
much sediment. Reaction, after ten days’ growth, neutral.

Glycerine bouillon.—Media with original reaction of +0.08. At 37° C. the
bouillon becomes slightly turbid in twelve hours and quite turbid in twenty-four,
with a fine,- whitish pellicle on surface, which does not extend to the sides of
the tube. If the culture is shaken, the pellicle deposits in flaky masses. The
reaction is +1.2. In thirty-six hours the turbidity clears, leaving the media
bright, with a smooth, thin, tenacious, and white pellicle on the surface. In
many cases the pellice becomes very wrinkled and greasy looking. At the end
of eight days the reaction is +2.2, and the bouillon is several shades darker in
color, but quite clear. The reaction after fourteen day’s growth is +4.2. At
22° CG. the same changes occur, but growth is slower. The bacilli are relatively
less numerous than in bouillon and are slightly shorter and thicker.

Glucose bouillon.—With a reaction of +2, at 37° C., the broth is more turbid
than plain bouillon after fourteen hours’ growth; and in twenty-four hours the
sediment is heavy and turbidity very marked, but no pellicle. In forty-eight
hours the media is opaque and cloudy, and the pellicle is beginning to form. In
ninety-six hours the broth is less cloudy, but the sediment is heavier, and a
white, thick pellicle is formed. -It is often wrinkled, but not quite so much so as
that on the glycerine broth. Reaction of broth after ten days’ growth, + 4.6.
The bacilli are occasionally clubbed, and Y-like forms may occur. They average
5 w in length and may be slightly curved.

Lactose bouillon.—With a reaction of +1.06, at 87° C., the growth resembles
that of plain bouillon for the first twenty-four hours; but at the end of forty-
eight hours, itis more turbid. In ninety-six hours, a tenacious pellicle forms, less
massive than that on glucose broth. Reaction after ten days’ growth, +2.4. The
bacilli average 3.5 uw in length.

Saccharose bouillon.—With a reaction of +1, at 37° C., the turbidity and
sediment are heavier than any of the other bouillons. In forty-eight hours the

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 45

broth is quite opaque and whitish looking. A heavy sediment is then present,
and pellicle formation is just beginning. In ninety-six hours the cloudiness is
about the same, but there is an increase of sediment, and the pellicle is thin and
membranous. Reaction of media after ten days’ growth, +4.04. The bacilli
average 5 uw in length.

Gelatine plates.—At 22° C., in twenty-four to thirty-six hours, the colonies are
small, round, oval, or lozenge-shaped, with peculiar projections or shoots from
one end of the colony, giving it a pear-shaped or tadpole-like appearance,
according to the amount of development of the projection. In many cases
several of these outgrowths occur from different portions of the colony. By
placing a cover glass on the surface of the gelatine and using objective 7, the
bacilli may be seen moving around and around the colony and to and fro along
the projections. At the end of forty-eight hours the colonies are larger. Fine
processes or projections are shooting out into the gelatine in all directions,
forming peculiar figures in circles or club-like forms. ‘It is impossible,’ says
Cheyne, “to give a proper idea of the appearance of the growth. The forms
assumed are the most beautiful shaped I have ever seen; but they are very
numerous, always retaining the tendency to form curves and circles.’ After a
time the gelatine is liquefied and the beautiful appearance of the colony is
destroyed by the liquefaction of the gelatin.

These peculiar shaped colonies are most typical when the germ is taken
from the diseased larvee. After prolonged cultivation on various kinds of media,
there is a tendency for the colonies to become round, and the peculiar branech-
ing forms are not seen in such numbers. The composition of the gelatine
also seems to make a difference in the appearance of the colonies. In gelatine
containing 12 per cent gelatine the processes are not so long. The same effect
may be brought about by using more peptone in the composition of the media.

Gelatine tubes.——In stick cultures at 20° C. growth occurs all along the line
of puncture. On the surface delicate branching or ramifying growth occurs in
three days. ‘These outgrowths soon run together and the gelatine is liquefied,
first around the line of puncture, and in five days extends over the whole
surface. The growth in the depth of the gelatine occurs as a whitish streak
all along the needle track, and from this numerous shoots and growths branch
out into the gelatine in all directions, giving a haziness to the appearance of
the gelatine, which then begins to liquefy. If the inoculation is a heavy one,
the shoots are coarse and may have club-shaped extremities, and from these
swollen ends fresh shoots may start. Cheyne obtained the most chuaracter-
istic growth in gelatine containing 3 per cent of peptone as well as 10 per cent
gelatine. The whole tube is liquefied in from two to four weeks’ growth. The
liquid becomes yellowish in color and gives off a peculiar odor. Klamann
states that in gelatin acidified with lactic acid the growth is slow and long
threads are formed.

Gelatine streak cultures.—In gelatine streak cultures the appearance is very
similar to what one sees in stick cultures. The bacilli first grow along the line
of inoculation, and then throw out shoots into the surrounding gelatine, pro-
ducing the appearance noted in the stick culture. The bacilli move to and fro
along the channels of liquefied gelatine.

Agar plates.—On agar plates at 37° C. the colonies at the end of eight hours
are small and burr-like, with spines protruding in all directions, giving the
colony the appearance of a sea urchin. In some cases the projections are
from one side or end. At the end of twelve hours the colonies have well-
defined projections, visible to the naked eye. The colonies in the depths of
the agar are more spiny, the processes being much shorter. On agar plates

46 MEETING OF INSPECTORS OF APIARIES.

streaked with a light inoculation most beautiful forms occur. The growth of
the bacilli spreads over the surface and branches repeatedly, giving the
appearance of seaweed. This appearance is distinctly characteristic; and as
the growth is very rapid, this method commends itself for making a quick
diagnosis of the presence of the bacillus in laryze supposed to be diseased.

Potato cultures.—On potatoes the growth differs considerably, according to
the reaction and age of the potato. Sometimes a brownish wrinkled growth
forms, which gives off a peculiar odor; at other times a dryish yellow layer
appears. “The bacilli grow very slowly indeed at 20° ©.” (Cheyne, Journal of
the Royal Microscopical Society, 1885, p. 381). Even at 37° GC. they grow
slowly.

Milk.—In milk at 37° C. coagulation of the casein occurs in three days. The
milk becomes yellowish and gives off a characteristic odor. After several
weeks’ growth the curd is digested and a whey-like fluid remains.

Blood serum.—On blood serum at 387° C. the growth is rather slow and
polymorphic forms are common. ‘ Very long filaments are formed” (Cheshire
and W. Cheyne, Journal of the Royal Microscopical Society, 1885, p. 381).
These long forms may be from five to ten times as long as the average bacillus
growing on gelatine, and consists of single cells. The filaments are often wavy
or twisted and of unequal thickness. The extremities of the long, bent rods
are often clubbed; and Y-like forms are numerous. Spores are formed very
sparingly, and the blood serum is liquefied.

Synthetic media (Uschinsky).—In Uschinsky’s medium no growth occur aa
but if the medium is neutralized, good growth ensues. The bacilli occur in
threads and a pellicle is formed.

Dunhan’s solution.—The bacilli are small when grown in this solution. No
threads form, but there is a slight indol reaction after nine days’ growth.

Relation to free oxygen.—Cheyne states that the germs grow most rapidly
on the surface of agar and arrange themselves side by side; and they produce
spores in this position after a few days’ growth. BHisenberg (Bakteriologische
Diagnostik, Hamburg, 1891, p. 298) says nothing under the head of aerobiosis.
Howard (Foul Brood: Its Natural History and Rational Treatment, Chicago,
189+) writes that, ‘It grows best under anaerobic conditions; is a facultative
aerobe; grows under the mica plate, and in the presence of oxygen the growth
is slight and slow.” Howard also states that under anaerobic conditions it
emits a foul odor resembling that of foul brood. It will be thus seen that
Cheyne and Howard do not agree on this point. The former author also says
that the characteristic odor is given off under aerobic conditions, whilst Howard
states that this smell is emitted under anaerobic conditions. Further, Cheyne
states that the bacilli grow with great rapidity on the surface of agar, whereas
Howard obtains his best growth under the mica plate, which does not give
complete anaerobiosis. Howard’s conclusions are thus at variance with
Cheyne’s, and my own results fully corroborate those of the latter author.

Howard states that the vitality of the spores of B. alvei is destroyed when
exposed to atmospheric air from twenty-four to thirty-six hours. In making
his experiments he took sterilized road dust and mixed it with the dry foul-
brood masses from several cells, which were previously dissolved in distilled
water. The mixture was worked dry and spread on sheets of paper, and trial
cultures were made immediately and at intervals of every twelve hours for
three days; and, according to his results, no growth occurred after thirty-six
hours. In giving these results, Howard does not state whether he exposed the
spores to sunlight or diffused light; nor does he mention the age of the dry foul-
brood masses, which he used from several cells. These are points of considerable

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 47

importance, for, as everyone knows, the disinfecting power of direct sunlight is
much greater than diffused light, and the vitality of the spores from foul-brood
masses of different ages varies considerably. This, I may add, has been clearly
shown by some of my experiments, subsequently described. In my experiments
the spores obtained from a pure culture on the surface of agar were spread on
cover glasses and placed in a glass chamber, so arranged that a current of air
was constantly circulating over them. This chamber was exposed to the ordi-
‘nary light of a room with six large windows, and a cover glass was taken out
every twenty-four hours and tested, to see if the spores would grow. This
experiment was continued for one month, and at the end of that time the spores
still germinated rapidly. In another experiment, spores spread on cover glasses
were exposed to a very diffused light, simulating, as far as possible, the amount
of light which would enter a hive. Cover glasses were taken out from time to
time and transferred to agar, in order to ascertain if the spores were alive or
not. The experiment was begun two years and four months ago, and from the
last cover glass taken and placed upon the surface of an agar plate a copious
and typical growth of B. alvei was obtained. Further, thin strips of filter
paper, plunged into a bouillon culture and allowed to dry, were threaded on a
wire suspended in a wire basket and so exposed that the air could freely cireu-
late around them in the ordinary light of a room. Trial cultures were made at
intervals, and at the expiration of six months the spores from the paper germi-
nated when strips were placed on the surface of agar.

Again, a drop of bouillon containing spores was placed in a sterile tube and
allowed to Gry; and at the expiration of one hundred and twenty-four hours
(thirty-six of which were in sunlight at a temperature varying from 30° to
37° C.) sterile bouillon was added. The tubes were then placed in the incubator,
and in less than twenty-four hours a good growth of the germs had taken place.

From these experiments it will be seen that the results are directly at
variance with Howard’s statement, as they go to show that the vitality of the
spores of B. alvei is not destroyed by exposure to atmospheric air, with or
without sunlight, for even a much longer time than twenty-four to thirty-six
hours.

With regard to the aerobiosis of this bacillus, good growth has been obtained
in an atmosphere of hydrogen by Novy’s method. Buchner’s method also gave
good results. The growths in the various media are very similar to those pro-
duced under aerobic conditions, but with this difference, that the surface
growths are, as a rule, whiter in the hydrogen atmosphere. In illuminating gas
(water gas) no growth occurred, but the spores were not destroyed by the
action of the gas; for when the gas was let out of the Novy jar, good growth
ensued on all cultures. In acetylene gas, a restricted growth occurred. In
fermentation tubes growth occurred both in the open and in the closed arm
of the tubes. No gas was formed, the bouillon in the closed arm was uniformly
turbid. Thus B. alvei is a facultative anaerobe.

Production of alkalii—in ordinary bouillon a slight amount of ammonia is
formed. Control bouillon did not give the Nessler test. In glycerine and the
sugar bouillons, there is no trace of ammonia. Cheyne’s cultures are faintly
alkaline, both before and after inoculation in meat infusion. Klamann states
that ammonia is produced.

Acids formed.—A varying amount of acid is formed. All the sugar bouillons
give an acid reaction.

Formation of pigment.—On potatoes a yellowish growth is produced; on all
other media, the surface growth is white.

Development of odors.—Cheyne states that gelatine cultures give off an

48 MEETING OF INSPECTORS OF APIARIES.

odor of stale, but not ammoniacal urine, or what may be better deseribed as a
shrimpy smell; and this peculiar odor has been found by Cheshire to be dis-
tinctive of diseased larvee, Klamann and Howard both state that a peculiar
odor resembling that of the diseased larve may be noticed in artificial cultures.

The effects of desiccation.—I have already noticed, under the head of ‘ Rela-
tion to free oxygen,” that the spores of B. alvei have considerable vitality in
withstanding desiccation. My experiments prove conclusively that the spores
are extremely hard to kill by desiccation and in this respect resemble those of
anthrax, which are known to resist thorough desiccation for a number of years.
One experiment which showed this characteristic was as follows: An agar
plate completely covered with a typical growth of B. alvei was allowed to dry
out completely, and was left exposed to the ordinary light of the room for seven
months, and at the end of that time, a portion of the film was scraped off with
a knife, placed on suitable medium and incubated, with the result that a typical
growth immediately ensued.

Spores on cover glasses were exposed to September sunlight (latitude 43°)
for varying periods of time, and growth occurred after four, six, and seven
hours’ exposure. The age of the spores varied from five days to eighteen
months; and spores three months old were not killed by seven hours’ exposure.

From the symptoms given in this paper the disease with which
Professor Harrison worked was doubtless American foul brood.
From the discussion of geographical distribution this is also evi-
dent, for he says: “ I have examined diseased larve from Canada,
from Europe, * * * Cuba, and thirteen States of the Union,
‘ranging from New ‘York to California and from Michigan
to Florida.” American foul brood is thus widely distributed, but
from all these specimens Professor Harrison obtained a_ bacillus
which he called Bacillus alvet. Since we now know that Bacillus
alvei is found in European foul brood and not in American foul
brood, it is evident enough that the germ must have been another
bacillus. European foul brood, as far as the author is able to learn,
is not found in Canada nor Cuba, and, although now found in
several States in the northeastern United States and spreading, is
not, as Professor Harrison would have us believe, widely distributed
in the United States.

How can this be accounted for? The only way open seems to be
in the identification of the bacillus. I do not feel qualified to pass
judgment on the accuracy of the description of Professor Harrison,
but the matter has been referred to Doctor White, and he assures
me that the description just quoted fits the bacillus which is
described as Bacillus A as well as it does Bacillus alvei. If this is
true, we can only conclude that Professor Harrison, not knowing of
the existence of two diseases, made a serious error in his identi-
fication. In no place does he speak of any difficulty in obtaining
cultures from American foul brood. For comparison, Doctor White’s
description of Bacillus A (possibly B. mesentericus) is here quoted.

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 49

BaciILitus A.
(B. mesentericus?)

Occurrence.—F ound very frequently on combs, on scrapings from hives, and on
the bodies of bees, both diseased and healthy.

Gelatin colonies.—Very young colonies show irregular edges, but very soon
liquefaction takes place and the colony gives rise to a circular liquefied area,
covered with a gray membrane, which later turns brown.

Ager colonies.—Superficial colonies present a very irregular margin consist-
ing of outgrowths taking place in curves. Deep colonies show a filamentous
growth having a moss-like appearance.

Morphology.—In the living condition the bacilli appear clear and often granu-
lar, arranged singly, in pairs, and in chains. The flagella are distributed over
the body. The rods measure from 3 to 4 in length, and from 0.9% to 1.2 in
thickness.

Motility—The bacilli are only moderately motile.

Spores.—Spores are formed in the middle of the rod.

Gram’s stain.—The bacilli take Gram’s stain.

Oxygen requirements.—Aérobic and facultatively anaérobic.

Bouillon.—Luxuriant growth in 24 hours, with cloudiness of medium; a gray
flocculent membrane is present. Later, the membrane sinks and the medium
clears, leaving a heavy, white, flocculent sediment, with a growth of the organ-
isms adhering to the glass at the surface of the medium. Reaction alkaline.

Glucose.—Luxuriant growth takes place in the bulb, with a moderate, floccu-
lent growth in closed arm. ‘The gradual settling of the organisms causes a
heavy white sediment to form in the bend of the tube. The reaction is at first
slightly acid, but subsequently becomes alkaline. No gas is formed.

Lactose.—Reaction alkaline.

Saccharose.—Reaction alkaline.

Levulose.—Reaction acid.

Maltose.—Reaction acid.

Mannite.—Reaction alkaline.

Potato water.—Reaction alkaline.

Agar slant.—A luxuriant growth takes place on this medium. The growth
gradually increases to a moist, glistening one, being then friable and of a grayish
brown color.

Serum.—A luxuriant, brownish, glistening, friable growth spreads over the
entire surface. No liquefaction is observed.

Potato.—An abundant fleshy growth of a brown color spreads over the entire
surface. The water supports a heavy growth. The potato is slightly discolored.

Milk.—Precipitation takes place rapidly, followed by a gradual digestion of
the casein, the medium changing from the top downward to a translucent liquid,
becoming at last semitransparent and viscid.

Litmus milk.—Precipitation of the casein takes place usually within 24 hours,
followed by a gradual peptonization. Reduction of the litmus cecurs rapidly,
leaving the medium slightly brown; later the blue color will return on exposing
the milk to the air by shaking. Reaction alkaline.

Gelatin.—An abundant growth takes place with rapid, infundibuliform lique-
faction. A heavy, white, friable membrane is formed on the surface of the
liquefied medium. A flocculent sediment lies at the bottom of the clear liquefied
portion. p

Acid agar.—Growth takes place.

Indol.—None has been observed.

Nitrate.—Reduction to nitrite is positive.

30547—No. 70—07 mu 4

50 MEETING OF INSPECTORS OF APIARIES.

Dr. William R. Howard, of Fort Worth University, Fort Worth,
Tex., has published several papers on the bacteriology of bee dis-
eases. In a paper published in 1894 (York Publishing Company,
Chicago) he attributed “ foul brood ” to Bacillus alvei. Evidently he
was dealing with American foul brood, and we now know that Baci-
lus larve is present in that disease.

The same author undertook to determine the cause of pickle broo«
and described a specific fungus, Aspergillus pollinis. No investi-
gator has since been able to find any such fungus in similar
specimens.

In 1900 (Gleanings in Bee Culture, p. 121) this author published
an account of some brief and entirely inadequate investigations made
on what he chose to call “New York Bee Disease, or Black Brood.”
A specific organism, Bacillus milii, is described, but the view is ex-
pressed that this is modified, perhaps, by Bacillus thoracis. During
the investigations of the Department of Agriculture it has been
learned from whom Doctor Howard got his specimens, and the same
men have furnished specimens which they declare to be of the same
diseased condition as those furnished Doctor Howard. ‘These, how-
ever, contain Bacillus alvei, and the disease is the same as that de-
scribed by Cheyne, now named European foul brood.

It is most unfortunate for Doctor Howard that in not a single
point have his descriptions been verified. Certainly it would seem
unwise in him to put out the names Bacillus milii and Bacillus
thoracis as new species without descriptions and after so short an
investigation. We can not, therefore, sympathize very much with
him when his views are overthrown.

The American bee journals and text-books on apiculture have
until recently contained statements to the effect that Bacillus alvet
is the cause of the disease which has been almost universally called
“ foul brood.” This is due not only to the publications of Mackenzie,
Harrison, and Howard, but very largely also to the attempt to deter-
mine Bacillus alvei by microscopic examination. The best-known
case of this is probably the examination of diseased brood made by
Mr. Thomas William Cowan, editor of the British Bee Journal.
On a visit to Medina, Ohio, Mr. Cowan was shown a sample of dis-
eased brood, and after a microscopic examination he announced that
he found Bacillus alvei, and that the diseased condition is identical
with that found in England. That this ropy type (for such it was)
is found in England can not be doubted, but that the germs which
Mr. Cowan saw were Bacillus alvei may well be doubted. I have
taken particular pains to ask Mr. E. R. Root, who was present,
whether Mr. Cowan made a cultural examination, and was assured
that the microscopic examination was the only one made. |

The announcement of this examination in Gleanings in Bee Cul-

PRESENT STATUS OF INVESTIGATION OF BEE DISEASES. 51

ture and the A B C of Bee Culture, coupled with the excellent repu-
tation of Mr. Cowan, made this appear convincing to American bee
keepers. It must be remembered, however, that at that time no one
had questioned the presence of Bacillus alvei i American foul
brood and on finding bacilli the conclusion that they were Bacillus
alvei was natural, even though erroneous.

Mr. Edward Bertrand, in his book “ Conduite du Rucher,” makes
a similar announcement, stating that he and Mr. Cowan examined
brood described as ropy and found Bacillus alvet.

[Mr. Dapanr: I have received a letter from Mr. Bertrand. He
informs me that they (Mr. Cowan and himself) had examined foul
brood, but I know from the tone of the letter that no cultures were
made. |

To indicate how much reliance may be placed in microscopic exami-
nations 1n the absence of cultural tests, let me quote from Sternberg’s
Text-book of Bacteriology, 1901 edition, pages 13 and 14. It should
be borne in mind that this refers to all microscopic examinations of
bacteria and not specifically to bee diseases.

The bacteria are also classified according to their biological characters, and it
will be necessary to consider the various modes of grouping them from different
points of view other than that which relates to their form. This is the more
important, inasmuch as we are not able to differentiate species by morphological
characters alone. Thus, for example, there are among the spherical bacteria,
or micrococci, numerous well-established species which the most expert micro-
scopist could not differentiate by the use of the microscope alone; the same is
true of the rod-shaped bacteria. The assumption often made by investigators
who are not sufficiently impressed with this fact, that two micro-organisms from
different sources, or even from the same source, are the same because stained
preparations examined under the microscope look alike, has led to serious errors
and to much confusion. As an example of what is meant, we may refer to the
pus organisms. Before the introduction of Koch’s “ plate method” micrococci
had been observed in the pus of acute abscesses. Some of these were grouped in
chains—streptococci—and some were single, er in pairs, or in groups of four;
but whether these were simply different modes of grouping in a single species, or
whether the chain micrococci represented a distinct species, was not determined
with certainty. That there were in fact four or more distinct species to be
found in the pus of acute abscesses was not suspected until Rosenbach and
Passet demonstrated that this is the case, and showed that not only is the
streptococcus a distinct species, but that among the cocci not associated in
chains there are three species which are to be distinguished from each other by
their color when grown on the surface of a solid culture medium. One of these
has a milk-white color, one is of a lemon-yellow color, while the third is a
golden-yellow.

This brings us down to the work of Doctor White. His investiga-
tions were begun with Dr. V. A. Moore in 1902, and in January, 1903,
a preliminary report was published. During the first year speci-
mens of “black brood ” were examined and to the surprise of the
investigators Bacillus alvet was found in every case. Obviously,

52 MEETING OF INSPECTORS OF APIARIES.

then, they were working with the foul brood of Cheshire and Cheyne
(iropean foul brood). In a second short paper by Doctér White
a brief note is given concerning some work done on the “ ropy type ”
of foul brood. He recognized that he was dealing with a disease the
cause of which had not been described and the disease is called for
the time “ X brood” and the bacillus, Bacillus X. The final results
of the investigation appear in Technical Series, No. 14, Bureau of
Entomology, under the title “The Bacteria of the Apiary, with
Special Reference to Bee Diseases.” Doctor White’s description of
Bacillus larve is as follows:

3ACILLUS LARVAE.

Occurrence.—Constantly present in diseased brood from colonies affected with
American foul brood.

Gelatin.—There is no growth.

Morphology.—Ilt is a slender rod, having a tendency to form in chains. This
is especially true when grown in bee-larve bouillon.

Motility —-Vhe bacillus is rather sluggishly motile.

Spores.—Spore formation takes place. This can be observed best in the dif-
ferent stages of the disease and decay of the larvie.

Oxygen requirements.—When Liborius’s method is used, the best growth
usually appears near to but not on the surface. After a few generations a
surface growth may be obtained.

Bowillon.—There is no growth.

Glucose bouillon—There is no growth.

Lactose.—There is no growth.

Saccharose.—There is no growth.

Agar plate.—There is no growth.

Bec-larve agar.—The inoculations must be made with the medium liquefied.
The growth takes place near to but rarely on the surface. Cultures must pass
through a few generations before a satisfactory surface growth can be secured.

Bee-larve agar slant.—On the surface of this medium a thin, gray, nonviscid
growth takes place.

Glucose agar.—Slight growth has been observed in the medium. No gas is
produced.

Potato.—There is no growth.

Milk.—There is no growth.

Litmus milk.—There is no growth.

Fermentation—In bee-larve bouillon no gas is produced.

Indol.—There is no growth in sugar-free bouillon.

To summarize, then, Bacillus alvet is found universally in Euro-
pean foul brood; Bacillus larve in American foul brood. No specific
micro-organisms have been found for the so-called pickle brood or
paralysis. Knowledge of the two werst brood diseases is accurate
enough to enable us to combat them by applying principles acquired
by comparison with results of work with other micro-organisms.

That our knowledge is complete is far from true. Not only is
there much to be learned which is of purely scientific interest, but
points of the highest practical importance are yet undetermined.

GEGGRAPHICAL DISTRIBUTION OF AMERICAN FOUL BROOD. 538

[ Nore.—Several other papers of importance have been issued on
this subject which were not discussed at the Inspectors” meeting.
They, however, have an important bearing on this subject. Lambotte
decided that Bacillus mesentericus vulgatus causes “ foul brood,” or,
in other words, that Bacillus alvei is but a variety of Bacillus mesen-
tericus. His work, however, is far from convincing. The principal
point of interest in this regard i is that he had oreat difficulty in get-
ting a growth from the ropy type of foul brood (American foul
brood ) on ordinary media. Burri in 1904 published an account of
his work and found Bacillus alvei in a few specimens from Switzer-
land (indicating that European foul brood is found there), but
found another organism which grows with difficulty; the latter is
undescribed and unnamed, and it is possible and probable that he
worked with Bacillus larve White. Maassen (1906) found the same
difficulty, isolating Bacillus alvei in only 13 specimens of diseased
brood out of 112 received. He, too, found an organism which could
be made to grow on ordinary media only with difficulty and called by
him Bacillus brandenburgiensis. It is undescribed, so far as is known
to the writer. He also claims to have found another organism, Sp7-
vochwte apis Maassen, but has not established any causal relationship.

These papers all tend to confirm the work of Doctor White. Baci-
us alvet is not found in the ordinary ropy type of “ foul brood,” but
another organism is; this is probably the Bacillus larve of Doctor
White.

In the face of all these facts several prominent bee men of Eng-
land are attempting to discredit all this work, the criticism, so far as
is known to the writer, being based entirely on comparisons of ltera-
ture and on an entire lack of investigation. They have, further, mis-
read the papers issued by the Department of Agriculture on this sub-
ject. It seems entirely unnecessary, therefore, to review the criticism

in detail.—E. F. P.]

EXISTENCE OF BOTH AMERICAN FOUL BROOD AND EUROPEAN FOUL BROOD
IN THE SAME COLONY.

Mr. Arcuitry. Do you think that both diseases, American foul
brood and European foul brood, could exist in the same colony ?

Doctor Puitiips. Reports are sometimes received that a colony is
infected with both diseases at the same time, but this is contrary to
the experience of those persons most conversant with these conditions.
While it may be possible for a colony to have the infection of both
diseases at the same time, it is not by any means the rule, and such
cases are probably not authentically reported.

Both diseases are found in New York State. The inspectors have
to treat both diseases and they treat both in the same way, but they
have never found both diseases in the same colony.

GEOGRAPHICAL DISTRIBUTION OF AMERICAN FOUL BROOD.
Mr. Dapanv. Is not the American foul brood spread more over the
world?

Doctor Puimurrs. It would seem so from the literature. It is
found in almost every State of the Union, while European foul brood

54 MEETING OF INSPECTORS OF APIARIES.

is, as far as is known, found only in the States mentioned a while
back.

Mr. Dapant. The American foul brood is characterized by the ropy
condition. The French name for their common brood disease is
“ loque,” meaning tatters, and this name therefore refers to what we
call American foul brood.

Mr. Cogesuauy. Which of these diseases is the one found in Cuba ?

Doctor Purures. That is the American foul brood. It has the
typical ropy character.

EFFECT OF CLIMATE ON VIRULENCE.

Mr. L. F. Junzau (Colorado). I would like to ask if brood diseases
are equally bad in all States or has climate anything to do with the
virulence of the diseases ? .

Doctor Pinus. Climate undoubtedly makes a great difference.
The American foul brood of California is not anything lke the same
disease in the East. It is simply terrible in California. Mr. Rankin
will tell us about that later in the day. It is the same disease, but
its ravages are much worse.

Mr. Juneau. Mr. T. L. Thompson (Colorado) sent some pickle
brood to Dr. W. R. Howard, and the latter called it * black brood,”
but said: “In your State it will not be so bad.”

Doctor Pumps. It was probably not European foul brood. That
disease has not been found west of the Mississippi River.

ASSOCIATION OF INSPECTORS OF APTARIES.

It was decided that it would be well for the inspectors of apiaries
of the various States to be organized in some way to bring about
greater cooperation in the work. After discussing the question it
was finally decided that Mr. N. E. France, inspector of apiaries for
Wisconsin, should act as chairman of a committee on organization
and appoint his own associates.

SECOND SESSION, HELD MONDAY AFTERNOON, NOVEMBER 12, 1906.

Doctor Pures. In the morning session we covered rather thor-
oughly the scientific side of investigations on bee diseases. That is,
of course, important; but when it comes to the practical work on bee
disease there are two subjects of much greater importance, namely,
methods of treatment and legislation. We will first discuss the treat-
ment of these two brood diseases and then take up the discussion, in
so far as we can, of the laws now existing, with suggestions as to the
form which a law should have to give the best results and the. powers
which should be given to the inspectors under the various conditions
which may arise. I have, however, a paper that I wish to read first,

APIARY INSPECTION IN NEW YORK STATE. 55

which was written by Mr. Charles Stewart, of Sammonsville, N. Y.,
one of the inspectors of that State. I had the pleasure and privilege
of spending four weeks in the field with Mr. Stewart last spring, and
I feel that I can say that if there are any good inspectors in the
United States one of them is Mr. Charles Stewart. Mr. Stewart 1s
exceedingly sorry that he can not be here, and he requested me to read
this paper to you.

APIARY INSPECTION IN NEW YORK STATE.

By CHAS. STEWART,
Inspector, Third Districi,.

BrorHuer Inspecrors: It is with a feeling of regret that I write
this paper, knowing that it will be impossible for me to be with you
at what must be both a pleasant and profitable meeting.

It is hardly necessary for me to describe European foul brood nor
to refer to its entrance into New York State, except to say that it was
brought in some years ago by a shipment of bees from one of the
Southern States, and just as we were feeling that we had nearly
stamped it out and were masters of the situation we discovered that
at least one if not two fresh importations had been made in a section
of the State where no-trouble of this kind formerly existed.

I wish to call your attention to the fact that no bee keeper can feel
reasonably safe from infection until every State in the Union is
under the surveillance of a keen-eyed inspector who knows every spot
of disease in his jurisdiction and allows no bees to be shipped out of
such territory. Had the inspectors of New York State not adopted
this rule, disease would have spread not only all over our State, but to
far distant points, as many, fearing the loss of their apiaries, were
eager to sell at a sacrifice. In order not to make this rule a hardship
to our people, we have made it a practice to find a buyer within the
diseased territory competent to cure the disease and so keep our
troubles within our own family.

IT wish I had the power to paint to you in words the pathetic
picture when four good men and true, who had been bee keepers from
boyhood and had large interests to protect, took up this work. I
have seen the faces of strong men blanch with fear or turn crimson
with anger at the first visit of an inspector, and later, when their bees
were saved and their product marketed, the young man sent back
to college, their little children cared for, or perhaps the home saved,
these same men with tears on their cheek would give one a hand clasp
that was far more eloquent than words and possesses a value beyond
gold. I question if there is an inspector present to-day who from a
mere money point of view would not be better off if he had given

56 MEETING OF INSPECTORS OF APIARIES.

his entire time to his own business; yet I honor this American spirit
you possess in that, having once started out to accomplish results,
you refuse to turn back until the end is attained.

I hardly feel like posing as an instructor to this gathering of in-
spectors, but will call your attention to a few important points. A
question often asked is, ‘* How does.the disease spread so rapidly?” I
would answer, “ By means of infected honey.” No field bee from an
infected colony goes out with its honey sack so empty of honey that
it contains no germs, and on their return many bees mistake their
hive and carry disease to their near neighbors in the same apiary, so
that it is a common thing to find a badly infected colony and those
in the same row infected in proportion to their distance from the
source of contagion.

How the disease spread from yard is yard when no robbing took
place was for a long time a puzzling question, until I found an
apiary of black bees 3 miles from an apiary of golden Italians that
were infected. In many colonies of the yard of blacks could be found
a sprinkling of the golden Italians and in nearly every case these
colonies showed traces of disease. Evidently bees are often driven
by stress of weather or some other cause to seek shelter far from
home, and thus disease may be spread.

We have found no bees immune from disease, yet some vigorous
strains of Italians are nearly so. For years we have recommended
the introduction of young Italian queens, but have warned the owner
of an infected apiary not to depend on that alone, as it would prove
disastrous in localities where the disease has just made its appearance
and the bees are mostly black. This method will often prove very
successful where European foul brood has existed for some time and
lost much of its virulence, but, like the use of drugs, it is not a safe
method for the inspector to advocate, while the shaking method has
never failed us if done in a thorough manner. Colonies that are
found to be diseased late in the season may be cured by taking away
all their combs after brood rearing has ceased and giving them clean
combs from a healthy colony, as any disease germs that are contained
in the honey sack will have been eliminated lone before brood rearing
commences in the spring.

In conclusion I would say that to be successful as an inspector a
man should not only be well versed in the management of bees and
bee diseases, but he must be broad minded, even tempered, possess a
liberal amount of tact and diplomacy, and be a shrewd judge of
human nature. Yes, and even more, he should be able to win the con-
fidence of others and share their burdens, and when the time draws
nigh when the working tools of life shall drop from his nerveless
clasp he may look back with satisfaction to a life well spent in the
service of others.

AMERICAN FOUL BROOD ON THE PACIFIC COAST. ie

Doctor Puitiies. When the notices were first sent out concerning
this meeting I wrote to Mr. Fred A. Parker, of Lompoc, Cal., under
the impression that he was inspector of Santa Barbara County. It
seems that he has resigned and Santa Barbara County now has no
inspector. He has, however, sent a paper, which will now be read:

AMERICAN FOUL BROOD ON THE PACIFIC COAST.

By Frep A. PARKER,

Former Inspector for Santa Barbara County, Cal.

During my term as inspector of apiaries for Santa Barbara County,
Cal., in the year 1905, 4,073 colonies of bees were inspected. I dis-
covered 47 cases of American foul brood and found 170 colonies not
on movable frames. These were ordered transferred, and in some
instances I did the work myself. Every case of foul brood was
either burned or treated by the shaking method. Five were burned,
being too weak in numbers to treat. Preparatory to burning, I would
dig a hole about 2 feet deep and build a fire in it, then throw in the
frames containing the diseased brood. After the fire did its work
the hole was filled with dirt to prevent bees from getting diseased
honey, if any might have been left unconsumed. If any comb was
attached to the hives the latter were placed on the fire in the hole
and when the interior was a mass of flames they were removed with a
shovel, hoe, or other long-handled tool and water thrown on to ex-
tinguish the flames if the wood had caught fire. The bees were
shaken into an empty hive and allowed to build comb on the hd for
three days, when they were shaken onto frames with starters and
allowed to proceed. The comb was scraped from the lid and the lid
scorched. This treatment, if carefully performed, is successful in
about nine-tenths of the cases treated. Bee keepers are generally
too careless in handling the diseased combs, thus giving other bees
an opportunity to steal a load of infected honey.

I have read many statements to the effect that queens do not carry
the infection, but my experience has convinced me otherwise. I had
shaken six diseased colonies in my own apiary in 1902 and four
were completely cured. While I was equally careful in handling
these cases the disease reappeared in two of them. I shook them
again, and again in due time the disease appeared. This caused me
to suspect that the queens were defective, and to test it I exchanged
them with the queens of two perfectly healthy colonies, shaking the
diseased stock again. In both cases the cure was complete, while the
(disease appeared in the brood of the formerly healthy colonies. This
appears to me to demonstrate beyond doubt that the ovaries of queens
are occasionally infected, that their eggs transmit the germs of Ameri-
can foul brood, and that the disease will develop in any colony to

58 MEETING OF INSPECTORS OF APIARIES.

which they are introduced. If this occurs in one-third of the cases
or one-tenth, it will pay to requeen in every case, unless you have an
especially valuable queen you want to save, in which case it may pay
to experiment. For that reason I now practice requeening every
colony treated for American foul brood.

My experience with drugs has been unsatisfactory in every case.
IT have tried carbolic, rosemary, Bingham’s sulphur plan (as outlined
in Gleanings in Bee Culture, April 15, 1902), the formaldehyde spray
plan (Gleanings in Bee Culture, December 1, 1903), and naphthol, but
while all these drugs have the effect of checking the disease and pre-
venting its spread over the combs as long as used, none of them cures
it, regardless of the duration or persistence of its application.

I have not tried the formalin-gas plan, nor do I intend to try it, or
any other drug treatment, so long as the shaking treatment will cure.
While destruction of frames and combs is expensive, it is to my mind
cheaper in the end than experimenting with every new cure that is
exploited in the bee journals. After trying these you are forced to
resort to the shaking treatment to make the cure complete, so why
not use it at first and save the trouble and expense? So long as honey
contains spores, so long will drugs fail, because they can not reach
and destroy the spores. Even if a temporary cure could be effected
the disease would reappear when the bees began feeding the larva
this germ-laden honey. Nothing short of removing all the combs
will make the cure permanent.

As an apiarist I have had experience in many infected apiaries,
and in every yard where the disease has ever been, with one excep-
tion, a few cases develop every season, and will continue to do so
until these old combs are retired. If a whole apiary is to be
treated, it pays to save the wax and honey, but I do not believe in
bothering with them if only a few hives are to be treated; it does
not pay to take the trouble. Of course progress is desirable, and I
would not discourage anyone who wants to experiment with drug
treatments, but I believe if any good is ever derived therefrom, it
will come from the work of experiment stations or trained scientists,
who have the means and time to devote to it and do not have to
depend on apiculture for a living.

American foul brood seems to act differently here than in most
places. The question may arise, Is it American foul brood? It
has the sunken, perforated cappings and the foul, glue-like odor,
and it ropes from one-eighth to several inches. I have seen many
cases where the brood chamber was badly affected with foul brood,
but when a honey flow came, the queen moved up and not one cell
of disease appeared. I have known these bees to cast strong swarms,
which proved to be entirely healthy.

Again, I have known American foul brood to disappear without

AMERICAN FOUL BROOD ON THE PACIFIC COAST. 59

any treatment whatever. Mr. B. Dickens, one of the most intelligent
and observing apiarists, had marked a colony for treatment. Not being
able to attend to it for several weeks, he was surprised when he did
open it to find every trace of the disease gone. I had the same expe-
rience this season. The most amazing case of this character, however,
was the experience of Mr. W. J. Oates (now my business partner) in
1903. He purchased an apiary of 30 colonies, nearly every colony
being badly affected with foul brood. The former owner, Mr. J. H.
McGee, desired to get rid of the colonies, not caring to go to the
trouble of shaking them. Mr. Oates treated the whole apiary by
the shaking treatment. As soon as there was sealed brood in the
hives, it was seen that disease had developed in about three-fourths
of them. I examined these colonies myself, and if they did not
have the disease after the shaking, then I never saw a case of Amer-
ican foul brood. Mr. Oates did nothing more to them, and, becoming
disgusted with the proposition, he sold out to Mr. F. S. Moorehead
and went to Nevada. The year 1904 was a poor season here, and
honey was extracted from these hives once, I think. Nothing was
done for the disease. In 1905 I inspected these bees, expecting to
find them reeking with disease, but to my surprise I could not find
a single case of foul brood; it had completely disappeared. Mr.
Oates was surprised when informed of this, but he managed this
apiary that season on the shares, and no disease developed. I had
occasion to look through this apiary just last week, and not one case
of disease exists there to-day. That is a case I can not understand,
unless it is that by the shaking the bees were relieved of all diseased
honey, and, being shaken in the fall, the queens ceased laying
entirely later, and the: bees cleaned out all infection. But I am
unable to account for the wholesale reappearance of the disease,
unless the treatment was carried out in a careless manner. I am
certain that the circumstances occurred just as related. The Simmins
plan is not a drug plan, and I intend to test it next season, if I find
any American foul brood.

Sometimes disease spreads quite rapidly here, infecting one-half
or more of the colonies in two seasons. Then I know of some apiaries
where a few cases have existed for years without any perceptible
increase. I know of one instance where an apiary was entirely
destroyed by the disease in one season. Whether our climatic con-
ditions have anything to do withthe matter I do not know, but it
is a fact that foul brood as it exists here is of a very erratic nature.
Furthermore, it is dangerous, and a relentless war should be waged
against it until it is exterminated.

A paper entitled “ The appointment of inspectors,” by Fred A.
Parker, of California, was then read, in which the writer showed

60 MEETING OF INSPECTORS OF APTARIES.

that the work of the inspector is far from easy. Many bee keepers
criticize the work of the inspector as soon as their apiaries are exam-
ined, and fault is found with inspection and the inspectors. It is the
duty of the bee keeper to uphold the inspector as long as he is doing
honest work for the bee-keeping industry. The salary paid an
inspector is in most cases smaller than the income he could make by
remaining at home and doing the required work in his own apiary,
so that inspection is usually done at a financial loss to the inspector.

Mr. J. M. Rankin (California). It has been my privilege since
May, 1905, to be in touch with bee-disease work on the Pacific coast.
During this time I have visited many diseased apiaries throughout
the State of California.

Few eastern people have an adequate conception of the bee-keeping
industry in California. It is not an uncommon thing for one man to
own 4,000 colonies of bees. This, of course, puts the business on an
entirely different footing than in the East. In the same way, condi-
tions of disease are also different. The control of American foul
brood among so many colonies becomes a much more difficult proposi-
tion than it is where the bee keeper owns only fifty to seventy-five col-
onies. There seems to be no doubt, also, that the American foul brood
is much more virulent in California than in the East. Whether this
is due to some climatic condition or not, I do not know. I have seen
an apiary showing only slight infection in February become almost
a total wreck in August. In California, also, the bees fly nearly 300
out of the 365 days in the year, and the honey flow in most parts of
the State is of comparatively short duration. This makes conditions
favorable for more rapid infection than in colder climates where the
bees are confined to their hives during about half of the year.

Under such conditions you can readily see that treating the disease
is difficult. It must be done at exactly the right time and under fay-
orable conditions or the treatment is worse than useless. Some of
the best inspectors in California use the shaking treatment, and all of
them shake twice, as well as disinfect the contaminated hives. There
ure some few men who do not believe in treating by this method and
who burn all diseased colonies, only saving hives when these are in
good condition. In counties where bees can be bought for 50 cents
a swarm it may not be a bad plan to destroy all diseased colonies, as
this is certainly an effective treatment if the burning is complete.

A treatment very favorably thought of by some is that of thor-
oughly boiling all diseased material. A large tank is used and the
diseased colonies, after having been sulphured the night before, are
carried to the tank and all the combs thrown in. After all the wax is
melted, the frames are removed from the tank and placed on the fire
under the tank as fuel. This is certainly an effective way of eradi-

TREATMENT FOR BEE DISEASES. 61

ating the disease and can be recommended more highly than the
burning plan, as by this means the wax is not destroyed.

California has the county system of inspection, and probably the
smallest number of colonies which one inspector has to look after is

30,000. From that number the colonies run up to 150,000 in a single
county.

Doctor Puimuirs. What Mr. Rankin has just said is in line with
my own observation during the middle of the summer. I visited one
apiary in Ventura County, with Mr. A. G. Edmondson, the inspector
for that county, and he showed me 151 hives. Two years ago this
aplary was in the hands of a competent bee keeper and no disease
was present. Ventura County is so large that the inspector can
cover only one half in one year and the other half the next year.
When we examined the apiary we found 15 healthy colonies and 136
hives in which the bees were dead or nearly so.

TREATMENT FOR BEE DISEASES.

In discussing the methods of treatment, it would be a good plan
to call on each of the inspectors present and get each one to tell what
method he employs. We should hear first from Mr. N. E. France,
inspector from Wisconsin. He is the oldest inspector in the United
States.in point of service.

Mr. France. Referring to the paper which was just read, I have
tried some of the methods of using drugs in the apiaries of competent
bee keepers and invariably all these methods are failures in Wis-
consin. The fumigating with formalin seemed for a time to check
the disease, as did also some of the other drugs, but in the end they
all are failures. The one method that has given universal satisfac-
tion we owe to the oldest inspector in America, Mr. William McEvoy,
of Ontario, and it has often been termed the “ McEvoy method.”
The plan is to remove the bees from the infection and keep them away
long enough to use up whatever infected honey there is in the stomach
of the bee.

IT am not satisfied to stop with finding disease in a yard and imme-
diately prescribing treatment. In fact, I seldom, after looking over
the yard and finding the disease, begin to prescribe treatment. for I
feel that we are not yet ready for it. What is the use of treating
when some neighbors might have diseased colonies? Take a wide
circuit; then treat at once all colonies having disease. This has some-
times vexed the bee keepers, for they want me to stay and show them
what to do at once, but I tell them that I see no good in treating
colonies while leaving another source of infection.

I try first to instruct the owner of the bees to be careful in his man-
agement. If, in my judgment. he is one who keeps the apiary clean,

62 MEETING OF INSPECTORS OF APIARIES.

and if I can depend upon him, I sit down and go over the “ McEvoy ”
plan with him very carefully, asking him frem time to time if he
understands it. If he says that he does, I say: “ Now, I am your
student; tell me what to do. When you can tell me what you are
going to do, I will trust it to you.” In nearly all such cases they have
treated it without my assistance, and cured it. I can not recommend
anything better than the ‘“ McEvoy ” plan.

Doctor Puiurps. There is just one thing I should wish to add to
that. The treatment of taking bees from the infected combs was
originated in 1769 by Schirach, as nearly as I can find out, and if we
are going back to give credit to the originator of this plan, Mr. Me-
Evoy is not the man to get that credit.

Mr. G. W. Yor« (Illinois). Was not the plan original with Mr.

McEvoy ?
~ Doctor Purures. It was probably original with him, but it was
advocated long before in many European works.

Mr. Suirn. The ground has been thoroughly covered by Mr.
France. Two years ago Mr. France said to me: “Now, Smith, I
have tried almost everything, but I find the ‘ McEvoy’ plan the best.
My advice is to use the ‘ McEvoy’ treatment, as I have done.” I have
only had one case this year where I have had:to make a second
transfer, and I found that to be due to infection from a neighbor’s
colony that I did not get to treat the first time, but which subsequently
was treated, and the bees were all right. I have no trouble, and I
have great confidence in shaking. I don’t alarm the bees. I shall
give my method. In treating a diseased colony I use an extra hive,
to which the bees are to be transferred, and an additional empty hive.
in which I place the infected frames after the bees are shaken from
them. The last mentioned is covered with a cloth to prevent other
bees from robbing. First I move the old infected hive back, and in
its place put a clean hive containing clean frames, with strips of
foundation. The frames are lifted from the old hive, shaken in front
of the new hive, and then covered up in the third hive, which is used
to store infected frames. This is all done in the middle of the day.
If there is no honey in the field, the colony should be fed well at
night.

Mr. J. Q. Svone (Illinois). How do you treat the old hive?

Mr. Smirn. I either burn out the hive, paint it with kerosene oil
and have it burned out, or wash it in strong salt water.

Mr. Frev Murs (Ohio). When you shake the bees, they carry over
honey, do they not?

Mr. Suiru. I set the hive right on the ground. I do not jar the
frames hard enough to jar out the honey.

Mr. Murn. You shake them off during the middle of the day. Is
it not better along toward evening ?

TREATMENT FOR BEE DISEASES, 63

Mr. Surrn. If you wait till evening you will never get through.

Mr. Murnu. Do you use smoke in that operation ?

Mr. Suirn. I use no smoke.

Mr. Mourn. How long do you keep the bees on the strips of founda-
tion; do you feed them right away ?

Mr. Smiru. Yes.

Mr. Mutu. You don’t believe in starving them at all?

Mr. Smirn. No, because the bees coming from the fields are loaded
with honey.

Mr. Mourn. Do I understand that the bees have these bacteria all
over them ?

Doctor Putuuips. Yes; they have the contamination on them.
When they are shaken they of course have it all over them, and when
they are shaken off they doubtless take the bacteria with them.

The McEvoy system is the radical treatment of shaking twice, which
the majority of bee keepers do not use.

Mr. Yorx. If I mistake not, Mr. McEvoy recommends the second
shaking.

Doctor Puitiies. He recommends the second shaking after the bees
begin to drop from starvation.

Question. What do you do with the unhatched brood m the infected
hive?

Mr. Smiru. My recommendation is to destroy the whole thing.

EFFECT OF REQUEENING ON DISEASE.

Mr. Dapanr. Has removing the queens any value in treating the
two diseases? Alexander, Simmins, and others have recommended
removing the queens. Is this of any value in either clisease ?

Doctor Puiures. As has been stated before to-day, I spent four
weeks last spring with the inspectors of New York State in the field.
Both American foul brood and European foul brood are found in
that State, but practically the same method of treatment is advocated
by the inspectors for both diseases. Colonies found to be diseased are
shaken according to the method which has been described several
times in this meeting.

In order to save any healthy brood which is found in colonies
infected with disease, the sealed brood from several colonies, four
to eight, is piled up in hive bodies above a weak colony which is
diseased. In seven to ten days all the brood which is worth saving
will have emerged and the weak colony will have been changed to one
strong enough to treat. This colony is then treated by the shaking
method as were the others. There is no necessity of waiting more
than ten days, for brood which was unsealed when the brood was first
attacked will scarcely be fed sufficiently to emerge.

In addition to this treatment, the inspectors recommend the in-

64 MEETING OF INSPECTORS OF APIARIES.

troduction of young, vigorous Italian queens from good stock. It
has been shown repeatedly that Italian bees are less liable to disease
than most of the black bees, especially of degenerate stock, as is so
much of the black stock when no attention is paid to improvement.
In a pamphlet issued in 1903 by the inspectors of New York the
introduction of Italian brood was recommended. This is not
advocated as a cure, however, but merely as a means of protecting the
colony against future infection.

Reference has been made to the introduction of Italian queens as a
method of curing disease, and to this method the name of Mr. Alex-
ander is attached. In the article in which Mr. Alexander first
advocated the plan he says, in part:

“How to rid your apiary of black brood” (By E. W. Alexander) .¢

This cure is on the line of introducing new blood into the apiary, * * *

Go to every diseased colony you have and build it up either by giving frames
of maturing brood or uniting two or more until you have them fairly strong.
After this, go over every one and remove the queen; then in nine days go over
them again, and be sure to destroy every maturing queen cell or virgin, if any
have hatched. Then go to your breeding queen and take enough of her newly
hatched larvee to rear enough queen cells from to supply each one of your dis-
eased queenless colonies with a ripe queen cell or yirgin just hatched. These
are to be introduced to your diseased colonies on the twentieth day after you
have removed their old queen, and not one hour sooner, for upon this very
point your whole success depends; for your young queen must not commence
to lay until three or four days after the last of the old brood is hatched, or
twenty-seven days from the time you remove the old queen. If you are very
careful about this matter of time between the last of the old brood hatching
and the young queen commencing to lay, you will find the bees will clean out
their breeding combs for this young queen, so that she will fill them with as
fine healthy brood as a hive ever contained. This I have seen in several hun-
dred hives, and have never seen a cell of the disease in a hive after being
treated as above described.

It is not necessary to remove any of the combs or honey from the diseased
colony; neither is it necessary to disinfect anything about the hive. Simply
remove the old queen, and be sure the young queen does not commence to lay
until three or four days after the old brood is all hatched. This treatment
with young Italian queens is a perfect cure for black brood.

In regard to those old queens that were formerly in your old hives, I think it
best to kill them when you first take them from their colonies—not that the
queen is responsible for the disease, for I am sure she is not; but a young
Italian queen that has been reared from a choice honey-gathering strain is
worth so much more to you that I can not advise saving these old queens.

I have experimented along this line considerably, and found, after the colony
has been without a queen twenty-seven days, as above directed, it will usually
be safe to give them one of these old queens, and the cure will be the same.
Still, there have been exceptions, so I advise killing them at once.

The essential point in the treatment is to allow several days to
elapse after the emergence of the last of the healthy brood before
the queen begins to lay.

aGleanings in Bee Culture, November 1, 1905.

TREATMENT FOR BEE DISEASHS. 65

There are several points in this treatment and its successful appli-
cation by Mr. Alexander which may well claim our attention. In
the first place, the scales formed by the dried larve of European
foul brood are less adhesive than are those formed when American
foul brood is present. It is therefore easier for the bees to clean
out the cells, and in most cases, at any rate, a strong colony would do
this. This is one point, then, in favor of the Alexander treatment
of European foul brood.

Mr. Alexander’s apiary is located in a portion of New York State
(Delanson, Schenectady County) where European foul brood has
been prevalent for several years. It is a matter of common observa-
tion that this disease becomes less virulent in any given locality
within a few years, and it is very probable that this plan might be
successful in Mr Alexander’s apiary and not in localities where the
disease is just appearing. At any rate, it is unwise to advocate its
use In new regions when there is an established remedy
method.

The hives used by Mr. Alexander seem to me to have a decided
bearing on this subject. They are several inches shorter than the
Langstroth hive, and,“as a result, in the spring, when European foul
brood usually appears, there is not a large supply of honey on hand.
This, taken into consideration with the fact that very little honey
comes in before August 1 in that locality, is very significant. The
hive is not full of infected honey, and consequently when the bees
clean out the combs they get all the infected material present. That
this method would be successful in a moderate-sized hive—e. g., a
10-frame Langstroth—may well be doubted, for in the twenty-
seven days during which the colony is left queenless many cells
containing contaminated honey would be left untouched. Either
we must advocate very small hives or advise against the Alexander
method as a cure.

The New York inspectors say that the publication of the Alexander
plan has been a great detriment to bee keepers.

Mr. France. I visited a yard last year where there were 22 infected
colonies. The owner wished to save some new drawn-out combs that
were on hives free from the disease. As an experiment we used foun-
dation with half of the colonies and in the others we put the new
combs. Eleven had to be treated again, while the others, right in
the same yard, did not. You can kill the germs in the honey, but
you have to boil it until it is as black as molasses to do it.

Mr. Louis Scuortni (Texas). I do not know that I can say much
about treatment in Texas. We do not rely on the shaking treatment
at all. Whenever we have had foul brood we have tried something
as radical as could be practiced—that is, the burning of the dis-
eased colonies. There is one trouble that we have here in shaking

80547—No. 70—07 m——)

the shaking

66 MEETING OF INSPECTORS OF APIARIES.

the bees, and that is that if we treat the bees during the honey flow
there is so much danger of shaking out the honey and starting the
disease again in that way. The other thing with which we have to
contend is robbing. During a honey flow there is a good deal of
inside robbing almost all the time. Until we find something that
is absolutely sure and absolutely a good thing we shall resort to the
burning of colonies whenever we find them infected. The way we
use the fire treatment is to inspect the yards and then toward evening
we dig a pit about 10 feet wide, according to the number of colonies
to be treated, and build a brush fire. By the time we have that
burning well we go to the colonies that are to be “ treated” and use
sulphur in a smoker. The entrance is smoked a little, and this kills
all the bees. We go from one colony to another to kill the bees, to
keep them from leaving the hives in handling; we know that no live
bees can escape from those colonies. We remove the combs and burn
them, then the bottom boards and the covers are treated over the
flames. The hive bodies are stacked on a single bottom board, and
from a small can of kerosene we pour just a little oil from the
top down the sides; by throwing in some dry grass or anything of
that kind, which has been hghted first, the fire will start at the bot-
tom and the hive bodies will act as a chimney. In that way we
scorch the hive bodies for a few minutes. As soon as these have
been scorched sufficiently we close up the top with a bottom board or
cover and close the entrance of the hive with earth; then we leave
them for a little while for what we call “ steaming.” -

Mr. Anperson. Is there any way of safely detecting American
foul brood before the cells are broken, and how long is it after it is
sealed before the cap is broken? That is a question I have been dis-
cussing at home, and I would like to know if there is a way that it
can be detected. For instance, if you have not treated a colony suc-
cessfully, or suppose American foul brood has been in your locality
and you are waiting for it, can you catch it before all the larve are
exposed ?

If there are only two or three diseased cells in a colony and if you
cut those out, will the disease go any farther? I have read that if
the cell cappings are broken and you take out those particular cells
you will never see the disease again in that colony. I have heard an
inspector say that he can tell the disease in his own apiary. He
claims that there is a way to tell it before the capping is broken, and
he says he can take away the disease then and it will not reappear.
I know he can, because he has proved it. He can tell where foul
brood is before he can actually see it. He further says that the
larve are killed, but do not show it for forty-two days afterwards.
Now, I want to know if anyone else has found such to be the case.

Doctor Puiuures. I think his record stands alone.

TREATMENT FOR BEE DISEASES. 67

Mr. Anperson. I know this: If you cut this foul brood out before
there is another exposure, you won't get it in that colony unless it 33
carried from somewhere else. I have proved that.

Doctor Puiiures. As far as the forty-two days’ time is concerned,
I have no faith in it, because in most cases inside of forty-two days
the colony would be dead. I have seen that demonstrated.

Mr. Hotexame. I might ask how early can the disease be dis-
covered ?

Doctor Puitiurps. Not sooner than the ropiness of the larve be-
comes evident. I never saw a sample of diseased brood from
Texas, but, assuming for the moment that the conditions in this
State are similar to those in California, the method described in the
East is not going to work in Texas. It will work where the disease
is not virulent. The same thing holds true for European foul brood.
Where it has existed for five years it 1s easily treated, and the Alex-
ander treatment is sometimes successful, but it is not when the dis-
ease first appears in a locality. As you know, European foul brood
started in New York and is spreading to the Vermont line. You
will find a great difference in the type of disease in Schoharie County
and on the Vermont line. The same thing seems to hold in a differ-
ent way for the American foul brood.’ The disease is much more
easy to combat in the East than in the West. I visited California
this summer. Inspectors there have proved to their satisfaction that
Eastern methods are not satisfactory, and they told me that it is
necessary to burn out the hives. Mr. Smith does not burn his hives,
and the inspector in New York does not burn hives; they insist, how-
ever, that no honey and no wax cells remain in the hives and that the
hives be clean. That does not prove satisfactory in California. We
know that this one disease is a very different proposition under dif-
ferent climatic conditions, and in discussing treatment it is neces-
sary to bring out this point. In discussing treatment in bee jour-
nals writers forget or do not realize that the plans which they advo-
cate may not do in different places. As Mr. Parker said in his
paper, the eastern treatment will cure nine-tenths, but the other
tenth has to be taken care of. The disease seems to be much more
virulent in the western part of the United States than in the east-
ern part.

Mr. L. Scuotn. Our conditions are the same here as in California,
Tam sure. We have tried some of the shaking treatments, but they
were unsatisfactory. On account of the character of the disease here,
we think we are on the safe side in using the burning method until
we can find something better. While Mr. Smith and others gave their
method of shaking the bees, I wish to put the question whether these
treatments would work west of the Mississippi River, and that is
why we have been practicing such radical measures here. My brother,

68 MEETING OF INSPECTORS OF APIARIES.

who is here from the Agricultural and Mechanical College of Bryan,
Tex., working for Professor Conradi, has been conducting experi-
ments on this shaking treatment, and you might get him to tell you
something about it.

Mr. Ernest Scnorn (Texas). Iam glad to be called upon, because
T have been paying close attention to the shaking treatment, and as
soon as Mr. J. Q. Smith mentioned his method of shaking but once,
I thought surely he is dealing with different conditions or it must
have been an accident that he succeeded. My work has been mostly
in the northern part of the State, but in one case I had some work in
the central part. I thought I would try some experiments. We
tried shaking once, but it would not work; the disease appeared just
as badly as ever. We tried shaking twice; that worked better, so
that shows that shaking once does not work here. I have tried many
other expériments, and am still on the go, but this is the only point I
want to bring out. Shaking once is not sucessful in Texas, and I don’t
think it ever will be. I don’t see how Mr. Smith can be successful
in treating, because the bees gorge themselves with honey. Down
here, as soon as you open a hive the bees will run to the cells and.
consequently, shaking once would not work; and, as my brother said,
there is always some honey taken up and the bees carry it into the new
hive.

Mr. Juneau. Mr. Smith’s plan is satisfactory in Colorado. We
shake our bees there, but we smoke them a little bit and we shake only
when a honey flow is on. The honey will sometimes drip on the
wings of the bees, but it is very seldom that foul brood starts again.
I have been an inspector there for a number of years, and the general .
way 1s to shake the bees hard. We shake them a little bit differ-
ently. We put paper down and we shake when the honey flow is on
and we save nearly all the brood—that is, the healthy brood—and let
it stay twenty-one days. The reason for letting it stay so long is be-
cause there is honey around and the bees hatching out will use it.
Not only do the inspectors instruct that shaking be practiced, but the
State association has issued pamphlets, in which this treatment is
explained, to be given by the inspector to each man who has foul
brood.

Mr. D. C. Miram (Uvalde County, Tex.). In our locality we are
governed by conditions. If the conditions are not favorable for
shaking, we burn the bees, frames, and all. If the conditions are
favorable, shaking is all right. Last May I shook five colonies in
one apiary for experiment, and week before last I went there and
they were all right, but honey was not coming in fast.

I wish to speak of another thing. In this apiary I watched espe-
cially to see if there was any disease of the unsealed brood and I

TREATMENT: FOR BEE DISEASES. 69

found none. Two years ago I found the disease both in the sealed
and unsealed brood, and the question comes up: Have we both dis-
eases, the European and the American foul brood? I began to hunt
for another disease attacking sealed brood, and I found it that year;
but I looked further, and in 65 colonies which I shook off last spring
you could not find disease in any unsealed brood. Last fall a
year ago I went to one apiary that had several colonies in which the
sealed brood was diseased. I told the family what they could do.
TI said: “ You will either have to fight this disease and take care of
the colonies through the winter or you can burn them up.” I will
say they were not bee keepers, and they said just to burn up every-
thing. I agreed to this, but said that there were two colonies in the
aplary that had only a few cells diseased, and I would experiment on
them—that J would take them under my own management. I
burned the rest, but I kept those two colomies until this spring. This
spring they became weak and I set one colony on top of the other.
Last week I went back there, although I had examined them some
time ago, and they had starved to death.

One shaking, I am sure, will do under favorable conditions, but if
the bees are not gathering honey, I would not advise shaking.

MEDICATION.

Mr. Dapanr. Has anyone ever tried feeaing medicated sirup?
The reason I ask the question is because some people succeed with
drugs.

Mr. Surrn. Mr. Reynolds was the first man in [lhnois who im-
ported Italian queens. He said that after foul brood got into his
bees and destroyed them he heard of a remedy that could be obtained
at the drug stores, and the next time he transferred his bees he used
this and he had good luck with them.

Mr. Dapanr. After shaking them ?

Mr. Surry. Yes, sir; and he ordered some of this drug from St.
Louis just a short time ago. He said he was going to feed it to the
bees next spring for fear they would develop the disease again.

Mr. Upo Torrrrrwetn (Texas). It is a good idea to feed the bees
sugar and naphthol beta.

Mr. Arcutey. As Mr. Scholl has already stated, I don’t believe
treatment will eradicate foul brood in Texas. I have seen a few
people that have experimented this season in shaking bees. We have
never been able to determine results in shaking in one season. I have
had the disease disappear in the summer and fall and the next year
the colonies would be diseased again. Another point, in Texas bees
are too cheap to treat. We can burn them and buy other colonies to
replace them with less expense.

70 MEETING OF INSPECTORS OF APIARIES.
PICKLE BROOD.

Mr. Suir. Is there anyone present whose bees have been suffering
from pickle brood ?

Mr. Dapanvr. Mine have, and I used oil of eucalyptus. I thought
IT had foul brood and I afterwards discovered that it was pickle
brood. About every four days I fed some oil of eucalyptus and in
three weeks there was no trace of the disease.

Doctor Puiiures. Would not that disease have disappeared without
the use of drugs?

Mr. Dapantv. I doubt it. I asked another bee keeper to try the
same thing and the result was the same.

Doctor Puitiurs. We have no proof that pickle brood is at all
infectious. Oil of eucalyptus is a disinfectant; therefore I was won-
dering what effect it had.

Mr. Horexamre. One of the members of the Missouri State Bee
Keepers’ Association, who was about 20 miles from St. Louis, asked
me to come over and help with his bees. He said last spring that his
bees were in a terrible condition; he was very busy and did not know
what to do. A good many of the colonies were in bad condition. He
put a tablespoonful of carbolic acid in a quart of water and sprinkled
his bees with this. He told me they were all well except two colonies.
He said he did not look at them. I looked at them and they were
clean. He told me that he had colonies that had gathered in five
days a super full of honey. He had about 10 square miles of Spanish
heather, but these colonies that had been affected did not make any
surplus, so there must have been some disease.

Doctor Puiuurrs. It might have been pickle brood. Pickle brood
is sometimes pretty bad, but it will disappear.

EXPENSE OF TREATMENT.

Doctor Pumures. Is it so expensive to treat bees? How much does
the colony lose by shaking during the honey flow ?

Mr. Anprrson. We lose a honey crop. Take all that brood away
from a colony and all that remains is the live bees. For ten days
there is no brood started to take the place of what has been removed.

Mr. Junzav. It is altogether different in our country (Colorado).
We shake bees, and they act just like a new swarm. I have had as
high as two or three swarms from those that have been shaken, if
they were ordinarily good strong colonies, and I believe it will do
just as well to shake a colony during a honey flow as any time. It
makes no difference.

Doctor Puinums. That is the point I was about to mention. I
know that in Colorado they sometimes shake bees whether there is
disease or not, because they claim the bees do better. You talk about

WR GIES &

TREATMENT FOR BEE DISEASES. G1

shaking bees being a very expensive operation, but you do not need
any brood during the honey flow, and the time makes a great
difference.

Mr. Yor. There is one thing to be taken into consideration. Bees
are worth less per colony in Texas and California than in the East.
When you talk to a man here about burning 30 colonies, it does not
take all his bees.

Dr. Pumurpes. I do not know about Texas, but I do know that
farther west an eradication of 50 colonies to many of the bee keepers
of the West is not a serious proposition. The western bee keeper's
normal increase is more than his loss, so it is not like the loss to a
small bee keeper.

Mr. Arcuiey. In Texas we hardly ever find an apiary in which
every colony has the disease; therefore, when we burn the affected
colonies we have enough left to rebuild the apiary.

Mr. Ranxrn. In considering this matter of bee disease and bee
inspection one sometimes wonders if the ideal inspector exists. It
would seem that bee inspectors are born, not made. The fact that a
man knows bee disease and its treatment does not indicate that he is
necessarily a good inspector. The most successful inspectors of
whom I know are men who not only know bee disease thoroughly,
but also have the ability to handle the bee keeper whose bees they
are inspecting. The successful bee inspector, then, must first of all
be able to diagnose the disease and know it under all its varying
conditions. Next, he must know its treatment and management
under every condition which may arise; he must know every con-
dition on which the success of the treatment depends. Then, last.
but also of vital importance, he must be able to use tact in the hand-
ling of the men whom he is appointed to help. He must know from
the appearance of a man and from the first words exchanged just how
to proceed with that particular individual to secure the best results.

Let me add a word in defense of the inspectors. I know 14 of
those in California personally, and among them are some very ex-
ceptional men. They are not all equally successful, although I believe
they all know bee diseases thoroughly, but among the entire number
I do not know a single man who is serving as bee inspector merely
for the money he receives for the work. Let us give credit to whom
credit is due. These men are doing good work, and it is through
these men that the bee keepers must look for the suppression of bee
diseases under the present system. Give them your support and en-
couragement, but never under any consideration criticize them in
public in a way which would interfere with the work on bee disease.
The laws provide for the removal of an incompetent man, and if a
man is not competent to serve as an inspector let him be removed and

72 MEETING OF INSPECTORS OF APIARIES.

4 man put in his place who is competent, but under no circumstances
subject an inspector to the criticism of the bee keepers of the com-
munity or of the bee-keeping press. This is unwise, for it gives the
public a prejudice against inspection rather than against the indi-
vidual inspector, while those few deserving of censure are perhaps
una ffected.

BOILING HONEY FROM DISEASED COLONIES.

Mr. Morn. Mr. France has said that you can not kill the germs in
honey until you boil’and boil until the life is all out.

Mr. Ranxin. All you have to do is to make a hot fire and the
honey will boil. Of course you have got to boil it sufficiently long
to kill the germs.

Mr. Murn. How large is the tank reservoir?

Mr. Ranxin. Big enough to hold your combs; as Abraham Lincoln —
said of your legs, they must be long enough to reach the ground. The
tank used by one bee keeper is 6 feet square and 4 feet high, and you
would be surprised to see the amount it will take care of.

Mr. Tues (Wis.). Are the frames destroyed then 4

Mr. Rannrin. Yes; we never use any secondhand frames.

Mr. J. A. Rouse (Mo.). I would like to ask if that water does not
get too thick ¢

Mr. Rankin. Not at all.

Mr. Rouse. How do you get rid of the honey? I tried that plan
and found that honey and wax hung with the frames until they did
not look like frames.

Mr. Arcuiey. Mr. Rankin’s treatment is similar to ours except that
we burn. The labor for digging ditches is very cheap. It would
only cost us $5 to get ten ditches, and in each ditch we can burn 30
or 40 colonies. Our treatment is something like your California treat-
ment, except that it is not so complicated and is less work.

Mr. Ranxrn. That is another phase of the proposition. Conditions
are different in that also. In California you can not hire a man to do
the work for less than $80 per month.

Doctor Puiniies. We have gone over the subject of treatment thor-
oughly, and I think all persons here have arrived at about the same
conclusion; that is, that it will not do for a man who has a few
colonies in one part of the United States to write to our bee journals
and tell us all what to do. We want to know what he is talking
about. The vast majority of the men who write to-day know
nothing about the varying conditions. What will work in one
little county in the East will not work in the West, and vice versa,
the methods of the West will not work in the East. Suppose that
Mr. Scholl should sit down here and tell everybody in the United
States to burn their bees. i

BEE DISEASE INSPECTION IN WISCONSIN. Uo

Treatment depends upon the locality. Locality is an important
factor, but what we have to do is to. find out in what respect the
locality is different, whether it is in climatic conditions or in the
conditions of the honey flow. Weare in just as much ignorance when
we attribute difference to “locality” as if we did not recognize
any difference. We must get down to the point where we know the
individual factors involved. I anticipate that when some of the
discussions that have been carried on this afternoon are read, they
will open the eyes of some people that think they have had some
experience with disease. We have men from the East and West who
have different conditions to contend with. That is one reason why
i have been in favor of an inspectors’ meeting. Here we get on a
common ground. Conditions from different parts of the country are
discussed in a way that you can not obtain practically in any
other way.

I have copies here of the laws relating to foul-brood inspection now
in force. Some of these are deficient and others have valuable points
which ought to be brought out. It seems that the best thing to do is
to put a copy of them in the hands of every man who is an inspector,
with a lst of questions taking up the points which are covered by
the laws, and ask each one to express an opinion concerning them.
Then all that expert testimony should be collected and put on record,
so that people interested in future changes of legislation may read
it. If there is anyone here that would suggest how this subject
should be handled, I should like to hear from him.

After some discussion, it was finally decided that the Bureau of
Entomology be asked to prepare a list of questions to be sent to all the
inspectors. (The future action in regard to this is discussed in the
preface).

Mr. France then read the following paper:

THE HISTORY OF BEE DISEASE INSPECTION IN WISCONSIN.

By N. H. FRANCE,

Inspector of Apiaries for Wisconsin.

From 1870 to 1886 bee keeping was one of the profitable agricul-
tural pursuits in Wisconsin. There was no limit to the bee pasturage
of white clover, besides miles of basswood timber and large areas of
wild flowers. Comb honey in all kinds of packages sold for from 25
to 30 cents a pound, queen bees for from $3 to $10, and full colonies
were from $10 up.

In 1886 one Wisconsin bee keeper received $10,000 in cash from his
1,400 colonies, and started the first bank of Jefferson, Wis. Another
apiary of 250 colonies yielded in 1882 29,000 pounds of honey; in

74 MEETING OF INSPECTORS OF APIARIES.

1884 30,000 pounds, and in 1886, with 240 colonies, it yielded 25,000
pounds. This bee keeper also became a partner in a bank. Another
apiary of 175 colonies in 1886 yielded 32,725 pounds of honey and in
1891 the bees were all dead with foul brood. Another apiary of 200
colonies in 1900 yielded 21,000 pounds of honey, but in 1904 all the
bees were dead. Another apiary of 50 colonies in 1897 yielded 3,500
pounds, but in 1900 all the bees were dead. Another apiary of 26
colonies in 1899 yielded 2,500 pounds of honey and in 1901 all the bees
were dead. Thus I could enumerate a pageful of similar sad results
of foul brood.

About this time the State Bee Keepers’ Association voted to delegate
to the president of the association the securing of proper laws for the
eradication of this disease. With little help from the bee keepers, he
had to see his efforts turned down. Two years later I was delegated
as before, but without the personal help of our members the bill was
ridiculed and lost. While before the legislative committee I learned
better what must be done, and two years later a committee of all the
officers of the State Bee Keepers’ Association was delegated to act,
with the promise that each State member would do his part. The
committee got figures of facts about Wisconsin bee keeping and fur-
nished each association member with copies of the same, with the
request that each one see personally the member of the legislature
from his district. Many members did as requested and our entire
committee appeared before the first legislative committee and made
good progress. When the last State committee on State appropria-
tions was to consider our bill I was alone. Several other bills call-
ing for aid were turned down before I had a hearing. I gave the
committee these facts to consider: (1) There are 10,535 farms in
Wisconsin, having 106,090 colonies of bees, which produce in one year
2,677,100 pounds of honey. (2) There are more than twice as many
pounds of honey produced each year in Wisconsin as there are head
of cattle or sheep. (3) One year’s honey crop in Wisconsin would
load 13 freight cars, or if placed all in full-weight pound section
boxes, touching each other, a sweet honey walk 44 inches wide would
reach 1814 miles—more than the distance across the State. (4) The
valuation of Wisconsin bees and products amounts each year to more
than the appropriation made by the State for several State institu-
tions. (5) The State Horticultural Society receives over ten times
more aid from the State than the bees do, yet over three times as
many pounds of honey as bushels of apples are produced. (6) Over
10,500 Wisconsin taxpayers and voters who send representatives to
the legislature are bee keepers and ask to be reasonably protected by
law to save the bees.

After I was excused from the committee room the committee voted
unanimously to recommend the bill for passage. It soon became a

BEE DISEASE INSPECTION IN WISCONSIN. 75

law, and an inspector was appointed in the person of the writer.
Owing to false statements in the papers regarding the new office
created, I met with all kinds of difficulties, such as being met at the
gate with a shotgun and bull dog. At other times I was ordered
from the premises with a pitchfork raised over my head, but each
time I quietly explained why I was there, what I intended to do, and
read the law, “or refuse to allow the inspector of apiaries to inspect
such apiary, honey, or apphances shall be fined not less than $50 nor
more than $100, or be imprisoned in the county jail not less than one
month nor more than two months.” Before going away I saw the
apiary cleaned up in proper shape, the owner well pleased, and was
requested, whenever in that part of the State, to call and see them.
Now, when I am called to inspect or treat an apiary the bee keeper
is glad to leave his other work and meet me at the train, take me to
the desired place, help me, and even take his team to aid inspecting
the neighborhood. Everyone who has once been through the treat-
ing process will never need State aid again, but will take care of his
own bees in the future.

Several times the disease has been almost stamped out of Wisconsin,
when newly imported cases have appeared, and before owners know
what the trouble is several apiaries are affected. I hope that soon
every State will have laws on diseases of bees, and that no one ean sell
or ship bees without a health certificate similar to that required for
farm stock in Wisconsin.

END EX.

Page

mexanger treatment tor bee diseases 202 ¢22-. fi S23 e ee es cee lee ace 64-65
manages: poninis, not found in ““piekle brood’? 2.222022. 5s lo 5.-- oe. 17, 50
Earnest ireeee Chl Viliny eget ae Meee e es ame On Re NS a ee 14
GETS (ITAL OUI 0) Gone ares mR Sect A Sy) cn at an ap Pa a ea le 11-12
[MOS ASTED YA TSG big LMR a a Re a MN Me a ae 12-14
MEE IVCERGUECAREH ty ear ag Se, TAMER Lo Rais teres oe cn heen Se ae 14-18
HN POURG eee ey a ee eee acl eee e te Nee ese eka, eee Me fisnais 10-11
THRO AMOROMeMNG een te ee ree men ce Seek ed We tess ES 14
Gigtientey Mees ae ee ine ee Ses ee cee ee 16-17
PM ACH eee ee tee ee ee time eC Ae ee Are Ss 14
GINO MC ES ere Cat ee ete oe ree Crees acu ae tee iG
RESUS RCUVE men ee neers. Sek OER Pe SN Saw ed Wo 14-16
otererin amy: OL Nee diseased. s+ 25u0 sere s tsa ot ieee oe co dono ee ts eS oe 10-18
Bacillus A (B. mesentericus?), description by White quoted.................- 49
on healthy adult honeybees and comb ..-..----- 16
perhaps mistaken for B. alvei by Harrison - ----- 48
Mackenzie ....- 35
alvei, can not be identified by microscope alone................----- 12
considered by Lambotte a variety of B. mesentericus vulgatus. - - 53
Cultinmlechanactens eae serene ee oe eae meee ny de oe eee 13, 14, 15
descriprensby-Cheyme quoted... 22022 Seo eee ee 28-35
cistribupion im iotected: hives. <7 oa Ae eh as a ity
presenti Waropean-toul proon 22-222. ee Soe ae alt ee 16
So-colledesiblackgpnoodhy ar se aha ecods On co eet penne 15, 50
B, in pollen and intestine of healthy honeybees .._.......-.-....--. 16
brandenburgiensis, supposed occurrence in ‘‘foul brood”’...........-- 58
cholerx suis, in intestine of healthy honeybee.............-.....---- 16
cloacx, in. intestine of healthy honeybee .--.-.--=-.-...-...=.. -.-..-.- 16
COURCOMUNUMMIS NCUA CH AnaClenG soe aes me ene ee ee 13, 14
in intestine of healthy honeybee. -¢e-..-........-..---- 16
SUING cee Ao ee eR eee es ee ee ce ee es ee 37
Be aintestine ot healthy homey bee’. 2/22. st od 2a. See sce ce ek 17
Lopes Cultura kenara Ghent sam ae ae meee Seite ek ene ee Se ek 13, 14
deschipmon«by. Winitee UOred eae fo eee ook tk eects 52
presenineAmentcan foul proodesas ane se ae seen IoOnoe,

mesentericus ? (See Bacillus A.)
AUCSCMLLCTUGUSIUULLG OLUUS pe ates is Xa are rare nee ee ee a ors aie a ee 18
considered by Lambotte cause of ‘‘foul brood”’ _- 53
milii, no such organism found in so-called ‘‘ black brood”’.........-- 15, 50
. subgastricus, in intestine of healthy honeybee ................------ 17
ULNDIREREL Sos cles tae otc BOG age PRE REE pcs aaa Re ge a 50
AS (Cs. wre) vil ea MOrieaT soul WroOdase = ooo 2. 22 Sena oe oan t obec 16, 52
Bacterium cyaneus, in combs of healthy honeybees................---------- 16
iin intestine On. healt mywNoney DOG. 25s .-0 6 .< 2-46 -edse= cas peak 16
HON ERIS, Soe eles Bs he PFN SE SSE RE NE ye Pe 33
mycoides, in intestine of healthy honeybee.....-.--.....-.-------- ily
ee aCieeaMeRe OM ChOTIOLOR ee > see nae See co he oe OE ee ee 10-18
effect of climate on virulence .....----- ry Re eo ante ale ee 54, 67
PRaleniseOL IME anmMel basen 2 eee aks. 5 Oasis. Scenic ae Sen eee 70-72
TONSA CNM: Sed eset eS Susy gm ae et earn aries ge 20275 is
publicationsoft Bureau of Entomology. -.-.......=-/..:-.252.2- 22
IGE NTO EM YS S'S eociao bs BES Soe Be Se oS Sie eee eee TNR ees oa 61-73
sees, healthy,adult, bacteria found’ externally ...-.-...-...2-20. 0.002222 2ele- 16

78 INDEX.
Page
Bertrand, Edward, regarding determination of Bacillus alvei by Cowan and
louac\:(:) ne eran We MME Ge a rain Slop mie Vay! Ty oe 51
> Blackibrood,”” -bacitusimnilin mo tale wile eee ogee a ee ee 15
= Huropeanstoull brood ee eee. ete ae ee ilig/
no. such Ciseaser Soo Se as REA pe oa een ee ee 21
Boiling :honey from diseasedreolontes™-- 2-02-22 Ronee eee 2 ke A See ee 72
treatment for bee diseases <. 2o50- 8215 a 3 ku ee a eee 60-61
Burning, ‘for destroyimgsbec diseases) fe =- aes =. a a eee 57, 58, 60
dna eas =F re SE een em sre ee 67-69
Carbolictacid Pacatins ties 1G ley Tc Clie eer eens ce ease a ee 70
Cheshire, Frank R., description of foul brood quoted ....................--. 26-27
Climate, effect/on virulence‘ of, bee diseases 2. = 22.02. eas = ee eee 54, 67
Combs,-of healthy apiary; bacteria.founmd: 2225.22. ar ee ane tee 16
Cowan, Thomas William, attempted determination of Bacillus alvei with micro-
SCOPE AONE Meee eee. ea oe Bn ets ae st ea ed ern es ale ae, de 50-51
Cryptococcus alvearis, formerly considered cause of ‘‘foul brood’”’.........-..-- 24-25
FOP TLONLUT Ue ee oa ae EES ins Meco tL a ee) ie arene 24
Drug treatment for bee diseases. (See Medication. )
Eucalyptus oil, against ““pickle-brood? Seiya ane a Eee ee 70
Formaldehyde. (See Formalin. )
Formalin; use against bee diseases. i wo teat ta miata ee 58, 62
Foul brood, Amenican, application’ of teria: a4 5.25 aise eee ee yao 16
Bacillisalanp eo resem eee eee serene ens ye 17
can it be detected before cells are broken? .........-- 66-67
description ‘quoted from Phillips 252.2. =... 224 ee eee 18-19
geographical distribution: 22 <2 nse ene ee eee 53-54
am Callin fo rrine, See Se Sos a eS ered sn een ee 57-61
OnWeaciic:COdstis.2 Ss ee eee Sere Se er ee 57-59
vitality Of SPORES 2s ante an cae ne ee eee a 22
are both diseases ever found in same colony ?............------- 53
companson of American‘and Huropeansc2.522- 955-245 eeesease 20
European, application: of terms 2-555 han oe Sa ee 16
Bacillus alven present *.- ge ee See eee ene ie
description.of Phillips, quotede:eaen 25224 e ae eee 19
geographical distributionrs:_ 25 a5. sss eee eee 19-20
im New-York Staten.2 51.822 S655 teeta ae eine ae ee 55-56
spread through anfectedshoney- oS <4 42". gone eee eee 20, 56
symptoms of “two diseases confused bya Cheshine = ese eee 26-27
France, N. E., paper, “The History of Bee Disease Inspection in Wisconsin”. 73-75
Geographical distribution, ‘of American foul¥prood 32 o-cen (ena eee 53-54
European toul brood. 2 3-0. sce eee eee eee 19-20
Harrison, F. C., description of ‘‘ Bacillus aivei’”’ (Bacillus A?) quoted ......-- 43-48
historical résumé of investigation of bee diseases quoted .... 23-26
Honey, irom-bealthy hive; usually sterile: 2252) 22-2 ee ee 16
ANE CUION, |S) adios Soe 62S Se be ee See ee Oe eae 20, 56
Howard, .W. (Re; work ‘on bee diseasesi. 232525. 240 22 oso ene, ae eee 15, 50
Tchnewmon apium mellificarium, formerly supposed cause of infectious bee disease. 24
infection, in ovaries Of queen. bees 220. <- 422 -e5 ee ay a 57-58
is it, carried to tlowers? bac chctee oe Be ee eee ee ee 21
of larvee ob honey bees-: wo. 3: Soe a ee ak wake ee ee ee ee 20-21
Inspection: of apiaries, jn vNew: York States 552. game esse eee eee ee 55-56
IWAISCOMSII Hg 22 set eys hk LEG re ae eg oe Pepe deere 73-75
On Paciic Coast | 22 sso i aes eee eee ieee tee ee 57-59
inspectors of apiaries, appointments, ¥.-f22- oes cece ee ee er 59-60, 61-62
OPSANI ZATION Me es ee ee a eer ae ee ee 10, 54
Intestine of healthy honeybee, occurrence of bacteria. ........-...---------- 16-17
fungiand yeactd = 225s. eee. 16-17
iarvee of healthy ‘honeybee, usually steriles.—°2_- ss eee ee aan oe 16
honey bees; infection 270 4 Joe Oa ae ce eee ete ate ee 20
Mackenzie, J. J., paper, ‘‘The Foul Brood Bacillus ( B. alvei); its Vitality and
Development;’’ quoted). hes Fi a Se ee le see ee 36-42
‘‘McEvoy”’ treatment for foul-brood diseases ..-..........----------- 40-41, 61-62
Media tor bacteriological: work \¢.62- 25 Jk seen ee ee 12-13
Medication for hée. diseases... 4. :us 0p) ch ue ie eee ee 56, 58, 69
Micrococcus C, in: combs, of healthy honey bees as eee eee ee ee 16

Moore, V. A., investigation of brood diséases'of bees!. 4-0 seen e ees a ee 51-52

INDEX. 79

Page.
Mea piiiolepeta mMOr Wee CIseases =. ..5 os ~ obser desc tibet etc occcee ec esos 69
IMBUREAbINEMtOMDeCeCISCASeSs 2-2 Nae aoe se Sooo sce asa tosses see 58
‘New York Bee Disease.’’ (See ‘‘ Black brood.”’ )
iechoL mectine Of imspectors Of aplaries: < -. = 2-24.22 5-- 25 esses et ees 9
rea ZAtONcOLMMSpeClOLs. CL apIalIGs. <b 2s -oce.c2 Je cecs ese ekkis ssc eec cece 10
Palsy. (See Paralysis.)
Panhistophyton ovatum, fungus formerly supposed cause of bee disease. ------- 24
Bandincisy nm osbactenialicauseknoOwMis-- 0.0 9- else ee ea! Se Sc earn ao 16
Parker, F. A., paper, ‘‘American Foul Brood on the Pacific Coast’’.......... 57-59
Phillips, EK. F., descriptions of American and European foul broods quoted... 18-19
paper, ‘‘The Present Status of the Investigation of Bee Dis-
DOV TSE 2 een A re a ea eh ee ne ee a I 22-53
Pabiekiewbroods, a no,pacterial cause: KNOW. s-scee Sa. ne os scat oe ole Se 16
ELCAUITVG IN oper ae eee pale ee eente ais ears A at eyes oe 70
Pseudomonas fluorescens liquefaciens, in intestine of healthy honeybee--.-.----- iy
Publications of Bureau of Entomology on bee diseases ........-.-.--.------- 22
MncenepcesOCCUrKEMCELOL DACLEMI Ai see eee te eA Se matte cs ioe one scenes Dil
Rankin, J. M., remarks on bee-disease work in California.....-...-.......-- 60-61
Requeening, as remedy for bee diseases............---.----------- 56, 57-58, 63-65
Root, E. R., regarding Cowan’s determination of Bacillus alvei in Ohio... ..-- 50
Saccharomyces F, in intestine of healthy honeybee ..--....-...-------------- 17
Gin Giseused, prood ol honeybees.) S2..- 2 2c ssc se 5-22 7
foscus» in. comibs or healthyshoneybees-----+.25-2+<----.-=---- 16
Schirach, originator of ‘‘McEvoy’’ treatment for bee diseases ..-........---- 62
Shakinestreatmenttoree Giseases=-..22 25-66-26 225- 525 l2522225c- 2s 56, 57, 58, 60
UI oc S the Menai eae creat eee eas A iG 67-69
Spirochxte apis, supposed occurrence in bee disease...-..........-----.------ 53
Stewart, Charles, paper, ‘‘Apiary Inspection in New York State”? .._........- 55-56
Percnnmentmol bee GIsedses aie oo SSE eh cis Aa cece eee fede asc owe seks oe c'ee 61-73
GX OCIIIOS Y See Se HESES eo eee REE ae ene mec serecte 10-72
Virulence of bee diseases, effect of climate.................---.....-------- 54, 67
Mitealiivwor spores.oibee disease bacteria. 2225.2: _- fic 2222 oy ec -ce tees cases 22
White, G. F., description of Bacillus A (B. mesentericus ?) quoted.........-.-- 49
lances quoteds= streak ee eee 52
investigation of brood diseases of bees ..........-..--.-------- 51-52
paper, The Bacteriology of Bee Diseases?” |... 2.25. 2-2-2... 2. 10-18
summaries to Parts I and II and Conclusions from Technical
Series, No. 14, Bureau of Entomology, quoted........--...- 16-18

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- U.S. DEPARTMENT OF AGRICULTURE,

BUREAU OF ENTOMOLOGY—BULLETIN No. 71.

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

[as ae
PERIODICAL CICADA.

¥ BY

‘ C. L:- MARLATT, M.S.

g Entomologist and Acting Chief in Absence of Chief.
ed

: Issurnp JuLy 18, 1907.

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hte et - WASHINGTON:
GOVERNMENT PRINTING OFFICER.
. 1907,

o tL. Manian, alone te and Acting Chief i in absen ce “
9 I) MONMMGaL SUN. Cha Che

SHrrENDEN, in charge of breeding Gpolnanie?
Hopxiys, im charge of Sorest insect investigations. act
Hunter, in charge of cotton boll weevil ‘investigations. ne 4 #
‘Wesster, in charge of cereal and. forage-plant insect ‘investigations
‘ Hehe in charge of deciduous- fruit inaget eee Va
HILLIPS, in charge of apiculture. SRL We aac eaten sis
Rocers, in charge of gipsy and brown:tail es work.
A. W. Morra, engaged in white fly ‘investigations. .

M.
1. §. G. Trrus, in charge of gipsy moth laboratory.
eB.

DD.

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Gants, engaged in silk investigations:
CURRIE, assistant in charge of balla ‘work.
“Mave Coxcorp, ate haste

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Bul. 71, Bureau of Entomology, U. S. Dept. of Agriculture. FRONTISPIECE.

C. V. Riley, del L. Sullivan pinx. A.HOEN @CO BALTIMORE

TRANSFORMATION OF PERIODICAL CICADA (TIBICEN SEPTENDECIM).

eo DEPART MEN FOF “AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 71.

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

THE

PERIODICAL CICADA.

BY

GC. L. MARLATT, M.S&.,
Entomologist and Acting Chief in Absence of Chief.

IssurEp JuLy 18, 1907.

WASHINGTON
GOVERNMENT PRINTING OFFICE.

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5

LETTER OF TRANSMITTAL.

U. S. DEPARTMENT OF AGRICULTURE,
BureEAv or ENTOMOLOGY,
Washington, D. C., April 16, 1907.

Sir: The subject of the periodical or 17-year Cicada has been
treated in two publications of this Bureau, namely, Bulletin No. 8
(old series), published in 1885, and Bulletin No. 14 (new series),
published in 1898. Both of these publications are now out of print,
and the accumulation of a large amount of new records of distribu-
tion and the increase of information on the habits of this insect call
for a new publication. Mr. C. L. Marlatt, who was the author of
Bulletin No. 14, has thoroughly revised that publication, incorpo-
rating all new records and information, and the manuscript is sub-
mitted for publication as Bulletin No. 71 of this Bureau.

In this publication the new numbering of the broods suggested by
Mr. Marlatt in Bulletin No. 18 (new series), of this Bureau, is fol-
lowed, so that now the designation of the broods indicates directly
their relationship to each other in time and distribution.

The writings on this species are voluminous, and the bibliography
published in Bulletin No. 14 has been extended to include the
important additions to the literature which have appeared since 1898.

A good deal of the matter from Bulletin No. 14 has been used with-
out change, but the brood records have been thoroughly revised and a
distribution map has been made for each of the known broods. Some
new photographs have ,been introduced to illustrate particular fea-
tures of the life history of the Cicada.

The periodical Cicada covers in its range nearly all of the United
States from the Mississippi Valley eastward, and has a very consid-
erable economic importance. The curious features of its regular
periodic appearances and its long subterranean life give it perhaps
the greatest popular interest which attaches to any insect whatever,
and lead to many inquiries with every recurrence of an important

brood.
3

4 LETTER OF TRANSMITTAL.

The present year will witness the recurrence in the Southern States
of the largest of the 13-year broods of this insect, and the prompt
publication of this Bulletin is therefore advised to meet inquiries
for information and to assist in the collection of accurate records of
this and subsequent broods.

Respectfully,
L. O. Howarp,
Entomologist and Chief of Bureau.
Hon. JAMES WIson,
Secretary of Agriculture.

CONE NES:

Summary of the habits and characteristics of the Cicada...................----
iieracesproods and: vanieiestot the: Cicadaias: 222... 2220502254522. 5 ee
Nel EVeatsrace, ANG als VearetACG. oo — . a. Sap cec ns asec eae e- 2 es
elamonon Chimate tosthe Faces: ccc. ac) bacucek les) ss seed ose fel. ees
Mihexdumarnpemodical Creatas sa... ufo u tase see ewe See ae vec weno
‘Phe broods of the periodical Cicada. ..-....-:22-2/...5---.--22. bak stetiaiel
Pero Mein cries MeO ds a ss oe ee eee ee Ie eT 8
ahevelassitication onthe: BhOOGS<*c.t.2. i622. se o-oo es 2s Lee
ae relationship ot the diferent broods:.. 22/2. 0..." 2 .e. 222.
Pae relationship of the (7-year broods...2...2.2 2.2.22. 25.2.8

The relationship of the 13-year broods...................--------
Pources'oLenrarsn the oldstedords.; ss-se252 1.222.122 222 Flee le
Brouclsoils lop OFmkGryCaruoeCrloas fos. fe fa oe ee eee ee
Brahe nO earaRCescs. fata. etna Seine SoS Comers ee ae
Pmercistrivutiom cb che periodical Cieada...:.:..-.240-1 2-0 .. 42.2 2-0-2-2252-
SOU ECesrn MIGhiea (HONS 84 jak ead ee ee os ee Sa
The general range of the species and of the two races.......--------------
The range of the well-established broods, taken in numerical order.....-- -
IBTOOdslOimune yi sy ean Cee seater sea ors sete ee St eee
Brood l—Septendectm—V91O. 3-252 22o ce eo hes eee s- ece see

Brood El——Septendecitm—l9le 2.2 see. Jose ck ss se eee

srcod: Ll —Seprendectm— 192. 28.2 oan ect nee tesla sents

Brood. FV—Sentendecim— 19s. -. 2-2 42-422e45-2- 42-22 See es

Brood, V—Seprendecvwi—19 14 os. ss oe ne ene eee ees as

Brood Vil Septendecim= Volo 22. —2 aces es ene bet ee tee

Brood Vil Sepiendectm—l916.5 oe ie ic ee

Broody VAll——Sepiendectm—il907 : 22.26.22 2-22 ote ees

Broad) LX Septendecum—l918. . 225.23 5 to. fan ne see nee ee

brood 2x-—=Septendecim—1010s s. 2. cs ee ot ue ne ees

Brood Xl —Septendeciin— 1920s... 2.22220) i 6222 bts et tla. bem

Brood: Xl —Sepiendectm-—190l sc oe le te as eg

Brood X1TV——Septendectm—1923 . 2... 220 ee
Brood xV—Septendecpm 10072 = noses. base Doce ae en ee
Brood XOVil-—Sepiendectm—1908 22... 2. 2 os ee ee

BTOOUmNOXe— seed cer — 90 Sere eee ee noe oe ee oe
STOOUMNONE——sircdeciin == see eee ee en an ae ee
Brood xexaltle—winedecrn—19ill Qmeameeeee ae ete 2 eee eae

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PAAAADP AP aaaan»nnapsp f& bP HP BP & ow W Ow OO & Ww OO
DHOMOOANENOSDUAMAROBSNSE RON DSHOKOARANA

6 CONTENTS.

The distribution of the periodical Cicada—Continued.

Thevange of the well-established broods, taken in numerical order—Cont’d.

Broods of the 13-year race—Continued.
Brood XXIII—Tredecom—1911......._.-

Brood! XOOx—iredecvin—1 9 Se eee

Systematic position: and structural details 225.5 Boe eee ee ee

AUMmveriaaKounilay Top nS, Ve leeVe eS e ae ee

We OVIPOSItOl sas ee haat eka ee ee

Nhemausteal: ap purataise- es 22 ee al ee ey cee eae kee een

The sone notes:of the periodical Cicadas. 2 402 -ce = sees ne Coe en ee

The:so-called sting \of the Cicada-/. 22-22) 22

Mransfiormation ‘to: the adult stages <= - Maen oe ee ee
Period’ of emiergente 2.) 22.7.2 So 7000 ee eer ere eee eee See
Duration, ofthe vadullit stages o2 st het nie ak Spee ce eee ne ee ET
Method of:emersences.. 2... ee toes oe. ee an ae a
Cicada huts, OLcomes. 2 sro. eke ae eee ys A ae
(heact of tra nstornmiation.. 7... 22°. Seen ols oe en Oe a eee

Oviposition and its effect on the plant.........-- SELES cra ISH LLY Beh eh act he

IPL AMIS SOUS Cero epee eet tea rea ah aR

Resaliito thesplant of oviposiiions:: 0-34.00 >) sa ee eee ae ae

Method of inserting the eggs...........-.-.------

The growth and hatching of the eggs.......-.-..-....-
The underground hfe of the Cicada..........-..-.....

Experimental proofs of the long underground life

A successiull 17-year ‘breeding record: -22- =: 220.02. ss =

History of the larval and pupal stages.......--.---

Technical description of the different stages... 2. = <2. 22 2222 Ga ee
inet damval stage. ¢ Gs i6 <': Bae: et en oe ge ee on ee

Second tarval-atage. 24.0 ES ete ae ee a eee

Third) terwaill piiees <>. 0s cee RO eee oe aan ee
Rourth larval diaper Sete oso ce ae as a neat ohn ieee ee

First pipal- Stages oe <8 - se erre cer ame
Second Soup e stele eee ae ee ee

The habits ef the Varva and ‘pupa.+-<.2-2<43-c2 ane oe wae Cee ae eee ee
The food of the larva and pupas 22 scos. 5 oe ee eee ee ee

Adanes koeannivorm. var wleves tolls 2 ee I

Phe maethod tof, burro willik es se sa e es e er eer

Damage occasioned by larvee and pupee.--------

Mhewnankuralmemenntesxos sulle 1 C1 sil eee re eee eae

Insect parasites: 5:2. 52 8. Sasa e ee eee es
Dipterous enemies, 2-222 eee
IHemipterous enemies. 5.2=.--- 22525-5528

Page

CONTENTS.

The natural enemies of the Cicada—Continued.
Insect parasites—Continued.

Paymenopterous memes. 2.2... 5285-2. -.s20-555-% aS Os pe
SAUVE SOE Wet CPUC CTAB OVEN 21s Ne geyser ee
siedanger Ger er WASPs---cateree se oe ee omens. shake nea
nie ams bes Onenne: CORRS ote ei Ree oe 2 oer ie ats Slam a Se
Mineman ai MMe seo, eI Pee OR ca en oe ae lg oe
Miscellaneous: predaceous miltés 22. .2< 2... Sos) 6 8s 25 se
Miecvetieurate GueMmueses Sou sms Nooo neat cc ent a aek sees oS eee
ithe dimmeauerdisease-ol Ge aaulisies = ose ee te one ap os whee ee Shee
Remedilecian dency cml vVekeeses-a-5-5 520 ee eR ch aoe mc coca ates owes Se
‘The ener) character of the problem. 22. 2:41.25... .-2-- 5255-22. ose ee S84
Means of destroying the emerged pupe and adults...............---------
ocho O ee Ae st ee rotate ek ah oe yo Reet est e DSSS
ID esgic ioLRe All UNSC VCNG les\e ee Boer Oucr DEneaee ace aa eee eee
Wo plleanOnE LOIPLe Vent OVEPOSIMION 25: © = 25. sss 225552 ee 22a she
PrecamilG Tamyann Gus Meda seer Meets Sena ee awk Sek ee etyreeloars = oe ee
Means against the Cicada in its underground life.............-.----.-.---
eyuenliot lean Cicada th Mabenabine=t So eee s ye i Sede a eee:
imiplarrapuy Of time wperoglery GleAdall 2 2.6255. ae ade seas - eect ewe
Appendix. Dr. Gideon B. Smith’s chronology of the periodical Cicada...... -
index oes Sees ee ea cia SEO SIRS Poe Robe Cee Rene oC eet Eis DEE SCaE re

y 7 to dy =

Ree Ts

7
*

PS EONS:

PLATES.

Page.
The transformation of the periodical Cicada ( Tibicen septendecim).....- - Frontispiece.
isenee WOK orgthe periodienl Cicada: 22.0 cso ns BR ke 12

Il. Photograph of chambers of the periodical Cicada, general view,
taken-ate New Baltimore, Ni» Y.;,/May,, 18945 222 3222 se. 2 So 92

III. Photograph of chambers of the periodical Cicada in woodshed,
WieshinigtomsDe Cet 902 tsp estes teehee es eet 2 Sle noe 94

IV. Chambers of the periodical Cicada, natural size, in woodshed,
Washimeton,. Ds Civ 1002) pre: -- oaeh sare Aten so Jail: Sb be 96

V. Empty pupal shells of the periodical Cicada clinging to leaves,
Brocdixe 1902 Washington, D. Gre siseh. Jes. S25 2s. ee Dae 98
VI. The transformation of the emerged periodical Cicada. .......-.-.- 100

TEXT FIGURES.

Fia. 1. The periodical Cicada, representing the typical form and dwarf form. . 21
2. Map showing distribution of the broods of the 13-year race.......-...-- 36
3. Map showing distribution of the broods of the 17-year race.........-.-. 37
45,Map showine distribution of, Brood J, 1910... 22. 25-52... 92228-4225. 39
5. Map showing distribution of Brood II, 1911........-. Laer. . Si aelw aes 40
GME showine distribution, of Brood LIT, 912... 2. 2 een 22 ce y scaess eee 4]
ie Mop showing distribution of Brood. TV, 1918... 2-22.22 t<ec -ncee Ss 42
8. Map showing distribution of Brood V, 1914...........-...--.-: Fee as 44
S.Map' showing distribution of Brood Vi, 1915. 52°.-.-. 2.2.5.2). .28622 5 45
102) Map) showine distribution of Brood VIL, 191G_..2. ....-.- 22-42-2246 47
iL Map showineidistribution of Brood VIIL, 1917 _....2....22 25. 22..,-45-2 48
i> Map showing distribution..of. Brood. LX, 1918. 4.222222. 2. 2. sc -s2ae-%< 49
i3Map showing distribution of Brood, xX, 1919)... 5... 5.5.22 252- sient 51
ifo Map ciowine distribution of Brood XU, 1920... .- 025-2. n2 2 54
15. Map showing distribution of Brood XII, 1921_...-. neat a hae Le eas ae te 56
lé. Map showing distribution of Brood X11L).1922. 22. ..0.22 222225 2G222 22 57
17. Map showing distribution of Brood XIV, 1923..........-......------- 59
1S Map showing distribution)of Brood XV, 1907. ..-: 2. 22-22. s.4-54-1)5:=2 61
19. Map showing distribution of Brood XVI, 1908 ..-...-..- 2. -...-+\--:- 62
20 Map showing distribution of Brood XVII, 1909. -.........:.22..2...2: 63
/ 21: Map showine distributionof Brood XVITE, 1919 =... 5..........0....<. 65
22. Map showine distribution of Brood: XTX. 1907 2... 22s)... +e 66
23. Map showing distribution of Brood XX, 1908.........-.-.....-------- 68
24. Map showing distribution of Brood X XI, 1909................-------- 69
25; Map showing distribution of Brood: XXII, 1910. ......-...<....-.------ 70
26. Map showing distribution of Brood XXIII, 1941.2)... 2... sll en cil

Fia.

. Map showing distribution of Brood XXIV, 1912
. Map showing distribution of Brood XX VI, 1914
. Map showing distribution of Brood X XIX, 1917
. Map showing distribution of Brood XXX, 1918
. Head and prothorax of Cicada, lateral view
. Head of Cicada, front view, right mandible and maxilla drawn out--
. Head and prothorax of Cicada, lateral view, with parts separated to

4. The periodical Cicada, side view, showing beak and ovipositor
. Abdomen of female, showing ovipositor and attachments ......-.----
b. Tip-of evipositor, much enlareedie 25.28.2522 ceee See ae ee ee
. Cross section of ovipositor
. The musical apparatus of the periodical Cicada
9. Pupal galleries ef tthe:Cieaday.\ 25) 22k ie ed es es
0: Sit holies ini ‘poids. 22 Saou ces Soo SS ened shee eee Sed Pee ee ee Ente
. Twigs showing egg punctures and illustrating manner of breaking

42.
. Cicada scars in hard-maple twigs after seventeen years........----..---
. The egg nest of the Cicada, showing nature of wound and arrangement

ILLUSTRATIONS.

SIO We BELT CULM fk cosa ec eee tee ee

Twig showing scars from punctures after the second year....-....-.----

OPERAS GE: Sele HS, SES =e. eS Mee ee nel hh oe eR Rete

5. Egg, much enlarged, showing young about to be disclosed. ......-...-
. Newly hatched larva, greatly enlarged...........-..-.-----+-------
. First larval stage, illustrating the larva at the beginning and end of

Ghai S/Staeet son See hes ieee Soe Se ee ee eed: Beene

. Second larval stage, illustrating the structure of the anterior leg. -......
. Third larval stage, illustrating the structure of the anterior leg... ..- .
. Fourth larval stage, illustrating larva and structure of anterior leg. -- -
51. First pupal stage, illustrating the structure of the anterior leg.....-..-.
2. Cecidomyiid ‘ege parasite of the Cicada =... 2. 2..2- oa see ene eee
3. Wipe parasite, Lathromems ercadac 2. .s 2 is. 2 222 ah eae oes
. Female Sphecius (digger wasp) carrying a Cicada to her burrow....-.
. Diagram of the burrows of the digger wasp........-.-----.----------
. Cicada tibicen with wasp egg attached to thorax..........--....--.--

. Full-grown larva of the digger wasp in its burrow feeding on a Cicada. . .
. Larva of digger wasp with anatomical details: pupa of same, front and

Tatteralivilewse. Dee oe Pee cme nee tne eee re he eer eee eee

. Digger wasp larva constructing its cocoon. -°>--- oo: - oe eee
. Cocoon of digger wasp, with enlarged section of breathing pore... -.-.
| Mite-ege parasite, Oribatella spsut 2-2. oe ee es ee ee
~Mite:ege parasite; Oripoda elongata. =) 322 ec ea ee ee
3: Mite ege parasite, Oppia pilosa: 32 --o 2 ae ee ce ae A ee
. Mite egg parasite, Pediculoides ventricosus. 2... <2 -- 5 22-- 2 ee
65.
66.
; Mite ege: parasite, Cheyletus spo 222e. See. s fee mtg ee ea ere
68.

Mite ege parasite, Tyroglyphus cocciphilus ....-.-.-..----------------
Mite ‘ego. parasite, Ponts Ovalis ic Stas ee ee eae ee ee ee

Mite ege parasites Bdella spe i252 eee sta ane oes ee eee

~I1

as
co 00

co CO CO 6
He eH Oo

82
83
92
100
106
106
108

109
i
112

118
119
120
120
PAL
129
130
130
ial
131
132

132
138
134
134
134
135
136
136
137
137
138

THE PERIODICAL CICADA.

SUMMARY OF THE HABITS AND CHARACTERISTICS OF THE
CICADA.

The periodical Cicada, often erroneously called the ‘17-year
locust,” or merely the ‘“‘locust’’—a term which should apply only
to grasshoppers *—is, in the curious features of its life history,
undoubtedly the most anomalous and interesting of all the insects
peculiar to the American Continent. This Cicada is especially
remarkable in its adolescent period, the features of particular diver-
gence from other insects being its long subterranean life of 13 or 17
years, during all of which time its existence is unsuspected and
unindicated by any superficial sign, and the perfect regularity with
which at the end of these periods every generation, though numbering
millions of individuals, attains maturity at almost the same moment.
To the naturalist, familiar in a general way with the peculiar habits
of this Cicada, its regular periodic recurrence always arouses the
keenest interest on account of the anomalous life problems presented.
To those unfamiliar with its habits, these sudden recurrences not
only startle but often excite the gravest fears for the safety of trees
and shrubs or even of annual plants.

In view of the damage often occasioned by unusual insect out-
breaks, such fears are not unreasonable, when, without warning, this
Cicada suddenly emerges over greater or smaller areas, filling the
ground from which it issues with innumerable exit holes, swarming
over trees and shrubs, and making the air vibrate with its shrill,
discordant notes. During its short aerial life it leaves very decided
marks of its presence in the egg slits which thickly fill all the smaller
twigs and branches, the killing or injury of which causes some tem-
porary harm and a sort of general twig pruning not especially inju-
rious to forest trees, but more so to fruit trees, and very undesirable

and appeared in the earliest published notice of the Cicada (1666), and the name locust
has ever since remained the popular designation of this insect. The sudden appear-
ance of the Cicada in vast numbers very naturally recalled to the first observers the
hordes of migratory locusts or grasshoppers of the Old World, as Say and Fitch early

pointed out.
Li

12 THE PERIODICAL CICADA.

Following briefly the history of the insect, the young ant-like larva,
hatching from the egg a few weeks after the latter has been laid,
escapes from the wounded limb, falls lightly to the ground, and
quickly burrows out of sight, forming for itself a little subterranean
chamber or cell over some rootlet, where it remains through winter
and summer, buried from light, air, and sun and protected in a manner
from cold and frost. It lives in absolute solitude, separated from its
fellows, in its moist earthen chamber, rarely changing its position
Save as some accident to the nourishing rootlet may necessitate its
seeking another. In this manner it passes the seventeen or thirteen
years of its hypogeal existence in a dark cell in slow growth and
preparation for a few weeks only of the society of its fellows and the
enjoyment of the warmth and brightness of the sun and the fragrant
air of early summer. During this brief period of aerial life it attends
actively to the needs of continuing its species, is sluggish in move-
ment, rarely taking wing, and seldom takes food. For four or five
weeks the male sings his song of love and courtship, and the female
busies herself for a little longer period, perhaps, with the placing of
the eggs which are to produce the subsequent generation thirteen or
seventeen years later. At the close of its short adult existence the
Cicada falls to the ground again, perhaps within a few feet of the
point from which it issued, to be there dismembered and scattered
about, carpeting the surface of the ground with its wings and the
fragments of its body. Such in brief is the life round of this
anomalous insect.

So far as is known, other cicadas appear every year, usually in
comparatively small numbers, and this yearly recurrence has led to
the belief that the larval existence of these species is much shorter,
if not limited to a single year. In the absence of direct experimental
proof, however, it may be true that all cicadas have a long larval
existence, and the absence of well-marked broods in other species
or the complete breaking up or scattering of these broods, so that
individuals emerge practically every year, have erroneously been
taken to indicate a much shorter term of underground life.¢

If we can not satisfactorily explain the reason for the long larval
life of the periodical Cicada or the conditions which led to the origin:

a The writer recalls that in the summer of 1885 a very large species of Cicada (C.
marginata Say) appeared in considerable numbers among the scrubby white oaks bor-
dering a stream near Manhattan, Kans., and filled the air with its very loud and dis-
cordant vibrations; yet, although familiar with and a frequent visitor of these woods
in earlier and later years, no other experience with this particular species was had.
It may be, therefore, that this species, which is more than twice the size of the period-
ical Cicada, has an even longer life period.

There are other western or Rocky Mountain species which give evidence of
paralleling very closely in periodicity and number the eastern periodical Cicada.
(See p. 36.)

PLaTeE |.

Bul. 71, Bureau of Entomology, U. S. Dept. of Agriculture,

a, Fresh wounds in maple
showing that the pune

from old trees; g, h

U

WORK OF THE PERIODICAL CICADA.

+ b, d, e, and f, Condition four months later in maple; ¢, Wounds healed in one season; ¢, Three months’ old scars in wild cherry,

Apple; j, k, l, Pear

‘tures may extend both toward the top and base of the twig; g, h, 7, j, &, J, Cicada sears seventeen years old on terminal branches

(After Hopkins. )

SUMMARY OF HABITS AND CHARACTERISTICS. 13

of this peculiarity, assuming it to be abnormal, we can at least see
certain advantages coming to the species therefrom. Among these
are the protection from attacks of parasitic enemies, since we can
hardly conceive of a parasite limited to this Cicada which could pos-
sibly extend its existence over an equal term of years. Its occur-
rence, also, in overwhelming numbers at almost the same moment
everywhere within the range of the brood prevents its being very
often seriously checked in its adult stage by the attacks of birds
and other vertebrate enemies, which fatten on it in enormous num-
bers. For this species this is a most important consideration, for it is
naturally sluggish and helpless and seems to lack almost completely
the instinct of fear common to most other insects, and this leaves it
an easy prey to insectivorous animals. The almost entire absence of
fear and consequent effort to save itself from danger by flight or
concealment is apparently a consequence of the long intervals between
its aerial appearances.

The greatest check on the species has been in the advent of Euro-
peans on this continent and the accompanying clearing of woodlands
and increase of settlement. The vast areas in the more densely popu-
lated East, which were once thickly inhabited by one or the other of
the broods of the periodical Cicada, are rapidly losing this character-
istic, and the Cicada will doubtless appear in fewer and fewer numbers
in all settled districts. A recent important factor which is assisting
in this particular is the English sparrow, and it has been shown by
Professor Riley and later observers that in and about cities nearly all
of the few cicadas which still emerge under these more or less unfavor-
able conditions are devoured by this voracious bird. On the other
hand, as stated (p. 58), the first brood of these insects to be noted
by the early New England colonists, namely, the swarm recorded for
Plymouth for 1634, was just as abundant in 1906, the year when it
last recurred, as ever. This is, however, not the normal condition,
the wooded areas having been considerably maintained in Plymouth
and Barnstable counties, whereas ordinarily such wooded areas have
been greatly reduced or obliterated, and the Cicada in consequence
slowly exterminated.

The rapid disappearance of the Cicada, as a result of the clearing of
forest areas and the conditions which accompany settlement, is nota-
bly shown in the case of Brood XI, which formerly occupied a compact
territory in the valley of the Connecticut River in the States of Massa-
chusetts and Connecticut. In a letter to the writer, Mr. George
Dimmock, who has made a special study of this brood in the northern
part of the town of Suffield, Conn., says: ‘‘ When I saw them in 1869
the cicadas were so abundant that small bushes and undergrowth in
the rather sparse woods in which they occurred were weighted down
with them.” In 1886 he was unable to visit the region, but was

14 THE PERIODICAL CICADA,

informed that very few of the insects appeared that year. In
explanation of this he writes: ‘‘ The woodland in the vicinity has
been steadily reduced and the cicadas, of which there are records
evoing back about a century, seem to be dying out. The owner of the
land where the cicadas appeared (a man born in 1815, died in 1892)
informed me that the rate of reduction was so rapid that he doubted
if any of them would appear in 1903.”

To the lover of nature there is something regrettable in this slow
extermination of an insect which presents, as does the periodical
Cicada, so much that is interesting and anomalous in its habits and
life history. During the long periods of past time the species has
recurred with absolute regularity except as influenced by notable
changes in the natural topographical conditions and the despoliation
of forests which has followed the path of settlement by the white man.
It is interesting, therefore, in thought to trace the history of this
species backward, taking, as time measures, its periodic recurrences,
until in retrospect it is possible to fancy its shrill notes jarring on the
ears of the early colonists or listened to in the woodlands bordering
the ocean by the still earlier discoverers and explorers. Still more
remotely one can picture its song causing wonderment to the savage
Indians who attributed to it baleful influences, and yet, less dainty
than their white followers, used the soft, newly emerged cicadas as
food; or further back in time, when it had only wild animals as
auditors. With these long-time measures our brief periods of days,
weeks, months, and years seem trivial enough.

THE RACES, BROODS, AND VARIETIES OF THE CICADA.

Much obscurity must always attach to the past history of this insect
and the origin of its peculiar habits, and notably the causes and con-
ditions which have led to the establishment of the long underground
existence and the equally extraordinary regularity in time of emer-
gence at the end of this period. Explanations may, however, be sug-
gested for some of its peculiarities as presented in its life at the present
time—as, for example, the origin of the two distinct races, one with a
17-year period and the other with a 13-year period, with both of
which a small variety occurs, and the existence of a multitude of dis-
tinct broods occupying the same or different territory and appearing
in different years but with absolute regularity of periods.

A SEVENTEEN-YEAR RACE AND A THIRTEEN-YEAR RACE.

One of the greatest difficulties in solving the problem of the broods
of this insect and their geographical limits was removed by the dis-
covery of the existence of two distinct races—namely, one requiring
seventeen years for its development and limited geographically, in a

A SEVENTEEN-YEAR RACE AND A THIRTEEN-YEAR RACE. 15

general way, to the northern half of the range of the species, and
the other requiring but thirteen years for its development and cover-
ing the southern half of the range of the species.

This interesting and very important fact was first discovered, it
seems, by Dr. D. L. Phares, then of Woodville, Miss., who announced
the 13-year period for the southern broods in a local paper—the
Woodville (Miss.) Republican, May 17, 1845. As this paper had only
a local circulation the significance of this discovery was lost sight of
and probably never came to the attention of naturalists: and it was
not until 1868, when Dr. B. D. Walsh and Prof. C. V. Riley arrived at
the same conclusion and published, in a joint article in the Ameri-
can Entomologist,¢ a mass of accumulated observations bearing
thereon, that the 13-year period for the southern broods came to be
generally accepted.

In Professor Riley’s first report on the insects of Missouri, pub-
lished the following year (1869), the joint article just referred to was
reproduced substantially without change, except for a revision of the
classification of the broods, based on data obtained chiefly from a very
valuable unpublished monograph entitled ‘‘ The American locust,”
ete., by Dr. Gideon B. Smith, of Baltimore, Md.

This manuscript paper, on the authority of Professor Riley, was
communicated to him by Dr. J. G. Morris, of Baltimore, some four
months after the publication of the existence of the 13-year race by
Walsh and Riley, but in time for use in the preparation of the article
for the First Missouri Report. In it the existence of the 13-year
Southern race, occurring in several broods, is fully recorded by Doc-
tor Smith in connection with the use of the specific name ‘‘tredecim.”
(See Appendix.)

After the existence of the 13-year Southern race was again brought
into prominence by Walsh and Riley, Doctor Phares published an
article in the Southern Field and Factory, Jackson, Miss., April, 1873,
in which he called attention to his earlier publication, cited above,
where he seems to have controverted the belief that there is no 13-
year brood, evidently entertained up to that time by Doctor Smith,
with whom Doctor Phares was in correspondence, and also to an
article published May 5, 1858, in the Republican, where he used the
title ‘Cicada tredecim.”” Doctor Smith later evidently accepted the
conclusions of Doctor Phares and introduced them in his last. revi-
sion of his manuscript memoir, which Professer Riley saw and used.
To Doctor Phares, therefore, belongs the honor of having made the
discovery of the 13-year period for the Southern broods. Neverthe-
less, but for the independent work of Walsh and Riley, the knowl-
edge of the 13-year broods might have been long lacking, and, in the

@Vol. I, pp. 63-72, December, 1868,

16 THE PERIODICAL CICADA.

nonpublication of Doctor Smith’s monograph,” these broods would
have failed ot the abundant proof on which they now rest. The race
name of tredecim for the 13-year broods was suggested by Walsh and
Riley without knowledge of its earlier use by Doctor Phares. The
latter’s early articles inthe Republican are lost altogether, the author
himself not being able to recover them in later years, and the credit
for the name tredecim for the 13-year race, following the customary
rules, should go to Walsh and Riley.

The discovery of the 13-year Southern race was of vast assistance
in clearing up the confusion which had attended the study of the
different broods of this insect and enabled Walsh and Riley to sepa-
rate some sixteen distinct broods, three of which belong to the tre-
decim race, and later enabled Professor Riley, with the aid of Doctor
Smith’s paper, to increase the number of tredecum broods to seven
and the total of the broods to twenty-two, twenty-one of which the
records of subsequent appearances have proved to be valid.

Doctor Smith’s remarks in his manuscript chapter on geographical
tribes and districts present the status of the 17-year and 13-year
races very clearly. He says:

There are two divisions or tribes, differing from each other only in the periods of
their lives; the one and much the larger division living 17 years, and the other
13; hence the impropriety of the specific name septendecim. * * * The anatomy
of the insects of both divisions is precisely the same, but septendecim does not of
course apply to the Southern division, whose lives are but 13 years. Shall we call
the latter Cicada tredecim? Why there is this difference in the periods of lives of the
two tribes we can not explain. It is not the climate that causes it, as a moment’s
reflection will prove. If that were the cause the difference would be more gradual.
For example, in northern New York they would have been, say, 17 years; in Pennsyl-
vania, 16; in Maryland and Virginia, 15; in North Carolina and Tennessee, 14, and
in South Carolina, etc., 13 years in completing their existence. But that is not the
case. The difference of years takes place abruptly on and about the line of 34°
and 35° of north latitude, on the north side of which the period is 17 years and on
the south 13 years.

While Doctor Smith is hardly justified in the last statement, it is
nevertheless true that the 17-year race is northern and the 13-year
race is southern. The territory of the two races is graphically shown
in figures 2 and 3, and is described in detail and mapped for all the
broods in a later section.

In this bulletin the two forms of the periodical Cicada have been
designated as ‘‘races,”’ adopting the position taken by Professor Riley
and the majority of the writers on this insect, rather than consider-
ing them to be distinct species, as is held by some specialists. Pro-
fessor Riley and others opposed the idea of their being specifically
distinct, not only because of their practical identity in general char-

aA summary, with extracts, of this manuscript made by Professor Riley is the
writer’s source of information on this valuable paper, which, while containing much
error and wrong inference, yet indicates careful study and accurate observation,

A SEVENTEEN-YEAR RACE AND A THIRTEEN-YEAR RACE. ii

acteristics and habits, but also on the ground of external structure,
no material difference in this respect having been noted between the
two races, although it was known that the individuals did not cross
when they appeared together. Doctor Walsh was very firmly of the
opinion, on the other hand, that they represent two distinct species,
yet in a letter to Mr. Darwin he described the 13-year race as an
incipient species, to which, for convenience, it is desirable to give a
distinctive name.? His published views on the subject, given in a
posthumous paper, are quoted below.? Reterring to the impossi-
bility of distinguishing species in certain genera by a mere compari-
son of the periect specimens, he says: .

Upon the same principle I strongly incline to believe that the 17-year form of the
periodical Cicada (C. septendecim Linn.) is a distinct species from the 13-year form
(C2 tredectn (Walsh and Riley¢) Riley), although it has been impossible for me, on
the closest examination of very numerous specimens, to detect any specific differ-
ence between these two forms. It is very true that the 13-year form is confined to the
more southerly regions of the United States, while the 17-year form is generally, but
not universally, peculiar to the Northern States; whence it has been, with some show
of plausibility, inferred that the 13-year form is nothing but the 17-year form accel-
erated in its metamorphosis by the influence of a hot southern climate. But, as these
two forms interlock and overlap each other in various localities, and as it frequently
happens that particular broods of the two forms come out in the same year, we should
certainly expect that if the forms belonged to the same species they would occasionally
intercross, whence would arise an intermediate variety having a periodic time of
14, 15, or 16 years. As this does not appear to have taken place, but, on the contrary,
there is a pretty sharp dividing line between the habits of the two forms, without
any intermediate grades of any consequence, I infer that the internal organization of
the two forms must be distinct, although externally, when placed side by side, they
are exactly alike. Otherwise, what possible reason could there be for one and the
same species to lie under ground in the larva state for nearly 17 years in one county
and in the next adjoining county to lie under ground in the larva state for scarcely 13
years? I presume that even the most bigoted believer in the old theory of species
would allow that, if it can once be proved to his satisfaction that two apparently
identical forms are always structurally distinct, whether in their external or their
internal organization, they must necessarily be distinct species.

The reasons urged by Doctor Walsh give a strong basis of proba-
bility to the theory of the specific distinctness of the two races, and
particularly the fact that where the broods overlap there seems to be
no interbreeding. Doctor Walsh’s position has been upheld by
Dr. Wm. H. Ashmead, who states that in a very careful examination

24See Index to Missouri Entomological Reports, Bul. 6, U. 8S. Ent. Comm., p. 58.

6 American Entomologist, Vol. II, p. 335.

¢Taking the ground that Doctor Phares can not be credited with the race name
‘‘tredecim”’ on account of the ephemeral character of the journal in which he employed
it, the credit should go to Walsh-Riley, since the article in the American Entomolo-
gist of December, 1868, where it was next suggested, was a joint or editorial one.
Professor Riley himself sanctions this course in the Bibliography of Economic
Entomology, Part II, p. 61, No. 474.

31117—No. 71—07

D)
_

18 THE PERIODICAL CICADA.

of the material in the National Museum he has observed small but
constant differences between the two races in the shape of the last
ventral segment of both the male and the female.

For the present purpose, however, it seems wiser to consider the
13-year broods as representing a race merely, or an incipient species,
as suggested by Walsh, because of the absolute resemblance in prac-
tically every feature of structure, coloration, and habit, in the two
forms, which exhibit the single important point of difference repre-
sented by the four years’ variation in the length of their subterranean
lives.

While in the matter of interbreeding they may be distinct, as the
records seem to prove conclusively, the two races represent one
species for all practical purposes and differ in a very striking manner
from all other species of the family Cicadidz. One race is unques-
tionably the offshoot of the other, the original differentiation being
probably caused by some variation in climatic conditions.

It is, perhaps, a hopeless task, and at best only a matter of conjec-
ture, to attempt to explain the phenomenon of what is practically the
same insect requiring in one part of the country seventeen years for
its underground development through its preliminary stages and in
another section thirteen years, in the face of the fact that while, in the
main, the two sections are, ee northern and southern, yet at
the point of juncture the broads of the two races overlap. That the
17-year period does not depend so much on the greater severity of the
northern winters is evident, protected as the insect is by the depth of
its burrows, and the natural explanation is that the longer period of
warmth in the South hastens the development of the insect, or, in other
words, that the difference in the length of the warm growing period
during which the insect can thrive and increase in size in the southern
half of its range enables it to go through its development in four years
less time than in the North, where shorter summers and consequently
shorter periods of growth occur. The chief objections to this theory,
but not necessarily controverting it, are those made by Doctors
Smith and Walsh in the quotations given. The problem is, however,
a very interesting one, and some light may be thrown upon it by
further experiments similar to those described under the head fol-
lowing.

RELATION OF CLIMATE TO THE RACES.

The anomaly presented of two distinct periods for the completion of
the adolescent stages of the periodical Cicada, exhibited by the 13-year
and 17-year races, and its apparent basis in climate led Professor Riley
to institute some careful experiments in transferring the eggs of the
13-year race, collected in various Southern States, to different locali-
ties in the North, and conversely, eggs of the 17-year race collected in
the North to localities in the South, to determine the actual influence

RELATION OF CLIMATE TO THE RACES. 19

of temperature or whether the 13-year race would maintain its normal
period in the North and the 17-year race in the South. The object of
the experiment, in other words, was to determine whether the differ-
ence in time of development between the two races is really one of
climate and temperature only or whether a fixed characteristic has
been acquired, not subject to much, if any, modification with changing
temperature conditions. That the separation was originally caused
by differences in climate in different parts of the range of the species
can not be doubted, but the fact that the two races often overlap in the
adjoining territory of their respective ranges would seem to indicate
that this time period has become in the course of ages a rather
permanent feature.

Doctor Riley’s early experiments in this direction were in 1881
with the 13-year Brood XIX, but the eggs distributed were in such
‘condition that it is doubtful if they hatched, and the effort failed.

A much more elaborate test was instituted in the summer of 1885,
in connection with the joint appearance that year of the 13-year
Brood XXIII, which returned in 1898, and the 17-year Brood X,
which returned in 1902. All possible precautions were observed not
only to collect the egg-bearing twigs at the right moment and to
distribute them in fresh, healthy condition, but to see also that they
were properly placed under suitable trees and that a record was made
in each instance of the exact locality. Furthermore, most of the
transfers were kept under observation for a time to see that the eggs
actually hatched and the larve entered the soil in their new situations.
The record of these transfers is given in detail in the report of the
Entomologist, Report of the Department of Agriculture for 1885,
pages 254-257, and was reproduced in Bulletin 14 as Appendix A.
The eggs of the 13-year brood were collected in Mississippi between
July 6 and 17, and distributed to entomologists in New York, Iowa,
Massachusetts, and Maine in eleven lots. The eggs of the 17-year
Brood X were collected in Indiana, Pennsylvania, and Michigan,
chiefly in the latter State, between July 6 and 21, and distributed in
seventeen different lots to correspondents or entomologists in Georgia,
Mississippi, Alabama, and Missouri. The preliminary report on the
condition of this material is given in the appendix cited of Bulletin 14.
The only positive record received was from Prof. Eugene A. Smith,
University of Alabama, who found in 1898 one pupal shell and
noticed several holes in the ground which answered to the description
of exit openings made by the Cicada. The pupal shell was sent to me
and proved to belong to the periodical species. That it comes from
the eggs planted in 1885 seems probable, from the fact that no brood
was due in this locality in 1898, and this would seem to indicate that
the 17-year brood may be greatly abbreviated or reduced to the
13-year term in a warmer latitude. Part of the eggs sent to Professor

20 THE PERIODICAL CICADA,

Smith came from Indiana and the rest from Michigan. Too muecn
importance, however, can not be given to this isolated experience.
Correspondence was kept up with as many of the points as could be
reached during the next four years, but no further records were
obtainable.

Nothing whatever came of the 13-year material sent to northern
localities.

The difficulty in an experiment of this kind lies in the long term
over which it extends, and the inevitable changes of local conditions
and the removal or death of observers intrusted with the experiment.
It is necessary, as demonstrated by a later test (see pp. 114-116) of
egg transfers, to have an enormous quantity of eggs to insure the insects
going through the entire term undestroyed by natural enemies or
accidents.

THE DWARF PERIODICAL CICADA.

In connection with the discussion of the 13-year and 17-year races of
the Cicada, it 1s interesting to note also that in both races the insect
occurs in two distinct types, viz, a large form and a small form,
the former comprising the bulk of the individuals of the brood and
the latter more rare and often unobserved. The existence of these
two types was commented upon as early as 1830 by Doctor Hildreth,
of Marietta, Ohio,“ and was especially remarked in the great Cicada
year 1868. The typical larger Cicada (fig. 1, A) measures on an
average 14 inches from the head to the tip of the closed wings and
expands over 3 inches. The underside of the abdomen is of a dull
orange-brown color and in the male four or five segments are of the
same color on the back. The smaller form is rarely more than two-
thirds the size of the larger, and usually lacks altogether the light
abdominal markings, although they are sometimes represented on
the edge of the segments beneath.

The small form (fig. 1, B) was described in 1851 as a distinct species,
Cicada cassinii, by Dr. J. C. Fisher.’ The contention that it repre-
sents a distinct species was urged particularly on the ground that there
exists a variation in the genitalia, but this variation has since been
shown by Professor Riley not to be constant, and specimens are to be
found in both sizes which present the same structure in these parts.

In view of the close anatomical correspondence, except in size, of
the two forms and the fact that they always occur together in the same
broods and have the same anomalous subterranean period of larval
and pupal life, the specific importance of the smaller Cicada has been
naturally open to question, and in Bulletin 14 the writer was inclined

a Silliman’s Journal, XVIII, p. 47.
b Proc. Acad. Nat. Sc. Phila., Vol. V., p. 272.

THE DWARF FORM. 21

to consider this small form as a mere variety or, more properly, a
dimorphic variety of the larger form.

On the authority of various observers certain divergences in habits
between the two forms were commented upon. It seemed to be the
general belief that the larger one appears somewhat earlier, from
eight to ten days, and correspondingly, also, the smaller form disap-
pears somewhat later in the season than the larger. The smailer
cicadas were also reported by various observers as being more or less
gregarious in habit, not always intermingling with the larger ones but
collecting in small companies in orchards or in thickets along streams
and moist places. Further, the song note of the small form was
somewhat different, but this last variation was not fully confirmed.

This small Cicada was particularly noted in the case of Brood X
at the time it was studied by
Walsh and Riley in 1868 and,
judging from the records ob-
tained of this brood of 1885
and 1902, Brood X seems to
be its particular stronghold,
although it occurs with other
broods, often very scatter-
ingly.

In 1902 the writer observed
this small form in great abun-
dance in and near the District
of Columbia, but, contrary to
the former belief, it appeared

Fig. 1.—The periodical Cicada: A, male of typical

nm large percentage during form, natural size; c, d, genital hooks enlarged; g,
the first week or ten days of singing apparatus, natural size: B, male of the small

the emergence of the Cicada, ney Netter Biles ona Hoses je ad
probably representing 50 per

cent of the specimens, and soon disappeared. Both sexes were rep-
resented, and mating and oviposition seemed to go on normally as
with the large form. The song notes of the dwarf Cicada were dis-
tinctly different from the common note of the large Cicada, namely,
a broken and chirping note, very shrill and loud. The abundance
of the small form in 1902 and the difference in its song notes were
observed by various persons throughout the range of the brood.

A careful statistical study of the variation in size and character-
istics of the large and small forms was made in connection with the
1902 appearance of this brood in Ohio by Prof. Herbert) Osborn. ¢
Professor Osborn examined and made careful measurements of some
800 specimens taken at random from various localities. The results

@QOhio Naturalist, III, pp. 323-326, December, 1902.

22 THE PERIODICAL CICADA,

of these measurements indicated a decided constancy for each variety
and for each sex of each variety in wing lengths and widths and body
lengths. The color variation was also very constant. The abdomen
of the cassinii form is normally entirely black beneath, only rare
specimens showing a narrow hind border of yellowish or orange
yellow. The cross veins also on the wing forming the W mark are
commonly less black, and the W therefore shortened. This point,
however, as in the normal form, seems subject to wider variation
than the other features.

There is a difference in genitalia, but apparently not enough to
exclude the idea of crossing, and, according to Riley, this difference
is not constant. In the mating, out of seventy pairs observed there

g,
was no instance of cassini pairing with the normal large form,
evidencing an apparently complete isolation by sexual selection.
Professor Osborn shows, therefore, that there is no ground for con-
sidering the small form as a dimorphic or seasonal stage of the large.

Professor Osborn infers that the cassiniz is a derived form, since it
appears less commonly than the other and probably has a more
restricted range, and suggests that it may be possibly a ‘‘depauperate
variety’? which has become in the course of ages fully established,
especially with Brood X, being very rare with Broods III and XIII,
which he had also studied. He concludes: ‘‘ Whether this form be
called a variety, subspecies, or species, is, it seems to me, of less
importance than a recognition of its distinctness, and the deterniina-
tion, if possible, of its phylogenetic relationship.”

The nomenclature of the species, variety, and races of the periodical
Cicada adopted by the writer is the same as that followed in Bulletin
14, namely, the Linnaean species Tibicen (Cicada) septendecim, with
the tredecim race of Walsh and Riley, and the variety cassinii of Fisher.

THE BROODS OF THE PERIODICAL CICADA.

The subject of the broods of the periodical Cicada presents a number
of interesting fields of inquiry, such as the consideration of the origin
of the broods, their chronological history and classification, and their
exact geographical limits or distribution. These topics will be taken
up somewhat in detail, with the exception of the chronological history
of the appearances during the last two hundred years and accompany-
ing voluminous historical records, which, for reasons to be later noted,
have been largely omitted.

THE ORIGIN OF THE BRoops.

It is not necessarily true, but it is a reasonable inference, that in the
early period of the existence of the periodical Cicada on this continent
it was represented by a single brood. Assuming this to have been the

THE ORIGIN OF THE BROODS. 23

ease, the Cicada would have appeared everywhere over its range in the
same year and probably at about the same time. In the long course of
ages, with the consequent important changes—geographic, climatic,
and topographic—this original brood became gradually broken up into
many broods, with constantly increasing divergence in the dates of
appearance, so that at the present time nearly every year has its
brood, or broods, each of which is limited, as a rule, to well-defined
districts, and each reappearing at the proper intervals with absolute
regularity. Of the upward of twenty broods which have been differ-
entiated, most of them have been carefully studied, chronological
records collected, and the limits of distribution fairly well determined.
For convenience of reference, these broods have been designated by
Roman numerals, as Brood VI, Brood X XVI, ete.

The origin of distinct broods in an insect possessing as long a
developing period as the one under discussion is not difficult of expla-
nation. It is a well-known phenomenon in connection with insect life
that, whatever may be the period of development of a species, certain
individuals will often, for some reason or other, such as insufficient
or unsuitable food, unfavorable temperature, or other conditions, be
delayed or retarded, while others, for reasons the converse of the last,
namely, conditions exceptionally favorable, will develop more rapidly
or will be accelerated and appear earlier. Therefore, under the former
conditions we have a longer and under the latter conditions a shorter
life period.

This is true to a slight degree at the present time of the periodical
Cicada, and especially with the larger broods has it been noticed that
scattering individuals appear the year before and others the year after
the great brood year. It is not difficult to imagine, therefore, that
under exceptional conditions some of the earlier appearing individuals
or the later ones may occur in sufficient numbers to establish a well-
marked peculiarity in this direction and form a new brood appearing a
year earlier or a year later than the original one. If in the long course
of years some accident should happen to the parent brood in that
portion of its range the derivative brood might be left to hold the
territory alone or to become the predominant swarm.

This explanation is supported also by the fact that it often happens
that the broods of two successive years occupy contiguous territory,
as, for example, the 13-year Brood XXII, which last appeared in 1897,
is distributed between Vicksburg and New Orleans, or just south of
the 13-year brood which appeared in 1898. It is reasonable to infer,
therefore, that Brood X XII is simply a strong, well-established colony
of accelerated individuals from the southern end of Brood XXIII,
with a 13-year period terminating one year earlier than that of the
parent brood. The conditions which led to the emergence of the
insect below Vicksburg in twelve years some time in the remote past

24 THE PERIODICAL CICADA.

being temporary, this portion of the old brood resumed the normal
13-year period.

Another marked instance of the same kind is shown in the relations
between Brood XI and Brood X, the former being merely an appen-
dix or a continuation in a northeasterly direction of the territory occu-
pied by the eastern branch of Brood X, which always precedes Brood
XI by one year. The interrelations of these and all the other broods
are indicated in the discussion of the distribution of the Cicada.

Local or temporary conditions which have caused a moderate
change in the time of emergence of the Cicada are on record, one
notable instance resulting from an artificial heating of the soil by hot
pipes (see p. 90). .

A similar instance of acceleration of Brood XIII, due in 1905, but
amounting to a full year, occurred in 1904 in a greenhouse at Belvidere,
Ill. The owner, Mr. B. Eldredge, writes that in 1888 he moved from
Chicago to Belvidere, and found everything covered with locusts,
and an enormous amount of damage to all kinds of shrubs and trees
wasdone. At the time he bought the place it was covered with an old
apple orchard, and the locusts worked very abundantly in these trees.
Some seven years afterwards these trees were grubbed out and the
ground covered with greenhouses, and the ground so protected had
been kept warm winter and summer ever since. Mr. Eldredge is
convinced, and he is undoubtedly right in this belief, that this con-
tinual heat and absence of frost accounts for the appearance of the
locusts in his greenhouses a year ahead of time.

He states that the locusts appeared in quantity. Before the mat-
ter was brought to the writer’s attention they had largely disappeared,
but two adult locusts were submitted and a lot of shed skins, which
fully confirmed the identification of the insect. It would be rather
interesting to know more about the local conditions to determine how
the cicadas were able to survive in soil from which the vegetation
must have been entirely removed.

An instance of a few weeks acceleration under outdoor conditions
is given by Mr. Schwarz.¢ Commenting on the shghtly earlier emer-
gence of individuals of Brood XIV near Harpers Ferry, W. Va., in
1889, in a small clearing surrounded by woods, Mr. Schwarz urges
that a clearing made in the midst of a dense forest forms a natural
hothouse, the soil receiving in such places much more warmth than
in the shady woods. That the cicadas should appear a little earlier
in such situations is not remarkable, and he suggests also that under
favorable circumstances the Cicada might develop on such cleared
places one or more years in advance of the normal time, and that
these precursors, if numerous enough, would be able to form a new
brood.

a Proc. Ent. Soc. Wash., I, p. 230.

THE CLASSIFICATION OF THE BROODS. PAS)

It is possible to conceive also of conditions which would result in
the acceleration or retardation in the development of an entire brood
or broods of the Cicada, such as variation in climatic conditions,
geological changes, or changed conditions of the topography of the
country, including the character of the vegetation.

In this or other ways, at any rate, the Cicada has become broken
up into a large number of distinct broods, often covering different
territory, but not necessarily so doing, each, however, maintaining
its regular time of appearance.

The sight but constant tendency to variation which has brought
into existence the broods now so well marked, continued indefinitely,
would so break up and scatter the present broods as to ultimately
obscure them altogether, and the overlapping of districts and the
variation in time of appearance would lead to a rather general occur-
rence every year of the periodical Cicada throughout its range, the
long period for development, however, still persisting. Anticipating
such an outcome from the intermixture and overlapping merely of
different broods, Doctor Smith (Smith MS.) rather mournfully says:
“Tn those times, if these sayings of mine should be thought of, they
will be ridiculed as a superstitious legend of the olden times.”’

THE CLASSIFICATION OF THE Broops.

In the first edition of this bulletin the numerical designation of the
broods of the two races suggested by Professor Riley was followed.
This numbering has, however, objectionable features and obscures
the relations of the broods of each race to each other. To overcome
these objections a new system of numbering was proposed by the
writer,” which has since been generally adopted. The reasons for
making this change and the numerical designations proposed are here
reproduced with little change from the publication cited.

The earlier writers, viz, Prof. Nat. Potter, Dr. William T. Harris,
and Dr. G. B. Smith, classified the broods solely according to the
years of their appearance. The unpublished register left by Doctor
Smith includes every important brood now known classified accord-
ing to race, and gives the localities for one additional brood, the
existence of which seems not to have been confirmed. Though
lacking any special designation for the broods, Doctor Smith’s classi-
fication is as complete and accurate as that published by Doctor
Riley and since followed by all later writers.’ Dr. Asa Fitch was the
first to introduce a numbering system for the different broods, enu-
merating nine altogether, but his data were very limited and he was
not aware of the thirteen-year southern period, and there necessarily
resulted no little confusion of the broods of the two races. The

@Bull. 18, n.s., Div. Ent., U. 8. Dept. Agric., pp. 52-58, 1898.
+b See Appendix.

26 THE PERIODICAL CICADA.

Walsh-Riley enumeration of 1878 gave the records for sixteen broods,
which were designated by Roman numerals from I to XVI, the enumer-
ation being based on the sequence of the different broods after 1868.
In 1869, in his First Missouri Report, Professor Riley, having in the
meantime secured the manuscript paper of Doctor Smith, added the
six broods from this paper not represented in the Walsh-Riley enu-
meration, increasing the number of the broods to XXII, and renum-
bered them again in: accordance with their sequence, beginning with
1869. ‘These broods vary enormously in their extent, some of them
being represented by scattered colonies, which perhaps have no real
relationship in point of origin, and others covering nearly uniformly
vast stretches of territory extending over several States together.
Several are rather unimportant, or lack confirmation, and one of
them, Brood III, was founded on an erroneous record and has been
dropped.

In the enumeration of the broods by Walsh-Riley, and later by
Riley, the two races are mixed together and a sequence of numbers
given which, after the first thirteen years, lost all significance as a
record of the order of the broods in time of appearance, and from
the first obscured the true kinship of the broods in each race. If, on
the other hand, each race be considered separately and its broods be
arranged in a series in accordance with their sequence in time, an
important natural relationship in point of origin and distribution is
plainly indicated.

Taking first the broods of the 17-year race, as Riley numbered them,
it will be seen from the subjoined table that if the enumeration begin
with Brood XJ, the 17-year broods follow each other in regular succes-
sion for eleven consecutive years, then after a break of one year fol-
low Broods V and VIII, and after another break of one year Brood
IX. Another break of one year precedes the next recurrence of Brood
XI, with which the series starts.

Chronological order of the Riley broods of the Cicada from 1898 to 1910.

pe 17-year | 13-year | rx 17-year | 13-year
Year race. | race. | Year. race. race.
|

ROSIE AR os ee ee pane ten FS eee SL | ese Val 1902s 1 Cee Sek eae | “OREN Tiy sees
TRO Dies Soe ed ae ne SEL LO NEWAITE 11/1 G O30 sere he nn renga i eR ES
II 2 5 aed ae ee ee Oa S| II NODAL RRs Et ah lesion 22 a | ees
SOG eso io te ops Yen a tees XIV 1\ | ROS tee. ke. ee amie Vee
Ihe tS ee eee ees eee | eee aoy Wali UN GOGSS 1-5. et Se eee On ee VII |) evil
ISOS BEN Se tans eee eee XVII VII LOOT 2,222 Shas Eel ake arte, ae ee | XVIII
S99 S Ca wae aaa ces Se XGA |e Pee 19082 2) Rees se | Ix I
1 Oe See Seer mee wate eser Ke Se Se eel 1909. Bar Rock sc ae eoe oe Ae EE Renee ae IV
LOD IME AD. Cael Baia co eee ae AIS lhe x TOL OLA: Pe Swi ae ew | a VI

Taking up the 13-year broods in the same way, it will be seen that
if the enumeration start with Brood XVI, a 13-year brood follows in
regular succession for six years. With the exception of the very
doubtful Brood X, which is separated from the last 13-year brood by

THE CLASSIFICATION OF THE BROODS. | 27

three years, there follow seven successive years in which no 13-year
broods occur.

Under the supposition that the different broods of the 17-year and
13-year races sprang in the remote past from an original brood of
each, it would naturally follow that the broods most closely related
in time would also present a closer relationship in their range, and
this, in fact, proves to be generally true.

To show this relationship and to indicate the natural order of their
occurrence, I have suggested a new enumeration of the broods in
which the two races are separated—the 17-year broods coming first,
followed, for convenience merely, by the i3-year broods. Thus
Brood XI of the 17-year race becomes Brood I, and the others are
numbered in the regular order of their occurrence, except that I
have assigned a brood number to each of the seventeen years. This
leaves Broods XIJ, XV, and XVII, as newly numbered, without
any definite colonies, so far accepted, as representatives of established
broods. As will be shown later, however, there are records which
indicate the existence of small or scatterme broods filling the three
gaps mentioned in the 17-year series.

In renumbering the broods of the 13-year race | have continued
for convenience from the end of the series of the 17-year race, the first
13-year brood becoming Brood XVIII, and I have assigned brood
numbers to each year of the 13-year period, making a total enumera-
tion of the broods of both races of XXX. As already indicated, six
of the numbers given to the 13-year race have had no brood assigned
to them, although records have been secured which seem to indicate
the existence of scattering broods filling some of the gaps, as will be
noted in the records given further on.

It does not necessarily follow, in fact it is quite unlikely, that Brood
I, as here designated, is the original or oldest brood of the 17-year
race. Undoubtedly some of the 17-year broods, perhaps half or more
of them, originated by retardation of individuals, and perhaps half
by acceleration of individuals; so that the original brood, if it still
exists, is more’ likely to be one of the intermediate ones. Brood X,
being the largest of the 17-year broods, perhaps has best claim to
this distinction.

For the same reasons an intermediate brood in the 13-year series
is doubtless the original brood of the 13-year race, and this title may
possibly belong to Brood XIX, which has the widest range of all the
broods of the 13-year race. The fewer number of broods in this race
would seem to indicate that it is of later origin than the 17-year race,
and this belief is further justified by the fact of its occupying, in the
main, a territory of later geological formation.

The following table, beginning with 1893, when the initial broods
of both the 17-year and the 13-year series appeared in conjunction,

28 THE PERIODICAL CICADA.

illustrates the new nomenclature suggested, and in parallel columns
also are given the corresponding nomenclatures proposed by Professor
Riley, by Fitch, and the year records in Doctor Smith’s register:

Nomenclature of the broods of the periodical Cicada.

Broods of the 17-year race. Broods of the 13-year race.

Year. Proposed| Riley Fitch | Smith | Proposed] Riley Fitch Smitt

enumer-| num- num- | regis- | enumer-| num- | num- saa fa
ation. | bers. bers. | ter. ation. bers. bers. | register.

0 ee eee ae eee eee I pain: © Oe ain (ee 3 aoe 1842 | XVIII SVE eeaeees 1854
sey ee Se cme oes ae Il XII 1 1843 XIX | XVIII 3 | 1842-1855

1SORo Matis oe oe Cen, TET Po] = NCTA Toon | eae eee 1844 | 1843

SOG sos 22 ae Ss OS: nese IV STV spall he Ae 1845 1844

INS ps Maia eS ae V SOY 5 | 1846 | 1845
LSOR FSA eek eee Seek VI SOVIET) 7 | 1847 | 1846-1859
SOON me ey eee em rae meas Vil DS Dae | ee ee 1848.) SXORMW eos
1900 ee 5. Satya te Seek VTE Tees |W ERERE D975 2=83 (11849 lite seas

OQ ae See pee slater IX OR eas See 1850 1849
HOO 2240. ¢ Seek yi ee ae x XXII Alle LSS lls SEARO W LAD | Fe ar Se aed |
NGO Soesaee ea Seek ene Sell I 9 1352) | EXSOV TE ee oe 2c eee eel eee
O04 2 See ae ee yao ee ROT tee ce ene, See ee 1853 ees See
O05 ee fins Jae eee XU \ 6 | 1854 Nes ean So

(Q0G=4- 2 ee ee ee sae XIV VIII 1855 1854
OO (eee cars Jase seeesiosee VE Se rete cay est erm | eee 1842-1855

19085 427205 2 Me eae xvi aU RN reese Pieris eae 1843

O09 Bare. soe caligene coc eee SREY TEL B\'- art eoee eeey| se, 2, em eee ee 1844

THe RELATIONSHIP OF THE DIFFERENT Broops.

As a rule, the relationship of the broods in point of distribution
agrees with their kinship as indicated by their sequence in time of
appearance. The relationship indicated by the latter, viz, their
sequence in time, is doubtless untrustworthy as indicating origin, in
some instances on account of the uncertainty arising from the action
of the principle of retardation on the one hand and acceleration on
the other in the forming of new broods.

In the case of a widely scattered brood, like Brood VI, it is quite
possible that certain swarms originated from a later-appearing brood
by retardation of individuals, and other swarms from an earlier
brood by acceleration in time of appearance of individuals.¢

This same condition may be true of other of the more scattered
broods, but with the broods presenting a compact range a singleness
of origin is evident.

Examination of the distribution of the broods in connection with
their sequence in time of appearance indicates, however, a certain

a Prof. W. E. Castle, Museum of Comparative Zoology, Cambridge, Mass., in a letter
to the writer July 20, 1898, suggested a plausible theory for Brood VI. The isolation
and wide distribution of this brood leads him to infer that it may be a relatively old
or “played out’’ brood, and if this be true it may be considered the parent of Broods
V and VII, the former an offshoot by acceleration and the latter by retardation of
development. He suggests, however, that the Pennsylvania portion of Brood VII
may have originated independently of the New York part, since it lies in the moun-
tainous country, where the broods would naturally be mixed up more than in any
other part of the range of the 17-year race. In Ohio he notes that the distinct areas

RELATIONSHIP OF SEVENTEEN-YEAR BROODS. 29

relationship between the different broods in point of origin, which
may be indicated as follows:

THE RELATIONSHIP OF THE SEVENTEEN-YEAR BROODS.

From the standpoint of distribution the broods of the 17-year race
may be grouped as follows: (1) Broods I and II; (2) Broods III and
IV; (8) Brood V; (4) Brood VI; () Broods VII, VIII, IX, X, and
XI; (6) Broods XII, XIII, XIV, and XV; (7) Broods XVI and
XVII, the last connecting again with Brood I.

Taking up these broods in regular order:

The main body of Brood I occupies territory immediately west of
the more important and perhaps parent Brood II, and also presents
a number of colonies extending westward to Kansas. Broods I and
II seem, therefore, closely allied in point of origin.

Brood III presents little, if any, relationship to Brood IT in point
of location and distribution, but is closely allied to the following
brood, IV, and the latter is evidently a retarded western and southern
extension of IIT.

Brood V presents little relationship with Brood IV in point of dis-
tribution and covers a very compact territory.

Brood VI, being a widely scattered one, and occurring usually in
small numbers, does not seem to present any particular relationship
with any of the preceding or following broods unless the explanation
suggested by Professor Castle be accepted.

Brood VII is local in distribution and not very important, and is
divided into two sections by the territory occupied by the following
brood, VIII, with which it thus seems to be closely allied. Brood
IX is very distinctly a southern extension of Broods VII and VIII.
These three broods seem, therefore, to be closely allied in their
origin, and, curiously enough, occupy territory which divides the
two main sections of the great 17-year Brood X, which next follows
in regular succession, and is perhaps the oldest or parent brood of
the 17-year race. Brood XI, following X, is evidently an extreme
northeastern extension of the latter.

Brood XII is represented by a series of very doubtful records,
which, if validated in future return periods, will connect the western

covered by Brood VI lie for the most part just outside the area covered by Brood V
and on opposite sides of the latter.

This interpretation by Professor Castle may be in part correct; but in view
of the wide range of this brood and the very scattering nature and separation of
the individual swarms, it seems to me more probable to account for it as a develop-
ment of scattering broods originating for the most part independently by means of
retardation or acceleration from other broods, and none of the colonies developing
enough to fill and hold any very large definite territory. In other words, most of the
colonies are probably of late origin rather than the remnants of an old, extensive,
worn-out brood,

30 THE PERIODICAL CICADA.

Brood XIII with group 5. Brood XIII is the principal western rep-
resentative of group 6, which, through the three broods XIII, XIV,
and XV, extends from the extreme western to the eastern limits of
the Cicada. Brood XIV has a very wide range to the eastward of
XU, and connects with the latter through the colonies in northern
Illinois and Indiana. Brood XV is limited to the Atlantic seaboard
and connects directly with the eastern colonies of XIV.

Brood XVI is based on somewhat doubtful records and is unim-
portant. Brood XVII is intermediate between Brood XVI and
Brood I, its western colonies connected with the former and the
eastern colonies with the latter.

THE RELATIONSHIP OF THE THIRTEEN-YEAR BROODS.

The broods of the 13-year race break up into the following natural
groups: (1) Related closely to Brood XIX, and comprising Broods
XVIII, XIX, and XX; and (2) related to Brood XXIII, and com-
prising Broods XXI, XXII, XXIII, and our new Brood XXIV.

The first of these broods, Brood XVIII, is a rather insignificant
one and is undoubtedly an eastern extension or offshoot of the great
13-year Brood XIX, which succeeds it. Brood XX, is undoubtedly a
section of Brood XIX retarded one year, just as Brood XVIII
consists of accelerated swarms of the same.

Brood XXI, separated from Brood XIX by two years, seems to
bear little relationship to the latter, and a more logical arrangement,
consists in connecting it with Brood XXIII through Brood XXII,
of which last it may be considered as an eastern and northern exten-
sion. Brood XXII is a very marked instance of the formation of a
new brood by an acceleration in time of the appearance of a portion
of a larger and older brood. Its relationship with Brood XXIII is
very marked and can not be questioned. Brood XXIII, the main
representative of this group, is followed by the new Brood XXIV,
which is evidently a retarded swarm of the preceding brood.

Of Broods X XVI and the new Broods XXIX and XXX, both of
which need verification, no significant relationship can be pointed
out. Brood XXIX is very doubtful, and the records are possibly
based on a confusion with the 17-year race.

Sources oF ERROR IN THE OLD REcoRDS.

In examining the records of the distribution of the broods of the
periodical Cicada, it is seen that considerable uncertainty attaches
to the data of certain broods, not only from the fact of their covering,
in greater or less degree, territory occupied by both races, but more
particularly because the records are frequently based on years in which
broods so overlapping have appeared in conjunction.

SOURCES OF ERROR IN THE OLD RECORDS. il

In the case of the broods of the 17-year race, the following extend
on their southern boundaries into the territory of the 13-year race,
and hence the records of the southern localities are open to some
question: Broods VI, X, XIV, XVI, I, IV, to a slight extent also
in the case of Broods IT and III, and doubtfully in the case of Brood
IX, the possibility of confusion in this last brood depending on the
accuracy of the extreme northeastern extension of the 13-year Brood
XIX.

The following broods of the 13-year race extend northward into the
territory occupied by the 17-year race, and hence are open to some
question: Broods XXIII, XVIII, XIX, and XX.

The records can not be questioned on this ground of the 17-year
Broods VII, VIII, XI, XIII, and V, and of the 13-year Broods XXIV,
XXI, and XXII, because these broods are limited in distribution
to the territory of a single race.

The most notable instance of the overlapping and consequent prob-
able confusion of the records is seen in the case of Brood X of the
17-year race with Broods XXIII and XIX of the 13-year race. The
remarkable feature in the distribution of the broods named is the
notable extension northward in Illinois and Missouri of the 13-year
Broods XXIII and XIX, which fills almost exactly a district which
would naturally be supposed to belong to the 17-year race and prob-
ably to Brood X. As pointed out in Bulletin 14, page 26, this cireum-
stance had special significance in view of the fact that the northward
extension of the 13-year race is based on Broods XIX and XXIII,
and that the records prior to 1898 of the former were collected for the
‘most part in 1868, when this brood was in conjunction with Brood X,
and of the latter in 1885, when Brood XXIII was also in conjunction
with Brood X, the limits of which, curiously enough, stop rather
suddenly at or near the eastern State line of Illinois. The possi-
bility was immediately suggested that the northern localities assigned
to Broods XIX and XXIII properly belong to Brood X.

The occasion of the reappearance of the 13-year Brood XXIII in
1898 without any important 17-year brood to confuse the records
and of the 17-year Brood X in 1902, also without a joint occurrence
of any important 13-year brood, gave the opportunity wished for to
determine the validity of old records and to fix more accurately the
distribution of the three broods concerned.

A very thorough canvass was made in 1898 of the territory covered
by Brood XXIII, and especially the territory in doubt, by calling
into requisition the very numerous county correspondents of the
Statistical Division of the Department of Agriculture and also of the
Weather Service in addition to the regular correspondents of the
Division of Entomology. Several thousand replies were received,

SP THE PERIODICAL CICADA.

negative and positive. Reports were also kindly submitted by
Professor Forbes, of Illinois, which added four or five counties to
the records obtained for that State, and other reports were received
from entomologists of other States covered by this brood. <A pre-
liminary report was published in Bulletin 14, and a full report in
Bulletin 18, of this Bureau. The records obtained confirmed the
general accuracy of the old belief of the distribution of Brood XXIII.
The occurrence of scattering colonies of the 17-year Brood VI over
some of the territory adds a slight element of doubt; but in the
main the records given for Brood XXIII, taken in connection with
older records, are probably correctly assigned.

The data obtained of the 17-year Brood X in 1902 is even more
satisfactory, inasmuch as in this case there was no 13-year brood to
throw doubt on any of the records. The same means was taken to
get full reports as were used in 1898; and, rather to our surprise, the
substantial correctness of the old records is strikingly demonstrated,
as seen on the map published in connection with the detailed discus-
sion of this brood. Thirteen-year Brood XXIII covers southern
{llinois, with a scattering outpost through southern Indiana. Brood
X stops, as hitherto believed, near the eastern line of Llinois, with
a few scattering outposts. There is overlapping, but, in the main,
south-central and western L[llnois and eastern and central Missouri
seem to belong to the 13-year race, as hitherto believed.

The recurrence this year of the great 13-year Brood XIX without
any 17-year brood to confuse the records will give an opportunity
to complete the data relative to the distribution of these three over-
lapping broods, but the records already obtained of Broods X and
XXIIT indicate very strongly the probable correctness of the old
records of Brood XIX.

Many of the other scattering records of 13-year broods northward,
or of 17-year broods southward, may possibly be based on similar con-
fusions, arising from the overlapping of broods of the two races.

The only way to accurately define the range of the different broods
is to undertake with each recurrence a thorough and systematic
investigation of all the territory open to the least doubt. Such work
has been repeatedly instituted, and particularly since 1868, and many
of the more strictly limited broods have been very carefully recorded,
and their distribution has been satisfactorily defined. Work of this
kind has been done for Brood III in Iowa by Professor Bessey, and
for Brood V in Ohio and West Virginia by Professors Webster and
Hopkins. Similar work has been done for Brood II in New York
and New Jersey by Doctors Lintner and Smith, and for X and XXIII
by Riley in 1885, and for Brood XIX by Walsh and Riley in 1868.

BROODS OF 14, 15, OR 16 YEAR PERIODS. as

The value of a thorough and systematic canvass of the territory
supposed to be covered by any brood is exhibited in much of the
work referred to above, and notably in the case of Brood V studied
by Professors Webster and Hopkins in Ohio and West Virginia. In
the case of this brood, however, there was no difficulty from an
association with any 13-year brood.

Broops or 14, 15, or 16 YEAR Periops.

The most notable thing about the periodical Cicada is the regularity
with which it has reappeared during more than 200 years of records
at the stated intervals of 13 years for the Southern race and 17 years
for the Northern race. If all the cicadas belonged to a 13-year or a
17-year period—in other words, if there were but one period—this
regularity would be less surprising. But the records are so complete
and full that there can be no doubt whatever of the absolute
uniformity of periods for the two races for the vast majority of the
individuals. That unusual conditions will, however, hasten the
development or retard it a year or more has been already indicated
on page 24, together with notable examples of artificial acceleration.
In view of these last instances there can be no doubt that this regu-
larity of appearance is governed more by the uniformity of tempera-
ture conditions over a long period of years than from any inherent
qualities in the insect itself. If these conditions are interfered with,
however, the Cicada becomes, as it did in the greenhouse at Belvi-
dere, Ill., accelerated one year; and if such conditions occurred in
nature over a large area, as already indicated, a new brood would be
established, but not a 16-year brood, because the climatic conditions
over the long period of seventeen years, would, and evidently have in
practically every instance, carried these accelerated or, conversely,
retarded individuals forward or back to the normal period. There
are, however, a few records which seem to indicate, and particularly
in the overlapping territory of the two broods, a variation in the
length of the subterranean period. These reports of 14-year, 15-year,
or 16-year broods have been so very scanty that it has not been possi-
ble to trace them out with any accuracy, but there seems to be no
reason whatever for doubting the possibility of swarms which have
actually developed and maintained for a time these intermediate
periods. In the course of years we may get enough of these records
to definitely map some of these variant broods.

31117—No. 71—07——3

34 THE PERIODICAL CICADA,
Future APPEARANCES.

During the next seventeen years broods of the 17-year and 13-year
races of the periodical Cicada will occur as follows:

Table of future appearances.

|
Year 17-year race. | 13-year race. Year. 17-year race. 13-year race.
| |
19072225) EXcVi New?....| XIX Major......- 1916...| VII Minor...) XXVIII] No record.
1908....| XVI | Minor....| XX Minor sees |} 1917...| VIDE |....do.:.| XXEX | New?
1 QUO SONS Wall |WIN erat same eNCICN Ieee doverees | TEA WSS [SRE RGIO) 2 Sal DOK Do.
191OE == il iMbbetonmes 3) 2.O;0U0 Boose 00s 2552 |) MOTOS Sa Xe Major-..| XVIII | Minor.
DOME = II Mayor= ao] Sexes Maton). sos | StO202e ENG ll SM Omen! Pee NGID NG Major.
TIDE eee lh ome 5) eel Ose | eNO NGA ING Wire ee | 1921...) XII | New?..- xX Minor.
LOIS ose) Ves aa Gor) eXOnays No record . .|| 1922...| XIII | Major..| XXI Do.
1914._.. V Eek Se olf DG-OVIT Pi huaoheye aa ee NlO23ce ENG: al eon dOL melee NGLIL Do.
1915 VI Minors 22 |) exe Vl ING ween ene | eee XV New?...| XXIII | Major.

In this table the large or important broods are designated as
major; the small or scattering broods as minor. In the latter class
the new and often doubtful broods suggested by the writer also fall.
In the case of a few numbers assigned to the 13-year race no records
of occurrence have been reported, but such may be forthcoming at
any time, although it is evident that the breaking up of the 13-year
race into broeds has not proceeded to anything like the extent that
it has in the 17-year race.

It will be noticed that as a rule a 17-year and 13-year race are
associated in the same year. This is purely accidental, and in point
of fact the same two broods could only come together once in 221
years. The greatest Cicada year of recent times was 1868, when
Brood X, the largest of the 17-year race, appeared in conjunction
with Brood XIX, the largest of the 13-year race. These two broods
will have their next joint occurrence in the year 2089, when perhaps
the increase of settlement and the changed character of vegetation
and superficial conditions over their respective ranges may have
entirely eliminated them except for stragglers.

THE DISTRIBUTION OF THE PERIODICAL CICADA.
SOURCES OF INFORMATION.

The records on which are based the present information of the dis-
tribution of the several broods of the periodical Cicada have been the
accumulation of more than two hundred years, and particularly dur-
ing the last fifty years they have assumed a most voluminous char-
acter, and any effort to discuss the subject at all minutely would
expand this publication beyond reasonable limits. Jt is impossible,
therefore, to detail the evidence which has been used in determining
brood limits or even to summarize the voluminous historical and
chronological records on which this distribution rests. All that is
possible is to continue the plan followed in Bulletin 14 of limiting

GENERAL RANGE OF SPECIES AND RACES. 35

the record to a brief description of the different broods and merely
noting the distribution by States and counties. The data for these
summaries is the rather full account given in Bulletin 8, old series, of
the Division of Entomology, supplemented, however, by the local
studies made by entomologists and others in various States, and par-
‘ticularly the voluminous records obtained by this Bureau, collated
and classified up to 1898 by Mr. E. A. Schwarz, who had long assisted
Professor Riley in collecting such data. Since 1898 this field of
inquiry has been under the charge of the writer, and a very thorough-
going effort has been made to get full and accurate data of the
broods which have appeared from year to year. The records for the
important 13-year Brood XXIII, which appeared in 1898, in conjunc-
tion with the 17-year Brood VI, and of Brood X, the largest of all
the 17-year broods, which appeared in 1902, were especially complete
and satisfactory, and are summarized under the accounts of these
broods. Particularly in later years, much exact information as to
local distribution has come from the active cooperation of State
entomologists, who have often been able to get more detailed and
accurate reports than was possible through the correspondents of this
office. The scant records, indicating perhaps scattering or incipient
broods, covering some of the blanks in the 13 and 17 year series, are
introduced in their proper order for future confirmation or rejection.
The records obtained by the Department of Agriculture, covering
nearly thirty years, have become very voluminous, and during the
last few years an effort has been made to go over all of these records
and transfer the important information to index cards, and all the
later records are being kept on such cards. It is expected also, as
time offers, to incorporate in this record all the data from experi-
ment station bulletins and other printed records. Ultimately, there-
fore, we shall have a classified card record which will be easily avail-
able for examination and study and which will assist greatly in estab-
lishing brood limits and determining the status of new reports.

THE GENERAL RANGE OF THE SPECIES AND OF THE TWO RACES.

Taking all the different broods together, this Cicada is known to
occur pretty generally within the United States east of the one
hundredth meridian and northward of latitude 30°—in other words,
east of central Kansas and north of northern Florida. No broods
have been found in northern New England except a doubtful record
in Vermont, nor west of the Mississippi above Iowa. The State of
Rhode Island, in which the Cicada was long believed to be absent,
proved to harbor a small brood, as discovered in 1903 (Brood XI).
The most eastward occurrences are the swarms occurring in Barn-
stable County, Cape Cod, Massachusetts, and on the island of Marthas
Vineyard. No colonies have been found on the peninsula of Florida,

36 THE PERIODICAL CICADA,

although the Cicada occurs in the northwestern portion of the State,
The western records reported in Bulletin 14 in Colorado, and doubt-
ful occurrences along the northern slope of the Big Horn Mountains
of western Wyoming and Montana, have been shown, with very little
doubt, to belong to another species of Cicada (Tibicen cruentifera
Uhl. and allies), very possibly also similarly periodic in reappearance.

The territory covered by the periodical Cicada is graphically illus-
trated by the two maps showing the range of the 13-year and the
17-year races, respectively (figs. 2 and 3). <A brief examination of
these maps develops the very interesting and suggestive fact that if
superimposed the areas occupied by the two races would, in a gen-

Fic. 2.—Map showing distribution of the broods of the 13-year race.

eral way, fit together along their adjoining sides. This was to have
been expected, but one would hardly have predicted the notable
northern extension of the 13-year race in Missouri and Illinois in the
Mississippi Valley, following, however, in an exaggerated way, the
isothermal lines of this region. The extension northward of the 13-
year race very greatly exceeds the limits of the Lower Austral zone,
as marked on Merriam’s map, and if this insect were taken as a basis
this zone would have to be very greatly extended northward in the
two States named. With this important exception, the 13-year race
is confined pretty closely to the Lower Austral and the 17-year race
covers the Upper Austral, with large extensions northward into the

GENERAL RANGE OF SPECIES AND RACES, 5)

Transition zone. The overlapping of the two races, discussed else-
where, is well illustrated by these two maps.

The range of the individual broods is undoubtedly much greater
than the limits now assigned, since the records until recent years
have been largely based on notable and dense swarms and have
rarely taken into account the scattering individuals, which undoubt-
edly extend over a much greater territory and usually pass unno-
ticed. The very careful records secured of the broods, including and
subsequent to 1898, have shown much of: this scattering occurrence
beyond the denser brood limits, as will be seen in the maps illus-
trating these broods. This indicates that the breaking up of the

HAMPS!

B Wiw

=
o
“)
Oo
7)
v
v

Fig. 3.—Map showing distribution of the broods of the 17-year race.

Cicada has already gone much farther than was hitherto supposed,
and points to the ultimate disappearance of great broods as such
and their replacement as scattering individuals every year. The dis-
appearance of the great broods, however, is not to be anticipated
in the very near future, and may not come about for a thousand or
even several thousand years. This is shown by the fact that the
broods first seen by the early colonists in New England on Cape
Cod, at Plymouth, and on Marthas Vineyard are, as elsewhere noted,
still practically unreduced in numbers and make just as startling an
impression as ever. This is due to the fact that much woodland
remains undisturbed in these localities. In other places, where the
woods have been largely removed as the result of settlement, the
Cicada has correspondingly disappeared.

38 THE PERIODICAL CICADA,

THE RANGE OF THE WELL-ESTABLISHED BROODS, TAKEN IN NUMERICAL
ORDER.

In the following description of the broods they are taken up in
their numerical order—first, the 17-year broods, I to XVII, and then
the 13-year broods, XVIII to XXX; that is, as many of the latter
as have definite records. The chronological order of the broods, show-
ing the broods of the two races which occur jointly in the same year,
is indicated in the table on page 34. This arrangement, rather
than a chronological one, is adopted for the reason that any chrono-
logical arrangement in the course of a few years becomes obsolete,
and for the same reason individual maps of the broods have been
made, rather than joining in one map the two broods that may hap-
pen to occur together on each of the next thirteen or seventeen years.
The maps of important broods which have been recently more care-
fully studied have been entirely revised, and the importance of the
records has been indicated by the size of the dots, the large dots
representing counties in which the brood occurred in one or more
dense characteristic swarms and the small dots, records of scatter-
ing occurrence or of doubtful validity. These same conditions are
more accurately shown in the State and county records, as described
under each brood. Such indications will be secured for all the broods
in course of time, and will give a much more accurate picture of
actual conditions than the old system of uniform dots for all records.
The maps of broods which have not been recently studied have also
been reengraved because of the discovery of new records—in some
cases few in number, in other cases of considerable amount.

Broods of the Seventeen- Year Race.
Broopv I—Septendecim—1910. (Fig. 4.)

Brood I is the first of the series of well-authenticated broods of
the 17-year race, and its main swarms occupy the territory immedi-
ately west of the more important Brood II, which follows the year
after. It mecludes also widely separated swarms extending west into
Kansas. It was established originally on data given by Dr. Gideon B.
Smith, but its distribution is now more definitely recorded as a result
of the study given it in 1893 by Professor Riley and of records which
have come to this Bureau in connection with the study of other
broods since that time. Several new counties for West Virginia
were added by Doctor Hopkins in Bulletin 68, West Virginia Experi-
ment Station (1900).

The doubtful records prior to 1893 were those relating to the occur-
rence of this brood in Kansas and Colorado. The localities in Kansas
received doubtful confirmation in 1893. The Colorado localities
remained unverified, although the district mentioned was visited

BROOD II—SEPTENDECIM—1911. 39

and special search was made for evidence of the insect. Undoubtedly
the Colorado occurrence relates to some other and probably also a
periodic species, such as that reported for another brood at Boulder,
Colo. (XVI), and for Brood VI in Montana.

The distribution, by States and counties, follows:

District or CotumBtA.—North of Washington.
Intrnors.—Madison(?).

InprANA.—Knox, Posey, Sullivan.
Kansas.—Dickinson, Leavenworth.
Kentucky.—Trimble.

MaryLanp.—Prince George, south half of St. Mary.

Fic. 4.—Map showing distribution of Brood I, 1910.

Norra Carortrna.—From Raleigh, Wake County, to northern line of State; Cabarrus,
Davie, Iredell, Rowan, Surry, Yadkin.

PENNSYLVANIA.—Adams, Cumberland, Franklin.

Vireinta.—From Petersburg, Dinwiddie County, to southern line of State; Bed-
ford, King William, New Kent, Rockbridge; valley from Potomac to Tennessee and
North Carolina boundary.

West VireaintA.—Grant, Hardy, Pendleton, Randolph.

Broop II—Septendecim—1911. (Fig. 5.)

This brood occupies, for the most part, territory immediately east
of Brood I, and is one of the best recorded of the broods, since its
almost exclusively eastern range brings it in the immediate vicinity

AQ) THE PERIODICAL CICADA,

of the large towns and more densely populated districts of the Atlantic
seaboard.

Fitch described its limits as his Brood No. 2, Walsh-Riley as Brood
VIII, and Riley’ as his Brood XII. It has been reported in Con-
necticut regularly every seventeen years since 1724, and in New
Jersey since 1775, if not earlier, and almost equally long records of
it in other States have been made.

On the occasion of its last appearance, in 1894, its distribution in
New Jersey was very carefully studied by Prof. J. B. Smith, confirm-
ing 1ts occurrence in every county in that State, and in New York

fn
!
o
o |
a
ae
v
\
a

Fic. 5.—Map showing distribution of Brood IT, 1911.

similar studies were made by Dr. J. A. Lintner. The Bureau also
received a vast number of reports from these and other States in
answer to a circular prepared by Professor Riley and mailed in May,
1894. Some of the southern records obtained in 1894 are doubtful,
and this applies especially to the localities in North Carolina, because
of the occurrence that year also of Brood XIX of the 13-year race.

The distribution as listed below is based on the old records given in
the circular cited, with such additions and corrections as the reports of
appearance in 1894 made necessary.

The distribution, by States and counties, is as follows:

Connecticut.—Fairfield, Hartford, Litchfield, Middlesex, New Haven.
Districr or CotumBrA.—Throughout.

BROOD IlI—SEPTENDECIM—1912. 4}

InprANA.—Dearborn, Posey(?).

Marytanp.—Anne Arundel, Calvert, Charles, Prince George, St. Mary.

Micnican.—Kalamazoo.

New Jersey.—Entire State.

New Yorx.—Albany, Columbia, Dutchess, Greene, Orange, Putnam, Rensselaer,
Rockland, Saratoga, Ulster, Washington, Westchester, and on Staten Island and Long
Island.

NortH Caro.tina.—Bertie(?), Davie(?), Forsyth(?), Guilford, Orange, Rocking-
ham, Rowan, Stokes, Surry, Wake(?), Warren(%), Yadkin(?).

PENNSYLVANIA.—Berks, Bucks, Chester, Dauphin, Delaware, Lancaster, Lebanon,
Lehigh, Montgomery, Northampton, Philadelphia, Pike, Potter, Schuylkill, Wyoming.

Vrrernta.—Albemarle, Alexandria, Amherst, Appomattox, Bedford, Buckingham,

! ei OAKOTA

Fiac. 6.—Map showing distribution of Brood III, 1912.

Campbell, Caroline, Charlotte, Culpeper, Fairfax, Fauquier, Fluvanna, Goochland,
Hanover, Henrico, James City, Loudoun, Louisa, Lunenburg, Madison, Page, Pitt-
sylvania, Powhatan, Prince Edward, Rappahannock, Spottsylvania, Stafford.

West Vrrerta.—Brooke(?).

Broop III—Septendecim—1912. (Fig. 6.)

This brood, described by Walsh-Riley as Brood IX (XIII of Riley)
is one of the more important of the Western 17-year broods, its
most compact body lying in the States of Iowa and Missouri. It is
closely allied in distribution to Brood IV, but shows little relation-
ship with Brood II. Records are given by Dr. G. B. Smith in both
Jowa and Illinois in 1844, and it has been regularly recorded since,

Arar THE PERIODICAL CICADA,

over at least a portion of its range. The Iowa distribution of the
brood was carefully studied by Professor Bessey in 1878.

The range of the brood as given below is based on the published
records, together with a number of additional localities collected from
the correspondence of the Bureau.

The distribution, by States and counties, is as follows:

InuiNors.—Champaign, Fulton, Hancock, McDonough, Mason, Warren.

Towa.— Adair, Adams, Audubon, Boone, Cass, Dallas, Davis, Decatur, Des Moines,
Greene, Hamilton, Henry, Iowa, Jasper, Jefferson, Johnson, Keokuk, Louisa, Mad-
ison, Mahaska, Marion, Marshall, Monroe, Muscatine, Polk, Poweshiek, Ringgold,
Scott, Story, Taylor, Union, Van Buren, Wapello, Warren, Wayne, Webster.

et
NS

WISCONSIN

JERMON?T

1
1
HLL} NOIS !

=
oy)
o
)
)
v
v

Fic. 7.—Map showing distribution of Brood IV, 1913.

Missourt.—Bates, Buchanan, Clark (?), Grundy, Henry, Johnson, Knox (?),
Lewis (?), Macon (?), Marion (?), Monroe (?), Putnam, Ralls (?), Randolph (?), Schuy-
ler (?), Scotland (?), Shelby.

NEBRASKA.—Johnson.

Oxnto.—Champaign.

West Virainta.—Mononegalia.

Broop I1V—Septendecim—1913. (Fig. 7.)

This brood, described by Walsh-Riley as Brood X (Riley XIV)
succeeds Brood III by one year, and in the main appears to be a
southwestern extension of the latter, covering a portion of south-
western Iowa, eastern Kansas, and Indian Territory, with detached
localities in Missouri and other States. Its original connection with

BROOD V—SEPTENDECIM—-1914. 43

Brood III is apparently well shown by the adjoining or overlapping
territory occupied by the two broods, together with the fact of their
separation by a single year.

This brood was well recorded in 1879, the data being published by
Professor Riley in Bulletin 8, old series, of the Division of Ento-
mology. A number of additional records were obtained at its last
appearance in 1896, and reports have been received since the publica-
tion of Bulletin 14 adding five new counties in northwestern Missouri.

The distribution of the brood as now determined is as follows:

ARKANSAS.—Hempstead (?).

InpraAn Trerrirory.—Muscogee, Tulsa.

Towa.—Adams, Cass, Dallas, Fremont, Mills, Montgomery, Page, Pottawattamie,
Taylor.

Kansas.—Allen, Bourbon, Chase, Coffey, Douglas, Greenwood, Jackson, Johnson,
Labette, Lyon, Marion, Morris, Osage, Pottawatomie, Wabaunsee, Wilson, Woodson,
Wyandotte.

Missourr.—Barton, Buchanan, Caldwell, Dekalb, Grundy, Henry, Holt, Jack-
son, Johnson, Lafayette, Mercer, Ray, Saline, Vernon.

NEBRASKA.—Otoe.

Trxas.—Cooke, Denton, Fannin, Kaufman, Wise.

Broop V—Septendecom—1914. (Fig. 8.)

Brood V covers in the main a rather compact territory and does not
connect directly with preceding broods, except possibly through
Brood VI, joining the following important series of broods of the
Alleghany region. Brood V was reported from Ohio as early as 1795.
Fitch described it as Brood 5, Walsh-Riley as Brood XI, and Riley
as Brood XV.

The limits of this brood as known prior to 1897, the date of its last
appearance, were given by Mr. Schwarz in Circular No. ‘22 of this
Bureau. In 1897 its distribution in Ohio was very carefully studied
and mapped by Professor Webster and in West Virginia by Professor
Hopkins. The distribution as listed below is based on the above in-
formation, together with numerous records which have since been
obtained by this Bureau in the investigation of this and other broods.

The distribution, by State and counties, of this brood as now known
is as follows:

Onto.—Ashland, Athens, Belmont, Carroll, Columbiana, Coshocton, Crawford, Cuya-
hoga, Delaware, Erie, Fairfield, Franklin, Gallia, Geauga, Guernsey, Harrison, Hock-
ing, Holmes, Huron, Jackson, Jefferson, Knox, Lake, Licking, Lorain, Mahoning,
Medina, Meigs, Monroe, Morgan, Muskingum, Noble, Perry, Pickaway, Pike,
Portage, Richland, Ross, Sandusky, Scioto, Seneca, Stark, Summit, Tuscarawas,
Vinton, Washington, Wayne.

PENNSYLVANIA.—Fayette, Greene, Washington.

Virainia.—Augusta, Caroline, Highland(?), Shenandoah.

West ViraintA.—Barbour, Boone, Braxton, Brooke, Calhoun, Clay, Doddridge,
Fayette, Gilmer, Grant, Greenbrier(?), Hancock, Hardy, Harrison, Jackson, Kana-

44 THE PERIODICAL CICADA,

wha, Lewis. Marion, Marshall, Mason, Mineral, Monongalia, Nicholas, Ohio, Pleas-
ants, Pocahontas, Preston, Putnam, Randolph, Ritchie, Roane, Summers(?), Taylor,
Tucker, Tyler, Upshur, Wayne, Webster, Wetzel, Wirt, Wood.

Broop VI—Septendecim—1915. (Fig. 9.)

This is an unimportant scattering brood designated as No. 7 by
Fitch, XII by Walsh-Riley, and XVII by Riley. It is difficult to
assign any very pointed relationship for this brood, either with pre-
ceding or following broods, unless one adopts the suggestion made by
Prof. W. EK. Castle that it represents a relatively old or played-out
brood, and may thus be considered the parent of Broods V and VII,

Fic. 8.—Map showing distribution of Brood V, 1914. .

the former the offshoot by acceleration and the latter by retardation
of development. (See pp. 28-29.) As stated elsewhere, however, it is
more likely to be an assemblage of swarms of diverse origin.

This brood, while not an important one, covers a much wider ter-
ritory than any of the other 17-year broods. With the exception,
however, of the two extremes of its distribution in the Northwest
and the Southeast, respectively, the records are of scattering indi-
viduals, in many localities only a few specimens being observed. To
illustrate this graphically on the accompanying map (fig. 9), the
small dots indicate localities where only a few specimens were
observed or captured or a doubtful record and the large dots localities

BROOD VI—SEPTENDECIM—1915. 45

represented by one or more dense swarms, such as are ordinarily
_ characteristic of the species. Some of these records of scattering
occurrence may be based on stragglers from preceding broods or
accelerated individuals from following broods and therefore may not
mean more than incipient swarms. Many of the records were secured
in 1898, when a very careful canvass of the whole Cicada region was
made by this Bureau with the assistance of the State entomologists.

The reports obtained in 1898, if they may be relied upon, extend
the range of the periodical Cicada in Wisconsin and Michigan much
farther north than any of the old records. The localities assigned
to this brood in North Carolina, South Carolina, and Georgia, and in

Fic. 9.—Map showing distribution of Brood VI, 1915.

eastern Kentucky and Tennessee are, in the main, in counties in the
elevated mountainous district, and the correctness of the reference
to this brood is established by earlier records as well as indicated by
the elevation.

Reports of the occurrence of this brood in Montana were sent in by
Mr. E. V. Wilcox, with the statement that the insect occurred in
smail numbers in the counties of Chouteau, Flathead, Gallatin, and
Missoula, and that in the latter county some damage was done to
young apple trees. This report was published in Bulletin 18 of this
Bureau, but doubts arose afterwards in the mind of the writer as to
the correctness of the determination of the Cicada, as the more

46 THE PERIODICAL CICADA,

recently acquired knowledge of the existence of another periodical
species in the northwestern United States threw some doubt on this
reference, and an examination of collected material from that region
indicates that the species referred to is Tibicen cruentifera Uhl.,
which apparently is also periodic and has other habits closely
resembling septendecim.

The records of distribution given below are as published in Bulletin
18 of this Bureau, with the exception of West Virginia, where a good
many counties have been added from Doctor Hopkins’s Bulletin 68
and from later records secured by him. The starred counties indi-
cate the occurrence of the Cicada in one or more characteristic dense
swarms; the italicized counties are confirmations of old records, and
the counties inclosed in parentheses are old records not reported in
1898. The distribution, by States and counties, follows:

DELAWARE.— Newcastle.

Districr oF CoLUMBIA.—Several localities.

GrorGiA.—Dade,* Elbert, Floyd, Habersham,* Hall,* Paulding, Rabun,* Spalding,
White.

Tuirnois.—Dewitt,* Douglas, Knox, McLean, Montgomery, Scott, Shelby,* Ver-
milion.

Inp1ANA.—Boone, Brown, Carroll, Grant, Johnson, Laporte, Wells.

Kentucky.—Letcher.*

MaryLanp.—Carroll, Cecil, Montgomery, Prince George, Washington.

MicuiGcAN.—Barry, (Cass?), Chippewa, Genesee,* Houghton,* Kent(?), Macomb(?),
Newaygo(?), Ogemaw(’), Otsego,* Shiawassee,* Washtenaw.

New Jersey.—Bergen, Cumberland, Esser, Hudson, Hunterdon, Mercer, Middlesex,
Morris, Passaic, Somerset, Union. ‘

New Yorx.—Greene, New York, Richmond, Schenectady, (Westchester).

Norru Caroriva.—Alexander,* Bladen, Buncombe, Burke,* Cabarrus, Caldwell,*
Catawba,* Henderson,* Iredell, Lincoln,* McDowell,* Macon,* Montgomery,
Moore, Pender,* Polk,* Randolph(?), Rutherford, Swain,* Transylvania,*
Union,* Washington (?), Wilkes.*

Oxnto.—(Ashtabula), Carroll, Champaign, Columbiana, Delaware, Madison, Mahoning,
Montgomery, Morrow, Pickaway, Shelby, (Summit?), Union, (Vinton?).

PENNSYLVANIA.—Bucks, (Dauphin), (Lancaster), Montgomery, (Northampton and
adjoining counties), (Philadelphia), Westmoreland.

SourH CaroLtrna.—Oconee.*

TENNESSEE.—Bradley, Greene, Hamilton, Jefferson, Knox, Meigs, Polk, Sullivan.

VireiniA.—Charlotte, Chesterfield, Fairfax, Powhatan, Prince Edward, (Smyth).

West Virainia.—Berkeley, Brooke, Clay, Fayette, Grant, Hampshire, Hancock,
Hardy, Jefferson, Marshall, Mineral, Monongalia, Monroe, Morgan, Ohio, Pendle-
ton, Pocahontas, Preston, Raleigh, Tucker, Tyler, Webster.

Wisconsin.—Burnett,* Columbia, Crawford, Dane,* Fond du Lac, Green Lake,* (La
Crosse), Marquette,* Sauk,* Sawyer, Washburn, Waushara.*

Broopv VII—Septendectm—1916. (Fig. 10.)

This brood was founded by Professor Riley in 1869 on Doctor
Smith’s register, in which it is recorded from 1797 to 1848 as occurring
in certain counties in western New York. As indicated elsewhere,

BROOD VIII—SEPTENDECIM—1917. AT

this brood is not very important and is divided into two sections by
the following brood, VITI.

The confirmations of the occurrence of this brood in New York in
later years are reported in Bulletin No. 8, old series, Division of
Entomology. The localities in Pennsylvania and West Virginia are
based on later Divisional records.

The distribution, by States and counties, is as follows:

New Yorx«.—Cayuga, Livingston, Madison, Monroe, Onondaga, Ontario, Wyoming,
Yates.

PENNSYLVANTA.—Allegheny, Washington.

West V1IrGINIA.—Summers?.

= me
wer Nr
>
C)

t
2°

ldd ISSISSIW

Fig. 10.—Map showing distribution of Brood VII, 1916.

Broop VIII—Septendecim—1917. (Fig. 11.)

This is Fitch’s second brood which he described as occurring in
western New York, western Pennsylvania, and eastern Ohio, and is
Brood XIV of Walsh-Riley, and XX of Riley. Dr. G. B. Smith also
gives valuable data relative to its appearance and distribution.

It is one of the smaller broods and did not attract much attention
on its appearance in 1883, but records of a number of additional
swarms were obtained on the occasion of its appearance in 1900.
The main territory covered by it is a rather compact one, lying in
western Pennsylvania, eastern Ohio, and the panhandle of West Vir-
ginia. The swarms in the area thus included probably originated

AS THE PERIODICAL CICADA,

by retardation from Brood VII, owing to mountain conditions as
affecting temperature.

The widely separated swarm occurring on Marthas Vineyard has
exceptional interest on account of the abundance of the insect and
its extreme eastern location. This swarm has been well recorded
since the time of Harris, and in 1900, when it last appeared, was
reported by Prof. H. T. Fernald as being as abundant as ever.

Of the other scattermg swarms the ones in western New York and
in northern Illinois and in South Carolina are old records but ex-
tremely doubtful, and possibly based on confusion of some annual
species of Cicada with the periodical species. No confirmations of

Fig. 11.—Map showing distribution of Brood VIII, 1917.

these records were obtained in 1900. New records were, however,
obtained for New Jersey, Maryland, and North Carolina considerably
away from the main body of the brood and very possibly having a
different origi. None of the records in these three States represents
important swarms, but merely scattering individuals. Some new
records were obtained also in Ohio, Pennsylvania, and West Virginia,
which, however, fall in with the general range of the main body of the
brood.

The county indications in the list below are as with other recently
studied broods, i. e., the star (*) means occurrence in swarms; italics,
confirmation of old records; and parentheses (), failure to secure such

BROOD IX—SEPTENDECIM—1918. 49

confirmation. The large dots on the map indicate starred counties -
and the small ones doubtful records or scattered presence.
The distribution, by States and counties, is as follows:

Intinors.—(Whiteside) (?).
’ MaryLanp.—Harford.

Massacuusettrs.—Dukes* ( Marthas Vineyard).

New Jersrey.—Essex.

New Yorx.—(Chautauqua) (?).

Norru CaroLina.—Moore (?).

Oxnto.—Belmont, Carroll,* Columbiana,* Hamilton, Jefferson,* Mahoning ,* Portage,*
Stark,* Trumbull.*

PENNSYLVANIA.—Allegheny, Armstrong,* Beaver,* Butler,* Cambria,* Clarion,*

eho
1
| :

=
2)
o
o
@)
v
v

Fig. 12.—Map showing distribution of Brood IX, 1918.

Crawford, Fayette, (Forest), Huntingdon, Indiana,* (Jefferson), Lawrence,* Mercer,*
(Snyder), Venango,* Washington,* Westmoreland.

SoutH Caroitrna.—Barnwell (?).

West Vrreinta.—Brooke,* Hancock,* (Marshall), Ohio.

Broop 1X—Septendecim—1918. (Fig. 12.)

In the main this brood (XV Walsh-Riley, XXI Riley) covers a
rather compact territory, extending from the southern part of West
Virginia across Virginia into North Carolina, and is the southern
extension of Brood VIII one year retarded. Some widely separated
swarms have been reported from Ohio and one from northern Vir-

31117—No. 71—07—4

50 THE PERIODICAL CICADA,

ginia, and one or two in northern West Virginia, but ir the main these
are doubtful records or unimportant, and may possibly not be con-
nected in origin with the swarms occurring in the main territory of the
brood. Since the publication of Bulletin 14 several additional coun-
ties have been reported for Virginia and West Virginia, the new
records for the latter State being chiefly from a very careful survey
made by Doctor Hopkins in 1901. Equally careful search would
doubtless show for adjoining States the wide scattering occurrence
which Doctor Hopkins has found in West Virginia. The unimportant
records are indicated on the map by the small dots.

The occurrence of a swarm on Marthas Vineyard in 1833 is recorded
by Doctor Harris, but the records of subsequent appearances of this
swarm have shown the date mentioned to be unquestionably an error
for 1832, which refers this swarm to Brood VIII.

The distribution, by States and counties, is as follows:

Norru Carouina.—Alleghany,* (Wilkes*).

Onto.—Cuyahoga, Madison?, (Medina?).

Virainta.6—Bland,* Buchanan,* Carroll,* (Craig),* Floyd,* Franklin,* (Giles),*
(Grayson),* (Henry),* Lee,* (Loudoun), (Montgomery),* Patrick,* (Pulaski),*
(Roanoke),* Smyth,* Wythe.*

West VireintA.—Barbour, Berkeley, Boone,* Braxton,* Clay,* Fayette,* Green-
brier,* Hampshire, Hardy, Harrison, Jackson, Jefferson, Kanawha, Logan, Marshall,
Mason, Mercer,* Monongalia, Monroe,* Nicholas,* Pleasants, Pocahontas,* Preston ,*
Putnam, Raleigh,* Randolph,* Roane, Summers,* Tucker, Tyler, Upshur, Webster,
Wetzel, Wood, Wyoming.*

Broop X—Septendecim—1919. (Fig. 13.)

This is the great 17-year brood occurring over the main areas
covered by it in numerous dense swarms and equaling, if not exceeding
in importance the largest of the 13-year broods, namely, Brood XIX.
It is Brood No. 4 of Fitch, XVI of Walsh-Riley, and XXII of Riley.
It has been well recorded, particularly in the East, from 1715 to 1902,
the date of its last appearance. It so happened, however, that on
each of the years (1868 and. 1885) when it was especially studied
prior to 1902 it appeared in conjunction with an important 13-year
brood, and as the territories of the two races overlap, there has always
been some doubt as to the correctness of the references of swarms in
such overlapping regions. In 1868, when it was studied carefully by
Walsh and Riley, it was in conjunction with the largest of the 13-year
broods, namely, Brood XIX. In 1885 it was in conjunction with the
second largest of the 13-year broods, namely, Brood XXIII. The

aCounty names in italics are confirmations of old records, names in parentheses
are old records unconfirmed, and starred names indicate occurrence in swarms.

b The old records for Virginia not confirmed specifically in 1901 are in the midst of
counties with large swarms and were confirmed in a general report covering the south-
western part of the State. ,

BROOD X—SEPTENDECIM—1919. 5l

year 1902 was not marked by the recurrence of any 13-year brood,
and hence the records for that year, for the first time, could practi-
cally all be assigned without question to Brood X. It had been
anticipated by the writer and others that many of the records in
middle and southern Illinois, for example, and northern Missouri,
which had been referred to the two large 13-year broods, might pos-
sibly belong to Brood X of the 17-year race. Rather to our surprise,
however, the old limits of distribution for the three broods in question
seem to be pretty definitely confirmed.

Very thorough plans were made early in 1902 to have the entire
territory over which the brood was expected fully and adequately

TLL NG!

Avr NS) LA iS} ae:
| An

YdgISSiISSiw

Fic. 13.—Map showing distribution of Brood X, 1919.

reported, and the responses received by this Bureau were numerous
and satisfactory for practically the whole area covered by the brood.
In addition to this several of the entomologists of the different States
within the range of the brood carried on independent investigations,
and the records obtained by them, most of which have been pub-
lished since, have been incorporated with the reports received by this
Bureau. All of the records agree in showing the substantial accuracy
of the limits of this brood as hitherto platted. The State records
available and used in the following list of States and counties, so far
as they represent counties new to our records, are those reported by
Pettit for Michigan, Smith for New Jersey, Sanderson for Delaware,

52 THE PERIODICAL CICADA.

Garman for Kentucky, Quaintance for Maryland, Felt for New York,
and Hopkins for West Virginia.

Leaving out the numerous scattering colonies the brood occupies
three important regions: (1) An eastern region, covering Long Island,
New Jersey, southeastern Pennsylvania, northern West Virginia and
Virginia, and most of Delaware and Maryland; (2) a southern region,
covering the Lower Alleghanies in northern Georgia, Tennessee, and
North Carolina; and (3) a middle western region, covering western
Ohio, southern Michigan, all of Indiana, except the lake shore
(Webster), and the eastern line of Illinois. Many reports of scatter-
ing occurrence or of chance individuals connect these three regions
and also extend the range westward as far as the Missouri River
(Iowa) and northward into New York and Vermont and _ possibly
Massachusetts. But in the main the three regions designated include
the abundant appearance of the brood in dense swarms.

Where conditions had remained at all favorable there was very
little falling off in abundance of the Cicada at the time of its last
appearance in 1902. In the District of Columbia, where the writer
personally observed the insect, emergence began with the second week
in May and was a fairly prolonged one, extending over three or four
weeks. In this brood the small variety, cassini, is perhaps better
represented than in any other, and this small variety appeared very
generally in 1902, and in great numbers. The deposition of eggs
began about June 1 and continued with considerable activity until
the middle of this month and was of sufficient amount to kill the
terminal branches of trees, in some cases almost all the outer branches
dying.

A big transplanting of eggs was made from the surrounding forests
of the District to the grounds of the Department, where very few
Cicadas had appeared, and all of them had been destroyed by birds
before any egg laying had been done. This planting was made in the
oak grove on the west side of the grounds, where similar experiments
had formerly been in progress, so as to afford material for study of the
development of the larve.

The records of distribution given below include all available data.
The starred counties are those in which the Cicada appeared in 1902
in one or more dense swarms. The italicized counties confirm older
records, and the counties in parentheses are old records which failed
of confirmation in 1902. This last does not necessarily mean that the
Cicada was absent from these counties, but simply the failure to
receive reports of occurrence. <A great many negative reports were
received, and, as platted on a large study map, confirmed the accu-
racy of the range of this brood as now given.

BROOD X—SEPTENDECIM—1919. 53

The distribution, by States and counties, follows:

AtaBAMA.—Cleburne, Jackson, Jefferson, Morgan, (St. Clair) (?).

District OF COLUMBIA.*

DELAWARE.—( Kent), Newcastle,* Sussex.*

Grorata.—Banks,* Chattooga, Dade, Dawson, Fannin,* Forsyth,* (Franklin), Gil-
mer,* Gordon,* Greene, Habersham,* Hall,* Jackson,* Lincoln, Lumpkin,* Murray,*
Newton,* Oglethorpe, Pickens,* Rabun,* Union,* Walker, Walton, White,* Whitfield,
Wilkes.

Inurnois.—Alexander, Clark,* Crawford,* Cumberland, (Dewitt), Edgar,* Edwards,
(Gallatin), Hamilton, Hardin, (Iroquois), Jackson, (Kane), Lawrence, Logan, (Pope),
Saline, Tazewell, Union, Vermilion,* Wabash, White,* Williamson.

Inp1ana.—Adams,* Allen, Bartholomew,* Benton, Blackford,* Boone,* Brown,* Car-
roll,* Cass,* Clark,* Clay,* Clinton,* Daviess,* Dearborn,* Decatur,* Dekalb,* Dela-
ware,* Dubois,* Elkhart,* Fayette,* Floyd,* Fountain,* Franklin,* Fulton,* Gibson,*
Grant,* Greene,* Hamilton,* Hancock,* Harrison,* Hendricks,* Henry,* Howard,* Hunt-
ington,* Jackson,* Jay, Jefferson,* Jennings,* Johnson,* Knox,* Kosciusko,* Lake,
Laporte,* Lawrence,* Madison,* Marion,* Marshall,* Martin,* Miami,* Monroe,* Mont-
gomery,* Morgan,* Noble,* Ohio,* Orange,* Owen,* Parke,* Perry,* Pike,* Porter,*
Posey, Pulaski,* Putnam,* Randolph,* Ripley,* Rush,* St. Joseph,* Scott,* Shelby,*
Spencer,* Starke,* Steuben,* Sullivan,* Switzerland,* Tippecanoe,* Tipton, Union,*
Vanderburg,* Vermilion,* Vigo,* Warren,* Warrick,* Washington,* Wayne,* Wells,*
White.*

Towa.—Woodbury.

Kentucxy.—Allen, Anderson, Barren, Bath, Bell, Boone,* Boyd, Breckinridge,*
Butler, Caldwell, Campbell,* Carroll,* Carter, Casey, Christian, Clay, Clinton, Crit-
tenden, Cumberland, Daviess,* Edmonson, Fayette, Fleming, Franklin, Gallatin,*
Garrard, Grant,* Grayson,* Green, Greenup, Hancock,* Hardin, Harrison,* (Hart),
Henderson, Hickman, Hopkins, Jefferson,* Johnson, Kenton,* Knox, Larue, Laurel,
Lawrence, Lee, Leslie, Letcher, Lewis, Lincoln, Livingston, McLean,* Madison, Magof-
fin, Martin, Meade,* (Mercer), Monroe, Nelson, Nicholas, Ohio,* Oldham,* Owen,*
Owsley, Pendleton,* Pike, Scott, Shelby,* Trigg, Trimble,* Unioh, Warren, Wash-
ington, Wayne, Webster, Whitley, Wolfe,

MaryLaNnp.—Allegany,* Anne Arundel,* Baltimore,* Calvert, Caroline, Carroll,*
Cecil,* Frederick,* Garrett,* Harford,* Howard,* Kent,*’ Montgomery,* Prince George,*
Queen Anne, Talbot,* Washington,* Wicomico.

Massacuuserrs.—(Bristol) (?), Worcester(?).

Micuican.—(Barry), Branch,* Calhoun,* Cass,* (Eaton), Genesee,* (Gratiot), Hills-
dale, Ionia, (Jackson), Kalamazoo,* Lake, Lenawee, (Livingston), Missaukee, (Mon-
roe), Muskegon, Newaygo, Oakland,* Saginaw, St. Clair, St. Joseph,* Van Buren,
Washtenaw ,* (Wayne).

New Jersey.—Burlington,* Camden,* Cumberland ,* Gloucester,* Hunterdon,* Mer-
cer,* Middlesex,* Monmouth,* Morris,* Ocean, (Passaic), Salem,* Somerset,* Warren.*

New Yorx.—Columbia, Kings, (Monroe), Nassau, (Niagara), Ontario, Queens,*
Richmond, Suffolk.*

NorrH Carortina.—Alexander, Alleghany, Burke,* (Caldwell) (?), Catawba, Chero-
kee,* Davidson, Davie,* Lincoln, Stokes, Surry,* (Wake) (2), Wilkes,* Yadkin.*

Outo.—(Adams), Allen,* Auglaize, Butler,* Champaign,* Clark,* (Clermont), Clin-
ton, (Columbiana), Crawford,* Darke,* Delaware,* Fairfield,* Franklin,* Gallia,
Greene,* Hamilton,* Hancock, Huron, Jackson,* Logan,* Ducas,* Madison,* Marion,*
Mercer,* Miami,* Montgomery,* Morrow,* Pickaway,* (Pike), Preble,* Putnam,* (San-
dusky), Seneca,* Shelby,* Union,* Van Wert, Warren,* Wyandot.*

PENNSYLVANIA.—Adams,* Bedford,* Berks,* Blair,* Bucks,* Carbon,* Chester,* Clin-
ton, Columbia,* Cumberland,* Dauwphin,* Delaware,* Franklin,* Fulton,* Hunting-
don, *Juniata,* Lackawanna,* Lancaster,* Lebanon,* Lehigh,* Luzerne,* Lycoming,

54 THE PERIODICAL CICADA,

Mercer, Mifjjlin,* Monroe,* Montgomery,* Montour,* Northampton,* Perry,* Philadel-
phia,* Schuylkill,* Snyder,* Somerset,* Union,* York.*

TENNESSEE.—Benton, Bledsoe, Blount,* Bradley, Carroll, Carter, Claiborne, Cum-
berland, Dyer, Gibson, Grainger, Greene,* Hamblen,* Hamilton,* Hancock, Hawkins,*

(James(?)), Jefferson,* Johnson,* Knoxr,* Loudon,* McMinn,* Montgomery, Obion,
Polk,* Rhea, Roane, Robertson, (Scott), Sevier,* Smith,* (Sullivan), Washington,*
Weakley, White, Williamson.

Vermont.—( Rutland.)

Virenta.—Alerandria,* Augusta, (Carroll), Clarke,* Fairfax,* Fauquier, Frederick,*
Grayson, Lee,* Loudoun,* Orange, Prince William,* Roanoke, (Spottsylvania),
Warren,* Wise, Wythe.

West Virerta.—Barbour, Berkeley,* Boone, Cabell, Grant,* Greenbrier, Hamp-
shire,* Hardy,* Harrison,* Jefferson,* Lincoln, Logan, McDowell, Mason, Mineral,*
Mingo, Monroe, Morgan, Ohio, Pocahontas, Preston,* Putnam, Raleigh, Roane,
Tucker,* Wayne.

Wisconsin.—Dane, (Sauk).

=
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0)
o
v
v

Fig. 14.—Map showing distribution of Brood X1, 1920.
Broop XI—Septendecim—1920. (Fig. 14.)

This is a small brood limited, for the most part, to the valley of the
Connecticut River in the States of Massachusetts and Connecticut,
with one colony in the vicinity of Fall River separated from the main
swarm. It is Brood I of Walsh-Riley and Brood 9 of Fitch, who
reports it as having occurred in 1818 and 1835. It was recorded also
by Dr. Gideon B. Smith from 1767 to 1852, and the genuineness of
the brood was fully established in 1869. Like most small broods in
settled regions, it is being greatly reduced in numbers, and in 1903

BROOD XII—SEPTENDECIM—1921. 55

Mr. Britton reports? that he was not able to secure any records for
Connecticut, although special effort was made to do so through
correspondence. A personal examination of the area was, however,
not made by the entomologist, and a clipping from the Hartford
Courant of June 6 reports them present.

In this year (1903), however, the first record of the periodical
Cicada from Rhode Island was obtained, no brood having previously
been reported from this State. The late James M. Southwick,
curator of the Museum of Natural History, Roger Williams Park,
reported under date of May 23 that a living specimen of the Cicada
was brought to him that day taken near the southwest corner of
Tiogue Reservoir, about a mile north of the New London turnpike,
an unsettled region with plenty of woods. The specimen was secured
by Mr. C. E. Ford, of Providence, who reported that the cicadas were
making so much noise that he thought they must be frogs or toads
having a late spring concert. Mr. Ford says, on the authority of his
mother, that some were collected there thirty-four years before. This
is a very interesting as well as unexpected record.

The distribution by States and counties is as follows:

ConnecticuT.—Hartford.

Massacuusetrs.—Bristol, Franklin, Hampshire.

Ruope Istanp.—Providence.

Broop XII—Septendecum—1921. (Fig. 15.)

The records on which this very doubtful new brood was based are
given in Bulletin 18, new series, of this Bureau, pages 56, 57. The
oldest record is that of Dr. Gideon B. Smith, who in his manuscript
reports the Cicada as occurring in 1853 in Vinton County, Ohio, and
Jo Daviess County, Ill. Neither one of these localities was con-
firmed, either in 1870, 1887, or 1904. In the latter year the writer
made special effort to have records secured if possible, but without
result, Professor Forbes particularly making inquiries for Jo Daviess
County, Il.

The other two records published in Bulletin 18 for this brood are
as follows:

Mr. J. R. Burke, Milton, Cabell County, W. Va., writing under date of May 22, 1897,
says: “‘The Cicada is not due here until 1904; its last visit was in 1887.”

Mr. W.S. Herrick, Thurman, Allen County, Ind., writes under date of June 10, 1898,
that ‘‘We had the 17-year locust in 1887, if I remember correctly.’’ This is also a
doubtful record, and it is possible that he referred either to Brood X XII, occurring in
1885, or Brood V, occurring in 1888.

No report whatever was received from Mr. Burke. Mr. Herrick,
under date of September 1, 1904, reported that he went through the
neighborhood where the locusts appeared in 1887, and failed to see

@ Report Conn. Exp. Sta. 1903, Part III, p. 214.

56 THE PERIODICAL CICADA.

any evidence of the occurrence of the brood. He states, however,
that they were quite numerous when they appeared before. The
possibility here, however, is pretty strong that there is a mistake in
the date.

Some unimportant new records were obtained in 1904. Mr. 8S. D.
Nixon found living cicadas, May 28, on a horse-chestnut tree in Mount
Olivet Cemetery, Baltimore, Md. Mr. Robert A. Kemp reports that
while collecting Lepidoptera in the woods at Catoctin Mountain,
near Braddock, Md., his attention was arrested by the unmistakable
cry of Tibicen septendecim. We was unable to secure the specimen,
which was safely hidden in a dense grove of young chestnuts. He
says:

=
o
o
o
o
v
v

- Fia@. 15.—Map showing distribution of Brood XII, 1921.

I was loath to leave him inasmuch as he gave me a parting ‘‘ Pharaoh” when I left
him alone in his glory. I have heard during the past week in this same woods several
specimens, and have not yet given up hope of securing one.

Both of these records may relate to belated specimens belonging to
Brood X of 1902. Mr. C. H. Bobbit, of Baltimore, Md., reports
that he heard twenty or thirty in a little piece of woods, and one
captured specimen was seen by Doctor Howard.

The records of this brood therefore are as follows, all very doubtful
or unimportant:

Intrnots.—Jo Daviess County.
InpiaAna.—Allen County.

BROOD XIII—SEPTENDECIM—1922. BY,

MaryLanp.—Frederick County and Baltimore.
Oxntro.—Vinton County.
West Virainta.—Cabell County.

Broop XIII-—Septendecim—1922. (Fig. 16.)

This very compact brood, described by Fitch as Brood No. 6, by
Walsh-Riley as Brood III, and by Riley as Brood V, covers in large

part a prairie or sparsely wooded region extending over portions of

several States in the upper Mississippi Valley.
A detached brood was formerly known in Pennsylvania, but seems
not to have been seen in later years. A few individuals were reported

SWWER MONT

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o
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v

D

Fic. 16.—Map showing distribution of Brood XITI, 1922.

from two counties in Maryland in 1905, and two very doubtful records
(1888) have been found for Kentucky and Virginia. Mr. Hopkins
in his Bulletin 68 gives records indicating possible swarms in Putnam
and Lincoln counties, W. Va. None of these eastern records can have
other than chance time relation with the main area covered by this
brood.

As the periodical Cicada is limited to forested areas, the broods
occurring in prairie districts of northern Illinois and adjoining States
are necessarily much broken and scattered, and Brood XIII occurs,
therefore, for the most part in small colonies in woods bordering
streams. No special effort was made to get records in 1905, and this

58 THE PERIODICAL CICADA,

brood therefore rests practically on the data secured in earlier years.
Reports came from eleven counties in [linois—all, however, included
in the region designated below—and from several of the counties in
other States, where they were expected. The italicized counties are
confirmations of old records.

The distribution by States and counties follows:

Intrvors.—All northern counties from Mercer southeast to Peoria, to Logan, Shelby,
Edgar, including Lee, Dekalb, Dupage, Kane, McLean, Rock Island, ete.

Inp1ana.—Lake, Laporte, Porter.

Towa.—Allamakee, Benton, Blackhawk, Bremer, Buchanan, Cedar, Chickasaw,
Clayton, Clinton, Delaware, Dubuque, Fayette, Howard, Iowa, Jackson, Johnson,
Jones, Linn, Thetis. Mitchell(?), Muscatine, Scott, Tama, iene dee <(?).

Kenrucky.—Lincoln.

Marytanp.—Baltimore, Frederick.

Micuican.—Berrien, Bench Cass, Hillsdale, Oakland(?), St. Joseph, Wayne)

PENNSYLVANIA. pent aster.

VIRGINIA.—Lee.

West Virarnta.—Lincoln, Putnam.

Wisconstn.—Crawiord, Dane, Grant, Green, Iowa, Jefferson, Lafayette(?), Mil-
waukee, Richland, Rock, Sauk, Walworth, Waukesha.

Broop XI V—Septendecim—1923. (Fig. 17.)

This brood, so far as our records go, is the one which was first
observed by the early European colonists on this continent. Two
important areas occur in eastern Massachusetts, one about Plymouth
and the other covering Cape Cod. The Plymouth swarm of 1634,
the first after European settlement, was noted by the early Puritans
and is referred to in the two earliest published notices of this curious
insect. (See Bibliography.) One of these records gives the definite
date of 1633, but, as shown by the subsequent appearances of the
swarm, this date is probably an error for 1634. No published records
have been found of the later appearances prior to 1789, but definite
records have been made of each return since that year. An interest-
ing account of the last appearance (1906) of the Cicada in Plymouth
County is given in a report received from Martha W. Whitmore,
Chiltonville, Plymouth, Mass. The near-by Barnstable colony was
also most abundant last year (1906) all along Cape Cod. As reported
by Miss Grace Avery, of Washington, D. C., the ground along the
coast was covered with the dead bodies and the trees in the forests
were all fired and brown from the egg-laying of the females.

Prof. H. T. Fernald reports (letter September 26, 1906) the dis-
tribution in Plymouth and Barnstable counties as in the following
towns: Plymouth, Wareham, Bourne, Falmouth, Sandwich, Mash-
pee, Barnstable, Yarmouth, and Dennis, being most abundant in
the three first named.

This brood, like Brood VI, covers a very wide range, extending
from Massachusetts westward to Illinois, with important groups of

BROOD XIV—SEPTENDECIM—1923. 59

swarms extending from Pennsylvania southward into northern
Virginia and in the Lower Alleghenies, covering portions of North
Carolina, Tennessee, Georgia, etc., and in the Ohio Valley region,
covering especially southern Ohio, Indiana, central Kentucky, and
western West Virginia.

Brood XIV has been carefully studied, notably so on the occasion
of its appearance in 1906, when a great many new records were
obtained by this Bureau and by the entomologists of the several
States included within its range.

Important new records were secured and kindly submitted to this

=
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o
)
v
v

Fic. 17.—Map showing distribution of Brood XIV, 1923.

office by Messrs. Garman, Fernald, Felt, Sherman, Howser, Bentley,
and Ramsay, for, respectively, Kentucky, Massachusetts, New York,
North Carolina, Ohio, Tennessee, and West Virginia.

The occurrence jointly with this brood in 1906 of the small and
rather unimportant 13-year Brood XVIII ieaves some doubt as_ to
the correct assignment of certain swarms in southern Illinois, western
Kentucky, and Tennessee.

The starred counties indicate the occurrence of the Cicada in one
or more characteristic dense swarms; the italicized counties are con-
firmations of old records, and the counties inclosed in parentheses are
old records not reported in 1898. The large dots on the map (fig. 17)

60 THE PERIODICAL CICADA.

indicate thé starred counties and the small dots the unimportant or
doubtful occurrences.
The distribution, by States and counties, is as follows:

District oF CoLnumBIA.—Throughout.

Grorata.—(Gordon), (Habersham), (Rabun), (Towns), (Union), (White).

Intino1s.—(Boone), Grundy(?), Jo Daviess, Johnson(?), (Lake), (McHenry)
(McLean), (Putnam), (Stephenson), Vermilion, Whiteside,* (Winnebago), Wood-
ford(?).

Inprana.—Boone(?), Brown,* Carroll(?), (Clark), (Clay), (Crawford), (Daviess),
Dearborn, Dubois(?), Floyd, Fountain, (Gibson), Grant, (Greene), Harrison, Jack-
son(?), Johnson, Knox, (Lake), (Lawrence), Monroe, Morgan, Orange, Perry,* (Pike),
(Posey), Putnam, Ripley, Scott, Steuben(?), Sullivan, Tippecanoe, (Vanderburg),
(Vigo), Warrick, Washington, Wayne.

Kentuckxy.— Adair,* Allen,* Anderson,* Barren, Bath, (Bourbon), Boyd,* Boyle,*
(Breckinridge), Bullitt,* (Carter), Casey,* (Clark), (Clinton), Cumberland,* Edmon-
son,* Estill,* (Fayette), (Fleming), Floyd,* (Franklin), Garrard,* Grayson,* Green,
(Greenup), Hancock(?), Hardin,* Harrison, Hart, Henry, (Jackson), (Jefferson),
Jessamine,* Johnson,* Knott,* Knox,* Larue,* Laurel, Lawrence,* Lee,* (Lewis),
Lincoln,* Logan,* (McLean), (Madison), Magoffin,* Martin, Mason, Meade, Menifee,
Mercer, (Metcalfe), Monroe,* (Montgomery), Nelson,* Nicholas,* Owen, Owsley,*
Pendleton,* Perry, Pike,* (Powell), Pulaski,* Rockcastle,* Rowan,* (Russell), Scott,*
Shelby,* Simpson, Taylor,* Trimble,* Warren,* Wayne,* Whitley, Wolfe.

Marytanp.—Allegany,* Frederick,* Howard, Montgomery, Prince George, Wash-
ington.*

Massacuusetrs.—Barnstable,* Plymouth.*

New Jersey.—Bergen, (Burlington), (Cape May), (Gloucester), (Mercer).

New Yorx.—On Long Jsland* and in Richmond County.

Norru Carotiwa.—Buncombe,* Caldwell,* Caswell,* Haywood,* Jackson, McDow-
ell,* Madison,* Mitchell,* Watauga,* Wilkes,* Yancey.*

Onto.—Adams,* Auglaize, Brown,* Butler,* Clermont,* Clinton,* Columbiana,
Cuyahoga, Delaware, Fayette,* Gallia,* Greene,* Hamilton,* Highland,* J ackson, Law-
rence,* (Meigs), Pike,* Preble, Ross,* Scioto,* Vinton,* Warren,* Washington.

PrENNsytvyanra.—(Adams), Bedford,* Berks, Blair,* (Center), (Chester), Clearfield,*
Clinton,* (Columbia), Cumberland, Franklin,* (Huntingdon), (Lancaster), Lehigh,
Luzerne,* (Lycoming), (Mifflin), Montour,* Northuwmberland,* Potter, Snyder,*
Schuylkill,* Tioga, Union,* York.

TENNESSEE.—Anderson, Bledsoe,* Blount,* Campbell, Cannon,* Carter,* Cheat-
ham,* Claiborne,* Clay, Cocke,* Coffee,* Cumberland,* Dekalb,* Fentress,* Frank-
lin,* Grainger,* Greene,* Grundy,* Hancock,* Hawkins,* Johnson,* Macon,* Marion,*
Morgan,* Overton,* Pickett,* Putnam,* Rhea,* Roane, Robertson,* Scott,* Stewart,
Sullivan,* Unicoi,* Union, Warren,* White.*

Virornta.—(Alexandria), Augusta,* Buchanan,* Dickenson,* Fairfax, Frederick,*
Lee,* Nelson, Tazewell,* Wise.*

Wesr Virarni1a.—Berkeley,* Boone,* Brooke, Cabell,* Doddridge, Fayette,*
Greenbrier, Hampshire,* Hancock, Hardy, Jackson, Jefferson, Kanawha,* Lincoln,*
Logan,* McDowell,* Mason,* Mercer,* Mineral,* Mingo,* Monroe, Morgan,* Pendle-
ton, Pocahontas, Preston, Putnam,* Raleigh,* Ritchie, Roane, Wayne,* Webster,
Wood ,* Wyoming.*

Broop XV—Septendecim—1907. (Fig. 18.)

This is one of the new 17-year broods indicated by the writer in
Bulletin 18 (new series) of this Bureau, and consists of retarded
eastern colonies of Brood XIV. The important colonies of this brood

BROOD XV—SEPTENDECIM—1907. 61

are in Dutchess and Saratoga counties, N. Y. All the records from
Bulletin 18 are reproduced below, with the exception of the report
from Indian Territory, which falls distinctly within the 13-year lines,
and has been transferred to Brood X XIX.

This brood is represented by the colony appearing at Tivoli,
Dutchess County, and Galway, Saratoga County, N. Y., in June,
1890, as recorded by Prof. J. A. Lintner in his Seventh Report,
pages 297-301. Mr. Davis records the occurrence of scattering
individuals the same year on Staten Island. In a letter of June 2,
1890, Prof. J. B. Smith, New Brunswick, N. J., reports that the

=
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v
v

Fig. 18.—Map showing distribution of Brood XV, 1907.

periodical Cicada had been taken by several Newark (Essex County),
collectors, and had also been observed at Anglesea, Cape May
County.

Another record which perhaps applies to this brood is given by
Mr. I. N. Smith, Scotland Neck, Halifax County, N.C., in letter of
June 22, 1885. He reports that his ‘‘ first recollection of the locust
was about the year 1839 or 1840, when the whole of the white-oak
lands were filled with them. * * * In 1855 or 1856 they appeared
again, but nothing to compare with the period first stated. The
locusts were all on the white-oak land and on the Roanoke River and
not on the pine lands.’’ Assuming the dates 1839 and 1856 to be the
correct ones, this would throw this swarm of Cicadas into Brood XV.

62 THE PERIODICAL CICADA.

and if there are any representatives left they should reappear in 1907.
The distribution, by States and counties, is as follows:
New Jersey.—Cape May, Essex.
New Yorx.—-Dutchess, Richmond, Saratoga.

Norte Caronina.—Halifax.

Broop XVI—Septendectm—1908. (Fig. 19.)

This old Brood IX of Riley (VII of Walsh-Riley) is a very small
and doubtful one, and represents a few isolated colonies in the
extreme western portion of the range of the species, possibly two
years belated swarms of Brood XIV. It was reported as occur-

=
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v

Fic. 19.—Map showing distribution of Brood X V1, 1908.

ring in 1857 in southeastern Nebraska, and a very definite record
for Franklin County, Ark., which apparently pertains to this brood,
was obtained in 1885. Mr. J. M. Pettigrew, writing under date of
July 1, states that the cicadas were numerous in that county in May,
1857, and in 1874, doing some injury to small branches of fruit trees,
especially apple. This record falls in the western central part of the
State, and is surrounded by 13-year records, but is at an elevation
of a thousand feet or more and, in view of the definiteness of the
report, does not seem to be open to doubt. There is a doubtful
record reporting the Cicada in Lee County, Iowa, in 1874, which
seems to belong to this brood.

BROOD XVI—SEPTENDECIM—1908. 63

In Bulletin 14 and older publications a western outpost in Boulder
County, Colo., was reported for this brood. This record is undoubt-
edly erroneous, and arises from the confusion of one or other of the
mountain species of Cicada which also have life cycles of several
years and duplicate somewhat the habits of the eastern species.
Prof. C. P. Gillette, in answer to an inquiry by the writer, states
that he does not believe that septendecim occurs in Colorado, inasmuch
as he has not found a single example of it in the course of the insect
collecting done there by himself and students during the last sixteen
years, and he suspects that the insect reported is Tibicen rimosa Say,
which might readily have been mistaken for septendecim.

=
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a

Fig. 20.—Map showing distribution of Brood XVII, 1909.

The distribution, by States and counties, is as follows:
ARKANSAS.—Franklin.

lowa.—Lee (?).

NeBRASKA.—Richardson.

Broop XVII—Septendecim—1909. (Fig. 20.)

This brood is a precursor of Brood I, and was indicated by the
writer in Bulletin 18 (mew series) of this Bureau. It comprises
small or doubtful colonies only. The records given in that publica-
tion are reproduced below without change except for the addition of
two new localities for Virginia, one in Appomattox County and
the other in the southwestern part of Washington County.

64 THE PERIODICAL CICADA,

A very definite record which may coincide with this brood is
furnished by Mr. Theodore Pergande, of this Bureau, who states that
Mr. Rosseau, of Charlottesville, Albemarle County, Va., informed
him that the Cicada was very numerous in that place in 1875. His
informant was positive as to the year from its being the one in which
he made a trip to Europe.

Another record is given by Mr. John D. Macpherson, Manassas,
Prince William County, Va., in letter of July 3, 1895. He writes:

I came here on the 23d of June, leaving the Cicada-in full song in Washington
(Brood X). Finding none here I made inquiry and was informed that they appeared
in full force in this county (Prince William) in the year 1875. This information I
regard as reliable, the date being fixed as the year following the marriage and arrival
of my informant in this county. :

Mr. J. R. Honley, of Spanish Oaks, in a report received in 1898,
states that the ‘locusts’ appeared in Appomattox County in 1892, and
Mr. A. M. Connell, in a postal of May 29, 1902, reports their appear-
ance in the southwestern part of Washington County in 1841, 1858,
1875, and 1892. These Virginia swarms are evidently precursors
of Brood I, with which they are therefore closely allied.

A western extension of this brood seems to be indicated in the
record furnished by H. J. Giddings, Sabula, Jackson County, Iowa.
He writes, ‘‘during last June (1892) the periodical Cicada was quite
common here. * * * I thought it was unusual to find them in
such numbers four years after their regular appearance. The last
regular year was 1883.” (See Insect Life, Vol. V, p. 200.)

If belonging to the 17-year race, the two records following should
also be assigned to this brood. Mr. A. J. Julian, Woolleys Ford,
Hall County, Ga., reports under date of June 14, 1898, that the
Cicada was present there in 1892. Mr. J. W. Seaton, Strasburg,
Cass County, Mo., writes under date of June 9 that the Cicada last.
appeared in that county in the summer of 1892 and in the summer
of 1896, being numerous both years. The 1896 record refers to the
17-year Brood IV, and hence the record of 1892 is probably also of
the 17-year race occurring in the same district.

The scattering specimens recorded by Mr. Davis as occurring on
Staten Island in 1892 may also be assigned to this brood.

The distribution, by States and counties, is therefore as follows:

Grorcr1a.—Hall.

Iowa.—Jackson.

MissouriI.—Cass.

New Yorx.—Richmond.

Vireinta.—Albemarle, Appomattox, Prince William, Washington.

BROOD XVIII—TREDECIM—1919. 65

Broods of the Thirteen- Year Race.
Broop XVIII—Tredecim—1919. (Fig. 21.)

This is an unimportant brood, most of the records representing
scattering specimens rather than dense swarms. It was originally
established by Professor Riley as Brood XVI on the testimony
of Dr. G. B. Smith, who gives in his Register a record of its appear-
ance in Cherokee County, Ga, in 1828, 184i, and 1854. Its appear-
ance in the same locality was also recorded by Dr. J. G. Morris
in 1867, and this seems to be the most important swarm of the brood.
The records obtained since relate to scattering occurrences in three
other States.

ERMON?
ae
HAMPSH

7]
Welw
\,

a

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>

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ANG
L

A ENNSYWWAN! AG

oS AS }M!1SSOURI

Fic. 21.—Map showing distribution of Brood XVIII, 1919.

This brood immediately precedes in time of appearance the largest
13-year brood known, namely, Brood XIX. The latter brood occu-
pies the Mississippi Valley in the main, but with scattermg swarms
extending well over the Southern States and into Virginia, and thus
overlaps the territory covered by Brood XVIII, indicating very
plainly the origin of the latter as accelerated swarms of Brood XIX.

The localities for Brood XVIII as listed in Bulletin 14 are those
given by Professor Riley in 1894.7 None of them was verified in

a Annual Report, U. 8. Department of Agriculture, 1893, p. 204. (The records on
which localities for this brood are based are given in an editorial note in Vol. V, Insect
Life, pp. 298, 299.)

31117—No. 71—07——5

66 THE PERIODICAL CICADA,

1893, but an additional and very doubtful locality (Montgomery, Ala. )
was reported that year. The records obtained in 1906 added three
counties for Georgia, six for Tennessee, one for North Carolina, and
one for South Carolina, but gave again no confirmations of old
records. The lack of confirmations, however, does not invalidate
these old records nor necessarily mean the dying out of the swarms,
as no particular effort was made to get reports from the exact
localities.
The distribution, by States and counties, is as follows:

AnABAMA.—(Lowndes), (Montgomery) (?).
Grorcta.—(Cherokee), (Cobb), Gordon, Oglethorpe, Screven.

Fig. 22.—

ie

fap showing distribution of Brood XIX, 1907

NorrH Caroiina.—Anson, (Lincoln), (Moore).
SoutH CaroLina.—Edegefield.*
TENNESSEE.—Carroll, Dyer, Lauderdale, (Lincoln), McNairy, Madison, Stewart.

3RoopD XI X—Tredecim—1907. (Fig. 22.)

This is the largest of the 13-year broods, and also the best recorded,
perhaps, from the standpoint of distribution of all the broods. It is
Fitch’s Brood No. 3, in part, XIII of Walsh-Riley, and XVIII of
Riley. Its existence has been known since 1803. Its limits were
most carefully studied by Walsh and Riley in 1868, particularly for
the Missouri and Illinois localities. As has elsewhere been explained
(p. 31), there is a possibility that some of the northern counties, at

BROOD XIX—TREDECIM—1907. 67

least of Illinois and Missouri, listed for this brood belong to the
17-year Brood X, which appeared with Brood XIX in the year men-
tioned. Some additional data were obtained in 1881 and published
in Bulletin No. 8, old series, of the Division of Entomology, and the
records were brought down to 1894 in the- circular issued by Pro-
fessor Riley in that year. The later records, mostly in reply to the
circular just mentioned, but including a good many reports received
subsequent to the publication of Bulletin 14, considerably modify
and extend the range of the brood. The record for El Paso, Tex.,
is open to much doubt. The relationship of this brood to Brood
XXII has been elsewhere discussed.
Its reported limits are as follows:

ALABAMA.—Autauga, Blount, Bullock, Cherokee, Colbert, Cullman, Dallas Dekalb,
Elmore, Etowah, Franklin, Hale, Jackson, Jefferson, Lamar, Lauderdale, Lowndes,
Macon, Marshall, Mobile, Montgomery, Perry, Randolph, Russell, St. Clair.

ARKANSAS.—Baxter, Benton, Boone, Carroll, Clark, Clay, Conway, Crawford,
Dallas, Drew, Franklin, Fulton, Garland, Grant, Greene, Hempstead, Hot Spring,
Izard, Johnson, Lawrence, Logan, Lonoke, Madison, Marion, Newton, Prairie, Pulaski,
Randolph, Scott, Searcy, Sebastian, Sharp, Stone, Van Buren, Washington, White.

GrorGia.—Campbell, Catoosa, Chattooga, Cherokee, Floyd, Fulton, Harris, Pike,
Polk, Rabun, Richmond, Walker, White.

Inuinois.—Adams, Bond, Cass, Champaign, Christian, Clark, Clay, Clinton, Coles,
Crawford, Cumberland, Douglas, Edgar, Edwards, Effingham, Franklin, Gallatin,
Greene, Hamilton, Hancock, Hardin, Iroquois, Jasper, Jefferson, Jersey, Johnson,
Lawrence, Livingston, McLean, Macon, Macoupin, Madison, Marion, Massac, Monroe,
Montgomery, Morgan, Perry, Piatt, Pike, Pope, Randolph, Richland, St. Clair, Saline,
Sangamon, Scott, Shelby, Union, Vermilion, Wabash, Washington, Wayne, White,
Williamson.

Inp1an Trerrirory.—Choctaw, Creek.

Inprana.-—Vanderburg.

Iowa.—Lee.

Kentucky.—Carlisle, Graves, Hopkins, McCracken, Marshall, Trigg.

Loutstana.—Bossier, Caddo, Claiborne, Morehouse, Washington.

Mississrppr.—Attala, Choctaw, Clarke, Copiah, Franklin, Jasper, Lauderdale,
Leake, Madison, Monroe, Oktibbeha, Scott.

Missourt.—Audrain, Barry, Barton, Benton, Bollinger, Boone, Butler, Callaway,
Cedar, Chariton, Clark, Cole, Cooper, Dade, Dallas, Douglas, Franklin, Gasconade,
Greene, Henry, Howard, Iron, Jasper, Jefferson, Knox, Laclede, Lawrence, Lewis,
Linn, Livingston, McDonald, Macon, Madison, Marion, Moniteau, Monroe, Morgan,
Newton, Oregon, Pettis, Phelps, Pike, Polk, Pulaski, Ralls, Randolph, Ripley,
St. Charles, St. Clair, St. Francois, St. Louis, Saline, Schuyler, Shannon, Stoddard,
Stone, Warren, Washington, Webster, Wright.

Norra Caronina.—Cabarrus, Caldwell, Cherokee, Clay, Graham, Haywood, Iredell,
Macon, Madison, Mecklenburg, Swain, Wake, Wilkes.

OKLAHOMA TERRITORY.—Payne.

Sourn Carorina.—Aiken, Anderson, Chester, Greenville, Laurens, Oconee, Orange-
burg, Pickens, Spartanburg, Union, York.

TENNESSEE.— Bedford, Blount, Cocke, Davidson, Gibson, Giles, Greene, Hamblen,
Hamilton, Jefferson, Knox, Lawrence, Loudon, McMinn, Marion, Monroe, Mont-
gomery, Rutherford, Sevier, Stewart, Wayne, Williamson, Wilson.

Trexas.—FEl Paso (?).

VirerntA.—Brunswick, Halifax, Hanover, Prince George.

68 THE PERIODICAL CICADA.
Broop XX—Tredecom—1908. (Fig. 23.)

This is a small brood, founded on records given by Doctor Smith.
Some of the localities cited were confirmed and others negatived on
the recurrence of the brood in 1869, as reported by Professor Riley in
Bulletin No. 8, old series, of the Division of Entomology. Since that
date three doubtful localities have been added, one each for Virginia,
North Carolina, and Georgia, possibly based on 17-year broods which
appeared in conjunction with this brood.

ERMONY

v

x

'
u=
ty
4’
?

ye

fa)

{MISSOURI

Fic. 23.—Map showing distribution of Brood XX, 1908.

The distribution, by States and counties, is as follows:
GrorGIA.—Banks, Greene, Jasper, Muscogee, Walker, Washington.
NortH Carotina—Wilkes (?).

VirGinia.—Wise (?).

3roop XXI—Tredecim—1909. (Fig. 24.)

This is one of the broods representing the extreme southern range of
the Cicada, and was recorded by Doctor Smith in Florida as occurring
in 1844 and 1857. Its existence was confirmed in 1870, when records
were obtained indicating its extension also into Alabama, Mississippi,
and Tennessee.

It is a brood which, according to report, does not seem to occur in
dense swarms, but scatteringly, at. least in its more northern range.

BROOD XXII—TREDECIM—1910. 69

No records of its appearance which have come to our notice were made
in 1883 nor in 1896.
The distribution, by States and counties, is as follows:

ALABAMA.—Lauderdale, Mobile.
FiLoripa.—Gadsden, Jackson, Washington.
Misstssrepr.—Jackson, Tishomingo.
TENNESSEE.—Hardin.

5

goat
{
! ;

Fig. 24.—Map showing distribution of Brood X XI, 1909.
Broop XXII—Tredecim—1910. (Fig. 25.)

This 13-year brood, which appeared last in 1897, is of small extent,
but well established by many reliable records, the oldest of which dates
back to 1806. It is Brood IV of Walsh-Riley and VI of Riley.

A summary of the distribution of this brood was given by Mr.
Schwarz in Circular No. 22 of the Division of Entomology, issued in
May, 1897. This inquiry resulted in the report of but one additional
locality. The distribution and relationship of this brood is given by
Mr. Schwarz in the circular referred to, as follows:

It is confined to parts of southern Mississippi and adjacent parts of Louisiana east of
the Mississippi, the particular localities being given further on. Dr. D. L. Phares, of
Woodville, Miss., has taken particular pains to ascertain the extent of this brood, and
his lucid and concise account, already published in 1885, in Bulletin 8 (old series) of
this Division, is herewith reproduced:

“Their western limit is the Mississippi River, the southern about 8 miles north of
Baton Rouge, the eastern about 4 miles west of Greensburg, the county seat of Helena,

70 THE PERIODICAL CICADA.

and 4 miles west also of Liberty, in Amite County, Miss., thus extendirg from 15 to 50
miles from the Mississippi River, and from the vicinity of Baton Rouge, 108 miles to
the northern limit of Claiborne County, Miss., perhaps even farther. They therefore
occupy East and West Feliciana, the northern part of East Baton Rouge, the northwest
corner of Livingston and the western part of St. Helena parishes, La., and Wilkinson,
Adams, Jefferson, Claiborne, and parts of Amite, Franklin, and possibly parts of one or
two more counties in Mississippi.”’

The reports received since 1885 are mostly confirmatory of Doctor Phares’s state-
ment, but Mr. Thomas F’. Anderson, of St. Helena, La., writes us that the parishes, or at
least parts of the parishes, of Tangipahoa, Washington, and St. Tammany had to be
added to the range of this brood. His statement is quite definite; still a confirmation
of these new localities is desirable. :

3rood VI [XXII] is evidently a forerunner of the very large 13-year Brood VII
[XXIIT], which will appear in 1898 inthe Mississippi Valley. The geographical

Srey Ane |

=
o
)
oy)
a)
v
w

Fig. 25.—Map showing distribution of Brood XXTI, 1910.

range of Brood VII [XXIII] was mapped out in the Annual Report of this Depart-
ment for 1885, and it will be seen from this map that the southern limits of Brood
VII [XXIII] almost precisely coincide with the northern limits of our Brood VI
[XXIT.]

One new locality in central Louisiana, in Catahoula County, has been
added.

The brood occurs in the following States and counties:

Loutstana.—Parishes of East Baton Rouge, Catahoula, East Feliciana, Livingston,
St. Helena, St. Tammany (?), Tangipahoa (?), Washington (?), and West Feliciana.

Misstsstppr.—Counties of Adams, Amite, Claiborne, Franklin, Jefferson, and Wil-
kinson. ;

BROOD XXIII—TREDECIM—1911. 7a
Broop X XIII—Tredecim—1911. (Fig. 26.)

This is Brood No. 5 of Fitch, Brood V also of Walsh-Riley, and
Brood VII of Riley. There are two large 13-year broods, which
honor Brood XXIII divides with Brood XIX. As indicated by Mr.
Schwarz in Circular No. 30, both of these broods occupy the Missis-
sippi Valley from northern Missouri and southern Illinois to Louisiana ;
but while Brood XIX occurs also in many other localities, Brood
XXIII is confined more strictly to the Mississippi Valley region. At
the time of the recurrence of this brood in 1898 a very careful investi-
gation was undertaken by the writer of its distribution.

Several thousand replies were received in response to a circular

S

W HAMPSH

We

A
at

Ly

Fig. 26.—Map showing distribution of Brood XXIII, 1911.

sent out, many of which were negative, the investigation being
extended throughout all States in which there was any likelihood of
the appearance of the Cicada, and necessarily covering many counties
and districts where the Cicada was not suspected. Local investiga-
tions were also undertaken by the official entomologists in several
States, Professor Forbes adding four or five new counties for Illinois,
Professor Garman adding six counties for Kentucky not previously
reported, all in the eastern end of the State, and Professor Stedman
adding one new county from Missouri. In ali three of these reports
our own records were confirmed for nearly every county. The
results of this canvass up to June 20 were recorded in Bulletin No. 14

(Gu THE PERIODICAL CICADA,

(new series), Division of Entomology, in an appendix. A large
number of replies were received subsequent to that date, and the full
report, with corrected list of localities, was published in Bulletin 18
(new series), pages 61-63. The State and county records given
below are reproduced from this final report.

The reports for 1898 nearly all indicate the occurrence of the insect
in enormous numbers. Unfortunately, however, there enters again
with the records of this year some doubt as to the correct reference of
some of the localities in Illinois, Indiana, and perhaps northern Mis-
souri, or, in other words, where the territory occupied by the two
races overlaps on account of the scattering presence of Brood VI. In
most of the records assigned to Brood cu however, in the States
mentioned, the evidence points pretty Jpn to the accuracy of the
reference. Where there is uncertainty a query follows the county.

The counties marked with a star (*) indicate those in which the
Cicada occurred in one or more dense swarms, in many cases several
reports being received from the same county. In the unstarred
counties the Cicada was reported in few or scattering numbers, or at
least as not abundant. The counties in italics duplicate old records;
the counties lacking confirmation by the records of this year are
inclosed in parentheses and incorporated with the others.

The State and county records follow:

ALABAMA.—Etowah.

Arxansas.—Arkansas,* Ashley, Calhoun, Carroll, Chicot,* Clark,* Columbia, Craig-
head,* Crawford, Crittenden,* Cross,* Desha,* (Franklin), Fulton, Garland, Hot
Spring, Howard, (Izard), (Jackson), Jefferson,* Lafayette,* Lee,* Lincoln, Logan,
Lonoke,* Marion, Mississippi,* Monroe,* Newton, Phillips,* Pike, Poinsett,* Prairie,*
Pulaski, Randolph, St. Francis,* Saline,* (Searcy), Sebastian, Sharp, Union, Van
Buren, Washington, Woodruff.*

Groroia.—(Cobb, Coweta, Dekalb, Gwinnett, Meriwether, Newton.)

Intrnots.—Alexonder,* Crawford,* Edgar, Edwards,* Gallatin, Hardin,* Jackson,*
Jasper,* Jefferson, Johnson, Lawrence,* Macowpin, Madison,* Marion,* Perry,* Pike,
Pulaski,* Randolph, Richland, St. Clair, Scott, Union,* Wabash,* W ‘ashington, Wayne,*
White, Williamson.*

Inp1ANa.—Bartholomew, Daviess,* Fayette, Floyd, Gibson,* Jackson, Jennings,
Knox,* Montgomery, Owen, Posey,* Putnam, Ripley, Spencer, Sullivan,* Vander-
burg,* Vigo,* Warrick.*

Kentucky.—Ballard,* (Barren?), Butler, Caldwell, Calloway, Carlisle,* Christian,
Clinton, Crittenden, Daviess, Fulton,* Grant, Graves,* Green, Hancock, Hardin,
Hickman,* Hopkins, Livingston, Lyon, McCracken, McLean, Marshall, Muhlenberg,
Ohio, Todd, Trigg,* Union, Webster, Wolfe.*

Loutstana.—Bienville,* (Bossier), Caldwell,* Claiborne, Concordia,* East Carroll,*
East Feliciana, Franklin,* Madison,* Morehouse, Ouachita,* Pointe Coupee,* (Red
River), Richland,* St. Helena, Tangipahoa, Tensas,* (Washington), West Carroll.*

Mrsstsstppr.—Adams, Alcorn,* Amite,* Attala,* Benton,* Bolivar,* Calhoun,*
Carroll,* Claiborne, Coahoma,* Copiah,* De Soto,* Franklin, Grenada,* Hinds,*
Holmes,* pe Itawamba, (ape, Jefferson, Eee, * Lawrence, Leake,

@ None of these localities, all of which were queried, was Senaraear in 1898, aa the
record of this brood in Georgia is undoubtedly erroneous.

BROOD XXIV—TREDECIM—1912. 73

Lee,* Leflore,* Lincoln,* Lowndes, Madison,* Marion, Marshall,* Montgomery,*
Neshoba, Newton, Oktibbeha,* Panola,* Pike,* Pontotoc,* Prentiss,* Quitman,*
Rankin,* (Scott), Simpson, Smith, Tallahatchie,* Tate,* Tippah, (Tishomingo),
Tunica,* Union,* Warren,* Washington,* Webster,* Yalobusha,* Yazoo.*

Missourt.—Audrain,* Barry, Benton, Boone, Callaway, Camden, Cape Girardeau,*
Cedar, Christian, Clark (?), Clinton, Cole, Cooper, Dade, Dallas, Dent, Douglas, Gas-
conade, Greene, Hickory, Howell, Iron, Jefferson, Johnson, Knox, (Lawrence), Linn,
Maries,* Miller, Morgan, New Madrid,* Osage,* Ozark, Pemiscot,* Perry,* Pettis,
Phelps, Polk, Pulaski, Reynolds (?), St. Charles,* St. Clair, St. Francois, we Louis,
Scott,* Taney, Texas, Warren, Washington,* Webster.

Onto.—Hamilton.

TENNESSEE.—Benton,* Carroll,* Chester,* Crockett, (Davidson), Decatur,* Dickson,*
Dyer,* Fayette,* Gibson,* Hardeman,* Hardin,* Haywood, Henderson,* Henry,* Hum-
phreys,* Lake,* Lauderdale,* Lewis, McNairy,* Madison,* (Maury), Montgomery,
Obion,* Perry,* (Robertson), Rutherford, Shelby,* Stewart, Tipton,* Wayne,* Weak-
ley,* Williamson.

=
Oo
a)
w)
Oo
v
v

Fia. 27.—Map showing distribution of Brood XXIV, 1912.

Broop XXIV—Tredecim—1912. (Fig. 27.)

This is one of the new ¢tredecim broods indicated by the writer in
Bulletin 18 (new series) of this Bureau on the strength of the following
records:

Mr. P. Lynch, Commerce, Scott County, Mo., under date of Decem-.
ber 24, 1874, reports that the Cicada appeared in the summer of me
in enn ole numbers, coming in June and remaining about tw
months. “Their eastern limit in this county (Scott) was the Miseis-

74 THE PERIODICAL CICADA,

sippi River, but they were as numerous on the opposite side of the
river in Alexander County, IIl.”’

Mr. W. 8S. Campere, Pickens Station, Holmes County, Miss., writes
under date of February 27, 1875, that the cicadas appeared in great
numbers in April, 1873. These two records would indicate a brood
originating doubtless by retardation of individuals of Brood XXIII.

Subsequent to the publication of the records for this brood in Bul-
letin No. 18 (new series) of this Bureau two additional localities have
been reported—one in Louisiana and one in Mississippi. Mr. Ben H
Brodnax, of Brodnax, Morehouse County, La., reported in 1899 that
the locusts first appeared in small numbers on May 2 and lasted only
about ten days. On inquiry he found that they were heard scattered
about the south Arkansas line (Ashley County) and down to the
lower line of Morehouse Parish. No specimens were collected. This
report carries the record of this brood into the edge of Arkansas.

Mr. George H. Kent, Meadville, Franklin County, Miss., in a letter
of May 30, 1899, reports the appearance of a small brood in the western
part of Franklin County between the latter part of April and May 15.
Both of these records, as with the earlier ones, are probably from
belated swarms of Brood XXIII, but may represent the start of a
new brood.

The State and county distribution of the brood is as follows:

ARKANSAS.—Ashley.

Inuinot1s.—Alexander.

LovuistanA.—Morehouse.

Mississipp1.—Franklin, Holmes.

Missouri.—Scott.

Broop XX V—Tredecim—1913.

(No Cicada records of the 13-year race have been reported corre-
sponding with this brood number.)

Broop XXVI—Tredecim—1914. (Fig. 28.)

This brood, No. X of Riley, was originally based on a very doubtful
record given by Dr. Gideon B. Smith, to the effect that he was
informed that the insect appeared in vast numbers in parts of Texas
in 1849, but that he was unable to get any particulars. No confirma-
tion of the occurrence of this brood in Texas has since been gained,
and its existence is very doubtful. A more definite record was
secured, however, by Professor Riley in 1875, from Dr. D. L. Phares.
A gentleman reported to the latter that a swarm of cicadas was heard
on the 10th of June in West Feliciana Parish, La., near the river and
‘south of Bayou Sara. Some specimens were secured of this brood,
all dwarfs. No other record seems to have been secured of this brood
until the year 1901. In that year the occurrence of the brood in West
Feliciana Parish was confirmed, locusts being reported by Mr. John

BROOD XXIX—TREDECIM—1917. 75

¥. Griffin, of Wakefield, as occurring in small numbers throughout the
parish between the 5th and 25th of May. An outpost in Mississippi
is also reported by Mr. George H. Kent, Suffolk, Franklin County,
who reports their appearance throughout the southwestern portion
of the county in the month of May.

The records for this brood, therefore, are:

Loutsrana.—West Feliciana Parish.

Misstssrppr.—F ranklin County.

TEXAS.— (?).

|M!SSOURI

iddISSiISSitw

Fig. 28.—Map showing @istribution of Brood XXVI, 1914.

Broop XX VII— Tredecim—1915.

A small brood was reported for Franklin County, Miss., as appearing
about May 20, 1902, by Mr. George H. Kent, of Suffolk.
3RooD XXVITI— Tredecim—1916.
(No Cicada records of the 13-year race have been reported corre-
sponding with this brood number.)

Broop XXIX—Tredectm—1917. (Fig. 29.)

Two records which can be assigned to this brood number were
reported by the writer in Bulletin 18 (new series) of this Bureau.

Mr. C. J. Wellborn, Blairsville, Union County, Ga., writes under
date of June 12, 1885, that ‘in May, 1878, locusts appeared south

76 THE PERIODICAL CICADA.

of this place, and the northern limit then was the present southern
limit of the territory covered now (by Brood X, 1885).”

Mr. James Pagon, Winnsboro, Fairfield County, S. C., writes that
locusts appeared in South Carolina in 1878, but does not give definite
localities. Both these records need confirmation.

A record submitted by the late W. S. Robertson, of Muskogee,
Ind. T., in a letter*dated June 17, 1879, of the occurrence of a brood
of cicadas in 1839 at that point, was assigned, in the publication just
referred to, to Brood XV, under the supposition that it probably
belonged to the 17-year race. This record falls, however, in territory

\N

W HAMPS!

=
o
o
oO
e)
v
v

Fic. 29.—Map showing distribution of Brood X XIX, 1917.

which is distinctly 13-year, and would seem to indicate that it
belonged rather to Brood X XIX.
The distribution of this brood, by States and counties, is as follows:
GEORGIA.—Union.
InpDIAN TERRITORY.—Muskogee.
SourtH Caro.iina.—Fairfield.
s3ROOD XXX—Tredecim—1918. (Fig. 30.)

This brood was established by the writer on a single record given
in Bulletin 18 (mew series) of this Bureau. This record follows:

Mr. B. H. Brodnax, Brodnax, Morehouse Parish, La., writes under
date of May 13, 1892, that cicadas are scatteringly present, and in a

SYSTEMATIC POSITION AND STRUCTURAL DETAILS. 77

later letter he asserts that the insect in question is the periodical
Cicada with which he is familiar.

An addition to this record was received in 1898 in a postal from
Mr. J. W. Seaton, Strasburg, Cass County, Mo., who reports that
they appeared there in the summer of 1892, as they did also in 1896
(Brood XX1I), being numerous both years.

The State and county records are:

LovuIstANa.—Morehouse.
Missourtr.—Cass.

=
Oo
o
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v
v

tig. 20.—Map showing distribution of Brood XXX, 1918.

SYSTEMATIC POSITION AND STRUCTURAL DETAILS.

The periodical Cicada belongs to the Homoptera, one of the two
divisions of the Hemiptera, or great order of sucking insects, familiar
to the public mind under the name of “bugs,” and including, in addi-
tion to the graceful and attractive species like the Cicada, such foul-
smelling species as the plant bugs, squash bugs, and certain animal
parasites. The members of the suborder Homoptera, to which the
Cicada and its allies belong, are, however, distinctly removed from
the lower suborder of “bugs” just referred to, namely, the Heterop-
tera, and as a rule lack the disgusting odor and habits of the latter
and less esteemed suborder of sucking insects. The Homoptera as a
rule comprise clear-winged insects, which subsist on the juices of

78 THE PERIODICAL CICADA.

plants, and are active usually in flight and often beautiful in form
and color. The cicadas are not only the largest and most striking
insects of their suborder, some of the species measuring over 6 inches
in expanse of wings, but in the male sex are endowed with the power
of song, which last characteristic has invested them with great pop-
ular ‘interest in all ages; and especially in the poetry of nature are
they noteworthy, from the time of Homer to the present.

The old genus Cicada is represented by species in all parts of the
world, over five hundred distinct forms being already known, and
they are especially abundant in North America, nearly one hundred
species having been described from the continent and adjacent islands.
The more familiar of these insects to the popular mind are the com-
Be) mon dog-day cicadas, or
harvest flies,represented
by several species, the
most abundant of which
is, perhaps, Cicada tibi-
cen Li. (pruinosa Say).
The sleepy droning of
these annually appear-
ing species in July and
August is commonly
taken as a harbinger of
greater heat and is a
most familiar character-
istic of midsummer.

The periodical species
is much more slender
and graceful than the
majority of the annual
visitors, but structurally
Fic. 31.—Head and prothorax of Cicada, lateral view, showing is not ve dissimilar.

parts in normal position. For description, see fig. 33. lit is medium sized, for

Geutkons Huse ten) the most part black in
color, with orange-red eyes and limbs, and with the margin of principal
veins of the four nearly transparent wings similarly colored.

In discussing the structure of this insect particular attention will
be given only to the important organs, viz, those for taking food, or
the beak, the instrument for piercing plants and depositing eggs, or
the ovipositor, and the organ of song in the male insect.

A cursory examination of one of these insects from above reveals
its rather robust body, covered by two pairs of transparent parchment-
like elliptical wings, which rest roof-like over the abdomen; the short
transverse head, with great oval prominent eyes at the lateral angles,
the three minute ocelli arranged in a triangle on top, and the very

THE MOUTH PARTS, OR BEAK. 79

short, thread-like antenneze projecting between the compound eyes.
Viewed from beneath, the triangular prolongation of the head into
the three-jointed beak it to be noted; the legs, not especially large
or strong except for the anterior femora, which are much thickened;
in the female the complex instrument for the deposition of eggs pro-
jecting from a fissure or slit in the lower surface of the abdomen, and
the blunter abdomen of the male without the fissure beneath, but
with two large ventral plates at the base of the abdomen covering
the sounding disks of the vocal apparatus. The latter is located on
either side of the base of the abdomen and appears as two inflated
ribbed drums of lighter color than the general body surface.

Fig. 32.—Head of Cicaga, front view, showing the normal position of mouth parts on the left, and
with the mandible and maxilla drawn out on the right. For description, see fig. 33. (Author’s
illustration.)

The structure and workings of the more important organs, namely,
the beak, the ovipositor, and the vocal apparatus, follow in some
detail.

THE MOUTH PARTS, OR BEAK.

In the order of insects to which the periodical Cicada belongs
it is possible to trace all the essential parts, though vastly modified,
found in the mouth of true biting insects, namely, the upper lip
(labrum), the main pair of jaws (mandibles), the second, or lower,
pair of jaws (maxille), and, beneath, the lower lip (abium). Within
also are the two tongues, one projecting from the roof of the mouth

SO THE PERIODICAL CICADA.

(epipharynx) and the other (hypopharynx) attaching to the upper
base of the lower lip. These tongues are short and of service, prob-
ably, in facilitating the suction necessary in raising the fluids of the
plant to the mouth. They do not extend beyond the mouth cavity,
and never enter the plant tissues.

The upper lip is comparatively short, and serves its normal purpose
as a covering for the adjacent parts of the mouth. What correspond
to the short, powerful biting jaws of gnawing insects are in the Cicada
greatly elongated and thread-like, and brought together to form a sort
of piercing and sucking apparatus, which is inclosed in the greatly
elongated lower lip. The latter is three-jointed and deeply grooved

Fia. 33.—Head and prothorax of Cicada, lateral view, with parts separated to show structure: I, a,
clypeus, b and c, labrum, d, epipharynx; I’, same from beneath; IJ, mandible, a, base, b, sheath for
seta, c, mandibular seta, c’, muscular base of latter; III, maxilla with parts §-milarly lettered; IV,
labium, with three joints as follows, a, submentum, b, mentum, c, ligula; the hypopharynx is shown
at d, from side, d’, from above, and d’’, from beneath; V, prothorax. (Author’s illustration.)

above so as to be almost tubular, and acts as a support and sheath for
the piercing seta-like jaws, and also assists in conveying the liquids
from the point of.contact with the plant to the mouth cavity. The
long lower lip just described is the piercing beak in popular belief, yet
in point of fact it never enters the tissues of the plant, the puncture
being made solely by the fine, stiff, needle-like jaws or sete, which can
be projected at will by the insect with great force from the tip of the
beak. (See figs. 31, 32, 33, and 34,a.)

The feeding habits of the adult Cicada are discussed on pages 101—
102. The main feeding is, however, during the long adolescent period,
comprising the larval and pupal existence of the insect under the soil,

~

THE OVIPOSITOR.

81

when the taking of food is a constant feature. The structure of the
mouth parts in these preliminary stages is identical in essentials with

that of the adult, and the characteristic features are
illustrated in the foregoing figures with subjoined expla-
nations.

In the taking of food by the larve and pup, as they
rest on the rootlets in their earthen cells, the tip of the
beak, namely, the lower lip, is brought to bear on the
root, and by alternating longitudinal thrusts of the setz,
especially the upper pair, which are the stronger and
which represent the great jaws or mandibles of biting
insects, the soft, succulent layers of the cambium
beneath the bark are reached, the slender setz being
supported, strengthened, and directed by the stronger
and encircling sheath-like lower lip. The irritation
caused by this
puncture induces a
flow of sap, which
passes up between

or throat

by suction,
Fic. 35.—Abdomen of female Cicada showing ovi- ine 2
positor and attachments: a, ventral, 6, lateral, as In higher
and c, dorsal view. (Author’s illustration.) animals.

THE OVIPOSITOR.

The ovipositor (fig. 34,5; fig. 35), or twig-piere-
ing and egg-laying organ, of the female Cicada
is also a very complex instrument. It issues -
from a groove, or fissure, on the underside of the
abdomen, and at rest is nearly concealed except
at the tip by the broad overlapping sides of the
eighth dorsal segment. The ovipositor proper
is protected and covered when at rest by two
valves, which form a sort of sheath, or scabbard.

Fic. 34.—The pe-
riodical Cicada:
Side view of fe-
male to show
beak, a, and
ovipositor, Db.
(After Riley.)

the sete to the lower lip and
within this along the basal por-
tion of the sete into the mouth

Fig. 36.—Tip of ovipositor

The inclosed ovipositor is a very tough, horny of Cicada, much enlarged:

instrument, spear-shaped, and serrated at the

a, from above, b, from be-
neath,with dotted portion

extremity, and consists of three pieces, namely, to show the alternating

a back portion (probably two pieces grown to-
gether), which acts as a supporting or connect-

motion of the side pieces.
(Author’s illustration.)

ing piece for the two lateral blades. These lateral pieces, or blades,
slide up and down in alternation on tongues projecting from the cen-

31117—No. 71—07——6 ‘

82 THE PERIODICAL CICADA.

tral or supporting piece, have serrated cutting edges, and are the chief
agents in piercing twigs preparatory to the deposition of eggs (fig. 36).
The relative position of the three parts of the ovipositor and the
nature of the locking tongues, grooves, and clasps, which make one
tube of the whole, are illustrated in the accompanying cross sections
(e237):

The different pieces of the ovipositor attach to flat plates partly
concealed within and attaching to the wall of the abdomen, and are
operated by powerful muscles both in making incisions in the twigs
and in passing the eggs from the oviduct (which opens at the base.
of the ovipositor) through the tube formed by the three parts of the
instrument, unti! they reach their final
lodgment in the twig. The act of oviposi-
tion will be described in another place.

THE MUSICAL APPARATUS.

Perhaps the most interesting feature of
the anatomy of the Cicada to the popular
mind is the musical apparatus, by means
of which it makes its peculiar note, or song.
This apparatus and the sounds produced by
its possessor have been studied and de-
scribed by many naturalists, beginning with
the very earliest, and, in fact, the fullest and
most aceurate description of the method of
of producing sounds and the anatomical
structure of the vocal organ in these insects
F1G.37.—Cross section of ovipositor is the one given, early in the last century,

Byes, Yee cok by that famous French pioneer in the study
to showinterlocking tonguesand of the biology and anatomy of insects,
grooves. (Author’s illustration.) Réaumur.4

The work of Réaumur was confirmed and added to a hundred years
later by a most painstaking study of living specimens by another
French student, Solier,’ and for a minute technical description of the
anatomy and workings of the sound apparatus the reader is referred
to these authors.

The special modification and structure of these parts in our periodi-
cal species have been studied by the more important older writers,
as Potter and Smith, and more recently by W. J. Burnett¢ and E. G.
Love.4

aHistoire des Insects, Vol. V (1740), pp. 158-170, pl. 17.
bAnn. Soc. Ent. France, 1837, Vol. VI, pp. 199-217.
cProc. Bost. Soc. Nat. Hist., 1851, Vol. IV, p. 72.
dJourn. N. Y. Micros. Soc., 1895, XI, pp. 39-42.

THE MUSICAL APPARATUS. 83

As already noted, the gift of song is found in the male insect only,
and the true sound apparatus consists of two small ear-like or shell-
like inflated drums situated on the sides of the basal segment of the
abdomen. These drums are caused to vibrate by the action of pow-
erful muscles, and the sound is variously modified by adiacent
smaller disks—the so-called ‘‘mirrors”’ or sounding boards—and issues -
as the peculiar note of the species, which once heard is never likely to
be forgotten, or, if heard again, mistaken for that of some other insect.
The true sound organs are entirely exposed in the periodical Cicada
except for the covering afforded by the closed wings of the resting
insect. In other cicadas these drums are usually protected by over-
lapping valves or expansion of the body wall.

The sounding drum,
or ‘‘timbal,”’ as Réau-
mur termed it, of the
periodical Cicada is a
tense, dry, crisp mem-
brane numerously
ribbed or plated with
the convex surface
turned outward. The
ribs are chitinous thick-
enings or folds in the
surface of the parch-
ment -like drum, and
strengthen the drum
while perhaps render-
ing it at the same time

é Fig. 38.—The musical apparatus of the periodical Cicada: a, view
more elastic. The trom beneath, showing the plates (light colored) covering the

sound is produced by sounding disks; 6, dorsal view, the timbals showing as light-
: : : colored areas; c, section at base of abdomen, showing attach-
the rapid vibration, or ment of large muscles to timbals; d, timbal greatly enlarged,

undulation caused by in normal position; ¢, same drawn forcibly in by the action of
a a one of the muscles, asin singing. (Author’s illustration.)
the springing or snap-

ping in and out of these corrugated drums. Two powerful muscles
of very peculiar structure situated within the base of the abdomen
set these drums in motion, producing the rattling so-called song of
the Cicada, very much, as has been suggested, as sound is produced
by pressing up and down the bottom of a tin pan which is somewhat
bulged.

Beneath each “‘timbal”’ in the base of the abdomen of the insect
is a large sound or air chamber, and a third occurs in the thorax
joining the first two. These are closed by the body walls and mem-
branes, and the two abdominal ones beneath by the very peculiar
‘‘mirrors,”’ or ‘‘spectacles’’—-the tense, mica-like membranes situated
at the base of the abdomen and protected and covered by the semi-
circular rigid disks projecting from the thorax. These transparent

84. THE PERIODICAL CICADA,

membranes are often mistaken for the true sound organs, but they
are rather sounding boards, or drums, to increase and transmit the
sound vibrations induced by the play of the timbals. That they are
not essential to the production of sound can be shown by slitting
them or removing them altogether without there being any cessation
of the note. Much more important modifiers of sound are the semi-
circular disks projecting from the thorax over the ‘‘mirrors,’’ which,
if closed artificially or by the insect, deaden the sound very much,
or if opened or cut off, allow it to escape in greater volume. In sing-
ing, also, the insect modifies the song notes and their volume by rais-
ing and lowering the abdomen, thus opening and closing these disks,
and the act of singing is also accompanied by a sort of trembling of
the thorax. The position assumed by the male when singing is always
with the head upward. The abdomen is then elevated and apparently
inflated, and with the beginning of the sound is slowly brought down
against the limb, when the note ceases. After a rest of a few seconds
this operation is repeated. These abdominal movements vary in
different species of Cicada and determine in a measure the peculiar
notes of each.

THE SONG NOTES OF THE PERIODICAL CICADA.

The song of the different species of cicadas is very distinctive, and
one familiar with the music of these insects can as readily recognize
the particular species by its peculiar notes as one knows the different
birds or mammals by theirs. The general character of the notes of
the periodical species has been thus described by Dr. G. B. Smith:¢

The music or song produced by the myriads of these insects in a warm day from
about the 25th of May to the middle of June is wonderful. It is not deafening, as
many describe it; even at its height it does not_interrupt ordinary conversation. It
seems like an atmosphere of wild, monotonous sound, in which all other sounds float
with perfect distinctness. After a day or two this music becomes tiresome and dole-
ful, and to many very disagreeable. To me it was otherwise, and when I heard the
last note on the 25th of June the melancholy reflection occurred—shall I live to hear
it again?

As one approaches a colony of these insects a peculiar roar, not
unlike the noise of a factory or a distant reaper, falls on the ears, and
this becomes louder and more intense as one draws nearer, having at
times to one standing in the midst of a colony a peculiar all-pervading
and penetrating effect. The individual notes are somewhat obscured
under these circumstances, but in the lulls the characteristic sounds
strike the ear, and the peculiarity is never to be forgotten, especially
the mournful falling note at the conclusion of each effort. Nearly
all the principal writers on the Cicada, and notably Potter, Smith,
and Fitch, have attempted to analyze the song note of this insect,

4 Scientific American, March 22, 1851,

SONG NOTES. 85

but the most careful study made is that by Professor Riley, who
distinguishes three important notes as characteristic of different sea-
sons or conditions of the aerial life of the male insect.

The loudest and most characteristic note, and the one which is per-
haps most familiar to the popular mind, is the note described by
Fitch as ‘represented by the letters tsh-e-e-H-E-E-K-e-ou, uttered
continuously and prolonged to a quarter or half minute in leneth, the
middle note deafeningly shrill, loud, and piercing to the ear, and its
termination gradually lowered until the sound expires.” The length
of this note given by Fitch is probably the maximum term and is
unusual. Ordinarily it is much shorter, ranging from two or three
to five or ten or even twenty seconds. This note is the character-
istic one of the height of the season, when great numbers of males
are singing together, and is rarely made by solitary individuals or
when there are only a few together. Some instinct also seems to
prompt the singing in unison, and as it rises at such moments the
intensity and volume of sound has a startling and weird effect.

The second important note is what is ordinarily known as the * Pha-
y-r-r-aoh” note, and is made early in the season, or when the males
are few in number and recently emerged. The termination of this
note is notable even more than the last for its peculiar mournful
cadence and lowering of pitch, which is very characteristic. It lasts
but two or three seconds. It has been compared, rather fancifully,
I think, by Professor Riley to the whistling of a train passing thr ugh
a short tunnel, or, when made by several individuals, more accu-
rately to the croaking of certain frogs.

A third, but less important, note is the clicking or intermittent
chirping, consisting of from 15 to 30 short, quick sounds, sometimes
double, the whole lasting about five seconds, and resembling the
sharp clicking of the chimney swift or some of the field crickets.

When disturbed and at the moment of taking flight the insect is apt
to make a short’ery or sharp chirp.

Allof these notes are said to occur in the small cassinii form, but of
higher pitch and less volume, but the clicking note seems to be the
characteristic one of this variety.

The strength and clearness of all the notes vary with the weather
conditions. They are loudest when the air is dry and warm and
clear, or between the hours of 11 and 3 o'clock. On wet days, or
when the air is unusually moist, the sound is much diminished, and
heavy or continued rains stop it for the time altogether.

While it is almost universally true that the song of the Cicada is
never heard between sunset and sunrise, they will, on very rare occa-
sions, when disturbed, start up singing in concert in the middle of

@ Science, September 25, 1895.

San THE PERIODICAL CICADA.

the night. Prof. A. D. Hopkins noted an instance or two of this
kind in connection with the brood of cicadas appearing in West Vir-
ginia in 1897. He says:

I was fortunate enough to hear the starting of one of these concerts on a clear, moon-
light night in June. One male in an apple tree near the house suddenly called out
as if disturbed or frightened. His neighbors in the same tree were thus apparently
awakened. One started the familiar song note, which was at once taken up by num-
bers of other males, and, like the waves from a pebble dropped into still water, the
music rapidly spread until it reached the edge of the thick woods, where it was taken
up by thousands of singers, and the concert was in as full blast as it had been the
previous day. This continued a few minutes, until all had apparently taken part
and the song had reached its highest pitch, when it began to gradually subside, and
in a short time silence again prevailed.

THE SO-CALLED STING OF THE CICADA.

With every general outbreak of this insect are associated many
accounts in local papers of its stinging human beings, the sting often
resulting, it is stated, more or less seriously to the person stung. Such
accounts were especially abundant in the great Cicada year 1868, and
in every important Cicada year before and since similar reports have
been made. So great was the fear in 1868, as noted by Professor
Riley, that in some cases fruits were avoided as being stung and
poisoned, and even drinking water was sometimes under suspicion.

So far as investigation of the reports has been possible they have
proved to be either utterly without foundation or much exaggerated.
Referring again to Doctor Smith’s manuscript, it is seen that he spent
much labor in carefully investigating such accounts, and found in
every case that he followed up, where death had been reported as
caused by the “bite” or sting of the “locusts,” that the story was
entirely fabulous. In the cases of apparent stinging he suggests that
the sufferer had probably been stung by a wasp, as will be later
explained, and soundly argues on the susceptibility of some veople to
whom the slightest scratch becomes a source of danger.

Professor Potter, referring to the Cicada, says in this connection:
“Tt can not defend itself against an ant or a fly. We have handled
them, male and female, time after time. We have mutilated them,
but never could provoke them’ to resentment.”’

Professor Riley says that of the thousands which he has handled,
and the hundreds of other persons, including children, who have also
handled these insects, not a single bona fide case of stinging has, to
his knowledge, resulted.

That the periodical Cicada can pierce the flesh with its sucking beak,
or, more properly, the fine needle-like filaments contained in it, or
perhaps extremely rarely with the ovipositor in the case of the female,
is quite within the bounds of possibility, and some apparently well-
authenticated cases or reports by reliable observers bear out this view.

THE SO-CALLED STING. 87

There is not a particle of evidence, however, to show that such pene-
trating is attended with the injection of any poisonous fluid, and the
injurious consequences which follow them in rare cases are evidently
due to unusual sensitiveness on the part of the individual, as suggested
by Doctor Smith, or a bad condition of the blood, which would cause
any wound to be attended with serious consequences. In this con-
nection it is to be remembered that there are well-authenticated
instances of most serious, if not fatal, results following the bites of
such insects as the mosquito, and other biting flies, the result of the
bites of which are very trivial in common experience.

With all the reports of stings by the Cicada which have been made
it is not to be questioned that some of them have a basis in fact. As
suggested by Doctor Smith, and afterwards fully elaborated by
Doctor Walsh,’ many of these reports are undoubtedly cases of wrong
determination, and the stinging had probably no direct connection with
the Cicada. There are, for example, severa! large digger wasps which
provision their larval galleries with adult cicadas for the maintenance
of their young. One of the commonest of the digger wasps is the
species S phecius speciosus Dru., described later on under the heading of
the enemies of the Cicada (pp. 132-134). As first suggested by Doctor
Smith, and afterwards more fully shown by Doctor Walsh, it is not
unlikely that this or some allied wasp, flying with its rather heavy
burden, might strike against or alight on some human being, and upon
being brushed off would retaliate by stinging the offender and then
flying away, leaving the Cicada behind. In the absence of the wasp
the Cicada would very naturally be accused of the offense. The usual
prey of this wasp, which appears rather too late in the season to
account for all the cases of stinging reported, is the later appearing
annual cicadas.

The rare cases of stinging by the Cicada that have any basis in fact
may be accounted for, as already suggested, by a thrust either of the
ovipositor or the sucking beak.

From the structure of the ovipositor, as already described, it will at
once be perceived that there is nothing impossible in a wound being
made by this instrument. The objections to this suggestion are that
the ovipositor when not in use in placing eggs in twigs is concealed in
a sheath in the insect’s abdomen, and also that the piercing of a twig
or other substance by the ovipositor is a slow and laborious process,
and therefore would not account for the quick sting usually described.
In no ease has an egg been found im the flesh, and in fact it is improb-
able that an insect would be allowed to rest long enough on the flesh
to accomplish the insertion of an egg. Furthermore, tests were made
and reported by Doctor Walsh? and later by Professor Riley, showing

a American Entomologist, I, pp. 7, 8; September, 1868. b Loe. cit.

88 THE PERIODICAL CICADA,

the absurdity of the theory that the stinging in question is done by
the aid of this instrument, the female not being able to puncture the
soft, yielding flesh at all. In one test reported by Professor Riley,
Mr. William Muir, of St. Louis, removed a female from a tree while
she was in the act of ovipositing, and placed her on his finger.
Although she instinctively endeavored to continue her work, she was
not able to make the least impression on the soft, yielding flesh. A
second experiment was made by Mr. Peter A. Brown, of Philadelphia,
who himself made several punctures upon his hand with the ovipositor
without experiencing any more serious results than would have
followed pricking with a pin or other sharp instrument. In a third
experiment, Doctor Hartman, of Pennsylvania, introduced some
moisture from the ovipositor into an open wound and it caused no
inflammation whatever.

The ovipositor having been removed as the probable source of sting-
ing, the beak only remains, and it is unquestionably by means of this
instrument that practically all the so-called stings of the Cicada are
made. The structure of the beak has already been discussed, and it is
not at all improbable, though certainly a rare occurrence, that the
Cicada, when held or caught, may thrust out the slender sets and
puncture the skin. Many other hemipterous insegts are known to
‘“sting’’ in this way and to cause some severe momentary pain. The
sensitiveness of the individual is, however, in the case of the Cicada,
the sole criterion of injury. The authentic reports of Cicada stings
show some variations in the effects, but as a rule the result is much
less serious than the sting of a bee and not much more than the punc-
ture of a needle, the wound usually healing immediately.

TRANSFORMATION TO THE ADULT STAGE.

PERIOD OF EMERGENCE.

The date of the issuing of the cicadas from the ground after their
long concealment varies a little with the latitude, being later in the
North than in the South. In the accounts of this sect published by
Professor Riley and most other writers up to the present time it has
been stated that there is very little divergence in the time of issuing
between the northern and the southern broods, the latter half, or more
strictly the last week, of May being the normal period for the emerg-
ence of the insect throughout its range. That there may be, however,
a considerable difference in time, depending on elevation and tempera-
ture, in a given district and in the northern and southern parts of the
country, also determined undoubtedly by temperature, has been fully
established. The variation in the dates of appearance is illustrated
by the following records:

PERIOD OF EMERGENCE. 89

Doctor Phares, writing of the occurrence of Brood XXII in 1871,
states that a few males began to appear about the 20th of April, but
that the bulk of the brood did not emerge until the 7th and Sth of
May, when they came forth from the earth in vast numbers, continu-
ing to emerge in diminishing numbers until the 18th of May. It will
be remembered that this is the most southern of all the broods—lying
in the southwest corner of Mississippi and the adjoining parts of
Louisiana.

Mr. John Bartram, writing of the brood appearing in 1749, states
that in the neighborhood of Philadelphia an abundance of these
insects which had just escaped from their skins was observed on the
morning of May 10 and that they continued to issue in great numbers
for a week or more, beginning to sing on the 13th and to oviposit on
the 16th, and disappearing altogether by the Sth of June.

In the great brood year of 1868 Professor Riley noted that in the
vicinity of St. Louis “they commenced to issue on the 22d of May,
and by the 25th of the same month the woods resounded with the
rattling concourse of perfect insects.’”’ At Washington, D. C., in the
Cicada year 1885, scattered individuals appeared on May 23, and they
issued, perhaps, most abundantly on the night of the 27th. Those
emerging within the city were somewhat earlier in appearance than
was the case in the neighboring woods across the Potomac in Virginia,
probably for the same reason that the trees in the city put out their
foliage a little earlier than in the near-by woods.

Mr. Davis, writing of Brood II as it appeared in 1894 on Staten
Island, New York, says that as early as May 19 many cicadas had
emerged, the first individuals of the swarm being noted six or seven
days earlier.

Mr. A. W. Butler, writing of the brood appearing in 1885 in Frank-
lin County, Ind., says that while in a few localities individuals were
seen as early as May 28, in other places not distant they did not
emerge until June 4, and later.

Mr. Hopkins made a careful study of the dates of emergence in West
Virginia in 1897 in connection with Brood V, and found very consid-
erable variation in time of appearance both between the northern and
southern borders of the brood and between the lowest and highest ele-
vations within the area covered by the brood. For the former a differ-
ence of nearly two weeks was indicated by the records, and for the
latter a difference of nearly four weeks. This variation, he says,
appears to be due to the difference of climate between the northern
and southern sections and between low and high elevations, in the
former case amounting to 34 degrees and in the latter to over 10
degrees in average summer temperature. He deduces from his obser-
vations, as a general rule, that there is about three and one-half days
difference in the time of the first general appearance of the Cicada for

90 THE PERIODICAL CICADA,

each degree of difference in the average summer temperature, whether
it be due to latitude or elevation.¢

An interesting case of artificial acceleration in the appearance of
these insects is recorded by Professor Riley as follows: Dr. E. S. Hull,
of Alton, Ill., having placed some underground flues for forcing vege-
tables, the unnatural heat caused the cicadas to emerge by the 20th
of March and from this time on until May. Other instances of accel-
eration are given in the discussion of the subject of retardation or
acceleration in times of appearance as an explanation of the forma-
tion of the different broods. (See p. 24.)

Notwithstanding the difference in time of emergence in the above
citations, the fact nevertheless remains true of the great uniformity
evidenced in the time of emergence, namely, the last week in May, for
the great bulk of the territory covered by the different broods of the
Cicada, and this fact is one of the noteworthy features in the life his-
tory of the insect.

The males precede the females by several days and disappear earlier
in the summer, both by reason of being shorter lived and also on
account of their earlier appearance, so that it often happens that while
the woods are still filled with females actively engaged in ovipositing,
the males are altogether absent and their songs are unheard.

DURATION OF THE ADULT STAGE.

Under normal conditions the Cicada remains in evidence in the
woods five or six weeks, occasional individuals occurring later, but as
a rule their disappearance is almost as sudden as their appearance and
is complete in the first weeks in July. Mr. Butler, writing of the 1885
brood in Indiana, says that twenty-three days after the appearance of
the Cicada a perceptible decrease in numbers was observed, chiefly
from a disappearance of the males. On July 15, nine days after they
had disappeared from the river valley districts, they were still abun-
dant and active in more elevated situations. Mr. Davis, writing of
the brood of 1894 on Staten Island, says that by the third week in
June the cicadas commenced to die of old age, and yet the males were
still singing and the females were abundant in certain localities as
late as the 8th of July, while by the 15th of the same month all had
disappeared.

Mr. Hopkins found on the hills near Morgantown, W. Va., that the
dates of the Cicada appearance were about normal, the first adults
appearing on May 20, the first general appearance not coming, how-
ever, until the 24th. Cold weather mtervening, there was a subsid-
ence again until the 30th, when they emerged again in enormous

@ Bulletin 50, W. Va., Agric. Exp. Station, p. 17.

CICADA HUTS, OR CONES. 91

numbers. Oviposition began on the 13th of June, and by the 17th of
the month the leaves on the wounded twigs commenced to wither.
All had disappeared by the 4th of July.

METHOD OF EMERGENCE.

In escaping from the soil the pupa burrows directly upward, but not
alwaysin astraight line, and under normal conditions emerges directly,
leaving a small round hole about the size of a man’s little finger.
While it is generally true that they do not pierce the surface at all
until they are ripe for transformation, they seem to have a frequent
habit of penetrating nearly to the top of the ground some time before
they actually issue and remaining usually within their burrows or
sometimes emerging, but concealing themselves under logs, stones,
etc., awaiting the proper moment to come forth. Usually throughout
the month of April they are to be found thus near the surface, as has
been recorded by many observers.

On the authority of Professor Potter the 10th of April is usually
the date for their appearance near the top of the ground. Here they
are often discovered by hogs and eaten with avidity, their holes com-
ing within a quarter of an inch of the surface and penetrating down-
ward from 6 to 12 inches.

CICADA HUTS, OR CONES.

Under special or peculiar circumstances, not always easily explain-
able, the Cicada pupx construct little cones, or chimneys, of earth
above the surface of the soil, continuing and capping their holes, sev-
eral weeks before the time of issuing. In addition to the names
Cicada “huts” or ‘‘cones,”’ these curious structures have been variously
termed ‘“‘towers,” ‘‘roofs,”’ ‘‘chimneys,”’ ‘‘turrets,’ and ‘‘adobe
dwellings.” .

The earliest reference to them, if the writer mistakes not the sig-
nificance of the language, and one which has hitherto been overlooked,
is by Professor Potter.* He refers to the ‘‘roofs of their tenements”
as being ‘‘neatly arched and so firmly cemented that water is never
found in them, although all of the surrounding grounds are over-
flowed and eee saturated,” and, stating that ‘‘the locust is not
singular in this provision,” he refers, in the same connection, to the
crayfish and other shellfish and some insects as building Pptiged alone
water courses, where the soil is wet, resembling ‘‘small chimneys,’
as a provision against ‘“‘inundation fod drowning.

The first definite account of the Cicada huts we owe to Mr. S. 5.
Rathvon, of Lancaster, Pa., who described them as occurring in
localities where the drainage was imperfect. He says

>)

a Notes on the Locusta, etc., pp. 17, 18 (1839).

eA

Q? THE PERIODICAL CICADA,

We had a series of heavy rains here about the time of their first appearance, and
in such places and under such circumstances the pupze would continue their galleries
from 4 to 6 inches above the ground, leaving an orifice of egress even with the sur-
face. In the upper end of these chambers the pupze would be found waiting their
approaching time of change. They would then back down below the level of the
earth (as at d, fig. 39) and, issuing forth from the orifice, would attach themselves to
the first object at hand and undergo their transformations in the usual manner.

Professor Riley had the accompanying figure (fig. 39) made from
one of the chambers furnished by Mr. Rathvon. This chamber meas-
ured about 4 inches in length, with a diameter on the inside of five-
eighths inch and on the. outside of 14 inches.

As will be later noted, the exit hole at the base of the turret in this
instance was probably abnormal, the insect issuing, as shown by later
observers, almost invariably from a hole clawed through the summit
of the cone.

The next instance of the occurrence of these cones of which we have
a record is a rather remarkable
one, and is given by Prof. J. S.
Newberry.“ These cones ap-
peared in May and June, 1877,
in a shallow cellar of a house
which had been erected on the
site of an old orchard at Rah-
way, N.J. The cellar had been
dug to the depth of about a
foot, and had been closed until
about the time of the emergence
of the cicadas, when it was
opened and the bottom was
Fic. 39.—Pupal galleries of the Cicada: a, front view; found to be thickly beset with

¢, ofifice; b, section; c, pupa awaiting time of mud cones or tubes, from 6 to 8

change;d, papa ready to transform. «(Aite: Fuley-) “inehes high.) lhe sesame re
for these curious structures suggested by Professor Newberry is that
the cicadas, finding a dark chamber, were apparently attempting to
work up to daylight. What is probably the true explanation of
their occurrence will be given later. An excellent photograph of one
of these structures, which considerably exceeds 6 inches in length,
accompanies Professor Newberry’s paper.

The references cited include all the records of the occurrence of
these cones up to 1894. In that year, however, these structures were
noticed in many localities in New York and New Jersey on the appear-
ance of Brood II, and excellent opportunities were afforded for their
study, advantage of which was taken by several competent obsery-
ers who were so situated that careful examinations could be made.

4 School of Mines Quarterly, vol. 7, January, 1886, 2 pp.

PLATE II.

ure,

a
rg

Bul. 71, Bureau of Entomology, U. S. Dept. of Agricul

PHOTOGRAPH OF CHAMBERS OF THE PERIODICAL CICADA, GENERAL VIEW

TAKEN AT NEW BALTIMORE

NL Veo

MAY

1894

(AFTER LINTNER

)

weak ah ehh
rin PES aes ALT’

CICADA HUTS, OR CONES. 93

The results of these investigations have cleared up much of the
obscurity which has hitherto surrounded these elevated burrows.

The first person to note these structures in 1894 was Mr. William
T. Davis, who reported their occurrence in April on Staten Island,
New York, stating that the pupx had been found on the 8th of that
month under boards on the edge of a meadow, where they had been
erecting cones of earth above the damp ground. In a later article
he says:

On the 22d of April many pup were found in the woods along Willow Brook under
stones, logs, and the chips about stumps of trees cut down in winter. Many more
were without protection of this kind, and their presence was indicated by the small
irregular cones of earth among the dead leaves. A heavy footfall near the cone was
sufficient to cause the insects to retreat, but if they were approached silently and
suddenly knocked over their constructors would be found within.

Some of the cones were 3 inches high, but they did not average
more than 2 inches. The experience of Mr. Davis corroborates the
theories of Professof Potter and Mr. Rathvon that the cicada cones,
occurring in moist situations, are designed to lift the insect above
such undesirable conditions.

Early in the spring of 1894 the attention of Doctor Lintner, the
New York State entomologist, was called by correspondents to the
occurrence of these cones and an investigation of the subject was
undertaken. A preliminary report was published in 1895,¢ but his
final report was not published until May, 1897.° In describing the
phenomenon in his Tenth Report, he says that the cones frequently
occurred in many thousands and occasionally hundreds of thousands
together, in some cases being intermingled with the ordinary open
burrows. At New Baltimore, N. Y., 16 miles south of Albany, as
early as the last week in April the pupz had brought up, apparently
from a considerable depth, masses of soft clay-like material and
molded it above the ground into conical and cylindrical structures
for their temporary occupancy. In places the ground was almost
covered with them, as many as twenty-five being counted to the
square foot. The cones inclined at a considerable angle from the
perpendicular and measured from 2 to 34 inches in height, and
the chamber within was uniform in diameter with the hole in the
ground. In emerging the pupa made a round opening in the upper
part of the chamber for its escape.

In the Twelfth Report cited, a long list of localities in New York is
given where the cones were found in 1894, together with notes on the
character of the chambers and accompanying conditions of the soil,
and also on the method of their construction. One of the pilates
illustrating this report is reproduced in this bulletin (see Pl. ID. It

@Tenth Report Insects New York, pp. 420-423.
6 Twelfth Report Insects New York, pp. 279-286,

94 THE PERIODICAL CICADA.

is a reproduction of a photograph of a small area of a cone-covered
district.

Two very elaborate accounts of these structures, by Mr. Benjamin
Lander and Dr. E. G. Love, were published in 1894-95, the authors
seeming very near the actual truth in their explanation of the
phenomenon. Mr. Lander describes the occurrence of the cones as
noted by him as follows:

On the 4th of May, 1894, while in the woods on the summit of South Mountain, at
Nyack, N. Y., I came upon a spot that had recently been burnt over. On this area I
observed vast quantities of the Cicada structures, entirely closed, averaging about 24
inches in height, the aggregation ending at the very edge of the burnt section. So
thickly studded was the ground thay often eight or ten would be found in the space
of a square foot; in one case I counted twenty-three in such a space. Subsequent
explorations showed that the Cicada city extended over an area of not less than 60
acres. Eight large aggregations were discovered by me on top of the Nyack hills and
the Palisades, covering many acres, and one near a stone quarry at a lower elevation—
none of them in a place subject to overflow. Later, when only the ruins of the domes
remained, I visited two areas where large numbers had been found, one in ground
thinly covering massive sandstone and another hard by a quarry, where the top soil
was thin.

The explanation offered by Mr. Lander is that the dome builders,
owing to the shallowness of the soil, determined either by the nearness
of the underlying rocks’ or of a subsoil of a character preventing the
insects working in it effectively, had responded more quickly to the
heat of spring and early summer, and the pupe coming prematurely
to the surface closed and extended their burrows as a means of pro-
tection while awaiting maturity. The extension of the gallery ehove
the ground, though not suggested by Mr. Lander, may be explained
by the same instinct which impels the insect to burrow upward from
its subterranean cell,

In substantiation of his theory, Mr. Lander calls attention to the
weather records for March and April, 1894, which indicate an unusually
high temperature throughout the region of the domed burrows,
causing wild plants to bloom a month before their ordinary season.
The occurrence of these structures over burned areas, which would be
acted upon more quickly by the sun, supports his belief. Additional
support of the same kind is an instance recorded by Prof. J. B.
Smith in a letter received from Mr. J. H. Willets, of Port Elizabeth,
N. J. The latter states that ‘On April 24 a fire from the South
Jersey Railroad burned over several hundred acres of woodland,
leaving the earth bare. Six days afterwards these fresh holes and
raised tubes appeared, and yesterday the whole surface was literally
covered with them.”’ In further description he says:

Imagine yourself standing out in the woods in south Jersey on 100 acres of recently
burned ground with millions and millions of raised tubes of new earth (clay ground)

aAnnual Report State Entomologist of New Jersey for 1894,

‘ZOBL “DO "G ‘NOLONIHSVAMA ‘GSHSGOOM NI VGVOID IVOIGOIYSd SHL 4O SYUSEWVHO 4O HdVYDOLOHd

(IVNIDIYO)

Bul. 71, Bureau of Entomology, U. S. Dept. of Agriculture.

PLATE III.

CICADA HUTS, OR CONES. 95

raised above the surface from 2 to 4 inchesrand from 14 to 2 inches in diameter, sealed
at the top, with a hole inside extending down in the earth 12 inches at least, * * *
and you will see mentally what I saw yesterday physically.

In this instance also, on the authority of Mr. Lander, the turrets
ended abruptiy at the edge of the burned area. The other instances
of these structures cited by Mr. Lander also bear out his theory. As
a rule, they were located on rocky cliffs with uniformly shallow soil or
in other situations where the soil in which the Cicada could work was
shallow. In the midst of one of the largest colonies a deep gully
occurred, 300 or 400 feet wide, in which the soil was a deep loam.
Here there were no domed burrows, although the hills on either side
were covered with them, and yet at the proper season the cicadas
appeared in the ordinary way in this gully in almost incredible num-
bers, leaving their customary small holes of exit even with the surface.

Some confirmatory records were obtained by Mr. Lander in 1898.4

The occurrence of these cones, as described by Professor Newberry
(p. 92), is confirmatory of this theory, a shallow covering of soil over
pupe of a few inches only being left by the slight excavation made.
A sunilar instance occurred in the District of Columbia in connection
with Brood X in 1902, and represented the only occurrence of these
structures observed that year in this vicinity. Mr. William Tindall,
living on Washington Heights, at the northwest section of the city,
discovered some of these curious structures in his woodshed, and an
investigation of the premises developed the fact that this woodshed
was studded with Cicada cones of perfect construction, varying from
1 inch to 6 inches in height. Evidently a tree had stood about
where the woodshed was built, and the cicadas had undergone their
development successfully in the ground beneath. All of those coming
to the surface outside of the shed escaped through simple holes with-,
out any structures above ground, but every individual which came
up within the shed built a turret or cone. The ground floor of the
shed was somewhat moist, rain running under, but it was rather drier
than the ground outside, so that the cones could not have been built
on account of the moisture. There had perhaps been a slight removal
of surface soil in this shed, bringing the cicadas nearer to the surface
and thus leading them to extend their galleries. Plate III is from a
photograph taken of the cones as they appeared in the shed, and
Plate IV illustrates half a dozen of these cones, nearly natural size,
two of which have been cut away to show the interior character of
the gallery.

Dr. E. G. Love, who also studied the problem of the Cicada huts
very carefully, agrees in the main with Mr. Lander, but differs some-
what in his explanation. As to the conditions of their occurrence, he
writes as follows:

@ Journal of the N. Y. Ent. Soc., Vol. VII, September, 1899, pp. 212-214,

96 THE PERIODICAL CICADA,

They are found in both wet and dry places; on the low and on the high ground;
singly and in colonies of many thousands. One hut, even in a damp soil, may be
surrounded by a dozen holes, from which the insects emerge without making any
huts, and often where we may expect to find them they are never seen.

Accepting the theory proposed by Mr. Lander for the condition
‘found to exist in the Nyack region, Doctor Love does not deem it
entirely adequate, as he says:

The huts are sometimes found in places in which the soil is of great depth and which
are not especially exposed. Such was the case at Baychester, where only a few huts
were found, and these in deep soil and so well protected that it was only after careful
search that they were discovered.

He offers the supplementary explanation that since it is hardly
possible that the Cicada larvee can determine instinctively the dis-
tance to be traveled in their upward journey nor the time required
to accomplish it, which will vary with the nature of the soil to be
tunneled and the directness of the line followed in their excavations,
it may often happen that individuals reach the surface before they
are prepared to assume the adult condition, and the number so doing
would be greater when the conditions all united to favor a short
passage. In protected localities where the soil is deep the larve lying
near the surface will be more: likely to emerge before their pupal
changes are complete, and would thus be led to the construction of
these cones. This, he says, would also explain their seeking tem-
porary shelter, as they do, under logs and stones, as has been pre-
viously noted.

The explanation offered for the construction of the Cicada cones by
Mr. Lander, as supplemented by Doctor Love, seems, on the whole,
satisfactory and adequate, so far as the conditions studied by these
writers are concerned. The conditions as described by Mr. Rathvon
do not inform us as to the nature of the soil, but both in the Rathvon
case and the later instance described by Mr. Davis, the wet character
of the ground would seem to indicate a soil of a considerable depth.
This would seem to give a basis of reason for the explanation suggested
by Mr. Rathvon and accepted by Professor Riley. A complete
hypothesis, therefore, seems to be in a union of the explanations
offered, namely, that the cone-building habit is induced either by a
shallow soil, proximity of the pup to the surface, or conditions of
unusual warmth which brings the pups to the surface in advance
of their normal time, and more rarely to unfavorable conditions of
excessive moisture. The mud caps are to protect the burrow from
cold until the time of issuing arrives.

The explanation of the occurrence of these structures on high
eround suggested by Professor Riley is certainly untenable. He
surmised that the individuals constructing cones in such situations
did so because impelled by habit that had become fixed and hered-

PLATE IV.

Bul. 71, Bureau of Entomology, U. S. Dept. of Agriculture.

PHOTOGRAPH OF CHAMBERS OF THE PERIODICAL CICADA, NATURAL SIZE,

IN WOODSHED, WASHINGTON,

D. C., 1902.

(ORIGINAL. )

Ae Ni

CICADA HUTS, OR CONES. Q7

itary in the course of a long period of existence in low wet situations.
The strict limitation of these cones to areas presenting peculiar
conditions thoroughly disproved this theory.

Some notes on the character of the huts may be appended. The
fact that there is no exit orifice at the ground, as described by Mr
Rathvon, is confirmed by the studies made by the observers cited
above, the insect invariably clawing its way out at the top. Mr.
Lander notes one instance where the pupal shell remained attached
and stuck in the summit of the burrow, the mature insect having
escaped. According to Mr. Lander, also, the huts are probably con-
structed at night, the insect taking advantage of the moist air, which
would prevent the too rapid drying of the earth used in making the
little tower and also of the delicate soft insect itself. The chambers
are constructed with soft pellets of clay or mud brought up from below
and pressed firmly into place. On examination it will be seen that
they are well rounded and rather firmly compacted within, although
the marks of the claws of the pupe are usually visible and leaves and
sticks are often incorporated in the walls. No one has actually
observed the insects while at work on these structures, and, although
Mr. Lander repeatedly broke off a number of cones to see if they would
be repaired, the insect failed to do so while being watched. Subse-
quently the broken portions were found to be recapped, but at some
ttle distance below the broken edge. In this connection may be
quoted the observation of Mr. Lawton, of Nyack, cited by Doctor
Lintner. Mr. Lawton found that in every case except one the pupa
repaired the cone soon after the injury by bringing up pellets of mud
and roofing over the broken portion about half an inch from the top.
The repairs were begun on one side and gradually extended over the
opening horizontally, there being no attempt to form a dome-shaped
roof. In some instances the repairing of the chamber began within
a quarter to half an hour after injury had been caused, and within
three or four hours the opening was entirely closed over. On one
occasion a pupa was caught with a pellet of mud in its claws.

The fact that these cones had been noted on only two or three occa-
sions prior to 1884 led to the belief that they were very rare and
abnormal. Their extraordinary abundance in 1884 in connection with
Brood II would seem to indicate that they are by no means as rare as
heretofore supposed, and it may be inferred that the absence of records
is simply due to the lack of examination, especially in localities where
the conditions would be favorable for their appearance. This view is
confirmed by the announcement ina recent letter from Mr. Davis of
the discovery of a cone April 30, 1898, on Staten Island belonging to
Brood VI, which appeared that year. He says that the cone was just
appearing above the dead leaves, which, with the ground also, were

31117—No, 71—07——7

98 THE PERIODICAL CICADA.

“soaked after the wet days just past.’”’ This belief is participated in
by Doctor Lintner in his last report on this interesting subject.* It
should not be forgotten, however, that the great mass of the insects
emerge without making any superficial construction whatever.

THE ACT OF TRANSFORMATION.

The phenomenon connected with the transformation of the period-
ical Cicada from the pupal to the adult stage is a very interesting one
and always fills the observer with considerable wonderment. As
remarked by Mr. Butler, when these insects emerge from the ground
it is usually with a rush, and a lively scramble ensues for each eleva-
tion near the point of their emergence. Trees, bushes, weeds, poles,
stumps, fences—in fact, everything upon which they can get above the
level of their recent homes is ascended. ‘The instinct which has
caused them to burrow to the surface of the ground still drives them
in the same direction upward, and they seem to make up in activity
for their long subterranean periods and their weeks of waiting near the
surface when the time has finally arrived for their emergence. The
different steps undergone by the insects in transforming from the
pupal to the adult stage have been perhaps most accurately described
by Professor Riley, as given below.? The plate accompanying his
description is reproduced in this bulletin as a frontispiece.

The unanimity with which all those which rise within a certain radius of a given tree
crawl in a bee line to the trunk of that tree is nost interesting. To witness these pupze
in such vast numbers that one can not step on the ground without crushing several
swarming out of their subterranean holes and scrambling over the ground, all converg-
ing to the one central point, and then in a steady stream clambering up the trunk and
diverging again on the branches, is an experience not readily forgotten and affording
good food for speculation on the nature of instinct. The phenomenon is most satis-
factorily witnessed where there is a solitary or isolated tree.

The pupe (frontispiece, figs. 1 and 2) begin to rise as soon as the sun is hidden behind
the horizon, and they continue until by 9 o’clock the bulk of them have risen. A few
stragglers continue until midnight. They instinctively crawl along the horizontal
branches after they have ascended the trunk, and fasten themselves in any position,
but preferably in a horizontal position on the leaves and twigs of the lowermost
branches. In about an hour after rising and settling, the skin splits down the middle
of the thorax from the base of the clypeus to the base of the metanotum (frontispiece,
fig. 3), and the forming Cicada begins to issue. be

The colors of the forming Cicada area creamy white, with the exception of the reddish
eyes, the two strongly contrasting black patches on the prothorax, a black dash on each
of the coxze and sometimes on the front femora, and an orange tinge at the base of
wings.

There are five marked positions or phases in this act of evolving from the pupa shell,
viz, the straight or extended, the hanging or head downward, the clinging or head
upward, the flat winged, and, finally, the roof winged. In about three minutes after
the shell splits the forming imago extends from the rent almost on the same plane with

a Twelfth Report Insects New York, p. 283.
b Annual Rept. Dept. of Agriculture, 1885, pp. 237, 238.

Bul. 71, Bureau of Entomology. U. S. Dept. of Agriculture. PLATE V.

EmprTy PUPAL SHELLS OF THE PERIODICAL CICADA CLINGING TO LEAVES, BROOD X,
1902, WASHINGTON, D. C. REDUCED. (ORIGINAL.)

NUMBERS AND LOCAL DISTRIBUTION. G9

the pupa, with all its members straight and still held by their tips within the exuvium
(frontispiece, fig. 4). The imago then gradually bends backward and the members
are loosened and separated. With the tip of the abdomen held within the exuvium,
the rest of the body hangs extended at right angles from it, and remains in this position
from ten to thirty seconds or more, the wing pads separating, and the front pair stretch-
ing at right angles from the body and obliquely crossing the hind pair (frontispiece,
figs. 5and 6). They then gradually swell, and during all this time the legs are becom-
ing firmer and assuming the ultimate positions. Suddenly the imago bends upward
with a good deal of effort, and, clinging with its legs to the first object reached, whether
leaf, twig, or its own shell, withdraws entirely from the exuvium, and hangs for the
first time with its head up (frontispiece, figs: 7 and 8). Now the wings perceptibly
swell (frontispiece, fig. 8) and expand until they are fully stretched and hang flatly
over the back, perfectly transparent, with beautiful white veining (frontispiece, fig. 9).
As they dry they assume the roofed position (frontispiece, fig. 10), and during the night
the natural colors of the species are gradually assumed (frontispiece, fig. 11).

The time required in the transformation varies, and, though for the splitting of the
skin and the full stretching of the wings in the flat position the time is usually about
twenty minutes, it may be, under precisely similar conditions, five or six times as long.
But there are few more beautiful sights than to see this fresh forming Cicada in all the
different positions, clinging and clustering in great numbers to the outside lower leaves
and branches of a large tree. In the moonlight such a tree looks for all the world as
though it were full of beautiful white blossoms in various stages of expansion.

A more realistic idea of the important stages in this transformation —
is furnished perhaps by a series of photographs kindly given to the
writer by Mr. Robert A. Kemp, of Frederick, Md. (see PL AVS Dyess
more natural position is given if figs. 1 and 2 are turned so as to make
the twigs perpendicular rather than oblique.

THE ADULT INSECT AND ITS HABITS.
NUMBERS AND LOCAL DISTRIBUTION.

In the case of a well-established brood under favorable conditions,
the enormous numbers of these insects in the soil is most vividly
conveyed by the accompanying photograph (fig. 40) taken by Mr.
Kemp in 1902 (Brood X), near Frederick, Md. Within the foot-
square area in the center of the photograph are no less than 84
openings, which would indicate for the ground surrounding a fairly
good-sized tree the emergence of from 30,000 to 40,000 Cicada pupe.

Mr. McCook took the trouble to count or estimate the burrows
under various trees. Under one tree he counted 9,000 burrows, and
under another, a small birch, the number of exit holes was esti-
mated at 22,500; and since many of the burrows interlaced under
ground and several insects emerged from the same opening, even
these figures do not indicate the actual number. In another case
668 openings were counted in a space 10 by 4 feet, and 17 distinct
openings in a space 6 inches square.

Mr. Davis, referring to Brood IIT on Staten Island in 1894, says:

About some of the trees the pupa shells became so numerous that they completely
hid the ground itself. At dusk the sound of the many insects climbing up the tree

100 ‘THE PERIODICAL CICADA.

trunks was quite audible, particularly vigorous pup ascending the trees to the
height of 30 feet.

Plate V, showing empty pupal shells clinging to leaves, still further
illustrates the large number of these insects which often emerge.

As noted by Mr. Farmsley, of Louisville, Ky., the cicadas do not
appear very numerously on tops of mountains within an infested area,
but gradually decrease in numbers as one ascends, the greater scarcity
being noticeable both to the eye and the ear, the rattling chorus
growing less and less strong.

On the authority of Dr. Hopkins, the diminishing of the Cicada in
numbers as one ascends to higher elevations is apparently not always
true. Dr. Hopkins describes driving for a day through the Cicada
district of West Virginia in 1897 on the occasion of the reappearance
of the 17-year Brood V, and states that as he approached the eastern

Fic. 40.—Exit holes of the periodical Cicada at surface of soil. The rule shows the large number o!
holes occurring, in this instance, in a square foot of ground

borders of Preston County the cicadas became more numerous, and
as the mountain west of Cranesville was ascended the Cicada was
found, at an elevation of 2,600 to 2,800 feet, to occur in far greater
numbers than at any point previously traversed. The leaves and
twigs of the trees were literally covered with the insects, and the twigs:
were bending from their weight. This point seems to have been the
eastern border of the swarm, and a few rods farther up the cicadas
became very scattered and soon ceased altogether.

They often also appear in greatest number in rather well-defined
districts within the general range of the brood, or, in other words,
are irregular in local distribution. This variation in abundance is
due in some cases to differences in the character of the soil and in
others, perhaps, to varying surface conditions, as of timber growth,
etc. They prefer, apparently, white-oak groves, and are most abun-

Bul. 71, Bureau of Entomology, U. S. Dept. cf Acriculture. PLATE VI.

THE TRANSFORMATION OF THE EMERGED PERIODICAL CICADA.

1.—Pupa ready for transformation. 2.—Adult beginning to issue from pupal shell. 3.—Adult
nearly free from pupal shell. 4.—Freshly transformed adult, the coloring immature.
5.—Adult, several hours after transformation, the coloring mature. About natural size.

«
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By
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FOOD HABITS OF THE ADULT. 101

dant where the land is high and well drained and the soil a rich,
sandy loam, with a sandy or soft clay subsoil. The irregularity of
local distribution is confirmed also by the experience of Mr. Davis on
Staten Island, who reports of the 1894 brood that the cicadas were
very rare in sandy districts, while in districts less sandy they appeared
by thousands. He says also that they occurred by millions on certain
hills and in certain bits of woodland, yet at a short distance away,
under apparently unaltered conditions, they were very scantily
represented.

The local abundance of the Cicada in well-defined districts is to
be explained by the fact, already noted, that the winged insect is
sluggish and scatters but little from the point of emergence, which,
with favoring circumstances, tends constantly to concentrate rather
than to scatter the species.

THE FOOD HABITS OF THE ADULT INSECT.

At the time of the writing of Bulletin 14 the observations of many
entomologists who had studied the periodical Cicada were practically
in accord that the taking of food in the adult stage was not a neces-
sary feature of the aerial life of the insect and was of comparatively
rare occurrence. Feeding to a limited extent had been shown, how-
ever, by the observations of Walsh and Riley,? and an-additional
instance is noted by Riley in Bulletin 8.? The observations by Mr.
Davis’ were referred to in Bulletin 14, reporting that the black
birch and sweet gum are favorite food plants, and that it is not
uncommon to see rows of cicadas along the branches of these trees
with their beaks embedded in the bark. Various other entomolo-
gists had noted a little feeding, but the opinion was general that the
feeding habit was unusual and not necessary to the insect. State-
ments had also been made that such feeding was limited to the
female, and that the male could take no food inasmuch as its digestive
organs were rudimentary. No special harm from feeding, at any
rate, had ever been noticed, even where the insect occurred in count-
less myriads.

With the recurrence of Brood X in 1902 Mr. A. L. Quaintance,
then entomologist of the Maryland experiment station, had _ his
attention called to the feeding of the periodical Cicada and made a
thorough study of the subject.? A correspondent called the atten-
tion of Mr. Quaintance to the feeding of the Cicada in his orchard,
and an examination of a local orchard confirmed this fact, which he
afterwards noticed in various localities in Maryland. Cicadas in

a American Entomologist, Vol. I, p. 67, 1868.

6 Bul. 8, o. s., Div. Ent., U. S. Dept. Agric., -p. 14.

¢ Natural Science Assn. Staten Island, 4, September, 1894, pp. 33-35.
@d Bul. 37, n. s., Div. Ent., U. S. Dept. Agric., pp. 90-94, Pl. I.

102 THE PERIODICAL CICADA,

numbers together were often observed with their beaks stuck straight
down against the bark in the attitude of feeding, and in numerous
instances the insects were observed when disturbed extracting the
thread-like setz from the plant tissue. Early in the morning or
late in the evening also the limbs of young apple and pear trees
were frequently quite wet with sap which had exuded from the
punctures made with the sete of the cicadas. This exudation of
sap was frequently noticed to immediately follow the withdrawal of
the sucking apparatus of the insect to such an extent as to run down
the trunk a distance of 4 or 5 inches.

Feeding was also observed in forest trees growing near the college
buildings by means of an opera glass. In the case of forest trees
the insects commonly go to the upper branches and hence are not
near enough for observation from the ground, a fact which may
account somewhat for the failure hitherto to have observed this habit
of taking food.

Professor Quaintance also made cross sections of the wood, show-
ing that the sete had actually penetrated deeply into the sapwood
of the trees. Both sexes were shown to feed to an equal extent,
and dissections of the insects themselves showed the stomach to be
distended to several times its usual size with sap taken from the trees,
and the alimentary canal was found to be perfect in both sexes and
not rudimentary in the male, as hitherto believed. The intestine
was very minute, but could be traced from the cesophageal to the anal
opening.

Professor Quaintance’s observations undoubtedly indicate that the
Cicada in the adult stage normally takes food in the same way as do
other hemipterous insects, and the fact that when these insects are
kept in confinement for a few days without food they invariably die
would seem to indicate the necessity of liquid food. Mr. Quaintance
himself, however, queried if the amount of feeding might not vary
with different broods; and that the Cicada necessarily and always
takes food has not yet been fully established.

The taking of food by the Cicada at any rate seems to cause the
trees normally very little injury and is not accompanied apparently
by any special poisoning of the wood which causes the death or
sloughing off of bark, as is more or less the case with the San Jose
scale, for example; and the belief expressed in Bulletin 14 may be
perhaps adhered to, that, so far as real injury is concerned, the feeding
in the adult stage is a negligible feature.

THE CICADA AS AN ARTICLE OF FOOD.

The fact has already been alluded to that the common name
“Jocust,’’ given by the early colonists to this insect, was undoubtedly
owing to a confusion of the Cicada with the migratory locust of the

THE CICADA AS AN ARTICLE OF FOOD. 103

Orient, which has been an article of diet from the earliest times, and
is so employed at the present day, in various places in northern
Africa and eastern Asia. A similar locust is also now highly esteemed
as a food article in the island of Madagascar. All of these locusts
belong, however, to the class of insects known as grasshoppers, and
on this continent the Rocky Mountain grasshopper or locust has also,
as is well known, been long used as an article of food by certain
Indian tribes.

That the Cicada was eaten by the red men of America, both before
and after the coming of the colonists, is indicated in a memorandum,
dated 1715, left by the Rev. Andreas Sandel, of Philadelphia, who,
referring to the use of locusts as food in eastern Asia, states also
that the Cicada is so used by the Indians. Dr. Asa Fitch corroborates
this statement, giving as his authority Mr. W. S. Robertson, who
informs him “that the Indians make the different species of Cicada
an article of diet, every year gathering quantities of them and pre-
paring them for the table by roasting in a hot oven, stirring them
until they are well browned.”’

No practical test was made with the Cicada as an article of human
food until the experiments instituted by Professor Riley and carried
out by Doctor Howard in the early summer of 1885. The following
is an account of Doctor Howard’s experiments:

With the aid of the Doctor’s (Riley’s) cook he had prepared a plain stew, a thick
milk stew, and a broil. The Cicadie were collected just as they emerged from pupe,
and were thrown into cold water, in which they remained overnight. They were
cooked the next morning, and served at breakfast time. They imparted a distinct
and not unpleasant flavor to the stew, but were not at all palatable themselves, as
they were reduced to nothing but bits of flabby skin. The broil lacked substance.
The most palatable method of cooking is to fry in batter, when they remind one of
shrimps. They will never prove a delicacy.4

Mr. T. A. Keleher, who sampled some of the dishes above described,
has informed the writer that he found the cicadas fried in batter to
be most palatable, and that he much preferred them to oysters or
shrimps.

The great liking manifested by various animals for the pupx before
and after they have emerged and for the transforming adults has
already been referred to. Doctor Hildreth, writing in 1830, says:

While here they served for food for all of the carnivorous and insect-eating animals.
Hogs eat them in preference to any other food; squirrels, birds, domestic fowls, ete.,
fatten on them. So much were they attracted by the Cicade that very few birds
were seen around our gardens during their continuance, and our cherries, etc.,
remained unmolested.

@ Proc. Ent. Soc. Washington, Vol. I, p. 29.
6 Journal of Science, 1830, Vol. XVIII, p. 47.

104 THE PERIODICAL CICADA,

He also states*that when the cicadas first leave the earth they are
plump and full of oily juices; so much so that they are employed in
making soap.

Mr. John Bartram, writing of the brood which appeared near Phila-
delphia in 1749, and referring to the pupe as they appeared near the
surface of the ground toward the end of April, says that they were then
full of a thick white matter like cream and that hogs rooted up the
ground a foot deep im search of them. Dr. Potter refers briefly to the
fact that great numbers of them are ‘devoured by hogs, squirrels,
all kinds of poultry, and birds, which live and fatten on them.”’

That they are sometimes considered to be poisonous when made an
object of food is indicated in the following quotation from Doctor
Phares. He says:

Many species of domestic and wild birds, quadrupeds, and other animals eat the
cicadas greedily and with impunity. In 1859 they were said to have killed a few
hogs in Amite County. They have no poison about them, yet it is not to be wondered
at that an occasional hungry hog or other animal, eating very largely of such food,
should become sick or even die. Dogs become very fond of them. One evening I
watched a bitch catching and eating so many that I expected her to become sick from
her rich feast of fat things, but she was in no way injured. Indeed, I have never seen
any animal injured or otherwise.

As has been indicated elsewhere, the liking of domestic animals and
birds, especially the English sparrow, for the cicadas, both in their
newly emerged condition and in the mature state, is one of the most
potent influences in exterminating or greatly reducing the abundance
of this insect in thickly settled districts.

The use of the newly emerged and succulent cicadas as an article of
human diet has merely a theoretical interest, because, if for no other
reason, they occur too rarely to have any real value. There is also
the much stronger objection in the instinctive repugnance which all
insects seem to inspire as an article of food to most civilized nations.
Theoretically, the Cicada, collected at the proper time and suitably
dressed and served, should be a rather attractive food. The larve
have lived solely on vegetable matter of the cleanest and most whole-
some sort, and supposedly, therefore, would be much more palatable
and suitable for food than the oyster, with its scavenger habit of
living in the muddy ooze of river bottoms, or many other animals
which are highly prized and which have not half so clean a record as
the periodical Cicada.

OVIPOSITION AND ITS EFFECT ON THE PLANT.

The Cicada becomes almost perfectly hardened and mature during
the first day of its aerial life, and does not wait many days before
beginning the important business of its existence in the perfect stage,
namely, depositing the eggs for another brood. Courtship occupies
a comparatively short time, and the sexes are found together usually

PLANTS SELEOTED FOR OVIPOSITION. 105

within a week after the emergence of the first individuals. Within
two weeks the egg punctures begin to. appear here and there in the
twigs. From this time on oviposition proceeds very rapidly, and
thousands of individuals may often be noted working at the same
time on the same tree.

PLANTS SELECTED.

The fact that the Cicada is not especially choice in its selection of
trees in which to place its eggs is patent to any careful observer,
although a preference is generally shown for oaks and hickories, and
the apple among the fruit trees. Any plant which presents itself is,
however, accepted, often herbaceous ones and occasionally evergreens,
although the sticky resinous sap of the latter seems to be distasteful
to these insects. No careful, complete list of plants in which they ovi-
posit has been made, although several observers have made rather
extensive lists, notably Mr. Butler and Mr. Davis, the latter having
observed the cicadas laying their eggs in between seventy and eighty
trees, bushes, and herbaceous plants on Staten Island in 1894, and
states also that he had evidently not nearly reached the limits of
plants. In some cases even the large petioles of plants, like the horse-
chestnuts, had been oviposited in. A list of plants could be given
which have been put on record, but it would have but little value, as
in every district in which es appear they will oviposit in practi-
cally all plants which come their way, with the exception of pines, as
already stated, which are ordinarily exempt.

That they are not very choice in this matter is shown by a case of
faulty instinct reported by Mr. Hunter Nichols, who observed a female
to alight on the iron rod of a bridge and attempt to insert her eggs,
even extruding them to the number of seven, some of which remained
attached to the rod and the others falling to the ground. Other
similar cases of error on the part of the insect are noted by Mr. Davis.
In one instance a female had attempted to insert her eggs in the
very hard stem of catbriar (Smilax rotundifolia) and in another place
had thrust her ovipositor entirely through the stem of a plant only to
find that it was hollow.

The part of the plant selected for a receptacle for the eggs is almost
invariably the twigs of the previous year’s growth. When larger limbs
are chosen, as occasionally happens, the female evinces her dislike for
them by constructing only a nest or two instead of the long series of
slits which are usually characteristic of her work on limbs of newer
growth.

106 THE PERIODICAL CICADA.

RESULT TO THE

PLANT OF OVIPOSITION.

The effect on the plant of the cutting of the smaller twigs by the
female in depositing her eggs has been often described, and is apt to

Fic. 41.—Egg punctures of period:cal Ci-
cada: a, twig showing recent punctures.
from front and side, and illustrating man-
ner of breaking; b, twig showing older
punctures, with retraction of bark and
more fully displaying the arrangement
of fibers. Natural size (after Riley).

the Cicada abounds, the injury is apt to be very con-

siderable.

newly cleared ground

be especially noteworthy and disas-
trous in the case of such favorite trees
as the oak, hickory, and apple, and in
the case of the latter, especially in
young orchards surrounded by woods,
or in recent clearings. (See fig. 41.)
The weakening of the twigs by the
punctures causes many of them to be
partly broken off by the winds, and
the brown, withered leaves are con-
spicuous for the remainder of the sum-

‘mer. Many of the twigs break off en-

tirely and fall to the ground, and the
general twig pruning which results is
often of considerable extent, giving
the forests, as sometimes described, a
eloomy appearance, or as though
scorched by fire, from the number of
the extremities of twigs thus injured.
With large shade trees,
and particularly trees
in forests, the damage
is not often excessive,
and the recovery is
usually complete, or
nearly so, within two
or three years. With
fruit trees and nursery
stock, on the other
hand, and especially on

or in the vicinity of
forests or groves where

1G. 42.—Twig show-
ing scars from

The following extract from a letter from Mr. Wil- _ punctures after the

second year. (After

liam G. Wayne, of Seneca Falls, N. Y., illustrates 3...)
the injury sometimes experienced. Referring to
the Hudson River Valley brood, appearing in 1826, he says:

They destroyed the fruitage of the orchards almost completely. Nearly all the
tender branches of the trees were so wounded in the deposit of the eggs that they

RESULT TO THE PLANT OF OVIPOSITION. 107

broke from the main stems in the following year and fell to the ground, thus completely
denuding the trees of their fruit-bearing branches.¢@

Peach, pear, and apple trees suffer most, and even grapevines are
often badly injured. With fruit trees in vigorous condition and grow-
ing rapidly, however, the wounds heal in a few years so that often the
sears can scarcely be detected; but, as shown by Dr. A. D. Hopkins,
with recently transplanted trees, the growth of which is slow, and
with the fruiting and terminal branches of old trees which lack vigor,
the wounds often do not heal for many years.

Another form of injury has been charged to this insect by some
of the earlier writers, viz, that after fillmg the twigs with her egg
clusters the female completely or partly severs it, causing it to break
off and die. This opinion is totally without foundation in fact, and is
undoubtedly based partly on the observation that many twigs are
broken by the winds and partly on a confusion of the work of the
Cicada with that of certain oak-pruning beetles, which after oviposit-
ing in the branches, cut them nearly off, causing them to fall to the
ground, thus furnishing their larve the dead or dying wood in which
they develop.

The absurdity of the theory that the Cicada purposely cuts the
limbs to weaken them and cause them to break off is shown by the
fact that wherever a limb is broken, through the weakening from
excessive puncturing or other causes, and falls to the ground, the dry-
ing up of the limb invariably causes the eggs to shrivel and die. The
breaking off of limbs, therefore, is purely accidental, and is confined,
so far as due to the Cicada, to the smaller terminal twigs which have
been too thickly oviposited in, the female by so doing defeating her
own object. The proportion of such broken and fallen twigs, while
often great enough to give the tree a deadened appearance, is small
in comparison with the many thicker and stouter limbs which remain
attached, and probably more than 90 per cent of all the eggs, and
more than 99 per cent of those that ultimately hatch, are laid in
twigs which never break off, though often much injured. A very
few young may come from twigs which are partly broken off, but
in such instances the flow of sap has not been entirely stopped.

The after effect of the egg punctures on the twigs is shown in the
deformity which characterizes their subsequent growth. In the
process of healing the punctures usually assume a wart or knot-like
appearance,‘as represented in the accompanying illustration of an
apple twig (fig. 42). The effect of punctures in hard-maple twigs after
the lapse of seventeen years is shown in fig. 43, and on various plants
in Plate I (see p. 12), these illustrations bemg kindly loaned me by Dr.
Hopkins.” Though ultimately healing over exteriorly with the growth

a Vintner, Second Report Insects New York, p. 177.
b Bulletin 50, W. Va. Agr. Exp. Sta., Pls. Il and IV.

108 THE PERIODICAL CICADA,

of the surrounding wood, there remains in the center of the twig a
dead spot, and the white glistening egg shells of the escaping larve
have been found in place six years after they have been inserted in
the twig by the fomale Cicada.

Considerable danger follows the work of the Cicada, in that as long
as the wounds remain open or as dead spots on the limbs they arenot
only a source of weakness in the case of winds, but they offer attract-
ive situations for the attacks of various wood-boring insects. — If left
to themselves the limbs might entirely recover, except for the scars,
but the borers gaining entrance through these spots complete the work

FIG. 43.—Periodical Cicada scars in hard-maple twigs after seventeen years. (Hopkins.)

of destruction which the Cicada began. Furthermore, such open
wounds or pockets in the twigs of fruit trees Doctor Hopkis has
shown to be favorite points of attack for the woolly aphis (Schizon-
~ eura lanigera Hausm.), the presence of which not only prevents the
wounds from healing but causes additional abnormal erowth, adding
considerably to the injury to the branches, and nike hea more
liable to the attacks of other insects.

METHOD OF INSERTING THE EGGS. ~ 109

METHOD OF INSERTING THE EGGS.

The work of the female Cicada in inserting her eggs is an interesting
subject for study, and so little does she mind the presence of an
observer that the operation can be closely watched without her exhib-
iting any alarm. The position taken is almost invariably with the
head upward or directed toward the tip of the branch, the work being
steadily prosecuted in that direction. When her course is interfered
with by the occurrence of side shoots, instead of moving to one side
or the other she reverses her position and thus extends her row of
punctures in a straight line completely to the base of the intervening
shoot. The branch selected is ordinarily of a size which the female
can surround and clasp firmly with her legs to give her the strong
attachment necessary to enable her to force her ovipositor into the
woody tissues.

The exact method of making the egg fissure and depositing the eggs
has hitherto, in the
main, been either
very briefly referred
to, or the actions of
the insect have been
inaccurately — inter-
preted. The descrip-
tion of this process,
hitherto generally ac-
cepted and quoted,
is that given by Doc-
tor. Harris, substan-

Cc
tially as follows: Fig. 44.—Egg nest of the periodical Cicada: a, recent puncture, front
ae view; 5, same, surface removed to show arrangement of eggs, from
Raising her body above; c, same, side view; d, egg cavity exposed after eggs are
somewhat above the removed, and showing the sculpture left by the ovipositor. All
enlarged (after Riley).

twig, the point of her
ovipositor is brought to bear on the bark at an angle of 45 degrees,
and is thrust slowly and repeatedly into the bark iia wood, the two

lateral saws working in alternation. When fully inserted he instru-
ment is pried upward by a motion of the abdomen, raising and loosen-
ing in this way little fibers of wood which, remaining attached, form
a sort of covering for the egg fissure or nest. The cutting normally
extends nearly to the pith or about one-twelfth of an inch in depth,
and is continued until space is made to receive from ten to twenty
eggs. After preparing the egg nest as described, the female moves
back to the point of commencement and again abaeste | in her oviposi-
tor, using the two side pieces as grooves or channels to convey the
eggs into the twigs, where they are placed in pairs, separated by a

central tongue of woody fiber, which has been left undisturbed, and
which is wider at the bottom than at the top. Two eggs having been

PO = THE PERIODICAL CICADA.

inserted in the portion of the fissure first made, the ovipositor is
withdrawn and again inserted, and two more eggs are placed in line
with the first; this operation being continued until the egg nest is
filled. A step or two forward is then taken, and after a brief pause
anew egg nest is begun. About fifteen minutes is occupied in pre-
paring and filling one of these nests with eggs.

The above account is substantially correct so far as the superficial
appearances are concerned. Instead, however, of first making an egg
nest and afterwards filling it with eggs in pairs, as described, the female
deposits the row of eggs on one side as she makes the original cutting
in the bark. She then moves back, and, swinging a little to one side,
inserts through the same hole the second row of eggs parallel with the
first, thus leaving a small bit of undisturbed wood fiber between the
two rows of eggs. This method of inserting the eggs corresponds to
that known to be true of allied insects which deposit their eggs in prac-
tically the same manner, and is confirmed also by the careful observa-
tions made by Mr. Ira H.. Lawton, of Nyack, N. Y., m 1894, and
reported by Professor Lintner.* Mr. Lawton found that the placing
of each row of eggs occupied a little over twenty minutes, or, for the
construction and filling of the double egg nest, some forty-five minutes.
During the cutting of the fissure the ovipositor made about eighty
strokes per minute, and after four chambers were made the female
would indulge in a short rest.

The number of nests made in a single twig varies from four or five to
fifteen or twenty, the latter number being not at all unusual, and as
many as fifty egg nests in a line, each containing fourteen to twenty
eggs, have been found in a single limb. The punctures are often
made so close to each other that they sometimes run together, so as to
form a continuous slit for 2 or 3 inches.

The Cicada passes from one limb or from one tree to another until
she has exhausted her store of eggs, which have been estimated to
number from four to six hundred. By the time the egg-laying is com-
pleted the female becomes so weak from her incessant labor that she
falls to the ground and perishes or soon becomes a victim to her various
natural enemies.

~ THE GROWTH AND HATCHING OF THE EGGS.

The eggs remain in the twigs for six or seven weeks after being
deposited. Professor Potter was one of the first to determine this
rather unusually long egg period by marking certain egg clusters and
watching them until the young larvee were disclosed. He reports that
from eges deposited on the 5th of June he witnessed the hatching of
the young on the 28th of July. This statement is also corroborated by

a Twelfth Report Insects New York, p. 275.

GROWTH AND HATCHING OF EGGS. hy

Doctor Smith. Miss Morris and others record a shorter period, and
there is undoubtedly considerable variation due to weather conditions,
but the normal period, as shown by the abundant records of this office,
and many observers, since those noted, ranges, as stated, from six to
seven weeks. Some interesting instances have been noted of retarded
development of eggs in plants yielding gummy exudations which had
hermetically closed the nests from the outer air. Professor Riley
notes a case of this kind where the eggs remained sound and un-
hatched until the end of the year, long after the trees had shed their
foliage. Except in the extreme south, where all of the periods are
somewhat earlier, the eggs are deposited chiefly in the month of June
and most abundantly about the middle of this month, and the hatch-
ing period ranges from the middle of July to the first of August.

The egg is a very delicate, pearly-white object, about one-twelfth of
an inch long, tapering to an obtuse point at either end and slightly
curved. The shell is very thin and transparent, the form of the larval
insect showing through some time before hatching. As is the case
with most insects that oviposit in the living parts of plants, the eggs
of the Cicada receive a certain nourishment from the plant and
actually increase in size before hatching,
by absorption of the juices from the a
adjacent plant cells. ei LETT | |

Discussing the development of the j,. 4; Se ope icat PO A ab
embryo, Doctor Potter says that on the enlarged, showing young about to be
fifteenth day a change in color in the egg “18°? Authors tlustration.
may be noted, and from this time on there is a gradual increase in
size, the embryo slowly assuming form—the eye becoming especially
prominent some ten days before hatching (fig. 45).

The larval Cicada makes its escape by rupturing the eggshell over
the back, from the upper end downward about halfway, by muscular
“movements, accompanied with an inflation of the head and forward
parts of the body. The rupture in the shell once made, the larva
works its way out by twistings and contortions until the tip of its
body only remains in the egg slit of the shell. The entire insect, how-

_ever, is still inclosed in an extremely delicate and almost invisible
membrane (amnion), and after resting a short time the violent move-
ments are again resumed, and by wriggling, twisting, and inflating its
head, thorax, and anterior parts the thin enveloping skin is burst open,
and by gradual efforts, coupled with contractions and expansions of the
body, the larva draws itself out, leaving the thin white skin held in the
tip of the eggshell. The larve nearest the opening come out first,
the others following in regular order, each usually pushing out the
abandoned egesheil of the preceding one, though commonly several
eggshells remain attached to the loose woody fibers of the egg nest.

-

I al 2s THE PERIODICAL CICADA.

Almost at the moment that it becomes free, the larva begins to
run actively about with the quick motions of an ant, but soon goes
to the side of the limb, loosens its hold, and deliberately falls to the
ground, its specific gravity being so slight that it passes through the
air as gently as a feather and receives no injury. The peculiar
instinct which impels this newly hatched larva to thus precipitate
itself into space without the least knowledge of the distance to the
ground or the result of its venture has been often commented upon,
but is not more remarkable than other features in the life history of
this species.

On coming to the earth the larva immediately penetrates it, usually —
entering at a crack or fissure, or at the base of some herbaceous plant,
and begins the leng period of its subterranean life.

The newly hatched larva (fig. 46) is about one-sixteenth of an inch
long and differs considerably in general form from the later larval
stages, while at the same time presenting the general structural char-
acteristics shown in the latter. It has, for example, a much longer
and distinctly eight-jointed antenna,
and the head is longer in proportion
to the body. It is yellowish white in
general color, except the eyes and the
‘ claws of the anterior legs, which are
Fic. 46.—Newly hatched larva, greatly reddish. It is sparsely covered with

SHEE Varner er minute hairs. In form it is quite elon-
gate and subcylindrical, and it is particularly notable for its very
prominent front legs.

THE UNDERGROUND LIFE OF THE CICADA.

EXPERIMENTAL PROOFS OF THE LONG UNDERGROUND LIFE.

The life and habits of the periodical Cicada above ground, which °
have so far only been discussed, are open to easy study and have been
fairly well understood, certainly since the time of Hildreth, Potter, and
Smith; but from the time of the disappearance of the young larva be-
neath the soil and thereafter, throughout its long hypogeal existence,
observations are difficult and with the earlier observers were limited to
the occasional or accidental unearthing of specimens, and no consecutive
series of observations of a definite brood or generation was attempted.
The proof for the 17-year or 13-year period for the development of
the Cicada was, therefore, based solely on chronological records, but
so noteworthy were the recurrences of the important broods and so
full and complete were the records, some broods having been regularly
recorded on the occasion of each visit for nearly two hundred years,
that there was no possibility of doubting the accuracy of the time
periods from such records alone; nevertheless, this unusual feature in

PROOFS OF LONG UNDERGROUND LIFE. Pls

the life of the Cicada always arouses skepticism in the minds of persons
who have not given the matter study and have not examined the
historical records. To silence such objectors, rather than because of
the need of experimental proof, Professor Riley was for many years
interested in demonstrating by actual rearing experiments the period
of underground development of this insect; in other words, to follow
a particular generation through its subterranean life of seventeen or
thirteen years, as the case might be, watching its development and
preserving examples of the different stages.

The great difficulty of conducting to a successful termination experi-
ments of this sort will be appreciated when the long period over which
the experiments must necessarily extend isremembered. The extreme
delicacy and softness of the larve themselves, especially in the first
years of their existence, introduces an additional difficulty, as the
slightest touch or pressure injures or crushes them and renders them
unrecognizable. It is therefore often difficult to find them, even when
the soil is very thickly tenanted.

The difficulty of carrying out breeding experiments with the Cicada
under any but natural conditions is illustrated by various efforts in this
direction undertaken by this Bureau. In one instance a number of
newly hatched Cicada larve were allowed to penetrate the soil about a
potted oak tree of small size. None of these larve survived for a
single year. In another instance the larvee were allowed to penetrate
the soil in large breeding tanks, each containing young trees, the tanks
being planted out of doors in the soil. These were left undisturbed for
a number of years, and although the conditions were seemingly very
favorable for a successful outcome, when an examination was finally
made, no traces of the larve were found.

The earliest systematic attempts. to follow the development of the
Cicada were made in the field in Missouri by Professor Riley, and
subsequently continued under the latter’s direction by Mr. J. G. Bar-
low, an agent of the Bureau. They consisted in making diggings
from year to year under trees which were known to have been thickly
stocked with eggs. The first records approaching in any way to
completeness were obtained with the 13-year Brood XIX, beginning
with its appearance in 1881. Observations on this brood were con-
tinued by Mr. Barlow at Cadet, Mo., with a fair degree of regularity
until July, 1891, when they unfortunately terminated.

During the ten years over which these observations extended the
insect had developed through all four larval stages and was ready to
enter the first pupal stage. The first molt occurred after a period of
from one year to eighteen months, the second molt after an addi-
tional period of two years, the third molt after an additional period
of three years, and the fourth molt after an additional period of

31117—No. 71—07——8

114 THE PERIODICAL CICADA,

three or four years, leaving in this 13-year brood three or four years
more for the pupal stages.

A much more careful series of experiments was instituted in con-
nection with the 17-year Brood X, beginning with its last appearance
in 1885. At the time that the eggs of the 13-year Brood XXIII were
being distributed to various points in the North, in order to determine
the effect of the temperature and climate (see pp. 18-20), quantities
of ege-laden twigs of the 17-year brood noted, collected in Virginia,
were distributed under certain linden and oak trees on the grounds
of the Department of Agriculture at Washington, D. C. Larve
came from these twigs in some numbers and went into the soil under
the trees, but not in such abundance as could have been wished for
the successful outcome of the experiment. This brood was followed
in its underground life from 1885 to 1896, at which time the speci-
mens had become so rare that extensive digging resulted in the dis-
covery of very few individuals, and further search was abandoned.
With this brood the first molt occurred after one year, the second
molt two years later, the third molt three or four years later, and the
fourth molt after an additional three or four years, thus occupying
upward of ten years with the four larval changes and bringing the
insect into the last larval stage with some six or seven years for the
subsequent larval and pupal life. If any adults emerged at the end
of the 17-year period in 1902 they were not observed.

A SUCCESSFUL SEVENTEEN-YEAR BREEDING RECORD.?

A much more promising experiment, because of more abundant
material, was instituted on the Department grounds in 1889 with the
17-year race which appeared in that year and which had its return
appearance during May and June of last summer (1906). This brood
is practically unrepresented in the District of Columbia, and did not
occur at all on the Department grounds. <A very large quantity of
egg-infested twigs was obtained from North Carolina, Long Island,
Kentucky, and Ohio, several cartloads altogether, and were distrib-
uted under oak and other trees on the grounds of the Department of
Agriculture. The eggs in most instances were hatching when received
and were placed under the trees in the very best condition for the
larve to enter the soil, and many thousands, probably hundreds of
thousands, of larve actually went into the soil under these trees. This
experiment was made during the first year of the writer’s connection
with the Bureau of Entomology, and the later examinations were
made chiefly under his direction. Three years after the planting the

soil under the trees where the egg-bearing twigs had been distributed

«The records of the plantings on the Department grounds of the eggs of Brood X
in 1885 and Brood XIV in 1889 are given in Appendix B, of Bul. 14,

A SUCCESSFUL SEVENTEEN-YEAR BREEDING RECORD. 115

was found to be thickly filled with larvee, so much so that a single
spadeful of earth would often turn up half a dozen or more. In the
spring of 1897 the larve had reached the fourth stage and were still
very abundant in the soil. Examinations were made from time to
time showing these larve to be still present in the soil about the trees
where the eggs had been distributed, going through the slow process
of growth and transformation which has been described elsewhere.
That a successful outcome was sure to be had in this experiment was
demonstrated in the early spring of 1906, the year for the appearance
of this brood, the ground about the planted trees exhibiting many of
the exit holes of the insects which are made to the surface long before
the insect emerges. These holes under certain trees were so numerous
as to indicate the emergence of thousands of cicadas. Under one tree
a count and estimate were made of more than five thousand openings,
and under other trees the openings ranged from a few hundred to from
one to three thousand. The actual emergence took place between
May 14 and 21. The writer visited the grove on two evenings and
witnessed the issuance of numbers of cicadas and collected some
specimens. In spite, however, of the considerable number of cicadas
which actually emerged, none was seen on the trees during the days
and weeks following emergence. Each morning about the planted
trees would be found a considerable group of blackbirds (Quiscalus
quiscula), which evidently had been feasting on the newly-issued
cicadas. The cast pupal shells were numerous on the trunks of the
trees and especially on the foliage, and also on the ground, but scarcely
a single Cicada escaped the sharp eyes of these birds, and the charac-
teristic song was not heard during June in this grove, although thou-
sands of adults had come forth.

At none of the examinations were Cicadas found of this brood under
any of the trees except where eggs had been distributed, and no
emergence holes appeared under other trees. The record from the
planting to the emergence of this insect is therefore complete, and
gives the demonstration by actual transfer and breeding record of the
long period of the 17-year brood, a demonstration which, as indicated
at the outset, was-entirely unnecessary to show the correctness of this
extraordinary hypogeal term.

The absolute failure of these insects to establish themselves when
planted in such enormous numbers, and even when the underground
period had been successfully passed, owing to the relentless onslaught
of birds, is a striking illustration of what is happening every year
with the different broods in nature, especially in thinly forested
regions, and accounts for their great reduction in numbers and the
practical disappearance of many local swarms formerly abundant.
It also shows that there may be emergences in considerable numbers

116 THE PERIODICAL CICADA,

without their being reported, unless some observer chances on a pupal
shell or notes the exit holes in the ground about the trees; hence
the slight value of a negative report as opposed to a positive one.

HISTORY OF THE LARVAL AND PUPAL STAGES.

A careful study of the material collected in the course of the experi-
ments described in the last section demonstrates the interesting fact
that this species, in spite of its very long period of growth, presents
the same number of adolescent stages as is found in insects which go
through their entire development within a single year or even of the
more rapidly multiplying species, which have many annual genera-
tions. But six distinct stages are found, four of which belong to the
larval condition and two to the pupal. In other words, the larval
and pupal changes in the periodical Cicada are normal and are not
increased by its long preparatory existence.

It has been inferred hitherto, and notably by Professor Riley, that
owing to the continual use of the claws in burrowing, this species found
it necessary to shed its skin and undergo a molting once or twice a
year, and instead of the normal number of changes or molts there
were probably from twenty-five to thirty. An examination of types
of the different larval stages which Professor Riley had provisionally
separated demonstrates that the differences on which these supposed
stages were based are either individual and exceptional or due to the
difference of age within the same stage, and that as far as structure
and size of the hard parts of the larva and pupa are concerned the
normal number of stages only is represented in this species.

For the separation of these different stages of growth useful char-
acters are found in the size and structure of the legs, and especially of
the anterior pair, in the antennz, and in the development of the wing
sheaths. It is the rule with insects that with each molt there is a
decided increase in the size of the head and hard parts generally, and
with the periodical Cicada especially it is also very doubtful if there
is ever a molt without a decided change of the sort indicated. Its life
beneath the ground in its moist cell over a rootlet is a very quiet one and
free from any of the wearing action of rain or the drying of the outer
air, so that the need of a molting or change of skin would apparently be
much less than that in an exposed or much more active insect. It prob-
ably also very rarely has occasion to burrow to any considerable extent
and probably often remains for years in the same cell, which it enlarges
from time to time without change of location. For these reasons the
writer is inclined to believe that moltings only occur when change of
form becomes necessary by the increased size of the insect, and this
seems to be borne out by definite structural peculiarities, which easily
permit us to recognize the different stages or determine the age of any
Jarva within a year or two. The larva of a particular molt or stage

HISTORY OF THE LARVAL AND PUPAL STAGES. el ¥4

of growth will vary considerably in size of the body and the softer
parts, representing perhaps a difference in age in some cases of one
or two years, but the hard parts will present a very uniform size and
character.

The peculiar structure of the enlarged anterior legs furnishes per-
haps the best means of distinguishing the adolescent stages of this
species from other cicadas and the modification which these limbs
undergo with the different molts the best means of determining the
age of the larve. The peculiarities of the anterior legs consist in the
enormous enlargement of the femora and tibiz and their development
into structures which resemble somewhat the cutting mandibles of
biting insects or recall the fossorial forelegs of the mole cricket. The
peculiar structure of these legs is in fact especially designed for dig-
ging, tearing, and transporting earth in the course of the insect’s sub-
terranean life. As already indicated, the amount of burrowing in
the early stages is not necessarily very great in any one year, but dur-
ing the entire seventeen years conditions may occasionally arise which
will demand a considerable activity on the part of the young Cicada.

The details of the structure of the front legs, which are given in the
technical description of this species, are quite characteristic and
diverge notably from the similar parts of other species. The anterior
tarsus of the periodical Cicada exhibits also a rather peculiar meta-
morphosis during the adolescent life of the species. In other words,
during the first larval stage and in the pupal stage it is similar to the
other tarsi but considerably longer, being attached to the inner side
of the greatly enlarged tibia and at a considerable distance from the
clawlike tip of the latter. The fore tarsi are of service to the young
larva in walking and climbing and in the same way to the pupa after
its emergence from the soil, facilitating its climbing trees or other
objects; in other words, covering the periods between the hatching
and entering the soil and between the emergence of the pupa and the
disclosure of the imago. During its long subterranean life, however,
these long, slender tarsi, being distinctly in the way in digging in the
earth and of no service, become rudimentary with the first molt and
nearly disappear in the subsequent larval stage. They reappear in
the first pupal stage, but in this and the subsequent pupal stage,
while the insect is still below the soil, they are folded back along the
tibiz, so as to be practically functionless (see fig. 51), and are only
unfolded and brought into service after the pupa has emerged from
the ground.

The more detailed description of the different stages which follows
will facilitate the easy recognition of any particular stage. The chie!
points to be considered in determining both the age of the larva anc
whether or not it belongs to the periodical species are the measur: -
ments of the corresponding parts of the legs and antenne, but pc

Ts THE PERIODICAL CICADA.

ticularly the variation in the structure of the peculiar comb-like
organ which is found on the apical margin of the front femora, together
with the important differences in the hairy covering of the body and
legs.

TECHNICAL DESCRIPTION OF THE DIFFERENT STAGES.

First larval stage-—The newly hatched larva (fig. 47, a) is about 1.8
mm. long from tip of head to the extremity of the abdomen, is rather
slender and of a nearly uniform thickness throughout, presenting,
however, the general characteristics of the later larval stages. The

body is clothed with numerous
scattering long hairs. The gen-
eral color is creamy white, with
h prominent, deep red, almost black,
eye spots. The antenne, beak,
and legs are, relatively with other
stages, very large in comparison
with the size of the body. The
anterior legs are developed in
general as in the later stages,
though lacking the femoral
comb-like organ which begins in
the second stage and the minute
second subapical tooth on the tibia
which appears in the fourth stage.
The anterior tibize are also
more slender and the mandible-
like tip is more sharply pointed.
The row of stiff hairs for retain-
ing the earth excavated in bur-
Fie. 47.—First larval stage: a, newly hatched TOW EEg [OUI in the
larva; b, antenna of same; c, larva eighteen later stages, 1s but sparsely rep-
months old; d, enlarged anterior leg of same. pegented. The anterior tarsus
(Author’s illustration.) : je 5 : 5
is inserted considerably within
the tip of the tibia, projecting beyond the Jatter, and is armed
at its extremity with two nearly equal, curved claws, similar to
those on the middle and hind tarsi. The basal joint of the two-
jointed tarsi in all the feet is very minute and with difficulty detected,
and in fact becomes still more inconspicuous in later larval develop-
ment. The antenne are seven-jointed, as in all the subsequent larval
and pupal stages (one of the characters distinguishing this species
from other allied species, particularly C. tibicen, which has an addi-
tional joint); but the presence of a very prominent antennal tubercle
gives an appearance of eight joints, the number which I have hitherto
assigned to it. The first true joint is robust and a little shorter than

TECHNICAL DESCRIPTION OF STAGES. 119

the second, the two following are subequal and sborter than the first,
the fifth is shorter than the fourth, and the sixth and seventh are
subequal and shorter than the fifth, the last tapering regularly from
the apex, which is armed with curved spines, one long and one short.
The terminal three joints form something of a club tip. During this
stage the larva increases in length to more than 3 mm. and the abdo-
men swells and becomes more robust. Thelength of the hard chiti-
nous parts remain, however, unchanged, as follows: Anterior femora,
0.27 mm.; anterior tibiz, 0.30 mm.; hind tibiz, 0.33 mm.

This stage lasts more than a year, the first molt usually occurring
during the second year after hatching. (See fig. 47.)

Second larval stage—The average length of the larva in this stage is
about 4mm. The more horny parts now measure: Anterior femora,
0.50 mm.; anterior tibix, 0.55 mm.; hind tibia, 0.80 mm. The
general appearance is unchanged from the later development in the
preceding stage. The eye-spots are still present, though reduced.
The under surface of the head is armed with some rather long hairs,
and a very regular row of minute spines occurs on the anterior face
of the hind and the middle femora.
The prominent apical tibial spur of
the middle and the hind pair ap-
pears with this molt, bemg previ-
ously represented, if at all, by a sim-
ple spine. The third joint of the
now distinctly elbowed antennz is Fic. 48.—Second larval stage: a, anterior leg,
as long as the second, and the three tee he ia ena bec ane
terminal joints are rather more com-
pressed into a club-like tip than in the first stage. The chief charac-
teristics of this second stage, however, are in the anterior legs (fig.
48). The femora now possess a rudimentary comb of three teeth, the
upper tooth being very broad and projecting beyond the two suc-
ceeding sharp ones, of which the lower is the larger. The central
tooth of the femora, which was rather minute, or, more properly, a
mere spine in the first stage, is now very much larger and broadened
at the base into a prominent triangular projection. The tarsus is
reduced to a horny rudiment about three times as long as wide, and
is closely applied to the inner surface of the tibial ‘“‘jaw” which
extends twice the length of the tarsus beyond the latter.

This stage, as already stated, is assumed during the first two or
three months of the second year of the insect’s existence and lasts
nearly two years.

Third larval stage —Length, 6 to 8 mm.; anterior femora, 1.20 mm.;
anterior tibiz, 1.35 mm.; hind tibiew, 1.85 mm. Eye-spots still more
reduced; numerous parallel rows of short hairs on the head are notice-

120 THE PERIODICAL CICADA,

able; hairy armature of legs more distinctly outlined; a row of small
spines on either side of middle and hind tibiz, while the rows of bristles
on the inner margins of the anterior femora and tibiz for holding the
excavated earth are well developed. Anterior tarsus reduced to a mere
tapering spur about two and one-half times as long as wide at base.
The femoral comb has one additional tooth, making four in all, count-
ing the blunt upper
one (fig. 49). The an-
tennal joints decrease
in length from the
basal to the terminal,
the basal two and the

terminal two _ being,

Fig. 49.—Third larval stage: a, anterior leg, outer face; b, same, however. of nearly
inner face. (Author’s illustration.) } .

equal length, respec-
tively. The wing cases are foreshadowed by minute pads. Sexual
differences very faintly discernible.

The larva is in this stage at the completion of the fourth year of its
existence.

Fourth larval stage.-—Length, 10 to 15 mm.; anterior femora, 2.40
mm.; anterior tibiz, 2.70
mm.; hind tibizw, 4 mm.
Eye-spots reduced to
from three to six minute
black points, rows of
hairs on head easily dis-
cernible and prominent;
spines on femora and
tibize of all legs, and par-
ticularly the anterior
pair, more numerous and
longer and stouter than
in the preceding stages.
The anterior tibia has a
small tooth within the
larger blunt subapical
one. The femoral comb
has again an additional Fia. 50.—Fourth larval stage: a, full grown larva, muchen-
tooth. making five in all. larged; b, outline of femoral coms, Cc, anterior leg, outer face;

) 3 d, same, inner face. (Author’s illustration.)
Antenne as in the preced-
ing stage. Rudimentary wing cases somewhat more prominent than
in the last stage, but still inconspicuous. (See fig. 50.)

The larva is in this stage at the completion of the eighth year of its
existence, and the stage probably lasts three or four years.

AMTARASANS

TECHNICAL DESORIPTION OF STAGES. {All

First pupal stage—Length in the early condition of this stage about
17 mm.; anterior femora, 3.30 mm.; anterior tibiz, 3.60 mm.; hind
tibiz, 5.80 mm.; width of head, 6 mm. LEye-spots entirely wanting;
eye prominences well developed, as in later pupal stages. Wing cases
extend to the tip of the third segment. Third antennal joint one-
third longer than second, fourth as long as second, others decreasing
in length. The anterior tarsi reappear perfectly developed, and are
nearly as long as the tibizw, and are folded along the inner face of the
latter; the first joint is very minute, and the second or last very long—
longer than the middle or posterior pairs—and armed with two curved
claws at the tip, of which one is rather longer than the other. Femoral
comb with an additional tooth, a very minute one being distinctly
separated from the large blunt
upper tooth. The anterior
tibiz have within the large
blunt subapical tooth, which
has occurred all along hitherto,
two minute saw-teeth instead
of the one present in the pre-
ceding stage (fig. 51). The
hairs of the legs and body are
arranged as hitherto, but are
rather more numerous and
longer, and this is particularly
true of the anterior limbs.
The sexual characters which
have been foreshadowed in the.
two later larval stages are now
distinctly defined.

Second pupal stage-—This
stage does not present any Fic. 51.—First pupal stage: a, anteriorleg, inner face,
differences from the last except showing tarsus bent Deck aeatust the tibia; b,
e same, outer face. (Author’s illustration.)
in the greater size of the speci-
mens, which is noticeable in the relative dimensions of the parts
hitherto measured for comparison. The length of the adult pupa
varies from 27 mm. in the case of the males. to about 35 mm. in the
case of the larger females. The adult pupa of the male presents the
following length of the parts referred to: Anterior femora, 3.80 mm.;
anterior tibiw, 4.30 mm.; hind tibiw, 6.70 mm.; width of head, 6.70
mm. In the case of the female: Anterior femora, 4.20 mm.; anterior
tibie, 5 mm.; hind tibiw, 7.50 mm.; width of head, 7.50 mm. The
anterior tarsus in all unearthed specimens is folded closely back
against the face of the tibia, but in all aerial specimens is unfolded
and projects forward to be of service in climbing.

122 THE PERIODICAL CICADA.

THE HABITS OF THE LARVA AND PUPA.

During its long life beneath the soil, in its small moist oval cell,
which at first is not larger than a ‘“‘birdshot,” but is gradually enlarged
to accommodate the slowly-increasing size of the inmate, little oppor-
tunity is afforded for much variation in mode of existence and habits.
The interesting features to be considered are the feeding and burrow-
ing habits, which together comprise the principal activities of its sub-
terranean existence.

THE FOOD OF THE LARVA AND PUPA.

The food taken by this insect beneath the soil is necessarily fluid, as
is also the case with the perfect insect, as well as with all other insects
of the order Hemiptera. That the Cicada should obtain its nourish-
ment in a manner different from the other members of its order would
not be anticipated, but, nevertheless, a good deal of difference of
opinion has been expressed as to the nature of the food of this insect
in its subterranean life, as also its method of feeding. Both Professor
Potter and Doctor Smith were of the opinion that the insect in its
underground life obtained its nourishment from the surface moisture
of the roots of plants through capillary hairs at the tip of the pro-
boscis—a curious misapprehension, as the hairs mentioned arise from
the sheaths, and have no connection with the true piercing and sucking
setz. Professor Potter expresses himself on this subject as follows:

In all places they are found attached to the tender fibrils of plants. When they
are disturbed or driven from them they seek for others the moment they are at liberty.
This is their only aliment, not the substance of the roots of the plants, which they can
not divide and comminute without teeth or jaws to use them, but the mere aerial
exhalation from their surface. This well-established fact would seem to account for
the slowness of their growth, and furnishes a reason for so long a subterraneous residence.

This absurd view of the method of nourishment of the larva and
pupa is on a par also with the belief of the same authors, reviving the
statement of Aristotle, that the adult insect subsists on “the dewy
exhalation of vegetable barks,” which was supposed to be swept up by
a brush of hairs on the tip of the proboscis. Doctor Smith claims a
basis for this theory of the feeding habits in perscnal observation, and
it has been supposed by others to be supported by the well-known
fact that the Cicada will occasionally issue from the ground that has
been practically cleared of timber and under cultivation for a number
of years, and that other species are known to issue from the prairies.
These facts lose much of their significance when it is remembered that
any vegetation, even annual, as of farm crops, would supply ample
root growth for the Cicada larva during the growing period of summer,
and in the colder months they undoubtedly lie dormant in their earthen
cells.

THE FOOD OF THE LARVA AND PUPA. 123

Perhaps the first writer to pomt out and demonstrate the true
method of feeding of the larva and pupa of this insect in their under-
eround existence was Miss Morris, of Germantown, Pa. That the
Cicada larve and pupz pierce small roots with their sucking beaks
and feed on the juices of the plant, as do other plant-feeding hemip-
terous insects, as their normal, if not their sole method of subsisting
was fully proved by her investigation, and has been confirmed repeat-
edly in the diggings made by the writer, and there can no longer be
any possibility of doubt in the matter. Jn practically every case, in
the writer’s experience, where the cell in which the larva rested was
taken out in condition for examination a small root, one-sixteenth to
three-sixteenths inch in diameter, was found to border usually the
upper end of the cell, and in several instances larve were found with
their beaks so securely embedded in the root that they were not easily
loosened. In other instances the roots showed, by the slight swell-
ing and reddish discoloration beneath the bark, unmistakable signs of
having been punctured.

The root-feeding habit can be best witnessed in light, rich soils, and
in the plantings of the brood of 1889 under oak trees on the Depart-
ment grounds the soil beneath these trees was so thickly inhabited
that between the depths of 6 and 12 inches every spadeful of earth
would throw out numbers of the larvee, and a most excellent opportu-
nity was afforded for the study of their habits. In hard, packed
soils, perhaps scantily supplied with roots, the difficulty of getting out
the cells in perfect condition is such that one might easily be led into
error, and the comparative rarity of the larvee in such soils adds fur-
ther to the difficulty of determining their feeding habits.

It is for this reason, I have no doubt, that the opinion has obtained
in some quarters that the larve subsist not on the roots of plants, but
on the nourishment obtained from the surface moisture of the roots,
or the general moisture of the earth, which might be supposed to
contain more or less nutrient material arising from the decomposition
of the vegetable matter. That the moisture of the surrounding soil
may, and doubtless does, supply the very delicate, thin-skinned larvee
and pupe with a certain amount of liquid by absorption through the
skin may be admitted, and in fact when the darve are taken from
their natural surroundings and exposed to the air they very rapidly
dry and shrivel. Larve are doubtless occasionally found in cells
away from roots, and this may be explained by the fact of their being
at that time either undergoing one of their long resting or hibernating
periods, which may be of frequent occurrence in such an extremely
long-lived species, or. they may be burrowing in search of roots on
which to subsist.

124 THE PERIODICAL CICADA,
THE LOCATION IN THE SOIL.

There has been great difference of opinion as to the depth beneath
the soil reached by the larve and pupe. In all of the extensive
excavations which have been made on the Department grounds in
following the results of the experimental plantings, specimens have
rarely been found at a greater depth than 2 feet below the surface and
usually between 6 and 12 inches, especially in the first years of the life
of the insect. This experience is corroborated by the examinations
made by Professor Riley in Missouri, and is fully confirmed by the
interesting manuscript notes left on this subject by Doctor Smith,
which are here reproduced:

The depth in the earth to which it descends depends upon that of the vegetable
soil, and its location is at the bottom of the soil, except perhaps in some of the deep
soils of the West and the alluvial soils, where the depth of its descent is probably
only sufficient to protect it against the inclemency of the weather. This is generally
from 12 to 18 inches and sometimes 2 feet. It never changes its locality from the
time it enters the earth till it emerges. The cells in which they shut themselves up
are, inside, well finished and smooth, of a sufficient size to accommodate them; but
outside they are mere lumps of clay and afford by their appearance no clew to their
internal character. It is this fact that has caused all the doubt and mystery about
their place of residence and habits during their long continuance in the earth. A
gentleman in the winter of 1850-51 was excavating on the side of a low hill for the
purpose of building a wall on West Baltimore street. The excavation was about 150
yards long and 6 to 18 feet deep to the level of the paved street. This hill had been
covered in former years with trees and shrubbery, and had been one of the fields of
observation in 1834. I watched this excavation daily and found the cells of the
locusts thrown down in the greatest abundance. The lumps of earth containing
the cells would roll down the heaps of earth just as others did, affording not the
slightest indication of their internal contents. But as the pick or the spade of the
workmen struck a cell in its place in the banks it readily broke open and the larva
was exposed. When the excavation was completed the observer standing in the
street had a fine view of the broken cells in the bank. From one end of the bank
to the other the cells were plainly visible, appearing like small augur holes, and all in
a regular stratum of earth about 18 inches below the surface of the earth, from 2 to 4
or 5 inches apart, and none more than 1 or 2 inches higher or lower than the others.
The internal size of the cells was from 14 to 2 inches long and about three-fourths of
an inch wide, forming an oblong cavity very smooth in its walls. The particles of
earth of which the cells were composed had evidently been agglutinated together
by some viscid fluid secreted by the insect. This is their habitation during the whole
seventeen years, or until they prepare for their ascent.

In the-face of the testimony given above there are records also by
apparently trustworthy observers which seem to indicate that the
larve are capable of gomg to much greater depths. An instance of
this sort is reported by Mr. Sadorus, of Port Byron, Ill., who built a
house in 1853 and found that they came up in his cellar in 1854.
Others have reported finding them at a depth of 10 feet or even more
below the surface. A rather remarkable instance is recorded by Mr.
Henry C. Snavely, of Lebanon, Pa., in which the Cicada pup are
reported to have worked their way through a hard mass of cinders
about 5 feet in thickness, which had been firmly compacted.

METHOD OF BURROWING. 125

It is difficult to say how many of these reported occurrences at
unusual depths are due to an unobserved tumbling of specimens from
higher levels, but where the insects have been observed to issue through
the bottom of cellars or similar situations the information would seem
to be reliable. The fact remains, however, that all of the extensive
diggings in the investigation of the early history of this insect here in
Washington and elsewhere have confirmed the statements of Doctor
Smith; in other words, the insects have always been found, as stated,
within 2 feet of the surface and in greatest numbers between the
depths of 8 and 18 inches.

A curious feature in connection with the underground life of this
insect is its apparent ability to survive without injury in soil which
may have been flooded for a considerable period. Doctor Smith
records a case of this kind where a gentleman in Louisiana in January,
1818, built a milldam, thus overflowing some land. In March of the
following year the water was drawn off and ‘in removing a hard bed
of pipe clay that had been covered with water all of this time some 6
feet deep the locusts were found in a fine, healthy state, ready to make
their appearance above ground, that being the year of their regular
appearance.’’ Another case almost exactly similar is reported by
Mr. Barlow. In this instance the building of a dam resulted in the
submerging of the ground about an oak tree during several months
of every summer, ultimately resulting in the death of the tree. This
went on for several years, until the dam was washed away by a
freshet, when digging beneath the tree led to the discovery of the
Cicada larve in apparently healthy condition from 12 to 18 inches
below the natural surface of the ground. In both of these instances
the ground may have been nearly impervious, so that the water did
not reach the insects nor entirely kill all of the root growth in the
submerged soil.

THE METHOD OF BURROWING.

The actions of the Cicada beneath the soil are not readily investi-
gated, the newly hatched and more active individuals disappearing
rather rapidly and seeming to be quite at home in the earth, as their
natural element. The method of burrowing of the larger and partly
grown specimens, as witnessed in captivity under fairly natural condi-
tions, is, as has been described in the manuscript notes of the Bureau,
as follows: The larva scratches away the walls of its cell with the
femoral and tibial claws, grasping and tearing the earth and small
stones just as one would do with the hands, bracing itself against the
sides of its cell mainly by its hind and middle legs, the former in their
natural position and the latter stretched out over the back. If it is
rising, so that the earth removed naturally falls to the lower end of
the burrow, it simply presses the detached portions on all sides, and

126 THE PERIODICAL CICADA,

especially on the end of the cavity, by means of its abdomen and
middle and hind legs. If the direction of the larva, however, is down-
ward, the loose soil has to be gathered and pressed against the upper
end of the cavity, which is accomplished by making the soil into little
pellets by means particularly of the front femora and placing these
pellets on the clypeal part of the head, carrying them upward and
pressing them firmly against the top of the cavity. The stiff hairs
that cover the head and border the inner side of the fore tibize and
femora assist very materially in securing the earth while it is being
transported. .

From time to time the burrowing insect rests and cleans the’
adhering earth from its forelegs very much as a cat washes its face
with its paws. The large, strong forelegs are moved over the rough-
ened front of the head, the stiff hairs springing from the latter acting
like a comb or brush to free the spines of adhering earth.

DAMAGE OCCASIONED BY LARVZ AND PUP.

During its underground life the Cicada has been charged with
damaging, and even killing, fruit trees. At first thought this is not
an unnatural inference when one remembers the immense numbers in
which the insect often occurs. The most specific charge brought
against them in this particular is the account published by Miss
Morris in 1846.“ Miss Morris having suspected for a number of years
that the failure of certain fruit trees over 20 years old was mainly due
to the ravages of the larvee of the periodical Cicada, had an examina-
tion made of one of them, a pear tree that had been declining for a
number of years without apparent cause. She says:

Agreeably to my expectation I found the larvee of the Cicada in countless numbers
clinging to the roots of the tree, with their suckers piercing the bark and so deeply
and firmly placed that they remained hanging for a half an hour after being removed
from the earth. From a root a yard long and about an inch in diameter I gathered
23 larvee; they were of various sizes, from a quarter of an inch to an inch in length.
They were on all the roots that grew deeper than 6 inches below the surface. The
roots were unhealthy, and bore the appearance of external injury from small
punctures. On removing the outer coat of bark this appearance increased, leaving
no doubt as to the cause of the disease.

In this particular instance there is some reason for believing that
the damage to the tree had been caused by the larve. The fact
remains, however, that no damage has ever been detected in forests,
where the Cicada emerges in countless myriads, the trees presenting
as vigorous and robust a condition as in other districts where no
cicadas occur, and this is true also of old original trees and planted
trees in parks or private grounds. In orchards also where the insects
have been so abundant that the ground was almost honeycombed after

a Proc, Acad. Nat. Sci. Phila., December, 1846 (1848), vol. 3, p. 133.

NATURAL ENEMIES. 127

their emergence the trees themselves exhibited a good state of vigor
and an inspection of the roots revealed no material injury save some
small swellings or callosities with slight discoloration which might
have resulted from the punctures.

The underground development of the Cicada is so very slow, thirteen
or seventeen years being occupied in attaining a size which with other
species is achieved in as many days or weeks, that the very slow
absorption of nutriment from the roots can scarcely have any effect
on them, and the only injury, and this is very slight, is probably due
to a poisoning of the roots, perhaps by the beak of the insect, as
indicated by the slight discoloration of the cambium at the point of
puncture. Callosities and other irregularities are, however, rare,
and have never been observed by the writer. Very often also there
are, undoubtedly, long periods of rest or dormancy, during which no
food at all is taken.

Referring to the injury noted by Miss Morris, it is a well-known
fact that fruit trees have a natural term of life, and after twenty years
they are very apt to show weakness and loss of vigor, and cease to be
profitable. It is possible, therefore, that this is the true explanation
of the condition of the trees noted by her rather than that it was due
to the presence of the larve of the Cicada.

THE NATURAL ENEMIES OF THE CICADA.

The fact that the periodical Cicada appears above ground so rarely
prevents its having any peculiar or specific parasitic or natural
enemies. We can not conceive of any parasite breeding solely, either
in the adult Cicada or in its eggs which could persist during the long
period of years when no host was available. Equally remarkable
also would be a parasitic insect the term of whose life should be so
extended that it could live in the body of the Cicada larva during
the years of its slow growth beneath the soil. Of the larger enemies
‘of the Cicada, such as birds and mammals, the habit of feeding on
the Cicada is necessarily acquired anew with each recurrence of a
Cicada year.

All these facts have a very potent influence in protecting the peri-
odical Cicada, which, as we have already pointed out, is particularly
helpless, and were it not for these natural protective influences the
very existence of the species would probably be early brought to
an end.

During their subterranean existence, the larve and pupxe, when
near the surface, are doubtless subject to the attacks of various
predaceous coleopterous larve, and many of them are unquestionably
destroyed by this agency. Upon leaving the ground to transform
they present an attractive food for many insectivorous animals, and
the pup and transforming adults are vigorously attacked by many

128 THE PERIODICAL CICADA,

different reptiles, mammals, and birds, and by cannibal insects,
such as ground beetles, dragon-flies, soldier-bugs, etc., while such
domestic animals as hogs and poultry of all kinds greedily feast upon
them. The preference shown by hogs running wild in woods for the
Cicada is especially marked, and we have elsewhere commented on
the fact of their rooting up the ground to get the pupx in April and
May, before the cicadas have appeared at the surface of the ground
for transformation. The birds are, perhaps, the most efficient
destroyers of the Cicada, and, as we have already noted, the English
sparrow is particularly destructive to them in and near cities, and,
indeed, bids fair to completely exterminate them in such locations.¢

In the perfect state they are attacked by at least one parasitic fly
(Tachina sp.), which lives internally in the body of its host. One of
the large digger wasps, to be later described, also preys upon the
adult, provisioning its larval galleries with the stung and dormant
cicadas. The Cicada is also attacked by a fungous disease, some-
times so abundantly as ultimately to destroy most of the male and
many of the female insects.

In the egg state the Cicada has many very effective enemies, com-
prising mainly parasitic flies belonging to the orders Hymenoptera
and Diptera, and also various predaceous insects belonging to the
orders Hemiptera, Neuroptera, and Coleoptera. A number of well-
known predaceous mites, and other mites whose habits seem to be
predaceous in this particular, are also found associated with the eggs
of the Cicada under such circumstances as to leave little doubt of their
feeding upon the eggs. All of these insect and mite enemies of the
Cicada are more or less general feeders, and are simply attracted in
numbers to the Cicada, and especially to the eggs in the case of the
egg parasites, on account of the abundance of the food presented. In
other words, we are furnished with a striking example merely of ready
adaptation to new and favorable conditions. This is true also of the
fungous disease of the Cicada, which is probably normally present in
other species of Cicada which are annual in appearance.

a This is well illustrated by the following experience in 1902 (Brood X) in the city
of Washington, as recorded by the writer: ‘Within the city very few of the cicadas
which came out survived more than a few hours, being quickly snapped up and
destroyed by the English sparrow. The numbers within the city were greatly dimin-
ished by the English sparrow at the appearance seventeen years ago, the destruction
by this bird at that time having been noted by Professor Riley and others to be very
considerable. The sparrows’ work this year, however, was much more effective,
the cicadas being fewer in numbers; and I doubt whether a single individual, certainly
very few, ever reached the egg-laying period. For two or three days in the midst
of the trees on the Museum grounds a few song notes were heard, but ceased very
soon. In the woods in the country about the city, especially out toward Chevy
Chase, the Cicada appeared in very considerable numbers, and here did not suffer
very much from the attacks of birds, and for the most part went through the, normal
aerial existence successfully.’’ (Proc. Ent. Soc. Washington, V, 1903, p. 24.)

DIPTEROUS ENEMIES. 129
INSECT PARASITES.

As already noted, among the more effective natural enemies of the
Cicada are the other insects which prey upon the eggs in the twigs, on
the newly-hatched larvee, and also, but to a much less extent, on the
adults. The more common and characteristic of the insect enemies of
the different stages of the periodical Cicada are given below:

DierrRous ENEMIES.

Some four species of two-winged flies have been found to subsist as
larvee on the eggs of the Cicada, but none of these has been reared
to the adult stage and, therefore, their specific identification is impos-
sible.

One of these bears some resemblance to an asilid, or, perhaps, more
remotely, to a bombylid larva, and was found by Mr. E. W. Allis at
Adrian, Mich., feeding on the contents of
the eggs of the Cicada, piercing the thin
shells and extracting the juices. These
larve are very minute, not much exceeding
a millimeter in length.

The most interesting of the dipterous egg
parasites is a cecidomyiid, which was found
in February, 1886, with eggs deposited in
sumac the previous season. When ex-
amined, all the eggs had hatched except in
some instances where they had been sealed
up by the rapid growth of the wood, so as to
prevent the escape of the larve. One of the Wis cosnacaaie ra eae
eggs thus inclosed was of an orange color, © of the periodical Cicada: Larva,
in distinction from the normal yellowish- = ™¢h enlarged, with anatomical

: q > details at side. (Original.)
white, and from it, on March 2, an orange-
colored cecidomyiid larva emerged. Other larve, apparently of the
same species, were secured in May from eggs in alder twigs. From
none of these, however, were adult flies obtained. The larve ranged
in length from 1 to 1.5 millimeters. Their general characteristics
are indicated in the accompanying illustration (fig. 52).

The fly parasite of the adult C’cada seems to belong to the family
Tachinide, which includes a number of species similarly attacking
grasshoppers as well as many other insects. The larve of these flies,
which have not been carried to the adult stage, sometimes to the
number of half-dozen or more, will occur together in the body of a
Cicada, which they have almost or quite completely eaten out.

51117—No. 71—07——9

130 THE PERIODICAL CICADA,
Hemiererous ENEMIES.

A few predaceous Hemiptera were found associated with Cicada
eggs under such circumstances as to leave little doubt but that they
were subsisting on them. Among these were two species of Thrips,
which were found both in the larval and adult stages in several
instances about the eggs on
which they had been feeding.
The material that has been
preserved of these Thrips is
not now in condition to be
worked up. Both species are
probably undescribed.

HyMeENorPreROoUS ENEMIES.

The hymenopterous ene-
mies of the Cicada comprise
Vie. 53.—Egg parasite, Lathromeris cicadx: a, female; a number of ese parasites,

b, abdomen of male; c, antenna of male; d, antenna of which are the more unportant

female. All greatly enlarged (original.) : : Cee Cyne ee

agencies in limiting the num-
bers of the insect, and the large digger wasp already mentioned.
The fact that the eggs and the newly hatched larve are much sought
after by various species of ants was early commented upon, Doctor
Potter stating that they are constantly infested by legions of ants,
both before and after they are
hatched. He says:

Even the little red species, the most
diminutive of the race, will shoulder the
eggs and the young and bear them off
to their cells. Inall our researches we
found them in battalions systematically
arrayed for wholesale plunder and de-
vastation.

Doctor Smith corroborates
Professor Potter, stating that he
has himself observed a small red
ant, scarcely as large as its in-
tended victim (a young Cicada
lat NE ); sees the latter, shoulder Fic. 54.—Female Sphecius carrying a Cicada to her
nibs and start off ata great speed. burrow. Natural size (after Riley).

THE PARASITES OF THE EGGS.

Several egg parasites were reared from the eges of the Cicada, but
with one exception were not abundant in the course of extensive
breedings. Single individuals were secured of a mymarid, a tricho-

PARASITES OF THE EGGS. 131

grammid, and two chalcidids. The excepted species, however, has
been reported as occurring in enormous numbers, and warrants a
more careful account.

Attention seems to have been first called to this parasite by Mr.
William T. Hartman in a letter dated October 5, 1868, to Doctor
Walsh. In this Mr. Hartman states that in getting some twigs, from

Fic. 55.—Burrows of Sphecius speciosus: e, e, ¢, main entrance; c, c, c, c, chambers for larve and their
food. Greatly reduced (after Riley).

which he hoped to obtain the larve of the Cicada, from an oak which
had been very thickly oviposited in, he found, after leaving the tree,
that his head and clothes were covered with what seemed to be small
red flies. The branches secured were kept in his office for several days
and the little red flies appeared again in
(7 countless numbers. The examination of
these flies under a microscope showed that
they were minute Hymenoptera instead
of Diptera, as he first supposed. He ob-
tained very few larve of the Cicada from
these shoots, and consequently inferred
that practically all of the eggs had been
parasitized by this insect. He states also
that a neighbor of his trapped thousands
of them in the soft paint which had been
newly applied to his window shutters, and
that by the middle of August this minute
parasite was ‘‘everywhere in force.”
What is probably the same insect (fig.
yA 53) was reared in some egg-infested twigs
Fig. 56.—Adult Cicada with Sphecius Collected by Mr. T. Pergande in Virginia
ee tacts ata. Naturalsize(aé in July, 1885. Doctor Howard has exam-
ined these specimens, and pronounces them
to be a new species of a European genus not hitherto recorded from
this continent, and has described them under the name Lathromeris
cicadx.* The life cycle of this minute parasite is evidently so short
that it is possible for it to pass through two or three generations

@Oanadian Entom., vol. 30, April, 1898, pp. 102, 103.

132 THE PERIODICAL CICADA,

within the egg period of seven or eight weeks of the Cicada, and this
accounts for its excessive multiplication, as described by Mr. Hart-
man, and probably makes it wherever it occurs one of the most
efficient agencies in keeping the Cicada in check.

THE LARGER DIGGER WASP.

I have already referred to the probability of the larger digger wasp
(Sphecius svneciosus Dru.) preying on belated individuals of the period-
ical Cicada. That
the bulk of the
brood has disap-
peared, however,
before this wasp
becomes at all
abundant has
been often pointed
out and is not to

be questioned, and
Fig. 57.—Cicada in burrow of Sphecius, with full-grown larva of latter it is well known

feeding. Natural size (after Riley). hat the toast ae
its work is with the later-appearing dog-day harvest fly (Cicada
tibicen Li.). With the assistance of Mr. Pergande and the writer, Pro-
fessor Riley worked out the natural history of this wasp in detail in
its relation to the dog-day harvest fly, and published a full illus-

Fig. 58.—Sphecius speciosus: a, larva; b, pupa, from below; c, same, from side, natural size; e, head
of larva; f, labium of same; g, maxilla of same. Enlarged (after Riley).

trated account of the species.? Its life habits when it preys on the
periodical Cicada are identical with its habits with the dog-day
species or any other annual Cicada with which it may store its bur-
rows. <A brief account of the habits of this wasp is here reproduced,
together with the figures illustrating its very curious and interesting
life stages. (See figs. 54-60.)

a Insect Life, Vol. IV, March, 1892, pp. 248-252.

THE LARGER DIGGER WASP. iss

This wasp and its near allies are the natural and perhaps the most
destructive of the insect enemies of the adults of the different species
of the Cicada, and their operations are often witnessed and are com-
mented upon in print nearly every season. In fact, no more curious
and interesting illustration of the wars which take place in the insect
world is afforded than the sight of one of these wasps seizing its victim
and silencing and paralyzing it with a sting, which, while throwing it
into a comatose condition from which it never recovers and suspending
or greatly reducing its vital functions, does not actually kill it, but
leaves it an unresisting, living prey for the delicate wasp larva.

The fact that some tragedy is being enacted is often brought to the
attention of the observer by the sudden cessation of the regular song
note of the unsuspecting Cicada. The song ends in a sharp cry of
distress, and if one is in position to witness the struggle the wasp may
be seen grasping its
victim and endeay-
oring to take flight,
the quick thrust of
its sting having al-
most immediately
quieted the Cicada.
Very often in the
first struggle the
wasp and the Ci-
cada fall to the
ground together,
and it 1s necessary
for ‘the former la-. .
boriously to cimb <*%%
the tree again, drag- Fia. 59.—Larva of Sphecius sp
ging the Cicada
with it, in order to take flight from an elevated point, the Cicada being
usually much heavier than the wasp and bearing the latter slowly
to the ground as it flies. For this reason it often becomes necessary
for the wasp to carry the Cicada several times up into near-by trees,
making repeated short flights before it reaches its burrow.

The latter is excavated with great activity by the wasp, the drier
and more elevated situations being usually chosen. The burrow
ranges from 18 inches to 2 or 3 feet in length and has three or four or
more branches of from 6 inches to a foot in length, each terminating
in a little oval chamber. Within each of these chambers is stored a
Cicada to which a single wasp egg is attached in such manner as to be
covered and protected by one of the middle legs of the Cicada.

The parasitic larva on hatching merely protrudes its head and
makes an opening into the body of its host at some suture where

Re re

inning its coccon. Natural size (after
Riley).

134 THE PBRRIODICAL CICADA,

entrance is easy, and slowly feeds on the soft, juicy interior. The
larva remains outside of the Cicada throughout its life, but by means
of its very extensile anterior segments, or neck, thrusts its small head
throughout the interior of the Cicada and gradually exhausts the soft
parts until the Cicada becomes a mere broken
shell. The wasp larva increases in size very
rapidly, ultimately attaining a length of 14 to 2
inches. It is then nearly white in color, with
the head and mouth parts remarkably well
developed and the anterior segments narrowed
and capable of very great extension. The whole
transformation from the egg to the full-grown
larva is comprised in a very brief period, the egg
hatching after two or three days and the larval
life not much exceeding a week.

Pe Ma eee yr When fully grown the larva constructs a co-
natural size; b, enlarged sec- Coon in a very peculiar manner. First a cylin-
Mon/of pore. (tery) der, open at both ends, is formed of earth awit

enough silk incorporated to form a rather dense and tough pod.

When the cocoon is nearly completed the ends are capped, and the

larva remains unchanged over winter and transforms to a pupa in the

spring or early summer shortly before the appearance of the mature

Fia. 61.—Mite egg parasite, Oribatella sp. Fic. 62.—Mite egg parasite, Oripoda
(Author’s illustration). elongala. (Author’s illustration).

insect. About the center of the cocoon are a number of very curious
structures which may serve as breathing pores until the larva has
become accustomed to its new conditions, since they are ultimately
sealed over, as represented in the illustration (fig. 60, 0).

MITE PARASITES OF THE EGGS. 135

Most of the fossorial wasps have habits very similar to this species,
but many of the other genera provision their nests with the larve of
Lepidoptera or with Orthoptera or sometimes with the larger spiders.

Mire PARASITES OF THE EGGs.

Of the mites found either preying on the eggs of the Cicada or asso-
ciated with them in such manner as to suggest a predaceous habit,
several represent species which are well known to subsist on soft-
bodied insects or other animal food. An almost equal number, how-
ever, belong to a family of mites, the Oribatide, which, so far as the
habits of the species are known, comprises, with few exceptions,
strictly herbivorous mites,
or such as subsist on vegeta-
ble decay. A few species,
however, of this family pos-
sess mouth structures which
indicate that they usually
prey on other insects, and
somc of them are known to
feed on decaying animal sub-
stances. In this country
two species have been re-
corded as being true insect
parasites, namely, Nothrus
ovivorus Pack. and Oribata
aspidiot, Ashm., the former
having been observed to
suck the eggs of the canker-
worm, and the latter to feed
on scale insects in Florida.
The types of these two spe-
cies have not been preserved,
and there is some doubt as
to their correct reference.

All of the mites associated with the eggs of the Cicada, both those
of doubtful and those of well-known predaceous habits, were invari-
ably found in the egg slits, down among the woody fibers, where
they could have little choice of food except that supplied by the
Cicada eggs. In no case were the mites actually observed to be
feeding on the eggs, but frequently the eggs were more or less shriveled
and the contents extracted.

All of the mites referred to below have been examined for me by
Mr. Nathan Banks, a specialist in this group, who has identified and
described the material as far as its condition, as balsam mounts, per-
mits. The accompanying illustrations are from very careful drawings

Fia. 63.—Mite egg parasite, Oppia pilosa. (Author’s illus-
tration).

136

THE PERIODICAL CICADA,

made several years since by Mr. Pergande, who collected several of
the mites and mounted and made preliminary studies of the others.

Fie. 64.—Mite egg parasite,
Pediculoides ventricosus.

Much of the material was collected by Mr.
E. W. Allis at Adrian, Mich., in 1885, the balance
by Mr. Pergande in the District of Columbia and
near-by Cicada districts in Virginia in the same
year.

THE ORIBATID MITES.

The members of the family Oribatidxe have
the popular designation of “beetle mites,”’ aris-
ing from their possessing a hard chitinous cover-
ing causing them to resemble minute beetles.
Some six distinct species were found in the adult
stage associated with the eggs of the Cicada, and
several nymphal forms—the latter being often
showily colored and the principal feeding stage
of these mites.

(Author’s illustration.)

The following are Mr. Banks’s determinations

of the oribatid material: (1) Oribata sp., collected by Mr. Pergande in

the District of Columbia in July, 1885

lected by Mr. EK. W. Allis at Adrian,
Mich., in October, 1885; (3) Oripoda
elongata Bks., MS. (fig. 62), collected
with the last; (4) Oppia pilosa Bks.
(fig. 63), also collected at Adrian,
Mich.; (5) Oribatula sp., collected
by Mr. Pergande in the District
of Columbia and in Virginia in
July, 1885; (6) Oribatid nymphs,
collected with the last and possi-
bly belonging to the same species;
(7) Hoplophora sp., collected by
Mr. Allis in Michigan in October,
1885.

MISCELLANEOUS PREDACEOUS MITES.

The following mites have well-
known predaceous habits and for
the most part are miscellaneous
feeders, subsisting on almost any
available animal matter, such as

; (2) Oribatella sp. (fig. 61), col-

Fic. 65.—Mite egg parasite, Tyroglyphus cocci-
philus. (Author’s illustration.)

soft-bodied insects, insect eggs, and various animal and also 7 eee

food products.

MISCELLANEOUS PREDACEOUS MITES. ley,

Perhaps the mite most commonly found with the eggs of the Cicada
is Pediculoides ventricosus Newp. ‘This species has a very general feed-
ing habit and is often an active agent in the destruction of the eggs or
young of insect pests. In breeding cages it is often a nuisance by
destroying the smaller insects being kept under observation. The
general form of the male and of the unimpregnated female of this
mite is similar to that of the next species listed. The gravid female,
however, develops an enormous globular extension from the tip of her
abdomen, as illustrated in the accompanying figure (fig. 64).

Another predaceous mite, not at all uncommon, in the egg slits of the
Cicada, both in the District of Columbia and in Michigan, is Tyroglyphus
cocciphilus Bks. (fig. 65), very near 7. longior Gerv., which species it

Fie. 67.—Cheyletus sp., mite egg
Fic. 66.—Mite egg parasite, [phis ovalis. parasite of Cicada. (Author’s
(Author’s illustration.) illustration.)

very closely resembles. The species named is a widely distributed one
and frequently occurs also in breeding cages, and often becomes very
troublesome from its presence in enorn.ous numbers on various food
substances in the larder. A smaller species of the same genus was
found with the Cicada eggs, but the material is not in good enough
condition to make its identification possible.

A species belonging to the family Gamasidz was found by Mr. Allis
associated with the eggs of the Cicada (fig. 66). It is apparently an
undescribed species and is certainly distinct from the half dozen
known from North America. Mr. Banks has suggested for it the name
Iphis ovalis. The family to which it belongs includes true insect
parasites which either live free or attached to their hosts, and there
is little doubt but that this mite was attracted by the Cicada eggs.

138 THE PERIODICAL CICADA.

Two mites, one belonging to the genus Cheyletus (fig. 67) and the
other to the genus Bdella (fig. 68), were found associated with the eggs
of the Cicada in Virginia in July, 1885. Both of these mites seem to
be undescribed, but the material is not in good enough condition to
warrant their description. Both genera are known to be carnivorous,
and the specimens secured had doubtless been preying on the Cicada
egos. :

THE VERTEBRATE ENEMIES.

Under this heading I will supplement merely the general state-
ments given elsewhere on the destruction of the Cicada by birds,
mammals, etc., by quoting the observations of Mr. A. W. Butler,
who devoted considerable attention to the natu-
ral enemies of the Cicada in 1885 in southeast-
ern Indiana. His lists and notes, which follow,
could be much extended and, if all the enemies
of the Cicada were known, would doubtless in-
clude all the insectivorous birds and mammals
occurring within the range of this insect. He
Says:

a Among birds the English sparrow, Passer domesticus
(Linn.), is perhaps its greatest enemy. Within one week
from the date of the appearance of the Cicada in Brookville
not one could be found, and I doubt if asingle specimen
was permitted to deposit its eggs, owing to the persistent
warfare waged by this garrulous sparrow.

Of native birds the robin, Merula migratoria (Linn.);

blackbird, Quiscalus quiscula xneus (Ridg.); catbird, Gale-
Fie, 68-—Bdella sp. mite Gscoptes carolinensis (Linn.); red-headed woodpecker,
parasite of eggs of Cicada. ¢ ¢ :

Greatly enlarged (author’s “Zelanerpes erythrocephalus (Linn.); flicker, Colaptes auratus

illustration). luteus Bangs; towhee, Pipilo erythrophthalmus (Linn.), and

orchard oriole, Icterus spurius (Linn.) were their greatest
enemies, Food of every other sort appeared to be neglected in order that they might
feast for a limited period upon the easily captured Cicada.

Of other birds examined, the following contained Cicada remains: Brown thrasher,
Toxostoma rufum (Linn.); Baltimore oriole, Jcterus galbula (Linn.); scarlet tanager,
Piranga erythromelas Vieill.; blue-gray gnatcatcher, Polioptila cxrulea (Linn.); worm-
eating warbler, Helmitherus vermivorus (Gmel.); purple martin, Progne subis (Linn.);
wood pewee, Horizopus virens (Linn.); wood thrush, Hylocichla mustelina (Gmel.);
yellow-throated vireo, Lanivireo flavifrons (Viell.); cardinal grosbeak, Cardinalis car-
dinalis (Linn.); tufted titmouse, Beolophus bicolor (Linn.); Carolina chickadee, Pen-
thestes carolinensis (Aud.); chipping sparrow, Spizella socialis (Wilson); downy wood-
pecker, Dryobates pubescens medianus (Swains); crested flycatcher, Myiarchus crinitus
(Linn.); indigo bird, Passerina cyanea (Linn.); cow bird, Molothrus ater (Bodd.);
white-bellied nuthatch, Sitta carolinensis Lath.; yellow-billed cuckoo, Coccyzus amer-

a The nomenclature of the birds and mammals mentioned in this extract has been
revised for this bulletin by Mr. Harry ©. Oberholser, of the Bureau of Biological Sur-
vey, U. 8. Department of Agriculture.

FUNGOUS DISEASE OF THE ADULTS. 139

icanus (Linn.); black-billed cuckoo, C. erythrophthalmus (Wilson); American gold-
finch, Astragalinus tristis (Linn.); crow, Corvus brachyrhynchos Brehm, and cedar
bird, Ampelis cedrorum (Vieill.).

But two specimens of all the birds examined showed no evidence of cicada-eating.
These were the cerulean warbler, Dendroica cerulea (Wilson), and the warbling vireo,
Vireosylva gilva (Vieill.). Most birds eat only the softer parts, but some species—the
robin, brown thrasher, towhee, and a few others—eat also the wings and legs, and
even occasionally the head.

I found fox squirrels, Sciurus rufiventer Geoffroy, eating them, the young showing
greater fondness for this food than did their parents. The ground squirrel, or chip-
munk, Tamias striatus Baird, was very fond of them. I have seen this mammal
climb to the highest limbs of an apple tree seeking cicadas.

When cicadas fell into our streams many of them became the prey of various species
of fish. Our fishermen complained of their inability to get fish to take the hook while
they were feeding upon this new food. The remains of this insect were found in black
bass, Micropterus salmoides Henshall; blue catfish, Jchthelurus punctatus Jordan, and
white sucker, Catostomus teres Le 8.

Rev. D. R. Moore, a valued fellow-worker, found two species of snails, Mesodon
exoleia Binn., and MV. elevata Say, feeding upon dead Cicadas. This fact was a great
surprise tome. But few instances were recorded of digger wasps killing these insects.
Stizus grandis Say 4 was the only species observed. Aside from the enemies men-
tioned above, there were many others to which I could not direct my attention. In
general, it may be said that beetles, spiders, and other insect enemies prey upon them
incessantly, while parasitic flies, scavenger beetles, and ants destroy great numbers of
their dead bodies.

THE FUNGOUS DISEASE OF THE ADULTS.

The peculiar fungous disease of the adult cicadas was noticed by
Dr. Joseph Leidy in the Proceedings of the Philadelphia Academy of
Sciences for 1851, page 235, and has since been described as Mas-
sospora cicadina by Prof. C. H. Peck. Mr. W. T. Hartman, of
West Chester, Pa., speaking of the occurrence of this fungus in 1851,
says:

The posterior part of the abdomen in a large number of male locusts was filled by a
greenish fungus. * * * The abdomen of the infected males was usually inflated,
dry, and brittle, and totally dead while the insect was yet flying about. Upon break-

ing off the hind part of the abdomen, the dust-like spores would fly as from a small
puffball.

One male specimen, received in 1868 from Pennsylvania, was
affected by the same or a similar fungus, the internal parts of the
abdomen being converted into what appeared to be a brown mold.
R. H. Warder, of Cleves, Ohio, in speaking of this mold, says:

I found that in many cases the male organs of generation remained so firmly attached
to the female during copulation that the male could only disengage himself by break-
ing away and leaving one or two posterior joints attached to the female, and it is these
mutilated males which I found affected by the peculiar fungus mentioned, and there-
fore conclude that the dry rot might be the result of the broken membranes.

@ Synonymous with Sphecius speciosus Drury.
b Thirty-first Rept. N. Y. State Museum Nat. Hist., 1879, p. 44.

140 THE PERIODICAL CICADA,

It is well established, however, that both males and females are
affected by this disease, the former, however, in the greatest num-
bers, and that it is by no means confined to injured individuals.

Professor Peck describes this disease in general terms as follows:

The fungus develops itself in the abdomen of the insect, and consists almost wholly
of a mass of pale-yellowish or clay-colored spores, which to the naked eye has the
appearance of a lump of whitish clay. The insects attacked by it become sluggish
and averse to flight, so that they can easily be taken by hand. After a time some of
the posterior rings of the abdomen fall away, revealing the fungus within. Strange as
it may seem, the insect may, and sometimes does, live for a time in this condition.
Though it is not killed at once, it is manifestly incapacitated for propagation, and
therefore the fungus may be said to prevent to some extent the injury that would

otherwise be done to the trees by these insects in the deposition of their eggs. For

the same reason, the insects of the next generation must be less numerous than they
otherwise would be, so that the fungus may be regarded as a beneficial one. In
Columbia County the disease prevailed to a considerable extent. Along the line of
the railroad between Catskill and Livingston stations many dead cicadas were found,
not a few of which were filled by the fungoid mass.4@

Professor Peck was not able to satisfy himself as to the time when
the Cicada is attacked by this fungus, suggesting the possibility of its
having entered the ground with the larva and slowly developed with
its host, or perhaps entering the body of the pupa at the moment that
it emerges from the ground, with the third possibility of its developing
annually in the cicadas which appear every year, and becoming much
more abundant, and therefore noticeable, in the years of the appear-
ance of the great swarms of periodical cicadas. The latter supposi-
tion is unquestionably the correct explanation. Mr. A. W. Butler
refers to this disease at some length in his notes on the Cicada in
southern Indiana in 1885, and is of the opinion that nearly all of the
male cicadas which are not killed by birds and other enemies ulti-
mately succumb to this disease.

REMEDIES AND PREVENTIVES.
THE GENERAL CHARACTER OF THE PROBLEM,

In discussing this subject it is well to be again reminded that the
fears aroused by the presence of this insect when in great numbers are
unquestionably out of all proportion to the real damage likely to be
done. While they are most abundant in old and undisturbed forest
tracts and confine their work for the most part to forest trees, it is true
also that in parks and lawns, especially such as contain trees of the
original forest growth or their natural and immediate successors, the
cicadas sometimes appear in scarcely diminished numbers. This is
true also of orchards located on cleared lands or in the vicinity of
standing forests, and under such circumstances instances of serious or
fatal results to cherished plants or fruit trees are not uncommon.

2 Loc. cit., pp. 19,20.

COLLECTION OF ADULTS. 141

Notwithstanding the occasional instances of serious injury by the
Cicada, it is probably still true that there is no other important
injurious insect in this country that is responsible for so little serious
damage in proportion to the fears aroused, and yet every recurrence of
this insect calls forth the most anxious demands for means of control
or extermination. The exploitation of the facts concerning this insect
is, therefore, more to allay such fears, and to supply the desire for
information concerning it which its presence always arouses, than
from the necessity of detailing elaborate precautionary measures.

It is, nevertheless, important to know what may be done in the way
of protection and control whenever occasion arises to make such
action necessary, as for the protection of young fruit trees which are
especially exposed to injury or trees and shrubs over limited areas, as
in parks and lawns.

Precautionary operations are necessarily against the adults chiefly,
as being the authors of the greater damage. Against the larve and
pupe in their subterranean life it is hardly worth while to take any
action unless it be deemed desirable to attempt to exterminate a brood
within a given territory or bit of woodland, in which case the remedies
commonly employed against other subterranean insects, such as the
Phylloxera or other root lice, will serve for this insect equally well,
especially in the first year or two of its existence.

The prevention of injury from the Cicada includes, therefore, (1)
methods of destroying the emerged insects, either mechanically or by
insecticide applications; (2) applications to the plant to prevent
oviposition; (3) certain precautionary measures which may be taken
to lessen injury; and (4) operations to destroy the larval and pupal
stages in the soil.

MEANS OF DESTROYING THE EMERGED PUPZ AND ADULTS.
COLLECTION OF ADULTS.

In some instances the hand collection of the insects is feasible and
will prevent damage. This method necessitates the continual driving
of the insects from the plants by fighting or collecting them in umbrel-
las or bags in the early morning or date evening when they are some-
what torpid and sluggish. If undertaken at the first appearance of
the Cicada and repeated each day, the work of control will be facili-
tated by the fact that most of the insects will be en the young trees or
shrubbery or on the lower branches of larger trees and within com-
paratively easy reach.

An instance of this kind of work is recorded by Mr. Abner Hoopes,
of West Chester, Pa.* The work reported was for the protection of
nursery stock on the edge of woods from the attack of Brood X in

4 Entomological News, Vol. XVIII, March, 1907, pp. 108, 109.

142 THE PERIODICAL CICADA,

1902. There were 240,000 peach trees in the field to be protected,
and seven men were kept at work in this field for over two weeks, and
by actual count it was found that these men killed more than 1,000
cicadas each per day by hand collecting. Seventy thousand cicadas
were collected in this field alone, and other men were employed in the
smaller fields. so that Mr. Hoopes feels sure that at least 100,000
were killed altozether. In spite of this work, however, a loss of 12,000
trees was sustained out of the 240,000.

DESTRUCTION WITH INSECTICIDES.

The various treatments aiming at the destruction of the insects
themselves lrave yielded satisfactory results, but to have any practi-
cal value it is necessary to continue them daily or as long as the
insects issue in any numbers. On a large scale, therefore, or over a
considerable territory, in the presence of immense swarms, work of
this sort will be ordinarily out of the question. The recommenda-
tions apply particularly, therefore, to small areas or orchards. Such
work may be directed against the Cicada the moment it emerges from
the ground, while still in the pupal stage, but perhaps more readily
and successfully against the insect after it has shed its pupal skin and
is still soft and comparatively helpless, and with less ease, but still
with some degree of effectiveness, after it has hardened and begun
its aerial duties.

Of the many substances experimented with few proved to be of
much value, the best results being obtained with (1) pyrethrum or
insect powder, using it both in the dry form and as an aqueous solu-
tion; (2) kerosene emulsions; and (3) solutions of various acids.
These substances either effected the immediate death of the insect,
or attained this end indirectly by preventing its transformation from
the pupal to the adult stage; in other words, rendering the last molt,
impossible.

Pyrethrum powder is a perfectly satisfactory destroyer of the newly
transformed and soft cicadas, and has considerable efficacy against
the mature and hardened individuals. The best results are obtained
in the morning, before the insects have gained full strength to ascend
and while the plants are still wet with dew. The powder may be
puffed on the insects while clinging to shrubbery or on the lower
branches of the larger trees.

Pyrethrum powder is absolutely worthless against the pupz, which
even when thoroughly coated with it, will often succeed in casting off
their powdered skins and escape uninjured. The winged insects are,
however, very sensitive to the powder, and after an Siplieoion soon
show signs of uneasiness and in the course of a few hours fall helpless
to the Saatchi. where, though they may continue to have the power of
motion for a day or more, a fatal termination is almost sure to follow.

APPLICATIONS TO PREVENT OVIPOSITION. 143

The pyrethrum and water mixture is prepared by stirring up as
much of the powder as the water will hoid in suspension, or a little
milk may be added to increase the holding power of the water. The
results obtained with pyrethrum in water against the transformed
insects are as satisfactory as with the dry powder, with the additional
advantage of its being possible to throw the water by force pumps to
parts of the plant where it would be difficult to place the powder.
Against the pup, the water solution is more effective than the
powder, but is less so than kerosene emulsion.

Kerosene emulsion, as an application for destroying the emerged
pupe and adults, is used in very strong solution, or at a strength
ranging from one part of the emulsion to one of water up to a dilution
of the emulsion with eight parts of water. The greater strengths were
more immediate in their effects, but even with the more diluted washes
very satisfactory results have been obtained. The emulsion at once
stops all molting or transformation. Applied to the partly trans-
formed insects, the soft wings harden into shapeless masses, and while
occasional individuals may survive the treatment for two days or
more, the application is usually fatal in the end. The treated pupz
are unable to transform to the adult stage and they eventually die or
are devoured by their natural enemies. The death of the mature and
hardened insect is caused by closing its breathing pores with the oily
mixture, and in the case of the partly expanded or soft, immature
individuals by the caustic effect it has on the forming wings and soft
body.

The experiments with acids demonstrated also that exuviation may
be prevented by spraying the newly emerged pupa with a 2 per cent
solution of carbolic acid or a 15 per cent solution of acetic acid.

APPLICATIONS TO PREVENT OVIPOSITION.

All the early experiments with washes or other applications to
prevent oviposition proved unavailing except such protections as
could be applied to small trees or shrubs, such as covering them with
netting. Professor Riley in 1868, and later, at his instance, Dr.
W. S. Barnard, tested a number of repellent substances, such as
kerosene emulsion, vartous oils, and carbolic-acid solutions, all pun-
gent and disagreeably smelling substances, with results either unsat-
isfactory or of negative value.

In the occurrence in 1902 of Brood X some indications were obtained
showing the possible* protective value of lime washes. Mr. Slinger-
land reports that spraying a heavy coat of whitewash on the trees
will keep the locusts away to some extent when there are other
trees in the neighborhood. He states that the reason for this seems
to be that the insects do not like to sit on a white surface. The

144 THE PERIODICAL CICADA,

females will, however, oviposit on whitewashed twigs if there is no
other place for them.

The experience reported by Mr. W. B. Alwood * would seem to
indicate that the injury from the Cicada in orchards is prevented by
the use of Bordeaux mixture. The cicadas appeared in tull force in
a young orchard which had been sprayed with this mixture, but
practically all migrated elsewhere without ovipositing in the trees.
Other orchards near the one referred to by Mr. Alwood were badly
punctured. In view of this experience it may be that Bordeaux
mixture or the lime-sulphur wash will prove a valuable preventive of
injury from this insect.

PRECAUTIONARY MEASURES.

In view of the difficulty of controlling this insect on a large scale
after it has once emerged, it is well to adopt any precautionary
measures that may tend to lessen or distribute the injury. The
advent of all the large and well-recorded broods is commonly heralded
in advance in the local papers by State entomologists or other per-
sons who take interest in such recurrences. [orewarned in this way,
much injury and loss may be avoided by neglecting all pruning
operations during the winter and spring prior to the expected appear-
ance of the Cicada in order to offer a larger twig growth and distrib-
ute by this means the damage over a greater surface. Another pre-
caution, when a Cicada year is expected, is to defer the planting of
orchards, especially in the vicinity of old orchards or forest land,
until the danger is past. The same advice applies to budding or
erafting operations in the fall and spring prior to the Cicada’s appear-
ance. Much disappointment arising from injury to orchards or val-
uable nursery stock may thus be avoided. Vigorous young trees
will, it is true, often recover in three or four years from the effects
of a loss of or injury to a considerable percentage of their branches,
but it is difficult to overcome the unsymmetrical appearance which
will commonly result from the indiscriminate pruning caused by the
work of this insect, and the gnarled and scarified branches will long
bear testimony to the industry of the female insect.

Much of the injury occasioned by the cutting of the twigs by the
female Cicada in depositing her eggs can be remedied by subsequent
proper treatment of the wounded plant. In the case of old trees, the
main object to be secured is the rapid healing of the wounds and the
prevention of their being used as points of secondary attack by other
insects. The worst injured limbs in such trees should be cut out, so
that all the vigor of the plant may be directed to the remaining wood.
Any treatment also, as of thorough cultivation or the use of ferti-

@ Proc. 15th Ann. Mtg. Assn. Economic Entomologists, Bul. 40, Bur. Ent., p. 75.

MEANS AGAINST CICADA IN UNDERGROUND LIFE. 145

lizers, which will give the plant a more vigorous growth, will hasten
the healing process. With young trees the worst affected branches
should be removed, and the less injured ones protected from other
insects while they are healing by coating the wounded parts with
grafting wax or a moderately hard soap. These protective coverings
should be renewed at least once a year, preferably in the spring, until
the wounds are entirely healed over. In the case of a badly injured
tree that has been recently budded or grafted, it may be well to cut
it back nearly to the bud or graft, so that an entirely new top may
be made.

MEANS AGAINST THE CICADA IN ITS UNDERGROUND LIFE.

While it is probably true, as we have already stated, that the Cicada
in its underground life does not work any serious injury to plants on
account of the very insignificant amount of nutriment which it annu-
ally draws from the rootlets, nevertheless in exceptional cases, where
the ground is suspected of being very thickly populated with the larve
and pupe of this insect, it imay be deemed desirable to undertake their
extermination. This may be accomplished, as suggested, by using the
remedies ordinarily employed against other subterranean insects, such
as the Phylloxera and the apple root-aphis, with this difference, that
the poisons will have to be introduced more deeply into the soil
unless applied in the first or second year after the larve have begun
their development.

If taken in time, the number of the larve in the soil may be greatly
reduced by cutting off the branches of the trees which have been
thickly oviposited in, thus preventing the hatching of the eggs. It
will rarely, however, be possible to so completely eliminate the eggs
from the tree as to prevent the entrance of the larve into the soil in
considerable numbers.

Of the means employed against subterranean insects two are espe-
cially suitable for the destruction of the larve and pupe of the
Cicada—namely, bisulphid of carbon injected into the ground and
tobacco dust incorporated in the soil.

Tobacco dust has a manurial value and is not at all injurious to
plants. Its value against Cicada larve is purely theoretical, but there
is little doubt but that if it can be incorporated in the soil some dis-
tance below the surface—namely, by first removing 6 inches or more of
the top soil—it will effect the destruction of many of the delicate larvee
and pup of the cicadas. This dust is a waste product of tobacco
factories and costs about 1 cent per pound, and is worth nearly its cost
as a fertilizer.

Bisulphid of carbon, the popular French remedy for the grape root-
aphis, will undoubtedly prove an efficient means against the Cicada in

31117—No. 71—07——10

146 THE PERIODICAL CICADA,

its underground life. It will be necessary, however, except in the first
year or two of the existence of the larvae, to inject it to a depth of at
least 12 inches below the surface. It should not be introduced into the
soil closer to the crown of young plants than 14 feet, and not more than
an ounce of the chemical should be introduced into each hole, which
should be immediately closed. An injection should be made to about
every square yard of surface. The bisulphid rapidly evaporates and
penetrates throughout the soil, and is very deadly to insects. It is
highly inflammable, and should not, therefore, be poured from one
vessel to another near a fire. It may be introduced into the soil by
means of injecting machines. This treatment is not expensive, and
will be valuable for orchards, small groves, or private grounds.

THE PERIODICAL CICADA IN LITERATURE.

As would naturally be inferred of an insect as interesting as the
periodical Cicada, the writings which have been devoted to it from
the time of its tie coming to the attention of the colonists to the
present have been most voluminous in number and extent; much
of this literature, however, is of a fugitive character and scattered
through publications not now obtainable.

The earliest published account of the periodical Cicada which
has come under my own observation was brought to my attention
by Prof. E. A. Andrews, of the Johns Hopkins University, Baltimore,
Md. It is contained in Volume I, No. 8, page 137, of the Phas
sophical Transactions of the Royal Society of desman published
January 8, 1666, and is reported, unsigned, by the ‘‘ publisher,”
Henry Oldenburg. The portion of ume communication relating to
the Cicada is quoted below:

SoME OBSERVATIONS OF SWARMS OF STRANGE INSECTS AND THE MIscHIEFS DONE
BY THEM.

A great Observer, who hath lived long in New England, did, upon occasion, relate
toa Friend cf his in London, where he lately was, That some few years since there was
such a Swarm cf a certain sort of Insects in that English Colony, that for the space of 200
Miles they poyson’d and destroyed all the Trees of the Country; there being found
innumerable little holes in the ground, out of which those Insects broke forth in the
form of Maggots, which turned into F'lyes that had a kind of tail or sting, which they
stuck into the tree, and thereby envenomed and killed it. * * *

The rest of the article referred to a plague of locusts (grasshoppers)
in Russia, with which the Cicada is confused. The brood referred to
here is very likely No. XIV, which appeared in 1651. No other
brood coincides with this narrative and No. XIV not very closely,
but as the quotation states the relation was “upon occasion,” &nd
was ‘‘some few years since,’ there is ample warrant for assigning
the account to the brood of fifteen years before.

THE CICADA IN LITERATURE. 147

Prior to the discovery of the above record the earliest published
account known was that referred to in Bulletin 14 (new series) of
the Division of Entomology, page 112, given in a work entitled
‘“New England’s Memoriall,’’ by Nathaniel Moreton, printed at
Cambridge, Mass., in 1669. I was unable to get the work cited,
but an account seen by me was a quotation from it published in an
editorial note to an article on the ‘‘Locust of North America,’ in
Barton’s Medical and Physical Journal of 1804. The brood referred
to by Moreton is undoubtedly the same one referred to above, but
the occurrence of seventeen years previous. Moreton, publishing of
an event happening thirty-six years after it occurred, evidently
made a mistake of one year, the occurrence not being 1633, as stated
by him, but 1634. We have records of this brood in New England
from 1787 to 1906. The records, if any were made of it after 1651
and prior to 1787, have not been discovered.?

The quotation from Moreton referred to follows:

Speaking of a sickness which, in 1633, carried off many of the whites and Indians,
in and near to Plimouth [Plymouth], in Massachusetts, he says, ‘‘It is to be observed,
that the Spring before this Sickness, there was a numerous company of Flies, which,
were like for bigness unto Wasps or Bumble-Bees, they came out of little holes in the
ground, and did eat up the green things, and made such a constant yelling noise as
made all the woods ring of them, and ready to deaf the hearers; they were not any
of them heard or seen by the English in the Country before this time: But the Indians
told them that sickness would follow, and so it did, very hot in the months of June,
July and August of that Summer,” viz. 1633. He says, ‘‘Toward Winter the sick-
ness ceased;’’ and that it was ‘‘a kinde of a pestilent Feaver.’’—New England’s
Memoriall, &c., pp. 90 and 91.

The fact noted, namely, that the native Indians associated the recur-
rences of this insect with pestilential diseases, is interesting, as showing
that the Cicada had probably‘long been sake observation by them
and had exerted a vivid influence on their imaginations.

One of the earliest references on this continent to the periodical
Cicada is recorded in Steadman’s Library of American Literature, vol-
ume 1, pages 462-463. It is from the writings of an individual signing
himself “'T. M.,”’ supposed to have been Thomas Matthews, son of
Samuel Matthews, governor of Virginia. It was written in 1705, and
refers to three prodigies which are said to have appeared in that coun-
try about the year 1675,° and which, from the attending disasters,
were looked upon as ominous presages. One of these was the appear-
ance of a large comet; another, the flight of enormous flocks of
pigeons; and the last, relating evidently to the periodical Cicada, as
follows:

The third strange appearance was swarms of flies about an inch long and as big as
the tip of a man’s little finger, rising out of spigot holes in the earth, which eat the

@See Proc. Ent. Soc. Wash., v, pp. 126-127, February, 1903.
b There is no recorded brood which could have appeared in 1675, and the year meant
is probably either 1673 or 1676, both of which were cicada years.

148 THE PERIODICAL CICADA, - f

new-sprouted leaves from the tops of the trees without other harm, and in a
month left us.¢

The next reference to this insect is in a journal, dated 1715, left
by the Rev. Andreas Sandel, rector of the Swedish congregation at
Philadelphia. It is important as giving the first reference to the use
by the native Indians of the cicadas as an article of diet, and has
been recently published in the Pennsylvania Magazine of History and
Biography. The note is as follows:

May 9 |1715].—In company with several English clergymen, Mr. Talbot, Guerney,
and Clubb,> I went up to Radnor, where we laid the corner stone of a church.

In this month some singular flies came out of the ground; the English call them
locusts. When they left the ground holes could be seen everywhere in the roads, and
especially in the woods. They were then encased in shells, out of which they crawled.
It seemed most wonderful how being covered with the shell they were able to burrow
their way in the hard ground. When they began to fly they made a peculiar noise,
and being found in great multitudes all over the country, their noise made the cow
bells inaudible in the woods. They were also destructive, making slits in the bark
of the trees, where they deposited their worms, which withered the branches. Swine
and poultry ate them; but what was more astonishing, when they first appeared
some of the people split them open and ate them, holding them to be of the same
kind as those to have been eaten by John the Baptist. These locusts lasted not
longer than up to June 10, and disappeared in the woods,

Specimens of this insect for scientific study were first carried to the
Old World by Pehr Kalm, a pupil of Linné, who was sent to America
by the Swedish Government and traveled extensively in the colonies
between 1748 and 1751. The account of his travels, published in
Stockholm between 1753 and 1761, contains much interesting informa-
tion relative to the common insects of this country at that early
period, and gives a brief statement of the habits of the periodical
Cicada. While this work was being printed, Pehr Kalm published
a more detailed account of the species in the Swedish Transactions
for 1756 (pp. 101-116). The account given in his travels (English
edition, 1771, Vol. II, p. 6), is as follows:

There are a kind of locusts which about every seventeenth year come hither in
incredible numbers. They come out of the ground in the middle of May, an make,
for six weeks together, such a noise in the trees and woods that two persons that meet
in such places, can not understand each other, unless they speak louder than the locusts
can chirp. During that time, they make, with the sting in their tail, holes in the
soft bark of the little branches on the trees, by which means these branches are ruined.
They do no other harm to the trees or other plants. In the interval between the years
when they are so numerous, they are only seen or heard single in the woods.

aSee Webster, Insect Life, Vol. II, p. 161.

b Rey. John Clubb, a Welshman, for some time was schoolmaster in Philadelphia,
and also assisted Rev. Evan Evans. He also preached to the Welsh settlers at Radnor
and vicinity, and became rector of Holy Trinity Church, Oxford. He died in Decem-
ber of 1715. (The Pennsylvania Magazine of History and Biography, Vol. XXX,
October, 1906, pp. 448, 449.)

THE CICADA IN LITERATURE. 149

The original scientific description of the species by Linné, based on
material collected by Kalm, followed in 1758.4 Fabricius afterwards
described the species in two or three of his works under the name
Tettigonia septendecim, reviving one of the old generic names of Aris-
totle for this class of insects, but Latreille, Lamarck, and subsequent
authors retained Linné’s name.

In his monographie work on the Cicadas of the world, 1788, Caspar
Stoll gives a figure and a short description of Cicada septendecim.

Some popular accounts of the species closely followed Linné’s
description. Under the title; “Som observations on the Cicada of
North America,’ Peter Collinson, esq., of London, England, gave a
rather full account of the insect as then known, assigning fourteen or
fifteen years as its life period, and published a plate illustrating the
adult insect and a twig lacerated by the female.? Shortly thereafter
appeared an article in Dodsley’s Annual Register (1767, p. 103),
entitled, ‘‘Observations on Cicada or Locust of North America, which
appears periodically once in sixteen or seventeen years, by Moses
Bartram, 1766, communicated by the ingenious Peter Collinson.”’

References to the periodical Cicada in American literature began
to be more abundant toward the end of the eighteenth century and
in the begining of the nineteenth, Thomas Say, in 1817, referring
to ‘numerous accounts of it in our public prints.’ Most of these,
however, were unimportant notices and are now lost or not easily
accessible,

The most interesting contribution to the American literature of the
Cicada of this period, comprising two papers with valuable editorial
notes, is contained in the Barton Medical and Physical Journal of 1804,
already cited. The first title reads: ‘‘Some particulars concerning
the locust of North America. Written at Nazareth, in Pennsylvania,
Aug. 27th, 1793. Communicated to the Editor, by the Reverend
Mr. Charles Reichel, of Nazareth.’’ The ‘paper gives a number of
dates of occurrence in Pennsylvania and some interesting notes on
the habits of the Cicada—some errors in which are corrected in a note
by the editor, who announces that he has ‘‘for several years, devoted
a great deal of attention to the natural history of this insect”? and
“designs to publish an extensive memoir on the subject,’ which,
however, he seems never to have done.

The second paper (pp./56-59) reads: “Additional Observations on
the Cicada Septendecim. By the late Mr. John Bartram. From a
MS. in the possession of the Editor.”’ The older paper indicated in
this title I have not seen, but it is evidently included in an account
of travels by Bartram in Pennsylvania and Canada, printed in Lon-
don in 1751. Under the title quoted are notes on the appearance

aSystema Nature, tenth-edition, 1758, p. 435.
b Philos. Trans. 1764, vol. 54, pp. 65-69.

150 THE PERIODICAL CICADA.

of a brood in the neighborhood of Philadelphia in 1749, which began
to emerge May 10, but ‘‘in the latter end of April * * * came
so near the surface of the ground, that the hogs rooted up the ground
for a foot deep, all about the hedges and fences, under trees in search
of them.’ There follow quite accurate notes on oviposition. The
editor concludes the article by the citation from Moreton which has
been already quoted.

Thomas Say, the father of American entomology, has one brief
communication on the periodical Cicada, in which he criticises the
use of the name “ locust,’’ and “gives references to earlier literature
and a brief note on habits.@

Another interesting communication of about the same period is by
Dr. J. F. Davis,? in which the author controverts the ‘14 or 15” year
period suggested by Collinson and quotes two letters, one from the
Hon. Judge Peters, of Belmont, Pa., and the other from Myers Fisher,
esq., of Philadelphia, to substantiate the 17-year period. Referring
to the noise of this Cicada, Judge Peters says:

One of your Spa-fields meetings can give you a faint idea of their incessant and
unmusical cheering and noise. If Hogarth had known these locusts, he would have
placed them about the ears of his enraged musician. Knife-grinders, ballad singers,
ete., would have been lost in their din.

Mr. Fisher gives a very accurate, though brief, statement of the
life cycle of the species (if his belief that they occur at great depths be
excepted), and adds the very significant statement that ‘‘there is
reason to believe that they appear every year in some part or other
of the United States, with the complete period of 17 years between
every local appearance.”

Dr. S. P. Hildreth, of Marietta, Ohio, made two very valuable con-
tributions on the Cicada to the American Journal of Science and Arts
(1826 and 1830), which are much more accurate than any of the earlier
papers and too long to be quoted in this place. In the second of
these papers he calls attention to the existence of the small form of,
Cicada, and gives a colored plate representing five views of the adult
insect. Doctor Hildreth published a third paper also in 1847.°

The first account of this insect to be issued as a separate work is the
memoir of Prof. Nathaniel Potter, of Baltimore, Md., entitled ‘‘ Notes
on the Locusts,” etc., written in 1834 and privately published in 1839.
This pamphlet of twenty-nine pages and one colored plate, represent-
ing the insect in both sexes and also the early stages, together with
the nature of its work on twigs, and anatomical details, was the chief
source of information for the account published by Harris in his
“Insects Injurious to Vegetation,’ and while containing some wrong

@Mem. Phila. Soc. Prom. Agric., 1818, v. 4, p. 225.
6 Jour. Sci. and Arts Roy..Inst., 1819, v. 6, pp. 372-374.
¢ Loc. cit., ser. 2, vol. 3, pp. 216-218.

THE CICADA IN LITERATURE. 151

inferences, gives with remarkable accuracy — detail observations on
practically all of the features of the insect’s life history and habits
which are open to easy study, not only in its underground existence
but throughout its transformation and aerial life. Professor Potter
was evidently fully aware, not only of the two distinct sizes or varieties
of the Cicada but also of the depth to which the larve penetrate and
the fact of their forming roofs or turrets over their burrows some time
before the period of fee emergence—a record which has been hitherto

overlooked and the credit for this discovery assigned to a much later
period.

In speaking thus most favorably of Professor Potter’s memoir it
must not be forgotten that probably much of the actual observation
and study upon which it is based are due to the research of Dr. Gideon
B. Smith, of Baltimore, Md., who is given full credit in one of the
introductory paragraphs, in these words:

As our professional avocations would not permit us to devote our whole time to the
pursuit, it became necessary to call in the aid of a colleague whose knowledge of ento-
mology and industry could be relied upon. These qualifications were found and well
exemplified in Mr. Gideon B. Smith. Should our labors reflect any light on so obscure
a subject, the credit is equally due to him.

These two men were the first to make a careful and at all complete
study of the periodical Cicada, Professor Potter’s interest in the sub-
ject dating, he says, from 1783, and great credit is due them, and
especially to Doctor Smith, whose later work will be subsequently
considered.

Several brief accounts of the Cicada appeared in American and for-
eign publications about this time, adding nothing, however, to the
facts already obtained, the most notable perhaps being the account by
J. O. Westwood in his ‘Classification of Insects,’’“ in which he refers
to the literature and habits of the species very briefly.

The next step of real importance was the discovery of a 13-year
southern brood by Dr. D. L. Phares, of Woodville, Miss., and the
publication of the fact in 1845 in the Woodville Republican.

Both before and after this time Doctor Phsres was in communica-
tion with Dr. Gideon B. Smith, referred to above, whom he evidently
ultimately convinced of the truth of the 13-year period for the south-
ern broods.

Doctor Smith continued for many years the work which he had
begun as the colleague of Professor Potter, keeping his notes in the
form of a rather voluminous manuscript, which was first prepared,
he states, in 1834, the date signed to Professor Potter’s memoir.
Doctor Smith twice entirely rewrote and revised his manuscript, the
title-page of the last copy reading as follows

The American Locust Cicada septendeciin, et tredecim. Embracing the natural
history and habits of the insect in its perfect state and while underground, with

a 1839-40, Vol. II, p. 426.

152 THE PERIODICAL CICADA.

drawings of its several organs and the perfect insects, the egg and the young taken
from life, with a register of the places and time of its appearance in every part of the
United States, by Gideon B. Smith, M. D. Originally written in 1834, transcribed
with additions 1851, and rewritten kee additions and illustrations in February, 1857,
in the sixty-fourth year of my age. Base

This manuscript is rae the paper by Professor Potter
revised, with much interesting matter added and particularly a regis-
ter of some 21 broods in many colonies, in which are separated the
two tribes, one of seventeen years, represented by fourteen broods
and the other thirteen years, represented by seven broods. Doctor
Smith’s classification of the broods under these two tribes undoubt-
edly resulted from his correspondence with Doctor Phares and perhaps
other observers residing in the South. Most unfortunately, Doctor
Smith failed to publish this very interesting manuscript and there-
fore never received due credit for the valuable work which he
accomplished.

Townend Glover used this manuscript to some extent in his article
on the Cicada in the Report of the U.S. Department of Agriculture for
1867 (1868), referring to Doctor Smith as having devoted much time
to studying the habits of the Cicada, and as the best authority on the
subject in the Middle States, and particularly as holding that there
are two tribes “‘differing only from each other in the period of their
lives, the northern being seventeen years, and the other, or southern
tribe, requiring only thirteen years in which they perform their trans-
formations.’’ The use of Doctor Smith’s manuscript afterwards by
Professor Riley, as will be subsequently noted, was not of such char-
acter as to bring into prominence the real value of Doctor Smith’s
contribution to science. Two minor notes only were published by
Doctor Smith. The first is his Scientific American note of March 22,
1851, which was afterwards communicated by Mr. Spence to the
London Entomological Society.” In this note Doctor Smith briefly
reviews and sums up the results of his seventeen years’ study of this
insect, and states that he has located thirty different locust districts,
occupying fourteen of the seventeen years. Since he does not men-
a the 13-year race he was evidently unaware of its existence as late

s 1851. The second is a brief note in the Country Gentlen:an for
ue 869, in which he mentions both races.

From We time on until the important publications by Walsh and
Riley a number of articles on the Cicada appeared, some of them of
considerable interest and value, and notably those by Miss Magaretta
H. Morris, of Germantown, Pa., on the habits, times of appearance,
and ravages occasioned by this insect, and by Prof. Joseph Leidy on
the fungous disease attacking the species.’ Dr. J. C. Fisher, in 1851,

@ Proc. Ent. Soc. London, April 7, 1851, Vol. I, pp. 80, 81.
b Described by C. H. Peck as Massospora cicadina in 31st Rept. N. Y. State Mus.
Nat. Hist., 1879, pp. 19, 20, and 44.

THE CICADA IN LITERATURE. 153

described as a distinct species Cicada cassinii, the small form referred
to by several of the earlier authors, and to this paper were appended
comparative notes on the habits of the two forms by John Cassin.¢
About this time, 1851-52, also appeared the very complete account
by Doctor Harris in his ‘‘Insects of New England,’ and also some
anatomical studies of the sexual system and musical apparatus by
Dr. W. I. Burnett. In 1856 Dr. Asa Fitch, in his first report on the
insects of New York, gives an extended account of the periodical
Cicada, classifying or listing some nine broods, but not adding other-
wise particularly to the knowledge of the insect. Several accounts
of the species followed, including the notice of a 13-year brood, which
Doctor Phares claims to have published in the Republican of Wood-
ville, Miss., May 5, 1858, under the title ‘‘ Cicada tredecim”’—the earliest
published suggestion of this name for the 13-year race. None of the
other communications, including papers and notices by Fitch, Walsh,
Glover, and Cook, is of great importance, if we except the reference
by Glover to Smith already noted.

The next step of real importance was the publication by Walsh and
Riley in the first volume of the American Entomologist of a very full
and illustrated editorial account, in which the 13-year species is char-
acterized and the 13-year period for the southern broods is fully estab-
lished and a register of some sixteen broods is given. Professor Riley
in his First Missouri Report reproduces this article with the additions
to the broods derived chiefly from the manuscript memoir by Doctor
Smith, which had been in the meantime communicated to him by Dr.
J. G. Morris, of Baltimore, Md. In this paper Professor Riley revised
and renumbered the broods, increasing their number to twenty-two.
Professor Riley’s classification of the broods, and the details of the
life history and habits of the insect, as given by Walsh and Riley in
the American Entomologist, and later by Riley in his report, have
been accepted as the chief source of information since.

From the date of these articles until 1885 the additions to the liter-
ature are chiefly of records bearing on the distribution of the broods,
furnished notably by Rathvon, McCutcheon, Riley, Le Baron, Glover,
Phares, Packard, Lintner, and many others.

The recurrence in 1885 of the great Brood X of the 17-year race,
im conjunction with the very important 13-year Brood XXIII, gave
again a great stimulus to the study of this insect. Professor Riley
published in June, 1885, as Bulletin No. 8 (old series) of the Division
of Entomology, an account of both races with a very full chronology
of all the known broods. These data were repeated in part, with
important additions, in the Report of the Department for that year,
published in 1886. Other general articles were published by Doctor

@ Proc. Acad. Nat. Sci. Phila. 1851, Vol. V, pp. 273-275.

154 THE PERIODICAL CICADA,

Lintner and many others. The output of literature on the peri-
odical Cicada since 1885, if one takes the daily press notices and
articles into account, has been enormous and particularly in the
special Cicada years. This has resulted from the fact that as the
dates for the appearances of all the broods are now well understood,
the recurrences have been foretold and looked forward to, thus vastly
increasing the popular interest. The new information gained has
related chiefly to facts of distribution. Some interesting data have
been given, however, on the subject of the peculiar huts or turrets,
which are sometimes constructed by the emerging pupx, and some
anatomical studies have been made.

For a description of these and other papers the reader is referred to
the bibliography of the writings on the periodical Cicada which is
appended. The important papers from the earliest times to the
present are listed, omitting much of the ephemeral and less valuable
matter which added little or nothing to the knowledge of the habits
and distribution of the species.

BIBLIOGRAPHY OF THE PERIODICAL CICADA.
[Chronologically arranged. ]

‘
1666. [OLDENBURG, HENRY, “ publisher’’ |.—‘‘Some observations of Swarms of Strange
Insects and the Mischiefs done by them.’’ < Philos. Trans. London, Vol. I,
No: 8) p. L3si7e
Includes first reference to periodical Cicada (Brood XIV).
1669. Moreton, NATHANIEL.—‘‘ New England’s Memoriall.’’ <Cambridge, 1669.
Refers to a “ kinde of pestilent feaver”’ fatal to whites and Indians in season of 1633 (1634?)
ascribed by the Indians to the ‘flies,’ which appeared that year and which are briefly
described.
1705. MarrHews, THomas (?).—Steadman’s Lib. Am. Lit., Vol. I, pp. 462, 463 (1887).
Quotes the writings of 1705 of ** T. M.,’? supposed to refer to Thomas Matthews, describing
the occurrence of a swarm of cicadas as one of three prodigies appearing about 1675. (See
pp. 147-148.)
1715. SanpeLt, ANDREAs.—Mitchell and Miller’s Medical Repository, Vol. IV, p. 71. ’
(Abstract.) (Memorandum dated 1715.) (See 1906.)
Refers to the use of the cicadas as food by the Indians.
1756. Kautm, Penr.—Beskriining pa et slagts Gras-Hoppor, uti Norra Americas ( Cicada
septendecim). <Vetensk. Acad. Handl. 17, pp. 101-116—German Transla-
tion 1767, t. 26, pp. 130-143.
1753-61. KatmM, PEur.—Travels in North America. Vol. IJ, p. 6.
Gives a brief account of the species, which is said to come about every 17th year.
1758. LiInNE, C.—Systema Nature, 10th Edit., p. 436.
Original description of the species.
1764, CoLLINSON, PETER.—Some observations on the Cicada of North America.
< Philos. Trans. Lond., vol. 54, pp. 65-69, 1 pl.
1767. Bartram, Mosres.—‘‘ Dodsley’s Annual Register,’’ p. 103.
Account in letter to Peter Collinson, London, England.
1775. Fasricrus, J. C.—Syst. Ent., p. 679, No. 6; Ins., II, 1781, p. 319, No. 6; Mant.
Ins., II, 1787, p. 266, No. 9.
Described as Tettigonia septendecim.
1788. Srott, Caspar.—‘‘ Der Cicaden,’’ etc.
Gives a figure and short description of Cicada septendecim.

BIBLIOGRAPHY. 155

1804. RetcHEeL, Rev. CHarLEs.—Some particulars concerning the locust of North
America. < Barton Med. and Phys. Journal, Vol. I, p. 52, ff.

1804. Bartram, Joun.—Additional observations on Cicada septendecim. From a MS.
in the possession of the editor. <Barton Med. and Phys. Journ., Vol. I, pp.
56-59. (Earlier paper prob. 1751, Miscl. in Travels in Pa. and Can. Lond.,
Sept., 1751.)

1818. Say, T.—Mem. Philad. Soc. Prom. Agric., vol. 4, p. 225.

Refers to earlier literature and gives brief note on habits.

i819. Davis, J. F.—On the Cicada septendecim. <Journ. Sci. and Arts Roy. Inst.,

vol. 6, pp. 372-374.

Criticises paper of Collinson; quotes letters by Hon. Judge Peters and Myers Fisher to
substantiate the 17-year period.

1826. Hinpretu, 8. P.—Am. Journ. Sci. and Arts, vol. 10, pp. 327-329.
Habits and appearances detailed with considerable accuracy.
1828. Hitpretu, S. P.—Ueber die americanische Cicada (Cicada septendecim). <Fror.
Not., Bd. 22, No. 426, pp. 33-35.
1828. Bootu, Jesse.—Ueber die Cicada septendecim. <Fror. Not., Bd. 22, No. 468,
pp. 84-87.
1830. Hinpretu, S. P.—Notices and observations on the American Cicada or locust.
<Am. Journ. Sci. and Arts, vol. 18, pp. 47-50.
Characters, habits, and appearances; refers also to existence of small forms.
1832. Hitpreru, 8. P.—Ueber Cicada septendecim. <Isis, 1832, pp. 1059, 1060.
1837. Cui~pREN, J. G.—Proc. Ent. Soc. London, vol. 1, p. xxx.
Exhibited specimens of the different stages and read extract from a letter from Doctor
Harlan, of Philadelphia, giving a brief statement on habits, 17-year brood, ete.
1839. Porrer, NaTHanreL.—Notes on the Locusta septentrionalis american decim
septima. < Baltimore, J. Robinson, 27 pp., 1 pl.
History, habits, descriptions, and figures of Cicada septendecim.
1839-40. Wrestwoop, J. O.—Classification of insects, II, p. 4.
Brief account of species and reference to the literature.
1843. Morton, 8S. G., and others.—Cicada septendecim, discussion and extracts from
minutes in reference to. <Proc. Acad. Nat. Sci., Phila., Vol. I, pp. 277-280,

June.
Notes on various broods of periodical Cicada.

1845. Puares, Dr. D. L.—Woodville, Miss., Republican, May 17.
Published fact of a 13-year brood in Mississippi.
1846. Forstey, C. G.—On the Cicada septendecim in 1835 in Louisiana. <Proc. Bost.
Soc. Nat. Hist., vol. 2, p. 162.
Gives habits and distribution.
1846. Morris, M. H.—Proc. Acad. Nat. Sci. Phila., vol. 3, pp. 132-134.
Ravages, habits, and the times of appearance at various places.
1847. Morris, M. H.—Apple and pear trees destroyed by the locust. <Am. Agric.,
March, vol. 6, pp. 86, 87.

Ravages, habits, and transformations. ~

1847. Hinpreru, 8. P.—Cicada septendecim in 1846. <Am. Journ. Sci. and Arts,
March, ser. 2, vol. 3, pp. 216-218, 1-2; Ann. and Mag. Nat. Hist., vol. 20, pp.
136-138. ;

Habits in Ohio.

1847. Morris, M. H.—Proc. Acad. Nat. Sci. Phila., vol. 3, pp. 190, 191.

Ravages of the larvee; enemies.
1847. Hitpretu, S. P.—Die Siebzehnjahr-locust (Cicada septendecim). <Fror. Not.,
Bd. 3, No. 60, pp. 241-245.

1848. Hitpreru, S. P.—Détail sur les moeurs de la Cicada septendecim. < Institut,
XVI, No. 744, pp. 107, 108.

156

1848.

1851.

1851.

1851.

1852.

1852.

1852.

1854.

1858.

1859.

1860.

1861.
1862.

1865.

THE PERIODICAL CICADA.

Morris, M. H.—Destruction of fruit trees by the seventeen-year locust. <Am.
Agric., September, vol. 7, p. 279.
Notes on injuries to forest and fruit trees by the larvee.
Smrru, Dr. GrpEON B.—On the American locust ( Cicada septendecim). <Scientific
American, March 22; presented by Spence in Proc. Lond. Ent. Soc. (n. s.),
Vol. I, 1851, pp. 80, 81.

Reviews his work; states that he has located thirty locust districts, occupying fourteen of
the seventeen years; 13-year race not mentioned.

. Spence, R. H.—Proc. Ent. Soc. Lond. (n.s.), Vol. I, pp. 103, 104.

Letter on the Cicada in Maryland in 1851.

. Lewy, J.—Proc. Acad. Nat. Sci. Phila., Vol. V, p. 235.

Characters of the spores of a fungus affecting the Cicada.

. Fisuer, Dr. J. C.—Proc. Acad. Nat. Sci. Phila., Vol. V, pp. 272, 273.

Description of Cicada cassinii as a new species hitherto confounded with C. septendecim.

. Cassin, Joun.—Notes on the above species of Cicada (C. cassinii), and on the

Cicada septendecim Linn. <Proc. Acad. Nat. Sci. Phila., September, Vol. V,
pp. 273-275.
Characters and habits of Cicada septendecim and of Cicada cassinii compared.
Burnett, Dr. W. J.—Proc. Bost. Soc. Nat. Hist., IV, p. 71 and p. 111.
Sexual system and musical apparatus; appearance in cleared lands.

Burnett, Dr. W. J.—Points in the economy of the 17-year locust (Cicada septen-
decim) bearing on the plural origin and special local creation of the species.
<Proc. Am. Ass. Adv. Sci., 6th Meet., pp. 307-311.

Witp, Ph.—Sur les moeurs de la Cicada septendecim. <Ann. Soc. Ent. France,
2d Ser. vol. 10., Bul., pp. XVIII, XIX.

Harris, T. W.—Insects of New England, pp. 180-189; Ins. Inj. to Veget., 1860,
pp. 206-217; do. Flint Edit., 1863, pp. 206-219.

General account of the species.

Evans, Gurpon.— Insects injurious to vegetation. <Trans. N. Y. State Agric.
Soc. for 1851, vol. 11, pp. 741-751.

Notes concerning various insects of Madison County, N. Y., including the 17-year Cicada.

Frrcu, A.—Report (on the noxious, beneficial, and other insects of the State of
New York). <Trans. N. Y. State Agric. Soc. for 1854, vol. 14, pp. 742-753;
Ist and 2d Repts., Albany, 1856, pp. 38-49.

Gives a general account of the species and enumerates nine broods.

57. Smrru, Dr. GipEON B.—The American locust, ete. <Last revision. February.

Unpublished manuscript used in part by Glover and Riley.

Puares, Dr. D. L.—Republican, Woodville, Miss., May 5.

Published a notice of the 13-year brood under the title ‘* Cicada ‘redecim.”

Smirn, Dr. G. B.—Country Gentleman, May 12, vol. 13, p. 308.

“Predicts the appearance of locusts the coming spring in Virginia, Maryland, North Caro-
lina, Tennessee, Georgia, and Mississippi. They belong to the tribe of thirteen and seventeen
year locusts.” (13-year Brood XXIII and 17-year Brood I.)

Fircu, A.—The Entomologist, No. 22—The 17-year Cicada. <The Country
Gentleman, March 29, vol. 15, p. 210.

Remarks on popular names for insects; regularity of appearance of Cicada septendecim;

necessity of ascertaining its distribution in order to predict its future visitations.
Srax, C.—Ann. Soc. Ent. France, 4th ser., Vol. I, Septer-ber, p. 618.

Reference of Cicada septendecim to genus Tibicen Latr.

Herrick, E. C.—Uprising of the 17-year Cicada in New Haven County, Conn.,
in 1860. <Am. Journ. Arts and Sci., 2d ser., vol. 33, pp. 433, 434.

Ritey, C. V.—Seventeen-year locust. < Prairie Farmer, August 19, vol. 32
(i ee yole 1G) eromeleie

Agrees with S. P. G. in doubting that Cicada septendecim lives seventeen years immature,
and gives reasons for his doubt.

BIBLIOGRAPHY. Loz

1865. Wausu, B. D.—Pract. Ent., December 25, Vol. I, pp. 18, 19.
Answer to inquiry of M.S. Hill; Cicada districts of the United States, as given by Fitch in
New York Report, I, p. 39; habits.

1866. Ritey, C. V.—Prairie Farmer, September 1, vol. 34 (n. s., vol. 18), p. 136.
Answer to inquiry of J. D. Swain; condensed account of Cicada canicularis; comparison
with Cicada septendecim.

1866. Watsu, B. D.—Pract. Ent., December, vol. 2, p. 33.

Answer to inquiry of M. 8. Hill; Cicada septendecim compared with an undescribed species.

1867. Lemwy, Dr. Joseph.—Proc. Ac. Nat. Sci. Phila., p. 93.

Mentions recent appearance in various counties in Virginia.

1867. Grover, T.—Rept. (U. 8S.) Comm. Agric. for 1866, p. 29.

Brief notes, with dates of appearance.

1867. Wausxu, B. D.—Pract. Ent., February, vol. 2, p. 56.

Answer to inquiry of M. 8S. Hill; variations in the imago.

1868. Watsu, B. D.—The 17-year locust. <Dixie Farmer, June 11.

Periodicity and local distribution of the various broods.

1868. RurHerrorp, H.—New York Semi-Weekly Tribune, June 27.

Records the appearance of Brood X in Rutland County, Vt., in 1851 and 1868, and also of
Brood XIV on Long Island in 1855.

1868. Riny, C. V.—The 17-year Cicada. <Prairie Farmer, July 4, vol. 38 (n.s.,
vol. 22), p. 2. :

Dates and localities of occurrences.

1868. Rey, C. V.—Prairie Farmer, July 11, vol. 38 (n. s., vol. 22), p. 10.

Occurrence in Michigan.

1868. GLtover, T.—Rept. (U. S.) Comm. Agric. for 1867, pp. 67-71, figs. (Published
after July.)

General account of the species, quoting Harris and Smith, and referring to the latter’s
13-year broods.

1868. WausH, B. D.—Study of periodical Cicada. <Am. Ent., Vol. I, pp. 7, 8.
September.

Records unsuccessful experiments to get the Cicada to sting the flesh, and urges that the
stings are probably by Stizus grandis.

1868. Watusu, B. D., and C. V. Riney.—The sting of the 17-year Cicada. <Aim. Ent.,
October, Vol. J, pp. 36, 37.

Communications from F. W. Collins, R. Richardson, and B. Borden on the reputed sting
of the Cicada and on the habits of Stizus grandis. a

1868. Watsn, B. D., and C. V. Ritey.—The periodical Cicada. <Am. Ent.,
December, Vol. I, pp. 63-72, figs. 58-64. Extract: Op. cit., June, 1869, Vol.
oee202%

Characterization of the 13-year broods of Cicada as a new species, Cicada tredecim; dimor-
phism of the same and of Cicada septendecim; seasons, natural history, transformations,
enemies, sting, and injuries of these species; chronological history of their several known
broods; figures the several stages of Cicada septendecim, the towers made by the pups, and
twigs with eggs.

1868. Cook, A. J.—Remarks on some insects injurious to vegetation in Michigan.
<7th Annual Rept. Secr. State Board Agriculture Michigan for 1868, pp.
163-170.

Habits and means against Cicada septendecim.

1868. Riney, C. V.—Entomology. < Prairie Farmer Annual (No. 2 for 1869), pp. 30-
41, 6 figs.

Includes an account of the periodical Cicada, with figures.

1869. Warper, R. H.—Notes on the periodical Cicada. It does oviposit in ever-

greens. < Am. Ent., February, Vol. I, p. 117.
Oviposition of Cicada septendecim in three evergreens; note on the fungus found in the
abdomen of the Cicada; injuries to young orchards.

a Synonym of Sphecius speciosus.

158

1869.

1869.

1869.

1869.

1869.

1869.

1869.

1869.

1870.

1870.

THE PERIODICAL CICADA,

Watsu, B. D., and C. V. Riley.—Am. Ent., February, Vol. I, p. 117.

Comments on the above.

Riey, ©. V.—The periodical Cicada. <Ist Rept. Ins. Mo: March, pp. 18-42.

General account following Walsh-Riley article in American Entomologist and incorporating

facts on distribution of broods from Doctor Smith’s manuscript, renumbering the broods
and ee them to twenty-two.

. Ratuvon, 8S. §8.—Hatching of the 17-year Cicada. <Am. Nat., April, vol.

3, p- 106.

. Watsu, B. D., and C. V. Ritey.—Out of evil there cometh good. <Am.

Ent., June, Vol. I, p. 202.
Probable abundance of the fruit crop in southern Illinois and in Missouri in 1869 due to the
pruning of the trees by Cicada septendecim in 1868.
Wausn, B. D., and ©. V. Riey.—The periodical Cicada. <Am. Ent.,
June, Vol. I, p. 202.
Request for records of appearance in 1869.
Wausu, B. D., and ©. V. Riney.—Belated indivicuals of the periodical Cicada.
—<~ Ame Emits. Jiuliys Wolk paeZ li
Occurrence of scattering individuals in years before or after their regular period.
Watsu, B. D., and ©. V. Ritey.—Eggs of periodical Cicada in savin-twig.
<Am. Ent., July, Vol. I, p. 228.

Answer to inquiry of J. A. Greason; Cicada septendecim ovipositing in twigs of Juniperus
sabina.

Watsu, B. D., and ©. V. Rmey.—The periodical Cicada; our first brood
established. <Am. Ent., August, Vol. I, p. 244.
Appearance in Connecticut of a brood in 1869.
Watsu, B. D., and C. V. Rirey.—Insectsnamed. <Am. Ent., August, Vol. I,
p. 201.

Answer to inquiry of D. L. Phares; irregular appearance of Cicada tredecim ( Tibicen
septendecim).

Ratuvon, 8S. 8.—Cicada notes. <Am. Ent., November, vol. 2, p. 51.
Gives habits and appearance.
Ritey, ©. V.—The periodical Cicada alias the 17-year and 13-year locust. <Am.
Ent. and Bot., May, p. 211.
Quotes from the lst Ann. Rept. State Ent. Mo., the localities in which Cicada septendecim
and Cicada tredecim will appear in 1870, with requests for reports of the occurrence of these
insects.

. Krre, Wm.—The 17-year Cicada. <Am. Nat., vol. 3, p. 106.
. Morris, Joun G.—Seventeen-year locust two years too late. <(Am. Ent. and

Bot., September, vol. 2, p. 304.
Occurrence of a retarded Cicada septendecim in Maryland in 1870; note on the year of the
appearance of the Cicada in York County, Pa.

. Wausu, B. D.—Am. Ent., October, p. 335. (Posthume 1s paper.)

Argues for the specific distinctness of 17 and 13 year races as illustrative of a general problem
in article ‘‘On the Grape Eurytomides,”’ etc.

. Raruvon, 8S. 8.—Periodical Cicada not in Kreutz Creek Valley. <Am. Ent.

and Bot., December, vol. 2, p. 372.
Notes peculiarities in local distribution.
McCurcuen, A. R.—Periodical cicadas in Georgia. <Am. Ent. and Bot.,

December, vol. 2, p. 372.
Occurrence in Georgia in 1866, 1869, and 1870.

. Le Baron, W.—Locust or periodical Cicada. <Prairie Farmer, April 29, vol. 42.

Natural history.

. Le Baron, W.—Prairie Farmer, June 3, vol. 42.

Occurrence of lary of Cicada septendeeim in southern Illinois,

1872.

1875.

1873.

1875.

1876.

1879.

1879.

BIBLIOGRAPHY. 159

Ritey, ©. V.—Fourth Rept. Ins. Mo., April, pp. 30-34.

Gives the data collected on the six broods which had appeared since the publication of
article in first report.

Dimmock, G.—Insects infesting apple trees. No. 1: <New England Home-
stead, June 1, vol. 5, No. 4, p. 25.

Treats of Cicada septendecim, ete.
Dimmock, (.—Insects infesting apple trees. No. 4. <New England Home-
stead, June 22, vol. 5, No. 7, p. 49.
Treats of Cicada (= Tibicen) septendecim, ete.
Howarp, J. W.—Phillips’ Southern Farmer, October. \
Reports the occurrence of the Cicada at Flat Bayou, La., in 1872.
Le Baron, W.—Second Rept. Ins. Ills., pp. 124-133.
General account from Harris, Fitch, and Riley.

Brown, J. J.—Coleman’s Rural World, January 1.

Records the appearance of the Cicada in northwestern Arkansas along the White River
and its tributaries; traces them back in 13-year periods to 1803.
GuLover, T.—Report of the entomologist and curator of the museum. << Rept.
(U. 8.) Comm. Agric. for 1872, pp. 112-138, 26 figs.
Appearance and ravages of Cicada septendecim.
Puares, Dr. D. L.—Southern Field and Factory, Jackson, Miss., April.
Refers to his previous publications in the Republican on the 13-year broods.
Puares, Dr. D. L.—Southern Field and Factory, Jackson, Miss., August.
Records of Brood XXII since 1806; its extent in Louisiana and Mississippi.

PackarD, A. 8.—Third annual report of the injurious and beneficial effects of
insects in Massachusetts. 20th Ann. Rept. Sec. Mass. Bd. Agric., pp. 16-20,
figs. 142, 143.

Includes general account of periodical Cicada.
PackarD, A. 8.—Am. Naturalist, vol. 7, p. 536, September.
Reprint with corrections of article in Third Annual Report.

Brruungs, C. J. 8.—Grasshoppers or locusts. < Ann. Rept. Ent. Soc. Ont. for
1874, p. 29, fig. 30.

In article on grasshopper ravages, etc.; discusses confusion in use of name Locust.

tiLEY, ©. V.—Periodical Cicada, ‘17-year locust.” <New York Semi-Weekly
Tribune, June 23, 3 figs.

Occurrence at Lexington, Va., in 1876; list of localities at which these insects will appear
this year; chronological history of a brood; figures of larva, pup, and imago.

Riuey, C. V.—Entomological notes. Trans. Acad. Sci. St. Louis, December,
vol. 3, pp. 217, 218; see Am. Nat., October, 1876, vol. 10, p. 635.

Includes correction of vernacular name of Cicada septendecim; occurrence of the same in
Virginia in 1876; yearly development.
Leipy, Jos.—Remarks on the 17-year locust, ete. <Proc. Acad. Nat. Sci.
Phila., pp. 260, 261.
tinEY, C. V.—The periodical Cicada. < Western Farmer’s Almanac for 1878, p.
48; Colman’s Rural World, November 28, 1877.
Popular description and natural history; chronology of twenty-two different broods.
Ossorn, H.—The 17-year locust. Western Farm Journal, July.
General account of natural history.

Bessey, C. E.—lowa Weather Bulletin, November.

Gives an account of the distribution of the Cicada in Iowa, illustrated by a State map.

Ossorn, H.—Report of noxious insects. < Trans. lowa State Hortic. Soc. for
1878, vol. 18, pp. 868-402:

Includes habits and natural history of Cicada septendecim.
Peck, C. H.—Thirty-first Rept. N. Y. State Mus. Nat. Hist., pp. 19, 20, and 44,

Description of Cicada fungous parasites as Massospora cicadina.

160 THE PERIODICAL CICADA.

1879. SrrecKER, HeErMANN.—The Cicada in Texas. <Science News, Vol. I, No. 16,
p. 206.
1879. Rinpy, C. V.—New York Tribune, 1879; Colman’s Rural World, June 25.
Boundaries of the areas in which Cicada septendecim is expected to occur in 1879; request
for information of its appearance.
1879. Osporn, H.—Seventeen-year locusts. < College Quarterly, September, vol. 2,
p. 58.
Occurrence in southwestern Lowa in 1879.
1880. Ritey, C. V.—The 17-year Cicada in Iowa. <Am. Ent., February, vol. 3 (n. s.,
Vol. I), pp. 25, 26.
Review of the above; limits of the broods of 1854-1871, 1861-1878, and 1862-1879 in Iowa;

occurrence of the last brood in Missouri; comparison of the distribution of these broods with
the distribution of timber trees.

1880. Bessey, C. E.—On the distribution of the 17-year Cicada of the brood of 1878,
or Riley’s Brood XIII, in lowa. <Am. Ent., February, vol. 3 (n. s., Vol. I),
pp. 27-30, fig. 7. -

Summary of replies to inquiries concerning distribution in lowa in 1878, with map.

1880. CHamBerRs, V. T.—Am. Ent., Vol. III, p. 77, March.

Occurrence in Cheyenne Canyon, Colorado, in 1876. (Probably erroneous.)

1880. Riney, C. V.—Fungus in Cicada. Am. Ent., June, vol. 3 (n.s., Vol. I), p. 14.

1880. Riey, ©. V.—The periodical Cicada. Am. Ent., July, vol. 3 (u.s., Vel. I),
pp: 172, 173, fig. 76.

Broods which appear in 1880.

1880. Rosinson, F. C.—Seventeen-year Cicada in Pennsylvania. <Am. Ent., July,

Vol: B.(i. 8+ Vole) sp. Lins:
Occurrence at Uniontown, Pa., in 1880.

1880. Barnes, Harury.—Periodical Cicada in Geauga County, Ohio. < Am. Ent.,

September, vol. 3 (n. s., Vol. I), p. 226.
Abundance in western Ohio.

1880. Barnes, H.—Seventeen-year Cicada in Ohio. < Am. Ent., September, vol. 3

(n. s., Vol. I), pp. 227, 228.
Seasons and injuries.
1881. Ritey, C. V.—Bul. 6, U. S. Ent. Comm., March, p. 58.

Orthography of name; quotes Walsh as to validity of tredecim as true species.

1881. Riney, C. V.—The periodical Cicada alias “17-year locust.’’ < Am. Ent., June
(May 19), v. 25, pp. 479-482, fig. 1. Correction ibid., July (June 22), 1881,
p. 578.

Extract from First Missouri Report. with additional notes; figures eggs, pup, imago,
and punctured twig.
1881. Riney, C. V.—The periodical Cicada alias “17-year locust.’’ <Farmer’s Review,
June 16, vol. 6, p. 370.
Extract from First Missouri Report, with additional notes and requests for further informa-
tion in regard to distribution of the broods which appear in 1881.
1881, —— ——.—Sci. Amer., vol. 45, p. 21, July 9.
Records the appearance of the Cicada in southern Illinois, Kentucky, Arkansas, and Mobile,
Ala., in that year.
1881. Riwey, C. V.—Selma, Ala., Times, July 19, Cicada tredecim, abundant in Ala-
bama, as predicted.
1881. Ritey, C. V.—The periodical Cicada. <Am. Agric., August, vol. 40, p. 132,
5 figs.
Brief sketch of the natural history, with figures.
1882. LintNeR, J. A.—The 17-year locust. < Ontario [N. Y.] County Times, July 12,
(N. Y.) vol. 28, p. 3.
Years of appearance during the present half century; broods in New York; injuries and
preventives. .

1882.

1884.

1884.

1885.

1885.

1885.

1885.

1885.

1885.

1885.

1885.

BIBLIOGRAPHY. 161

Rmey, ©. V.—Cicada septendecim. <Gardener’s Mo. and Hortic., September
b] p ’

a4

vol. 24, pp. 274, 275.
Orthography of the names Cicada tredecim and Cicada septendecim; dimorphic forms;
Massospora cicadina parasitic on Cicada.

2. Osporn, H.—Insects of the forest—Cicada septendecim. <Iowa State Leader,

December 2.
Food habits of Cicada septendecim; life history.

. Rivey, C. V. —Am. Nat., March (February 21), vol. 17, p. 322.

Instinet of Cicada and sense direction in insects.

. SaunpDERS, WmM.—Ins. Inj. to Fruits, pp. 35-39.

General account of the species.

. Besspy, C. E.—The periodical Cicada in southeastern Massachusetts. <Am.

Nat., October (September 17), vol. 17, p. 1071.
Abundant in June, 1883, in Marthas Vineyard: notes by C. V. Riley.
Ossorn, H.—Insects of the orchard. < Bul. Iowa Agric. College, August, No. 2,
pp. 87-97. Trans. Iowa State Hortic. Soc. for 1883 (1884), vol. 18, pp. 510-521.
Includes reference to periodical Cicada.
Unter, P. R.—Cassino’s Stand. Nat. Hist., II, p. 227.
Natural history and distribution.

Rizey, C. V.—Destroying Cicadas. < Rural New Yorker, May 23, vol. 44, p.353.
Reply to letter of J. A. K. "
Riney, C. V.—Expected advent of the locust. <Sci. Am., May 23, vol. 52, p. 320.

Farmer’s Home Journ., June 13, 1885. Orange County (N. Y.) Farmer, May
28,1885. See Sci. Am., June 20, 1885, vol. 52, p. 389.
Simultaneous appearance of a 17-year and a 13-year brood; localities of the two broods; life
history and habits.
WereEp, C. M.—The coming locust plague. < Prairie Farmer, May 23, vol. 57,
p- 329.

Notes on Cicada septendecim.

. Ritey, C. V.—The periodical or 17-year Cicada. <Harper’s Weekly, June 6,

vol. 29, p. 363, 4 figs.

Distribution of Brood X and of Brood XXIII; habits, enemies, ete.

_ Riizy, C. V.—The periodical or 17-year Cicada. < Am. Grange Bulletin, June 11.

Chronological record, natural history, and popular names.

. Warp, Lester F.— Premature appearance of the periodical Cicada. <Science,

Vol. V, No. 123, June 12, p. 476.

. Rey, C. V.—The periodical Cicada. An account of Cicada septendecim and

its tredecim race, with a chronology of all broods known. < Bul. No. 8 [old
series], Div. Ent., U.S. Dept. Agric. (June 17), 46 pp., 8 figs. Second edition,
July 13, 1885.
Weep, C. M.—The 17-year locust. <Prairie Farmer, June 20, vol. 57, p. 393.
Brief account of the habits, ete.
Ritey, C. V.—Notes on the periodical Cicada. <Science, June 26, vol. 5, pp.
518-521. Reprint: Sci. Amer. Suppl., June 27, 1885, vol. 19, pp. 7905, 7906.
Distribution of Brood X and of Brood XXIII in 1885; life history, ete.
Ritey, C. V.—Periodical Cicada in Massachusetts. < Science, July 3, vol. 6, p. 4.
The occurrence in southeastern Massachusetts needs confirmation.
Ritey, C. V.—The influence of climate on Cicada septendecim. <Ent. Am.,
August, Vol. I, p. 91.

Records the transfers of eggs of Brood X to the extreme Southern States where no sep-
tendecim brood is known to occur and of Brood XXIII to Northern States where no tredecim
brood is known to occur. °

Davis, Wu. T.—The periodical Cicada on Staten Island. <Ent. Am., Vol. I,
August, No. 5, p. 91.

Pit No, 7107 ——F

1885.

1885.

1886.

1886.

1888.

THE PERIODICAL CICADA.

35. Ritey, C. V.—The song-notes of the periodical Cicada. <Science, September 25,

vol. 6, pp. 264, 265. Reprint, with additions: Proc. Am. Assoc. Adv. Sci. for
1885, August, 1886, vol. 34, pp. 330-332. Translation: Stett. Ent. Zeit., 1886,
Jahrg. 47, pp. 158-160. See Science, September 11, 1885, vol. 6, p. 226.
Kansas City Review, October, 1885, p. 171.
Description of the three prevalent notes.
Lintner, J. A.—The 13-year Cicada.< Argus (Albany), October 11, p. 4.
Notice of the life history.

5 (?). Harwaway, G. H.—Sci. American.

Gives records of Brood XI for 1818-1869. (See Bul. 8, 0. s., Div. Ent., U. S. Dept. Agri-
culture, p. 18.)

Lintner, J. A.—The 17-year locust, etc. <(Second report on the injurious and
other insects of the State of New York. Albany (February, 1886), pp. 167-179,
figs. 43-47.

Brief bibliography and general account of the species, with special reference to the broods
occurring in New York.

. Newserry, J. S.—Turrets of periodical Cicada in a cellar. <From School of

Mines Quarterly, Science, Vol. VII, March 12.

§. Howarp, L. O.—Proc. Ent. Soc. Wash., I, June 4, 1885 (March 31, 1886), p. 29.

Edibility.

Ritey, ©. V.—The periodical Cicada, etc. < Report of the Entomologist, Ann.
Rept. U. S. Commissioner Agric. for 1885, pp. 233-258, 1 map, 2 pl., Separate:
Washington, June 8, 1886, pp. 27-52.

Reproduction and revision of Bul. 8, with important additions, map and plates.

Burier, A. W.—The periodical Cicada in southeastern Indiana. < Bul. No. 12,
Div. Ent., U. 8. Dept. Agric. (July 13), pp. 24-31.

Contains many interesting observations on habits, enemies, etc.

Ritey, C. V.—Some popular fallacies and some new facts regarding Cicada sep-

tendecim L. < Proc. Am. Assoc. Adv. Sci. for 1885 (August), vol. 34, p. 334.

Variety cassinii is not the race tredecim; twigs with eggs do not necessarily break off or die to
insure the hatching of the larva.

_ Apcar, E. A.—Some observations on the anatomy of Cicada septendecim. <Jour-
’

nal Trenton Nat. Hist. Soc., January, pp. 43-46.

Mechanism of the genitalia.

. Lintner, J. A.—The 17-year locust, Cicada septendecim. <The Owl, May, vol.

2, pp. 17-19, figs. 1-5.
Life history.

. Lintner, J. A.—An experiment with the 13-year Cicada. < Report of the

State Entomologist to the Regents of the University of the State of New York
for the year 1885. 39th Ann. Rept. State Mus. Nat. Hist. for 1885 (July 6),
Dp. WE we:

Planting of eggs of a 13-year brood in New York.

7. Rocxwoop, C. G., Jr.—An insect fight. <Science, vol. 10, No. 237, August 19,

p. 94. (Sphecius and Cicada.)

. Scuwarz, E. A.—Cicadas at Fortress Monroe in June, 1886. <Proc. Ent. Soc.

Wash., Vol. I, July 8, 1886 (March, 1888), p. 52.

. Woopworts, ©. W.—Synopsis of North American Cicadide. <Psyche, Vol.

V, June, pp. 67, 68.

Tables for determining genera and species.

. AtexaNnpeER, A. G.—After-effect of the oviposition of the periodical Cicada.

<Insect Life, Vol. I, July. p. 15.
Tells of injury to apple trees by falling of fruit.
Riney, C. V., and L. 0. Howarp.—The periodical Cicada in 1888. <Insect
Life, Vol. I, July, p. 31.
Appearance of Brood XIII, septendecim, and Brood X XVI, tredecim, this year.

1888.

1888.

1889.

1889.

1889.

1889.

1889.

1889.

1889.

1890.

1890.

1890.

1891.
1891.

1891.

1891.

1892.

1893.

1893.

1893.

BIBLIOGRAPHY. 163

McNrat, J.—Insect Life, Vol. I, p. 50.
Notes of precursors of Brood XIII in 188s.

Weep, ©, M.—Cicadas or harvest flies and beetles. <Pop. Gard., November,
p. 45, 1 fig.
Marcu, J.—Wisconsin letter on Cicada septendecim. <Insect Life, Vol. I, Janu-
ary, p. 218.
Brief note on their habits, food, and enemies.

. Rimey, C. V., and L. O. Howarp.—The periodical Cicada in 1889. <Insect

Tite, Vol. 1; April, sp: 298.
Localities of expected appearance of Brood XIV, septendecim, in this year.

Ossorn, H.—Notes on destructive insects. <Ann. Rept. Iowa State Agric.
Soc. for 1888, pp. 670-680.

Contains a brief general article on the 17-year Cicada.

Morrretpt, M. E.—Rept. U. 8. Dept. Agric. 1888, p. 135.

Trees killed in Illinois.
Lintner, J. A.—An experiment with the 13-year Cicada. < Fifth Rept. on the
Injur. and other Insects of the State of N. Y., pp. 276-278.
The planting of eggs for experiment.
Suirn, J. B.—The periodical Cicada. <“Garden and Forest, p. 436.
Lecalities in New Jersey for Broods XIV, II, VI, and X.

Wesster, F. M.—An early occurrence of the periodical Cicada. <Insect
Life, II, November, pp. 161, 162.

SmirH, Joun B.—The periodical Cicada. <Ann. Rept. of the Entomologist
[New Jersey] for 1889, p. 270.

On Brocd XIV and other New Jersey broods.

Bucknout, Wm. A.—The periodical Cicada in Pennsylvania. <Report

Penna. Agric. Expt. Sta. for 1889, pp. 182-187, 1 map.
Distribution of broods in the State.

Scuwarz, E. A.—Notes on Cicada septendecim in 1889. <Proc. Ent.’ Soc.
Wash., Vol. I, pp. 230, 231 (May).

New localities in 1889 of Brood XIV; Iceal distribution and reasons for accelerations, ete.

Packarp, A. S8.—Insects injurious to forest and shade trees. <Fifth Rept.
U.S. Ent. Comm., Washington, pp. 95-97.

A general acecunt.

Ritey, C. V.—Periodical locusts. <Sci. Amer., May 16, p. 313.

Lewis, R. T.—The song of the Cicada is appreciated and listened to by other
insects.< Nature News, August, 1891, abstracted in Nature, 44, No. 1140,
September 3, p. 437.

Koni, W. M.—Seventeen-year locust. <Farmer’s Review, November 4, p.
700.

Lintner, J. A.—Cicada septendecim Linn. The periodical Cicada. <Seventh
Rept. on the Injur. and other Insects of the State of N. Y., pp. 296-301.

Additions to the bibliography and account of a new or unknown brood.

Wesster, F. M.—Cicada septendecim. Notes. <Ohio Bul. No. 45, Dec., p. 210.
Brief note on 17-year Cjcada.

Lintner, J. A.—The periodical Cicada. <Count. Gentl., March 23, p. 226.
Reply to inquiries that it will not be a locust year in New Jersey or New York.

Riey, C. V.—Circular Div. Ent. U. S. Dept. Agric., June.

Localities for Broods XVIII (tredecim) and I (septendecim); request for confirmations, ete.

Rey, C. V., and L. O. Howarp.—The present year’s appearance of the peri-
odical Cicada. <Insect Life, Vol. V, July, pp. 298-300.

Localities for Brood XVIII, tredecim, and Brood I, septendecim, in this year.

164

1893.

1893.

1893.

1895.

1893.

1894.

1894.

1894.

1894.

1894.

1894.

1894.

1894.

1894.

1894.

1894.

. 1894.

1894.

1894.

1894.

1894.

THE PERIODICAL CICADA,

SLINGERLAND, M. V.—The “17-year locust” in itshole. <Rural New Yorker,
July 29, p. 509. /
Life history and habits.
Ritey, C. V.—Periodical Cicada.- <Science, August 18, p. 86.
Localities for Brood XVIII, tredecim, and Brood I, septendecim, in 1893.
Ossporn, H.—Trans. Iowa State Hort. Soc., 1892.
Brief account.
McCartuy, G.—The periodical Cicada. <Bul. No. 92, N. C. Agric. Exp.
Sta., August, pp. 108, 109, 1 fig.
Brief account.
Smiru, J. B.—The periodical Cicada. <Bul. No. 95, N. J. Agric. Exp. Sta.,
September, pp. 6, 1 fig.
The expected appearance of Brood II in the next year in New Jersey.
Ritey, C. V.—Circular Div. Ent. U. 8. Dept. Agric., May 5, pp. 4.
Gives distribution by counties of Broods XIX and II, and asks for confirmations.
SLINGERLAND, M. V.—The periodical Cicada or locust. <The Farmer’s
Advocate, June 1, p. 225.
Brief account, and the broods that will appear in 1894.
Davis, Wm. T.—The 17-year locust on Staten Island. <Proc. Nat. Sci.
Assn. Staten Isand, IV, No. 4, February 10, pp. 13-15.
Davis, W. T.—The 17-year Cicada on Staten Island. < Journ. N. Y. Ent.
Soc., II, No. 1, March, pp. 38, 39.
Records of cicadas observed since 1877.
Davis, Wm. T.—The harvest flies (Cicada) of Staten Island, N. Y. <Amer.
Nat., Vol. 28, No. 328, April, pp. 363; 364.

Various notes on Cicada tibicen L., song, time of appearance, and capture, and on C
marginata Say and C. canicularis Harr. Mere mention of the periodical Cicada. |

RinEy, C. V.—The periodical Cicada. <Rept. of Ent. in Ann. Rept. U. S.
Dept. Agric., 1893, pp. 204, 205.
Records of broods for 1893.
Davis, W. T.—The 17-year locust on Staten Island. <Amer. Nat., Vol. 28, No.
329, May, p. 402.
Cicada observed since 1877.
Smirn, J. B.—The periodical Cicada. <Entom. News, May, Vol. V, p. 145.
Notice of the brood to appear in 1894, and the relation of the English sparrow to this insect.
Lintner, J. A.—The periodical Cicada, or the 17-year locust. <Circular,
Albany, N. Y., June 19, p. 4.
Habits, turrets, questions regarding the present appearance.
SLINGERLAND, M. V.—The periodical Cicada, or 17-year locust. < Rural New
Yorker, July 28, p. 470; August 4, p. 488.
General popular account, habits, and broods.
Davis, W. T.—The 17-year locust on Staten Island in 1894. < Proc. Nat. Sci.
Assn. of Staten Island, Vol. IV, No. 9, September, pp. 33-35.
Suuretpt, R. W.—The 17-year Cicada and some ofits allies. < Popular Science
News, Vol. XXVIII, October 10, pp. 154, 155.
Lanper, B.—Hut-building 17-year cicadas. <Sci. Amer., October 13.
Full account, with explanations and figure.
Smiru, J. B.—Insect Life, Vol. VII, October, pp. 192-195.
Under ‘‘ Notes of the year in New Jersey,” gives distribution of Cicada in 1894 in that State.
Krom, O. S.—The hut-building Cicada. <Sci. Amer., November 10, p. 295.
Criticises theories of Mr. Lander, and suggests other explanations.

Lanper, B.—Cicada hut-builders. <Sci. Amer., November 24, p. 327.
Replies to Mr. Krom, and gives additional proof in support of his (Lander’s) point of view.

1894.

1895.

1895.
1895.
1895.
1895.

1896.

1896.
1896.
1896.

1896.
1896.

1897.
1897.
1897.
1897.
1897.
1887.

1898.

BIBLIOGRAPHY. 165

Smrru, J. B.—The periodical Cicada. < Rept. Entom., Ann. Rept. N. J. Agric.
Exper. Sta., pp. 582-591, figs. 52-57.
General articles relating especially to Brood II in New Jersey; Cicada towers discussed.
LANDER, Bensamin.—Domed burrows of Cicada septendecim. <Journ. N. Y.
Ent. Soc., III, March 1, pp. 33-38, plate.
Love, E. G.—Notes on the 17-year Cicada, Cicada septendectm. <Journal N. Y.
Micros. Soc., XI, April 2, pp. 37-46, plate 49.
Original observations on structural details, the turrets, etc.
Martarr, C. L.—The Hemipterous mouth. <Proc. Ent. Soc. Wash., Vol. ITI,
No. 4, pp. 241-249, June 22, figs. 21-23.
Detailed study of the hemipterous mouth from dissection of the periodical Cicada.
Lintner, J. A.—The 17-year locust in the State of New York in 1894. <10th
Rept. Inj. and other Ins. N. Y. From 48th Rept. N. Y. State Mus. Nat. Hist.,
pp. 420-425, 2 figs.
Quotes circular (see above) and gives the results.
McCartuy, G.—Bul. No. 120, N. Car. Agric. Exp. Sta., November.
Contains brief notice of Cicada.
OsBorn, HerBERT.—Observations on the Cicade of lowa. < Proc. Iowa Acad.
Sci., III, 1895, pp. 194-203. Cicada septendecim.
SLINGERLAND, M. V.—The apple crop and 17-year locusts. <Rural New
Yorker, January 25, p. 53.
Doubts that the Cicada has any influence on the crop.
Hyart, J. D.—Cicada septendecim; its mouth parts, and terminal armor. <Am.
Mo. Mic. Journ., February 17, p. 46.
Mouth parts and ovipositor of the Cicada; method of oviposition (illus.).
SLINGERLAND, M. V.—On what do 17-year Cicadas live? <Rural New Yorker,
May 23, p. 351.
Food habits of Cicada.
Bruner, L.—Seventeen-year locusts. < Nebraska State Journal, June 19.

SLINGERLAND, M. V.—Seventeen-year locusts not poisonous. < Rural New
Yorker, July 11, p. 464.
Food habits of Cicada.
Scuwarz, E. A.—The periodical Cicada in 1897. <Cir. No. 22, s.s., Div. Ent.,
U.S. Dept. Agric., May, pp. 4.
Localities for Brood V, septendecim, and Brood XXII, tredecim, in 1897.
Wesster, F. M.—The 17-year locust in Ohio. <Ohio Farmer, May 20, p. 40, 1
map.
Expected occurrence in the State.
SLINGERLAND, M. V.—Do 17-year locusts damage fruit trees? <Rural New
Yorker, July 3, p. 487.
Damage not of great importance.
Wesster, F. M.—The San Jose scale and the periodical Cicada. < Newspaper
bulletin, Ohio Agric. Exp. Sta., November, pp. 2.
Relates to Brood V in eastern Ohio.
Wesster, F. M.—Brood XV [V] of Cicada septendecim in Ohio. <Can. Ent.,
October, pp. 225-229.
Distribution of Brood V in Ohio in 1897.
Wesster, F. M.—The periodical Cicada, or so-called 17-year locust in Ohio.
<Bul. No. 87, Ohio Agric. Exp. Sta., November, pp. 37-68, 11 figs.
Distribution in Ohio, habits, natural enemies, etc.
Hopkins, A. D.—The periodical Cicada in West Virginia. <Bul. No. 50, W. Va.
Agric. Exp. Sta., January (April), pp. 46, 23 figs., 1 map, 4 pls.
A general account with detailed records of all broods in West Virginia.

166

1898.

1898.

1898.

1898.

1898.

1898.

1898.

1898.

1898.

1899.

1899.

1899.

1899.

1899.

1899.

THE PERIODICAL CICADA,

Scuwanz, E. A.—The periodical Cicada in 1898. <Circular No. 30, s. s. Div.
Ent., U. S. Dept. Agric., April, pp. 4.

Location of Brood XXIII, tredecim, and Brood VI, septendecim, in 1898.

Howarp, L. O.—A new egg-parasite of the periodical Cicada. <Can. Ent., Vol.
XXX, pp. 102, 103, April.

Description of Lathromeris cicadz, new species.

Lintner, J. A.—The periodical Cicada. <12th Ann. Rep. State Entom., New
York, 1896. (May, 1898), pp. 272-289, Pls. [X-XIIT.

Gives additions to bibliography, a general account of insects, with original observations on
habits, and especially on the Cicada chambers, chiefly based on Brood II in 1894.

Marutart, ©. L.—The periodical Cicada. An account of Cicada septendecim, its
natural enemies and the means of preventing its injury, together with a sum-
mary of the distribution of the different broods. < Bul. U.S. Dept. Agric.,
Div. Ent. No. 14, n. s., 148 pp., 4 pls., 57 figs.

Habits, varieties, distribution, systematics, structure, development, enemies, and remedies.

. Fores, S. A.—The seventeen-year Cicada. < Prairie Farmer, June 25, p. 9.
. SanperRson, E. D.—Entomology.<Country Gentleman, July 21, pp. 573, 574,

1 fig.
Note on Cicada tredecim.

DE VARIGNy, H.—La cigale de dix-sept ans. Histoire d’un insecte. <Rev.
scient. (4) T. 10, pp. 353-365. C. septendecim. Q

A lengthy review, with extracts from Bulletin 14, n. s., Division of Entomology.

ScHENKLING, Steamunpb.—Die siebzehnjihrige Cikade. <Die Natur, Jahrg. 47,
pp. 447-451. C. septendecim nach Marlatt.

Hopkins, A. D.—Some notes on observations in West Virginia. < Bul. No. 17,
n.s., Div. Ent., U. 8. Dept. Agric., November, pp. 44-49.

Includes a reference to work on the periodical Cicada, and especially the relation of tempera-
ture to appearance.

Maruartr, ©. L.—A new nomenclature for the broods of the periodical Cicada.
<Miscl. Results Work Div. Entom., Bul. U. 8. Dept. Agric., Div. Ent., No.
18, n. s., pp. 52-58. C. septendecim.

Maruartt, C. L.—A consideration of the validity of the old records bearing on
the distribution of the broods of the periodical Cicada, with particular refer-
ence to the occurrence of Broods VI and XXIII in 1898. <Miscl. Results
Work Div. Entom., Bul. U. S. Dept. Agric., Div. Ent., No. 18, n. s., pp.
59-78. C. septendecim.

Saso, Karu.—Die siebzehnjihrige Cikade ( Cicada septendecim). <Prometheus,
Jahrg. 10, pp. 388-393, 401-406. 13 figs. (Z. T. nach C. L. Marlatt).

Account of habits and life history based largely on Bul. i4, n. s., U. S. Dept. Agric., Div.
Entom.

Wesster, F. M.—Distribution of Broods XXII [X], V [XIII], and VIII [XIV]
of Cicada septendecim in Indiana. < Proc. Indiana Acad. Sci. 1899, pp.
225-227, 1 map.

Surrn, J. B.—The periodical Cicada. <Report of the Entom. of the New
Jersey Agric. Coll. Exp. Station for 1898, pp. 447-450, May 1.

New Jersey records of Brood VI, 1898, with map.

Lanper, B.—Note on the seventeen-year Cicada. <Journ. N. Y. Ent. Soc.;
Vil, pp. 212-214.

Notes on the Cicada huts—shallow soil believed to be cause.

Wesster, F. M.—Entomology. <(Ohio Farmer, August 31, p. 152.
Notes on Cicada septendecim.

Fer, E. P.—Notes of the year for New York. .<Country Gentleman, Sept..

14) p. 733.

Note on Cicada septendecim.

1899.

1899.

1900.

1900.

1900.

1900.

1900.

1901.

1901.

1901.

1901.

1901.

1901.

1901.

1901.

1902.

1902.

BIBLIOGRAPHY. 167

Fert, E. P.—Notes of the year for New York. <Bul. No. 20, n.s., Div. Ent.,
U.S. Dept. Agric., November, p. 62.
Record for Brood VII in western New York, 1899.
ALLEN, GRANT.—“ The day of the cankerworm.’’ <Strand Mag. Oct.
Relates chiefly to the periodical Cicada, with many illustrations from Bul. 14, n. s. (Div.
Ent., U. 8S. Dept. Agric.), given as new! Reprint, Sci. Am. Sup., v. 49 (1900), pp. 20122-20124.
Frur, E. P.—Fifteenth Report of the State Entomologist on the injurious and
other insects of the State of New York. <Bul. N. Y. State Museum, vol. 6,
No. 31, June, p. 544.

List of localities in western New York for Brood VII, 1899.

Wesster, F. M.—The 17-year locust in Ohio. <Ohio Farmer, July 5.

Relating to Brood VIII in eastern Ohio.

Hoprxins, A. D.—The periodical Cicada or seventeen-year locust in West Vir-
ginia. <Bul. No. 68, W. Va. Agric. Exp. Sta., September, pp. 259-330,
3 pls., 4 figs., 9 maps. A revision of Bul. 50.

A full account of the Cicada, with descriptions cf all the broods occurring in West Virginia,

with maps.
Luacer, O.—Bugs injurious to cultivated plants. <Bul. No. 69, Minn. Agric.
Exp. Sta., December. ;
Periodical Cicada discussed, pp. 102-3. Illustrations. Does not occur in Minnesota.
Wesster, I*. M.—Notes on the occurrence of Brood XX [VIIT] of the period-
ical Cicada, Cicada septendecim, in Ohio in 1900. <Entom. News, Vol. XI,
December, pp. 638-640, 1 fig.

Report on Brood VIII in Ohio in 1900, with map.

SLINGERLAND, M. V.—Work of the 17-year locust. < Rural New Yorker, Octo-
ber 12, p. 690.

Letter by ‘‘S. B.” and not by Slingerland. Tied green rye straw around the trees and left
two trees unprotected. It was several years before they fully recovered (the two trees).
Says that trees that exude gum are not much harmed by the locust. Speaks of sparrows
eating Cicada.

Jounson, W. G.—Timely warning to fruit growers. <American Agricultur-
ist, July 13; p: 32; 1 fig.

Notice of coming brood of seventeen-year Cicada.

SLINGERLAND, M. V.—Seventeen-year locust. Watermelon bug. <Rural
New Yorker, July 13, p. 484.
Letter and reply, N. Y. .
Fernatp, C. H., & H. T. Fernatp.—Report of the entomologists. <Thir-
teenth Annual Report, Hatch Exp. Sta., Mass., January, p. 86.
Brief notes on Marthas Vineyard swarm (Brood VIII) in 1900; as much in evidence as ever.
Wesster, I’. M.—Report of the committee on entomology. <Ann. Rept.
Ohio State Hort. Soc. f. 1900, pp. 1, 2, 7 pls., 2 figs.

Report on Brood VIII.

Feit, E. P.—Seventeen-year Cicada. <Country Gentleman, November 7,
p. 902.

Hopkins, A. D.—Circular of warning. <W. Va. Agric. Exp. Sta., Oct. 15,
1 p., 1 map.

Relating to the 1902 swarm of the periodical Cicada.

. ie Fe: 5
Sanperson, E. D.—Three orchard pests. <Bul. No. 53, Delaware Agric. Exp.
Sta., January, 1902 (Dec., 1901), pp. 13-19.

Includes an illustrated account of Cicada septendecim L., pp. 13-19, with special reference to
Brood X, 1902, in Delaware.

Fexr, E. P.—Spraying for Cicada. <Country Gentleman, March 13, p. 219.
Mechanical oil emulsions not advised. :

Hunter, W. D.—The periodical Cicada in 1902. <Circ. No. 44, s. s., Div.
Ent., U. 8. Dept. Agric., March 13, pp. 4.

An inquiring circular sent out for Brood X, 1902.

168

1902.

1902.

1902.

1902.

1902.

“1902.

1903.

1903.

1903.

1903.

1903.

THE PERIODICAL CICADA.

SLINGERLAND, M. V.—Whitewash for 17-year locusts. < Rural New Yorker,
March 15, p. 189. ;
Heavy coat cf whitewash on trees fairly protective.
Lowe, V. H.—Miscellaneous notes on injurious insects. <Bul. No. 212, New
York State Agric. Exp. Sta., April, pp. 1-15, pls. I-IV on Cicada.

Contains an advance warning of Brood X of the periodical Cicada, with some good photo-
graphs.

. Maruarr, C. L.—A new nomenclature for the broods of the periodical Cicada.

<Cir. No. 45, s. s., Div. Ent., U. S. Dept. Agric., pp. 8, May 1.

2. Auwoop, W. B.—What to do with locusts. < Rural New Yorker, May 17, p 351._
2. Horxins, A. D.—What to do with locusts. < Rural New Yorker, May 17, p. 351.
2. Hoopes, ABNER.—Entom. News, XVIII, Mch., pp. 108, 109.

Record of hand collecting in nursery, June 18,1902. Brood X.

2. QuarinTANCE, A. L.—The seventeen-year locusts; how the adults feed. < Rural

New Yorker, July 26, p. 511.

. OsBorN, H.—Insects affecting forest trees. < Proc. Columbus Hortic. Soc.,

vol. 17, September, pp: 79-92.

. Quarntance, A. L.—On the feeding habits of the periodical Cicada. Bul.

No. 37, n. s., Div. Entom., U. 8. Dept. Agric., November, pp. 90-94. 3 figs.

First careful observation demonstrating feeding of adult cicadas.

. OsBorN, H.—Some notable insect occurrences in Ohio for the first half of 1902.

<Bul. No. 37, n. s., Div. Entom., U. 8. Dept. Agric., November.
Brief reference (p. 116) to periodical Cicada, Brood X, 1902, in Ohio.
SanpEerson, E. D.—Notes from Delaware. <Bul. No. 37, n.s., Div. Entom.,
U.S. Dept. Agric., November.
Brief record for Delaware for Brood X, 1902 (p. 101).
Quatntance, A. L.—The periodical Cicada or seventeen-year locust. <Bul.
No. 87, Maryland Agric. Exp. Sta., November, pp. 65-116; 2 pls., 17 figs.
Full account of Brood X, 1902, in Maryland, with valuable new observations on habits,
etc., and map.

. OsBorN, HEeRBERT.—A statistical study of variations in the periodical Cicada.

<The Ohio Nat., III, Dec., pp. 323-826.
A careful study of the cassinii form.
OsBorN, HERBERT.—Statistical study of variation in the periodical Cicada.
<(Amer. Assoc.) Science, new series, vol. 16, pp. 345, 346.
Synopsis of paper.
Sanperson, E. D.—Report of the entomologist: ~<Delaware Agric. Exp.
Sta., Report 1902, pp. 137-139 (3 plates).
Brief notes on periodical Cicada in Delaware, Brood X, 1902.
Maruatr, ©. L.—Notes on the periodical Cicada in the District of Columbia
in 1902. < Proc. Ent. Soc. Wash., vol. 5, pp. 124-126.
Maruatr, C. L.—An early record of the periodical Cicada. <Proc. Ent. Soc.
Wash., vol. 5, No. 2, pp. 126-7.
Author’s extras published February 4, 1903.
Perrit, R. H.—Mosquitoes and other insects of the year 1902. <Special Bul.
No. 17, Mich. Agric. Exp. Station, January.
Brief note on periodical Cicada (pp. 21, 22) with records for 1902, Brood X.
Smiru, J. B.—Report of the entomological department for 1902. <New Jersey
Agric. Exp. Sta., February.
Cicada septendecim, pp. 470-489—record of Brood X, w ith notes on other New Jersey broods
and map of all broods occurring in the State.
Garman, H.—Seventeen-year locusts in Kentucky. <Bul. No. 107, Kentucky
Aer. Exp. Sta., pp. 81-100, 4 pls., 3 figs. (May 23)

General account. Special reference to Brood X, 1902; maps, 1902 and 1906.

1903.

1903.

1903.

1903.

1904.
1905.

1905.

1906.

1906.

1906.

BIBLIOGRAPHY. 169

QuatntTaANceE, A. L.—Entomological notes from Maryland. <Bul. No. 40, Div.
Ent., U. 8. Dept. Agric., August, p. 47, 1 map.
Includes reference to periodical Cicada, with map for Maryland of Brood X, 1902.
Atwoop, W. B.—A note on the oviposition of the seventeen-year locust ( Cicada
septendecim Linn.) <Bul. No. 40, Div. Ent., U.S. Dept. Agric., August,
pp. 70, 76.
Indicates the possible repellent action of Bordeaux mixture, and gives list of 33 plants
oviposited in.

3. Meek, W. J.—On the mouth parts of the Hemiptera. <Bul. Univ. Kansas,

il, pp. 257-277, pls. vii-xi. November.
A letailed anatomical study based on the periodical Cicada.
Britton, W. E.—Third Report of the State Entomologist. <Report Conn.
Agric. Exp. Sta. f. 1903, p. 214.
Brood XI, 1903, expected but no records obtained. (Was reported for Rhode Island.)
Fett, E. P.—Eighteenth report of the State Entomologist on injurious and other
insects of the State of New York. <Bul. 64, N. Y. State Museum, Albany,
jee
Brief report of Brood X, 1902, in New York.
McCook, Henry.—{The periodical Cicada.] <Harpers’ Mag., June, 1904.
GArMAN, H.—The seventeen-year locust will not appear in Kentucky this year
(1905). <Kentucky Agr. Exp. Sta. Bul. 120, pp. 74-76. May.
Distribution of Broods XIV, 1906, and XIX, 1907, from Kentucky Bulletin 107.
[Surrace, H. A.]—The seventeen-year locust in Pennsylvania. < Penna. Dept.
Agric. Mo. Bul. Div. Zool., vol. 3, No. 6 (Oct.), pp. 174, 175.
Advance warning of Brood XIV, 1906.

Maruatt, C. L.—The periodical Cicada in 1906. <U. 8S. Dept. Agrie., Bur.
Ent. Cir. 74, pp. 5, figs. 3, Apr. 16. :
[Surrace, H. A.]|—The Cicada or seventeen-year locust in Pennsylvania.

<Penna. Dept. Agric. Mo. Bul. Div. Zool., vol. 3, No. 12, pp. 369-377.
Various records of appearance in Pennsylvania.
[SANDEL, Rev. ANDREAsS.]—‘‘ Extracts from the Journal of.’’ <Pa. Mag. of
Hist. & Biog., Vol. XXX, Oct., pp. 448-449.

APPENDIX.

DR. GIDEON B. SMITH’S CHRONOLOGY OF THE PERIODICAL
CICADA.

[From a copy, by Dr. J. G. Morris, of Doctor Smith’s unpublished manuscript.]

It is proper to remark in relation to the districts in this tribe or
division that there is some uncertainty in relation to some of them
(as well as to those of the northern division) that have their borders
on the great line that separates the two divisions, owing to the fact,
remarked upon in another place in this work, that the districts often
interlock, those of the northern running down into the territory of
the southern and those of the southern running up into that of the
northern division, sometimes for hundreds of miles. A remarkable
instance of this will be found in the case of the southern Illinois dis-
trict, which ascends to the north nearly three degrees of latitude above
the regular line of division; and also to the lapping of one district
over another on their respective boundaries, elsewhere noticed. The
reader will therefore make due allowance for such errors as he may
find in the dates of appearance.

REGISTER OF THE SOUTHERN TRIBE (THIRTEEN-YEAR LOCUSTS).

1842. Iilinois—In Washington, Jefferson, Franklin, Perry, Randolph, Monroe, St.
Clair, Madison, Bond, Clinton, Edwards, Marion, and adjacent counties in
the southern end of the State, in 1829, 1842, 1855, and again in 1868. Of this
there is great doubt whether it belongs to the seventeen-year tribe, as is
indicated by the following paragraph from the Baltimore Sun of June 13, 1859:
“The locusts have made their appearance in Egypt, in southern Illinois,
and cover woods and orchards in swarms.’’

1842. Kentucky.—Northwest corner of State, about Paducah and adjacent counties
in the south, in 1829, 1842, 1855, and again in 1868.

1842. Alabama.—Russell and adjacent counties on the east side of Black Warrior
River, in 1842, 1855, and again in 1868.

1842. Lowisiana.—Morehouse Parish, Caddo, Claiborne, Washita, and adjacent
parishes, in 1855, and again in 1868.

1842. Arkansas.—All the northern counties in 1842, 1855, 1868.

1842. South Carolina.—Chester district and all adjoining to the Georgia line and to
North Carolina north[ ward] in 1816, 1829, 1842, 1855, 1868.

1842. Tennessee.—Montgomery, Bedford, Williamson, Rutherford [and adjacent
counties], in 1842, 1855, and again in 1868.

1842. Georgia.—Cherokee County in 1816, 1829, 1842, 1855, 1868.

1842. North Carolina.—Mecklenburg County in 1816, 1829, 1842, 1855, 1868.

170 :

e

. Louisiana.—East and West Feliciana in 1806, 1819, 1832, 1845,
. Georgia.—Gwinnett, Dekalb, and Newton counties in 1846, 1859.

DR. GIDEON B. SMITH’S CHRONOLOGY. Wel

2. Missouri.—All southeast part in 1829, 1842, 1855, 1868.
3. Georgia.—Habersham and Rabun(?) counties in 1843, 1856, 1869.
. Georgia.—Muscogee, Jasper, Greene, Washington, and adjacent counties in 1843,

1856, 1869.

. Florida.—Jackson, Gadsden, and Washington counties in 1844, 1855, 1870.
. Mississippi.cFrom the Mississippi River east to a ridge that divides the State

north and south, 45 miles from the river, and north and south to the bound-
aries of the State, in 1806, 1819, 1832, 1845, 1858.
1858.

Tennessee.—Northern part in 1846, and again in 1859.

. Mississippi.All the east of the State, from the ridge 45 miles from the river

on the west to the east boundary, in 1820, 1833, 1846, 1859.
Texas.—Appeared in some parts in vast numbers; unable to get any particulars.
If true, will appear again in 1862.

4. Georgia.—Cherokee County, northern part, in 1828, 1841, 1854, 1867.

55. North Carolina.—Buncombe and McDowell counties in 1855.

1843.

1843.

1843.

1844.
1844.

[N. B.—Doubtful whether this is a southern or northern district. They
appeared in 1855, at all events, and will again in 1868 or 1872.]

. Louisiana.—Carroll Parish, May 1.
. Arkansas.—Phillips County, May 10.
. Tennessee—About Memphis.

REGISTER OF THE NORTHERN TRIBE (SEVENTEEN-YEAR LOCUSTS).

. The locust appeared in North Carolina from Raleigh to near Petersburg, in

Virginia, and will appear again in 1859.

. They appeared in the valley of Virginia from the Blue Ridge on the east, the

Potomac River on the north, to the Tennessee and North Carolina lines on
the south, and several counties in the west, in 1808, 1825, 1842, and will
appear again in 1859.

. Illinois —About Alton, and again in 1859.
2. Maryland.—Southern part of St. Mary County, dividing the county about mid-

way east and west. Appeared there in 1825, 1842, and again in 1859.

2. North Carolina.—Rowan, Davie, Cabarrus, Iredell, and adjacent counties in

1825, 1842, and will appear again in 1859.

. Indiana.—Sullivan and Knox counties in 1859.
3. New York and Connecticut from Long Island Sound, west side of Connecticut

River, north on both sides of the Hudson River to Washington County, N. Y.,
and west to Montgomery County on the Mohawk River. Appeared there in
1809, 1826, 1843, and wiil again in 1860.

3. Michigan.—Kalamazoo; appeared in 1843, and will again in 1860.
3. Indiana.—Dearborn County; will again in 1860.
3. North Carolina.—Caldwell(??), Rockingham, Stokes, Guilford, Rowan, Surry,

and adjacent counties; appeared in 1792, 1809, 1826, 1843, and will again in
1860.

Pennsylvania.—Bounded by Peters Mountain on the south, Mahonlago (?)
Mountain on the north, and extending from the Susquehanna to the Dela-
ware River; appeared there in 1843, and will in 1860.

New Jersey.—Whole State, in 1775, 1792, 1809, 1826, 1843, and again in 1860.

Maryland—From Anne Arundel County to the north part of St. Mary, from
the Potomac to the Chesapeake Bay, in 1809, 1826, 1843, 1860.

Tilinois—In Warren County, and will again in 1861.

Jowa.—In various parts, and will again in 1861.

1846.
1846.
1847.
1848.

1849.

1849.

1850.
1851.

1851.

THE PERIODICAL CICADA.

Missouri.—All the western part of the State from Saline County west, as far as
heard from, north to the boundary of the State and south to Arkansas in 1845,
and will again in 1862.

}. Ohio.—Fastern part, extending west to Scioto River and Sandusky on Lake

Erie, extending over twelve counties in 1829, 1846, and again in 1863.

Virginia.—Southeastern part in 1829, 1846, and will in 1863.

Virginia.—Lewis County, in 1795, 1812, 1829, 1846, and will in 1863.

About Wheeling, in Virginia, in 1830, and will again in 1847, 1864.

New York.—In Monroe, Livingston, Madison, and adjacent counties in 1797,
1814, 1831, 1848, and will in 1865.

Pennsylvania.—In Armstrong, Clarion, Jefferson, Chemung, Huntingdon, Cam-
bria, Indiana, Butler, Mercer, Beaver, and in nearly all the western counties
in 1832, 1849, and will in 1866.

Ohio.—In Mahoning, Carroll, Trumbull, Columbiana, and adjacent counties,
especially in Columbiana in 1812, 1829, 1846, the eastern district lapping over
this in that county; appeared in this district in 1815, 1832, 1849, and will
in 1866.

Virginia.—County (?) and adjacent territory in 1833, 1850, and will in 1867.4

Maryland, Pennsylvania, Delaware, Virginia.—Beginning at Germantown, Pa.;
south to the middle of Delaware; west through the eastern shore of Mary-
land, upper part of Anne Arundel; west through the District of Columbia,
Loudoun County, Va., where it laps over the south Virginia district from the
Potomac to Loudoun County some 10 to 20 miles in width, and [in] this strip
of territory they appear every eighth and ninth year. Thence the line
extends through the northern counties of Virginia and Maryland to the
Savage Mountain, and thence along the southern tier of counties in Pennsyl-
vania to Germantown. The whole territory embraced in these boundaries is
occupied by the locusts. Appeared here in 1766, 1783, 1800, 1817, 1834, 1851,
and will again in 1868.

Ohio.—Cincinnati; Franklin, Columbus; Piqua, Miami County. This district
extends into Indiana to New Albany, Madison, Indianapolis, to the Wabash
River, Terre Haute, and to Louisville, Ky., in 1834, 1851; will again in 1868.

Massachusetts —Bristol County, Dearfield, Hampshire, and to Fall River in
1767, 1784, 1801, 1818, 1835, 1852, and will in 1869.

Ohio.—Vinton County in 1853, and will in 1870.

. Illinois.—In Jo Daviess County, and will in 1870.
. Illinois —In Winnebago, Menard County, and neighborhood in 1854; again in

1°71

Maryland.—On the old Liberty Road leading to Carroll, and Adams County,
Pa., and on the Winden (?) Mile Road extending to Carlisle, Pa., in 1838,
1855, and in 1872.

. Kentucky.—About Frankfort, Lexington, and Flemingsburg, extending to

Meigs and Gallia counties, Ohio, in 1838, 1855, and in 1872.

Maryland.—Eastern Shore from Cecil County to Worcester in 1838, 1855, and
in 1872.

Massachusetts.—Barnstable County, in 1770, 1787, 1804, 1821, 1838, 1855, and in
1872.

Virginia.— Kanawha County, extending only 15 miles each way, in 1838, 1855,
and in 1872.

. North Carolina.—In Buncombe and McDowell counties in 1855; again in 1872.

[N. B.—There is some doubt whether this district is not a 15-year district.
The locusts appeared there in 1855, at all events. |

a This evidently refers to Brood IX, which is known from many counties in Vir-
ginia (see pp. 49-50).

DR. GIDEON B. SMITH’S CHRONOLOGY. LS

[Nore on THE SmirH ReGisteR.—An examination of the above register of appear-
ances, prepared by Dr. Gideon B. Smith, at once indicates the painstaking care which
Doctor Smith must have devoted to the subject, and surprises one with the accuracy
and completeness of the records. All of the important broods known until recently
are designated more or less completely in Doctor Smith’s register, namely, the seven
13-year broods and the fourteen 17-year broods mentioned in Bulletin 14.

Taking the records in the order in which they are given in Doctor Smith’s register,
and beginning with the 13-year race, it will be seen that the localities listed after 1842
and 1855 refer to Brood XIX, after 1843 to Brood XX, and similarly 1844 to Brood X XI,
1845 to Brood XXII, 1846 and 1859 to Brood XXIII, 1849 to Brood XX VI, and 1854
to Brood XVIII.

Comparing in the same way his register of the northern tribe, or 17-year race, it is
seen that his localities listed after 1842 apply to Brood I, after 1843 to Brood II, and
similarly 1844 to Brood III, 1845 to Brood IV, 1846 to Brood V, 1847 to Brood VI, 1848
to Brood VII, 1849 to Brood VIII, 1850 to Brood IX, 1851 to Brood X, 1852 to Brood
XI, 1854 to Brood XIII, and 1855 to Brood XIV. The records given after 1853 are
probably erroneous, as indicated in the discussion of my Brood XII (pp. 55-56).—
¢.L. M.]

aa
acne a8
eee aT

Meee acid. avast periodical! Cicada... =. ..2-2-4-.--2-2

Ampelis cedrorum, enemy of periodical Cicada..-........--

Auiavenemies Ol periodical Cicada. .....+..---£--f2:-2-0-6

Aphis, woolly. (See Schizoneura lanigera. )

Apple;-food plant of periodical Cicada...........-...------

injury through oviposition of periodical Cicada. - .---
Astragalinus tristis, enemy of periodical Cicada. .......-.----
Bzxolophus bicolor, enemy of periodical Cicada ....-.-.-----

Bass, black. (See Micropterus salmoides. )

Bdella sp., enemy of periodical Cicada...........--...-----
Birch, black, food plant of periodical Cicada..........-----
Bisulphid of carbon, against grape root-aphis..........-----

periodical Cieada, 2... -2...+

Blackbird. (See Quiscalus quiscula and Q. q. xneus.)

Bordeaux mixture, repellent against periodical Cicada. -..--
Canker-worm, Nothrus ovivorus an enemy .......----------
Carbolic acid, against periodical Cicada. ..-0...-.02-+---.--+

Cardinal grosbeak. (See Cardinalis cardinalis.)

Cardinalis cardinalis, enemy of periodical Cicada ......-.--- -

‘atbird. (See Galeoscoptes carolinensis. )
Catbriar. (See Smilax rotundifolia.) -
Catfish, blue. (See Ichthelurus punctatus.)

Catostomus teres, enemy of periodical Cicada. .........----.
Cecidomyiid, enemy of periodical Cicada..........-...-.--

Cedar bird. (See Ampelis cedrorum.)

Chalcidids, parasites of periodical Cicada.........--...-!--
Cheyletus sp., enemy of perijodical Cicada.....--...--- ee

Chickadee, Carolina. (See Penthestes carolinensis. )
Chipmunk. (See Tamas striatus.)

Oicada cassinit, a variety of periodical Cicada ............--
= Tibicen septendecim, var. cassinit.......---
dog-day. (See Cicada tibicen.)

MANGURALE, PIOWAWLY PCH OMIC = —o-.en<<-.-5--4-4-- = =:
periodical, ‘‘adobe dwellings.’’ (See cones. )

aculd <aratiGm Of StACE - 52.26. Soe ceo s

1S] OFFS pee SENS et ope a a ge ee

advantages to species from long larval life

appearance regarded as ominous presage. .

appearamees We TWHUTe. 06... 5 - gees ee ee

ARMOPOCIOL AMINMAIS =, 2 Se oe eee sl win

TTS Ves CAS A ea a ee

| OED SSSR I ee A eC ener ee

Bo aE A ale 102

bid See eas eee et 139

ite See en Oise 101
Be ele aN ae 145
ee Soe 145-146
Sot ee See aes 144
be ee cece 135
ST ae ee 143

Leh, Fe DC rT: 138

SE ae Seton ae a 90-91
Seen sees 13
Evie ae OS 103-104

Be oe: Ie 102-104, 148
Sete eh state oe a Sets 79-81

176 THE PERIODICAL CICADA.’

Page.

Cicada, periodical: bibliography ee - oa csee en oe ae ee 154-169

birdiememiles; > 2524, 2. ceyh2c1s Sor cits Ree ee eee ee 138-139

breeding record confirming 17-year period...........---- 114-116

Brood I-=septendectm=——V91O- =) See ae a oe ee ee 38-39

Ti—septendectm—l9M so. sas eee Sea 39-41

ITT—septendecrm—1912 s,s et ig ee ee 41-42

LV—septendectn=1913' = 2-22: alee eee Bae See eee 42-43

V—septendecim—1914) 2.2 ice 8 2A ees. eee 43-44

Vi-—septendecim= 9. 2 atc Soe ee Be ae ere 44-46

Vil—septendecii— 19062. to sn oe a Seer 46-47

Vill—séptendeciin—19NT 23 30 o= ecb ae Se eee 47-49

TX se piendecvi—lOnee 2 3 eee ee Se 49-50

K—septendecin— 1919. oo ea). hoc icie nae eis oe bee 50-54

XT septendectmi—=l9 200 an. P 2a cee oy te ee ee 54-55

R-—sepiendectim—V02 shes cee 5 eee eee 55-87

XI ——sepicndenm—l922 - eet os eee ee 57-58

MEV —sepierdecim—V9 aoe.) Fo. S Se eee = a ee 58-60

XV—septendectm—1907 —. . 2. shine See ee See 60-62

DOV 1“ sentendecim— AGUS se pe etere hee ee 62-63

XK Vil=—septendecrm—I909 2 ese. 8-422 re eee 63-64

EV tredecim— 199 ee ees oe eee a eee 65-66

XEX—tredectm==1901 sone sone ee ae eee ee 66-67

XK predeotm== S908 e ee soos ts eae een a> Sipe ee 68

NEXT iredecin— 1909 i aceon ete pone 2 eee eer 68-69

XR redecim 19 lO eee eee Soe: eee eee 69-70

SERS tredecti—1 OI See eases ee a eer eee 71-73

XXIV predegim— 1902 tse sn dee se see 73-74

SOXV = iredecim=—19 13 a Fe ee ee 74

RONG V 1 tredecvin=— NOMA ee tee eee eee eee 74-75

XV tredecim= 9 1b Sse ee eee eee 75

XV Li tredecim = 1916 2 oso ae ia eee ee 75

RUN tredacin 9072 eee ne eee 75-76

XOX tredecim NWS ae ee ae eo ee eee 76-77

Pproodes foes a Se Re ee eee ee 22-34

’ classifi calion ie) 2o weet oar eee eee 25-28

. fuisire SppearaMees. 5 deo. one ae oe Fe ee 34

Ob 14.015. or kG. sieuit penlOds. 52a enema ae eee 33

lf-year race, Tange: << * 22: . ee ee ee eee 38-64

1B-yearrace, TAM. — 7p. ce ae cn eer eee 65-74

OMIM SAE tne. ole yee adie Cee ee ere eee 22-25

PAROS Cee oe ems oe oe ee ee 38-77

relationships © 0... tcp aes vee ee Se ieee 28-30

sources: OF error in Old MECORGHes-.c os--e- ee = ee 30-33

DilImOwiiG: 45.25 fee a2 eee ee a eee ee ee 125-126
“‘chimneys.’’ (See cones.)

chronology of Dr. Gideon B. Smith.-...--.:-...=-------- 170-173

Glinmate imemel aieione nos ea Cis ee eee eee 18-20

GOTLES so esos are ote Se ee ree re a en 91-98

confusion with locust or erasshopperes-=--—-=- 2-2 = == === 1d

damace by adullisi: tee ee yee ae goo 11, 106-108

Jarvie' and pllpe ss... ce eee eee eee eee 126

Gecimiakionn Ca SES a= eee eee ee eee eee 13-14

description of stages...---..-25)-38 225-8 see een e ee . 118-121

INDEX. ay

Page.
awdia~ periodical, diptcrous enemies): +420 see. -- 222-6 l lls lst ase .ke 129
dustriputiones = scale cha... : ee rae aera Sete ee 34-77
LI ELOT' ere p ly Ets ag ea a 99-101
SOUECESON MmlonmMabioniaa-]---.22..--..25s25225- 34-35
durationroiwaduiliistavemseemne rete. et oe 90-91
CAI Oe Eee Maree MR eas a 2 ns cnc og ace oe see 20-22
Sanlyene Cones mre settee See SH Sea 146-150
COADARARMLeOM areas Means een eee oy ee a Sa ee 130-132
eboe cd CSRhipbIOni emia woes ne eee ed ol 111
erowmiitand hatebing sss. o2 26... ico eke 110-112
(CUYSCST AH CTI se 38 ky cS Sp 109-110
retarded development: :2: 822022 2! St. ace eee 111
emergence, acceleration artificially... ._._- hg et eter 90
VEEYEL OV. 2 cc Mee) ar aa ge ced ae a 91
| SCEUTLATG A epee ag Rs, a REee 88-90)
food habits of adult. .....-.-. SCORE oe een 6 Pee ee tae 101-102
IarVvagne pipe. 22. yo. 5 Lee ie Sl 122-123
POMPOM SeASes Sater ne nee SUG Sb ces 139-140
babitsjand | ‘chisracteristies, samimary... 0/2... 2.2... 2. ee 11-14
Ob adultes. e222 PEO ee en ens oo Se. BS: ---- 99-102
IEP EXPoDS Ei 107 ST OF ie te a a 122-127
Hemipteroussenemiles: - 22280. Seer noe Se A 130
INISLOR RE BeSaretrre te ee Sn ek ee ons A de 146-154
“huts.’’ (See cones.)
nymiewopterous enemies: 22522 i. 22S s.r. ek. 130
THOT 2) pete Ue Sess See a pce eM i a ey 146-154
USCS, UB O22 TS) le net Eg i eS 129-138
Jaryas toodvnabitse:s-to-e- ce ee eee a 122-123
ise inidcee eee eer ees Sane Jes rem | 122-127
newly hatched, actions and description.............. 112
Sigees eseriponee seas. S25 ee ee oo oe 118-120
locationter stages im-seiled=* 220 Ses so 124-125
TMC MV ARANUOR TCO sree pee a ee eee eee 135-138
OULU ANB ar nme Cee eMC res ae ee ann oes oe 79-81
TUS CHA Ap PArabUe we es oe ek SL ae 82-84
HaLUrab CHE UIeds eee sett as Caw a MM MCs 127-140
UCIT X20 9ai8 roe) 5 cnet chs cede ag > ep Unt eae a 99-100
OMiposiionmemodude ts see tees ee oe SPT. 109-110
Tenino Np lambe ess ceca eke lo ee 106-108
CURPIPORILON pester a ees SG MIS. Se ee edo 81-82
plants oviposited in..........- Fe SiStae ek geri eater | eee 105
precautionary measures against injury.................-- 144-145
papa GceeriuOnee asset k! a oo a ence ee et 121
toocshallaieuere. ee Ree eS os aAash eS Ae 122-123
ITY OPTS Ro) Se Oa ca el a oh gS Dae ea cl OE ae 122-127
TACOS EE tsar ee pene 5 oem Se ew Foe eee PLES SS 14-20
FANE C10 Le TOOUS ere Mee Sek cers eee re teers La 2 38-77
of seventeen-year race... ..:.. 222. 52222255: 38-64
thirteen-year race! 2.22... f2k.. 2222.02. 65-77
species and -the-two races. ...::-...:----..-------- 30-37
remedies and -preventives...-..2.-.2.---...--.2--.52----- 140-146
repellents........- Pisin boca ea Ae rea AR es fe 143-144

‘roofs.’ (See cones.)
31117—No. 71—07——12

- 178 THE PERIODICAL CICADA,

Page.
Cicada, periodical, seventeen-year breeding record.........-..--....-----+-- 114-116
broods; relationships sass.) ee eee eee 29-30
PAN SC re 5526 8a55-8 staan seen eee 38-64
TEC OM Pane, 2 PREM RIE S 8 etree en tr RNS aid = 14, 16-18
SONG MOLES? 24 22s Be a aie Pata ha eat ees eee 84-86
Stages, Geacriptions 4c... <8 tien secre ee 118-121
histOby:<.2 2226 heradens Jed oil aoe tains eer 116-118
Leeation) tarsOpliies,. 2 ac-4 epee eae as eee 124-125
sting > so-callledinics2<..2 296.2010 a binge ae eee cericio sce 86-88
structural details: 2:2... 4e4eseeehomee ass eee Ree ee 78-84
differences between the two races. ..........-.-- 17-18
superstitions..........- Sirsa k fale Dee SE Ee 147
systematic positionc.-5$2203.-2.2g—- 2 Secs queen oes ee eee 77-78
thirteen-year broods, relationship...................------ 30
TANGO sO ae cee eee oe ee hee 65-77
race. (See also Cicada tredecim).....- 14-18
‘‘towers.’’ (See cones.)
transtonmationite adulitees << e 5.5.55 see te one eee 98-99
‘‘turrets.’’ (See cones.)
cand exonound lites ak. esas eee At ae espero i ago 112-121
experimental: jpiroais: 25.0 Saar. Sac nie 112-116
USE AM: Makino SOap 5.52 saa eee eee eee 104
MATLED Ys ere eo a ae din tants cee eee ee eee 20-22
Dros —C Cada TD tent eis Ee Nap eee ee ee 78
septendecim. (See also Cicada, periodical, and Tibicen septendecim. )
considered by Walsh a species distinct from C. tredecim.. -. Lz,
= Tibicen septendectw = 52-2 asc. 2244 5 Hatguaie Ho 8 3 ae 22
hbicen, Sphecvus speciosus anienemiy.-)- 2222. 2S aisieae ne 132
tredecim. (See also Cicada, periodical, thirteen-year broods and race.)
considered by Walsh a species distinct from C. septendecim. . 17
=Tibicen septendecvm, Race Megeclm...... 2 2. . 22 ss eee 22
first: publication oh aines 228 o5 eee eeeeeer 15-16, 153
Cicadas, yearly recurrence not necessarily indicative of life round limited to a
MEO. ofan nasa Se sais non le Saisie oe ce oie Ses = oi epi eee eee 12
Climate, in relation to races of periodical Cicada........----.-.---------+---+--- 18-20
“Coccyzus americanus, enemy of periodical Cicada......--..------+---+---+----- 138-139
erythrophthalmus, enemy of periodical Cicada...........------------- 139
Colaptes auratus luteus, enemy of periodical Cicada........--....-----+-+-+-+- 138
Corvus brachyrhynchos, enemy of periodical Cicada........-...-------+-------- 139
Cowbird. (See Molothrus ater.)
Crayfish, house-building habit..........-. Hannes! NE Cee a ETE SSO 91
Crow. (See Corvus brachyrhynchos.)
Cuckoo, black-billed. (See Coccyzus erythrophthalmus.)
yellow-billed. (See Coceyzus americanus.)
Dendroica cerulea, not found to eat periodical Cicada.......-..--- chs Sates Leyes 139
Development, retardation in eggs of periodical Cicada.......-----.------------ 111
Digger wasp, larger. (See Wasp, larger digger.)
Disease, periodical Cicada imagined cause.........-.--------------+----+-+---+- 147
Mishribution, of periodical Cicada- tse... acs = seine ee ees 34-77, 99-101
Dogs, enemies of periodical Cicada..............--.------ BN apr ee eo 104
‘Dragon-flies, enemies of periodical Cicada... ....-...--,<--++--2-==~++--++-+;-"- 128
Dryobates pubescens medianus, enemy of periodical Cicada....-....-.-.---- Pegs 1c

Evergreens, seldom oviposited in by periodical Cicada.....-..-.--.----------- 105

INDEX. 179

Page
Flicker. (See Colaptes auratus luteus.)
Flycatcher, crested. (See Myiarchus crinitus.)
Proit tees, Injury to roots by periodical Cieadats..2...2--...2...-.5--2--5-2- 126
Pumeous didesseron periodteal* Cicada. 225 00b 22-82... = 2 eee 139-140
Galeoscoptes carolinensis, enemy of periodical Cicada...............5.-+-2--.---- 138
Gnatcatcher, blue-gray. (See Polioptila cxrulea.)
‘Goldfinch, American. (See Astragalinus tristis.)
Grape root-aphis; bisulphid of carbon a remedy. .-.-...-..-.-.---+--------.--- 146
Grapevines, injury through oviposition of periodical Cicada..............-.-- 107
Grosbeak, cardinal. (See Cardinalis cardinalis.)
Ground-beetles, enemies of periodical Cicada:.-¢...-.1.--...........-2.----- 128
Hand collection, remedy for periodical Cicada...........-..--------------- 141-142
Harvest fly. (See Cicada.)
Helmitherus vermivorus, enemy of periodical Cicada....................-.----- 138
irekories- oviposition of periodical Cicada. -....2.-...--2----..-22s.22-.--- 105, 106
ieeewenemiesof periodical Cicadas. ..2.....2-.--2-2:-2.0+--.----- 91, 103, 104, 128
Hoploprora sp., enemy of periodical Cieada:-_.+.....-..-.---.....2--5---2-2-- 136
Honcapus virens, enemy of periodical Cicada:......-2.-....2-----2-..---02--: 138
Hylocichia mustelina, enemy of periodical Cicada.-..........-.--.----~.---.-- 138
Ichthxlurus punctatus, enemy of periodical Cicada.................-.-....-.-- 139
Percrus-galoula, enemy or periodical Cicadals.. 2... ssh 2-2 ee eee 138
SMiavus senouLy Ol perodical Cicada...-22. 2.2500). . e ke lee hee e ee 138
Indigo bird. (See Passerina cyanea.)
Mpursnovris, enemy o1 pemodieal Credda>)s0 02. 2255.22. ee ee Pe ST. 137
Kerosene ‘emulsion, against periodical Cicada.............--2----.....0..-- 142, 143
Lanivireo flavifrons, enemy of periodical Cicada....-...-..------ eS ee 138
Lathromeris cicadx, parasite of periodical Cicada..........----.------------ 131-132
Lime-sulphur wash, possible value as repellent against periodical Cicada... ... 144
washes, repellents against periodical Cicada.......:-...-.....-........ 143-344
‘*Locust, seventeen-year,’’ erroneous name for periodical or 17-year Cicada... . Lil
JLyOCRUSIS, BYS EW AFUE SISO C6 01 eee gra eC ore i ara 102-103, 148
Maple, hard, injury through oviposition of periodical Cicada........-..-.-.-- 107-108
Martin, purple. (See Preane subis.)
Massospora cicadina, fungous disease of periodical Cicada.....-......-----.-- 139-140
Melanerpes erythrocephalus, enemy of periodical Cicada...........-.......-.... 138
Merula migratoria; enemy of periodical Cieada_.....-2......2-----------...-.- 138
Mesodon elevata, feeding on dead periodical Cicada.......22..--.----.--------- 139
eroleta; ieedine on-dead periodical Cieaday.:°. 2... 2.2.2.2.) 228.2222. : 139
Micropterus salmoides, enemy of periodical Cicada.......-...-.-.------------- 139
moras dicen enemy ol penodical Cicada.: 222022 2.2... b2bc 2.2. oee te SSe4 138
myrarches crimcus,- enemy ol periodical’ Cicadat -:....2.052552.52.2-2--022 224 - 138
Memmarid:- sparasiteror perrodical:Gicadas». i222. -2.-2 2222222022. 2 Senses 130
Wiathrus omvonis, enemy of canker-worm:-..2-22.--2--.-2s.0-- 2-2-2222 eee 135
Nuthatch, white-bellied. (See Sitta carolinensis. )
Pere ovaporruomor periodical Cieaday 2.22 2 2)2 5222... - 2.20.2 S82 oe 100, 105, 106
pina pilosa, enenty of periodical Cicadat...2..-.. 2-222-022-2222. ee eee. 136°
euaimaspioiow, enemy of seale insects... =). 22-2. 202.228. .e e: 135
Sp etlemyron periodical Cicadas ec le .ce2. 2. ee ap Soe ee one ee 136
Pe batcllasp.. enemy of periodical’ Cicada... 22-22... . 22. ts eee ee 136
Gribatid mites, enemies/of periodical Cicada... .--...-2 2-2-2222 -2 lee 136
er euimiarsp> enemy’ of periodical Olcada: 9.222.222... 2-2-2 see) ee- 2% 136

Oripoda elongata, enemy of periodical Cicada.........-.....---.---.-+-2+-4--- 136

180 THE PERIODICAL CICADA.

Oriole, Baltimore. - (See Icterus galbula.)
orchard. (See Icterus spurius.)

Passer domesticus. (See Sparrow, English.)

Passerina cyanea, enemy of periodical Cicada

Pear, food plant of periodical Cicada

Pediculoides ventricosus, enemy of periodical Cicada
Penthestes carolinensis, enemy of periodical Cicada
Periodical Cicada. (See Cicada, periodical. )
Pewee, wood. (See Horizopus virens.)

Pipilo erythrophthalmus, enemy of periodical Cicada
Piranga erythromelas, enemy of periodical Cicada
Polioptila cxrulea, enemy of periodical Cicada
Poultry, enemies of periodical Cicada
Progne subis, enemy of periodical Cicada

Robin. (See Merula migratoria.)

Seale insects, Ortbata aspidioti am enemy 22." -..... con case ee eee
Schizoneura lanigera, injury to fruit trees following oviposition of periodical

Cicada he. 22 een SSO ee Ce, Bee ee ene
Sciurus rufiventer, enemy of periodical Cicada......-..-.------- aes

Seventeen-year Cicada. (See Cicada, periodical.)

locust, erroneous name for periodical or 17-year Cicada
Sitta carolinensis, enemy of periodical Cicada.............-.-.-.--..--
Smilax rotundifolia, oviposition of periodical Cicada attempted

Snails. (See Mesodon exroleta and M. elevata.)

Soldier-bugs, enemies of periodical Cicada...........--..-----------

_ Sparrow, chipping. (See Spizella socialis.)

English, enemy of periodical Cicada... :.....-----.-+---+-=:

Sphecius speciosus. (See also Wasp, larger digger.)

enemy of Creada tibicen 28-2 Se2 see eee
Sitizus- Grands a, SyMONYM: © 524 ee See ee
Spizella socialis, enemy of periodical Cicada.........----------+-----

Squirrel, fox. (See Sciurus rufiventer.)
ground. (See Tamas striatus.)

Squirrels, enemies of periodical Cicada........--------.--+---+-----+++---+---
Stizus grandis==S phecius spectosus 2.3.2.5 --22 32 2-eeees eee

Sucker, white. (See Catostomus teres.)

Superstitions, regarding periodical Cicada.......-...--.------------
Sweet gum, food, plant of periodical Cicada.............------------
Tachina sp., parasite of periodical ‘Cicada l= 32-20) <3) -2 eo. oe eee
Tamias striatus, enemy of periodical Cicada.....-.-....--.-------+----

Tanager, scarlet. (See Piranga erythromelas.)

Tettigonia septendecim, name of Fabricius for periodical Cicada

Thrasher, brown. (See Toxostoma rufum.)

Thrips, enemies of periodical Cicada.........---<.- -2 22.5222 stein =< oi

Thrush, wood. (See Hylocichla mustelina.)

Peach, injury through oviposition of periodical Cicada.........-..-.--
injury through oviposition of periodical Cicada..............--
to roots by periodical Cieadai 222-23. 52.< ste pea ee oe

Pycethrum, against) periodical Cicadas: s2.- 2.2. hee eee eee
Quiscalus quiscula exneus, enemy of periodical Cicada............-----

enemy of periodical Cicada.......-.-.- eee ee
Retarded development, in eggs of periodical Cicada.........-..--.---

Page.

" 138
107

104, 128, 148

138

103, 104

139

2 INDEX. 18]

Page.
Tibicen cruentifera and allies, mistaken for periodical Cicada.............-- 36, 45-46
FUL OS CARLIN S La eral O Rae en Se DUC ILOCCUTI ee ee tee ee eso ao ae oe ot eye ielge 63
septendecim. (See also Cicada septendecim and Cicada, periodical. )
correet name-for periodical Cicada.......2....----..+--.- PD,
race tredecim, correct name for 13-year race of periodical
(CEU oA ae ae a Ot Ae a 22
var. cassinii, correct name for Cicada cassinii........--.-- 22
Titmouse, tufted. (See Bxolophus bicolor.)
Mebacco Gist, apaiust, perlodieal Cicadas. <2. =o. cscaice ne osc ecce sae eee cence 145
Towhee. (See Pipilo erythrophthalmus. )
Hazossomaruyum, enemy of periodical Cicada... ---..5c%s. sek esee ese ee 138
Prichorrammid parasite.of periodical’ Cicada-<2-..252- 22.22.26 e532 eens. 130-131
Tyroglyphus cocciphilus, enemy of periodical Cicada........-.....--..-------- 137
Vireosylva gilva, not found to eat periodical Cicada.........-...--..-..------- 139

Vireo, warbling. (See Vireosylva gilva.)
yellow-throated. (See Lanivireo flavifrons. )
Warbler, cerulean. (See Dendroica cerulea.)
worm-eating. (See Helmitherus vermivorus.)
Wasp, larger digger, enemy of periodical Cicada.....................--- 132-135, 139)
Woodpecker, downy. (See Dryobates pubescens medianus.)
red-headed. (See Melanerpes erythrocephalus.)

O

U.S: DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 72.

L. 0. HOWARD, Entomologist and Chief of Bureau.

INFORMATION CONCERNING

THE NORTH AMERICAN FEVER TICK,

WITH NOTES ON OTHER SPECIES.

BY

iW. D. HUNTER anp W. A. HOOKER.

IssuED NOVEMBER 2, 1907.

WASHINGTON:
GOVERNMENT PHINTING OFFICE.

1907. Mae eA

U. S. DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY—BULLETIN No. 72.

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

INFORMATION CONCERNING

THE NORTH AMERICAN FEVER TICK,

WITH NOTES ON OTHER SPECIES.

1B Ne

W. D. HUNTER anp W. A. HOOKER.

IssuED NOVEMBER 2, 1907.

ees

IN

WASHINGTON:
GOVERNMENT PRINTING OFFICE.

OTE.

BUREAU OF ENTOMOLOGY.

L. O. Howarp, Entomologist and Chief of Bureau.
C. L. Maruarr, Entomologist and Acting Chief in absence of Chief.
R. 8. Currron, Chief Clerk. ¢

. CHITTENDEN, in charge of breeding experiments.
. Hopxins, in charge of forest insect investigations.
. Hunter, in charge of cotton boli weevil investigations.
. WEBSTER, in charge of cereal and forage plant insect investigations.
. QUAINTANCE, in charge of deciduous fruit insect investigations.
. Puruies, in charge of apiculture.
. RocErs, in charge of gipsy moth and brown-tail moth work.
. MorRILL, engaged in white fly investigations.
. Fiske, in charge of gipsy moth laboratory.
. Hooker, engaged in cattle tick life hisiory investigations.
. MorGan, engaged in tobacco insect investigations.
. WoeivM, engaged in hydrocyanic acid gas investigations.
. GILLISS, engaged in silk investigations.
R. P. CurRIE, assistant in charge of editorial work.
Mase Coucorp, librarian.

2

On> gar unr aap
Pe ais areas ote

LETTER OF TRANSMITTAL.

U.S. DEPARTMENT OF AGRICULTURE,
BurEAU OF ENTOMOLOGY,
Washington, D. C., June 19, 1907.

Srr: I have the honor to transmit herewith a manuscript prepared
by Messrs. W. D. Hunter and W. A. Hooker of this Bureau. The
manuscript is a study of the life history and habits of the North Amer-
ican fever tick, together with notes on other species. The work upon
which this bulletin is based was begun in July, 1905, after practically
all of the directors of the southern experiment stations had brought
to the attention of this Bureau the necessity of additional work on
the important parasite which transmits Texas or splenetic fever of
cattle. Prof. H. A. Morgan, director of the Tennessee Experiment
Station, has given valuable advice during the progress of the work.
The paper contains information of great value in the practical work
of tick eradication. I therefore recommend that it be issued as
Bulletin No. 72 of this Bureau.

Respectfully, C. L. Maruatr,
Acting Chief of Bureau.

Hon. JAMES WILSON,

Secretary of Agriculture. -

a el

Sa

oe Mp gaa ee

CONArE NGL Sy,

CONCERNING THE NORTH AMERICAN FEVER TICK.

Page

UPCINET ORCUTT oe ee Pho ya eae EAS 9

Poser occasioned: Dy. Une, Gapule Weekes 2. oe Ue ee oes ee og eee ee 11

ine eshisiory, Obdeks wm generals. )-2-es22 252. 2222 ss de aoe. t 2-2 22.4 12

The life history of the cattle tick... .-.-. og Sy I IE aR OE se 13

Rested previous Low POSItOMe 2 0 bac ks Gee be es oe an ihe Pascale 14

SSL LINC TNON 02) Los Lae SRE EC AE onl ae ae ae LE 14

SEES (SES a 2 che I ae yah tg at Sd ee et ee 15

TC Ufo GLO Mee rree er tae toe eee ee ane tend, See sche ee cate Saas M7

Relation: of temperature to imcubabion...:..2222..:..-22----+---2.-: 19

Biecv.ot heatand.coldsonveresss 22220 F552 04. .o\ oe oae cee Shae 21

Sipmercence of eges) iawaler..- 2 sc. 222 oo... o2 oto 2 oso ads sos ons 22

Percentageor eres banehines 2.2222 es So on eee Sea cle 23

Mermabnr Nea -llekr same ass) ens she aah ore es se BSG as eR tne 23

ROU paeAsiules OMude mare ey a me sawn rote oC Lae we oak adie. Sats 23

ee Or WalOrOMm need UbCkS: ear se See SA es ie bak 2 oe: ae 24

Gn evita OUSCER LICKS). pices. J Suse kee asian Sees oe ha 25

PST RESUS OVS a Leek NL SL, Me am A RA a ea pee ere 27

De alovmment On NOS asa. o. aees Ole ee AI Sy tothe wa CES Sah 8 a als 28

oho TDSLLIS EIRP Sap LS aT en ne eee Ge ee ere en 30

Effect of continuous cold and heat on engorged females...........------- 31

BiteePorndimeehsUnMont, Ol-AC UGS S. 05228 s 6 ee er oe Falak a a clea slate a2

Effect of submergence in water on engorged adult ticks. ......-..----.--- 32

Hihres pune isi MOS yor see od kee et Se tee els ae ow 33

Hsiccomm ere ergs ee SEP eps eee cee ay ee Piacoa eLearn 33

IGHtenelamonn ar the eattle tie: wie "0 nee) oe ec Sees kee oboe 34

Relation between ration and tick infestation.............-....-------------- 35

OHM ROt aC kee amaiti< 9b Gum e eee a rar eas tee Lal OEE Ye! ce Nero eae 36

The practical application of the information recorded in this bulletin... ...--. Mijn MO
NOTES ON VARIOUS SPECIES OF TICKS FOUND IN THE UNITED STATES.

ie anneaLion: sau IU OL tikes a5 eevee coe ee So ae eS Sees eee wee wee 40
Key to families, subfamilies, and North American genera of ticks

(Ixodoidea).. - - - bg Datel eG ae a ee Ge ee a a ge El ae oe 40

IS Arnal Stu ge Feyd ee SM oe Oe Ami OR ee ee ee eee ee 41

CSTETS LATCH ays 2 a pa ad neal alk ice) Og SO ER Rees ee Ee 42

Genus Ornienae OfOsuee ei es SG eo oe we Shae ewe as ae Be eae 45

SST ISS 2 alt A gel ge 46

Subiamullyenonipicepnalitce:. 22.422. 8.2) 2lne Sse 32 ce See eS = 47

USrne) MTES al no] TIO, OST GI Ls FS i oR ee eR eR IE he 47

GenignpNanmaTO mus aa ten eee a els eA ne se oe Ae ee ee 49

Ce TPCT AC CMLBOE yo aye es Mal 2 oc A a aie bd SN aS ye ogee 49

Gonnsrlcemap iy Ralint cose 048 ts naa ra ns ees y eee cae 52

SHU Saeraren Aya oC ees SS i a ge ee er aM 54

(GReTET SCAU ch) haa So ee ey ry ar ee oe Se ee 54

Gemuse Nam phy OMAeyes yee sae eee ass 2 ra)2 Bala = Salsa sl acale Sierinr whe Eee 58

Mera ini seo G asec et fae ctl 8 Sk 533) bes ead - seins 64

TEL RpMROIN ere, Seas Tee er eS Re Pen SSE (banal eS oie eo cle aladia a eee iif

Prats I.

Fia.

Ie

Ill.

TVs

—

oS OF ke HS bo

ee
HEH Oo oN

eae
vw bo

ILLUSTRATIONS.

PLATES.

The North American fever tick. Fig. 1.—Eggs of the tick, Marga-
ropus annulatus, deposited under stable litter. Fig. 2.—Seed
ticks of Margaropus bunched on grass and stake.......--------.-

The North American fever tick. Fig. 1.—Steer used in experimental
work. Fig. 2.—Arrangement for obtaining data on incubation. .-

The North American fever tick and other species. Fig. 1.—Marga-
ropus annulatus, male. Fig. 2.—Hxemaphysalis leporis-palustris,
female. Fig. 3.—Stigmal plate of Margaropus annulatus, male.
Fig. 4.—Mouth parts of Ixodes cookei. Fig. 5.—Stigmal plate
of Rhipicephalus sp., male. Fig. 6.—Stigmal plate of Amblyomma
LOCLULCL LOD RENIN Al Coates Nip yep! We eee) tie So mae WE Tt sep a cle

Stigmal plates of ticks. Fig. 1.—Stigmal plates and anus of Derma-
centor nitens, male. Fig. 2.—Stigmal plate ofsame. Fig. 3.—Stig-
mal plate of Amblyomma cajennense, male. Fig. 4.—Same, female.
Fig. 5.—Stigmal plate of Dermacentor variabilis, female. Fig.
6.—Stigmal plate of Dermacentor occidentalis. .......-.------------

TEXT FIGURES.

Margaropus microplus: Genital apparatus..........-.-----------------
. Graphic table for the separation of the families and genera of ticks -- - -
REnemeephatus spe Capitulum ob fenmile}: 227-2)... 5252-2 - s22s-
. Rhipicephalus sp.: Coxe of male and female ..........-.-.-----------
. Dermacentor mitens: Capitulum. of female.....-.+.--.....--------+--- :
Dermacentor ‘nitens: Coxe of male and female..................------
. Hxemaphysalis leporis-palustris: Capitulum and scutum of female. ...--
. Hxmaphysalis leporis-palustris: Coxe of male and female .....-.------
. Amblyomma cajennense: Mouth parts of male......--..---------------
. Amblyomma cajennense: Coxee of male and female..........-----------
. Amblyomma maculatum: Mouth parts of female.....-.-.--.-----------
. Amblyomma maculatum: Coxe of male and female.....----------.-----
. Amblyomma maculatum: Scutum of female......----------------------

Page.

16

16

48

48

INFORMATION CONCERNING THE NORTH AMERICAN FEVER TICK,
| WITH NOTES ON OTHER SPECIES.

INTRODUCTORY. ‘4

It is safe to state that no more important problem than the eradi-
cation of the cattle tick (Margaropus * annulatus Say) confronts the
farmers of any country. Not only the cattle-raising industry, but
the whole economic condition of a large section of country is affected.
The tick, without any but the most limited power of locomotion,
and for all practical purposes dependent upon cattle for its existence
and dissemination, presents a problem in eradication of a hopeful
nature. Cattle are under the control of man. Therefore, the prob-
lem is quite different from that involved with other pests, like the
boll weevil, which by flight spread over large areas of land. In the
one case absolute eradication is possible and in the other it is out of
the question. In fact the possibility of the total extermination of
the tick in this country is by no means visionary. It was foreseen
originally, probably, by Dr. Cooper Curtice, who wrote as follows in
1896: ‘“‘L look most eagerly for the cleansing of even a certain portion
of the infected territory under the direct intention of man, for it
opens the way to pushing the tick back to the Spanish Isles and
Mexico, and liberating cattle from disease and pests and the farmer
from untold money losses. Let your war cry be, Death to the
ticks.”’> ;

In view of these facts it is evident that the most complete knowl-
edge of the habits and life history of the tick is of the utmost impor-
tance. All means of eradication must depend upon such knowledge,
and improvements in present methods must depend upon additional
information regarding the tick. Dr. Cooper Curtice, who will be
quoted frequently, because he has been among the foremost in the
study of the problem, has written as follows: ‘‘To the scientist
studying the tick to learn its life history, habits, form, and anatomy,

@ Neumann has shown that the generic name Boophilus of Curtice must, in obedi-
ence to the zoological law of priority, fall as a synonym of the earlier name, Marga-
ropus of Karsch.

6 Journ, Comp. Med. and Vet. Archives, Vol. XVII, p. 655,

10 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

the fact that these animals are pests to the stockman throughout
the greater part of the year is of very little importance, while the
latter cares little about such matters if he can only learn how to rid
his cattle of them. Yet it is only by learning the life history that
remedies to prevent them-can be applied intelligently, and the fact
that the knowledge attained is of practical value adds a double
interest to their chek a

In view of the evident importance of the work it is surprising that
so little has been done in this country. In 1892, about a year pre-
vious to the issuance of Smith and Kilborne’s epoch-making bulletin
denvonstrating the tick transmission of fever, Dr. Cooper Curtice
published the first data regarding the life history of the cattle tick
as Bulletin 24 of the Texas Agricultural Experiment Station. It
-was accompanied by excellent illustrations. The value of this work
will be understood from the fact that it was of a pioneer character,
and that all subsequent work has depended upon it. Nevertheless,
it was of a preliminary nature and merely outlined matters that
must eventually receive the most careful investigations.

Prof. H. A. Morgan, principally in bulletins 51 and 56 of the Louis-
iana Experiment Station, has added greatly to our knowledge of the
cattle tick as well as other species. His work ‘has such practical
bearings that it has been the chief indication of the value of life-
history studies in pointing out successful methods of eradication.
Recently Messrs. Wilmon Newell and M. S. Dougherty, of the Louis-
lana crop-pest commission, have published a valuable contribution
which still further shows how every fact relating to the tick can be
utilized in combating it.

The above are the principal publications by American workers.
There are many others which also contribute important facts. Among
these are Connaway’s, Schroeder and Cotton’s, Ransom’s, and others.
In other countries excellent work has been done on related forms. In
South Africa Prof. C. P. Lounsbury has made scholarly studies of
Margaropus (Boophilus) decoloratus and many other species. In
Argentina, Dr. F. Lahille has recently published the results of some of
the most exhaustive work on ticks that has been done. These works,
with others, are listed in the bibliography at the end of this bulletin.

Notwithstanding the studies that have been conducted in this
country, it must be stated that our knowledge of the tick is far short
of what it should be. There is a lack of knowledge of local variations,
due to climatic influences, as well as such matters as dissemination.
To supply this deficiency, the Bureau of Entomology, in cooperation
with the officers of several experiment stations, has undertaken a

4Tex. Agr. Exp. Sta. Bul. 24, p. 238.

LOSSES OCCASIONED BY CATTLE TICK. 11

careful study of the tick. Some of the results of practical bearing
are given in the following pages and others will be published from
time to time.

The writers desire to express their thanks to Prof. H. A. Morgan,
director of the Tennessee Agricultural Experiment Station, for many
most valuable and courteous suggestions in the course of this work.
He has turned over to the writers many of his original notes and has
generously assisted in numerous other ways.

LOSSES OCCASIONED BY THE CATTLE TICK.

Undoubtedly the popular idea of the damage caused by the cattle
tick concerns itself with the actual death of cattle from the disease
transmitted by the tick. Although this is a very important matter
and would. fully justify the most energetic attémpts toward the
eradication of the tick, it is really unimportant in comparison with
the other losses. Mr. August Mayer, a practical cattle breeder of
Shreveport, La., and Dr. J. R. Mohler, of the Bureau of Animal
Industry of this Department, have made most careful, comprehensive
estimates of the losses caused by ticks. The following summary is
taken largely from their writings: ;

1. Loss by death from disease in young animals and those removed
from temporarily tick-free localities (as, for instance, in cities) to
places where they become infested. The enormous loss under this
heading will be understood when it is recalled that every bovine
animal in the tick area must suffer an attack of fever if it becomes
infested with ticks. In an instance that came to the attention of the
writers, 39 out of 40 calves dropped in a city died of tick fever when
removed to an infested pasture.

2. Loss in weakened condition and stunted growth caused by the
fever.

3. Loss by gross tick infestation. At the present time (March,
1907) hundreds of cattle in south Texas are dying from gross infesta-
tion resulting from a mild winter. In extreme cases, Mr. Mayer
estimates that as many as 200 pounds of blood may be withdrawn
from the host during a singie season. This makes a gain in weight
impossible even in the best of pastures. Moreover, Prof. H. A.
Morgan and other observers believe that gross infestation and the
consequent general debility induce acute attack of fever even in
animals ordinarily immune.

4. The tick makes hazardous the importation of pure-bred cattle.
This prevents the upbuilding of southern cattle and at the same time
largely deprives the northern breeder of a market that he should have.
Moreover, the inability of the southern breeder to exhibit his stock in

12 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

the north and of the northern breeder to exhibit his in the tick area is
a handicap, the importance of which will be readily seen.

5. The necessary restrictions in the shipping of southern cattle
also handicap the breeder and affect the price.

6. The maintenance of the quarantine involves considerable annual
expense for the protection of the cattle owners north of the line.

7. Minor losses may be grouped as follows: (a) In Texas, especially,
the tick induces the attack of the screw-worm fly (Chrysomyia macel-
laria Fab.); (6) there seems to be, as pointed out by Mr. Mayer, a
considerable interference with the fecundity of infested cows; (c) the
railroads are put to the expense of disinfecting cars and maintaining
separate pens and the stockman to the expense of dipping—items
which react on the price that southern cattle bring.

All the losses that have been mentioned total approximately
$100,000,000 each year. At present the loss, as indicated by Doctor
Mohler, amounts annually to at least 10 per cent of the value of the
cattle. The quality of the animals is the lowest and the loss is greatest
in the regions where the natural conditions without the tick should
produce the finest cattle with the least loss. But the damage may be
better expressed by the statement that the tick makes profitable
production practically impossible in the South. Any successful
system of agriculture must rest upon a diversification of crops, and
this, in turn, depends upon animal husbandry to maintain the fertility
of the soil. Therefore, until the tick is eradicated or placed under
control, a rational system of agriculture in the infested area is out of
the question, and that achievement would mean almost as much to
the North as to the South.

THE LIFE HISTORY OF TICKS IN GENERAL.

The following general statement regarding the life history of ticks
is taken from Salmon and Stiles: “

Ticks are temporary parasites, attacking mammals, birds, and reptiles. They do
not appear to be so strictly confined to certain hosts as do parasites in general. Stiil,
this may be more of an apparent than a real rule. Certain it is that, although a given
tick may be found occasionally on animals which are very dissimilar (dog ticks have,
for instance, been found on snakes), still the various species show a decided predilec-
tion for certain hosts.

The parasites copulate during the period of parasitism > and suck the blood from
their hosts. The female grows to a large size and eventually drops to the ground and

a Seventeenth Ann. Rept. Bureau of Animal Industry, U. 8. Dept. Agric., p. 398.

+ This is not invariable. Amblyomma americanum sometimes copulates soon after
the second molt, but before it has gained a host. It is likely that other species also
occasionally do so.—W. D. H. and W. A. H.

LIFE HISTORY OF CATTLE TICK. tS

lays numerous eggs, which are usually more or less clustered together. The larva
upon hatching possesses three pairs of legs, the fourth pair being added during the
first molt. Either the hexapod or the octopod form may attack its host.

From the foregoing it will be seen that the cattle tick, like other
ticks, passes through the following stages: Egg, larva (six-legged
form), nymph, and adult.

The eggs are nearly round, dark brown in color, and deposited in
large masses, held together by the gummy secretion with which the
female coats each egg as it is deposited. The next stage, known as
the seed tick, differs remarkably from the later stages in the fact that
six instead of eight legs are present. The stigmata are located be-
tween the second and third coxe, just anterior to the third coxe,
and problematic indications are seen between the first and second
coxe. No distinct genital or anal opening can be seen in this stage.
The anterior legs are much larger than the others. They are waved
violently through the air when the seed ticks are disturbed either by the
approach of a host or in any other way. After some time the seed
tick molts and the next, or nymphal stage, is provided with eight
legs. The absence of the genital opening will differentiate this stage
from the following one. Ticks in the nymphal stage are frequently
referred to in the South as “yearling ticks.’”? After a second molt
the adult form is reached. Copulation then takes place, and after
engorgement the female drops to the ground for the purpose of depos-
iting eggs.

THE LIFE HISTORY OF THE CATTLE TICK.

As pointed out by Morgan the most important fact about the cattle
tick (Margaropus annulatus Say), from the standpoint of practical
control, is that the time of development on the animal is always shorter
than the total of the preoviposition, oviposition, and incubation
periods. This gives the farmer an opportunity to free his cattle and
pastures of ticks by the same process of rotation. As a foundation
for the surest and most economical procedure an accurate knowledge
of the variations of the periods in the life history of the tick under
different conditions is absolutely essential. Our effort in this bulletin
is in a measure to supply this information. The work has been prin-
cipally to obtain data necessary in the pasture eradication and feed-
lot systems of eradication. We have consequently studied the devel-
opment of the tick both during its existence on the animal, by means
of a steer procured for that purpose, and during its life, under various
conditions, when not attached to the host.

14 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

PERIOD PREVIOUS TO OVIPOSITION.

The cattle tick, like other species, passes through a distinct period
between the time of dropping from the host and the beginning of ovi-
position. When the tick drops, the eggs are not ready to be depos-
ited, but must pass from the ovary through the oviduct. (See fig. 1.)
Thus there is a definite physiological basis for a period which has a
very practical bearing on plans of eradication that depend upon a

a

Fig. 1.—Genital apparatus of Margaropus microplus: a,
Position of eggs at time of dropping of tick from host;
b, position of eggs when oviposition begins. Highly
magnified (redrawn from Lahille).

knowledge of the exact time
to be allowed in removing
cattle from one inclosure to
another. lLahille has used
the term “prootoquie”’ for
this period, but we shall refer
to it merely as the preovipo-
sition period.

_As will be seen from Table
I the preoviposition period
ranges from 2 to 40 days,
depending upon tempera-
ture. In the summer it
averages between 3 and 4
days, and in winter over 20
days.

It niight be supposed that
the data in the table referred
to show a preoviposition pe-
riod longer than normal on
account of the removal of
the ticks artificially. How-
ever, only ticks about to drop
were selected, and repeated

tests with ticks actually dropped showed that the method followed

gives the natural preoviposition period.

OVIPOSITION PERIOD.

As will be seen from Table I, the period occupied in oviposition
ranges from 6 to 70 days, depending upon the temperature. In the
summer it averages 10 or 11 days, while in the winter it is two or

three times as long.

LIFE HISTORY OF CATTLE TICK. pd

TasBLE I.—Oviposition of Margaropus annulatus, July, 1905, to July, 1906, at

Dallas, Tex.
= Preovipositign Oviposition aes OU OL: Number of eggs per
S period. period. position. tick.
fe) e = so
1 = ; r : an ; l
When collected.) © E sh lseye |) AE fee) Mshee Se lord cs 3 Sr all rses
| ke 5 of | 3 | 28] 3 Sh easy oe a a oie
Poors ee eee metall wey sla |g) e. lige
Simalioceee |e Henene eal e eye ISOs lies Oe Tapes | oe a= OS
5 is Ah sie |e Sh | Poi | Sioa iva a fe o>
3 A E Ss |ta & | 2s s i =e
4 = Shes = = = a |= = =
|
1905. Days.|Days.|Days.|Days.|Days. Days.|Days.|Days.|Days.| Eggs. | Eggs. | Eggs
REEL Weoley St oe 2 18 2) 229 13 6] 8.2 14 8 | 11.0 | 3,806 515 1,185
PATI NIS te ae ol < 4 5 3} 3.5 11 7} 9.0 13 11 | 11.8 | 2,228 | 1, 466 1,971
So] 0X hom cee ea 10 4 Sul 3.4 18 12' | 15.0 20 TV at SWE (Oe le Seal gee See ee sageel ga
Si this Wioroseeene 6 3 3] 3.0 19 19 | 19.0 21 21 | 21.0 | 3,875 | 2,365 2,576
(O Yeh in) ee ee see 15 6 4| 5.1 44 21 | 30.7 48 21 | 32.0 | 2,311 694 | 1,795
INOWalOmes.ccce sss 15 15 10 | 11.9 64 21 | 39.9 79 33 | 50.8 | 2,689 805 1, 891
WCGHOE Sere: ae 14 41 21 | 28.6 70 20 | 42.6 90 55 | 71.0 | 3,946 464 1,779
1906.
HED: Ons= seo: ua 2 8 39 18 | 21.8 47 8 | 29.8 58 26 | 50.5 | 2,134 147 695
Relive Zima te soe 9 40 13 | 18.7 41 30 | 38.9 53 38 | 47.8 | 3,496 10 2, 009
1 Is eR 10 10 9 | 9.8 29; 12) 23.4 38 21 | 30.2 | 2,437 | 1,118 1,941
PATST VOL seo ete tan. < 10 6 3) 4.8 207) Li 2168 30 20 | 25.7) 2,260 | 1,281 1, 658
i:\ OY lp’ gee Seas 10 6 Di Onde finn LS 11 | 19.6 22 16 | 20.0 | 3,412 | 2,197 2, 891
Miya sea c\a= 5 = 10 10 6] 7.8 17 11 | 14.6 24 18 | 21.4 | 2,676 | 1,701 2,251
Misi 22 les Si ote Hee LO 6 4] 4.3 15 10 | 12.6 18 15 | 16.4 | 3,180 | 1,391 2,250
mame S a. ss. 2e. 10 5 3] 3.5 13 9 | 10.9 15 12 | 13.4 | 2,881 152 1, 802
Iota?) 0 Ee ea here) 4 3 | 3.5 14 ON ase, 16 8 | 14.2 | 2, 467 843 1,950
PRY eke oe so xe | 10 5 Ceca) 14 9 | 11.0 16 12 | 13.9 | 2,292 | 1,135 1, 837
ADT 5 la seer ie 10 4 Bia eae 13 7} 10.4 13 8 | 12.5 | 2,397 | 1,666 2. 068
Gey Oph oat CRs oN a De a em rel ye See ak) PRP PRE ea 32, 499
PATENT: ETC ee a Leeper | Denia | pp MeN WUT |e LSU | Cea SOE ENE Oe Pe hae oe aes PAu?

From Table I the following important, practical data are obtained:

1. The preoviposition period ranges from about 3 days in summer
to as many as 28 days in winter.

2. The oviposition period ranges from between 8 and 9 days in
summer to 42 days in winter.

3. The total period from dropping to the end of oviposition ranges
from 11 days in summer to 71 days in winter.

It should be noted that Table I gives the total period from drop-
ping to the end of oviposition based upon the weighted averages
of the preoviposition and the oviposition periods. Therefore the
maximum total period may be somewhat longer than indicated, as,
for instance, in cases where either the preoviposition or the oviposi-
tion for some.reason are prolonged beyond the average.

EGG STAGE.

The eggs are generally elliptical, but vary in shape on account of
pressure and drying. In color they are at first honey-yellow, but
soon change to a deep yellowish brown. They are shiny and
smooth. The average size in a lot of 10, measured by a micrometer,
was 0.54 by 0.42 mm. About the middle of the incubation period
in many species a whitish spot appears on the eggs and becomes more
conspicuous as the time for hatching approaches. This spot is

16 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

located toward one end and seems to be due to the excretion of- the
embryo. In Margaropus annulatus it is very conspicuous and is a
_certain indication of viability.

The act of oviposition is most interesting. This process was
referred to by Dr. Cooper Curtice.¢ An analogous operation in
Ixodes ricinus has recently been carefully described and illustrated
by Wheler,? and was earlier noted by Lewis. One of our asso-
ciates, Mr. R. A. Cushman, has observed the operation. The follow-
ing description is based upon his notes:

When oviposition is about to take place the capitulum is bent
downward toward the genital aperture. This exposes a delicate,
viscid membrane between the capitulum and the scutum. ‘The
membrane becomes distended and is projected out over the capitu-
lum in two rounded lobes, practically covering it. This process is
repeated several times before the egg is finally ejected, the mem-
brane being extruded and retracted alternately while the capitulum
is lowered and raised. Finally the white, membranous ovipositor is
extended, turning inside out, until it Hiclitee the distended mem-
brane. The capitulum is now completely hidden. As soon as the
ovipositor and membrane have come in contact the former slowly
recedes, leaving the egg adheri ing to and partially enveloped by the
membrane. The egg remains in this position for a varying length of
time. Then the membrane is withdrawn, rolling the egg along for a
short distance on the dorsal surface of the capitulum. At the same
time the capitulum is raised. Then the processes of distention and
contraction of the membrane and lowering and raising of the capitu-
lum are repeated several times, the egg being finally completely
coated by the viscid substance from the membrane and being finally
pushed back and deposited on the anterior edge of the scutum.
Each egg is laid in this manner, the tick backing slowly away and
leaving the mass of eggs in front of her. The actual time consumed
by the tick in laying a single egg is about 30 seconds, while the
removal of the egg and the resting period consume from one to sev-
eral minutes, a much longer resting period being taken at intervals
between lots of from 10 to 50 eggs. It has been impossible for us
with the means at our command to demonstrate the ‘paired, race-
mose glands” of the membranous sac referred to by Curtice. As far
as we have been able to see, the substance with which the eggs are
coated is secreted from numerous minute glands scattered over the
surface.

a Tex. Agr. Exp. Sta., Bul. 24, p. 242.
b Journ. Agric. Sci., 1906, p. 405.
¢ Proc. Roy. Micros. Soc., 1892.

PLATE |.

Bul. 72, Bureau of Entomology, U. S. Dept. of Agriculture.

Fi

g. 1.—Ege:

sof

Margaropu

s annulatus de

THE NORTH AMERICAN FEVER TICK.

posite

d under stable

(

litter.
Original

Pig
-)

(py)

.—Seed ticks of

Marg

ar

opus bunche

Sk ed

d on grass and stake

PLATE II.

Bul. 72, Bureau of Entomology, U. S, Dept. of Agriculture.

if

t
fs

a

i

THE NORTH AMERICAN FEVER TICK.

Fig. 1.—Steer used in experimental work. Fig. 2.—Arrangement for obtaining data on incubation.

(Original.

—_

LIFE HISTORY OF CATTLE TIOK. 17

In the manner described a mass of eggs grows steadily in front of
the tick, while its body becomes correspondingly smaller as the
process proceeds. The gummy secretion holds the eggs together so
that the mass looks not unlike a large accumulation of minute brown
beads.

The number of eggs deposited varies greatly with the size of the
female. The highest number recorded in our experiments was 3,806.4
The average of 189 ticks under various conditions was 1911.7, and
this probably very closely approximates the average under natural
conditions. The daily average, of course, varies also. The maxi-
mum is generally reached from 7 to 9 days after deposition begins.
The highest number for any 24-hour period was 826. The average
for 20 ticks was 144.

INCUBATION.

Most important means of control of the cattle tick depend upon
taking advantage of the fact that eggs remain on the ground for a
considerable time before hatching. Provided there are no seed ticks
present, it is perfectly safe to allow cattle in areas in which ticks may
be dropped from them, as, for instance, in fields under cultivation
for one crop season, if the animals are removed before hatching takes
place. It will be seen that this has an important bearing on the
process of relieving cattle of ticks by placing them for limited periods
in different tick-free inclosures. Accordingly we have made an
especial effort to obtain data regarding the period occupied in incu-
bation under different conditions and in different seasons.

In 1905 a number of experiments to determine the length of the
incubation period were conducted, the eggs being placed in paper
pill boxes. Subsequent work showed that this arrangement gives
more rapid development of the embryo than takes place under normal
conditions. Especially is the period shortened when an abundance
of moisture is furnished. In these experiments in July and August
the eggs hatched in from 17 to 21 days, and during September in
from 25 to 44 days. These results are of value only in showing how
the incubation period may be shortened under extreme conditions,
which must rarely or never occur in nature.

No eggs deposited in October, 1905, hatched before April 10, 1906,
a period of over 170 days.

a The greatest number found by H. A. Morgan was 3,198 (La. Agr. Exp. Sta., Bul.
d1, p. 242), but Newell and Dougherty record 4,124 in one instance (La. Crop Pest
Comm., Cire. 10, p. 23).

5795—No. 72-07

9
Z

18 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

In 1906 a more extended series of experiments was instituted. In
the first series, consisting of 59 lots deposited from February 20 to
September 21, eggs were kept in pill boxes. As has been pointed out,
this method accelerates incubation further than natural conditions
are likely to. The following is a summary of the results of these
experiments:

Eggs deposited in March hatched in 74 days. .

Eggs deposited in April hatched in 43 to 53 days.

Eggs deposited in May hatched in 24 to 33 days.

Eggs deposited in June hatched in 20 to 22 days.

Eggs deposited in July hatched in 19 to 22 days.

Eggs deposited in August hatched in 19 to 21 days.

Eggs deposited in September hatched in 23 to 154 days.

Eggs deposited October 1-7 hatched in 135 to 139 days.

Eggs deposited October 15 and later have not hatched to date (February 20, 1907).

In the second series of experiments relating to incubation conducted
in 1906 a special effort was made to provide conditions that would
give approximately the same period that must occur under normal
conditions. (See Table II.) To do so some eggs were placed on soil
in open-bottom glass tubes in the open air (see PI. I, fig. 2) and shaded
for a portion of the day. Others placed under the same conditions
were exposed to the sun at all times. In the third series the eggs
were located in an outdoor thermometer shelter where they were pro-
tected from sun and rain. The separate lots consisted of several
hundred eggs. They were deposited by different females in the
laboratory and placed together for the observations under considera-
tion. In this way a large number of eggs deposited within the same
24-hour period was obtained. It is supposed that these conditions
approximate closely to those which surround the great majority of
eggs in pastures. The following is a summary of the data contained
in the table:

Eggs deposited in April hatched in from 39 to 54 days.

Eggs deposited in May hatched in from 27 to 33 days.

Eggs deposited in June hatched in from 21 to 28 days.

Eggs deposited in July hatched in from 22 to 26 days.

Eggs deposited in August hatched in from 23 to 32 days.

In September eggs deposited prior to the 18th hatched in from 23
to 76 days. Eggs deposited after September 18 have not hatched up
to February 20, with but two exceptions. These were eggs deposited
October 3 and October 7, which began to hatch on February 18.
These eggs were placed in such a way that they were more exposed to
the sun than other lots that have not hatched, deposited before and
since the dates mentioned.

LIFE HISTORY OF CATTLE TICK. 19

TasLe I1.—Period of incubation of Margaropus annulatus at Dallas, Tex., 1906, under
various conditions.

| Minimum | | Minimum
Eggs deposited. Hatching. | incubation || Eggs deposited. Hatching. | incubation

period. | | period.

|) Days. || Days.
AD Ke per assess < VIC See eee a= | Gia AUIS Alas ars eee 28
‘Neves OS Beene I Migiv; 30 ooh 8-5 Onan | eA ee Ace ce) Se aa 3 b24
Apr. 15 Dv i iy A a ee CALA ATI ES a eee 3 b 23
PATINA yeti Sa. A eb ovetw: Vee ea ae GOON PAUP See az eet a 32
Misa sien nek Ja Jumesl4-* oe Ai Gon |e aslo sey e aaa b23
Wa Gl ee ee [fybire ve bauer een pe weal UDO) Ni ACO) ee Se c24
IMIG ORL ae eer nigeaes JUNC QE eae 2s). H WON ANG. Dee Ales a23
EIN C 1 eae FUMEWG Ss eles | GORE AN Gy 22 oe ae atsae c24
Misvaalec< ees. ine ease an | GOR || mAu ee 4 a oe b24
UTI Oe eee ate oe NbN asi8 Je So toe ee GD Teal AUS Omnis ce te oe b24
Tit: Sik an ae Ti liya bee ae Sone | CIGu eos ee c25
STOLL = ate i oss Dall varieees =e eee os | DO Tale ANE) 2S Sp eee ae a27
PURLEL Al ee = ses </> 5 TUL VALS eee COOSA WTS (:5 0) reall Ie ea ee ¢ 23
MMC oe 5 Jule Se es = | OBS Steyoll te Ne eee Beet | b 24
lbs (2, ul yes Gee seers @23\| Sept. 1.---.---.--. a27
pum wes. es Sek Oily pul Geese ees GQat He SODibe ass 2s. scene c25
dime s0u. fs. 2-222 Jilly 20 Meee ee BOT Sop tuoee ten 2 sate. c29
Dyes hese Sade te Daily Zoe eee B22 SOP Ono: seca es b 32
Pinliygs 20S Se orsectecis © Anes Oe Sates b 22 | Sept. 9....2....... a37
‘Gila Osta Hgneee Tues STS | COG NeSep tiltecenes eee c41
JUlye QOS oh ese ce PAC eA Sota Sr c24 | Sept. tse cane eces b 43
NEU OE) a aes ae TA 2D o seeps WOR RISI=| Olin te ae Bee sere a53
iibthy Biles eee ae Aiea OO Meta See COPA NSepital Saeeen tee we b76
SUbY tell mS ae Aug. 237 - 2a) Sept. (8.2205. 2s. a76
VHS a a ee Sept: 4e-cos e222: OP ipsa Pia la eo we b148
ENTE 0) sei eae Sep tesseeee se = c23

a The eggs were placed in open-bottom test tubes in sand in the open air. They were protected from
the direct rays of the sun at all times by a cheese-cloth screen. Up to 11 a. m. they were
within the shade of the house.

b In open-bottom test tubes in soil exposed to sun at all times.

¢ In open pil! boxes outdoors protected from sun and rain at all times.

RELATION OF TEMPERATURE TO INCUBATION.

In the series of experiments just referred to accurate data on tem-
perature were obtained. Standard maximum and minimum ther-
mometers kept in an instrument shelter were used. In Table III
the records of temperatures are given together with the average
incubation periods for the various lots of eggs under observation
during different months. The data show that there is an intimate
relation between temperature and the period of incubation. The
shortest average minimum incubation period (23.4 days) occurred
when the average temperature was highest (80.2° F.). The longest
average minimum incubation period (137 days) occurred with the
lowest average mean temperature (53.2° F.): Between these ex-
tremes there is a graduated correspondence between temperature and
incubation.

20) NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

TaBLE II1.—Relation of temperature to period of incubation in Margaropus annulatus
‘ at Dallas, Tex., 1905-1906.

| Tneahation 1 ae
THenbesion Re = aoe He DESH:
| No. period. | iat e
Month deposited. | of | Mean. Total effective.
hlOts=) |e sen aes | A Ae ir eterna a
Maxi- Mini-| A ver- Maxi- Mini-|Aver-- Maxi- | Mini- | Aver-
mum. mum.| age.@z);mum.|/mum.| age. | mum. mum. | age.
APSE eS A orn e = ae aie: |p iat | P SARE
1905. Days.|Days.|Days.| °F. | °F. | °F. | °F. oF. oF.
Sep tere = tase eee hee ees 2 56 43 | 49.5 | 62.6 | 62.3 62.4 | 1,083 | 840.5 | 961.8
1906. -
1:Nj Of Se eter Rien ge Sect ae pals Mgt. oT t 54 39-| 46.7 | 72.6 | 69.7 | 70.8 | 1,475.8 | 1,139.1 | 1,286.4
IMD ype a1 con se eae a ee 5 33 27 | 29 CO oso Lt aOuk ty lOse2 le cose 1,023
Jiinies a2 ee Pian yee 8 | 28'| 21 | 24.8 | 80.5 | 79.8 | 80.2 | 1,046.6 | 782.7 920.7
UU Visors ee, eee eee een Meee 7 265ey, 220 2ae4-NeeUron toe Sal SOs, 962.1 824.3 | 866.5
Aug ieee ee Mtn. A Nails aye 14 32 23 | 24.9 | 79.7 | 78.9 | 77.8 | 1,169.3 | 826.7 | 904. 4
1S (2) 0) pene Sere me ate ee ores ae. aa 13 154°; 23 46.2 | 79.4 | 54.6 | 70.7 | 1,907.9 837.6 1,127
Qt = etwte rey cee paee ned we 27) Soo eau ley 53.4 | 53 5322) | 1; 6006.4)" 351085) welaoaenn
| |

a Weighted.

It will be noted that the total effective temperature necessary to
cause eggs to hatch varies from 866.5° to 1,555.7°. It would be of the
greatest practical importance to formulate a rule which, on the basis of
effective temperatures, would show the time when eggs will hatch at
different seasons of the year. It is probable that the present data are
insufficient for this purpose, and special efforts will be made to add
to them. However, with the data at present available the following
tentative law may be proposed: ¢

When the average daily mean temperature ranges less than 53.2°, at
least 1,510.8 degrees of effective temperature must accumulate before
hatching will take place. When the mean daily temperature averages
from 61.4° to 77.8°, from 840.5 to 1,159.1 degrees of effectiwe temperature
will be required for hatching. When the mean daily temperature aver-
ages higher than 80°, between 782.7 and 824.3 degrees of effective tem-
perature must be accumulated before hatching will take place.

It is not our purpose to advise an attempt at the practical appli-
cation of this rule at the present time, but it is supposed that such
practical application can be ultimately made. To do so it would
only be necessary for the stockman to have a set of self-recording
maximum and minimum thermometers, such as can be purchased
for $3. Or the data might be obtained from the nearest Weather
Bureau station. By either of these means the average daily tempera-
ture could be easily obtained. By summing up the daily effective
temperatures—that is, the number of degrees above 43—the stock-
man, by reference to the minimum amount of accumulated effective
temperature necessary for hatching with various average daily tem-
peratures, could determine at least within certain limits what time

a This rule, of course, may not apply in rare cases in which eggs are deposited where
they will be subject to artificial heat rather than to weather temperature, as in manure
piles. It will apply, however, to the great majority of pasture conditions throughout
the infested area.

LIFE HISTORY OF GATTLE TICK. 21

it would be necessary to remove cattle from pastures in which tick
eggs might be to avoid danger of infestation by seed ticks. Possibly
a more feasible application would be the collection of the necessary
data from many localities by the State entomologists and the publi-
cation of predictions based upon them from time to time.

A seasonal arrangement of our data for 1905_and 1906 shows the
following:

Eggs deposited in June, July, August, and up to September 15
require from 824.3 to 840.5 degrees of accumulated effective tem-
perature for hatching.

Eggs deposited from September 16-30, in October, and in later fall
and winter months, require an accumulated effective temperature of
from 837.6 to 1,510.8 degrees.

Eges deposited in April and May require from 981.6 to 1,139.1
degrees.

Naturally an arrangement by months as above must be defective,
since no two seasons are exactly alike. The only accurate method
must be based upon a knowledge of the temperatures that are accu-
mulating in any particular season.

EFFECT: OF HEAT AND COLD ON EGGS.

In experiments to determine the effects of heat upon eggs a con-
tinuous temperature of 100° was maintained by means of an incu-
bator. The period of application of heat was 15 days. In one series
no moisture was provided, and in this case no hatching took place.
In another series in which abundant moisture was furnished, hatch-
ing took place and the incubation period was reduced to 15 days.
It will thus be seen that a moist atmosphere is essential to the hatch-
ing of eggs under a constant high temperature.

In experiments relating to the effects of low temperatures on eggs,
by means of a refrigerator a mean temperature of about 45° was
maintained, with a minimum of 32° and a maximum of 65°. The
eggs were kept in pill boxes with gauze tops to allow free circula-
tion, except during the period of refrigeration, when the ordinary
covered pill boxes were used. The eggs from about 20 engorged ticks,
collected on July 26, were placed in the refrigerator on August 4
and remained for 30 days. Hatching began on September 23, and
about 60 per cent of the eggs were viable.

In the case just referred to, the normal period of incubation was
increased, as the result of refrigeration, by 8 days. In a long series
of similar experiments, however, in which the period of refrigera-
tion ranged from 1 to 21 days, the period of incubation was not
appreciably lengthened and the normal percentage of hatching took
place.

22 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

In a series of experiments with alternate cold and normal tempera-
tures the eggs were kept in a refrigerator exposed to a mean tem-
perature of about 45° F. from 8.30 p. m. to 8.30 a. m. During the
day the eggs were removed from the refrigerator and remained at
the temperature of the air. _ After six consecutive nights of exposure
in this way from 5 to 10 per cent of the eggs hatched. It is probable
that the necessary manipulation in these .experiments interfered
with the viability of the eggs and that normally a considerably greater
percentage would hatch under the same conditions.

SUBMERGENCE OF EGGS IN WATER.

Sixteen different lots of eggs were used to determine the effect of
submergence in water. One-half were submerged for from 10 to 24
days, and most of the eggs hatched. In another lot submerged
for 25 days 33 per cent hatched. The experiments were _per-
formed from June to September and the incubation period under
water was not appreciably different from the normal at the different
periods at which submergence took place. In all these experiments
complete submergence was secured by means of a screen obstruc-
tion below the surface of the water. Our results agree with those
recently published by Messrs. Newell and Dougherty and with unpub-
lished data obtained by Prof. H. A. Morgan, who suggested our
experiments.

The practical importance of these experiments is to show that the
flooding of pastures would have no effect whatever on the viability
of tick eggs on the ground. Not only would the great majority
hatch, but the time of hatching would not be materially different
from that in case no water whatever were present. As a matter of
fact the flooding under some conditions, as, for instance, during a
drought, might hasten incubation.

These data, taken in connection with data mentioned elsewhere,
showing the remarkable resistance of seed ticks to water, indicate
clearly the reason why pastures overflowed for considerable periods
have repeatedly been found to furnish tick infestation.

A series of eggs varying from those recently deposited to others
about to hatch were submerged in tube-form vials. To keep the
eggs submerged absorbent cotton was pressed down into the water.
None of the eggs hatched, and we suppose this was due to insuffi-
cient aeration.

Several lots of eggs that were kept in pill boxes until they had
become thoroughly dried were placed on water in Stenter dishes to
determine if hatching would follow. After submergence for a short
period the eggs in large part filled out and appeared viable, but did
not hatch. |

LIFE HISTORY OF CATTLE TICK. 23
PERCENTAGE OF EGGS HATCHING.

To determine the exact percentage of eggs hatching is a rather
difficult matter for the reason that manipulation in counting inter-
feres greatly with their viability. In June, July, and August, 1906,
a number of observations were made showing the percentage of
viable eges in different lots from 30 to 90. In these cases the eggs
were counted, and it is likely that the percentage hatching under
natural conditions is considerably higher than that indicated. The
examination of masses of eggshells where hatching has taken place
normally reveals few unhatched eggs.

Messrs. Newell and Dougherty* have recorded a percentage of
hatching during the months of April, May, June, July, August, and
September of from 61.6 to 92. These experiments were performed
at Baton Rouge, La., under conditions which approached the natural
ones very closely, although the eggs were counted as in our experiments.

LARVAL OR SEED-TICK STAGE.

The larval tick isa minute 6-legged creature without distinct genital
opening, and with indistinct stigmata between the second and _ third
pairs of coxe. As Salmon and Stiles state, larval ticks frequently show
indications, at least, of stigmata between the first and second pairs
of coxe, and behind the posterior coxz in addition to those between
the second and third pairs. Only the pair first mentioned seem to
be functional. The color of the larva at first is whitish but soon
becomes dark brownish.

NONPARASITIC PERIOD.

‘For a few hours the larva remains about the shell from which it has
just emerged, but later makes its way upward on the first blade of
grass, stick, post, or other support that presents itself. Professor
Morgan informs us that he has seen seed ticks on the tips of sugar
cane about 8 feet from the ground. By placing a pole in the vicinity
of millions of seed ticks we have observed them to reach a height of
about 6 feet in a surprisingly short time; but the tendency is strongly
to remain not more than about 4 feet from the ground. In the absence
of some vertical object the seed ticks do not seem to scatter to any
great extent, but collect on the highest immediate point, even if it
is only a small clod or stone.

On whatever support the young ticks happen to be, they collect
in masses often nearly an inch in diameter. (See Pl. I, fig. 2.) Here
they remain for weeks or months awaiting a host. The front legs,
which combine the functions of antennz and legs in the insects

@La. Crop Pest Comm., circ. 10, p. 24.

24 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

proper, are waved through the air more or less constantly, and vio-
lently when a moving object approaches. The ticks attach them-
selves to any animate or inanimate object which touches them. Only
those that happen to attach themselves to cattle (and rarely a few
other animals) ever develop; the others either die or, dropping off,
become widely scattered.

Heavy rains wash the seed ticks to the ground, and it is possible
that violent winds may also serve to disseminate them. In our
experiments they have thus been spread to a distance of 5 or 6 feet.

It is noticeable that the seed ticks shun direct sunlight. We have
repeatedly seen bunches move halfway around the support with the
shade. In the morning they would be on the west and at night on
the east side.

EFFECT OF WATER ON SEED TICKS.

Interesting data having a bearing on the dissemination of the cat-
tle tick through the agency of water courses have been obtained. It
has been noted that heavy rains wash seed ticks from their supports
to the ground. In a considerable number of experiments seed ticks
were found to endure submergence varying in different lots from
10 days to 157 days. The latter record was obtained in an experi-
ment in which seed ticks were first placed in water in a Petri dish
and a few days later removed to a tube with earth on the bottom.
The details of these experiments are given in Table IV.

It is doubtless true that dissemination by water courses is not
quite as important as these results would indicate. Of course it is
possible that seed ticks may be carried many miles and deposited
on grass or bushes, from which they may reach cattle. The enor-
mous scattering of ticks so submerged in water, as would be the case
in floods, would undoubtedly greatly reduce the chances of infesta-
tion in pastures.

Taste 1V.—E fect of water on seed ticks.

f
; | Seed ticks | Longevity,
Eggs hatched. SuPer SUN Remarks.
Days. .

Septy2seeeeenss Sept: ye sete 47 | Eight inches of water in tub.

Sepusic0 025s" 157 | In Petri dish; Oct. 9 removed to tube with dirt on bot-

tom.

Octo4-11. 222.5: INOVigllie teen 14 | Alive on Nov. 25, and may have lived longer.
Sepbtlercsc. ce Septrilacet a 22 12 | In tub with sand and vegetable matter.

Ap yas oe ete 41 | In Petri dish on porch, with alge in water.
Aug. 16-17...... PATE Speers 32 | In Petri dish on laboratory desk.
Aug. 21-25.....- ANS AI ers =e 17 | Submerged as eggs.
Aug. 4-11....... Septs4: ssn 4c: 10-17 | Submerged as eggs. Culicid larva may have interfered.

Some years ago Professor Morgan obtained interesting results on
the effect of low temperature on seed ticks. In brief, he found a
temperature of 15° or 16° F. for a short period did not kill many seed

=,

LIFE HISTORY OF CATTLE TICK. 25

ticks, but that such a temperature continued for 24 hours resulted in
the death of practically all. Our experiments have been of two
classes—(1) in water and (2) without water. In both cases the seed
ticks were subjected to a temperature somewhat less than 32° F. In
the case where water was supplied solid ice was obtained. Both in
and out of water the seed ticks survived a freezing temperature of one
hour’s duration, although there was perceptible mortality among

them.
LONGEVITY OF SEED TICKS.

The time that seed ticks may survive without a host is a most
important matter in plans for control. Our experiments on this
point have been of two kinds—(1) with seed ticks from eggs deposited
by females placed on the ground in favorable circumstances, thus
giving absolutely natural conditions, and (2) with seed ticks in glass
tubes, where they could be observed more closely.

In the first series, which was instituted at the suggestion of Pro-
fessor Morgan, several hundred engorged ticks were placed on the
ground at regular intervals. The only inclosure was a cylinder of
2-inch-mesh wire screen about 4 feet in diameter to prevent disturb-
ance. The seed ticks from females placed under these conditions

.accumulated in enormous numbers on the blades of grass or stakes

provided for the purpose (see Pl. I, fig. 2). Table V gives the num-
ber of days the seed ticks survived in these experiments, together
with other data. It will be seen that the shortest period was 49
days and the longest 159+ days. However, the important period
is from the time of dropping of the adults to the death of the resulting
seed ticks, since the farmer must always suppose in rotating his cat-
tle that adults were dropped on the day of removal from the pasture
which it is desired to clean. This period ranged from 91 to 175+
days. It will be noted that there is considerable variation in the
period of survival, even at the same season of the year. This seems
to depend upon two factors, namely, the number of seed ticks in the
bunches and the amount of rainfall. The larger masses survive
longer, perhaps because the moisture is better retained, and heavy
rains scatter and reduce the masses. It is probable, on account of
the very large numbers of seed ticks in our experiments, that the
periods given in the tables are somewhat longer than normally occur
in the field. However, such excessive numbers do sometimes occur
in nature. For instance, when an animal dies of gross infestation
thousands of ticks deposit in a very restricted area. Mr. J. D.
Mitchell has seen cases of this kind in which the bunches of seed ticks
were fully as numerous as in our experiments.

‘

26 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Taste V.—Longevity of seed ticks of Margaropus annulatus Say, at Dallas, Tex., 1906.

Period from dropping Maximum
to completion of ovi- period Period
When dropped. position. When _ | Seed ticks | from drop-| of life of

hatched. | all dead. ping to seed

; ant is death ofall] ticks.
Minimum. | Maximum. SeadniGia

1 4

Days. Days. ‘ Days. Days.

VINE ZO Pea ae wich geen 12 185) Aue. 122 32! OCtSiecce 110 57

PUINION2S | = Reena ines meee tel oe 12 18°) Auge 15-7. 2) Oet. 215225 120 a67

14 Beta allt epee ae eee ie eee 14 PA) wba ase Oct. 205252 154 107

Mithloygelteye 3 ee oh ae rae eee 12 19 Avie e142 OCt. pose. = 86 62

DOME ete On Ge Se ace heck 8 16% Sept-wi-c--| Ochw2os-- 2 91 49
UNirec au hk see ee ener ORL An Mba 8 16 | Sept. 14... (b) 175+ 159+

v

aIn this experiment the seed ticks were accidentally disturbed. They would have lived some days
Py gome alive Feb. 20, 1007. |

In the second series of experiments to determine the longevity of
seed ticks the eggs were placed in glass tubes with open bottoms.
As will be seen from the footnotes to Table VI certain seed ticks in
tubes were shaded at all times, others in tubes were exposed con-
stantly to the sun, while the remainder were placed in pill boxes
protected from sun and rain. It is supposed that these diverse con-
ditions give an average length of survival that approaches closely to
that occurring under natural conditions. The detailed results follow:

Taste VI.—Longevity of seed ticks of Margaropus annulatus Say, Dallas, Tex., 1906.

| Period | Period |
Eggs deposited. Hatching. @ Seed ticks dead. Peas Renee Cones
: to death.| to death.
a Ses oe h

Days. Days. |

AME QBs ners ae Ne sce 84 138 (b)

Ao R LOUIS, © eens Tif 124 (b)

UN (Se en ak 83 | 130 (0)

AMIE HAS eee eee 85 | 124 (0)

Sen tc hi oy see peer 98 131 (>)

Chae ae ce 115 144 (b)

sna Coss et Masser 104 131 (0)

Lat pact Ma, TOR eee OBES ee Roa essa ly seec ast bl)

ATE OE eed Dee Bais oe 43 71 (b)

Vests) Oi Fast? Opes ert ce meager 83 110 | (b)

()

(b)

(0)

(¢)

(b)

(e)

(3)

(b)

(¢)

(b)

(d)

(¢)

(d)

aS eoo)s eee eee etna ()

a For details regarding oviposition of these ticks see Tables I and II.

b The seed ticks were in open-bottom test tubes in sand in the open air. They were protected from
the direct rays of the sun at all times by a cheese-cloth screen. Up to 11 o’clock a. m. they were also
shaded by the house.

c¢In open-bottom test tubes in soil exposed to sun at all times.

dIn open paper pill boxes outdoors, protected from sun and rain at all times.

Considerable numbers of seed ticks in all lots hatching after August
23, 1906, are still alive (February 20, 1907). .

LIFE HISTORY OF CATTLE TICK. 27

PARASITIC PERIOD.

The data already given regarding the periods of preoviposition,
incubation, and survival of seed ticks have an important bearing on
the time required to free pastures or other inclosures from ticks pro-
vided the cattle are removed. The data given under the present head-
ing, on the other hand, show the time required at different seasons
to free cattle of ticks by placing them in inclosures from which the ticks
have been eliminated either by systematic starvation or by the use of
naturally tick-free areas, as, for instance, fields that have been in culti-
vation for one crop season.

In this work we have utilized a grade Durham steer, 17 months
old at the beginning of the experiments. (See PI. II, fig. 1.) By
means of kerosene emulsion he was carefully cleaned of the thou-
sands of ticks infesting him when obtained. Thereafter he was thor-
oughly washed to remove traces of the insecticide and hundreds of
seed ticks were applied. Under proper precautions to avoid the steer’s
accidental infestation, these ticks were allowed to reach maturity.
After the ticks of each infestation became adult the steer was thoroughly -
cleaned and placed in another inclosure, which in each case had been
carefully disinfected by means of sprays. This process has now been
repeated until ten infestations have been reared covering the period
between August, 1905, and March, 1907. The details are given in
Table VII. In rotation systems the minimum developmental period
is the most important, because the cattle must be removed before
the earliest developed ticks have had offspring to reinfest them.
Therefore special reference is made in the table to the shortest periods
found, although the longest and the average are both given.

The following deductions may be made from this table:

1. The period from attachment to dropping ranges from 21 to 58
days. It should be noted that in the longest periods the limit was
reached by only one or two belated ticks, the majority approaching
the average.

2. The average period ranges from 26.5 to 43 days.

3. The average parasitic period is normally some days longer in
winter than in summer. But warm winter weather, as happened in
infestation No. 9, may reduce the period even below the average for
the summer.

4. The slowest developing ticks of one infestation may occupy from
10 days (in the summer) to 32 days (in the winter) longer than the
most rapidly developing ones. The rapidity of development of the
ticks of the same infestation depends somewhat upon their location.
Those on the portions of the body where the blood supply is most
abundant develop most quickly. In general it seems that heavy infes-
tations develop a little more quickly than light ones. This may be

-

28 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

due to the fact that, in light infestations with widely scattered ticks,
fertilization is less likely to take place than in other cases where the
males may find the females more readily. We have beer unable to
determine that unfertilized females occupy longer in development
than those that are fertilized, but our impression is that they do.

The table also shows that the principal variation in the time of
development of the ticks of the same infestation takes place not in
the larval or nymphal but in the adult stage.

Tasuie VIL.— Development of Margaropus annulatus on steer at Dallas, Tex.
i q I

| Adults dropped. ere iNtam-
\iNtinai | eee ee ea Re Mini- Wor

|

Period from

Ap- ntetead| | |
: Mini- | | | mum | attachment to
es When First | mum |} Second se ta pe- | Es dropping.
-*" | applied. | molt. {larval} molt. oh ena riod | =|
tion| “PI phal | First. Last. Gaute| ed or =

stage. | |
No. | 8 | stage. stage. 17P- Maxi- Mini-| Aver-

ped. mum.mum.) age.

|= |
| | \

1905. Days. Days. | \Days.| \Days. Days. Days.
1 | Aug. 16 | Aug. 28 12 | Sept. 2 | 5 | Sept. 15 | Sept. 274 WS] BON sD 30 36
2 | Sept.27 | Oct. 4 FEM (Oye Ay 8 | Oct.' 21 | Nov. 3} 9 10 42 24 33
3 | Nov. 11 | Nov. 22 11} Nov. 28 6] Dee: 8) Jan. 9)" 10 13 59 27 43

| | |

1906. | | |
4| Jan. 16] Jan. 25} 9) Feb. 3] 9 | Feb. 17 | Mar. 8 | 14 11 dl 32 41.5
5 | May 22 | May 31 9| June 6] 6 | June 14} June 30 | 8 95) < 39 23 31
Gol we dull 9) | eee ee ee dnalyaedG | eee July 24] Aug. 5 8 | Sal aaron 25 31
7} Aug. 2 |Aug.10-11 8-9 | Aug. 18 7-8 | Aug. 25) Sept. 11 7 14} 40 23 31.5
8 | Sept. 5] Sept. 12 7 | Sept. 22 10; Sept. 26 | Oct. 6 4} 236 31 21 26. 5
9 | Oct. 6} Oct. 14 | 8 | Oct. 21 7 | Oct: 305) Now: 12) 09) 200) |) 37) )|ee4 aso n
10 | Nov. 29 | Dec. 8 | 9 | Dec. 14 67); Dees 22) Jam.) 8 55 33 23 28

1907. | | |
ITE AE oe IEP AE Hoke ts) 7| Jan. 14 6| Jan. 24 | Feb. 3 | 10 35 33 23 27.5

|

a Removed.
DEVELOPMENT ON HOST.

When the larval ticks find themselves on the host they rapidly dis-
appear in the hair and attach themselves to the skin. They are
principally found on such parts as the legs, belly, and dewlap that
come in contact with the bunches on the grass, but may be found on
any part of the host. In cases of severe infestation they practically
cover the entire surface of the body, even the eyelids being infested.

In from 7 to 12 days the larval ticks molt and enter the nymphal
stage, in which they have eight instead of six legs. The nymphal
stage is further distinguished from the larval stage by the presence of
a pair of large stigmata quite in contrast to the rudimentary organs
of respiration of the larva.

The second molt (from the nymphal to the adult stage) occurs in
from five to ten days after the first. _The nymph can be distinguished
from the adult, which it resembles very closely, by the absence of any
genital opening. The process of both molts is undergone by the
females while the hypostome is firmly inserted in the skin of the host.
The shed skin splits open along either side and drops off in two scale-
like pieces. A portion of skin from the capitulum is also shed at the

LIFE HISTORY OF CATTLE TIOK. 29

‘same time. In the case of males, while during and after the first molt
the tick remains fixed to the host, after the second molt it detaches
itself and travels in search of a female. As a rule the males molt two
or three days ahead of females and frequently two are found attached
to the skin of the host directly beneath nymphal females, awaiting
the molting of the latter.

When the adult stage is reached the development is very rapid.
In from as few as 4 to 14 days the females become fully engorged
and fall to the ground to deposit their eggs. They die when the
operation is completed.

Although female ticks are somewhat more easily removed at the
time of molting than at other times, repeated careful observations
show that they do not actually detach themselves at either molt.
However, Mr. B. H. Ransom has shown that, when detached arti-
ficially just before or after the second molt, females will reattach if
placed upon another animal.“ Likewise he found that ticks removed
just after the first molt would reattach after 24 hours. Specimens
detached just before the second moit transformed and lived without
host for two weeks. Several experiments have led us to the con-
clusion that only in the rarest accidental cases can reattachment
normally take place. In many cases we have attempted to cause
detached ticks to reattach, but in only two cases did we succeed.
The following are some of the particulars:

February 23, 1906, eight ticks, ranging in size from nymphs to one-
half engorged adults, were placed on the shoulder of a steer. The
next day all but one of these ticks had disappeared. One, however,
which was about one-half engorged, had fastened to the skin about
6 inches from the point where it was placed. This tick was found
detached on March 1 and trying to crawl out from the hairs which
had been glued together to hold it in place. At this time it was not
fully engorged. It began depositing eggs on March 18, and con-
tinued oviposition until April 26, reaching a total of 523 eggs. An
attempt was made to cause these eggs to hatch, but without success.
It is not likely, however, that the failure to hatch was due to the
experience of the tick. It is probable that the state of engorgement
may have had something to do with the matter, and, moreover, at
the time of the year when the eggs were under observation it is
exceedingly difficult to cause them to hatch. After the experiment
that has been mentioned repeated attempts were made to obtain
other cases of reattachment. Ticks at various stages were placed on

a ahille (Contr. l Etude Ixodidés Argentine, p. 112) had previously detailed a num-
ber of experiments in the reattachment of Margaropus (Boophilus) microplus. Tt was
found that immediately after molting the ticks would more or less frequently reattach
after being removed artificially. Nothing, however, was found to indicate that the
parasites naturally detach and reattach on the host,

30 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

the steer and confined to limited area by means of vaccination
shields. In no case under this manipulation did reattachment take
place. Recently, however, a tick removed just before the second
molt became adult in a pill box and reattached after 25 hours.

ADULT STAGE.

Adult females are the ticks that are generally seen, and their
appearance is familiar to most persons. The males (which do not
become engorged) are generally overlooked, although they may be
easily found’ attached to the skin of the host directly beneath the
females. This gives rise to a rather prevalent popular idea that
females carry young with them.

The following descriptions are taken from the work of Salmon and
Stiles: 7 ;

Male.—Body oval, narrowed on front, broadest (1.3 mm.) at stigmal plane, 2.15 to
2.35 mm. long. Scutum reddish brown, covering entire dorsal surface, prolonged in
front by two pairs of projections—one pair of more prominent dorso-lateral projections,
dorsal of anterior projection of coxee I, and one pair somewhat less prominent and
more median, ventro-median of first pair and nearer the neck. Two cervical furrows
shallow, extending more or less distinctly to the posterior border; may be somewhat
interrupted in the middle; a median furrow present in posterior half, may be very
indistinct; posterior margin of body divided into festoons, which may be only slightly.
marked. Relatively large circular pores, with extruding short bristly hairs, scattered
over entire surface. Eyes small and pale, often problematic, at I intercoxal space.
Ventral surface lighter than dorsal, all portions provided with short stout hairs; geni-
tal pore, broad, transverse, between coxze II; anus slightly posterior of stigmal plane;
two pairs of anal plates (clypei): one pair elongate, rectangular to triangular, close to
anus, in some cases extending cephalad to middle of coxee IV, and caudad to near or
beyond posterior margin, the anus being about at the middle of the length, in other
cases extending from height of middle of stigmal area to beyond posterior margin of
body; the median border longer than lateral border, the former prolonged into a point
posteriorly, the postero-lateral margin may be nearly straight, or somewhat curved, or
irregular in outline, thus presenting broad tooth-like projections; lateral and contigu-
ous to each of these shields is found another shield somewhat similar in form, but
smaller in size. Median caudal appendage absent. Capitulum 450 to 500 long, base ~
similar to that of the female, but a little straighter, longer, more salient in front of
dorsal shield, into which it penetrates by a sort of rectangular neck, lateral projec-
tions not very prominent. Mandibles 600 long, digit about 90”; internal apophysis
with straight base and broad bifide point; external apophysis bidentate, the terminal
subventral tooth may be very small while the proximal tooth is strong and large, or
both may be large. Hypostome similar to that of female, four distinct rows of teeth on
each half. ‘Palpi about 190 long, similar to those of female. Legs strong; coxe large,
those on each side contiguous, as broad as long; coxze I triangular, apex may be pro-
longed anteriorly beyond the corresponding anterior point of dorsal shield, reaching
anterior angle of base of capitulum, or may be very short, base posterior and more or
less distinctly bidentate, the teeth short, often slightly pronounced, or quite promi-
nent, the lateral tooth in some cases prolonged into a well-marked spine. Tarsi like
those of female.

a‘ The cattle ticks (Ixodoidea) of the United States: 17th Ann. Rept. Bur. Animal
Industry, U. 8. Dept. Agric., pp. 420-424, 1901.

oxox LIFE HISTORY OF CATTLE TICK. 31

Female.—Body elliptical, as broad in front as in back, usually somewhat con-
stricted in middle, near IV pair of legs; may attain 13 mm. long, 7.5mm. broad. Color
exceedingly variable; live specimens vary from a tawny yellow (younger forms) to an
olive green (very old specimens), alcohol specimens from yellow to red or black; the
excretory system often shows through the cuticle as tortuous whitish canals. Dorsal
shield (scutum) very small, visible as a dark brownish spot in a depression at anterior
end of median line; usually about 1.1 mm. long by 0.8 to 0.9 mm. broad, decidedly
emarginate anteriorly to receive capitulum; lateral borders nearly straight and par-
allel in anterior portion, from antero-lateral points to eyes, then convergent from
eyes caudad, forming a more or less bluntly rounded posterior angle in median line;
cervical grooves divide the anterior half of scutum into three more or less equal lon-
gitudinal fields, and diverge posteriorly; surface of scutum provided with short bris-
tles which are more numerous near anterior border and near the eyes than elsewhere.
Eyes rather small near anterior third of margin of scutum. On nearly the entire
length of dorsal surface of body are two antero-posterior grooves, interrupted or nearly
effaced near plane of IV pair of legs, and ending a short distance from the shield and
from posterior margins of body; also, an unpaired median groove in posterior half of
body; all three vary with the muscular contractions and may more or less com-
pletely disappear when body is replete. Ventral surface shows four pairs of more
or less distinct marginal constrictions corresponding to the four pairs of legs, the IV
constriction being most marked, antero-median region also depressed at insertion of
capitulum; vulva small, median, at plane of cox I; sexual grooves corresponding
to the paired dorsal grooves; but showing some variation in different specimens;
median groove extending from anus to posterior margin. Anus about on border of
second and last thirds of body. Stigmata short oval; stigmal pore slightly crescentic,
convexity lateral;, stigmal field with numerous larger and smaller wart-like strictures,
forming a zone near the margin. Cuticle of entire body finely wrinkled, bearing short
hairs. Capitulum very short, about 800s from posterior dorsal margin to anterior
end of hypostome; base of capituluni hexagonal, enlarged on its dorsal surface; in-
serted in emargination of scutum; lateral projections not very prominent. Mandibles
860 long, digit 120”. Internal apophysis conical (Neumann), bidentate (Fuller),
with its base near the terminal extremity; external apophysis with three successive
teeth, one terminal, subventral, small; the second stronger; third large. Hypostome
rather spatulate, broad, a little longer-than the palpi, provided on each half with
four rows of nine to ten nearly regular denticles, which do not extend to the base.
Palpi very short (3102), subconical, articles at least as broad as long; first article par-
tially hidden under the antero-dorsal border of the base of the capitulum; second
article pedunculate, dilated in a salient crest in its middle portion, thus forming a
prominence inward (toward median line) and outward, and provided with strong
hairs, especially on the inner prominence; third article smaller, subtriangular on its
dorsal surface, where it forms a projection in and out; fourth article small, cylindrico-

_ conical, infero-terminal. Legsrather thin, short (pair I, 2 mm.; pair II, 2.6 mm), yel-

lowish brown, first articles darker than the others. Coxee: pair I subtriangular, pos-
terior border bidentate or biundulate, the division in many cases indistinct. Tarsi
I unicalearate, 11 to IV bicalearate. Pulvillum about half as long as claws. Stiff
bristle-like hairs on all articles.

EFFECT OF CONTINUOUS COLD AND HEAT ON ENGORGED
FEMALES.

Twenty-five ticks were used in the experiments with cold, which
were conducted during the month of August, 1906. A mean tem-
perature of about 48° F. was maintained, with extremes ranging

ay NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

from 34° to 53°. With exposure up to 300 hours practically all ticks
recovered and in most cases deposited viable eggs. In cases of more
than 300 hours’ exposure practically all ticks survived, but none
deposited viabie eggs, although in many instances oviposition took
place.

In a number of experiments with heat a mean temperature of from
98° to 102° was maintained. Up to 103 hours of exposure to this
temperature practically all ticks deposited eggs that were viable.
With exposure at the same mean temperature of from 144 to 218
hours’ duration, eggs were deposited, but were found not to be viable.
They were dry and shriveled when deposited.

Some of the females survived heating for the longest period, namely,
218.5 hours. With an exposure of 103.5 hours or more, however, at
least one-half succumbed.

EFFECT OF DIRECT SUNLIGHT ON ADULTS.

~

Eleven unengorged females placed in a box exposed to the direct
rays of the sun in September died in three days. Seven unmatured
females in direct sunlight from morning until noon seemed dead at,
noon. They did not survive until the next day, although they were
removed from the direct sunlight at 2 o’clock. Similar experiments
showed that death resulted in the case of engorged females after a few
hours’ exposure to the sun. In experiments with eggs, tubes were
subjected to direct sunlight for one day. When moistened while
kept in these tubes, hatching seems to take place normally, and
hatching followed in similar experiments in which the eggs were
kept dry.

EFFECT OF SUBMERGENCE IN WATER ON ENGORGED ADULT
TICKS.

Adult ticks have remarkable resistance to the effect of submergence,
as has been pointed out to be the case with eggs and seed ticks. The
immediate effect of submergence is to cause a cessation in the activ-
ity of the ticks, while they become somewhat distended apparently
from the absorption of water. In August and September, 1905, a
considerable number of experiments were conducted in which the
adult ticks were submerged in water from the city mains at Dallas,
Tex. Judging by the experiments with seed ticks and eggs mentioned
elsewhere it is not likely that water impregnated with foreign matter
would have changed the results. During the months mentioned a
period of submergence of 24 hours did not result in the death of any
appreciable number of ticks used\in repeated experiments. After
one or two hours the specimens recovered from the immediate effect
of submergence and proceeded to deposit eggs which were found to be.

LIFE HISTORY OF CATTLE TICK. 83

viable. With between 24 and 48 hours’ submergence the number of
ticks that recovered diminished rapidly. Occasionally specimens
recovered after a period of submergence of over 48 hours. For
instance, the specimen collected on July 12 and submerged for 50
hours survived and deposited viable eggs. Later in the season (that
is,.in October) somewhat, different results were obtained in experi-
ments in the submergence of adults. In this month many ticks
recovered after from 50 to 90 hours of submergence. In fact, fully 50
per cent of the ticks submerged between these extremes regained
their full activity. In one experiment two out of five ticks sub-
merged in October for 91} hours recovered and deposited viable eggs.
Nevertheless a number of ticks submerged for 115} hours did not
recover.

The results that have been mentioned above indicate that where
engorged ticks fall from cattle that are standing in permanent pools
of water none will survive to deposit eggs. At the same time the
results show that temporary flooding of from 24 to 100 hours’ duration
would not in all cases prevent ticks from depositing eggs. It must
be noted in this connection, however, that the vitality of eggs depos-
ited by ticks just prior to temporary flooding would not be interfered
with by the water, although, of course, they might be washed away.

DROPPING FROM HOST.

It is a more or less prevalent popular idea that ticks have some
sense which enables them to drop from a host in places favorable
for oviposition. A few observations have shown, as was supposed in
the beginning would be the case, that there is probably no such
power of perception present in the cattle tick. The popular impres- —
sion probably had its origin in the fact that bunches of seed ticks are
found most numerously in the places where the cattle collect for the
benefit of shade or water. This phenomenon obviously is due merely
to the collecting and standing of the cattle.

Many observations seem to show clearly that there is no preference
as to the time of dropping. In our experiments many ticks have
been known to drop from host during the night and many others to
~ drop during the day.

LOCOMOTION.

Some attention has been paid to obtaining data on the distance
engorged ticks may travel after dropping from the host, since this
has a practical bearing on double fencing in eradication work. In
some experiments specimens were placed on the floor in the laboratory
and allowed to move at will. The total movement varied from 24
inches to 123 inches, the latter distance being covered in 52 minutes.

5795—No. 72—07

3

34 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

The effect of antiheliotropism was always in evidence in these exper-
iments. The light was admitted from different quarters at different
times and the ticks always changed their course so as to travel away
from it. These results are very similar to those published by Lahille
from experiments with Margaropus (Boophilus) microplus in Argen-
tina. The distance engorged females will travel depends upon how
soon they can reach shelter. If they obtain protection under débris
of any kind they seem to be disinclined to go farther. The greatest
distance traveled on the ground out of doors was only 24 inches.

It is popularly supposed that engorged female ticks often burrow
into the ground. In a number of experiments with light barnyard
trash in tubes it was found that females will work their way down
from one-half to 1? inches. In such cases the masses of eggs assume
various shapes on account of the surrounding débris. (See PI. I,
fig. 1.) It was found that seed ticks from eggs so deposited had no
difficulty in reaching the surface.

HOST RELATIONS OF THE CATTLE TICK.

Margaropus annulatus was described in 1829 by Say “‘from speci-
mens taken on a Virginia deer in Florida.’’ Since that time we
have been unable to find records of the occurrence of the species on
deer. However, a number of instances have come to light recently
in which undoubted specimens of Margaropus annulatus have been
found on deer. The first of these cases was the result of an examina-
tion made by Mr. R. C. Howell on a herd of tame deer in a park at
Mount Pleasant, Tex., in October, 1905. Since that time Mr. J. D.
Mitchell has found specimens on a deer at Oakville, Tex. Mr. T. R.
Coker has sent specimens from the same locality and host. In both
of these instances the ticks were found on wild deer that had just
been shot. In February of the present year Mr. F. C. Pratt collected
a few specimens of Margaropus annulatus from a dry deer hide at
Kerrville; Tex., and in December he examined a fresh hide on which
considerable numbers were found.

The matter of the possible development of Margaropus annulatus
on various animals, among them guinea pigs, rabbits, dogs, and cats,
has been studied by various investigators. Dr. J. W. Connaway’s
experiments in Missouri in 1897 showed that the ticks would not
attach to rabbits or guinea pigs. They did attach to dogs in consid-
erable numbers, but only one of them ever matured. Recently
Mr. B. H. Ransom has repeated the experiments with rabbits, dogs,
and cats. On rabbits and dogs the ticks attached, but remained so
only a short time. On a cat a female tick remained attached from
July 25 to August 30 and molted on the host. Nevertheless, it did
not reach engorgement.

HOST RELATIONS OF CATTLE TICK. BD

In our experiments with ticks we have kept several dogs primarily
for experiments with species other than Margaropus annulatus. At
the same time we have repeatedly sprinkled thousands of seed ticks
of Margaropus annulatus on these dogs, but in no case have we noticed
that attachment took place. This, with the work of Connaway,
Schroeder, and Ransom, seems to indicate that dogs can play but a
very unimportant part as hosts for the cattle tick. Ransom men-
tions the fact that the collection of the Bureau of Animal Industry
contains specimens of Margaropus annulatus collected from a dog,
from which host Francis (1894) seems to have been the first to report it.
- One of the writers has had one case of an attachment of Margaropus
annulatus to his person. This was a male specimen that attached
between the fingers of the hand. It was removed after about half an
hour. Mr. Ransom mentions a similar case in which, however, the
specimen was a female and remained attached to his hand for twenty-
four hours before it was removed. Attachment to human beings must
be very rare. The junior author had worked with thousands of
specimens of ticks before and after the single case of attachment that
has been mentioned.

In addition to the abnormal and unusual host relations mentioned
above, there are not infrequent instances in which Margaropus annu-
latus has been found to occur on horses, mules, and asses.

All this work shows clearly the remarkable host restriction of the
cattle tick that is most important from the viewpoint of its attempted
eradication.

The early records of Packard, showing the occurrence of the cattle
tick on a porcupine and a similar record of its occurrence on the
rabbit in Idaho, must have been due to a misidentification of the
species.

The strict instinct for the proper host in the cattle tick is shown in
the extreme infrequency of attachments of ticks to each other.
Thousands of ticks have been sent to the laboratory alive inclosed
together in tin or wooden boxes. In only one case was it found that
a tick had inserted its rostrum in another. This happened in a lot
collected in southwest Texas by Mr. J. D. Mitchell on April 19. A
female had inserted its beak firmly on the lateral dorsal portion
behind the posterior coxe. The two specimens were placed in
alcohol and still remained connected.“

RELATION BETWEEN RATION AND TICK INFESTATION.

For several months after experiments were started at Dallas in
placing seed ticks on the steer used for experimental purposes it was
found that a surprisingly small proportion ever became adult. In

@Since the above was written important discoveries regarding the occurrence of
Margaropus annulatus on sheep have been made bythe Bureau of Entomology. See
Circular 91, Bureau of Entomology, U.S. Department of Agriculture, issued July 3)1907.

36 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

many cases from 5,000 to 10,000 seed ticks were placed on the animal,
but only from 1 to 20 adults ever developed. Prof. H. A. Morgan
made the suggestion, based upon the observations made by him in
Louisiana, that the ration the steer was receiving was responsible for
this remarkably small proportion developing. At his suggestion we
changed the ration. The steer had been receiving daily 4.76 pounds
of corn chops and 5 pounds of prairie hay. The inspection tag on the
chops guaranteed not less than 9 per cent protein crude and not less
than 4 per cent fat crude. At Professor Morgan’s suggestion the
corn chops were eliminated. Immediately a much larger percentage
of seed ticks developed to adults on the animal, although his general
condition did not seem to have been changed materially. While
before a dozen adults from many thousand seed ticks was the maxi-
mum, after the change in the ration hundreds developed from no
larger numbers of seed ticks applied.

ENEMIES OF TICKS.

At one time it was supposed that sowbugs may sometimes be
important factors in the destruction of tick eggs. A number of
observations have shown that the greatly preferred food of these
isopods is vegetation either live or decayed. In laboratory experi-
ments Armadillidium vulgare was found to feed on dead ticks and
also to devour the eggs whenever no other food was provided. Thirty-
eight sowbugs, furnished with 897 tick eggs, consumed 366 at the rate
of 3 eggs per day each. In another case two sowbugs devoured 159
tick eggs at the rate of 15 each per day. These results hardly seem to
substantiate the impression that sowbugs may be of considerable
economic importance. It should be emphasized that the experiments
referred to were conducted in the laboratory, and the sowbugs were
deprived of other food. Under natural-conditions the results might
have been quite different.

The little “‘fire-ant”’ (Solenopsis geminata Fab.), which has recently
been found to be acquiring a special taste for the boll weevil, undoubt-
edly destroys many engorged ticks that have dropped to the ground.
Experiments performed by placing ticks in the immediate vicinity of
nests of this ant show that under such circumstances they must
invariably be killed. The nests of this ant are found throughout the
pastures in the South, and the total of the work done by them must
be considerable.

A number of dipterous larve have been found feeding upon tick
eggs and an undetermined species of Phoridz has been bred.

At one time we were inclined to believe that a chalcidoid parasite
of the cattle tick had been reared. Early in 1906 such a parasite
was found in a pill box with the remains of an engorged tick placed
there the fall before. Upon sending the specimen to Dr. L. O. How-

ENEMIES OF TICKS. ot

ard it was learned that it belongs to a new species and genus. Chalci-
doids of the subfamily to which it undoubtedly belongs are known to
be parasitic on various dipterous larve. Upon reexamination of the
remains of the tick a portion of a dipterous cocoon was found. Con-
sequently the hymenopteron was probably not a parasite of the tick,
although the interest remains, since the dipteron was probably para-
sitic on the tick.

A number of observations have been made showing that domestic
fowls frequently learn to remove ticks from cattle in barn lots. Mr.
F. C. Pratt observed a case in which the fowls regularly visited hides
hanging up to dry for the purpose of picking up the ticks which
dropped from them. Mr. J. D. Mitchell has witnessed ‘“ jackdaws”’
(Quiscalus major macrourus) picking ticks from cattle near Victoria,
Tex., and the farmer informed him that he believed these birds kept
the cattle practically free of engorged or nearly engorged specimens.
Mr. S. E. McClendon, of the Louisiana Experiment Station, informs
us that he has repeatedly seen kingbirds (Tyrannus tyrannus) engaged
in the same work.

In connection with the enemies of ticks it may be stated that it
seems likely that mice are of some importance. In the laboratory
it was found that the best bait for mouse traps that could be used was
engorged cattle ticks. It seems likely that in the pastures field mice
may frequently devour ticks. Dutton and Todd (Human Tick fever,
p. 17) write as follows: ‘‘Ticks are not without natural enemies.
Rats eat adults with avidity, and ants carry off young ones and
eges. We have lost ticks in both ways. One occasion over two
hundred young ticks were carried off in a single night by small
ants.’ These remarks apply to Ornithodoros moubata in West Africa.

THE PRACTICAL APPLICATION OF THE INFORMATION RECORDED
IN THIS BULLETIN.

In the preceding pages at different places the special practical
importance of the data discussed has been mentioned. As a matter
of fact the work upon which this bulletin is based has been planned
to accumulate additional information for use in the practical work of
tick eradication. Some methods of control are satisfactory in certain
districts, but much less so in others. Plans that would be feasible
along the northern border, for instance, where the tick is on a rather
delicate equilibrium and is never found on the cattle for months during
the winter, would not be applicable to the moist regions along the
Gulf where the cattle are infested throughout the year. Of the
various methods of eradication undoubtedly those of the widest
utility are the ones which prevent the development of the ticks by
breaking up the relation between them and the cattle. Of these, the
more important are the feed-lot or soiling system for relieving the

38 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

cattle of ticks and the pasture-rotation system (to be used in con-
junction with the former) for freeing the pastures.

In the feed-lot or soiling system the basis is the time occupied for
development on the host in connection with the time from the drop-
ping of engorged females to the hatching of seed ticks from eggs
deposited by them. In a tick-free inclosure, as a feed lot, cattle
may be left to drop their ticks until it is time for the eggs from the
first-dropped individuals to hatch. Information as to exactly when
it will be necessary to move the cattle to avoid reinfestation is given
in Tables I and II, which show the numbers of days at different seasons
before ticks begin to oviposit after dropping and the time before the
first eggs deposited will hatch. In July, 1906, for instance, this was
from 25 to 27 days. The time required for all ticks to drop. from
cattle, which indicates how long they must remain in one or more
tick-free areas, is shown in Table VII. The period is from 31 to 59
days.

The data necessary for an intelligent plan of freeing pastures, by
removing the cattle until the death of the ticks, are given in Tables
V and VI, showing the period from dropping to the death of all the
resulting seed ticks. The former table shows that if cattle were
removed from a. pasture on June 20, 110 days later it would be per-
fectly safe to consider it tick-free.

The data referred to above, together with the results of other
experiments, have been arranged to cover maximum preoviposition
and maximum oviposition periods, for convenience, in Table VIII.

Taste VIII.—Periods in the life history of the cattle tick wpon which means of control

are based.

| eee from POPP IRE Minimum | Maximum

0 oviposition, aoe period from period from
Minimum 4. : Adan ha alae p

When ticks dropped. incubation dr GEping ot BR droppers

Minimum. Maximum. period. hatching death of

of eggs. seed ticks.

1905. Days. Days. ‘Days. Days. Days.
[STE OU Sn ce eet feline eet 2 3 5 - 43 46 | June 22 293
1D) OL aE a Ss. Boats a obese 3 5 56 59 | May 31 271
WECH2B» Roe Pores fee naan 21 41 54 75 | Aug. 28 248
Meck 24d. ois f so keeles 21 41 47 68 | Aug. 15 234
WECH25 Aa oe Oo ee ae 21 41 47 68 | Aug. 22 240
1906.

MEDD celta ate eee ola tain een 9 10 39 | 48 | Aug. 28 161
YNioy oy Cae oe Ge Mae oe ohne 1 Bee 5 6 33 38 | Sept. 20 154
ATOR ollie an ee Seeds 5 Aas 5 6 29 34 | Oct. 5 167
Maye dir cate Pace Se Bee he e 2k 6 | 10) 27 | 33 |...-do 157
IM Siva thre ove rete ak eee oe 6 10 | 28 34 | Aug. 9 97
Miivel O)5e = 952. SOAs wee 22 nee 4 6 | 27 31 | Sept. 26 130
I iii Oleg Se eee Be aN Pe 4 | 6 | 27 31 | Sept. 22 124
LITO YS eee oe ae oe eed Se 3 | Sy 28 31 | Aug. 15 73
MUNG 3 oc E KEE econ e aetees 3 | 5 | 23 26 | Aug. 26 a 83
A AIMGY Ge eat Se Sear nee Ace see 3 Sa 23 | 26 | Aug. 6 a 61
DO eee ae eee 3 | 5 | 23 26 | Oct. 6 122
Af ubach Pere, ae eas See eam dl 3 4 | 21 24 | Sept. 26 106
JUNC AS aaa Wrewace ease ezee at 3 4 | 22 25) |Eae doe 105
RUT yaa) rhe A eR a eae 3 a 22 25 | Oct. 20 ail
ULL ail eee eee oe oes Oe soe 3 4) 23 26 | Feb. 18 a 220
DO} ees a Se Be Re Woe eS ee eee 3 4 24 27 | Jan. 23 194

a These ticks were kept in open-bottom test tubes exposed to sun at all times.

APPLICATION OF INFORMATION. 39

In the above experiments (Table VIII) the eggs and seed ticks,
except those marked (7), were kept in cylindrical tubes of }-inch
diameter inserted for about 14 inches into sand or soil in receptacles
on or below the surface. These were kept in a cage on the west side
of the laboratory so that they were protected from the sun until 11
a.m. Assoon as the eggs commenced to hatch, there was placed in
the tube a stopper of absorbent cotton which remained unremoved
until the death of the ticks.. In cases marked (“) the conditions were
the same, except that the tubes were exposed to the sun at all times.

The conditions furnished included a humid atmosphere and a min-
imum temperature as compared with the normal. The enforced»
bunching and protection from wind and rain furnished additional
favorable conditions. It would seem that the periods are maxima
and hardly to be met with in the work of extermination. One unfa-
vorable factor met with, which could not be prevented, was the
growth of algz on the inside of some of the tubes. This accounts for
some of the variation in the longevity.

In addition to the main practical data given the following are of
importance:

All experiments and observations show the close host restriction
of the cattle tick. Though the occurrence of the cattle tick on deer,
horses, mules, asses, and some other animals is comparatively rare,
it must be taken into consideration in the practical work of tick eradi-
. cation.

Eges are but little affected by water. When submerged, they
hatch in about the normal time. Seed ticks are also resistant to water.
In one case seed ticks survived submergence of 157 days. Adults
are less resistant, but in the summer they frequently survived sub-
mergence of 48 hours. Later in the season the resistance seemed to
be greater, some females surviving after more than 90 hours. It is
evident that water courses must play a very important part in tick
dissemination. .

The adult cattle tick has only the most limited means of locomotion.
After it drops from the host it crawls but a few feet at the most.
On the ground in our experiments the greatest distance traversed
was only 24 inches.

On the whole, no very important enemies of ticks have been found.
Domestic fowls frequently devour considerable numbers of them, and
some wild birds also render valuable assistance in picking them off
animals or from the ground.

NOTES ON VARIOUS SPECIES OF TICKS FOUND IN THE
UNITED STATES.

r

The role that Margaropus annulatus Say was found by Smith and
Kilborne to play in the transmission of splenetic or Texas fever in
cattle impressed upon investigators the importance of ticks as carriers
of disease. Since that time the study of these creatures has progressed
rapidly. Smith and Kilborne, Lounsbury, Theiler, Marchaux, Salim-
beni, Dutton and Todd, Motas, Kossel, Ricketts, eal King are among
oe who have sain aoe that ticks are the agents through Deh
various diseases of man and other animals are transmitted. What
is greatly needed in this country at the present time is a convenient
means of identifying the various species. This the writers have
attempted, in a measure, to supply in the following pages, in which
will also be found notes on the life history and habits of such species
as they have encountered.

CLASSIFICATION AND HABITS.OF TICKS.

The following key will enable one to determine the genera of the
various ticks found in this country:

KEY TO FAMILIES, SUBFAMILIES, AND NORTH AMERICAN
GENERA OF TICKS, (IXODOIDEA).4

Seupuin ‘Absentee ss ote Oye aha, de Ee res ie Mk ee a Family ARGASID#.
SS CUCU OLESEN Fess tte artis ne eee op ee ere ae ees we Family [xoprpz.

Family ARGASID.-

Capitulum at least its length from the anterior margin.........-. Genus Argas (p. 42).
Capitulum under a beak-like projection, close to anterior margin.
Genus Ornithodoros (p. 45).
Family TIXODID.

1. Palpi short, not or only slightly longer than broad; capitulum short.
Subfamily RurpicePpHALIN#®, 2
Palpi plainly longer than broad; capitulum longer ......-- Subfamily Ixopin~#, 5

Subfamily RurprcePHALIn ©.

2. Dorsal surface of capitulum hexagonal, the sides projecting in angles; male with
analiplates. <7. Y72223_ 2 2 ee ea oe 0 le ae 3

a This ae is based upon those of Salon onal Stiles (1901) ea ‘Banks (1904). “The
genus Ceratixodes is not included (see p. 54).

40

CLASSIFICATION AND HABITS OF TICKS. 41

Dorsal surface of capitulum rectangular, sides straight; male without anal plates. .4
. Second and third palpal segments extend laterally into sharp points; stigmata
fay eenyal ses SC Ra eA ae Riel i ene a aero Genus Margaropus (p. 49).
Second and third palpal segments even; stigmata comma-shaped.
Genus Rhipicephalus (p. 47).
4. Eyes present; external border of palpi straight; coxee I bidentate.

(oN)

Genus Dermacentor (p. 49).
Eyes absent; external border of palpi uneven; coxe I not bidentate.
Genus Hemaphysalis (p. 52).
Subfamily Ixopin#.

5. Anal groove surrounds anus anteriorly and opens posteriorly; eyes absent; stigmal

plate Mean yeCwreWlars ko. aaa ae hein ae a. 2 en = = Genus Ixodes (p. 54).
Anal groove surrounds anus posteriorly and opens anteriorly; eyes present; stigmal
FULD MSOs ey. LORI a tpt os ier ae ee cera ee Genus Amblyomma (p. 58).

Lahille has recently ® published an ingenious graphic table for the
separation of the families and genera of Ixodoidea. It is reproduced

SuB-ORDEN: ARPAGOSTOMA
SaaS Apoes

Argaside ant a aan
Heo PRIOR sr ARs Hee maphysalis”

SGERe ee ;
~Neumannielia

Amblyomma *

FAMILIAS

Anopli oe al

Cat CaP lois | Oe Oe Dermacentor *
| Hyalomma

pe Ae Mia © “s @e Rhipicephalus”
ae ; 26 @---Boophilus’

eis DN AOE S
Per ssoph —
Se \Anommata, PA) ee E schatocephalus
‘Ge a

_ Ceratixodes’

fig. 2.—Graphie table for the separation of the families and genera of ticks. (From Lahille.) The
underscored genera are represented in the United States.

lxodide

here as figure 2. The genera underscored are known to be repre-
sented in the United States. The suborder Arpagostoma of Lahille
is the same as the superfamily Ixodoidea of Banks, used by us.

Family ARGASIDZ. '

The family Argaside is represented by two genera, Argas and
Ornithodoros. They are readily distinguished by the characters
given in the table. ate species are peuoalae on imanabaals and birds.

Gye iene Argent., 1905, p. a1,

42 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Genus ARGAS.

The species belonging to the genus*Argas are nocturnal parasites
of chickens, pigeons, and other birds, and occasionally attack mam-
mals, man included. Two species are represented in this country—-A.
miniatus Koch and A. sanchezi Dugés. The former has been found
by Marchaux and Salimbeni to transmit spirillosis of fowls in Brazil.
There is some evidence of the occurrence of this disease in the
United States.

ADOBE TICK.

(Argas sanchezi Dugeés. )

This species was described in 1891 by Dugés from larve collected
at Guanajuato, Mexico, on a dove (Zenaidura macroura). In struc-
ture it is very similar to A. mimatus. Neumann’s description is
based upon a number of specimens collected at Mariposa, Cal., upon
a quail, and at Santa Agueda, Lower California, upon a wild turtle
dove. Mr. Nathan Banks has specimens from Deming, N. Mex., and
from Arizona, and it is quite probable that it will be found in Texas,
and possibly farther north. Mr. Banks states that in New Mexico it
is found in houses and is there known by the common name used
above.

FOWL TICK.

(Argas miniatus Koch.) @

This species was described in 1844 by Koch from Demerara. The
name Argas americanus, applied by Packard in 1873 and used largely
by writers in this country, is a synonym of A. miniatus. For excel-
lent illustrations of this species see Salmon and Stiles, [xodoidea of
the United States, Plate LX XIT and text figures.

The species is well distributed over the world. The records include
Persia, Algeria, Russia, India, Australia, South Africa, Central and
South America, and, in the United States, Florida, Texas, New
Mexico, Arizona, and California. In southwestern Texas it is found
in large numbers in and about chicken houses and out-door roosts,
hiding away in crevices by day and coming out at night to engorge
on the fowls. This species seems occasionally to attack mammals,
as Mr. J. D. Mitchell has taken it once in Texas from the rabbit.

As pointed out by Lounsbury, the sexes are so much alike that,
except by the size, the only safe way of separating them is through
the form of the genital orifice. To quote from him, ‘‘This orifice
[genital] is situated just behind the mouth parts on the under side of
the front of the body; that of the male is relatively inconspicuous

aThe name of this tick is in great confusion. The one used by the writers is that
recently adopted by Banks.

CLASSIFICATION AND HABITS OF TICKS. 43

and is surrounded by an oval ring, and that of the female appears
as a transverse slit in the leathery surface. Both sexes are of the
same dimensions when they become adult. The male, however, does
not perceptibly increase in length and width by feeding, whilst the
female does, and hence amongst specimens collected about a fowl
house the females are generally larger than the males.”’

Its life history has been worked out by Lounsbury at Cape Town.
As brought out by him, the life cycle and habits are strikingly dis-
similar in some important respects from those of the true ticks, the
general habits being found much more like those of the bedbug
(Cimex lectularius L.) As might be expected, we have found the
stages in the life cycle to vary somewhat from those he found. Unlike
the ticks of the family Ixodide, this species feeds for but a few hours
at a time, and then always at night, excepting in the larval stage,
when we have found it to remain attached for five to eight days
before dropping. There is also an extra nymphal molt. Unlike the
ixodid ticks again, these do not die following engorgement, but live
on, ovipositing repeatedly. Within a week or ten days from one
feeding in warm weather they again find a fowl and engorge. <As
many as five different feedings as adults have been recorded by Louns-
bury, each followed by the deposition of eggs.

In our experiments seed ticks were placed upon a fowl and obser-
vations made to determine the period of larval attachment. In
about three days from attachment they became rounded and black
from the engorgement of blood. A few hours before dropping they
commenced to flatten and assume the typical Argas shape. Attach-
ment continues for from five to eight days. In September and Octo-
ber fourteen days were found to pass after dropping before molting
took place. The attachment for second and third engorgements
Lounsbury found to last but a few hours, about two weeks to pass
after the second, and a like period following the third engorgement.
before molting.

TaBLeE 1X.—Oviposition of Argas miniatus at Dallas, Tex.

First oviposition.

First engorgement Num-

When collected. recorded: Wx. |) berior

From— To— ber of | €888-

days.
1906. |

IER Se AMS ee ee ee a en ge When collected..........- June 23) July 4 12 113.
DON eee eee aaa eta ATSC AIS 5 2 0) Se Res se May 17 | May 30 14 274
IDOE RE Sse Sat Nae CO Inns Suge ni aie CO ee ce seen ee Guek May 16 May 23 8 158.
ED) OS te ep ON AS ORG LT eet |Get Ove Mer es et cee corte May 17 | May 29 13 169
ID RSAC SUE IRE eee, Jee mere [eer Ov ce ee aise May 18 June 3 17 194
LOA ley aaS ache RARE ess ss aaa e ill beaten C0) See o eee nSaA Ese aaEe May 18 May 24 7 50

1D a) Gare BC Oe SDE BABS Aer aS Tnee: Semele ster Oo c ee toa May 16) May 22 7 32
RNR s DAE RO RNS ofS no SR ean ee A ADI TelGpacn eek esse May 2) May 20 19 130
MD) eerste nes aaa sree ate Sete met | Japs 0 Oe Soe Bee ee Aug. 1) Aug. 10 10 83

44 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

TaBLE LX.—Oviposition af Argas miniatus at Dallas, Tex.—Continued.

| Second ov. iposition.

Second _ = — Num- Third | | Total

When collected. engorge- | | Num-| ber of SUEOTES: \ Reroit
ment. | From—j| To— _ | ber of| €88s. ment eggs

days ,

BF oS ara Toe eS Se ete ciate eae leva | bistanla tals otay| le atte eae i US Been Benen ne cern 113

EaeAuiese8= || | Aug. 1 ‘| Aug. 24 11 180 | Oct. 22-23 454

.-| Aug. 9-10 | Aug. 15 | Aug. 21 | 7} 199 | Sept. 3a 357

‘| Aug. 10-11 | Aug. 16 | Aug. 25| 10 | pilOSa |e Soe ee 362

Aug. 15-16 | Aug. 21| Sept. 4| 15] 148] Oct. 17-18 342

| Aug. 17-18 0 Leet | Oil rephO! eeeprne ase eee / 50

| Aug. 3204p aes DR SR | ees: ore eeeereeee 32

| Aug. 7- 8 | Aug. 17 | Aug. 24 8 55 | Sept. 28-29 185

ID Yo vie AS Tae Ss (pa aR er ee SYNE oo | Aug 18-19 | Aug. 24 | Sept. 2 10:|. 154) Oct. 16-17 237

a Dead. b Last engorgement.

Adult ticks weighed before and after engorgement were found to
increase in weight more than 300 per cent.

In order to determine the incubation period, 35 daily lots of eggs,
deposited between May 16 and September 1, were recorded. Of
these, four lots commenced hatching in 14 days, 26 in 15 days, and 5
in 16 days. In the incubator eggs deposited August 21 and subjected
to a mean temperature of 99.8° hatched on August 29, the maximum
temperature being 108°. From experiments carried out by placing
eggs and seed ticks in an ice box and exposing’ them continuously,
vhs were found to be exceedingly resistant to cold. Eggs deposited
August 27 were exposed from September 8 to October 1 to a mean
temperature of 48.9°, a maximum of 67° and a minimum of 37°.
These commenced hatching October 6. Two lots of larve, one of 13,
that hatched September 2, and a second of 30, that hatched Septem-
ber 8, were exposed in the ice box from September 8 to October 22
to a mean temperature of 45.9°, the maximum being 67° and the
minimum 36°. These were all alive when removed and were as active
as ever October 25.

At Dallas larve kept submerged in meter to a depth of about an
inch lived for 11 days.

The length of life of this tick and its capacity to exist in the absence
of a host are surprising. At Dallas larve kept confined in summet
in pill boxes immediately after hatching lived about two months,
some surviving somewhat longer. Larvee of Margaropus annulatus
kept under similar conditions live for but two or three days at the
most. In Australia Robertson found the nymphs to live in pill boxes
for about the same period as we have found the larve to survive.

The longevity of the adult, however, is most remarkable. Riley
reports an adult specimen as remaining alive i in a corked vial without
food for five years.¢ Robertson has found them to remain alive for
two years and three months and Dr. Cooper Curtice informs us that
he has kept them alive without food for more than two years. In
our experiments adults collected in March, 1906, and kept in corked

a@Proc. Ent. Soc. Wash., III, p. 121.

CLASSIFICATION AND HABITS OF TICKS. 45

vials are still alive, March 1, 1907, although a number have succumbed.
Not less surprising than the longevity of the adult is its resistance to
insecticides. Lounsbury has kept adults confined for three months
in a box nearly filled with flowers of sulphur with no apparent effect
on them. He has also exposed them for two hours to hydrocyanic-
acid gas at the strength of 1 ounce of potassium cyanid to 150 cubic
feet of space and found that this scarcely served to decrease their
activity. Further, many individuals survived for some days after
treatment with paraffin and various oils.

Genus ORNITHODOROS.

Ornithodoros, the second genus belonging to the Argaside, is
represented in the United States by two species, O. megnini Dug.,
and QO. turicata Dug., both known to attack man.

A species widely distributed through Central and South Africa,
O. moubata, was reported by Dr. Cuthbert Christy (1903), of the
Liverpool School of Tropical Medicine, as the probable transmitter
of tick-feverin man. In 1905 Dutton and Todd,* not knowing of the
work of the other investigators, demonstrated that ‘‘tick-fever’’ in the
oriental province of Kongo Free State is a relapsing fever produced
by a spirillum, probably Spirillum (Spirachexte) obermeveri, and that
this organism can be transmitted by the bite of the tick.

The life cycle of members of this genus has yet to be followed.
Lounsbury states that O. savignyi begins to engorge at once when
applied to a host, and that generally it is off again inan hour. After
an engorgement, he states, it rests for many weeks or months and, gen-
erally, at least, sloughs its skin if immature or lays eggs if a mature
female before again seeking an animal.

It is suspected by Mr. Nathan Banks that a species of the genus
transmits a disease of cattle in California.

SPINOSE EAR TICK.
(Ornithodoros megnini Dugées. )

This tick was first described in 1883 by Dugés, from Guanajuato,
Mexico, as a species of Argas. It has been reported in the United
States from New Mexico, California, Kansas, and Nebraska, and is an
important tick in Texas. The writers are informed by a rancher in
the western part of the State that considerable injury is due to the
irritation produced by it in the ears of cattle and that its presence can
often be told by the rough appearance of the hair. A prominent
stockman in Dewitt County states that, in his opinion, it is second to
the fever tick in importance. In addition to cattle it is found upon
horses, asses, dogs, and sheep, and has been reported several times
fromman. Mr. J. D. Mitchell, of the Bureau of Entomology, reports
two cases at Victoria, Tex., in which specimens were taken from
human ears by a physician, following prolonged severe pain. Mr.

4Memoir XVII of the Liverpool School of Tropical Medicine.

46 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Mitchell informs the writers that he has known this tick to climb 3
feet from the ground and deposit its eggs in a crevice on the corner of
a house and in other cases in cracks in the bark of trees at about the
same height. From the action of specimens confined in nearly verti-
cal vials he supposes that the species normally climbs some distance
upward for the purpose of depositing the eggs.

The nymph and adult are quite different in appearance, so much so
that Neumann states that he would hesitate to consider the two forms
as belonging to the same species had he not found a cast skin about a
female. Marx figured the nymph as a new genus and species, Rhyn-
choprium spinosum. |

Stiles and Hassall have described and figured a pupa-like stage of
this tick. This, however, does not seem tobe a true resting stage, as
specimens run very actively when removed and placed in boxes. The
present writers, from live material in hand, are inclined to consider
the so-called pupa-like stage as merely the engorged larva.

The longevity of O. megnini seems to be much like that of Argas. A
specimen collected by the writers in July, 1905, lived to beyond Jan-
uary 12, 1907, having been kept in a pill box for a year and a half.

TURICATA TICK.
(Ornithodoros turicata Duges. )

This tick was originally described from Mexico. It attacks man,
the punctures causing large swellings that remain for several days
and are followed by severe pain. It has been reported from South
America on the llama; from the United States, in Florida, on a land
turtle (Gopherus polyphemus) and in a snake’s burrow, in Texas on
hogs, and in California. We have a specimen taken at Burnet, Tex.

In the Canadian Entomologist for 1900, page 20, Lounsbury men-
tions the possibility of the African species O. savignyi being intro-
duced and identical with O. turicata. He states, ‘‘ Neumann in his
monograph does not give extensive ground for separating O. sawignyi
and O. turicata. In this country [South Africa], natives are known to
carry the tick unintentionally with their belongings from place to
place. It might easily have been introduced into America with slaves
in the last century or earlier, just as negroes returning to Africa are
said to have introduced here the jigger flea (Sarcopsylla penetrans L.).
This latter insect continues to spread and is now found as far south as
Durban, Natal.”

Family IXODIDZE.

The members of the family Lxodidx from their structure naturally
fall into two subfamilies as suggested by Salmon and Stiles. The
first, Rhipicephalinz, comprises the forms with short, more or less
conical palpi, represented in this country by the genera Rhipicephalus,
Margaropus, Hemaphysalis, and Dermacentor; the second, subfamily
Ixodinx, includes forms with long palpi and is represented in the
United States by the genera Ixodes, Amblyomma, and Ceratixodes.

CLASSIFICATION AND HABITS OF TIOKS. AT

Subfamily RHIPICEPHALINZ.

Genus RHIPICEPHALUS.

The genera Rhipicephalus and Margaropus (formerly Boophilus)
are structurally so similar that Neumann and Fuller consider our M.
annulatus as belonging to the genus Rhipicephalus. All species of
Rhipicephalus, so far as studied, including five South African forms
investigated by Lounsbury and one taken by us, drop from the host
for the purpose of undergoing at least one of the molts. The three
species placed under Margaropus that have been studied pass both
molts upon the host. This would seem to supplement the structural
differences in indicating the generic validity of Margaropus.

g

Fig. 3.—Rhipicephalus sp.: Capitulum of female, ventral view. Greatly enlarged (original).

BROWN DOG TICK.
(Rhipicephalus sp.)

Salmon and Stiles in 1901 stated that they were not acquainted
with any North American species, although they called attention to
the fact that R. sanguineus had been reported by Neumann from
Panama, Ff. bursa americanus from Jamaica, and an undetermined
species from Porto Rico. Banks records an undetermined species
from Colorado. What appears to be a new species has been taken
quite generally from dogs in the southern part of Texas and also
at Tlahualilo, Durango, Mexico (see figs. 3 and 4 and PI. III, fig. 5).

.@The Arachnida of Colorado. Ann. N. Y. Acad. Sci., Vol. VIII, 1905.

48 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

It may be called the brown dog tick. Our specimens are from nine
different localities and were all taken on dogs. Unlike Dermacentor
variabilis, Mr. Mitchell in-
forms us, this species occurs
on all parts of the body of
dogs. Nathan Banks in-
forms us that this form is
closely allied to R. sangui-
neus. It is probably the
same as was referred to as
R. sanguineus in the annual
report of the Bureau of Ani-
mal Industry for 1905, page
30.

Lounsbury has found five
species belonging to the
genus Rhipicephalus that
transmit African coast fever
in cattle. The possibility
€ of the transmission of dis-
Fig. 4.—Rhipicephalus sp.: Coxe of male and female. ggse by the species we have

Greatly enlarged (original). : ‘
found remains to be inves-
tigated. 'The following are notes we have made on the life history
of this form:

TaBLE X.—Oviposition of Rhipicephalus sp., from dog.

Period of Period
First eggs deposited. Oviposition completed. ovi- from

position. dropping.
Days. | Days.
34 | 37
31 | 37
25 | 33
20 29
20 39
26 35

In the above lot of five ticks collected March 29, as will be seen, the
maximum period of oviposition was 34 days, the mimimum 20, with
an average of 26. The maximum number of eggs deposited in a lot
of seven ticks collected July 22 was 1,270, the minimum 91, with an
average of 636. An engorged tick collected April 30 commenced ovi-
position May 11, continuing for 12 days, as follows:

TasBLe XI.—Rate of ne m Sena Sp., Ove dog.

Number of eBES deposited—

Total.

May! Ma; May May! May| May may ey

May May| May May| May May
16: sllvas | slS. | Le: | 20. |

May May May
1 OB ts Ie La a bY le Se ale ae pd gees 4. | 25.

| | Se ce
0| 73 ae 181 Ist 107 eae ho. 70 2 | 20 2) 0 0 0) 1,300

a Dead.

Bul. 72, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE III.

THE NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Fig. 1.—Margaropusannulatus, male. Fig. 2.—Hemaphysalis leporis-palustris, female. Fig. 3.—Stig-
mal plate of Margaropus annulatus, male. Fig. 4.—Mouth parts of Ixodes cookei. Fig. 5.—Stig-
mal plate of Rhipicephalus sp., male. Fig. 6. —Stigmal plate of Amblyomma maculatum, female.
Figs. 1, 2, much enlarged; fig. 4, more enlarged; figs. 3, 5, 6, highly magnified. (Original. )

nna

Ma

ha
uhiEG:

of

‘edith hao —
UY) pea
1) CAR a
Me Sees

7

Bul. 72, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE IV.

STIGMAL PLATES OF TICKS.

Fig. 1.—Stigmal plates and anus of Dermacentor nitens, male. Fig. 2.—Stigmal plate of same.
Fig. 3.—Stigmal plate of Amblyomma cajennense, male. Fig.4.—Same.female. Fig. 5.—Stig-
mal plate of Dermacentor variabilis, female. Fig. 6.—Stigmal plate of Dermacentor occidentalis.
Highly magnified. (Original.)

CLASSIFICATION AND HABITS OF TICKS. 49

The incubation period of eggs deposited during the middle of April
was 6 weeks and of those deposited at the end of that month, 33 days;
seed ticks that emerged the first of June lived for 10 weeks, when kept
in test tubes on sand.

Attempts were made to rear this species by placing seed ticks on
dogs, but with poor success, as few seemed to attach. A dog was
infested with seed ticks on May 29, but none could be found attached.
On July 13, however, three adult ticks, two partially and one nearly
fully engorged, were found between the toes on a front foot. One of
the small ticks dropped July 14 and the engorged one on July 15.
In the middle of October several males of this species appeared upon
the dog, and these must have come from the above lot of seed ticks.
They were found to change the location of their attachment from
day to day. Our conclusion is that this species drops to the ground

for both molts.
Genus MARGAROPUS.

Neumann stands alone in suppressing Margaropus (Boophilus)
under Rhipicephalus. Although the two groups are closely related,
it seems evident, not only from their structure but from their habits,
that they form two distinct genera.

Neumann’s latest arrangement of the forms is as follows:

R. (M.) annulatus (Say). Southern United States.

R. (M.) annulatus var. australis (Fuller). Australia, the Antilles,
and South America.

R. (M.) annulatus var. calcarata (Birula). North Africa.

R. (M.) annulatus var. decoloratus (Koch). South Africa.
~ R. (M.) annulatus var. caudata Neumann. Japan.

R. (AL) annulatus var. argentina Neumann. Province of Buenos
Ayres.

Neumann considers that R. (J.) microplus (Canest.) is very prob-
~ ably identical with R. (M.) annulatus var. australis.

The first portion of this bulletin deals with the sole North American |
representative of this genus, M. annulatus (see Pl. III, figs. 1, 3).

Genus DERMACENTOR.

This genus is characterized by the presence of cleft front coxe in
both sexes, the fourth pair of an immense size in the male but normal
in the female. The structure of the stigmal plate furnishes valuable
characters in specific determination in this genus as do the porose
areas.

Salmon and Stiles in 1901 had but three species before them from
the United States, which they identified as D. electus Koch (varia-
bilis Say), D. reticulatus Neumann, and D. variegatus Neumann.

5795—No. 72—07——4

50 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

The species which they then listed as D. reticulatus Neumann is now
considered by Doctor Stiles and Mr. Banks as Neumann’s D. occi-
dentalis, described from specimens collected in Sonoma County, Cal-
ifornia, and labeled D. occidentalis by Marx. The D. reticulatus of
Salmon and Stiles is now considered by Banks as albvpictus Packard.
While D. reticulatus Fabricius is widely distributed, being found in
Europe and Asia, so far as known it has not been taken in this coun-
try. D. parumapertus, described by Neumann in 1901 from 4 female
specimens taken at Lakeside, Cal., labeled as taken on a man and in
a chicken house; and D. bifurcatus Neumann, from a wild cat in
Texas, described as Ixodes and later referred to the genus Derma-
centor, seem to come close together, although they may be distinct
species. According to Mr. Banks, Jxrodes nigrolineatus Packard is a
Dermacentor. To those referred to can now be added Dermacentor
nitens Neumann, which has been collected by Mr. J. D. Mitchell, of
this Bureau, making a total of 7 described species so far known to
occur in the United States.

AMERICAN DOG OR WOOD TICK.
(Dermacentor variabilis Say.)

Synonymy (on the authority of Mr. Nathan Banks): D. americanus authors (not
L.); D. electus Koch, 1844; Ixodes albipictus Pack. (Ist Peabody Acad. Rept., p. 66,
not Guide and Am. Nat.); J. quinquestriatus Fitch, 1871; I. robertsoni Fitch, 1871; J.
punctulatus Say, 1821(?).

This species is distinguished by the finely punctate stigmal plate
(see Pl. IV, fig. 5). Itis widely distributed over the country, and has
been taken commonly in northern and southern Texas and in Florida
on the dog. In some sections of Texas Amblyomma maculatum and
Rhipicephalus sp. are the common ticks on the dog, which is also the
case with J. scapularis in Florida. Neumann records a male taken
on a rabbit, Lepus callotis, by Dugés, at Guanajuato, Mexico. Cattle
also serve as hosts.

Prof. H. A. Morgan records 7,378 eggs as deposited by a single tick
between May 8 and 26. These eggs commenced hatching on August
20, an incubation period of 27 days. Our records include data on the
deposition of eggs by a tick collected April 30, oviposition commenc-
ing May 8. The details follow:

Taste XII.—Oviposition of Dermacentor variabilis.

Number of eggs deposited—
| ; feet ae V ce feat (ceo) Caunimice ela adn 2c) ae ea
SSS SISA SS SSS SAN AA A lalala ee
Be |e |. be | Br] eae) Be] bs fe tt bbe | Bes des | eae endl enc le es ee acai
AlS/S 2/25 /S/S/S/a ale alas lala aj |3/3/¢
32 | 72 ce 232 2ol 320 ie 246 217 |257 |235 |253 153 35 104 34 | 6;| 01 }-0 | 0 | (a) 2802 z
(ecm | Pere eal

aDead.

CLASSIFICATION AND HABITS OF TICKS. 51

In five lots of eggs deposited in April the incubation period varied
from 37 to 43 days. Four lots deposited during May prior to the 10th
hatched in 33 or 34 days. Seed ticks that hatched from eggs deposited
May 10 lived until November 8 or 10, being alive November 6, but
all were dead on November 10. The period of survival was thus six
months from deposition.

Under summer temperature Prof. H. A. Morgan found engorgement
of the adult to take place in from 5 to 8 days. He concludes that the
larve and nymphs attach to mammals other than cattle, as the species
has only been found on cattle in the adult stage, and attempts to cause
seed ticks to attach failed.

NET TICK.
(Dermacentor occidentalis Neumann.)

This species (see Pl. IV, fig. 6) was received by Marx from Occi-
dental, Cal. He determined it as a new species, labeling it D. occi-
dentalis. Several writers have made use of this name, but it remained
for Neumann to describe it for the first time in 1904, placing it as a
variety of D. reticulatus Fab. Curtice in 1892 referred to it briefly as
D. americanum (variabilis). It is now considered by Banks to be a
distinct species and is the one referred to by Salmon and Stiles as
reticulatus. The true D. reticulatus Fab. is not represented in our col-
lections, although it may possibly be found to occur here when a
thorough tick survey is made.

The species seems to be a western one, being found in the region of
the Rocky Mountains especially. The Bureau of Entomology and
Marx collections contain specimens from California, Washington,
British Columbia, Colorado, New Mexico, and Texas.. Salmon and
Stiles also record specimens from Oklahoma and Tennessee. In a
specimen taken from a deer skin at Kerrville, Tex., we have what is
apparently this species, there being some doubt because of the poor
condition of the individual. The recorded hosts include cattle,
horse, sheep, deer, and man.

ELK TICK.

‘( Dermacentor albipictus Packard.)

Dermacentor albipictus Packard, Am. Nat., II, p. 559; Guide to the Study of In-
sects, 9th ed., p. 662. Not Ist Rept. Peabody Acad., p. 66. (See Banks, A Cata-
logue of the Acarina or Mites, <Proc. U.S. Nat. Mus., Vol. XXXII, 1907, p. 608.)

This species is found commonly on the wapiti or “elk’’ (Cariacus
canadensis) in the States of Washington, Montana, Nebraska, Nevada,
and New York. Salmon and Stiles report that game keepers on the
reserve of the Blue Mountain Forest Association complain of its being
very common on the wapiti and that it kills numbers of them.

52 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

They suggest the possibility of disease being transmitted by it. The
specimens in the Bureau of Entomology collection from Nebraska
were taken on the beaver.

TROPICAL HORSE TICK.
(Dermacentor nitens Neumann. )

This species is readily distinguished by the,characteristic structure
of the stigmal plate. (See Pl. IV, figs. 1, 2, and text figs. 5 and 6.) By
collections by Mr.
J. D. Mitchell from
the ears of horses at
Brownsville and at
Harlingen, Tex., this
species has been
added to the list of
ticks found in this
country. The spe-
cies was described
by Neumann in
1897 from specimens
in the Marx collec-
2 & tion, locality un-

Fig. 5.—Dermacentor nitens: Capitulum of female. Greatly enlarged known, and from
(original).

ra
4,

a)
Kare

2S >

fe

Soy
ol
SER Tre"
oS

SHS

ae
<cr2

SA

SS
SG

FN

RS

me
oS

specimens from San
Domingo and Jamaica on the horse. In 1901 Neumann listed it
from Guatemala, Venezuela, and Porto Rico.

A single specimen, appar-
ently of this species in the
nymphal stage, was taken at
Kerrville, Tex., by Mr. F. C.:
Pratt from a deer skin that
had been removed in January.

Genus HE MAPHYSALIS.

The ticks belonging to this
genus may be readily recog-
nized by the presence of con-
spicuous lateral prolonga-
tions on the second palpal
segments. The eyes are ab-
sent; the coxe of the male
are all provided with spines,
those of the female withsmall]l F16- 6.—Dermacentor nitens: Coxe of male and female.

' Greatly enlarged (original).
tubercular processes. Neu-
mann mentions two species from North America, H. leporis-palustris

CLASSIFICATION AND HABITS OF TICKS. 53

and HH. concinna, but Banks informs us that he has not found the
latter in this country. He recognizes H. chordeilis Packard as the
form occurring in the eastern United States.

A South African species, H. leachi, has been determined by Louns-
bury to transmit malignant jaundice of dogs. The larva and nymph
‘both drop from the host to molt. Both engorge quickly, sometimes
in less than 48 hours from the time they attach; usually, however,
remaining from 65 to 75 hours.

RABBIT TICK.
(Heemaphysalis leporis-palustris Packard).

Synonym: Gonizxodes rostralis Duges.

Packard described this species (see figs. 7, 8, and Pl. III, fig. 2) in
1869 from a female specimen collected at Fort Macon, N. C., on a

Fig. 8.—Hemapiysalis leporis-palustris: Coxe of
maleand female. Greatly enlarged (original).
rabbit, Lepus palustris.. Marx
reported the species as quite com-
mon in Kansas, Texas, and Cali-
Fie. 7.—Hexemaphysalis leporis - palustris: Capit= fornia. Dr. Cooper Curtice has
ulum and seutum of female, dorsal view. taken an engorged female from a
oe ahaa horse in Texas, and it has been col-
lected in Mexico, the host not being given. Neumann mentions an
engorged iemale in the museum of Paris. labeled ‘from Brazil” and a

54 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

molting nymph taken on a Paradoxurus by Forbes in the Malaysian —
archipelago. Lahille reports it from Argentine Republic. During the

past two years it has been collected in Texas by agents of the Bureau at

eight different times from rabbits and ares. The species seems to be

generally restricted to the genus Lepus, but two instances of other

hosts having been recorded and these probably accidental. On the

rabbit the ear seems a favorite place for attachment. As a more

extensive collection of the tick is made it will undoubtedly be found

to occur in a much greater territory than isnow known. A few notes

on the life history of this species have been made by us.

TaBLe XIII. ives of eee es: ee

Number ofe eggs deposited—

May | May | May | May | May | May | May May | May Mary | May May | June| June
I 23. 2 I 27. | 29. 30. 31. 1 2-3.

260 | 110 | 228| 146| 131| 98| 72| 63 | 4 0 0 0 0} (2) | 1,112
0 0,| 2) 39 8 7 3 O} 0) @) |... beac learners as 57
0| 47/ 70], 48| 43| 29 Aes hemes ©) Pi | Tiles 7| (a) | 326
Oy 50) 0 0 TA, Eee Doe eo leeds 4 | 9 5 1| (a) 99

a Dead.

TaBLE XIV.—Period of incubation and longevity of Hxemaphysalis leporis-palustris.

Eggs dence: | Hatching | Minimum Remarks
ited | commenced. ition period. ; :
May 20-23... --. | jiune20es- eee 231 | March 5, dead.
May 24-29...... JUNE Zon saecee 39 | March 5, dead.
September 2-5 | September 26. 24 | May 11, dead.

Subfamiiy IXODINZ.

Three genera of this subfamily, Ceratixodes, Ixodes, and Ambly-
omma, are represented in the fauna of the United States. They may
be readily distinguished. The Ixodes have long club-shaped palpi,
the form of the third segment being typical of the genus (see Pl. III,
fig.4). The male and female have only the first coxz provided with a
spine of varying size; the anal groove surrounds the anus anteriorly,
opening posteriorly; eyes are present. In Amblyomma the palpi are
long but cylindrical; the male has a long spine on the first and fourth
cox, while in the female only the first coxe have spines, the others
possessing tubercles; the anal groove surrounds the anus posteriorly
and opens anteriorly. Ceratixodes, with pointed palpi, occurs on sea

birds.
Genus IXODES.

(The Castor Bean Ticks.)
Previous to 1796 all ticks were placed under the Linnzan genus

Acarus. In that year Latreille erected the genus Ixodes, giving J.
ricinus as the type species. For many years following all ticks

OLASSIFICATION AND HABITS OF TICKS. 55

were described as belonging to this genus, as is the case with the
species of Say, Fitch, and Packard.

Of the genera represented in the United States this has the largest
number of species. Fourteen are’ recognized, as follows:¢

I. ricinus L., I. frontalis Panzer, I. scapularis Say, I. fuscus Say,
I. brunneus Koch, I. urize White, [. cooket Packard (Synonyms: J.
cruciarius Fitch, and J. heragonus of S. & S., on the authority of
Nathan Banks), J. arcticus Osb., I. diversifossus Neum., [. dentatus
Neum., 7. angustus Neum., J. inchoatus Neum., I. sculptus Neum., and

I. californicus Banks.

Of these, four have been taken in Texas by agents of this Bureau—
ricinus, ek el, scapularis, and sculptus.

Coamperatieels little is known of the life cycle of the species of
Ixodes. The work of E. G. Wheler on ‘‘Louping Ill and the Grass
Tick” U/. ricinus) is about all that has been done. It seems prob-
able that all species drop to molt.

As Mally ascertained to be the case with /. pilosus in South
Africa, we have found engorged females of J. scapularis to dry up
in captivity before ovipositing. From Wheler’s studies and our
experience moisture seems to be a necessity in order that molting
may take place.

The longevity of the larve of ticks of this genus must be excep-
tional, even when some of the long-lived species of other genera
are considered. Wheler mentions larve (J. plumbeus?) which
hatched on October 9, 1898, from eggs laid in August, that lived
until the beginning of August, 1899, or about 10 months. They
were kept in a bottle with a sprig of moss and some damp sand.

AMERICAN CASTOR BEAN TICK.

(Ixodes cookei Packard. )

Synonymy (on the authority of Nathan Banks): Lvodes cruciarius Fitch; I. hexagonus
8. &S. (not hevagonus Leach, 1815); J. hexagonus var. longispinosa Neum.

This species(PI. III, fig. 4) was described by Packard from speci-
mens on a woodchuck, Arctomys monax, at Salem, Mass. Neu-
mann, in 1899, placed the American form with the European J.
hexagonus, but in 1901 he separated the two as varieties. Salmon
and Stiles, in 1901, followed Neumann, who at that time had all their
specimens. Banks has examined the National Museum material
and considers longispinosa the same as I. cookei of Packard, but
states that there may be a true hexagonus in this country; though he
has not seen it.

@ According to Nathan Banks, J. nigrolineatus Packard is a Dermacentor.

56 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

~

We have taken this species in Texas from a goat, a raccoon, and
a skunk. Neumann records it as taken in the United States from
the otter, mink (Putorius vison), sheep (Texas), spermophile, domestic
cat (Maine), fox (Colorado), weasel, porcupine, and marmot.

a

EUROPEAN CASTOR BEAN TICK.
(Ixodes ricinus Linnzeus. )

From its general distribution throughout Europe this species has
been called the European castor bean tick.

Mr. Banks informs us that in this species the front tarsi are
longer than in any other species of the genus found in the United
States; that in structure it is very similar to J. scapularis, but the
porose areas are larger and closer together and the scutum is more
angulate on the sides than in that species. Neumann, in his list
of hosts of the adult stage, includes sheep, goat, cattle, horse, stag,
roebuck, dog, cat, fox, ferret, hedgehog, and man; the nymphs and
larve having occasionally been found on lizards, birds, hares, rab-
bits, squirrels, polecats, ferrets, hedgehogs, mules, bats, and mice.
In the United States he lists it from Maryland, ‘‘ Carolina,” Florida,
California, Kansas, and Texas, on Lepus sylvaticus, Felis pardalis,
cattle, opossum, gray fox, panther, and wild cat. While we have
expected to find it frequently in Texas, in but a single instance has
it been taken. This was by Mr. F. C. Pratt, at Mountain Home,
from a dog. In Louisiana Prof. H. A. Morgan reports it as found
on mink in all its stages, but on cattle only in the adult stage.

Although this is an old and widely distributed species, but little
study has been made of its life history, that of E. G. Wheler, of
England, in 1899, standing practically alone. His most valuable
studies were made to determine the relation of the tick to ‘“‘louping
ill’? of sheep, with which, in the light of present knowledge, it seems
to have only an accidental connection.? The following is the sub-
stance of Wheler’s observations made in England and published in
1590;

The aduit females are readily recognized before they become dis-
tended by their deep-red bodies, dark-brown iegs, shield, and other
points. The males are of a uniform dark brown. <A record of 2,050
eggs from a single female is given, and a very interesting account of
the remarkable process of oviposition. Larve, upon finding a host,
attach, and remain for about 2 days, by which time they are distended,
black, and globular. Fully distended larve received February 7 and
kept in a bottle became hard, dry, and torpid, but on April 29, after 11
weeks, were found to have changed into nymphs and resumed active
habits. After molting the nymph takes up its position on the herb-
age, just as the larve had done, for a chance of attachment to a host.

@ Journal of the Royal Agricultural Society of England, Vol. X, pt. 4.

CLASSIFICATION AND HABITS OF TICKS. Da

Whereas adults seem to confine themselves mostly to sheep, cattle,
and deer, the larve and nymphs attach themselves very readily to
various hosts, such as horses, dogs, and human beings. About a
dozen distended nymphs taken from sheep May 29, though kept
moist, had the same dry appearance as the larve, as before deseribed.
These molted about July 19, about 11 weeks after removal from
the host. For some time previous they had appeared dead, no motion
of the extended legs being perceptible. Of these about half proved
to be males. About 10 days passed before the sexes attained their
proper color and strength. On reaching the adult stage both males
and females again wait on herbage for a passing host. At this time,
as well as after distention of the female on the host, an action which
appears to be sexual intercourse freely takes place even in confine-
ment. The rostrum and other mouth parts of the male are inserted
in the genital opening of the female, which is situated between
the bases of the posterior pair of legs. On the host the females grad-
ually distend, and in the course of so doing vary much in color and
appearance, so much so in this case that it is difficult to believe that
they are of the same species. Of the exact periods of engorgement
we are not informed. Under unfavorable conditions larve have been
kept alive for 4 months. Nymphs were kept alive for 6 months and
adult males and females for 4 months, being still alive at the time of
writing (1899). Without moisture, when kept in a dry empty bottle,
neither larve, nymphs, nor adults survived more than 2 or 3 days.
Females exposed to 25° F. for a night were found to be but little
affected.

This species, according to recent investigations of Kossel and others
in Germany, transmits European piroplasmosis of cattle. The fact
that the organism Piroplasma bigeminum is the same as is found in
this country lends great interest to investigations to determine
whether J. ricinus may not transmit Texas fever in the United States.

BLACK-LEGGED TICK.
(Ixodes scapularis Say.)

Say states that this species is rather common in forests, and is fre-
‘quently found attached to different animals. Neumann has not
recognized the species, but Banks has identified it with a form com-
mon in some parts of the South. In Texas we have collected it from
deer and dogs, and in Florida from dogs. In the latter State it seems
to be very common, and was taken at Hawthorn, Orlando, and Fort
Myers.

This species is remarkable for the size that the engorged seed tick
reaches. These are as large or larger than the engorged nymph of
Margaropus annulatus, although the adult is not as large as the adult
Margaropus. Large numbers of seed ticks and adults have been

58 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

taken on dogs, though as yet not a single nymph. This suggests the
remote possibility that the species may pass from the larval to the
adult stage at a single molt.

SCULPTURED TICK.
(Ixodes sculptus Neumann.)

This speciés was described in 1904 from a female specimen taken
with a female 7. ricinus in the Santa Cruz Mountains of California.
So far as known it had not been taken since until Mr. F. C. Pratt col-
lected it on prairie dogs at Sherwood, Tex., November 2, 1906.
More extensive collection will undoubtedly show a wider distribution

than is now known.
Genus AMBLYOMMA.

The species of the genus Amblyomma are distinguished by the
palpi, which are long and cylindrical. The male has the first and
fourth cox armed with a long spine; the female has only the first
cox with a spine, the others with tubercles. As far as known all the
species drop from the host for each molt. Lounsbury has found
Amblyomma hebreum to be the transmitter of heartwater in cattle,
sheep, and goats, and has carefully worked out its life history. .

The genus is represented in. the United States by four species—
americanum, cajennense, maculatum, and tuberculatum. A fifth
species, A. multipunctum, is described by Neumann from two speci-
mens taken on a tapir and an antelope (Dicranocerus furcatus) in
‘“‘North America.” These are reported as collected by Donckier.
As no species of tapir is found north of Nicaragua it seems probable
that A. multipunctum must have been taken from that section of
North America.

TABLE FOR SEPARATING THE SPECIES OF AMBLYOMMA OF THE UNITED STATES.

(Adapted from Neumann.)

MALES.
1. Marginal groove extending around the posterior border..........-..--..--------- 2
Marginal groove not extending around the posterior border... ...- . 4 A. tuberculatum.
Bi Come I bictspid). 5523.2 fvd oe 7 oer Se tn ea eas GN 3
Coxze’l armed: with one long spines. 5-0 ase ee oe ite ae oe ee A, maculatum.
3. Punctations cf scutum lacking from the triangular projections, flat, radiating on
posterior hallby- fj. c).cniseian sic pe Ph ee Arete See verake Sel aie te eae A. cajennense.
Punctations distributed oyer entire surface of scutum......-...-..- A. americanum.

FEMALES.

Pe Coxse Tbietspids! oF 8 see tr ee a fe te ee ta S Selae Sane ee ae 2
Coxe IT armed with:one:very long spine. S525 oe cba ee toes ees A. maculatum.
2. Scutum triangular, posterior-lateral borders nearly straight........-..---------- 3

Scutum cordiform, oval, or pentagonal, posterior-lateral borders convex.
A. tuberculatum.
3. Eyes in front of anterior third of scutum................+.-----.-- A, ‘cajennense.
Eyes at plane of or behind anterior third of scutum.........-...--- A. americanum.

CLASSIFICATION AND HABITS OF TICKS. eS

LONE STAR TICK.

(Amblyomma americanum Linneeus. )

The lone star tick derives its name from the bright-silvery spot on
the scutum of the female. It is widely distributed, having been
reported from Labrador to Florida, and also from Guatemala and
Brazil. After Margaropus annulatus, it is the most important tick
in the United States. Though found more commonly on cattle we
have taken it from man, horse, mule, dog, goat, hog, deer, squirrel,
and wolf, and it appears to attack mammals generally. In portions
of Louisiana and Texas it becomes a pest of considerable importance
to moss gatherers and other persons who spend much time in the
forests. It has been repeatedly taken in Texas during the summer of
1906 on the clothes or attached to the body of agents of the Bureau.
Packard mentions a case in which one buried itself in the arm of a
young girl, producing a raised tumor.

In May it was found on a herd of dairy cows near Dallas in large
numbers, though only an occasional specimen of Margaropus annu-
latus was present. The proprietor of the dairy stated that they were
very annoying through their attaching to milkers. Mr. J. D. Mitchell
has found it in the vicinity of Kerrville and Llano, Tex., to be the
most important species, being much more numerous than the fever
tick. In that region the cattle suffer greatly from it. Its abundance
seems to be due to the large numbers of sheep and goats kept in that
section. These serve as hosts, spreading it broadcast, at the same
time reducing the bunches of Margaropus annulatus seed ticks.

TaBLeE XV.—Oviposition of Amblyomma americanum.

ge Period of PTopping| Number

Collected ame evipo. | ip end | of exes

Pe eee ae OS aattion= |) siGadl

Days. | Days.

5 | Apr. 20 16 BiH seca
5 | May 18 14 | 22 2, 444
13 | May 21 9 | 20 984
25 | June 5 12 22 2, 508
27 | June 7 12 20 5,040
...| June 6 11 19 2,659
-| June 9 14 22 1,736
.| June 5 10 18 950
-| June 10 15 23 1,510
.| June 7 12 20 1, 306
STIS CU Bye i hee a anh aA Rc oe TE ee ee de Ba | ig) See 19, 137
TANTS es ep ees RIO berg Sen tr Rhea 2k Pane eel [ee Oe Se i DR a mere 12.5 21.2 2,126

Our records regarding oviposition, as shown in the above table, are
from 10 engorged ticks. It will be seen that the maximum number
of eggs deposited by an individual was 5,040, deposition continuing
for 12 days; the minimum, 950, with a deposition period of 10 days;
an average of 2,126 eggs deposited in 12 days. Morgan records as
many as 6,519 eggs.

60 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

TaBLE XVI.—IJncubation and longevity of Amblyomma americanum.

| | period of —

|
+ Minimum eit ce eH
Eggs deposited. menced BES Been Bu _ Oviposi- | Hatchin
to hatch. period. | : tion | rea ec
| ; to death. | :
Days. Days.
Apr DRT es Skat ae see eat ie mci eel -, June 3 103
ig STIG! OF |e AD ee es alee a ee ee
2) UMC 8) |r et ADel aae A OaEe Die aes (ee
Ap ailtvily 7 seek 129+
July 1 145

a One or more alive.

From the above table it will be seen that the incubation period in

April and May is about 7 weeks.

of May hatched in 5 weeks.

Eggs deposited in the latter part

Prof. H. A. Morgan has found some specimens to pass the first

, g

Fig. 9.—Amblyomma cajennense:
Greatly enlarged (original).

were again submerged for 45 hours.

specimen, survived.

Mouth parts of

molt on the host, drop-
ping in about 10 days fol-
lowing that molt, or just
previous to the second
molt. The greater num-
ber dropped in from 4 to 7
days. The molting of
the nymph was found to
last 6 weeks. Engorge-
ment of the adult in March
and April occupied from
7 to 11 days.

The resistance of the fe-
male to water seems to be
similar to that of the fever
tick. Seven females were
submerged for 18 hours,
all becoming active fol-
lowing removal from the
water. The next day all
Only one, an unengorged

male.

In the adult stage both sexes will reattach, as seems probable in the

larval and nymphal stages.

Mr. Mitchell has observed adults on grass in the act of copulation.

CAYENNE TICK.

(Amblyomma cajennense Fab.)

Synonyms: J. crenatwm Say (Banks), A. mixtum Koch (Banks), J. herrere Duges

(Neum.), A. sculptus Berlese (Neum.).

This species was described from Cayenne in 1794, no host being

CLASSIFICATION AND HABITS OF TICKS. 61

given. The writers’ opinion of the specific identity of specimens from
Venezuela, on cattle, with Texas specimens, on horse and peccary, is
confirmed by Mr. Banks. The species may be distinguished from
maculatum readily by the characters given in the table. (See also
figs. 9 and 10 and PI. IV, figs. 3, 4.)

The species has been reported from, and seems to be generally dis-
tributed through Mexico and Central America. It has been reported
from Colombia, Venezuela, Brazil, and Argentine Republic in South
America. Neumann also reports it from Cuba and Jamaica. In Texas
Mr. J. D. Mitchell has taken it in Live Oak County from the peccary
and horse. Mr. Banks reports specimens from Louisiana, Missouri,
and Florida. In addition to the hosts mentioned it has been reported
from toad (Bufo agua), capy-
bara (Tydrochxrus capybara),
an ant-eater, and man.

Stoll, in the Biologia Cen-
trali- Americana, states that
this species is the most com-
mon of all [xodidz in Central
America, and gives some in-
formation concerning its hab-
its. He has never found the
male in a parasitic state, but
has found it free on grass and
bushes in Guatemala. The fe-
male, which he states abounds
in the woods and fields on
grass and bushes, is occasion-
allv brushed off by horses, ecat- Fic. 10.—Amblyomma cajennense: Coxe of male and

A = female. Greatly enlarged (original).

tle, or dogs, and even man.
It adheres tenaciously to the skin, remaining when undisturbed for
several days until filled with blood. If forcibly removed, the beak
breaks off and remains in the wound, causing a disagreeable and
sometimes painful inflammation. The young, which are distinguished
by the inhabitants of Guatemala by the name of ‘‘ mostacilla,”” hang to
the grass in clusters of thousands, especially during the dry seasons.
By their creeping on the skin and frequent biting they form one of the
greatest plagues of travelers.

Ina letter accompanying specimens of this species from Venezuela the
writers are informed that the ticks do great damage by producing fever
in cattle, which become weak and in many cases die. It hardly seems
possible that the malady can be Texas fever; nevertheless this species
may possibly transmit some disease.

62 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Taste XVII.—Record of deposition of an engorged tick, Amblyomma cajennense, col-
lected on peccary April 29.

Number of eggs deposited—
|)3/]a ¢/2/4/2|s ¢lalglalalals|aialslala|s a!
bm | b& = eS itsse itso elves tesel letesy lie fex |
2|2\2/4/2/3/9 2 2/3|2/2|2|2|s g\sisleeisa
er 8 212 278 P| 200 289) 213 336 307 206 119 bal ai 33 a3 23)) 3 ‘ 4 ee 2,650

|

a Dead.

It will be seen that oviposition continued for 20 days from May 9,
a total of 2,650 eggs being deposited. Eggs deposited May 14, com-
menced hatching June 21, an incubation period of 38 days. Two
seed ticks from eggs de-
posited May 14-25 were
alive November 6, a pe-
riod of more than 5
months from deposition.
The fact that this pro-
longed survival included
summer months shows
remarkable vitality in the
seed ticks of this species.
Seed ticks of Margaropus
annulatus seldom survive
over 3 months in the
summer.

GULF COAST TICK.

(Amblyomma maculatum
Koch.)

—— Synonyms: A. tigrinum
Fig. 11.—Amblyomma maculatum: Mouth parts of female. Koch, A. tenellum Koch, A. ru-

Greatly enlarged (original). bripes Koch, A. ovatum Koch,
A. triste Koch, A. complanatum Berlese (the preceding on the authority of Neumann),
Dermacentor occidentalis Marx of Niles (Morgan).

This species was described by Koch in 1844 from ‘‘Carolina,’”’ no
host being given. Prof. H. A. Morgan calls our attention to the fact
that it was referred to as Dermacentor occidentalis Marx by Niles.@

It seems to be the common species along the Gulf coast of Louisiana,
and Texas. (See figs. 11, 12, 13, and Pl. HI, fig. 6.) It was taken
by Mr. J. D. Mitchell of this Bureau from cattle, horses, dogs, and
man in Cameron Parish, La., and Calhoun, Jackson, and Victoria
counties, Tex. In the Marx collection there is a male specimen

a Bul. Va. Agric. Exp. Sta., VII, No. 3, pp. 28, 29, Pl. IV. .

CLASSIFICATION AND HABITS OF TICKS. 63

recorded as taken from cattle at Memphis,.Tenn. It seems quite

probable that the animal f

bred in the Coast sec-
tion. There are also
several specimens in
the Marx collection
taken in Texas. Neu-
mann reports it from
Paraguay, Uruguay,
Brazil, Mexico, and in
the United States from
California, Texas, and
Tennessee, the latter
based upon. the tick
before mentioned as
collected at Memphis.
He mentions 2 males
and 1 female as being
taken on a coleopteron,
Cercus campestris, at
Buenos Aires, Argen-
tina. lLahille reports
it from Argentine Re-
public, where the fa-
vorite host is the dog.
He mentions the fact

rom which the tick was collected had been

~

\

1 &

Fic. 12.—Amblyomma maculatum: Cox of male and female.

Greatly enlarged (original).

that itis used by the Indians as a leech in certain cases of inflammation.
The male is especially large, much more so and more elongate than

Fic. 13.—Amblyomma maculatum: Scutum of
female. Greatly enlarged (original).

either A. cajennense or A. ameri-
canum. Mr. Mitchell reports that-
while he has observed the sexes in
close proximity on the animals he
has as yet to find them in coitu. He
has frequently noticed them to copu-
late after being removed from the
animal. However, in Argentina,
Lahille states that several males
are usually found attached in the
immediate vicinity of each female.
Mr. Mitchell states that on one oc-
casion he found 7 females clustered
on a cow between the ear and the
horn with no male near. In another

case he found a cluster of 5 females on the neck of a dog with no male
in the immediate vicinity. He has also seen instances where as

’

64 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

many as 5 males were located very close together without any
females, and other cases on various parts in which both sexes were
found together. This species is very firmly attached to the host.
While Rhipicephalus sp. can be removed easily A. maculatum can not
be removed without evident pain.

Attempts were made to rear the species upon dogs. On October
11 seed ticks were placed upon two small dogs. Eight were found
attached on the 15th. For the first five or six days after attach-
ment the seed ticks increase slowly in size, remaining of a light color;
then in a few hours they seem suddenly to engorge with blood.
The color of the body becomes’ purplish and afterwards still darker.
On the fifth day from placing seed ticks on the dogs 8 were found
still light in color and 1 dark. On the sixth day 2 were taken that
had just detached themselves, leaving 2 dark and 4 light on the host.
On the seventh day 3 were found to have dropped during the night;
2 dropped and the remaining 1 engorged during the day. On Octo-
ber 19 the eighth tick was found to have dropped during the night,
making a minimum period of 5 days and a maximum period of 8 days
for attachment of larval stage.

Several of the engorged larve were kept on moist sand to molt,
but shriveled and died.

The incubation period for eggs deposited from the Ist to the 11th
of September varied from 26 to 31 days. Seed ticks from these eggs
were still alive on March 1, 1907.

LAND TURTLE TICK.

(Amblyomma tuberculatum Marx. )

This species was described by Marx in 1894 from specimens col-
lected by Hubbard at Crescent City, Fla., from the Florida land tur-
tle, Gopherus (Xerobates) polyphemus. Neumann reports a male
specimen from Cuba.

We have received 3 specimens collected in February, 1907, by Dr.
A.W. Morrill in Florida on a land turtle (Gopherus sp.), presumably
the same as the host of the original specimens.

BIBLIOGRAPHY.

No attempt has been made to make this bibliography complete.
In Section A, relating to the cattle tick of North America and the
problems caused ‘by it, the literature is scattered through many
experiment station bulletins, veterinary journals, bulletins of the
Department of Agriculture, and other publications. Only the more
important are included. In Section B, relating to foreign disease-

BIBLIOGRAPHY. 65

transmitting ticks, the literature is found for the most part in the
governmental publications of South Africa and Australia, although
recently important memoirs have been issued in Germany and France.
As in the preceding section, we give only the more important publi-
cations that are now available. For a very complete bibliography
of this subject down to 1903, containing 221 titles, see A. Schmidt
below. In Section C, relating to ticks as transmitters of human
diseases, there is at present but a scanty literature. It is practi-
eally all referred to by us. In Section D, relating more particularly
to the classification of ticks, practically all of the works dealing
with North American species are listed, together with the more impor-
tant foreign publications. For a more extended list,see Salmon and
Stiles, 1901, below.
SECTION A.

RELATING TO THE NortH AMERICAN FEVER-Tick PROBLEM.

Butter, Tair.
1902.—The breeds of beef cattle and beef production in North Carolina <N. C.
State Bd. Agr., bul. 23, no. 7.
Pages 48-51 deal with: ‘‘ Cattle tick an obstacle to the development of the cattle
industry.”
1903.—Report of State Veterinarian in Rept. Comm. Agr. N. C. for 1902, pp. 40-47.
Regarding the cattle quarantine, the extermination of the cattle tick, losses from
tick fever, and the tick an obstacle to the improvement of the quality of the cattle
of the State.
1903.—The cattle tick and its relation to the cattle industry of North Carolina
<N. C. State Bd. Agr., bul. 24, no. 5, pp. 28, figs. 2. .
1903.—The cattle tick and the quarantine restrictions <N. C. State Bd. Agr.,
bul. 24, no. 10, pp. 30-37. p

1906.—Progress made in exterminating the fever tick (Boophilus annulatus) in
North Carolina <N. C. Dept. of Agr. circular (unnumbered) issued
January 1, 1906, pp. 4. Reprinted in part in Farm and Ranch, vol. 25,
no. 15, p. 7, April 14.
This circular deals with the most successful work that has been done in the practical
eradication of the cattle tick.
ConnaAWAY, J. W.
* 1897.—Texas fever or acclimation fever <Mo. Agr. Exp. Sta., bul. 37, pp. 81-139,
figs. 11, April.
Connaway, J. W., and M. FRANcIs.
1899.—Texas fever <Mo. Agr. Exp. Sta., bul. 48, pp. 66, figs. 11.

CurRTICE, COOPER.

1891.—The biology of the cattle tick <Journ. Comp. Med. and Vet. Arch., vol. 12,
no. 7, pp. 313-319, July.

1892.—About cattle ticks <Journ. Comp. Med. and Vet. Arch., January.

This and the preceding paper were issued together with independent continuous

pagination.

1892.—The cattle tick <Texas Agr. Exp. Sta., bul. 24, pp. 237-252, 2 pls.,
December. Appendix on preventive measures by M. Francis. Ab-
stract, Insect Life, vol. 5, p. 294.

This bulletin gave the first details of the life history of the cattle tick. It was a
pioneer work the value of which has been appreciated by all subsequent workers.

5795—No. 72—07 3)

66 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

CurtTIcE, CoopErR—Continued.
1896.—On the extermination of the cattle tick and the disease spread by it
<Journ. Comp. Med. and Vet. Arch., vol. 17, pp. 649-655.

On page 655 Doctor Curtice gives the first reference we have found to the possibility
of totally exterminating the cattle tick from the United States. ‘‘ 1look most eagerly
for the cleansing of even a certain portion of the infected territory under the direct
intention of man, for it opens the way tc pushing the ticks back to the Spanish isles
and Mexico, and liberating cattle from disease and pests and the farmer from untold
money losses. Let your war cry be: Death to the ticks!’’

1897.—The cattle tick and what may be done to prevent it <The Southern
Planter, vol. 58, no. 1, pp. 24-27, January.

This paper contains the original suggestion of the pasture eradication and feed-

lot systems of tick eradication.

1897.—Texas fever <The Southern Planter, vol. 58, no. 3, pp. 116-117 , March.

1899.—Cattle quarantine, ticks and distemper <N. ©. State Bd. Agr., bul.
(unnumbered), July 1.

1899.—Cattle quarantine line <N. C. Agr: Exp. Sta., spec. bul. 52, pp. 1-28.

1905.—The cattle-tick plague <The Southern Planter, vol. 66, no. 5, pp. 376-378,
May.

A report of progress in extermination and a continuation of the two articles pub-

lished in the above Journal October, 1896, and January, 1897.
DauryMP_e, W. H., H. A. MorGan, and W. R. Dopson.
1898.—Cattle tick and Texas fever <La. Agr. Exp. Sta., bul. 51, pp. 230-282,
figs. 2, pls. i—vil.

Detailed account of many experiments relating to the life history of Margaropus
annulatus. Includes studies of the effects of water, heat, cold, and light on the eggs,
seed ticks, and adults, and much information on pasture eradication. The plates
are excellent. This publication adds greatly to the data furnished by Doctor Curtice
in Texas Bulletin 24. '

1899.— Immunization against Texas fever by blood inoculation <La. Agr. Exp.
Sta., bul. 57, ser. 2, pp. 146-185, pls. 2, chts. 3, figs. 3.

Dinwippre, R. R.
1892.—Parasitism of domesticated animals < Ark. Agr. Exp. Sta., bul. 20, pp. 14-31.
Treats of mites, cattle ticks, etc.
1896.—Texas cattle fever in various localities <Ark. Agr. Exp. Sta., report, 1896,
pp. 36-40.
A reprint from bulletin 40 of the station.
1898.—Methods of combating communicable diseases of farm animals < Ark.
Agr. Exp. Sta., bul. 51, pp. 35-46, May.
Francis, M.
1889.—Report of the veterinarian, Tex. Agr. Exp. Sta., second annual report,
1889, pp. 55-60.

A brief account of the southern cattle plague and its treatment, with notes on joint
experiments by the Texas and Missouri Stations.

1892.—The cattle tick, preventive measures for farm and range use <Tex. Agr.
Exp. Sta., bul. 24, pp. 253-256, fig. 1, December.
1894.— Veterinary Science <Tex. Agr. Exp. Sta., bul. 30, pp. 436-458, March.
Treats of animal parasites, device for destroying ticks, and influence of ticks in the
dissemination of Texas cattle fever.
Francis, M., and J. W. ConNnaway.
1899.—Texas fever <Tex. Agr. Exp. Sta., bul. 53, pp. 53-106, figs. 13.

Lewis, L. LL.
1897.—Texas fever <Okla. Agr. Exp. Sta., bul. 27, pp. 8-15, fig. 1, June.

a BIBLIOGRAPHY. 67

Lewis, L. L.—Continued.
_ 1899.—Texas fever. An account of experiments <Okla. Agr. Exp. Sta., bul. 39,
pp. 28, figs. 5, May.
1900.—Means of preventing Texas fever <Okla. Agr. Exp. Sta., report, 1900,
pp. 26-28.
Mayer, AUGUST.
1906.—The cattle tick in its relation to southern agriculture le S. Dept. Agric.
farmers’ bul. No. 261, pp. 1-22.
Anexcellent essay on the broad aspects of the tick problem.
Mayo, N.S.
1897.—Texas fever <Kans. Agr. Exp. Sta., bul. 69, pp. 124-134, June.

McCuttoca, C.
1899.—The prevention of Texas cattle fever and the amended laws controlling
contagious and infectious diseases <<Va. Agr. Exp. Sta., bul. 104,
pp. 167-180, September.

Me .vin, A. D:
1906.—How to get rid of cattle ticks <U.S. D. A., Bur. A. I., cir. 97, pp. 1-4,
1 pl.
Deals with the hand-picking and greasing, the tie-rope or picketing, and the two-
field methods.

MinuER, W. McC. N.
1895.—Texas cattle fever <Nev. Agr. Exp. Sta., bul. 31, pp. 11, December.
Mouter, J. R.
1906.—Texas fever (otherwise known as tick fever, splenetic fever, or south-
ern cattle fever), with methods for its prevention <U. S. D. A., Bur.
A.I., bul. 78, pp. 1-48, pls. 3, figs. 3.
A comprehensive treatment of the whole subject, including symptoms, pathology,
therapeutics, and methods of tick eradication.
1906.—Texas or tick fever and its prevention <U.S. D. A., farmers’ bul. 258,
pp. 1-46, figs. 1-6.

This is a somewhat condensed re-edition of the preceding.

Morean, H. A.
1899.—Ticks and Texas fever <La. Agr. Exp. Sta., bul. 56, pp. 128-141, pls. 9,
September.
This is the only American work on the life history of species other than Margaropus
annulatus. It deals with Amblyomma americanum, Dermacentor variabilis (electus) ,
both of which were found to be nonpathogenic as far as splenetic fever is concerned,
and Irodes ricinus. Many details of life history are given. The plates are most
valuable. Marx’s paper on the classification of the Ixodide is reprinted as a part of
this bulletin.
The starving-out method of eradication is again emphasized. (See Dalrymple,
Morgan, and Dodson, 1898.)

t 1903.—How can we exterminate the cattle tick? <Proc. La. State Agr. Soc. and
Stockbreeders’ Assn., 1903, pp. 77-79.

Emphasizes the necessity for cooperation among the farmers.

The cattle-tick situation <Proc. Soc. Prom. Agr. Sci., 1903, pp. 72-74.
Notes are given on the life history of the cattle tick.

1905.—Texas fever cattle tick: pasture methods of eradication <Bul. of Agr.

Exp. Sta. Univ. Tenn., vol. 18, no. 1, pp. 1-10, figs. 3.
Also published as bul. 82 (second series), La. Agr. Exp. Sta., pp. 1-15.

This publication urges the feasibility of the practical application of a knowledge
of the life history of the cattle tick in feed-lot and pasture-eradication systems of
eradication and centered public attention. It gives full details as to procedure.

See also Dalrymple, Morgan, and Dodson.

eee.) le a

yee"

68 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Newe i, Witmon, and Douauerty, M. 8.
1906.—The cattle tick. Studies of the egg and seed-tick stages. A simple
method of eradicating the tick <La. Crop Pest Comm., cir. 10, pp. 1-32,
figs. 1-8.
Contains records of very careful work on the egg and seed-tick stages. The prac-
tical importance of the work is well demonstrated.

Nines, E. P.
1897.—The cattle tick in Virginia <Va. Agr. Exp. Sta., bul. 76, pp. 45-50,
May. Southern Planter, July, 1898, pp. 326-327.
1898.—A preliminary study of ticks <Va. Agr. Exp. Sta., bul. 86, pp. 25-30,
4 pls., March.

NOrGaAaArD, Vicror A.
1898.—Cattle dipping, experimental and practical < U.S. D. A., yearbook, 1898,
pp. 453-472.
Experiments, principally with saturated solutions of sulphur in extra dynamo oil
Ransom, B. H.
1906.—Some unusual host relations of the Texas fever tick <U. 8. D. A.,
Bur. A. I.; cir. 98, pp. 1-8.

Details experiments in attempting to cause Margaropus annulatus to develop on
horses, mules, asses, rabbits, dogs, and eats. On the last a specimen developed to
the adult stage, but did not engorge. Shows that under certain conditions the cattle
tick can reattach after being artificially removed. Refers to recent European investi-
gations that show that Jrodes ricinus transmits European piroplasmosis of cattle,
and suggests the possibility that Margaropus annulatus may transmit its disease in
the nymphal or adult states.

Repprina, R. J. ;
1889.—Cattle ticks and Texas fever <Ga. Agr. Exp. Sta., bul. 49, pp. 228-229.

Ritey, C. V., and L. O. Howarp.
1899.—The Texas cattle tick <Insect Life, vol. 2, July, 1889, p. 20. Habits
and remedies.

‘

RoseErt, J. C.
1897.—Acclimation fever, or Texas fever < Miss. Agr. Exp. Sta., bul. 42, pp. 32,
figs. 4.
1901.—Texas fever <Miss. Agr. Exp. Sta., bul. 69, pp. 1-15, figs. 4, November.
1902.—Tick fever or murrain in Southern cattle (commonly termed Texas fever)
Miss. Agr. Exp. Sta., bul. 73, pp. 1-24, figs. 2, July.

Saumon, D. E.
1884.—Geographical distribution of Southern cattle fever <In report of the
Chief of the Bureau of Animal Industry. U.S. Comm. Agr., report,
1884, pp. 252-258, pls. 3.
Discusses occurrence of fever in Va., N. C., 8. C., Ga., Tenn. Maps show limits
east but not west of Mississippi River.
1904.—Relations of Federal Government to control of contagious diseases of
animals <U. 8. D. A., yearbook, 1903, pp. 491-506; pp. 505 et seq.

Deals with M:rgaropus cnnul<tus.

Saumon, D. E., and THEOBALD SMITH.
1892.—Southern cattle fever (Texas fever) <U.S. D. A., Bur. A. I. special
report on diseases of cattle and cattle feeding, pp. 428-438, pls. 42-44.
1904.—Texas fever, or Southern cattle fever <(U. S. D. A., Bur. A. I., cir. 69,
pp. 1-13. (Reprint from special report diseases of cattle, revised, 1904,
by Salmon and Mohler.)

BIBLIOGRAPHY. 69
Saumon, D. E., and C. W. Srizzs.
1901.—The cattle ticks (Ixodoidea) of the United States <U.S. D. A., Bur. A. L.,
17th ann. report, 1901, pp. 380-491, pls. 74-98, figs. 47-238.
This is a most valuable work, particularly rich in bibliographical references and
illustrations. It is an absolute essential in the study of the ticks of this country.
ScHroeper, E. C.
1900.—A note on the vitality of the Southern cattle tick <U.S. D. A., Bur. A. L.,
16th ann. report, pp. 41-42.
ScHROEDER, E. C., and W. E. Corron.
1900.—Growing noninfected ticks and afterwards infecting them <U.S8. D. A.,
Bur. A. I., 16th ann. report, pp. 33-41.

Smith, THEOBALD, and F. L. Kinporne.
1893.—Investigations into the nature, causation, and prevention of Texas or
Southern cattle fever <<U. S. D. A., Bur. A.I., bul. 1, pp. 1-301,
pls. 10, figs. 7.

This scholarly work demonstrated the transmission of fever by Margaropus annu-
latus. It suggests much of the work since done in the study of disease transmission
by insects and ticks.

Suiru, T., F. L. Kizporne and E. C. ScHRoEDER.
1893.—Additional observations on Texas cattle fever <U. 8. D. A., Bur. A. I.,
bul. 3, pp. 67-72.
VINCENHELLER, W. G.
1906.—The cattle tick in Washington and Benton counties< Ark. Agr. Exp. Sta.,
bul. 90, pp. 131-141.
Wittovuauey, C. L.
1904.—Cattle ticks and Texas fever; immunizing experiments in Georgia <Ga.
Agr. Exp. Sta., bul. 64, pp. 143-182, figs. 9, August.

SECTION B.

RELATING TO FoREIGN DISEASE-TRANSMITTING TICKS.
Banks, C. 8.
1904.—The Australian tick (Boophilus australis Fuller) in the Philippine Islands
<U.S. D. War, Bur. Govt. Laboratories, bul. 14 (Biological Laboratory,
Div. Ent., bul. 2), pp. 13-21, figs.. 16-23, pls. 9.
Bruce, D.
1905.—The advance in our knowledge of the causation and methods of prevention
of stock diseases in South Africa during the last ten years <Science,
n. 8., vol. 22, pp. 289-299 and 327-333.
Epineton, A.
1904.—Note on the co-relation of several diseases occurring among animals in
South Africa <Agr. Journal Cape Good Hope, vol. 25, pp. 139-152.

Froae@att, WALTER W.
1901.—The fowl tick (Argas americanus Packard) <Agr. Gazette N. 8. Wales,
vol. 12, pp. 1349-1352, pl.

FULLER, CLAUDE.
1896.—Ticks, a fowl-infesting tick (Argas sp.) <Agr. Gazette N.S. Wales, vol. 7,
pp. 590-597 (reprint, pp. 1-8).
1896.—The bovine tick fever <Agr. Gazette N. S. Wales, vol. 7, pp. 760-787,
pls. 1-5. .
1899.—The common blue tick <Agr. Journal Cape Good Hope, vol. 14, pp. 363-—
369, March.

70 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Gray, C. E., and W. Roprrrson.
1902.—Report upon Texas fever or redwater in Rhodesia <Argus Printing
and Pub. Co., Ltd., Cape Town, pp. 27, pls. 22.
1902.—Redwater in Rhodesia <Agr. Journal Cape Good Hope, vol. 21, pp. 435—
458, November.
Herrera, A. L.
1905.—Las Pardsitos del Ganado <Com. Parasit. Agricola., cir. 8, pp. 22-25,
figs. 31-34.
Hunt, Smney. 4
1898.—Tick fever Queensland Agr. Journal, pp. 235-236.

HurcHeon, D.
1900.—Redwater and its history <Agr. Journal Cape Good Hope, vol. 17, pp.
331-339, 395409.
1900.—History of heartwater <Agr. Journal Cape Good Hope, vol. 17, pp. 410-
417.
1903.—Virulent redwater in the Transvaal <Agr. Journal Cape Good Hope,
vol. 23, no. 1, pp. 39-60.
JOBLING, J. W., and P. G. Woo.ey.
1904.—Texas fever in the Philippine Islands and the Far East <U. 8S. D. War,
Bur. Govt. Laboratories, bul. 14, pp. 5-11, pls. 15.
Kosssrt, H., A. Weser, Scuitirz, and Minssner.
1903.—Ueber die Himoglobinurie der Rinder in Deutschland <Arb. K.

Gesundheitsamte, no. 1, pp. 1-77, pls. 3.
The blood parasite is the same as that of Texas fever. Ixodes ricinus (reduvius)
‘ acts as a transmitter.

,

LanItieE, F.
1905.—Contribution a l’étude des Ixodidés de la République Argentine, pp.
1-166, pls. 1-18.
This paper contains a great deal on the biology of Margaropus microplus.
LIGNIERES, J.
1900.—La Tristeza ou Malaria bovine dans la République Argentine, pp. 1-172,
pls. 14.

Lounssury, C. P.
1899.—The bont tick Amblyomma hebreum Koch <Agr. Journal Cape Good
Hope, vol. 15, pp. 728-743.
1900.—Life history of a tick <Ent. News, vol. 11, pp. 336-340, January.
Life history of Amblyomma hebreum Koch.
1900.—Tick-heartwater experiment <Agr. Journal Cape Good Hope, vol. 16,
pp. 682-687.
1900.—Insect bites and the effects thereof <Can. Ent., vol. 32, pp. 17-24.
Argas and Ornithodoros spp.
1900.—Notes on some South African ticks <U. 8S. D. A., Bur. Ent., bul. 26,
N. 6., pp. 41-49:
1902.—Report of Government Entomologist for the Cape of Good Hope for 1901.
Includes ‘‘Tick heartwater investigations,’ pp. 29-73, pls. 4-6.
1903.—The fowl tick. Studies on its life cycle and habits <Rept. Agr. no. 20,
pp. 1-15, pls. 3. Reprint Agr. Journal Cape Good Hope, September.
1903.—Report of the Government Entomologist for the Cape of Good Hope for
1902.
‘“Ticks and Rhodesian cattle disease,’ ‘‘ Ticks and malignant jaundice,’’ ‘‘ Ticks
and heartwater,’’ pp. 16-41.
1903.—Ticks and African coast fever <Transvaal Agr. Journal, vol. 2, NO: 70%
pp. 4-13.

BIBLIOGRAPHY. 71

Lounssury, C. P.—Continued.
1904.—External parasites of fowls <Reprinted from the Agr. Journal, pp. 7,
November.
Relating to Argas persicus.
1904.—Persian sheep and heartwater <Agr. Journal Cape Good Hope, vol. 25,
no. 2, pp. 175-186, figs. 3.
1904.—Distribution of coast fever ticks < Agr. Journal Cape Good Hope, vol. 25,
no. 3, pp. 268-270, pl. 1.
The distribution of Rhipicephalus appendiculatus, R. simus, R. evertsi, and R.
capensis is briefly outlined.
1904.—Transmission of African coast fever <Rept. Agr., no. 5, pp. 1-7, pls. 3.
Reprint: Agr. Journal, Cape Good Hope, April.
1904.—Report of the Government Entomologist for the Cape of Good Hope for
1903.
Includes ‘‘ticks and malignant jaundice’’ and ‘‘ticks and heartwater,”’ ‘‘ticks and
South African coast fever,” pp. 11-45, pls. 7
1905.—Tests of substances for tick destruction <Agr. Journal Cape Good Hope,
vol. 26, pp. 387-395, March.
Report of the Government Entomologist for the Half Year ended June 30,
1904.

Contains a special report, ‘‘ticks and African coast fever,” pp. 10-25.

1905.

Matty, C. W.
1904.—Notes on the so-called paralysis tick, [vodes pilosus <Agr. Journal Cape
Good Hope, September. Reprint by Dept. of Agric., no. 17, 1904.

Marcuaux,. E., and A. SALIMBENI.
1903.—La Spirillose des Paules < Annals 1’ Institut Pasteur, vol. 17, pp. 569-580.

Spirillosis of chickens and other fowls transmitted by Argas miniatus.
Moras.

1903.—The role of ticks in the development of carceag <Compt. Rend. Soe.
Biol., Paris, vol. 55, no. 14, pp. 501-504. é

The writers have seen only a review of this paper in Experiment Station Record.

Pounp, C. J.
1899.—Notes on the cattle tick. Its development, hfe history, habits, and geo-
graphical distribution <P. Soc. Queensland, vol. 14, pp. 28-38.

Ropertson, F. H.
1905.—Fowl tick experiments <Journ. Dept. Agr. West Australia, vol. 12, no. 6,
pp. 561-563.

It was found that fowl ticks remain alive at least 2 years and 3 months without the
presence of any fowls from which to derive nourishment. In these experiments the
ticks were kept in small pill boxes which were practically air tight. In the nymph
; > stage ticks may live for 2 months without food.

ScHMIDT, A.
1904.—The tick disease of cattle (hemoglobinzemia ixodioplasmatica boum) in
German and English East Africa and Uganda <Arch. Wiss. u. Prakt.

Tierh., vol. 30, nos. 1-2, pp. 42-101.

The literature of this subject is discussed with references to a bibliography of 221
titles. We have not the original work at hand and refer to the translated title in
the Experiment Station Record, XVI, p. 201.

STOCKMAN, STEWART.
1904.—Rhodesian redwater, vel East African coast fever, vel tropical piroplas-
mosis <Report of the Transvaal Dept. of Agric., 1903 to 1904, pp.

40-66.
Includes history of invasion and spread in Transvaal, permit system, dipping,
immunity, preventive inoculation, prevention and eradication, and transport experi-

ments.

72 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

THEILER, A.
1903 and 1904.—The Rhodesian tick fever <Transvaal Agr. Journ., vol. 1
(1903), no. 4, pp. 93-110, pl. 1; vol. 2 (1904), no. 7, pp. 421-488, pl. 1.

1904.—The transmission of East Coast fever by ticks <Transvaal Agr. Journ.,
vol. 3, no. 9, pp. 71-86, October.

1905.—Further notes on piroplasmosis of the horse, mule, and donkey <Trans-
vaal Agr. Journ., vol. 3, no. 12, pp. 706-716.
1906.—Transmission and inoculability of spirillosis in cattle <Transvaal Dept.
Agr., ann. rept. Dir. Agr. 1904-1905, pp. 123-151.
The writer shows that the natural transmission of spirillosis is by the progeny of
Rhipicephalus decoloratus, which have developed on cattle affected by spirillosis.
WHELER, E. G.
1899.—Louping ill and the grass tick <Journ. Royal Agr. Soc. England, ser. 3,
vol. 10, pt. 4, pp. 626-644.
See note under following title.
1902.—Parasitically inoculated diseases <Trans. Highland and Agr. Soc. Scot-
land, ser. 5, vol. 14, pp. 16-35, figs. 2.

Surmises that ‘‘louping ill’’ is transmitted by Ixodes ricinus. Later investigations
have negatived this.

SECTION C.
RELATING TO TRANSMISSION OF HuMAN DISEASE By TICKS.

CHRISTY, CUTHBERT.
1903.—‘‘ Tick fever’? in man <The Thompson Yates and Johnson Laboratories
Report, vol. 5, n. s., part I, pp. 187-189.

Duron, J. E., and J. L. Topp.
1905.—The nature of human tick fever in the eastern part of the Congo Free State,
with notes onthe distribution and bionomics of the tick <Liverpool

School of Tropical Medicine, memoir 17, pp. 26.
Includes paper by Robert Newstead, ‘“‘On the external anatomy of Ornithodoros
moubata (Murray).’’

Kine, W. W.
1906.—Experimental transmission of Rocky Mountain spotted fever by means of
the tick <U. 8S. T. D., Public Health and Marine-Hospital Service, .
Public Health Reports, vol. 21, pp. 863-864, July 27.

NewstTeEaD, R.
1905.—On the external anatomy of Ornithodoros moubaia <Liverpool School of
Tropical Medicine, memoir 17, pp. 21-26, November.

RickHnre He i.
1906.—The study of ‘‘Rocky Mountain spotted fever” (tick fever?) by means
of animal inoculations <Journ. Am. Med. Assn., vol. 47, pp. 33-36,
July 7.

1906.—The transmission pf Rocky Mountain spotted fever by the bite of the
wood tick (Dermacentor occidentalis) <Journ. Am. Med. Assn., vol. 47,
p. 358, August 4.

Stings, Cu. W.
1905.—A zoological investigation into the cause, transmission, and source of Rocky
Mountain ‘‘spotted fever’? <<U. 8. T. D., Public Health and Marine-
Hospital Service, Hygienic Laboratory, bul. 20, pp. 1-121.

BIBLIOGRAPHY. 73

SECTION D.

RELATING TO THE CLASSIFICATION AND DISTRIBUTION OF TICKS.

Banks, NATHAN.
1895.—The Arachnida of Colorado <Ann. N. Y. Acad. Sci., vol. 8, pp. 417-434.

Dermacentor americanus L. and Rhipicephalus sp. are listed.

1899.

Reports upon the insects, spiders, mites, and myriapods collected on the
Commander Islands Expedition (The fur seals and fur-seal islands of
the North Pacific Ocean, pt. 4, pp. 328-351).

Lists Ixodes borealis, supposes J fimbriatus Kramer and Neumann to be the male.
1901.—Acarina in ‘‘Some spiders and other arachnida from southern Arizona”’

<Proc. U. §. Nat. Museum, vol. 23, p. 590.

Mentions Argas sanchezi Dugés from New Mexico and Arizona; also a species

of Ixodes from Arizona.

1902.— Papers from the Hopkins-Stanford Galapagos Expedition, 1898-1899,

vol. 7, Entomological Results (6), The Arachnida, Proc. Wash. Acad.
Sel.) vol4, p: 705 ply 2,ifie: 9:

Argas transversa, n. sp., and mention of Amblyomma pilosum Neum.
1902.—Some Arachnida from New Mexico <Proc. Acad. Nat. Sci. Phila., 1901,
pp. 568-596.

Lists Argas sanchezi Neum., Ixodes diversifossus Neum., Dermacentor reticulatus _

Fab., Margaropus (Boophilus) annulatus.

1904.—The Arachnida of Florida <Proc. Acad. Nat. Sci. Phila., pp. 120-147.
Margaropus (Boophilus) annulatus Say, Ixodes scapularis Say, Dermacentor
variabilis Say, Amblyomma tuberculatum Marx.

1904.—Some Arachnida from California <Proc. Cal. Acad. Sci., vol. 3, ser. 3
no. 13, pp. 331-369, pls. 38-41.

1904.—A treatise on the Acarina or mites <Proc. U. 8. Nat. Museum, vol. 28,
pp: 1-4:

?

Brruta, A.
1895.—Ixodidze nove vel parum cognitze Musei Zoologici Academie Czesareze
Scientiarum Petropolitane < Bull. Acad. Imp. Sci. St. Petersbourg, ser.
5, vol. 2, no. 4, pp. 353-364, pls. 1-2.

Frrcn, Asa.
1872.—Fourteenth report on the noxious, beneficial, and other insects of the
State of New York, pp. 363-373.

Contains descriptions of the following species: Ixodes americanus, quinquestriatus,
robertsoni, cruciarius, Ixodes (?) odontalgiz.
Hassatu, ALBERT. A
1900.—Note on the chicken tick (Argas americanus) U.S. D. A., Bur. A. L., 16th
report, pp. 496-500, figs. 16-22, pl. 16.

Kocu, C. L.
1847.—Uebersicht des Arachnidensystems, vol. IV, pp. 1-136, pls. 1-30.

Marx, GEORGE.

1893.—Note on the classification of the Ixodide <Proec. Ent. Soc.:.Wash., vol. 2,
pp. 232-237.

Contains tables of North American genera.

1893.—On the morphology of the ticks <Proc. Ent. Soc. Wash., vol. 2, pp.
271-287.

1894.—Plate illustrating following species published in connection with obituary,
Proc. Ent. Soc. Wash., vol. 3, pp. 195-201: Rhynchopriwm spinosum,
(=Ornithodoros megnini, nymph), Ornithodoros americanus, (=O. meg-
nimi, adult), Argas americanus.

74 NORTH AMERICAN FEVER TICK AND OTHER SPECIES.

Murray, ANDREW.
1877.—Economic entomology, Aptera, pp. 180-204.
Deals with the Ixodide.
NeEuMANN, L. G.
1896.—Révision de la famille des Ixodidés, I <Mem. Soc. Zool. France, TX,
pp. 144, figs. 1-36.
1897.—Révision de la famille des Ixodidés, Il. Ixodine <Mem. Soc. Zool.
France, vol. 10, pp. 324-420, figs. 1-45.
1899.—Révision de la famille des Ixodidés, III <Mem. Soc. Zool. France, vol.
12, pp. 107-294, 63 figs. in text.
1901.—Révision de la famille des Ixodidés, IV <Mem. Soc. Zool. France, vol.
14. pp. 249-372, 18 figs. in text.
This important monograph of the ticks of the world, unfortunately, is obtained
only with considerable difficulty.
1902.—Notes sur les Ixodidés, I <Arch. Parasit., vol. 6, pp. 109-128, figs. 6.
1904.—Notes sur les Ixodidés, II <Arch. Parasit., vol. 8 (1904), no. 3, pp. 444—
464, figs. 2.
1905.—Notes sur les Ixodidés, III <Arch. Parasit., vol. 9 (1904), no. 2, pp.
225-241.
‘Nurras, G. H. F.
1899.—The réle of insects, arachnids, and myriapods as carriers of human and
animal diseases which are produced by bacteria and animal parasites
“Hyg. Rundschau, vol. 9, no. 5, pp. 209-220; no. 6, pp. 275-289; no. 8,
pp. 393-408; no. 10, pp. 503-520; no. 12, pp. 606-620.

Osporn, H.
1896.—Insects affecting domestic animals < U.S. D. A., Div. Ent., bul. 5, n. s.

Pacxarp, A. S., Jr:
1869.—List of hymenopterous and lepidopterous insects collected by the Smith-
sonian expedition to South America, under Prof. James Orton; appen-
dix to report on Articulates, Ann. Rept. Peabody Academy of Science,
pp. 1-14.

Contains descriptions of following new species of American forms: Lrodes perpunc-
tatus, unipunctata (=Amb!yomma americanum), leporis-palustris (Hemaphysalis),
chordeilis, bovis (= Mzrgaropus annulatus), bibronii, naponensis (S. A.), albipictus
(N. A.), nigrolineatus, cookei.

1873.—Descriptions of new insects. Arachnida <U. 8. Geological Survey of
Mont., Idaho, Wyo., Utah, 1872; pp. 740-741.

Includes Izodes bovis Riley (= Margaropus annulatus) and Argas americanus N.sp.,

with figures of each.

Rinny, ‘©. Vi. i
1893.—Report on a small collection of insects made during the Death Valley
Expedition <U. 8. D. A., Biol. Surv., N. A. Fauna, no. 7, Part II,

p- 202.
Lists seven species of Ixodidee—determined by Doctor Marx.

Satmon, D. E., and Cu. W. STiuzs.
1901.—See above.

Say, THOMAS.
1821.—An account of the arachnides of the United States <Journ. Acad. Nat.
Sci., vol. 2, pp. 59-83; Leconte edition, vol. 2, pp. 9-24.

Descriptions of the following species of Ixodes: annulatus ( Margaropus), errati-
cus, scapularis, orbiculatus (=A mblyomma americanum ?), variabilis (Dermacentor),
fuscous (fuscus), crenatum (=A mblyomma cajennense), punctulatus (= Dermacentor
variabilis ?).

BIBLIOGRAPHY. 15

STOLL, Orro.
1886-1893.—Arachnida Acaridea < Biol. Cent.-Am., pp. 18-24, pls. 12-14.
Five species are discussed.
Warp, Henry B.
1899.—The ticks of Nebraska <Nebraska State Board of Agriculture, annual
report for 1899, pp. 193-205. Reprint: Studies from the Zoological
Laboratory, no. 38.
WHELER, E. G.
1906.—British ticks <Journ. Agr. Sci., vol. 1, pp. 400-429, pls. 5-10.
WituraMs, S. R.
1905.—Anatomy of Boophilus annulatus Say <Proc. Boston Soc. Nat. Hist.,
vol. 32, pp. 313-334, pls. 18-22.

*

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Page.

Amblyomma americanum, bibliographic reference.............-..------------ 67

copulation before gaining host occasional... ...- ~~ -- 12

Ienlowitisy ehavdl linia louistioraiee SAE uy eae SUS a 59-60

trodes orbyculatus a synonymi(?). 4 2/2 ve wean ed 74

UNAPUNCLOLG B SYNONYM. «<i = 2 = ae eo oe 14

outranks Margaropus annulatus as a pest, locally... - 59

egjennense, Nalbtts: lite history; syMomyms . 2... 26 22) c-!) x's Sx=> o/- 60-62

COULD ALOE I— A a TLACUUDOLEU ote ota Men Pe oh ot ere oom ee A 62

Lae] Ovi YS i EG, ie eR ay ba ea eee tat ee eR ae rea le 58

heprecuine -Dipliopraphic reterenee to. ni siets 3) a0) kee ea 70

transmitter of heartwater in cattle, sheep, and goats... . 58

TEL ACILN s MOVES! a YANO TWINS ee i nr Seca ene NN Wace 62-64

ADT Osa eke eS NS IE MART ar ol tee Pe Lae eg) ia eee 7th 50

TI VOCET ATU ———veatte CCL COUILETES CH ers Ty ee te rat Bee Ree er eee (eh 60

muliipunctum, on antelope and tapir-......-...-----.---------- 58

BUGLE Ar TCM LORIE ete eae ype et See ey he es, Mi A ao 62

pilONuima it Galapagos, [slander Sega ee 8. = Mr, else eh 73

RUDE aCe AES TAS CUOUIDGT TI Uy Oe hh I rae Pe a) ae ae x ta ee 62

BRUM OTN Creme URRY ES OFM cl OCR een deo: 9 Sh = aay a A Nea ae = 60

ABOCIes I NOt nnAImenl cata maton oe Ae ee rose tue tus. fed ae 58

BUC CURE erat oo Meher elt ee pete och leprae SoM Le Sal Nee 54, 58

fblevotepecics im United Statest 052.2682 oe Po. See : 58

LETC =— Ale PIRES TTY Peers ee oe ae A ey 2 tee A 62

PUGET — AL TMICITOTLINY <Ses Sie ee PML e 3s 2t ie ee 62

ipiste—— A THOCULALUM, 5 2. hit aoe eect 5 Sea tee ears 62

TUDELCULaTU Dalits amy cistniO Uhl OMmees = See eee eee ye 64

UIST ESOT cen UA ae OA le ie eee neat eo fi:

wnt eCALeL NOStOLPATILON OMNIA, COACMTLENSEs etn eae ee 61
Antelope. (See Dicranocerus furcatus. )

epee ORO pena sit CatL le WhCK. van at as eae SD te 34

Margaropus (Boophilus) microplus .........:----'..:---%- 34

Amite wenemilesion Oniithodoros OWlalOs 2225-2 ae ee See ee ek 37

PaELOUa WOME BOs Ol TEOGES COURCT We /6 ) ose a eh be eee lene 5d

Argas americanus, bibliographic references ............-----.------------: 69, 73, 74

SUN CO Res eS ASL Ie els Li Pa RES, eC Rm ee Regen 42

[NOV ae a Shae eed Sects AS ee ek ty NG BARE ene She ena 42

nis HA eile WistOr ye. SVMOUYIA e222 ft bel ese e nes 42-45

tranemitier of spirilosis,or towls.« 2.222.202. 5 2) s. ie ee 42,71

persicus Dioliopraphie relerenee: say. 5 2. os) See te See eae ten 71

CPL OM UB (52 | THiS tee Se ga RY OR A Ops ation a ae 42

DeATZona and. New Wexleo. 6 20.0. he 22 7k oe 73

Spy Cea Ean GTRS: GIA OT Ke SoMa rl Lee Ge 2 ets pee a ae aco el a aaa 70

fronsversa 10 Galapagos Islands. 2 2.2 2. l oa ol ote tke Ee 73

78 INDEX.

Page
Argasides, habits: 21: A doe ae eee ns Shes ee ere eet Skee ee ee ea 41
table of eenera 2 tet. errs: SONS Ie Ate bine eh a ne ee 40
Armadillidium vulgare, eating eggs of cattle tick.................2.25.-2------ 36
Arpagostoma. ( See Ticks.)
Ass; “host of, Margaropus aunilarus: > 2a. 328 e 22 Sas eek ee eee 35, 39
Ornithodorog mneqnintie: 2. ie PRs A AEE en ee ed 45
Bats hosts sof Wroteswriciiiisss soca wee Ae te Oe be one a ee ey 56
Beaver, ‘host of Dennacerton albipietus.. ke ore hie once oe ae eee ae 52
Birds, enemies of cattle tick.........- Pd oA A) oR NEN C5) Sas nO 8 ences Ee oe 7, 39
ests: Of, Areas: sec tee sw Saas Derk ae ee ee a 42
PANG 212) 16 02 ee Ry a eM OER CT Bh EC Panlg Ne. gwar? EA Se oo 41
TBOMES TICERUAY o.oo tee oan Hoa eae od Coe ER eee ee eee 56
aes, mosis of Geratixades: se) cue el cee oe ee ee cane ee 54
Boophilus annulakis—= Margoropus annulate. 2. 220. 2-22. se Ss ee ee 9
auetralis. in Philippine Walandsss enn 2) ee ee . 69
= MAPPALO DUB Nee eo css eteia cin 2 Manes pe yikes Aes Se te ee aos oe ee 9
‘Bufo agua, host ot Ammbigommacajenmenses 20s 5. 422 22 ss ees ceu ewes ae en 61
Capybara. (See Hydrochxrus capybara.)
Cariacus canadensis, host. of Dermacentor albipictus:......122..2-22.22-..0.2.-2. 51
Cate host OlelnOdes COORCE ATIC) [he ntCUntGs ee nae ate pt etna ee eee 56
EG OOPS TALULVONLRe iys Me rete eee Ae Yee 34, 68
wald.host/ol Dermacentor ibifurcatig. <2 22. a. 5 oh Mae oes oe ose eee 50
TRORCEATCITINIS 2 SANs Clan oie So eek ie ie ae a ee ee ae 56
Gattle hosts of Amblyonimalamericamian 2% 226.2. ee ee ea eee 59
COP ENTLONSC athe a icp ner. Sere ee Oe ear ee eee 61
GOULOLUT ee eas eRe kee a2 SI hye eae 62
Chrysomylanmacelonia. <>... 25242 3028 eee ee 12
PerpiGcenitOn OCCLMCTILOLIS 920.5 Es a EAT a ge el ee 51
MCPLD ULL oo Si et Se = Eee Ont ae ee ee 50
UGOdes TACT nds Ree x oe Mele ee ee Oo a ene te Oe aes ee 56
Ornithadoros gnegiint.. 205058. oe he ee ee eee deat ge ae a 45
infestation by cattle tick affected by ration........- Bic pees erat ee eee 35-36
tick. (See Tick, cattle, and Margaropus annulatus. }
Geratixodes siractune and hostSesa-4- scenes oe eee eee ee eae eee 54
Cercus campestris, host of Amblyomma maculatum.........-..----------------- 63
Chalcidoid, with remains of cattle tick, probable parasite of dipteron.........- 36-37
Chicken. (See also Fowls.)
house, occurrence of Dermacentor parwmapertus.......---------------- 50
Chrysomyia macellaria, attack on cattle following that of tick ........-, Rs ol 12
Deer, host-of Amblyonvma americanum: on nee- oe as Se te et ee a eee 59
Dermacentor nitens o.oo. ee eee ee ee eee ee 52
OCCULCTUECLIS 2 Re eer is ihre, ee pe ea 51
Riodes Sta PUlanisere enc Das oe Bote ea NEE ga eka te eee ee 7
Margaraipus anmiulatis. 3223: Montel oe ee she 2 eng 34, 39
Dermacentor americanum authors, not Linnwus,==D. variabilis ..........-.------ 50
Init COLOLAG ON: cee eie eee cess ae ae ee Pen te eee 73
(veriabilis) Curtice=D..octidentalis’..-.. 2.50.24 ea an ee 51
albipictus, distribution and hoste-. 2 -2c-ee 2 <-see ta ee 51-52
Packard (Am. Nat., and Guide}, D. reticulatus Salmon
and Stiles (ini pant)'a' synonym: : 262 -¢ ace eo eons 50
bifircutus, ow wild=¢at =. 285-5 mete eee nee eee ie eee 50
elecitis==D Varad ilis ee eee Pe eee eee 49, 50

migrotineatus, in United (States? i. s-.. wen = rare aie ete eee eee 50

INDEX. 79

Page.
Dermacentor nitens, distribution and hosts. ..........-.--.------- yee Me eee 52
occidentalis, distribution, hosts, synomyms-..-.-.-.--..-.---------- 51
Neumann, D. reticulatus Salmon and Stiles (in part) a
SHYAMO MN ON Ss eens eRe tare eer ee en ee ee 49-50
INiles—Amiblyommarmaculatuny. = = 2282 255.2 a5 ao = 62
transmitter of Rocky Mountain spotted fever.....---. 72
parumapertus, in a chicken house and on man.......-.-.--------- 50
reticulatus Fabricius, not yet found in United States........-...- 50-51
Salmon and Stiles (in part)=D. albipictus........---.- 50
Salmon and Stiles (in part)=D. occidentalis. ......- 49-50, 51
Species. In United Siabes, SEMUGHITE 25225.) caste a cn ceeek .s0% 20 49-50
Dariausiis: heats amd: Wier WIstonys.. 2k ta3 es geen oO Se 50
ae JE op MGS SS tN he eS eek Re Ae en eas ee oye 73
Sid) oh 00S Slee Sees eee ee eee ee Mate a Ea TACs 50
variegatus, listed from United States by Salmon and Stiles... --.- 49
Dicranocerus furcatus, host of Amblyomma multipunctum ......-.....--------- 58
rnLerou,, probable: parasite ot cattle tick:.2..05. Ss 4.40 $242 5.25,0 425k eee 37
Dinteraus lakves, enemies of Cattle tick « 4.4.02 2454s Joys Joe aek alee eek 36
Disease, human, transmission by ticks, bibliography...-..............-.--.--- 72
possible transmission by Amblyomma cajennense....- - - Poel PA Nis Se 8S 61
DCRR ACETIOT GUDLDUCIUS ss: ah. atk oe Sessa 52
transmission by foreien ticks, bibliography .....5.:-2----.-:--:--2-- 69-72
LRG Sepia 28 Pe eeemyea tN ok So oe fannie A 40
Woe shost ol AmplyommaanvericenuMis= <i ic. 72S. SoM hye hous seek os Aas oF 59
CO TOM TCTs mete eek See Rie BR Ste ee oe eee 61
LST CCRITTIT One ORS MOR Bn Neel Sey Ete SN CE, Le eae 50, 62
Dep QcentonrLONDEONASia Mae amet a beets, tnd td See ee ky es 50
SOG CR GICEICUS rt opt omeue A eae Se at Pl ST pe ete ea Oe 56
CYS TOL) (0) epee eee MGS es sc a OER ARP AY aac OE eae 50, 57, 58
POL POLO PULs LAAT EVEL LUG = as nee meme Pocaieer, at Ae Ty ae rae oat 2 BE 34-35
OnnibhOG OLOSINEGNU ieee Sosa SN a ose ey BR Laee omar c ahead 45
LIPO A EL OTL TIN STIS SOOM Se th Aa eo ie ea a PEP ee ee 47-48, 50
Dove. (See Zenaidura macroura. )
wild turtle, host of Argan sanchect. os. 2222 2432-2 BR Re ei Rte cB 42
“Elk.”’? (See Cariacus canadensis. )
Heed-lot system jor eradicating cattle tick <2 <. 4. 42.4450.5 Shc s0de5-454-: 37-38, 67
Felis pardalis, host of Ixodes ricinus... . - SEC Pe SE eeteps nee ee aT ey aw 56
Herre tel Os itOlel@OUeSuTUCLIVUS = a eeeie er 4 et Ok ae eM) Res eta tasty 56
Fever, African coast, of cattle, transmission by five species of Rhipicephalus.. 48, 70
ie an. sraviSOONSSl ON OY bICKSs oy. yaa ween We J oe ee ds te
in cattle, said to be produced by Amblyomma cajennense...-..-------- 61
Rocky Mountain spotted, transmission by Dermacentor occidentalis - ..-- 12
splenetic. (See Fever, Texas.)
ER EScr 8 LOSS GR i ee eee Senge ak SMB Sad bs Pree BM 8 Sayre ep keIy k eae
may it be transmitted by Ixodes ricinus? .......-----.---+-- ne 57

transmission by cattle tick ( Margaropus annulatus)...- 10, 11-12, 40, 69
“Fire-ant.”’ (See Solenopsis geminata. )
Flea, jigger. (See Sarcopsylla penetrans. )
Flooding. (See also Submergence. )

effectontseedmicks\omeattlertielko: 22) es a os koe 24-25
Haws wenenmlessor cattle tickers ees eerie esheets 37, 39
Fox, host of Izodes cookei and J. ricinus......----.-.c+-c+--2+se+e ae rete te 56

Freezing. (See Temperature.)

80 INDEX.

Page

Goat, host‘of Amblyomma. amerveanium «20 2095: aN ee ee eee 59
Ixodes ‘cooker sae Tricuius 2 ie SIRE Sel eee eae oe 56

Gonixodes rostralis=Hexemaphysalis leporis-palustris.......-.-.-----.--+-+------ 53
ropherus polyphemus, host of Amblyomma tuberculatum.........--.---------- 64
Ormuthodaros turteaia 282 oe Rae eke ee eee 46

sp.; host of Amblyonma tuberculatim. 2c i720 oS eee Se ey Oe 64
Greasing,-method of eradicating esttle ticka!..000 2 s6 2 2 Ae ee eee 67
Guinea pig, not a host’of Margaropus annwlatus...-2.. 22.22.22 02.22 34
Hemaphysatis chordethis,-in United States. ..02 0.2, 222. - 2 22 eo tee -- eee 53
COTLELTUNOMIN SUMNCEd SLATES eee reels eee ee ee 53

leachi, transmitter of malignant jaundice of dogs_............-. 53
leporis-palusiris; habits, -symomym.... 022.0: 52 a: 62 ace oo 53-54

structure and United Statesispeciess: 220.5. 2 - ase ee eee es 52-63
Hand-picking, method of eradicating cattle ticks.............22.-.-22-26e2:- 67
Hares; hosts of Hamaphysalis leporis-patusims 2... 820.2 2222s asc eeeoeee 54
LOCOS RUCERVS (28. 2 E1 acta apes See te eye PE oe ey ey nS bet ee eg 56

Heartwater, in cattle, sheep, and goats, transmission by Amblyomma hebreum. 58
Hedeehoo mhiosholelrodesnicinitises He-Ne ee eee ae ee ee ee 56
Hop host; of Amblyomma aMerichnuent..<e sates ed te eee eee eee 59
OP CUROM ORES UUPIRECO Ce eke iae Ss Se ote heeds etek ee al ae eel ae 46

Horse; host of Ambiyonmaainertcanwn! 2.5.0 2 eh eae ees eh ee le ads 59
COP ETATCTIRCSS SS Ae ere het an ae ea ers ee ieee eee 61
PUDONG LS No eee es yi Sy Cee ee See 62
DD ErMOCENtOTUGEMS asco) 3 Sem ett Pave Seale ante eee ee eet 52
GCCUICNUMIAS OER OR Se FREE rain = iets eee Ce eee 51
Hamaphysalis leporis-palustriss. - 2.2 (0. Doo - 20 noes eee 53
LE OUES FOCUS! Fook an a SN 2 a np Anata eR esate Ce nae IE ae ee 56
Mar garo pues WR RALLOCUR. so NS SY ANN Le NE st cee eee 39, 39
Ormnithodoros Iveco: 2) BIS So Sate rae ee oe ees 45
Hosprclaitonstol HCKS 3550) 0) 6 a a a sec Mee ee oo eee ee 12
TFGUSEH, OCCULFENCE Ol, ATGas s0NUChEZ, ..2 Nose 32 Soe Lote be eee ees 42
Hydrochexrus capybara, host of Amblyomma cajennense..... ....--..---+------- 61
Insecticides. resistance of, Argas miniaius.. 220.) a24.02 == se ees ee eee 45

Ixodes albipictus Packard (Am. Nat. & Guide)=Dermacentor albipictus.
Packard (Ist Peabody Acad. Rept.)=Dermacentor variabilis. - - 50

reierence to original ‘description. !.. 2:22.22. ecko. eee 74
americanus, reference to original description... ........-..----------:- 73
Crug@istiie, 1s WWmibed Iba beS 202 tease ee ee ee 55
annulatus= Margaropus annulatus.........------+--.----++-+++++++---- 74

feierence to\original description (2s. fesse 2. ese eee eee 74
arclicus. an WimitedsStabesac! 2 oc.8 Wyss se ele ne = eo reee eck fee eee 55
bibroeni, reference to original deseription:. <2 222 es) Sc 2 e cece ee 74
bifurcatus—= Dermacentor bifurcatus 2: of hye. sae seis eae eee ee eee 50
borealis, bibliographie reference: 020. 1. S20 ' ee eee > eee oe eee 73
bovis, bibliographic references... 5.22 seein sane = Nee oe eee ree 74

= Mergaropusvaniulatis: "1.05 228) 0 see ae ead eee ee 74
brunneus. in United, Stabes = 22-5 .c.o. memo ee eee ee eee eee es 55
californicus, in’ United Statese.0. o. 1/5 i Eke Seca eee 55
chordetlis; reference to original description..°.....-....05..2 92205. 2ceeee 74
cooker, hosts, Synonyms... 250s ose inet awe kee eee 55-56

reference. to original description: 2. Soe/.ti2s22e ele eee 74
crenatum= Amblyomma cajennense.......---------- ie meyule shoelace ee anemone - 60, 74

reference to original description... 0 Jp he ses. e eed eee 74

INDEX. sl

Page
LANL CONN — PENG REN re te eA) eas NA fa ne Be boob be Bie oe 55
fererenceoreneiaall Gescripulian:s = 2.2) ot 2 as- ed hod se 52 es ie
GG OS. way Uhanietel Sis eo seee oeese ete she Ge eee mee er eens eee 55
RUIBEE SU OSBING IA NC WEN GSI OMe e503 Hac forn oises. aia) = cho ae ba ee ee Sy is 73
erraticus, reference to original description.........-.------:------------ 74
WU LUP GI e —= PrOGes GOMER Ns ANALOG 22 alo SSeS. ee Sco ale ves wend ii
qrontarisein WMUbCG miates ssa S owes 2. Loe a ln 5d
fuscous ( fuscus), reference to original description.............-...-..---- 74
FMS CUS THU MILE Sbabee scatters cee oa a 6 cacy pda eee Me) age 55
PROT TSe one ne er See cea henley ce. eas Lyk Bok ae 55
error? — Ami bly Onvind, AIbeRICOMT oo Si eie << nla saa eats Waa fae siden +s lel 60
PRC TCL COIR) on can Syl OOOO ee en che each See cigs ay Mla toe tata 55
SV RIC MOTUSES DUR OSI — lt NOD KEN. te a5. akin a Sas hh a Bes eS 55
ACNOALUS Til UMIEe GM ohates emer at a eee Sia) ee See 55
leporis-palustris= Hemaphysalis leporis-palustris............--.-------- 74
rererence fo original degcriptiony. .-< <2 5.04 4u- 4 656 - es aes 74
naponensis, reference to original description...............-.-.-.------- 74
nigrolineatus= Dermacentor nigrolineatus.....- 7, NERC Ie ES eee 50, 55
reference to original description. ......-..------ Rata ne “74
(?) odontalgix=, reference to original description .........-....-------- 73
orbicilatus— Amblyomma americanum (7)... ..-2 s22-2---2a.2e216 tee. 74
teference tovoriginal deseription. 3.2.2.2. 4.442 ¢s9se2clayees- 74
perpunctatus, reference to original description....-...--..------------- 74
Pilosis sbiplioptapute TekereO ee 2 secs aos lar eo Saki e een ok 71
drying up of engorged females in captivity before ovipositing. -- 55
PlamnpeUus: (1) AOREeVNbY OManVee 5 boot ae face Sees seed Lacan 22's late 55
DUTCH ALNS — DETMALERTOR UORAUULES (f) 8c aoa eis ae obese 2 =e 50, 74
felekence te original deseripmMon.-. oe... 2s oes ee 74
quinquestriatus=Dermacentor variabilis ....2..-.-:.-------.--2-2--++-- 50
reference to original description ...........--------.---- 73
PCIe INIA ——RTTCUITURS arin te 5 ohn Be het Ae Bae ly th We ca By 5 E'S a 8 70
nmeimus.. bipliosraphic references 4. - 2 ac ches anata 22 se 2 <3 spe 67, 70, 72
connection with ‘‘louping ill” of sheep only accidental... ..--- 56
esp hs cone ne MIRLORY +. faces sets Oe es alee Sia eagle 56-57
InjSantaeruzeMountaims. Caliiormiavssss: -- sso see ee ee eae 58
may it transmit Texas fever in United States ?...............-. 57
OWEPOSIGION: process a6. i-th asd ee Sass a eke 2 als bia ale 16
transmitter of European piroplasmosis of cattle.......-......---- 57, 68
Ly E Ole Re US ie Se a Se Shia ee ye es et ES as sic oh ks ae 54
Raventsani——) erinigcentonsvOnvauvaas ao Met Ne aaah sk 8. SE ee eee sss 50
fererencesto oricinal desceriptlomec. cs a4 S262. ek 73
scapularis, drying up of engorged females in captivity before ovipositing- 55
TNE SG BSE ROPE See Se ee et Rey en Sar 57-58
Tan) BUST GA AEE a. APSF aR. oi) 2 ae pan SNe A Coie EN ea geRgP Rare 73
SIMCOE erent weet ANON EN ENS o> MO se Lay 50, 57, 58
Fererenes LoO.orie inal desempiom ss 45.05 Ji y.ess. se. Ss. S 74
Sch pints. cistribubionvand NOSh a. eoies 2 Feet eos so 8s. 2-2 2S e = 4 OS
Be ci ce pinyin’ Crabens a2 abs teens hates ci sey Aaa oat sas Sica 55
ERO en RGD LeN tein Ae ary ee aE I SES Se ey a2) na iS ols IE 54a eee 7
SETUP LOMA EMNIS= tome terete OS PO 8k Jag sae abe bee 54
UNUM CHLA— AMD YONA CIMEPLCAMUM a. or ea ae «= 2 2 oe ge ee 74
reference to original descriptions. 9..\°,..-------+-2+222)--- 74

5795—No. 72—07——6

82 INDEX.

Page.

Teodes urtz, in. United States? see ng ee 55
variabilhs—Dermacenter. varnidonise 54s: ss2. eet ee ee ee ee ee 74
reference ‘to onipinal descriptions 22.22.25. 2 hes saat see oe eee 74

Ixodidie, subfamilies and: genera:= 2.3 22j04382 sch Po a ae ees 46
tablejotisubiamilitesiandiceneraa es ae ee ee ee 40

txodinte; structural (characterand generavs:c.. ao eee eee 46
structure of generar. sleet Seo cig seek BEE ene bo ee 4 A 54

table of genera ss? ds ae hse as AA eee eee a eer et ce eee as 4]

Ixodoidea. (See Ticks.)

“Jackdaw.’’ (See Quiscalus major macrourus.)

Jaundice, malignant, of dogs, transmission by Hainiaphysalis leachi..........-- 53
Jigger flea. (See Sarcopsylla penetrans.)

Kingbird. (See Tyrannus tyrannus.)

Leech, use of Amblyomma maculatum by South American Indians.......-...- 63
hizards. hosts ofinodesiriemue. Vee fas Se Ps Se See a ee ee 56
Wamashosto Onnithodonosi Cunvcaui= sence haere ree saan rae eee eet 46
Hepus'callotis, host of Dermacentor vaynaviiga. . Seo en Re ees ee 50
palustris, host of Hemaphysalis leporis-palustris.........-..------+------ 53
syluatious, host of Srodes rics? 62. 2 hse oe 8 ee BT'S stony FA 2 ky tegs 56
honrevity of Argasminidtuss. 2. /i28e Nese se Phe ee EE Se ae Sa eae see 44-45
Tod sete fe aN he Os a SR ae ee ee 55
Ornithodoros megniny 2220 2 eee eae eee eee ee oe 46
“TLouping ill,’ of sheep, connection of [xodes ricinus only accidental .......-.- 59
Mammals, hosts of Amblyomma americanum. .....---------------------+--++-- 59
Ave Gp. OS Sse tase Se OR AE Sot BEE SSIS 2 Biyori eter 42
Arpasidee: 5229 nse a5kjr tet sa Rees Atle cae eas ee ene 41
Mak: hhost:of Amblyonina amencanmim:. =. 53.2 -)2.52 22 sae eee Beet ane 59
COJeNTIENBES 18 25. HSPs 32 eats SNe ee ieee eel nie ear 61
Maculatumiss ele 5292 sss Aen ee Sees ae ae ee eee 62
ic Se Ae ne rae et AIS foeig eS oth Rhea nae ed er Sr oh on 42
Dermacentor occidentalis: 2 pu.+ tone te sn 22S oe a tee 51
PONUMULPET UB: «2. ial, SAN ee tk ee eee nee ce owe 50
LLOCS TLCS se ee Ae De ae Ee hs Bae to Rie ene Nee ee 56
Margaropus onnwlatius® 22225. 522 c8 eee ee eee eee oar 35
Ornithodoros megnint. 228.222 Sec che TE ae Be ee eee 45
EOPICGLON SIDS Se et a Le as dole ae 46
Margaropus annulatus. (See also Tick, cattle.)

hg bitat= 223 ofl hades aks See eae gst aoe Reo ee eee 49
hostirelationss.. ¥+<sssnh sees nae oe oe nee eine 34-35, 39
in Florida. <0. t22sca2 ee 2 se) Pe ee ae ee 73
New Mexi®0: i sj2222s¢ eae hs eee cee eee 73
Erodes bovis & SyHotiyne 165222 - 2d. bteeek at FOB 74

outranked as a pest, locally, by abet amert-
CONMUMs 225 528 a LES SE Re ae aS oe ree eer eae re 59

placed in genus Rhipicephalus by Netimann ae Ful-
lene. js safe cub Se I EIS, SE Se AE Speen 47
transmitter of splenetic or Texas fever.........------- 40, 69
ver: argentina, habitat. 052 22. Acidk.. Alsen 2 Sees 49
australis; habitatsss0-<% s2cncs = saee See eerie eee 49
M. microplus probably asynonym. ....--. 49
calcerata; habitat? <5.) ce ch oe ee. Se eee 49

éaudata; Habitaby)\:.2 due ees eee 49

INDEX. 83

Page.
Margaropus annulatus, var. decoloratus, etc., studies by Lounsbury.......-..-- 10
lOMOW oceecooe Seb ere ioe a2 Lee area 49
VigeE) ay US Um gee 2 lee ae aa 15-16
correct generic name for cattle tick (Boophilus annulatus) ......-- 9
habits and structure distinguishing it from Rhipicephalus......--- 47, 49
) MiPOplys, AULIBEMOMOmIs Ne wees sheet se Se lo ue Sek... 34
bibliographic wrelerences-2s222 .. Sis. ee 70
SemMig Aamir lese cea Ma he tes sce SEINE Lh 14
Loc OnMMsensererse s Nareiae <2 xt A ac) el Se Rey 34°
probably= M. annulatus var. australis.............--.-- 49
FEAAGHIM en TOROS eset aye eit) ie 8 Beats) has sre A 29
Marmot. (See also Arctomys monaz.)
OST AO el Et6' ARC TIN 77 SEES eee TAY Des ile ae MO ne Pe ce a 56
Miceshostsiot Uodesmciiiuss] 5. seen see ee eee Ree Bee at eke a eee eee a 56
jLOVAPIG Chemies OCAthle HemKes. Ami sase lick S Socee ll Paella 37
Mink. (See also Putorius vison.)

GSE OD LON cS TCC Ma air rs oa enn Seis op bia ore SE ba SRA Sk 56
“*Mostacilla,’’ Guatemalan name for young of Amblyomma cajennense........-- 61
Maite: ost ot Amblyomma americanum 2-8 es 3 YS 98 Sees 2 3s oe be 59

ce ICH y BRED Ei uated rete ae OS Seb held 22 eek 56
ORDER AN A IMRIGID = tee A cake wow sk te ee oe SUS PLL Bae 39, 39
ier eeluas) SOMO, PS ESI ETA eas es) ee ey ew os ers oes EMIS Sal Sik 56
Onnithodorosiamenicanius—O--megnini Aguile. ee se2 eee ee eee eee 73
Waites e bc My MO AL) ies ML has steer Dae bt Beis I Ae 45
AGW Lo AE) ONS eee et A Bee aes neti ee chyna es eS teh ale nae 45-46

Ornithodoros americanus and Rhynchoprium spinosum
SY MONMMIS san sae REN sys eee Se eee ee SM SE SS kB 7/2
moet, bipliceraphicrelerence. +2.2% 2529s. 22-2 ee es Pea. sce oe 72
PP IMNEH 1) Hie Te ee ee SOR ead SS Be he he 37
transmitter of human tick-fever.........-.......---.-.- 45
TOE ALERTED | SS fe ae OL RIAU AAI A aR oa AN Sa 45
is.if identical -qvith: Oy ¢uracala? so. e242 soe. Ls Se 46
BOD ewe cia Ol! PILES. WS. 4 24,/ seis pemege Mason weet 6 oa SS 2 BY Le 70
sp., suspected transmitter of cattle disease.............-........ 45
OUAYOTIA TH el ON OTN s\n DON pea EE Ge ES Bee ak og! oe ee 46
isaividentical with. O: samgnen? ss 522 oe SS 46
Ge tereh ost Ole 700 Cs) COOKE as ar oe etces ose SS br a ee a 56
Berio et lum roccos IM (CKae 2. sun e se ot ae pae Ses SPE tee Set aE 16-17
Airs Osi OleebOdEs TICHIUStec re mee cn cone <a Re Es See eA eee ide: 56
Paradoxurus, host of Hxmaphysalis leporis-palustris.............--.---------- 54
Pasture-rotation system, for eradicating cattle tick.........................-. 38, 67
Beccary, font of Amblyomma cajenmense....:-...-..2228) 2222S. vies oe 61
Pionds:4 species bred from cattle tieke 23520-0212 k Oe eee 36

Picketing, method of eradicating cattle ticks. (See Tie-rope. )

Bae IED WAI SEK OLY NEAR fe te te 2. ear ee ee Ma eR shel. a pb oe 42
Piroplasma bigeminum, found in both Europe and America.......-.-.-.------- 57
Piroplasmosis, European, of cattle, transmission by [vodes ricinus............--- 7, 68
RalecaisuhosisoleZOucsmeCiniisie- my eee AO eee ae NS eG lly ye 56
MCC OMINE erin OE OOF OORLG Jel wou) eaten eet Ae. leak ky ROE 56
record as host of cattle tick probably erroneous........-.......--- 39
Praerical app iicavoniaiMiORMallute — 22%) o6 oe. 6. we ee ww 37-39
Prairie dogs, hosts of Irodes scapularis............-- SES ae LAS eRe See 58

84 INDEX.

Page
‘* Prootoquie,’’ term used by Lahille for period, in tick, previous to oviposition. - 14
PALCONULS, UUSOT™ DOSULOleLDOUESECOOREI= een natn eee eee 56
Quail; host’of Argas sanchesi. 2S-S 2s. lake 5 Pee eee oe ee ee 42
Qiuscalus major macrourus, ‘enemy ol cattle ticles 4. - scene nee ee eee ee ee 37
Rabbit. (See also Lepus callotis, L. palustris, and L. sylvaticus.) :
host of Hamaphysalis leporis-palusiriss...- 2.2.2: 0s bane tp e200 2- 22 53-54
Tpodes tennis) ae ie Sere ee ee Vel ayes ck toa 56
Margaropis-aniilotus. 0.) Oe See ee sr ea a lee 34
record as host of cattle tick probably erroneous..-................-. 35
IRACCOOT MOS isOLLLBOMES COO CII ne coos Se TT ere Teen nD ee re pe nee 56
Rats senenilestomOnnittnOdOnOs WiLOLLU CLG eae) ayer ee ee 37
Rhipicephaline, structural character and genera...........-.--------------- 46
tableoneen eras: Sn TO ee eee ye ese icreaear ore, Nee cgay: emer ae 40-41
Rhipicephalus appendiculatus, bibliographic reference..............---------- 71
bursa americanus, reported from Jamaica..............-.---.-.- 47
capensis, bibliographic TeterencGe sn. = 4b eeee eee ee nee val
decoloratus, transmitter of spirillosis in cattle.................-- 72
evérisz, bibliographic relerence st Oy 2) kG MOL) ety oe 71
habits and structure distinguishing it from Margaropus......-..--- 47, 49
sanguineus (Ann. Rpt. Bur. Animal Industry f. 1905, p. 35)=
Rhipicephalus sp. of present bulletin. .....-.-...--- 48
Ln. /Panamauy } oc heeds Leal ae tS eee oe 47
SIMS oilbluocra phic relerenCessasas sae a ee ee 71
Ape, wabite and lifevhistory.-s45.- 0." > os. eee a2 ae 47-49
mt Colorado.) da Fee Soe sels eee Oe eS ees ene 73
DY LO Gene AEH eV me re GE a 2k ea 50
same as R. sanguineus in Ann. Rpt. Bur. Animal Industry f.
LOOD Spt Sb. 2s ss Sie ee eR he eee aca 48
undetermined species from Colorado and Porto Rico.....--- 47
Rhynchoprium spinosum=Ornithodoros megnini, nymph.....-....-...-.------- 46, 73
ROE OILGKwhOst Ole! LOGeSChUCHILUS == a" rae st en Se eee eee eee 56
Sarcopsylla penetrans, introduction into South Africa..................---.---- 46
Screw-worm fly. (See Chrysomyia macellaria. )
Sheep; host ot Dermacentor, ocedentalhissst >. 22/e 22020 2h. ls8- ee ee ee Dill
Trodesicookenandelam cuutisee ser se see eee ee eee eee 56
Margaropus ann ulatusods ta. Shee et one ae er 35, 59
Ornithodorastmegnint B22 hace. ds Aaa ee A ee 45
Slsimik: host:of Trodess cooket ss 2.0/3 ae eo sect oe Oe he ees 56
NTA Ke MORNIN OMOTOSUL MICO LGENO! D LIETO Wine ceee ee ae eee eae eee 46
Soiling system, for eradicating cattle tick. (See Feed-lot system. )
Solenopsis geminata, enemy of cattle tick......-.-.... a een SS, 36
Spermophile, host of Yeodes: cookers. 2/2 ee Se aa ee ee 56
Spirillosis of fowls, transmission by Argas mintatus.............---------------- 42
Spirillum (Spirachexte) obermeiert, probable cause of human tick fever ....-.--- 45
Splenetic fever. (See Fever, Texas.)
Squirrel, host of Amblyommaiamertcanum. . 2222. .5-- 2. 79292. 2 2S eee eee 59
Trodés Menus: a2. 225-3258 AO ok Bee ee ee 56
stag, host of Ivodes ricinus. 5222/2 0..2 Shae eae ee ee ee ee 56
Steer, used in experiments onscattlerticke 22-32 ase) ae et ete 27
Submergence, in water, effect on engorged adult cattle ticks...........-.--- 32-33, 39
IMeUbaAtiOniOl Cattley tC kee eee ee 22, 39
Sunlicht, direct, ettect on adult (cattleiticks ssa us ee ee tee eee 32

Tapir, host: of Amblyomma muliipunciin ss. <2) 5-2 ee aa ee ee ees 58

INDEX. 85

Page.
Temperature, effect on engorged females of cattle tick..............-.------- 31-32
Reetsulena Of Cattle iCK. saree) 225.2 Se ee 24-25
inrelationiia iiteubearion ol eathle tieks 2:22.22: 22 2s--. e255 2 19-21
Texas fever. (See Fever, Texas.)
Tick, adobe. (See Argas sanchezt.)
Australian. (See Boophilus australis.)
black-legged. (See Ixodes scapularis.)
bige.common,, bibliosraphie reterence =. 2042202. So e2 oh eee ee 8k 69
bont. (See Amblyomma hebrxum.)
castor bean, American. (See [vodes cooket.)
European. (See Ixodes ricinus.)
cattle. (See also Margaropus annulatus.)
adnlttenmale deseripmonsie suse ep tei. 2s oe ores. 3t tes 31
Tito eRe PlONe Snes sae loss os ol MESS A eS 30
BIR e eth ol od eke Ck oS eRe Bone aoe eee nae eee esek 30-34
Sadia as iected by: direct sumlight!- 2700022 2.e etek. . jokes 32
submergence in water.............----- 32-33, 39
attachments of specimens to each other...............-.------- 35
pills prep yee eee cee ee ons ose te ee Se aS eee 65-69
control. (See eradication methods.)
Geseri Wings OAUWMe se Mee tte Mere BND Pl Se Loe 30-31
development as affected by ration of host................-.-- ..- 35-36
ONUDOSLS tess See eet wen eee ee eee te rs 28-30
Crop pig InompHOnt esau. teen ots tee Sone, ce ad] ee te 33
COURS TAO Ch senate a ael tat Perr ter Oetstee a ere ert AG soe ac ae ara ee ee 15-23
eesnas atectedipy heat and coldie. osetia h Sees dels 21-22
BubmMercenceiniwatere n= ssn sss =e eee 22, 39
InNdICAtloOnvOmwwablithvessee sas meee eee = oe ere 15-16
Miumben deposited!: 2 xo2-ce5-26 soe sae. laste tee fake ons se 17
metcentacephate him pee. eee see ee ae gas at oy ec ane 23
engorged adults as affected by submergence in water... ----- 32-33, 39
continuous cold and heat... ...-- 31-32
(GU SGISIAS 5 ok Se OR Rae Sel el ioe eR EE 36-37, 39
eradication, from United States possible................:.-.--- 9
HUAWEI CROCUS ee ere Mena ets 3 en es rat ee yc SRR eel 37-39
MOO STE TONY OLA MF MCKCO Ce Senin see ee a eres ieee 13
HOStETelantOnis ter cece tes ieee re ee I PR are ESE 34-35, 39
importance of knowledge of life history................-.-----.- 9-10
PCM DLL OME Tate Taeet ove sia EO PRE ee OES J et IO, 17-23
CROC ace meson Pee sere me mses Mee WAM eee oe es 17-21
infestation affected by ration of host..............------------- 35-36
knowledge of variations of periods in life history essential... ..--- 13
lack in knowledge of local climatic variations and dissemination . 10
LLG SLAC ee aaa enon ee ead eh Fe to Ios Nae soe ses asieaiae te 23-30
nonparasiirerpeciod.: 2 Sie Ile Sls s eh ee 23-26
ParAMtOLGL PRETIOU cere! oy Are ck ays Sb ee Meee 27-30
LITE LST RGUAE Rls Eek Ae reeks ee Si age ee Oe AED 8G 13-33
POCORMO TOME Hes ee eee eet Ft ern SS. yy te Tete 33-34, 39
lone vinyeOlnOCd MAGA mmr as! Se Se oe SUL et Bee 25-26
Lens et eee ae SUS 2s eM EY Degree a DE ie 11-12
male, position on host with relation to female..............-.-- 30
mOlisvOlplarvalystaresas 7215 sh) ss 5 va Sass ols ae es 28-29

most important factor from standpoint of control.............--- 13

86

Tick,

Ticks,

INDEX.
Page.
cattle, nymphalistage': 9... 2: [Siete as Seer Sa oye ee ae eee 28-29
OVEPOSIELONGMENION A. os bs rere Pie es he ee oP ae 14-15
PFOCERS ssc ctv) salt es Eke 5 theater sie eh me iapberee: Raaiete reeet ne 16-17
periods in life history upon which means of control are based. . 38
practical application of data(-2 20 eee ie 37-39 ©
preoviposition period... ist eo soc 2 en o ke eee 14
previous work onlite: Ristoryie: 57.6. Seas ae See eee 10
reattachment Nost=s: +. seg. 0520 hc eth peek ee Ue Nae eee 29-30
seed ticks as affected by submergence in water.........-..-- 24-25, 39
ION BEVILY <2 occ. ce totes SO OAU te wero teea ae eee 25-26
tuickestage tse sae yes eee a eae ie A dt 23-30
nonparaditie period -22)2g-2..08-</44-nees oe 23-26
PATASLPLC PETION! =. [eines as hee ce es ee ee 27-30
submergenceras alleching eprstas sees nee ee ee 22, 39
engorged adults............:.-...: 82-383,39
sunlich tas eitec tin os cilities te ees soe ae ee ore ne By
temperature as affecting engorged females................----- 31-32
Incubation ges es a ea oe be ee 19-21
Seearticls: Niet Sie Sele se eee 24-25
inca shoounnereront “Memes ere, 6 9 oe ea oma akooseneslcckesse 10.1112
Cayenne. (See Amblyomma cajennense.)
dog, American. (See Dermacentor variabilis. )
brown. (See Rhipicephalus sp.)
elk. (See Dermacentor albipictus.)
fever, human, /biblicpraphic reference:<2- 2-2-2255. 2. eee eee 12
transmission by Ornithodoros moubata.......-.-.---+---+-+. 45

North American. (See Margaropus annulatus and Tick, cattle.)
fowl. (See also Argas miniatus.)
bibliographicsreterences: se. jo" een nee es eee. eee 70
Gulf coast. (See Amblyomma maculatum. )
horse, tropical. (See Dermacentor nitens. )
land turtle. (See Amblyomma tuberculatum. )
lone star. (See Amblyomma americanum.)
net. (See Dermacentor occidentalis. )
paralysis. (See. Ixodes pilosus.)
rabbit. (See Hemaphysalis leporis-palustris. )
sculptured. (See Lvodes sculptus.)
spinose ear. (See Ornithodoros megnini.)
turicata. (See Ornithodoros turicata.)
wood. (See Dermacentor variabilis.)

nibligoraphiv 22. As oo oe Sete skeet ae ieee ee eee eee 65-75
castor bean. (See Ixodes.)
classification and distribution, bibliography.........-.-.-.----.-------- 73-75
Rabitge oye oe See ee ee ee 40-64
foreign, that transmit disease, bibliography............-.------------- 69-72
graphic table for separation of families and genera, from Lahille.......- 41
Hatta tes oe ol. te pies ane ORE In hee 3k pee 40-64
ost relations: s..... 4. -...2. weth, Wes emereet ee ence et ge eee eee 12
key to families, subfamilies, and North American genera.......-..----- 40-41
fife history, general statement). 28340 222 eee eee = ol eee 12-18
transmitters:of diseases) 5.5) ee Ae ee ee ee 40, 65-72
work by Labille:. 222% 3 .ces de> 6 oe = so ee i eee ee fe 2 eva 10

‘“‘vearling,’’ name given to ticks in nymphal stage.........-......-- Syeiae 13

INDEX. 87

Page
aaeerape, metho or eradicating cattle-ticks 252-6. 2... asic sec nceecce oS ais 5 67
Toad. (See Bufo aqua.)
Turtle, land. (See Gopherus polyphemus.)
ivwo-teld method of eradicating cattle ticks. :22221..--2-...5-02 55.2 be.ce bees 67
Wapiti. (See Cariacus canadensis. ) :
Wiican menos th OlesGOdLSuCOOKEY ian eee tee eres eek eine nis ye SS hee eles 56
Molt, host ol Anbijomma amercanunt. 22602-0222 2222 2 2 2 ode ee 59
Woodchuck. (See Marmot and Arctomys monaz.)
Xerobates polyphemus. (See Gopherus polyphemus. )
‘Yearling ticks,’’ name given to ticks in nymphal stage.................-..-.- 13
ema MUCrOlnd, ost OL “ATGas SONChEr. 6.222. -eotoce ke ete 2 ot ees bbe lee 42

O

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U.S. DEPARTMENT OF AGRICULTURE,

EP OCHOW ARD: Entuncloget and Chich of Bureau,

BY

i
a W. DWIGHT PIERCE,
: Special Field Agent.
4 ,
me Issuep JANUARY, 21, 1908.
‘ .

i} K
ih pea
q
as WASHINGTON:

GOVERNMENT PRINTING “OFFICE,

1908:

BUREAU OF ENTOMOLOGY—BULLETIN No. 73.

STUDIES OF PARASITES OF THE
COTTON BOLL WEEVIL.

NEB UREA U OF ENTOMOLOG y

A O. Howarp, Entomologist and Chieg of Birra ,
tes La ee ea Entomologist and Acting Chief in absence oF Chief.
Re Tipe TRS. CLIFTON, Chief Clerk.

. CHITTENDEN, in charge of breeding experiments. 9 7.
. Hopxins, in charge of forest insect investigations.
* M. WEBSTER, in charge of cereal and forage-plant insect ‘investigations;

shady

i QuAINTANCE, in charge of deciduous fruit insect Pe ha pete

. Rogers, in charge of gipsy moth and brown-tail moth work,
. Me RRILL, engaged in white fly investigations. Wis
( W. * Be: Fisker, in charge of gipsy moth laboratory, =
Ww. ’. A. Hooxsr, engaged in cattle tick life history investigations. :

ORGAN, engaged in tobacco insect investigations.

af
n

: . Woanum, engaged 1 m hydrocyanic acid gas” investigations.

Corron Bou WEEVIL IsvEstiamions.

W. D. Howren, im bt

: _Srooxr, W. W. Yorines, aacatae field dcths.
‘ ~ \ ‘ ,

“a

} i
Wa 3h |) ) ii MIAN OITA i)

U.S. DEPARTMENT OF AGRICULTURE,

BUREAU OF ENTOMOLOGY—BULLETIN No. 73.
L. O. HOWARD, Entomologist and Chief of Bureau.

STUDIES OF PARASITES OF THE
COTTON BOLL WEEVIL.

BY

W. DWIGHT PIERCE,
Special Field Agent.

IssuED JANUARY 21, 1908.

ANN

ZA

Fel AN NYS
e/g AN)
w/a) H.

a Las

\ A$

Canaan
NY Z2 a

WASHINGTON:
GOVERNMENT PRINTING OFFICE.
1908.

LETTER OF TRANSMITTAL.

U.S. DEPARTMENT OF AGRICULTURE,
BurEAU OF ENTOMOLOGY,
Washington, D. C., July 29, 1907.

Str: I have the honor to transmit herewith a manuscript entitled
“Studies of Parasites of the Cotton Boll Weevil,” by Mr. W. Dwight
Pierce, a special field agent of this Bureau engaged in cotton boll
weevil investigations. Some of the preliminary results of these
studies were published as Bulletin No. 63, Part II, of the Bureau of
Entomology, under the title ‘‘ Notes on the Biology of Certain Weevils
Related to the Cotton Boll Weevil.”” The boll weevil in the United
States is now known to be attacked by several species of parasitic
insects. These have not been introduced with the boll weevil from
the country from which the latter spread, but they have been here
for years as parasites of other species of weevils occurring in the
territory now infested by the boll weevil. The present manuscript
contains a record of investigations of these parasites and deals
especially with the possible practical applications which may be
made of the results gained. I recommend its publication as Bulletin
No. 73 of this Bureau.

Respectfully, L. O. Howarp,
Entomologist and Chief of Bureau.
Hon. JAMES WILSON,
Secretary of Agriculture.

9

“

PREEACE,

The boll weevil has been present in the United States for fourteen
years. At this time it is doing serious damage to cotton in four
States, and during the present season undoubtedly will invade a fifth.
In regions like Central America, where the insect has existed for
many years, there are various influences that serve to hold it in
check. In this country similar influences are beginning to be in
evidence. Among the direct factors in the natural control of the
boll weevil that are now at work are the following: Heat and dry-
ness, the native ant Solenopsis geminata,* winter temperatures,
proliferation, parasites, defoliation by the cotton leaf-worm, deter-
minate growth, and birds. Of these, proliferation and birds have
already received careful special attention, while the work of the
native ant has received preliminary notice. The other factors have
also been more or less discussed in the publications of the Bureau
of Entomology and elsewhere, and more extensive studies of certain
ones will be discussed by Dr. W. E. Hinds in Bulletin No. 74 of this
Bureau.

While the work of parasites, discussed in this bulletin, is not at
present one of the most important factors in the control of the boll
weevil, the indications are clear that its importance will grow rapidly.
Moreover, the matter has special interest for the reason that it is
not unlikely that practical means may be devised to increase the work
of the parasites.

In the United States fifteen species of insects which seek the weevil
in its immature stages are known. These insects have not been
introduced artificially from other countries, nor have they followed
the boll weevil into this country. They have all been present for
years, and have been actively parasitic upon other species of weevils
found within the boll-weevil-infested territory. In fact, there are
a great many closely related weevils of the genus Anthonomus, some
of which are very common in parts of the cotton belt. These weevils
are frequently held completely in control by parasites, and for the
most part the parasites are not confined to the attack of a single
species of weevil. In many cases the boll weevil may appear in a

@Not to be confused with the kelep, the introduction of which has proved unsuc-
cessful.
3

4 PARASITES OF THE COTTON BOLL WEEVIL.

region in which the parasites have reached the critical point of
maximum: parasitism, and in such cases the overflow of parasites
must attack some other host, which naturally will be the predominant
insect of the same group as their former host. It is therefore not
at all surprising that when the boll weevil appears by millions in a
new region these same parasites in the very first generation of the
boll weevil become adapted to it as a new host.

The distribution of the other species of weevils is limited to floral
or geographical regions, and consequently, to a less extent, the para-
sites are also limited. This condition is illustrated by the distribu-
tion of the species of parasites which have already attacked the boll
weevil. It is safe to say that as the pest advances to the East and
North still other insects will adopt it as their host in the same meas-
ure as it invades regions inhabited by other species of weevils. In
many transitional regions there is a strong possibility that several
species of weevils may be very highly parasitized and that all of them
will lend parasites to the attack of the boll weevil. In such regions
especially there are to be expected instances of very high percentages
of parasitism. At Waco, Tex., where two regions border upon each
other, a parasitism of over 40 per cent occurred at one time during
the season of 1906. :

There are two possible practical applications of the information
obtained and recorded in this bulletin, both, however, requiring ex-
pert entomological knowledge and experience. These are: (1) The
propagation and collection of parasites, and their distribution in re-
gions where the same species are either present in but small numbers
or altogether absent, and (2) the elimination of related weevils by the
destruction of their food plants in or about cotton fields, thereby
forcing the parasites to transfer their attention to the boll weevil.

Under the plan of artificial propagation by picking large numbers
of squares in fields where the weevil is highly parasitized and placing
them in cages adapted for such breeding, a large number of parasites
might be obtained. The weevils should then be killed. These para-
sites could be released in fields with a low percentage of parasit-
ism, and the results under favorable conditions should be apparent
after one generation of the boll weevil. In this bulletin will be found
an account of an apparently very successful experiment of this kind,
in which, at Dallas, Tex., the percentage of parasitism was brought
up 9.1 per cent by the introductions of parasites from Waco, about
100 miles away.

Under the plan of eliminating, more or less, the natural hosts of
parasites that will attack the boll weevil (by the destruction of their
food plants) no experiments have yet been performed. As an exam-
ple of the possibilities, however, it may be mentioned that there is a
common but easily controlled weed (Croton spp.) grown in and about

PREFACE. 5

cotton fields throughout the South the seed pods of which are infested
by a weevil (Anthonomus albopilosus) closely allied to the boll weevil.
This weevil has three different parasites, which also attack the boll
weevil. The croton weevil does not feed upon cotton or any other
cultivated plants. By merely mowing down or otherwise removing
the croton plants at the proper time there could be no danger from
the croton weevil, while its parasites would be forced to turn their
attention to the boll weevil. It is conceivable that the encourage-
ment of the croton plants or the actual planting and later removal
might be undertaken in order to obtain the best possible results.
Other results of practical application are recorded in this bulletin.
Among them may be mentioned the fact that fallen forms exposed
to the sun show higher parasitism than those shaded, due undoubt-
edly to the light-loving character of the parasites. This gives another
reason for the wide spacing of the plants and the use of varieties of
cotton with the minimum tendency to form leafage and the greatest
tendency to shed their leaves in the fall.
W. D. Hunter,
In Charge of Cotton Boll Weevil Investigations.

CONAN ES:

Rhicmmonicran jarasitesiim: W906). 28.2) ee es Laci ee aise ececccs 2 veetake
lee MAMMA GOM WORK seat ene eke es tes eS LE Thue Be)
Records prior rocbo0 bees: see: Shae Bo ek ol ee ek
Breedimeazeconds ol 190G.)-20 2 es ssacsee coe se. Vee ee es hess

Most favorable plant conditions for parasitism of the boll weevil. .

Biel LCOndiitonsen re ener ae Serie. Se id SAN Bo
Geographical conciderabions-¢21 4 sss sesh)
OMaweevdeotaiUsa: seers soo spleen. EY. DARN nT ks ts

Boll weevilechronglopyt esse. ens Pt ee
Womehusionssserssas ict eee te en A oT ee Aap

Riese roparatlomworkey.< ses. anaes e ose Seed ORE
1. Interior work—transfer or artificial propagation of parasites... . . -

2. Field work—release of parasites...........- UR IN heal)
iiabearssite breed ae: works ssc. sesh. te sere peels Se TS) Behe
MC CUrLeN Ce’ OR SPECIES. Nasa Soe ee eo. ah ee Ne
Geographical and seasonal distribution of parasites...............-

Parasites known to attack Rhynchophora......................---
Biologies of the weevils contributing parasites..................----
OLALIONUOeMOSERO rose oc aks eerste ees PN Se ooh as ok ee ee

PULUS TREES:

PLATES.

Puate I. Parasites of weevils. Fig. 1—Eurytoma tylodermatis, pupa. Fig.
—Catolaccus incertus, pupa. Fig. 3.—Cerambycobius cyaniceps,

pupa. Fig. 4.— Microdontomerus anthonomi, pupa. Fig. 5.—Larva

of Bracon. Fig. 6.—Bracon mellitor, pupa. Fig. 7.—Larva of chal-

CLOGIG 58 kee S02 Sec Seed She ee eet on he wee a eee er

II. The purple mallow (Callirrhoe involucrata), a food plant of two wee-
vils related to the boll weevil. Fig. A.—The buds and flowers, which

are subject toattack by Anthonomus fulvus, and the capsule, subject

to attack of Macrorhoptus estriatus. Fig. B.—Bud infested by
Anthonomus fulvus. Fig. C.—Flower injured by adult weevil

Of Anthonomius fultus occa. oft ake eee toe eee ee

III. Biologies of weevils. Fig. A.—Pepper, showing egg punctures of
Anthonomus eugenii. Fig. B.—Head of Sideranthus rubiginosus,
showing cell of Desmoris scapalis. Fig. C.—Pepper, showing larva

of Anthonomus eugenii in situ. Fig. D.—Head of Sideranthus
rubiginosus, with larva of Desmoris scapalis in its cell. Fig. E.—
Section of cotton square, showing pupa of Anthonomus grandis

and pupa of primary parasite Catolaccus incertus. Fig. F.—Earthen

cell of larva of Desmoris scapalis. Fig. G.—Desmoris scapalis,

adult -weevilsiccg. .025: G55. ae eet eee Stes oan eee

TEXT FIGURES.

Fia. 1. Bracon mellitor, parasite of boll weevil......--- Pt iene
. Pediculoides ventricosus, mite enemy of boll weevil..........-.--------
SS TGAL PRUs CUICULIONIS =. 725652 caeeen oe ek ee eee ee :
. Map of Texas, divided into geographical regions and illustrating average
percentage of parasitism of the boll weevil in all cotton forms... ..--

. Diagram of cotton plantation of H. O. Samuel, Dallas, Tex., of 64.4
acres, showing location of plats where observations and experiments

on boll-weevil parasitism were made-:......-+--.2 62-005 4ss-+5-5

6. Occurrence of Catolaceus incertus, Bracon mellitor, Cerambycobius
cyaniceps, and Eurytoma tylodermatis in Texas and western Louisiana.

8

m co bo

or

Page.

46

46

10
10
11

19

25

30

SUIS OF oeARASI LES OF THE COTTON
BOLL WEEVIL.

INTRODUCTION.

Although the boll weevil (Anthonomus grandis Boh.) has shown
adaptation to many diverse conditions, it is not free from parasites.
From the work detailed in this bulletin it has become apparent that
certain enemies attack it in arid regions and certain others in humid
regions; that some species attack principally larve in dried or sun- °
exposed forms, while other species attack the weevil stages in moist,
decaying forms; that some species work most readily in prairie
country, while others prefer woodland. The weevil does not escape
parasitism by dispersion, for the local parasites are capable of
‘‘adaptation and attack the boll weevil in its first generation.

Being a species with an all-season food plant, Anthonomus grandis
has made a decisive gain over most of its near relatives, which are
confined to more or less limited periods because of the shorter sea-
sons of their host plants. The most active weevil parasites in
the South have become adjusted to one host after another until in
most localities they have a regular seasonal rotation of hosts. From
this multiplicity of parasitic relations it is but natural to evolve a
new tendency and to attack the most abundant species of the locality,
namely, the boll weevil.

As the native weevil hosts are more or less limited in distribution,
it is found that the boll-weevil parasites are likewise geographically
restricted. The combined activity of two or more species in certain
favored districts has been the cause of forming three known centers
of intensive parasitism in Texas, of which the most important is
near Waco; the next around Goliad, Cuero, and Victoria, and the
third in eastern Texas. The agencies forming these centers are in
no case identical. Catolaccus incertus is the most active parasite at
Brownsville, Tex., and Orange, La. Bracon mellitor (fig. 1), which is
the predominant parasite for the entire infested area, is the most
active in the entire western half of Texas. Cerambycobius cyaniceps
is predominant in northeastern Texas. Hurytoma tylodermatis shows
its greatest activity at Overton and Dallas.

9

10 PARASITES OF THE COTTON BOLL WEEVIL.

HISTORY.

Prior to the year 1906 very little information concerning the para-
sites of the boll weevil had been accumulated. In 1895 Townsend

Fic. 1.—Bracon mellitor, parasite of boll weevil.
enlarged (from Hunter and Hinds).

mentioned a small hymen-
opterous parasite; also re-
corded the suspicious oc-
currence of several species
of Scymnus in the squares,
and mentioned that a fun-
goid parasite, a species of
Cordyceps, ‘‘was found
growing out of a dead pupa
in its cellin a boll, Novem-
ber 26, in a field in San
Juan Allende, Mexico.”
(Townsend, 1895.)¢ In
1896 Doctor Howard
stated that the parasites
were only abundant late
in the fall and that from
“15 to 20 per cent of the

weevil larvee in fallen squares in November at Beeville and Kenedy

were destroyed by parasites.”’
(Howard, 1897.)

In 1901 Professor Herrera
published a preliminary note
concerning Pediculoides ven-
tricosus Newp. (fig. 2) (Ran-
gel, 1901b, p. 206), and in the
same year a more extensive
note on the work of this mite
was published by Rangel. In
testing the mite’s ability to
propagate, 250 squares were
divided into lots of 50 each
and an infested larva placed
with each lot for four days.
One hundred check squares
were used. At the time of
examination the check squares
were free from the mites, while
the others contained 193 wee-
vil stages, of which 61 were

Fig. 2.—Pediculoides ventricosus, mite enemy of boll
weevil. Much enlarged (adapted from Brucker).

attacked by Pediculoides; in other words, 31.6 per cent were parasit-

a Bibliography, pp. 50-51,

HISTORY. tt

ized. A small field test proved that the mites increased and spread
(Rangel, 1901¢).

In 1902 Dr. Wm. H. Ashmead described Bruchophagus herrere,
from Coahuila, Mexico, as a primary parasite of the boll weevil (Ash-
mead, 1902). In the same year Prof. F. W. Mally recorded the
fact that Bracon mellitor Say and Cerambycobius (Hupelmus) cyani-
ceps Ashm. had, since 1899, been bred by him in considerable num-
bers from the weevil. He also recorded a species of Eurytoma
(Mally, 1902). In 1904 Hunter and Hinds recorded additional pri-
mary parasites as follows: Sigalphus curculionis Fitch (fig.3), Catolaccus
yncertus Ashm., Urosigalphus (robustus Ashm.), Bracon (dorsator Say),
and Hurytoma tylodermatis
Ashm., as well as an ento-
mogenous fungus, Aspergil-
lus sp. (Hunter and Hinds,
1904, pp. 104-110). The
determination of this Uro-
sigalphus has been found
incorrect. It has just been
described as Urosigalphus
anthonomi Cwfd. (Crawford,
1907a). The form known
under the name of Bracon
dorsator is merely a small,
melanistic, fall form of Bracon mellitor Say. Finally, Banks has
described a mite, Tyroglyphus breviceps, collected at Victoria, Tex.,
from boll weevil larve (Banks, 1906, p. 17).

Fig. 3.—Sigalphus curculionis: a, male; b, female; c,
antenna. All enlarged (after Riley).

THE WORK ON PARASITES IN 1906.

As the work of the year naturally divided itself into distinct sec-.
tions, it may be thus divided for treatment in this report:
I. Examination work—ascertaining the general status of boll weevil parasitism.

II. Propagation work—attempting to increase the percentage of parasitism by
release of parasites.

III. Breeding work—study of the biology of the parasites. ;

IV. Source work—study of the surrounding field conditions in order to ascertain
the hosts which have contributed the parasites, and to learn the biology of
each of these hosts.

VY. Conclusions—a review of the situation as a whole.

I. EXAMINATION WoRK.

In order to arrive at a proper knowledge of the status of boll
weevil parasitism large collections of infested squares and_ bolls
under various plant and field conditions were made in many parts
of the weevil territory. This material was immediately shipped to
the laboratory and examined by the various members of the labora-

12 PARASITES OF THE COTTON BOLL WEEVIL.

tory force. Messrs. Hinds, Bishopp, Crawford, Cushman, Jones,
Morgan, Pratt, and Yothers were engaged at various times in the
collection of infested material and the more important work of
examination. Upon their accuracy of observation the finding of
the parasites was dependent, and to them belongs credit for the
amount of figures herewith presented relating to the percentage of
parasitism. A careful tabulated record was kept of all stages of
the weevil, alive or dead, and all parasites were isolated in tubes.
Kach- polleeamns lot acest a number and each individual a sub-
number, and its stage and the nature of its parasitism were recorded.
As each parasite matured the record was placed opposite its number,
and the insect was either used in experiments or laid aside for the
collection. In this manner there could seldom be an error as to the
nature of the parasitism, as all evidence was in the tube.

RECORDS PRIOR TO 1906.

In previous years no regular method of making the parasite rec-
ords was in use, and consequently there is difficulty in the comparison
of the former conditions with those now prevalent. In some cases
the percentage was obtained from the breeding records only, and
at other times by an examination for the total number of stages
present.

In 1902 three observations were made between July 31 and August
11 at Calvert, Guadalupe, and Victoria. The figures obtained
included only bred stages, omitting consideration of stages killed by
heat, fungi, ants, crushing, etc.

In 1903 Dr. A. W. Morrill first pointed out in his notes that para-
sites occur in dry hanging forms,? and also in bolls. No other
mention of the former condition can be found prior to 1906. Only
two observations were made, both being at Victoria, in June.

In 1905 four examinations of bolls were made during March, at
Calvert, Palestine, Runge, and Victoria, in order to ascertain- the
condition of hibernating stages. These examinations consider the
total number of stages found hibernating in the bolls. The figures
are important, as they prove that the parasites also hibernate in bolls.

During August of the same year nine examinations were made at
Gurley, Quinlan, Victoria, and Waco. These figures were obtained
from fallen forms and included the total number of stages in the
forms.

In 1902 and 1903 the figures obtained must give a higher per-
centage of parasitism than actually occurred, as no account was taken
of the stages dead from other causes. It is also very unsatisfactory
to take the entire percentage of parasitism in all forms on account

aThe ap of a ms”? ae been used to designate a mixture of squares and bolls.

RECORDS PRIOR TO 1906. 138

of the differences which prevail between squares and bolls and
between fallen and hanging squares or bolls. However, taking the
figures as they are, we have the following results:

Taste I.—Total percentage of parasitism of the boll weevil, by years, prior to 1906.

Weevil | Percentage

ar. Months. ie Parasites. | of para-

Year | stages. | fel ae
OEE ee eee ne eee DUly Sb OPAMP USHA Scere oat eee eae 601 6 | 0.99
MOOS S55. ease nae oes JUNETS COOL leans sales cece sees ween | 178 6 3.37
GOS eee ee teen ec bes March 1 to 23 (hibernating) .............. 1,005 32 3.18
OOS Sg Skates ace See ATIZUSEHLZ GONSLS estonia one sioteeiee inceees 1, 702 21 | 1.23

|

In 1905 some of the records differentiated between squares and
bolls, so that a comparison may be given between the percentages
of parasitism in each.

Tae 11.—Percentage of parasitism of the boll weevil in fallen forms at Gurley, Quinlan,
Victoria, and Waco, Tex., in 1905.

| ea | Percentage

Form. ieee Parasites. | of para-

Nata es | sitism.
CUBR TGS ameter ee ee craic mae SORE pee seins ac cob Se cs enti sane 852 | 17 1.99
BONS epee nee neta te ose oaiee oar-ctee Rome aceploc cone eeistiace wo c/s Ae 123 1 81

BREEDING RECORDS OF 1906.

Although the work of 1906 was very incomplete in some respects,
it is representative of the conditions. The facts have been ascer-
tained one at a time, and as each one has been proved, it has been
found that more complete records should have been taken earlier in
the season. The material was collected at 25 representative locali-
ties in Texas, 2 in Indian Territory, and 5 in Louisiana. Seventy
collections of infested forms were made, comprising about 87,000
bolls and squares. The result is that these infested forms contained
39,183 weevil stages, of which 1,689, or 4.31 per cent, were para-
sitized. The material collected was separated into lots, according to
location and the nature of the forms.

A brief classification of the results, by months and by the four
plant conditions, gives some very important information. In the
first place, the percentage of parasitism is, to all intents and pur-
poses, about equal throughout the season and not highest in the fall.
It must, of course, be borne in mind that the collections were not
made at regular intervals at stated places, but are from different
localities each month and include different factors. During Sep-
tember and October a number of localities on the extreme border
line were examined, but the percentage is not appreciably altered.
The plant conditions were altered also, so that one month there
might be more squares than bolls, and another month more bolls than

14 . PARASITES OF THE COTTON BOLL WEEVIL.

squares, examined. A correct view of the monthly variation may be
obtained by an examination of the complete tables of observations,
which are given on succeeding pages. It will be noticed in that con-
nection that there is no graduated seasonal wave of parasitism.
Other and still unknown factors enter into the problem.

The following summary also discloses the fact that hanging forms
provide conditions superior to those afforded by fallen forms, and
that squares are more favorable than bolls.

Finally, it appears by comparison with the table of results for pre-
vious years that there has been a gain in the total percentage of
parasitism as well as in the two classes of forms.

Taste I11.—Percentage of parasitism of the boll weevil, by months, in 1906.

Hanging Hanging

Fallen squares. squares. Fallen bolls- isis. Total.
Montl sleet giual léle alee = alee
= aaltepet MT ae ey le | eee later hae re a heer lop hdl oe
BB es [me |” Sey cada ROE, SMa eS eg) Pee eer aes an
oe A pete aes Ea aes ce sae ete a alleacs 2 a 5
mo |u| & nm || & a || A | m |A| a D a | A
wars | as dae 3A. pea RSS te Peel ute
1906
Dts 5 tas eee .| 3,881} 118} 3.08} (2) |@| @ | @ |@| @] @ |@! © | 3,831] 118) 3.08
nly Aare eee. a 4,621| 210 4.54| 247) 76] 30.76] 169] 0] 0.00/ 22] 0! 0.00| 5,059 286) 5.65
AUS te soon eee 10, 342} 174! 1.68] 2,973) 348] 11.70] 2,490) 20) .80| 2,925) 192) 6.54/18, 730] 734} 3.91
September... .... 5,665} 286) 5.04] 1,883) 117] 6.26] 1,591) 23) 1.44] 1,140| 67| 5.87|10,279| 493) 4.79
October cere: ...| 347] 8} 2.30) 20) 0} 0.00] 57] 1) 1.75) 860) 49) 5.69) 1,284) 58) 4.51
| | | |
Entire season. ./24,806| 796) 3.20) 5, 123 541| 10.56| 4,307| 44) 1.02| 4,947] 308| 6.22/39, 183] 1,689) 4.31

a No records made.

The observations for June were made only at Victoria; those for
October were made at Dallas for hanging forms and at Mineola for
fallen forms.

Owing to the extensiveness of the tables the figures are consider-
ably condensed, but still retain the value of their records. Numerous
cases of high parasitism were found, of which the following may be

cited as notable examples:

Number of Per cent
stages. parasitized.
Fallen squares:

Roosevelt,-Tex.,¢ September 24... 22 ash .24.2 5. ee nee eee 69 14.4

Brownsville, @exs¢ September 62) 22-55. ere ee eee 1, 147 12.4
Hanging squares:

Waco, Tex. dully 2o..5a2 28. 22sec. bee nes See eee aS 99 52.6

Waco; Tex.; September 20.22. 1:24. ese eee ee 109 23.8

Cuero, ‘Tex. August 3,-2.6 222s. ae te eee eee 347 74 ee}
Fallen bolls:

Corsicana, Tex. -Septentber 18: 22a: ener ae ee en eee 34 rays!)

Junction; “Tex. Sepvember 242 asus aye oe ae oe ii 5.8
Hanging bolls:

Marshall Tex., (August 22 .:22akece ihe ao kee hare ene tee 52 13.5

Trinity; Dexs, Ampust) see 5 8228 oe ee eee 142 12.0

Waco; @ex.,; September 20s... geet oe eee eee een 303 11.8

@ In both of these localities fallen squares would naturally be dry.

BREEDING RECORDS OF 1906.

15

Numerous other high percentages were found, but these will con-
vey an impression as to the promises of success obtained.
In the table herewith presented only the totals for each locality
are given, and in more specific tables the seasonal variation at given
localities will be recorded separately.

MOST FAVORABLE PLANT CONDITIONS FOR PARASITISM OF THE BOLL WEEVIL.

TasLEe IV.— Most favorable plant conditions for parasitism of the boll weevil.

Locality.

TEXAS.
Brownsville...

Beeville.......
Corpus Christi.
Goliad: -=5 25 =.
San Antonio...
Kerrville

Home.
Lula
16 miles south

of Roosevelt.

Palestine......

Trinity
Dallas

Overton
Marshall

INDIAN TERRI-
TORY.

Fort Towson. .
Kosoma

Johnsons
Bayou.

Motale see

Squares Squares Bolls Bolls :
fallen. hanging. fallen. hanging. Totals.
ae Salle cle ee 34
Season. | g 28 g 22 2 a: é 2s g ae
i JE IBS| & |S (SS a ESS] a [HIBS] a | & ES
S |3/88| 8 |Si(58] S (S|SS| S |SlBS] S | & IER
nD |e wm |Hial nm Alsal wm |B al a Ay | ey
|
1906.
J ay 28-Septem-| 3, 095/202) 6. 5).....].-- 384/13] 3.3 |... -. 3,479) 215) 6.1
er 29.
aly 12-Septem-| 2, 660) 36) 1.3)..... 324] 0/ 0.0 |....- 2,984] 36] 1.2
ber 3. |
UbIhy Ms Sosqose | 438) 39] 9.0|..... selEAsel) 22.0)| CUSO) bec ee =|-<=-|| 458]? 30)) 885
August 7-Sep- | 821) 70) 8.5) 209) 29/13.8) 133) 3] 2.2 | 265] 13) 4.9] 1,428} 115) 8.0
tember 3.
MUSIStLOS see ss ee Ol Os Olpeceree | eeeeel| © aoe aera tec Goal emetel ae 33 0) 0.0
August 26-Sep- 38] 4/10. 5)... . AD) Tl 750) |S 2 Aee| ee = 333} 11) 3.3
tember 3. 7 |
September 24...) 69) 10/14. 4).....|.-- TNO) CRONE RARE ee 143 10] 6.9
ee GOs see sree 29), RSE Ble Sree ss IgA all Sate h |e ed 146) 12) 8.2
September 13254|) | S93 MONOL0l ss 52 5 S22 nab nee pa oe eat 93 0} 0.0
September 26...| 95] 0| 0.0|.....|...|....|.-.-- “Mewettabe ? be 95) 0] 0.0
September 25...| 81 2) De Al eee aeons G} COMOROS ee ale oe 90 2| 2.2
August 9-31 ....| 1,059) 11) 1.0 439) 87/19.9| 80, 0, 0.0! 189) 2| 1.0 1,767; 100) 5.5
August 9-30 ....| 1,040) 5) 0.4) 401) 37) 9.2) 93) 1) 1.0) 295) 11) 3.7] 1,829) 54) 2.8
June 16-Sep--| 6, 557/203) 3.0} 802} 82)10.2| 162) 0} 0.0 |” 201} 1) 0.5] 7,722) 286) 3.7
tember 1. | |
August 16...... 153) 1) 0.6) 160) 8] 5.0} 63) 1) 1.5 40) 1) 2:5] 416) 11) 2.6
Jeep chem 1, 508/128) 8.4} 460/126/27.3] 283) 5] 1.7 | 746} 83/11.1] 2,997) 342/11. 4
er 20.
August 22-Sep- | 1,054 15) 1.4) 81] 8] 9.8) 347) 0) 0.0 | 181] 12) 6.6) 1,663} 35) 2.1
tember 13.
August 23-Sep- 16} 0} 0.0) 102) 12/11.7) 34) 2) 5.9 | 176) 6] 3.4) 328) 20) 6.0
tember 18.
August 10-Sep- | 1,054] 14) 1.3) 414) 48)11.5) 209) 1) 0.4 211) 7) 3.3) 1,888) 70) 3.7
tember 4. |
August 9-30 ....| 1, 554) 20/ 1.2) 169) 21/12. 4) 949) 4) 0.4 | 269) 28/10.3) 2,941) 73) 2.4
August 17-0 c- 75} 0) 0.0) 150} 7} 4.6) 74! 0} 0.0 1,068} 64) 5.9) 1,367) 71) 5.1
tober 6.
September 19.-.| 53) 0 0.0)_.-.. AGITO|NOXO} | Eee! see ee 99} 0} :0.0
August 10-O c- 445) 9] 2.0)....- GyA\ alll alee Bee Salisoalmee 502} 10} 2.0
tober 2.
August 23......| 197} 2)1.0) 45} 1} 2.2) 37) 0] 0.0 89} 4) 4.5) 368 7 1.9
August 22...... 23/ 0} 0.0 6) 116.6 18) 0} 0.0 §2| 7/13. 5) 99 8} 8.0
|
September 12. ..|...... YAN) OOO) eocllaelseeed| aaoee Ve 521 0} 060
September 14...|_..... GAO MONO | Rees es) Sots Seer [eee 64 0) 0.0
August 24-Sep- 491) 3] 0.6) 249) 7] 2.9) 211) 0} 0.0} 626) 37} 5.9].1,577) 47) 2.9
tember 29.
Aveust 232 2. :| 1,315) 6) 0.4} 633] 66110. 4/ 321) 5! 1.5] 539! 32) 5.9) 2,808) 109! 3.8
September 23-30} 435) 5) 1.1).----|--- eee G7 OR OFOM | Eee sae te 502 5) 0.9
September 8..../......]..- seal) eZ2k NOS Boeas|lealesses |eseaeleee 218 1} 0.5
ATICUSt 22 eee) eeeo5 lee Ol ONE ole ee Se eles el (egret sea (ene 225) 0} 0.0
pe Pee oe Cha 24, B06)796 3) 215, 123 541 10. 5/4, 307 44| 1.02 4, 947 308 6. 2:39, 183/1, 689) 4.3

The records from only five points extend into three months, and
these may be taken as indicative of the seasonal variation at a given

locality.

16 PARASITES OF THE COTTON BOLL WEEVIL.
TaBLeE V.—Parasitism of the boll weevil in fallen squares, Brownsville, Tex.
: Percentage
Date. | Stages. | Parasites. parasitized.
1906.
Ly 28 Also se = aR ne in Pee CER lun view Wo weet nen al eee 1, 568 50 3:2
TAVUE ed 0 (| ic aS et er Pere este ee ee eet on AM itr enn SN os Oe Na et 115 2 ils 7/
September 5s, soto) at. Sas 5 reek eee pet et Mens eed OL eC 1, 147 142 12.4
Sep temlperi2g of crt: Pace Sees cee eee eee oer Seni Ste De men ae 265 8 3.0
TasBLE VI.—Parasitism of the boll weevil in fallen squares, Beeville, Tex.
, Percentage
Date. Stages. | Parasites. parasitized.
1906.
A AOUn AYE Senne Aaa ee Re Se Ine Pee tee ee, Reece ey ed 656 26 4.0
PANISUISTIS S O> rae Neyo Steyn = Se ce tee nse oe et ae nee ee ee eae ee 442 2 ~ 45
PAMUIUIS TNS Ein 35 tae area Se tors Bee Ae Ee eT ae ae Se ee ee es 884 2 22
DEPLOMIbONS shoe ie cc see Noses ere ae esas eee ee eee eee 678 6 9
TaBLeE VII.—Parasitism of the boll weevil in fallen squares, Victoria, Tex.
Date Stages Parasites Percentage
: eS * |parasitized.
1906.
MEIC NEG = vceaerstae Ua) 2 eee ee Retin s = Bb n uae es Sg eee EI 864 15 1.8
AUTOS CASED eee rae pera nid crit aoe Ne RRS Ae Hy eg Serta meer, tat 667 9 1.3
S10 TR Oe bet OORT SE ee Seana pete a See eres NE 396 11 3.0
AUT GG Sa rose ee ree ae aoe ae ee eee 652 26 3.9
SIUTIO V2 Ue ro fterarars chs 2 Berets Se Se fee a at A te REO em ore ee 829 33 4.0
HUME DZS re ne orate cee a eR eae ee cers eee ae ec ene 423, 24 5.7
JELLY RCP oe ye Re ce On ete ee eta fe R= Ae ee ee ae 535 19 3.5
EST 0 eee oS rare rete ia coe Ree ere ee rece Oe ee eee eee 560 13 2.3
al ya 20 eee rat ere Jia BL er aban lg nt vos toe ape) McA ae On 518 32 6.1
MAE ADD ete Ree MS ore ae See See ayn an et ne Sta mS A ca 87 5 5.7
Septembordenes asa o5 ee ses ee nt See ancl me epe ire ey aaa 1,026 16 1.6

TaBLE VIII.—Parasitism of the boll weevl in all forms, Waco, Tex.

Fallen squares. | Hanging squares. Fallen bolls. Hanging bolls.

og og oq og

Date. On| Se eee a) a a eee

| ¢)2/88| 2) 3/84) ¢| 3 |82| 3 | a |3a

of) s 3 1) S s =f) Ss 3 1) S 3

= Bence es a |Sa} 8 a | oe VS a 188

a BA |e) om | & |aa| om | ow [eal om | & lee

1906.
MIU 2 25 Ree Be 259 26 | 10.0 99 52 | 52.6 44 0; 0.0 22 0 0.0
RUPUISE B72... oe. 82 eee 32 2) (622 22 3 | 13.6 24 0; 0.0 16 0 0.0
INUIP USD 28 Eee ee 2 eee 1,217 | 100} 82) 230 45 | 19.5} 215 5| 2.3) 405 47 | 11.6
September! 20: 25h. - 5 2) stiontecl Mee seal -,- eee 109 7a PS syel teciel eerseere| aeaae 303 36 | 11.8
| |
TasLe [X.—Parasitism of the boll weevil in hanging forms, Dallas, Tex.
Hanging squares. Hanging bolls.
Date.
: Per cent of F Per cent of
Stages. Parasites. parasitism. Stages. | Parasites. parasitism. ~
1906.

IIPS 29> oo. cenoris ee dicceeece 12 1 8.5 26 0 0.0
September12--2..cee sce. eee 118 6 5.0 182 15 8.2
October 2.52.9 esse 8 0 0.0 250 10 4.0
OctoberiOr ress ceaseneeceece 12 0 0.0 610 39 6.3

BREEDING RECORDS OF 1906. 1
. J

The only one of these tables in which there appears any definite
progression is the Victoria table, in which there is a regular increase
of percentage throughout the last half of June.

FIELD CONDITIONS.

In order to arrive at an explanation for the obviously irregular
conditions of parasitism, the figures have also been rearranged
according to field conditions. It was found that generally hanging
forms in wooded or cleared country were more highly parasitized
than those on prairie land. The only definite comparisons which
could be found were as follows:

Taste X.—Percentage of parasitism of the boll weevil in forms in woodland and on

prairie.
Woodland. Prairie.
Locality. Date. Hanging| Fallen | Hanging| Fallen
forms. forms. forms. forms.
1906.
Giolla dist rer ssc rac tees ee SeptLeMpeCld asa ee eee ae 11.2 8.2 | 6.0 6.4
WHC ONS esmcteee ao Slee icicle Sacra soe os | Neptember 192222 ccs---- | 1 Oe eaeseeeee | TANG) |e eee

Stages in fallen forms on prairie land were found to have a higher
general percentage of parasitism than those in woodland, although
the definite case of Goliad gives a different condition. It is only nec-
essary to quote the highest percentages reached in each condition to
give support to these statements.

The highest percentage in hanging forms on woodland was 42.6.

The highest percentage in fallen forms on woodland was 8.3.

The highest percentage in hanging forms on prairie was 14.6.

The highest percentage in fallen forms on prairie was 10.1.

Considerable contradictory matter was brought out in comparing
the parasitism in fallen forms in shade and in those exposed to the
sun. The weight of evidence supports the statement that those fallen
and exposed to sunlight are more highly parasitized than those which
fall on moist shaded soil. Only five definite comparisons can be
presented to show the irregularities which exist.

TaBLeE XI.—Parasitism of the bobl weevil in fallen forms in sun and shade.

: Woodland. Prairie.
Locality. Date. = aor ==>
Shade. Sun. Shade. | Sun.
1906.
Mansfield Ware aces sesso seaeee ND Feg DES) P-L Ue eo en eae 0.54 VOOM | esas =A ee
Raby lor exe esc eee pee PACS tin Oper eee ene i ery ee SSS nee ite 0. 69 i133}
print yew ROX. he Se hase ees a ISIS t Sees by eee 2.3 LOT ance ot ene eee
Wan Web eine SEU ty ee ee ASLO PAO) hs PN cles ci, ee a liaee lle SE ae ary a ma St 6.5
WictorianiRexs 4. oe seer tat UU ges ey rn me Se eros ye Lee 20%5)|Sae ae
NVi8LCO, ME OX: tee 5. cine ete gerry. Ale 8 AUSUST AO ec eA ee econ Soe 8.7 GiSiene a2 See WE = See ane emer

10292—Bul. 73—08——2

18 PARASITES OF THE COTTON BOLL WEEVIL.
+

The highest percentages obtained in each class are as follows:

Tm ghadeon woodland 2) tse = as Fe ee cee ne eee eee ees Oe 8.7
insu on woodlands a... sceu Sette: He eee oo ey ee oe ee eee 8.3
Im-shadeofiplantea op prairie. 3... 22) 2s. shee ep i a 8.2
[ns On praities:. 0. oes seen te ae ee en ee ea ee 10.1

Should further inquiry prove that sun-dried squares are the most
highly parasitized, another reason is presented for wide rows.

GEOGRAPHICAL CONSIDERATIONS.

By far the most important facts established by the table of per-
centage of parasitism (Table IV) were elicited by using the geograph-
ical map of Texas, published in the census of 1880, as a base.

I. The alluvial prairie of the Rio Grande is represented by Browns-
ville with an average percentage of parasitism in all forms of 6.1.

II. The coast prairie is represented by Corpus Christi with 8.5 per
cent, Goliad with 8.0 per cent, Cuero with 5.5 per cent, and Victoria
with 3.7 per cent.

III. The Edwards plateau is represented by Junction with 8.2 per
cent, Roosevelt with 6.9 per cent, and 16 miles south of Roosevelt
with 2.2 percent. This latter case was a farm with very rank growth
of cotton, from which shaded squares were picked on the ground.
The weevil had not been present long.

IV. The black prairie is represented by Kerrville with 3.3 per cent,
Taylor with 2.6 per cent, Corsicana (IVb) with 6.0 per cent, sal
Dallas with 5.1 per cent.

V. The cross timbers are represented by Waco with 11.4 per cent.

VI. The eastern hardwoods are represented on the south by
Beeville with 1.2 per cent, Hallettsville with 2.8 per cent, each at the
tip of an extension of this region, and by Calvert with 2.1 per cent,
Palestine 3.7 per cent, Mineola 2.0 pe cent, Overton 1.9 per cent,
Marshall 8.0 per cent, Mansfield, La., 2.9 per cent, Many, La., 3.8
per cent. Marshall is probably pore ented by a field on a red-land
knoll, which would account for the discrepancy.

VII. The eastern pine country is represented by Trinity with 2.4
per cent, and Orange, La., with 0.9 per cent. ‘Trinity is in a transi-
tional region of pines and hardwoods, but seems to belong more
typically 1 to the hardwood region.

It will be readily observed that regions I, II, III, V, with Corsicana
in IVb, which really belongs to the brown oa region in the eastern
hardwoods, and Marshall in VI, which belongs to che red-land areas
of the eastern hardwoods, are the most highly parasitized, while the
regions IV, VI, and VII are the lowest parasitized.

The map on the opposite page (fig. 4) is presented to illustrate these
statements.

While it may be merely a coincidence that the places 1 in the same
belt have about the same pr Speman of parasitism, it is nevertheless

BREEDING RECORDS OF 1906. ag)

worthy of some attention. It is undoubtedly true that each of
these floral regions is also a distinct subfaunal region. It may there-
fore be expected that different weevils will exist in the different
belts and different parasites operate upon these weevils. Some
of these parasites may be able more readily than others to adopt
the boll weevil as a host, or in certain regions there may be more
species of parasites capable of doing this. In the discussion of the
geographical distribution of the parasites more light will be thrown
upon this question.

BOT, TOMS

Fia. 4.—Map of Texas, divided into geographical regions and illustrating average percentage of para-
sitism of the boll weevil in all cotton forms: I, alluvial prairie; II, coast prairie; III, Edwards
Plateau; IV, black prairie; 1V>, brown loam prairie; V, cross timbers; VI, oak, hickory, and pine;
VII, pines. (Original.)

BOLL WEEVIL STATUS.

That there should be some definite relation between the percent-
age of infestation and the percentage of parasitism was expected,
on the ground that a condition favorable to the weevil should be
favorable to the parasites. It was, of course, conceded that climate
would be another agency to figure in the question. The year’s
work is deficient in that coordinate examinations of infestations and
parasitism were not taken. In another year this will be necessary
in order to show whether the parasites are reducing the infestation.

At present no relation can be found between parasitism and infesta-
tion or climatology. Beeville, Goliad, and Victoria have a mean

2() PARASITES OF THE COTTON BOLL WEEVIL.

annual temperature of 70° and a precipitation of 30 inches. At
Beeville the average percentage of parasitism in fallen squares was
1.3 per cent, at Goliad 8.5 per cent, at Victoria 3 per cent. This
condition occurs throughout the entire State. At Calvert the per-
centage of infestation of squares on August 28 was 55.6 per cent and
at Goliad it was 20.9 per cent, and yet the average percentage of
parasitism in fallen squares at Calvert was 1.4 per cent and Goliad
8.5 per cent. A reverse condition may be cited as follows: At
Cuero the percentage of infestation of squares on August 28 was
74 per cent and at Victoria it was 32.7 per cent, and the average per-
centage of parasitism in hanging squares at Cuero was 19.9 per cent
and at Victoria 10.2 per cent. Table XII gives in detail the irregu-
larity displayed between these three factors.

TasLe XII.—Boll weevil status and parasitism.

First exami- :
eis Weevil status, ed eRe ee Se aire
nega Jee een Aug. 28, 1906. Average parasitism, 1906.
an | nual a
Locality. ne pee AV ere’ per qT orenrage in- Squares. Bolls.
pera Ap Bee se : ; .
ure. | 4;

. tion. | _ Tee- aioe An
Plants. Me Squares. | Bolls. | Fallen. ee Fallen. nce
Brownsville, Tex......... 72 Did |e ae eel Stee ea ciara oe See 625 kee 353); eeceer
Corpus Christi, Tex...... 70 30 | 9,412 212 71.0 | 84.0 S50R Re eenee OSO; See ee
Beeville; Mexeoi see ane 7 30 8,859 300 218 | Sak Si er eee 0:0: Seeee ee
Goladn@extaesceceecace 70 30 | 10,169 305 20.9 1.6 8.5 13.8 2a) 4.9
WMictoria, exe so. 4555 pes 70 30 | 8,334 119 S207) Lon0 3.0 10. 2 0.0 Fi)
San Antonio, Tex........ | 68 25 | 12,186 101 38. 0 574 (OK Os sete | oor Sie Besta
Wornktowmhexn. = eee 68 SOB ieee cl secee 67525520! Sr 4 ae hese tee Sal See ne See
Cuero wexsss ee a eee 68 3 7,225 135 74.0 | 59.0 1.0 19.9 0.0 1.0
Hallettsville, Tex......... 68 | 30] 8,056] 289 7810) Ba SAN Oye TOM ee resad
PAtIstine Dex ceases eee 68 30 | 15,580 73 19. 2 DU Ne eee, ool eal ee ee
Giddings, Nex* 2255 -.--2 68 30 | 15,600 76 BAL uted ho Se | 9a ae Bean ee ec |e
Whulas Dexa. see esse 66 Ol eee ecneell YU acts lb Sp pan to ea Ree (Ce OLOM ec bee ee ae ere

Sixteen miles south of
Roosevelt, Tex........- 66 P25 el eek Acts | nee agement eee rer ged [Perey aaa Da sense |S ees SeeeeerS
Roosevelt, Dex2) 2 ese.-- 66 PS Ve Ae | ein a aire eee ete |e a TAS 4a eee | O20; |2=2eeee
UNCON AENe xe ae ae 66 Dive esee etl eet eal eee cee ae Su0n | Lames et Set |e ae
Mountain Home, Tex....| 66 yA RS Spe | eed | yes oka Va cont OL Oi Sass. c alee ee |e ee
Kerrville exec ae nese | 66 DS eaten Po ee | es ee | ere ee IO SGIE Seaee DSi) Ee eee
iDTryans Mex sence e see 68 40 | 10,170 38 52.3 Aid Nes. 4, todos) eae See
Hempstead, Tex...-.---- 68 40 | 12,255 84 V4. O50 Allan os Sella Sonree oll Seeeeee eee
Eagle Lake, Tex.......-- 68 40 | 5,735 89 2028 ISE0" Ik: Shee |b asa eee eee
Wiharton, Mexes.-.-. =f. 2 68 40 | 5,127 93 5400 | abi8 4 Ae ees Seen eeeeeee
ayvlon TEx caer -cceone. 66 SOY | SRS ee | es ol ee Cee eee ee oe .6 5.0 | 15 20
Camoron-"Nex-ee- 5. a-e 66 30 | 16,317 67 73;.3°)'-4650) ||eccoze 4a beetten be aee eee eee
Wisco Lex: 2570 ae eee oe 66 30 | 13,800 49 68. 7 Ga 8.4 27.3 67) nbieal
EinlishonoWLexares-esee oe 66 80 | 18,200 0 AGM 8. 4y 0) 22s ot |boe-g2 |e eres
Corsicana hex. snot ee 66 30 | 20,524 0 38.0] 1.4 COM) ali 7 5.9 3.4
Mexia Dexa sto 4 2a-2 oars 66 30 | 18,520 0 -8 OHO! Oh ceseyeeel Sees | er aa | eee re
Calvert: Texs25)) ae oe 66 40 | 11,377 186 | 55. 6 8.0 1.4 9.8 0.0 6.6
Palestine; exes. eseees— 66 40 | 12,712 61 52.8 | 14.4 12, 11.5 .4 30)
Athens? Wex 55-7852 Sek: 66 40 | 11,568 21 | PANG Tol ie Noy ite (eer Meare IAs 55 5/8
Rroupe; Mexss. 2225. =e 66 40 | 8,126 390, UPAR Giger = |lyseasoee Pees see||hoossess|-on..55
Jacksonville, Tex........- 66 40 | 10,963 160 AUS SP UOY «|(Be eee Ee ep Pe ete Seles hn ca
Henderson, Tex.......--- 66 40 | 7,536 564 SWAG al bb25 Sinaloa | Soa ohsacllrere Seyaall ae ees
IDallass Mex sehen aes 64 40) ||. eee. |Meemee PPE) 0.0 0.0 4.6 0.0 9.9
Terrell; Nex #5. <8. 52 64 40 | 14,000 0 0.0 0.0 O00 eee Ql0n bessere
Mineola, Texc2 5.222252. t8 64 40 1,607 Ode emcee ee ee ZAO | Zeme mee 1.3) |e eens
Marshall Mex. sess 66 50 9,480 69 78.0 | 29.0 0.0 16.6 0.0 1 S25
Rrinity, “Lexa: ion lee sce - 68 50 9,990 182 50.8 | 11.0 M2 12.4 4 10.3
Watkins Text ae ee eee == 68 50 | 11,3290 36 DSF 2 LOU Alo Sees Oo eects |e eee engi tate
Overton ex eae ae 68 50 7,780 69 78.0. | 29.0 1.0 2.12: 0.0 4.5
Fort Towson, Ind. T...-- 64 DO! syeceere 0 | (ObO WO" eestor 0:00) S3-eeee 0
Kosoma, Ind: 2s. 222.252 64 HOM re eters 0 0.0 OF Os | eeenace O20 |= eeeee 0.0
Mamsfield, Wais2.2--s252.- 68 LV es ae ee Sees sae ace ae ss 2a aaa 6 2.9 0.0 5.9
Many sala eee as eee 68 E10] Meee al ieee a(S ag er Nice cS 4} 10.4 1.5 5.9
Orange, Wha acest eee ee 68 ee ae WE Seg RS \ ii, Oreste =e |e iP Ll eyes Se OO see
Minden anne seen aeees oe 66 OO soe ees 0 | ONO} ONOF seeaeee es 2D albsee aera peeerent
Johnsons Bayou, La....- 70 HOME See elt cee eral’ soe eee eo Gee eee O05) -feceeei|¢ estoecgeceaas-

BREEDING RECORDS OF

BOLL WEEVIL

1906.

CHRON OLOGY,

21

The boll weevil has been present in the United States since 1892.
Year by year it has extended its ravages, although during the sum-
mer of 1896 and the winter of 1904—5 it received severe setbacks.
Its territory now extends into four States, and in three of these

observations as

to parasitism were made.

Several examinations

were made immediately after the dispersion of August in newly
infested fields, and at Minden, La., it was actually found that para-
sitization had commenced within two weeks of the weevil’s arrival.
The parasites do not follow the weevil; they are already present and

native to the country invaded.

Thus it is that in some parts of

Texas where the weevil has been present ten or more years the
parasites have not as great a hold as they have at Waco, which has

only had it four years.

sites at Waco than at other points.

There are probably more sources for para-

Taste XIII.— Weevil chronology and parasitism.

Average parasitism.

Locality. Chronology. aes Bolls.
Fallen. Banging Fallen. | Hanging.
Mindenwsluaies cans sce cn. Weevil arrived in the August disper- |.....--- Eby (Ro ae Pe wear eee’
sion of 1906. First generation in
green squares collected.
Hort Rowson; ind: T. -2.|5. 25s Ose Sica toma see a eee ole meen ae O2Q) lease oases
LSToys(oya ays )p 3) bars od Se ee ee eel LOPES eee eee ea ae cee eet nes meter ORM Mee Set S20 ee ae
la Mext e225. enseces Weevil arrived in 1906, probably....-.-- CSOT | Bseaes etitere)| tek Sn ys Aloe oe eee
Sixteen miles south of | Weevil arrived in 1905 or 1906........-. | Re ae [Epeheek 2 SOE ee Bo
Roosevelt, Tex.
Roosevelt; Tex. f2-..-. 5. Wieeviliarrivedin 190522. 2secesee-see|) PAS cee Sco QAO! Se eee rae
Wimchon lex. =e sess Ose ne oe ee Ee Wei (Sv erecta Bish [ie eae
Mountain Home, Tex....|..... OSes ee ae ee eee | QUO eae, Ad S eLe BO ae
iKernvillewtex 2 2-.- Weevil arrived in 1904 or 1905.........- TORS) Rae See eS PAST a ihn
Dallas Nex = oe Weevil arrived in 1904; set back Feb- 0.0 4.6 0.0 5.9
ruary, 1905; rearrived 1905.
Mineolan Nex: 2c). 5.2. acoce (SC \a dies Sa ae ae eee a ae ed Eo DON Saree eee IE ed Nee
MenrelleMmexe. foe 0 SE a ee GOES eae aes eens eee eee ame ae (U0) i] | ee ee | 0:10) S38 eee
Johnsons Bayou, La...-.- Weevil arrived in the August disper- OLO) Pao ee teeter wre ted ea
sion of 1904.
Mamnstieldeean= sas ss. os ac OS tee ee: eae ee ee .6 2.9 0.0 5.9
Monshalleiexs 5: oss Os Soe ee Seen ee RE Se 2 0.0 16.6 0.0 1325
Overtonwexsofte-2 2 ye slooe ee GOSS ane ee eae Cee 1.0 2.2 0.0 4.5
WIE ALE Sa eoe oS ee eee Weevil arrived in the fall of 1903....... 4 10. 4 1G) 5.9
Corsicanas hexes- 9 5. 22- Weevil present in 1903................- 0.0 MSZ 5.9 3.4
WiLCON Dex 29 - ooo Weevil present in 1902................. 8.4 27.3 aE ileal
MrimityeeVexccao. a6. aoe Weevil present in 1901............ ...- 115? 12.4 .4 10.3
palestine Texe cae so) sie ass CORR aS ere ee EE One ee 1) 11.5 m4 3.3
Calvert {Uexes--peen ae eeleens GOS tae A ce erat ae ee eS 1.4 9.8 0.0 6.6
MA VOL CKsss ea 5 See mens GOR Sea a ee ee -6 5.0 1S 250
Hallettsville, Tex........ Weevil present in 1897................- .4 9.2 1.0 Bhi
San Antonio, Tex....._.. tae arrived in 1894; set back until QhOM Ss dre iseiecars |e sin, wia:=| ere eee
1897.
Crlero WL exeeee sce eee Weevil arrived in 1895; set back until 1.0 19.9 0.0 1.0
1896.
WietOrinn lemmas. = eel o-oce LOM eee nee Se ened een aI Soe 3.0 10.2 0.0 a)
Goligda Pex te, en 2 Weevil present in 19942 AB. eee S55 13.8 PLP) 4.9
Corpus Christi, Tex...... Weevil present in 1803.........-..----- ON Ol Peaeeeeeae 0:08 |S eee
Beeville; Mex ten ees al5a sos US RA = Gaara San a eee ee eae SG id baer ace Of0"|.a3:ee2eee
Brownsville, Tex. ...--.- Weevil) present Anvl892)- 322225. 2s. - 5 GxO)|Eeeeceeiee= 35.0) | see eeeeee

CONCLUSIONS.

A few points of practical importance have been deduced from the

foregoing general

tables.

vp PARASITES OF THE COTTON BOLL WEEVIL.

1. Inasmuch as the parasites are known to attack the weevil in
its first generation in a newly infested locality, it may be expected
that by releasing the proper parasites in a given locality the per-
centage of parasitism may be increased in a very short time. Such
a case was actually obtained by the release of Bracon mellitor at
Dallas, as described in the section on propagation.

2. Dryness and sunlight assist attack by the chalcidoid and bra-
conid parasites of the weevil, as proved by the following considera-
tions: (It is, of course, to be understood that this conclusion may hold
only for those species and regions studied. In fact two or three of
the minor species, such as the tachinids, give promise of doing most
favorable work under directly opposite conditions in regions so far
unstudied. )

a. Stages in hanging forms are, as a rule, parasitized to a higher
degree than those in fallen forms.

b. Stages in fallen forms on prairie land are generally more highly.
parasitized than those in wooded country, although frequently hang-
ing forms in wooded country are more highly parasitized than those
in prairie land.

c. Fallen and hanging forms on unshaded ground are more highly
parasitized than those on shaded soil.

These facts give an added importance to certain cultural methods
already advocated, viz:

a. That the rows should be far apart, in order to allow the sun to
dry the squares on the ground.

b. That determinate varieties should be planted in order to give
additional heat and light for the parasites during the cooler autumn
months.

That, as indicated by the present studies, dryness seems to be a
most favorable condition for attack by hymenopterous parasites is
quite natural, as these delicate little insects are very fond of sunlight
and warmth.

II. Propagation Work.

1. INTERIOR WORK — TRANSFER OR ARTIFICIAL PROPAGATION OF
PARASITES.

It has not been a difficult matter to breed the parasites of the boll
weevil. There have been used in this laboratory four distinct
methods of obtaining parasites, all of which served the purpose for
which they were used. In all cases where definite records of per-
centage, length of stages, or nature of parasitism were required
there has been a careful examination of each form (square or boll),
and those forms containing parasites, or sometimes only the para-
site and its host, were placed in individual pill bottles, numbered,
recorded, and placed in trays for daily observation. A somewhat

t

TRANSFER OR ARTIFICIAL PROPAGATION. ; ey

less exact method has been to place a limited number of squares in a
tumbler on moist or dry soil in order to make the time of develop-
ment more nearly normal. These tumblers were covered with
cheese cloth and tagged.

When definite records were not requisite large quantities of squares
or bolls were placed in Riley breeding cages and the parasites were
obtained in larger quantities with less mortality, which is frequently
caused by rough handling in the closer examinations. Closed boxes
with numerous tubes on one side, after the pattern of the parasite
breeding cages adopted by the California board of horticulture,
were also used for small quantities of squares. These cages probably
hasten development by increasing the heat and moisture.

There was, however, another problem which proved beyond solu-
tion for this season. It may be described as the effort to induce
the parasites to attack the stages in forms placed in cages of various
kinds. No positive results were obtained, but considerable experience
was gained in the matter of breeding-cage technique.

The first work was done with various kinds of glassware, closed
with cheese cloth. As fast as males and females of the same species
could be bred they were isolated in pairs and placed with a limited
number of infested squares. These squares were fresh, flared, or
fallen, in order to test all conditions.

Glass tumblers covered with cheese cloth proved too dark. When
dry earth was placed in the tumbler or when there was no earth, the
squares rapidly dried and became very hard, and the parasites
quickly died. When moistened earth was used the squares quickly
molded-and the parasites were killed by the fungus. When a water
reservoir was sunk in the earth the parasites drowned.

When glass lamp chimneys were placed on clay saucers filled with
moist soil there was plenty of light and the material remained in
good condition. The objection to this method is that the parasites
can not be easily handled.

Erlenmyer flasks gave plenty of light and were easy to handle as
they have a small mouth, but they sweat profusely and the parasites
were caught in the moisture on the glass. Blotters, absorbent
cotton, and corks with large wire-covered openings failed to prevent
the sweating.

The last attempt on this line was with mica lamp chimneys such
as are used with the ordinary incandescent gas burners. These
were covered at both ends with cheese cloth. They are of light
weight, give plenty of sunshine and sufficient circulation. For a
small breeding cage they are very handy in many ways. But the
parasites did not attack the weevil even in these cages.

In all of the above types of cages the life of the parasite was
hardly a day. In wire types of cages the parasites lived several
days. In the following, several of each sex of the parasites were used.

24 ; PARASITES OF THE COTTON BOLL WEEVIL.

The first type was a cubic cage made very easily by using strips of
cork for the framework and fine meshed wire (known as 50 to the
inch, but really 35 and 45 to the inch) for the sides. One side was
fixed with two openings which were closed on the inside by a tin
shutter. The lower opening was round, and in this a cork with a
glass tube through it was placed before the shutter was raised.
Thus the parasites, being attracted to light, could be quickly removed
by darkening the sides and allowing the light to enter only through
the tube. When the shutter was raised higher it exposed a larger
opening through which material could be passed.

With this first cage as a type various modifications were contrived,
all with wire fronts, shutters of various kinds, and corked holes for
the admission of a tube or for passing in small objects. Cigar boxes
became the bases for these cages.

Still another modification was a wire cylinder corked at one end
and with a smaller cork centered in this for the removal of material.
The other end was covered with cheese cloth.

It was found impossible to provide plant conditions in a small
cage unless the plant were actually transplanted or grown from
seed. “A branch of cotton withers so quickly that experiments of
this sort were of no value.

Both kinds of tubes were tested on the plants, but the mica tube
caused a heavy sweat and killed the branch it was on; the wire tube
was too heavy. All of the plants placed under the large parasite-
tight cages with glass sides died before results could be expected.

2. FIELD WORK—RELEASE OF PARASITES.

The release of parasites in the field was not commenced until
September 12 on account of the small amount of material gathered
prior to that date. Notwithstanding that fact the results give an
indication of success. On the laboratory farm at the top of the hill
there were released 35 parasites on September 12 and 38 on Septem-
ber 15. These parasites were of three species. An examination of
hanging forms was made in this part of the field and another down
the hill at the opposite side and about 350 yards distant. At this
time the check area (‘‘B,” fig. 5) showed 1.9 per cent higher para-
sitism than the parasite area (‘‘A,” fig. 5). Twenty days later, on
October 2, like examinations were made with the result that the
percentages were reversed; ‘“‘A’’ showed an increase of 2.6 per cent
and ‘‘B” showed a decrease of 6.5 per cent; that is, “A” was 7.2
per cent more highly parasitized than ‘‘B” and showed a real gain
in parasitism of 9.1 per cent. On October 6 this examination was
followed by another, and, although the difference was less, “A” was
3.9 per cent more highly parasitized than ‘“B.” Plat “A” was next
to a fence and separated from another field of cotton by a road and a

FIELD WORK—RELEASE OF PARASITES. 25

high hedge of Ambrosia trifida with scattered Solanum rostratum.
As a further check on the experiment the nearest part of this field
(‘“D,” fig. 5) was examined and proved to be parasitized by 2.1 per
cent less than ‘‘A.”” On the contrary, the farther part of the field
(‘‘B,” fig. 5) showed 5.8 per cent higher parasitism than “‘A.” The
conclusion was that ‘““E”’ and ‘“‘D” did not influence “A.”

The results are given in Table XIV for the entire experiment, and
also according to species. It is believed by the writer that it was the
release of Bracon mellitor which accomplished the change.

The evidence upon which this belief is founded is both positive
and negative. Bracon mellitor was not found breeding upon any
other species of weevil in the vicinity, and yet the examinations
showed an increase of parasitization by this species in the ‘‘ parasite

oAD J
eS

ge COTTON

8
x
S
g
nN
&
K
8
Ss
cS
e

Fig. 5.—Diagram of cotton plantation of H. O. Samuel, Dallas, Tex., of 64.4 acres, showing location
of plats where observations and experiments on boll-weevil parasitism were made. (Original.)

ce

area”’ opposed to a decrease in the ‘‘check area.’’ Catolaccus incer-
tus, although released in large numbers, did not take hold at all,nor
was it found breeding upon any other species of weevil in the vicinity.
A few Cerambycobins C yaniceps were released, but the only trace of
parasitization by this species was found in the original check. And
yet this species was bred from Lirus musculus in the stems of Poly-
gonum pennsylvanicum near by, and also from Trichobaris texana
Lec., in the stems of Solanum rostratum adjacent to ‘‘A’’. No
Eurytoma tylodermatis were released, although this species was abun-
dant throughout the field. It is known as a parasite of Lixus scro-
bicollis Boh., which breeds in Ambrosia trifida stems. Immense
numbers of this plant were growing along the road adjacent to “A.”
In order to show that plat ‘“E,’’ which was highly parasitized by

PARASITES OF THE COTTON BOLL WEEVIL.
aN ”)

1es 1n.

the fact may be cited that the higher percentage of Hurytoma tylo-
Taste XIV.—Release of parasites of the boll weevil—Summary of all species.

dermatis in ‘“‘K” evidently had no influence upon the presence of

26
Bracon mellitor, did not influence the numbers of that species in ‘‘ A,”

that spec

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gaa “TIA00M “On SS | Ras “‘TIAvoM A 14 = cole “TIA00M ‘
= ay 10 sesvig eae S |S 941 Jo sosRqg : Bes ee e244 JO sass i
ad “paztqis bo = | ad “paztqis Hau S | ad *poziqis :
2 3 -eied o8B{UddIEg nig = ES -pied o8¥8jU00I0g ‘ a! 3 = S -eied 0384000107 7
= = ) no yee pa '
< is *peziqisered er = zy ss | ‘pezipiseied : Hehe S . is *pozipis vied i
S z so3eB4s JO loquINN Sa = Soe $98eB4s JO l9GQUINN ; eee 8 Bae so3B4s JO JoquInN ;
a A “TIAveM 10} § a 5 *[TLA00M. : ei a 3 *TIA0OM ;
eS 9447 JO soseyg Bite = ~ 8S |a49 JO sosuqg : > PbS 944 Fo sosyjg|] |
a “paziqis sg = | w "paziyIs in =| "poziys
ee -Bied 088) Ue010g : flatness s Bred (988400010 © Ra Mi 3 § ad -eied a3 UI010g ' '
S 3 ‘peziqisvied ; ae She % os ‘poziqisvavd F ana "S Oe “poziqiseied : f :
28 sasBq4s JO laquInN ‘ arid = a8 Sese4s JO TaqUINN ; Pad > ae $9584s JO LoquUInN ; 5
26 iaere Q ’ 7 4 3 2 in = ‘ .
ov “TAv0mM He &, © pee Ns : Her See Ages i ‘S
= o9U1 JO soseqig oN SS = 949 JO soseig ' 1A z - ay? JO sosv4g ‘ 1A
dis iS Boy,
a "paziyis b NN | S| 4 “paztys x as S| x "poziys A
3 -eied o8vquo010 N no > = -eied 038} U00I10 : ano 2 = -Bied 9384U90.10, lor
g 4 di nk = & ai H edits se SN, :
S) g *pozipisvied ae an S Os *poziqisBaied ~t pa ae = Og *peziyiseied <t
7H sasRys JO IoquInNN = 5 > =H soseqs JO IaqUINN | "> Bis SoBR1s JO IaqGUINN |
oa] ‘TIAGO 3 a9 RS | ‘TIADOM | 3s 9 ji eo ‘TLA2o MN 3
: a74 FO soseig | a4 art) | : 943 JO sesByg | 4 i ; = e9u%d Jo sosuqig | a
4 one > > er Cae ae
- *peztqIs Cor Of wt rs = aDeZiaIs So AA iF = ‘Bee oho
a -e1ed osvyuedleg | in oN: x a -Bled o8B1U00I9g | 4H ised p< z -wivd osejuedleg | So 6
. N oD i! rm hom] [+f
Ho: —— & HH: - : Hu:
aS *pezipisvied mS TS wm): - as *poziqisvied av wo fe aS *peziiseied ov 3
4 5 | sodeqs Jo roqumN Bie % a Slee | mY & | _seseys Jo 1equinn oie = m4 & | sods Jo toquinNn ZG.
5 oe : z S% s ere)
S 5 d Yo : 2, : ® ss Se cmAay Le ate)
x4 TAeom Bato ameeie| ef 4 [Aeon e088 erie [Aaa Raasns
: e444 Jo sosyg| “RA SB ) : 94% JO sesyg| “a 7 g 944 JO seduyg | “AR fe
) i vee * ' A ‘
= ' Jo na o . o ‘
Ss Se yo ae = Ss aS 3 Se
A ~ ASS ra ACs A ‘
yy BSB 4 $4545 ‘oy fey
oy a 92.9 o RRS es) ®
n Rn OOO opt ooo wn n

FIELD WORK—RELEASE OF PARASITES. it

TasLeE XVII.—Release and parasitism of Cerambycobius cyaniceps.

“A? P . “ ” “Ana i cc ” CO Ree AST OLa
: arasite B.”’ Check C.”? Check D.” Area 100 yards Total
area. area. area. across road. mail D.” 3
© n \ o D 1 o n L o |m 1 o n n ) De lis
Fst
Date. ~ sala. 1? jsdla.]P edie |r jessie PP iigea |? sega.
=| es oS |e ene oS jam bee os jw an ov Hm ere os Jam pares oD
of }OS] MR [oF lS] MS] oR SG) WS Oo R/OF| WS oR l|OR| WR | oR [Oe WS
olkal ss | Seales] SiRalss] Sesiss| olkalss] Slyalss
ns Hl ga laFliSslanlazl2alaglas|Shl ag lo BiSelaaloelPates
ro) 2s 7A or l@aloaW@la 2sl|o2\o 2s ola 2a\o eo” |£ aio"
ao |£ael © wo |8ae}o | |8e2)5 |b Blo | |Ga So |e |e alo
s 3 5 S |Isei5 |S |saIs |S BS SSM Wiese esl anes
nD Zz my an IB | m i&@ | im 1 |e |m |4 | m IA |
Sept. 12...... 1205 Os | OnO leony i Olbole eee Secelbeaee Hepes eee | eects 300 1 5083
Released 3
Sept d55ecca- Released 5 9, |
1 | |
(OVO |B Sa ea 95 0 0 | 168 (Oh MOOR Se eee ook ean eealleeellatcse Sede | Seis ., Maras 258} 0} 0.0
OcgivGeetee: 188 0 oO) 125 0 Ol aes See aoe 218} 0/| 0.0} 96; 0} 0.0 | 622] 0} 0.0
Oci al OMessealecotee tera sts slarotall averete Se Seer 247 OM MORON Sores | teeilecrec | Gator See | reser aad 0 | 0.0
|
Taste XVIII.—Parasitism of Eurytoma tylodermatis.
| “ee ”?
“A.” Parasite | “B.” Check | “C.” Check | “D.” Area en ane Total
area. area. area. across road. from <D ” Sn
oO n L oO n 1 o n I oO n joie oO n U o n i
= je |2 |2 8/8 |2 lasl@ 2 Sle & |8)/é |2 [8
Date. * fos] eis ei les erie sie. iesia. |" egle
=| us) r= N| oT SS lace] oF eS ly] oD lH ue) =| Nilo
oe (Sal 8 |S les| GR) Ce SS| HS [SESE He SelSS| Hel Ce lsslhe
Sika ss | SRG SS) SlealSs| SealSs| Salsa] sip alss
BF os) Ba | oF OR Ba | oF eal b a |e eal a ler cal aa) oP okie
mw |8a}| 9 a0 RS) tm |salo | [sails | |8alo tw |8ajo
s |5e5 |s Ble) Nee. ween Sace “la SS eae lie rene
Chen Zi CE Zit |e Ze OSC ee Eilon 91 2ip Pan ace all Za 84
Sept.nl2:-2 =. 120} 0/ 0.0 180 Sa lsGGi eee Sec eheceees Sees pees Sr ese | ese 300 3) 120
Oe Qe 9A 2 | 2:10) 163 0 OM as—5- Sal Sete accel cao eagee See Sera tear 258 ZAOnt
Octibn eas 3 18. 1 54 | 125 1 BOM ere See seas | 218} 2] 0.91) 96 6 |6.25 | 622 | 10 | 1.6
ORs iae Raaahe isin ais anal) 247/20) 10301 eases ee Bipese. [ease 247 | 0 | 0.0
|

III. ParastreE BREEDING Work.

Four new primary parasites of the boll weevil have been bred this
year, and several definite cases of predaceous attack have been
observed. There are now known 11 primary hymenopterous para-
sites, 1 primary dipterous parasite, 2 acarid parasites (at least), 2
coleopterous enemies, and 2 ant enemies which attack the weevil
stages in the cotton forms. They are as follows (insects which
attack the adult weevils are omitted from the consideration):

PRIMARY PARASITES.
Hymenoptera.

Chalcidoidea.
Torymide, Monodontomerinz.
1. Microdontomerus anthonomi Cwid. (Texas).
Eurytomide.
2. Eurytoma tylodermatis Ashm. (Mexico, Texas, Louisiana).
3. Bruchophagus herrere Ashm. (Mexico).
Encyrtide, Eupelmine.
4. Cerambycobius cyaniceps Ashm. (Louisiana, Texas).
Pteromalide, Pteromaline.
5. Catolaccus incertus Ashm. (Texas, Louisiana).

28 PARASITES OF THE COTTON BOLL WEEVIL.

Hymenoptera—Continued.
Ichneumonoidea.
Braconidee, Sigalphine.
6. Sigalphus curculionis Fitch (Texas).
7. Urosigalphus anthonomi Cwid. (Texas, probably Mexico).
8. Urosigalphus schwarzi Cwid. (Guatemala).
Braconine.
9. Bracon mellitor Say (Mexico, Texas).
10. Bracon dorsator Say (Texas).
11. Undetermined braconid (Texas).

Diptera.
Tachinide.
12. Myiophasia xnea Wied. (Texas, Louisiana).
Acarina.

13. Pediculoides ventricosus Newp. (Mexico).
14. Tyroglyphus breviceps Banks (Texas).

PREDACEOUS ENEMIES.
Coleoptera.
Cleride.
15. Hydnocera pubescens Lec. (Texas, Louisiana).
Cucujide.
16. Cathartus cassix Reiche (Texas).
Hymenoptera.
Formicoidea.
Myrmicide, Myrmicine.
17. Solenopsis geminata Fab. (Guatemala, Texas, Louisiana).
Formicide, Formicine.
18. Formica fusca subpolita perpilosa (Mexico).
HYPERPARASITES.
Hymenoptera.
Microdontomerus anthonomi Cwid.
ELurytoma tylodermatis Ashm. Accidental on Bracon mellitor Say.
Cerambycobius cyaniceps Ashm.

POSSIBLE PRIMARY PARASITES.
Hymenoptera.
Catolaccus anthonomi Ashm. (Texas).
Two species of entomogenous fungi, Aspergillus sp. and Cordyceps sp., have been
recorded as probably parasites of boll-weevil larvee.

OCCURRENCE OF SPECIES.

1. Microdontomerus anthonomi Cwfd.* This species was collected as
follows: Goliad, Tex., September 3, 1906, 1 bred from Bracon mellitor
cocoon; Cuero, Tex., August 31, 1906, 1 bred from Bracon mellitor
cocoon; Hallettsville, Tex., August 30, 1906, 4 females, primary para-
sites upon A. grandis; Waco, Tex., August 28, 1906, 2 females, pri-
mary parasites upon A. grandis; Waco, Tex., August 29, 1906, 1
male, 3 females, primary parasites upon A. grandis; Waco, Tex.
August 29, 1906, 1 male, bred from Bracon mellitor cocoon.

@ Crawford, 1907a, b.

OCCURRENCE OF SPECIES. 29

The range of the Microdontomerus is evidently western. ‘T'wenty-
three per cent were secondary parasites.

2. Eurytoma tylodermatis Ashm. This species, while not veryimpor-
tant in any part of the State, is very well distributed. It was recorded
continuously after July 12, 1906. Specimens were bred from mate-
rial collected at Beeville, Brownsville, Calvert, Cuero, Dallas, Goliad,
Hallettsville, Overton, Palestine, Trinity, Victoria, and Waco, Tex.,
and at Mansfield and Many, La. No records were obtained from
Corpus Christi, Corsicana, Junction, Kerrville, Lula, Marshall, Mine-
ola, and Taylor, Tex.; nor from Orange, La. Specimens were sent in
by Prof. A. L. Herrera from Allende, Coahuila, Mexico, with the note
that it was abundant at that place.

3. Bruchophagus herrere Ashm. This parasite has not been taken
in the United States. It was described from Coahuila, Mexico.

4. Cerambycobius cyaniceps Ashm. This parasite has been taken
- at all points investigated except those in the Edwards Plateau region.

5. Catolaccus incertus Ashm. This parasite has not been taken at
Overton or Marshall in northeastern Texas, but is common at all other
points investigated.

6. Sigalphus curculionis Fitch has not been recorded since the first
specimen was bred at Calvert, Tex.

7. Urosigalphus anthonomi Cwfd.t One female was bred from a
cocoon in a weevil cell from material collected September 5 at Browns-
ville, Tex. As the former record wasalso from Brownsville, this species
may be taken to be Mexican in origin.

8. Urosigalphus schwarz. Cwfd.t From 300 squares Mr. E. A.
Schwarz and Mr. H.S. Barber bred 5 females and 1 male of this newly
described species at Cacao, Finca Trece Aguas, Alta Vera Paz, Guate-
mala, during April, 1906.

9. Bracon mellitor Say was bred from all points, except Overton and
Marshall in northeastern Texas. The form known as bracon xan-
thostigma has only been bred from the weevil between September 16
and April 28 in the various years of this investigation. A specimen
of this species is recorded on the Bureau of Entomology files (D. A.
6424 (412)) as bred from the boll weevil at Coahuila, Mexico, Novem-
ber 15, 1902, by Professor Herrera. Specimens were sent in by Pro-
fessor Herrera from Allende, Coahuila, Mexico, during 1906, with the
notes that they were quite common. -

10. Bracon dorsator Say was not bred until October in 1906. It is
only known to occur between August 2 and December 17 in Texas.
It is probably another instance of seasonal dimorphism and identical
with Bracon mellitor.

11. Braconid. One female of this species was bred September 1
from a cocoon in material collected on the ground at Victoria, Tex.
The specimen was lost in transit.

@ Crawford, 1907a.

PARASITES OF THE COTTON BOLL WEEVIL.

30
Pupe of this species within the lar-

12. Myiophasia exnea Wied.
val skin of the weevil were taken from material collected at Many,

La. (3 specimens), and Waco, Tex. (1 specimen); also one specimen
bf

a. ‘cimens),
in imperfect condition was found in a small boll collected at Victoria
Its puparium was within the skin of a weevil larva.

Tex., July 12. It
13. Pediculoides ventricosus Newp. is a common weevil parasite in

Mexico.
14. Mites were frequently found to have destroyed the weevil

stages in fallen squares, especially at Calvert, Tex.
T yroglyphus breviceps Banks was described as a weevil enemy from

Victoria, Tex.

PS
Sea et
anon een Sas
i ' H ' i
' ' ' } H
‘ i t i t H
ae Fem \~-= +: \
' 1 ' icant :
i ! \ 1 t 1 '
So eee sone th ai
(ape FR as
etree ee eres abe
esate nS So Sear
point
aoe ek [erate Mes alone!
Sa (=
i ' J 1 \
Rp ere ee me mare eet OU Lae
i hs ' | , ! 1 1 ‘ '
me REAR NNT ier Gath aM Tey
Se St nae ‘
Hole ewes ga pen tera teed eee
H ~ Nicos ad mm
Saat Tanase
ie CAIN ' i H , Bile 4 ws
Pa ~ Be ee ' ' '
- \ SaaS Re ae foes
ae See \ HH E =
; ‘: ‘. Ht
ea > re
. ; + s S
! \ ae oe i .
! \ ® : \ .
: ine OMENS
: aldara TS
, ~ HHA ua H
} elessrs : .
/ vad <j a. ; — :
— PHO TE a G
. ‘t
He t
tt om j
cr 5 v
t
: ; S
=
it
imed
F
/

Fig. 6.—Occurrence of predominant parasites of the boll weevil in Texas and western Louisiana
e showing-—

||| Occurrence of Catolaccus incertus. = Bracon mellitor. \\ Cerambycobius cyaniceps. _ |||| Eury-

toma tylodermatis. || Limits of predominance of single species. /Limits of two predominant
species. (Original.)

15. Hydnocera pubescens Lec. was found frequently to have killed

Records of this predatory habit are from Browns-

weevil stages.
In a great many other cases

ville and Waco, Tex., and Many, La.

evidence of its predatory nature was less reliable.
16. Cathartus cassix Reiche was found several times to be preda-

ceous in its larval stage upon the weevil
Catolaccus anthonomi Ashm., determined by J. C. Crawford. Four

Y
Ja nth
specimens practically identical in character with the type of this
species were bred from cotton squares collected October 12, 1906, at

Waco, Tex.

OCCURRENCE OF SPECIES.

bl

Although a large quantity of these squares were exam-

ined, no specimens of this species were obtained except from the

general breeding

cage.

The species is known as a parasite of an

estivating weevil (Anthonomus signatus Say), and it is possible that
in the fall the parasites may attack Anthonomus grandis.

GEOGRAPHICAL AND

SEASONAL DISTRIBUTION OF

PARASITES.

Only four of these parasites are at present of any great importance.
In arranging the percentages of each species with regard to the total
number of parasites the following table (Table XIX) has brought
out sufficient data for the production of a map (fig. 6) which shows
the area over which any two of them are predominant.
the entire western half of the area studied Bracon mellitor and Cato-
laccus incertus are the most active, while in the Northeast and East
Cerambycobius cyaniceps shares the importance with the other three
in more or less limited belts:

TasBLE XIX.—The seasonal distribution of the parasites.

Thus, over

Eurytoma | Ceramby-
Bracon | Catolaccus F pra
Total : . tyloderma-}| cobius cy-
uous || mellitor. | incertus. hie: sulesas.
Locality. % Date. ber of > z = 7

para- _| Per- | _| Per- _| Per- | _| Per-

sites. hem cent- pies cent- poe cent- Neu cent-

SLage. * | age. ~ | age. ealkaee-

1906

Beeville Mex. == ence so 5 iTliygl Oe ome mieaeeee 26 8 | 30.7 Dane PA 76) 1 3.8
IDO) Seeme eer ane eae ee AUIS TIS GIS jesse 2 PATON)! | Secs kel MARE sa Geasee boeesel Heese lootbes

DOM CE eisisieste eens Augustils Se <<. 2A | eaters eee fel | ee ae ee a ET Lee oe ee

ID ORS eae hen ake September 3.....-- 6 Gp LOO3O} | Ree RSet | S528 2 eae tes per ere lece ace
Brownsville, Tex......-- Wiijnesemecaseees 43 38 | 88.3 £5 Wt ees PE ees EG sel Semi
ID) Of ea reicrs we ete nice sre IuiliyA28 eyevers avieeeer 119 39 | 32.7 77 | 64.7 or eral pee eeelisase ce

DORM tases ose secd AUPUSE: 3.22 ccc a. 2 | acne Pea ee ee tase chen mee cll uo one toe se ae
DONC s Steen cot eceae September 5..-..--- 155 50 | 32.2 85 | 54.8 1| 0.6 2 1.2

IDO) eR ROME eee September 29....-- 53 12 | 22.6 AQ Glos Al = aca pte elle oe sre ee Sek
Serer, i <>: co ea August 28.......-- 13 3 | 23.0 3 | 23.0 LN Go 4| 30.7
Drees Se Se ame September 13.....-| 21 Boe eo oes acl Gennee acres Meene 4) 19.0
oeus Christi, Tex.-... Jiiby Ose ee eset 39 9 | 23.0 12 | 30.7 LS Peron | retain eee
Corsicana, Tex Se TS ees August 23....-..-- 19 9 | 47.3 4| 21.0 in|) bs Diy be eo eee
1D Xoy testers enwie eee September 18. ...-- 2 TSI ESCO CO) ACR | eae ge ete Pea ie tl (ct
Cuero ext as c.g ske% AUP USE Oe) ieee 13 3 |: 23.0 GAe| alse eee) Lesa 1 7.6
D Omri See asses oe AU SUSU Slee eas 7 45 | 51.7 25 | 28.7 6} 6.8 4 4.5

iD) alles Meee 2 eae Seisre.2 August 29.......-- UD eee eS ace anes Heh See [Bec Sell eetnsaa| ee ee loser
iD Te le om a see September 12......] 21 6 | 28.5 4) 19.0 3 | 14.2 1 4.7

ID) OM Sts hekiccuso-cieece October\2-=23.----- 10 Cal GOLO; Ee ee alee a. 2 20! ON Se eee

ne SSA oe pene October'6.2-22_7- =< 39 19 | 48.4 tS | a) OF 523: ON Eee eee eeeeee

ete a idislele as eerein lous Octoberjl0= =e -= 5 Gy IPA ORO) ects ececel Sree eee ect [Aa
Goliad, TOE we eee INOUE se po ceca ie | ee Seas ee cine ice oc avec | ans Sollee tele
DD Ones ae ate Seen esa September 3......- 108 73 | 67.5 22 | 20.3 3] 2.7 2 1.8
Hallettsville, Tex....... AMSUSU. OS Sales. aie = 15 3 | 20.0 3 | 20.0 2 | 1353 1 6.6
Ope a nss eos tee AUEUSt.30 Ls 5255 2- 39 7 | 17.9 4] 10.2 || te®) 8} 20.4
JUNCHION, Nexo. 55s ose September 24......} 12 10 | 83.3 al ay (el tees ere ee econ ona oe
iKernyvalle, Mex. o.25525.5 ATIGUSt262 255 52.60 6 3 | 50.0 2h GSOROi | Sea se| cee cee eee cae | eee
DOeie tems soe sass e< September 3....... 5 2 | 40.0 UP ZORO)| Eo en Eee ne, | oer
Mansfield, La..........- August 24... 7 3.5.5 47 5 | 10.6 Di 7452 a eae | 36 | 76.5
Many, [ie aap nea on ON IUISG 238 eae ote 109 Galeeot 5: 12 | 11.0 8] 7.3 45 | 41.2
Marshall) exis 2-225. Angus 225-- ee FS | rete | Oe 2 IS ee | Serer Pe ee 6 | 75.0
Mineola,Mex. 25222525. 28 Aust On Seen Ty. arc [LS ete ai a VO) Re eS EL em oreal h oe
Dae secs ocd-e -tisecac October 25.22-<- sec 9 4 | 44.4 7 AY Ao eee Senee a Peciseal Paecos
Orancen(aeeress: saen eee September 30...._- Bison ees ete ae APSO 40 i |Ss ec. | Sansa eeeeer cece
Overton), Nex: .4.- 222-52 /Mpisuist, PRicecacseee (One 2 cel eee eesee Renee 1 | 14.2 2| 28.4
Palestine, Tex 222.2257 Arist lO oe. 70 TPA Alea 10 | 14.2 6} 8.5 73) 1020
Roosevelt, Tex.......... September 24...__- 10 DaleOOV Ol |WoSt ac lle see bees ence eee eel peters
Sixteen miles south of | September 25...... | ee | ener ee | (DONO NER aie Sees eae ens eee

Roosevelt, Tex.

Reavlor Nex oes ses. AUISTISGG Se eee <= 11 SF lee ieen le sneee pecs Quilt VS zis Sere ere eee ata
Trinity, POX toe cckcee ses AtisISt. QS ee 69 A eS 9 | 13.0 | LOST 12} 17.3
DOseste cc euceeneee August 30......... 14 Hb il SMP Sasa) ese naclioece as 2) 14.2

32 PARASITES OF THE COTTON BOLL WEEVIL.

Taste XIX.—The seasonal distribution of the parasites—Continued.

| Bracoueel(Gatolacens Eurytoma | Ceramby-

Total) mellitor. | incertus. |*Y!oderma-| cobius cy-
num- tis. aniceps.
Locality. Date. [srr ———— = =

para- Per- Per- | Per- Per-

sites Ao cent- eats cent- eal cent- Nu cent-

line age age. | ~~ * | age "| age

|
1906. | |

WiACtOnia Bexar ess ase e Afibbarsy Wy es eo aooane Ne Eee (teresa [ae tera Is eee ecto ce later Be cee ena ae
One eere eciee cee see JUNE Zoe see eee 9 | 5 | 55.5 DAN a? 7A ete aol We A eee So
DO ees sock esses JUNG 20 eee sci s itil 2 | 18.0 CR (ea Faye lene | [as aad ee le ea
Dion Beta teeta cee UME 268 See eee 20 10 | 38.4 WN 2639) Wize epee es eel sree coe | eee
1D XC Rea sas Se UME: 2S yn eee Si) |) aly |) 22! 8 | 24.2 1 B30) hese eee ae
DOx Az 4a. sSactecereee JUNCWS ee eee ee DAN eS Soe cael te SN Sa Le De |
DOM ee pee ee liven Pete eee PN este ea| posal be ay st tek Coma fons al Se ee IR a
DOR eee ae tees ully/5 22a peste OE TN Fe ares] Peres [ese ariel eneteeees (esters es |S Ses ee
DOS e ees cee cee tly sO tes eee ee re 13 | Sezai |e 2 ele EM eee ee ees eee
DOueesn Soe sere ces July 852) 222235 32 | 18 |} 56.2 2) 6.2 dW eho Oe ete leet ns
ID NOYE ot As ae en euaianae at Jnalye22n. eee se ee 5 34 OOO | ayer | rape | rctvcees)| eee | eee ne | a
DOS ee ete ce NEE September 1...-... 75 45 | 60.0 3 | 4.0 3} 4.0 a fs ale 3)
WISCOM Meme) fad = ohne Julbyg 20s eects ne | 78] 25 | 32.0 6726 Si isis} 4 5.1
1 Da} ees wee ee INpesuSHE 7/2 = 5 5 ee ee ates} 1 | 20.0 1 | 20.0 ||: 20501)) 5 sees |e eee
Oars ee etn ee eee August 28........- 83 Aa D4. 2 lore cess aco. | S824 25 | 29.8
DO eee se eres Musust2 oe ee | 114] 64] 56.1 11} 9.6 3] 226 12) 10:5
IL) OS er crsiae eee ae September 19...... 36 19 | 52.7 Z|) eD20 8 | 22.2 20} 55.5
DOS Sas sera cakee October l2easesee=- irae) oe (ees oer bee esr ems tee (Mea ft gop el | oe

These figures show very plainly the range of each species and
have therefore been used to form a map of the two predominant
species throughout the State. The centers of the areas predominated
by Catolaccus are Brownsville, Tex., and Orange, La. The influence
of Bracon radiates from Goliad and Corsicana, Tex. The center of
predominance for Cerambycobius is Marshall, Tex. Eurytoma
becomes an active agency at Dallas and Overton, Tex.

BIOLOGICAL NOTES ON THE PARASITES.

1. Microdontomerus anthonomi Cwfd. The Torymide have always
been considered by Dr. William H. Ashmead as parasitic on Diptera.
In addition to the records of parasitism upon the boll weevil, which are
given on a preceding page, one other specimen was bred September
13, 1905, at Mexia, Tex., by F. C. Pratt from Brachytarsus alternatus
Say, breeding in the stems of Sideranthus rubiginosus. The species
is not at all abundant, and has only been collected in cotton forms
between August 28 and September 3. It is perhaps a one-genera-
tion species, although it may have some other and still unknown
host. There is but one individual to each host, and the host may
be in the larval or pupal state when attacked. The sexes occurred
in the proportion of 2 males and 11 females.

In order to ascertain the length of the developmental periods, the
following scheme had to be used: The maximum period from col-
lection to maturity is the nearest approach possible to the total
developmental period, and next to this is the total period in a Bracon
cocoon in case of hyperparasitism. The maximum period from the
observation of the larva to maturity is the nearest approach possible

BIOLOGICAL NOTES ON THE PARASITES. 33

to the total period passed as larva and pupa. The exact length of
the pupal period may be defined as between the maximum period
from observation of the pupa to maturity and the minimum period
from observation of the larva to maturity. Thus it will appear
from the following that the total developmental period is over
twenty-three days, and the pupal stage lasts from six to nine days
during August and September.

TaBLeE XX.—Length of developmental periods in Microdontomerus anthonomi.

In
Bra-
Collec- | Pupa :
tion to Larva to | +o ma-|COnCco Development.
: maturity maturity. turity coon
Locality. Date. to ma-
jturity.
Maxi- | Mini- | Maxi-| Maxi-| Maxi- Larva + |.
mum. |mum.|mum.|mum.|mum. | 7°%!- pupa. Fupa.
1906. Days. | Days.| Days.| Days.| Days.| Days.| Days. | Days.
Cuero; Tex.255 Js. - August 31...... MANS oes Se esis seine aie Oho saacce| Sees el: See
Goliad Tex 22-25) -: September 3... Dies ae alice ate es eee iG} nary. 3 3 le eae or el (Gs
Hallettsville, Tex...) August 30...... 14 9 | 9 | GP ees SS | ah ene eee eV eee 6
Wiaico, Tex Js. 452)" August 28, 29... 10 | 9 | 9 53 (3) eee Se eee ee lee ah de
roe Nai foreras | Neate ral oeisiniavne > arnavelels| ne acai sies 23+ 23 6-9

It was impossible with a lens to find any characters to distinguish
the larvee of this species from any of the other chalcidoids, as all of
the chalcidoid larvee concerned in this report are finely, transversely
lineolate and clad with a few hairs regularly placed in a line around
the middle of each segment. The pupa is nearest in appearance to
that of Cerambycobius. The female pupa is robust, brown until
almost mature, with no darker spots on the dorsal abdominal seg-
ments, but with three longitudinal white lines and a transverse
white line on each segment; the ovipositor is appressed to the dorsum
and proportionately longer than in Cerambycobius. The exuvium
is easily recognized by the form of the ovipositor and its brown color.
(See figure of pupa and of chalcidoid larva, Pl. I, figs. 4, 7.)

A most interesting point in the biology of this parasite is the
fact that a number of individuals were bred from Bracon cocoons.
This is one of those phenomena which are here designated as accidental
secondary parasitism—that is, where a natural primary parasite finds
its host consumed by another parasite and in order to retain its hold
on life is compelled to attack that parasite itself.

10292—Bul. 73—08——3

34 PARASITES OF THE COTTON BOLL WEEVIL.

The breeding records of the parasite are as follows:

TaBLE X XI.—Breeding records of Microdontomerus anthonomi.

As primary As secondary
Total. : F :
Toate Date. parasites. parasites.
Male. | Female. | Male. | Female. | Male. | Female.
1906.

Cuero Dex oi-e se ee eee ANIPUStOl Saco ce sab sae sse Tl ete eal San gs seal erotica 1
Goliad exes sees ese ee SeptemiberiSseeena|a ae soe- | yy ee ee Se eee ee 1
Hallettsville; Texe~ 5. o=. 9 August 30....-.-- Ee eet oe (PPE Alin oe
ACOMMERA acs e a ees August 28....-..- eer Ieee | PA es, BES AE Dye ws sa aealeeee neue
DOR scsecaask ee nee = August 29.......-. 2 3 1 3 Lal. aosoneee
2 11 1 9 1 2

Thirteen parasites were bred, of which 23 per cent were secondary.
Owing to numerous expressions of incredulity concerning this dual
parasitism it may be proper to state the proofs. In the first place,
primary parasitism has been thoroughly established in the case of
this species and also in the case of Eurytoma tylodermatis and Ceram-
bycobius cyaniceps. Adults have been bred from larvee which were
actually observed to be feeding upon the weevil larve. The devel-
opmental period of those adults bred precludes any arguments that
they were bred from unobserved eggs of parasites on the larva. In
the second place, the Bracon and chalcidoid larve are very easily
distinguished, so that the notes in most cases stated whether the
larva observed was a Bracon or a chalcidoid. In no case was one of
‘the latter bred from a Bracon except when the Bracon cocoon was
already formed at the time of the observation. Finally, secondary
parasitism was proved because the parasites were actually bred from
isolated Bracon cocoons, which were subsequently opened and from
which the known exuvia of the secondary parasites were removed.

2. Eurytoma tylodermatis Ashm. This species was bred by Town-
send in 1895 and by Mally during several years from the boll weevil.
Ashmead described it as a parasite of Tyloderma foveolatum Say
(Ashmead, 1896), a weevil breeding in the stems of Onagra biennis
and also in a species of Epilobium. It was reared by F. H. Chitten-
den from the larva of Tyloderma foveolatum Say in stems of Onagra
biennis, September 23, from material collected at Rosslyn, Va.

This species was bred by the writer from Anthonomus disjunctus
Lec. breeding in the heads of Heterotheca subaxillaris at Jacksonville,
Tex., October 13, 1905 (1 male); from Anthonomus squamosus Lec.,
breeding in the heads of (rindelia squarrosa nuda at Clarendon, Tex.,
September 22, 1905 (6 females), and September 26, 1905 (2 males);
from Orthoris crotchii Lec. breeding in the seed pods of Mentzelia nuda
at Clarendon, Tex., October 2, 1905 (1 male); and from Lixus mus-
culus Say, which forms galls in the stems of Polygonum pennsylvani-
cum, at Clarendon, Tex., October 2, 1905 (2 females), and October 17,
1905 (1 male). Mr. W. W. Yothers bred this Eurytoma from Lirus

Bul. 73, Bureau of Entomology, U S, Dept. of Agriculture, PLATE ],

PARASITES OF WEEVILS.

Fig. 1.—Eurytoma tylodermatis, pupa.
bycobius cyaniceps, pupa. Fig. 4,—
Bracon. Fig. 6.—Bracon mellitor,
(Original. )

Fig. 2.—Catolaccus incertus, pupa. Fig. 3.—Ceram-
Microdontomerus anthonomi, pupa. Fig. 5.—Larva of
pupa. Fig. 7.—Larva of chalcidoid. Much enlarged.

4 “ap
‘7

Le

BIOLOGICAL NOTES ON THE PARASITES. 35

scrobicollis, which breeds in great abundance in the stems of the com-
mon road weeds Ambrosia trifida and A. psilostachya, at Victoria,Tex.,
April 5, 1905 (1 female). In a personal letter dated August 30, 1906,
Dr. Wm. H. Ashmead writes of this species: ‘“‘ Hurytoma tylodermatis
Ashm. is another similar case [referring to his remarks on Ceramby-
cobius, given later]. I have had several species of Eurytoma bred
by Hopkins, Marlatt, etc., from beetles, and I can only quote their
records. The genus seems to be primary and secondary as well as
phytophagous, unless we can find characters to still further subdivide
it generically.”’

This parasite is quite abundant throughout the season and is a con-
tinuous breeder. The number of generations is probably little less
than that of the weevil. There is but one individual to each host, and
the latter may be in the larval or pupal state when attacked. The
sexes are in the proportion of 29.1 per cent male and 70.9 per cent
female. The females are very much larger than the males.

Proceeding in the same manner as for the preceding species it is
ascertained that the total developmental period is over 12 to 15 days,
that the larval plus pupal period is over 11 to 14 days, and that the
pupal period alone is at the minimum 6 days and maximum 13 days.
Judging from this last period, for which the figures are definite, it
may be safely said that the estimates for the larval and egg periods
are very low. The increase in the length of the pupal period in
October is noticeable.

The data upon the biology of Eurytoma tylodermatis are compara-
tively continuous from July 18 to October 19. In studying the length
of the various stages the same methods were used as outlined for
Microdontomerus. The only stage which could be definitely limited
as to the period of development was the pupal stage, which was found
to last from 7 to 8 days in the latter half of July, 6 to 9 days during
August, 7 to 9 days in September, and 11 to 13 days in October.

The larvee of Eurytoma were described in the notes as smooth,
transversely lineolate, and with wrinklings at the sutures. The
pup are white, pink-eyed, turning black toward maturity. They
may be easily distinguished, by the lateral compression of the abdo-
men and the straight venter, from the pup of Catolaccus with their
dorso-ventral compression and obtusely angulated venter. The
exuvium is cast in parts and is never found intact as in the case of the
other chalcidoids studied; the remains are yellowish transparent.
(See figure of pupa, Pl. I, fig. 1.)

Secondary parasitism: One male was bred from material collected
August 9 at Trinity, Tex., and two males from material collected
September 19 at Waco, Tex., as secondary parasites in the cocoons
of Bracon mellitor. In each case the cocoon was isolated when first

36 PARASITES OF THE corron’ BOLL WEEVIL.

found, so that no doubt could be expressed as to the source of the
parasites. Corroboration was obtained by examining the cocoons.

3. Bruchophagus herrere Ashm. Considerable doubt has been raised
by very eminent parasitologists as to the actual parasitism of the
boll weevil by this species and some doubt has been raised concerning
its generic location. No subsequent corroboration has been obtained
to prove parasitism of the boll weevil.

4. Cerambycobius cyanicepsAshm. The first published record of the
parasitism of the boll weevil by this species is found in Mally’s Report
(Mally, 1902), in which that writer states that he had bred it since
1899 at Austin, Tex. Only four records were obtained in previous
years by the boll-weevil laboratory of the Bureau of Entomology; 1
female bred July 7 at Calvert, Tex., by G. H. Harris; 1 male and 1
female bred July 25 at Victoria, Tex., by W. E. Hinds, and 1 female
bred March 11 at Corsicana, Tex., by C. M. Walker.

Dr. F. H. Chittenden has reared it from the larva of Tyloderma
foveolatum breeding in the stems of Onagra biennis, September 11,
from material collected at Chevy Chase, Md. He has also reared it
from Bruchus eriguus Horn (Chittenden, 1893b). In a personal letter
dated August 30, 1906, Dr. Wm. H. Ashmead said of this species:
“Tt is not rare, has been bred from cerambycids and other beetles,
and I have always supposed it to be a primary parasite. All of these
beetles, however, have braconid parasites, and it may yet prove to
be a secondary parasite as other of the eupelmines are said to be.”

One female was bred by the writer October 17, 1905, as a probable
parasite of Lizus musculus Say, breeding in stem galls of Polygonum
pennsylvanicum collected September 19 at Clarendon, Tex. On Octo-
ber 26, 1906, a female was bred from a Lixvus musculus pupa collected
in the same plant at Dallas, Tex., opposite the farm on which release
experiments were conducted. On September 11 and 12, 1906, 2
females were bred from Anthonomus albopilosus breeding in seed of
Croton engelmanni collected August 26, 1906, at Johnsons Bayou,
La., by J. D. Mitchell. On October 30, 1906, 1 female was bred from
Trichobaris terana found breeding in stalks of Solanum rostratum
by R. A. Cushman and the writer, along the road close to the plat
upon which parasites were released at Dallas.

This parasite is very abundant in eastern Texas, diminishing to
the Southeast, and is entirely absent at Brownsville and through
western Texas. It is a continuous breeder, attacking the weevil
larvee and pupe, one individual for each host. The sexes are in the
proportion 19.7 per cent males and 80.3 per cent females. The
females are several times longer than the males. -

The developmental periods for Cerambycobius cyaniceps have been
worked out from data covering the period between July 12 and Octo-
ber 4. The pupal period in July is 10 days or less, in August is from

BIOLOGICAL NOTES ON THE PARASITES. 37

8 to 11 days, and in September from 9 to 12 days. During August it
was ascertained that the entire developmental period exceeds 26 days.

The larve of Cerambycobius are described in the breeding notes
as white, transversely lineolate, with a row of prominent hairs around
the middle of each segment. The pupe are elongate brown, with a
short, obtuse, erect ovipositor, and with the dorsum of the abdomen
marked with two dark spots on each segment. The brown exuvium
is easily recognized from the ovipositor. The pupxz may be easily
distinguished from those of Microdontomerus after a little practice.
(See figure of pupa, Pl. I, fig. 3.)

Secondary parasitism: On September 14 a female Cerambyco-
bius was bred from a larva which was found feeding on a Catolaccus
pupa in material collected August 28 at Calvert. Although numer-
ous cases of parasitism of Bracon mellitor were found, only 11.5 per
cent of this species were secondary parasites. It was found as a
secondary parasite on Bracon as follows:

Beeville, Tex., July 12, 1 female.

Corpus Christi, Tex., July 10, 1 female.
Cuero, Tex., August 31, 1 female.

Mansfield, La., August 24, 2 females.
Trinity, Tex., August 9, 1 male, 1 female.
Victoria, Tex., September 1, 1 female.

Waco, Tex., August 28, 3 males, 10 females.
Waco, Tex., September 19, 1 male, 2 females.

In one case the attack of the Bracon is known to have been prior to
spinning.

5. Catolaccus incertus Ashm. This species has been one of the most
important species mentioned in the literature of the boll weevil.
It was described as a primary parasite of Anthonomus signatus
(Chittenden, 1893a). It was reared by F. H. Chittenden from Apion
decoloratum Sm. breeding in Meibomia paniculata at Washington,
D. C., from Apion griseum Sm. breeding in Phaseolus perennis at
Washington, D.C., and from Bruchus exiguus (Chittenden, 1893b).

Two females were bred by the writer October 16, 1905, from Antho-
nomus disjyunctus breeding in the heads of Heterotheca subaxilaris
collected at Jacksonville, Tex. From Anthonomus eugeni breeding
in peppers at San Antonio, Tex., F.C. Pratt bred 3 females on
October 18, 1905. From <Anthonomus albopilosus breeding in the
seed of Croton engelmanni at Johnsons Bayou, La., the writer bred
2 females on September 6, 1906, and from the same weevil breeding
in Croton capitatus at Leesville, La., the writer bred 1 female on Sep-
tember 29, and subsequently 3 females and 3 males.

Catolaccus incertus is very common in all parts of the State and is
well established as a boll weevil parasite. (See Plate III, #.) It is
a continuous breeder, attacking the weevil larve and pup, one

38 PARASITES OF THE COTTON BOLL WEEVIL.

individual for each host. The developmental period seems to be
considerably shorter than in either of the species preceding. The
data upon which the following figures are based cover the period
from June 25 to October 28. During June and July the total develop-
mental period is known to be over 11 or 12 days, during August it is
over 14 days, and during September and October over 18 days.
The pupal period in June and July is only 4 to 6 days, in August 6
to 7 days, in September 5 to 9 days, but in October 13 to 15 days.

The larve of Catolaccus were described in the notes as having
short hairs in a row on each segment. The pup are white to yellow,
the thorax turning black first as they approach maturity. The
abdomen is very flat below and bent at an obtuse angle to the thorax.
The head is very broad, and the eyes are pink. The exuvium is
yellowish and resembles a little pointed cap. (See figure of pupa,
Plea pie .2-

6. Sigalphus curculionis Fitch. No corroboration has been recorded
of the parasitism of the boll weevil by this species. This parasite
is very commonly bred from the plum curculio (Conotrachelus nenu-
phar Hbst.). Chittenden also records it as a primary parasite of
Trichobaris trinotata Say, which breeds in the stems of Solanum
rostratum (Chittenden, 1902).

7. Urosigalphus anthonomi. Cwfd. This parasite was again bred
September 20 from material collected September 5 at Brownsville.
The period in the cocoon was at least 9 days. The cocoon with
weevil larva’s head attached was half in a cotton seed in a boll. It
was very much finer meshed than that of Bracon mellitor and easily
broken. In addition to this species and (8) Urosigalphus schwarzi
Cwfd., both bred from Anthonomus grandis, Urosigalphus armatus
Ashm. has been bred from Balaninus, and another species ( Urosigalphus
bruchi Cwfd.) from Bruchus prosopis Lec. taken at Harlingen, Tex.,
in beans of Prosopis glandulosa. Hence it is very probable that the
species of this genus are normally weevil parasites.

9,10. Bracon mellitor Say. Bracon dorsator Say. Bracon vantho-
stigma Cress.

These are all forms of a single very variable species, of which the
entirely red form holds the name B. mellitor, that with black sternum
and metathorax the name B. xanthostigma, and that with the thorax
almost entirely black and the vertex of the head also black, being
also smaller in size, has been known as B. dorsator.

A peculiar record is furnished by F. H. Chittenden who reared this
species from the strawberry leaf-roller Ancylis (Phoxopteris) comp-
tana Frol., from material collected at Cabin John, Md., July 9, 1899.

In a personal letter dated August 30, 1906, Dr. Wm. H. Ash-
mead writes: ‘‘ Bracon mellitor Say is avon ic a most important
parasite, widely distributed in the United States, and must affect

BIOLOGICAL NOTES ON THE PARASITES. 39

very many different beetles, although usually rhynchophorous
beetles.”

The typical form has been bred from Anthonomus grandis at. all
times and in all places between June 15 and October 4. The form
xanthostigma has only been bred from the boll weevil between Sep-
tember 16 and April 28, while dorsator was bred August 2, 1903, at
Victoria; November 20, 1895, at Goliad, and December 17, 1895, at
Beeville. B. mellitor was bred June 19, 1905, at Dallas by W. W.
Yothers from Anthonomus fulvus Lec., which breeds in the buds of
Callirrhoe involucrata, and on June 14 he bred dorsator from the same
weevil. On September 20, 1905, F. C. Pratt bred male and female
dorsator and female mellitor from Desmoris scapalis Lec., breeding in
the flower heads of Sideranthus rubiginosus at Mexia, Tex.; on Sep-
tember 23 he bred the typical form from the same weevil collected
at Calvert, Tex.; on September 29 both the typical form and dorsator
from Mexia material; and on October 27 he again bred the typical
form from Mexia material. On September 30, 1905, the writer bred
mellitor and xanthostigma from Anthonomus squamosus Lec., breeding
in the heads of Grindelia squarrosa nuda at Clarendon, Tex.; on
October 2, 4 of the typical form and 1 xanthostigma were bred; on
October 3, 3 typical, and finally on October 16 a xanthostigma, were
bred. On October 18 and 23 dorsator was bred by F. C. Pratt from
Anthonomus eugeni Cano, breeding in peppers at San Antonio, Tex.
On September 6, 1906, J. D. Mitchell bred a typical form from Antho-
nomus albomlosus Dietz, breeding in seed of Croton capitatus at
Victoria, Tex.

The data upon the life-cycle of Bracon mellitor cover the period
from June 25 to November 15. The entire developmental period
covers at least 21 days in June and 33 in October, but probably con-
siderably more. The period within the cocoon has been limited to
between 5 and 6 days in June, to 7 days in July, 6 to 11 days in
August, 3 to 7 days in September, and 15 to 27 days in October.
The minimum of 3 days in September is based upon an observa-
tion of the spinning of the cocoon and the time of maturity, and the
record of 5 days in June was obtained in the same manner.

The larve of Bracon can be immediately separated by the absence
of hairs and by the body being punctate instead of lineolate. The
pupe are yellowish, loosely constructed, with the appendages very
fragile and inclosed in a stout one-meshed silken cocoon, which varies
from pure white to golden yellow or dark brown. In several cases
where particular note was made of the brown color of the cocoon
hyperparasites were bred, but this was not a constant indication.
The species varied greatly in size, depending upon the amount of
food available. (See figures of larva and pupa, PI. I, figs. 5, 6.)

40 PARASITES OF THE COTTON BOLL WEEVIL.

No dark forms of this species were bred until September 10, 1906,
except a single 6. dorsator which was bred from Cuero material col-
lected August 31. The rest were all bred from material collected
at Brownsville, September 29; Dallas, October 2, 6, 10; and Waco,
October 12. The record of all Bracon bred after October 10 is as
follows:

October 10, Bracon mellitor, 3 males, 5 females.

ae Do. 1 male.

12: Do. 1 male.

113}, Do. 1 female; mellitor xanthostigma, 2 females.
15, Do. 1 female.

18, Bracon mellitor xanthostigma, 3 females.

20, Do. 2 females.

22, Bracon mellitor, 3 females; mellitor xanthostigma, 2 females; mel-
litor dorsator, 2 males, 1 female.
24, Bracon m. xanthostigma, 1 female.

Die Do. 2 females; m. dorsator, 1 female.
ol, Do. 2 females.
November 1, Do. 3 females; m. dorsator, 1 male.
5, Do. 1 female.

6, Bracon mellitor dorsator, 1 female.
14, Bracon mellitor canthostigma, 1 female.

11. An undetermined braconid of the size of Bracon mellitor, but
with the base of the abdomen black and belonging to a different sub-
family, was bred September 17 from a yellow cocoon in a fallen
square collected at Victoria, Tex., sabe 1. Unfortunately this
was lost in the mails.

12. Myvophasia aznea Wied. ‘This species is recorded as a parasite
of Balaninus nasicus Say, Chalcodermus sp., Conotrachelus juglandis
Lec., and Sphenophorus parvulus Gyll. (Coquillett, 1897), and also of
Ampeloglypter sesostris Lec. (Aldrich, 1905). One specimen was bred
July 6 by the writer from a larva of Conotrachelus affinis Boh. found
in a hickory nut collected June 8 at Logansport, La. As parasites
of the boll weevil from fallen squares collected at Many, La., August
23, 1 male was bred September 5 and 1 female September 8, while
1 failed to mature. From a hanging square collected at Waco,
Tex., August 29, a puparium was obtained but the fly not bred.
From a small boll collected at Victoria, Tex., July 12, a deformed
fly was bred.

Parasitism by this species can always be positively proved if the
puparium is found, as it is inside of the stretched skin of the weevil
larva. The affected larva is of a tawny parchment color and shows
the projections of the appendages of the fly puparium.

The developmental period is undoubtedly in excess of 28 days, as
shown by the first record above.

13. Several attempts have been made to introduce Pediculoides
ventricosus Newp. into Texas as a parasite of the boll weevil, but.

BIOLOGICAL NOTES ON THE PARASITES. 41

with no success. Much has been written upon this mite by Professor
Herrera and his staff.

14. There are, however, native mites, including T'yroglyphus
breviceps Banks (Banks, 1906, p. 17), which in some cases are a con-
siderable factor in the destruction of the immature weevils. At
Calvert, Tex., in moist fallen squares collected September 13, 11 out
of 18 cases of parasitism were mites. 7. breviceps was described from
Victoria.

15. Hydnocera pubescens Lec. This clerid was found to be pre-
daceous in its larval stage upon the boll weevil and all of its parasites.
The egg is probably inserted into the cell of the weevil, and the
young predator immediately starts its attack, consuming whatever
it finds in the way of insects, and finally spins a loosely meshed,
single-layer, silken cocoon, and pupates therein. From data of mate-
rial collected at Waco, Tex., August 28, an approximation of the
developmental periods was obtained. The longest period from col-
lection to spinning was 34 days, the period in the cocoon was 12
days, the adult being formed 3 days before leaving the cocoon. In
order to prove the predaceous habit live weevil pupze were fur-
nished individual clerid larve, and in most cases were completely
consumed. In several cases large larve were found in Bracon
cocoons which were punctured with holes large enough only for the
‘very smallest of the larve.

16. In a number of cases, material collected at Victoria, Dallas,
and Brownsville gave evidence of a predaceous attack upon weevil
stages by Cathartus cassix Reiche.

IV. THe SoOuRCES OF THE PARASITES.

In the preceding sections it has been pointed out that the para-
sites of Anthonomus grandis are also common to various other spe-
cies of weevils. There is a more or less clearly defined theory in
parasitology that the parasites of a particular genus or tribe are con-
fined to the insects of a particular family or order. With this idea
in mind a list of the known parasites of the Rhynchophora has been
compiled, in order to show the characteristic groups and the possible
sources of boll-weevil parasites.

PARASITES KNOWN TO ATTACK RHYNCHOPHORA.

Parasite. Host.
Fungi.
PAS PORGMULUs Spm te ee nas, Soe 2S Se bs Anthonomus grandis Boh.
COnincepaspe aston foo 28 oo ve ocd Anthonomus grandis Boh.
Empusa (Entomophthora) sphxrosperma..Phytonomus punctatus Fab.
Entomophthora phytonomi...........---- Phytonomus punctatus Fab.

Sporotrichum globuliferum ......---.----- Epicxrus imbricatus Say.

49 PARASITES OF THE COTTON BOLL WEEVIL.

Parasite. Host.
Acarina.
Tarsonemide.
Pediculoides ventricosus Newp ......-----4 Anthonomus eugeni Cano.
Anthonomus grandis Boh.
Tyroglyphidee.
Tyroglyphus breviceps Banks .........-- 4 Anthonomus grandis Boh.
(ii Re RRR ae Late Fa Ey Sr era Anthonomus eugenvi Cano.
Diptera.
Dexide.
Metoderin-basatis ra We 32 eee tae ee Conotrachelus juglandis Lec.
Tachinidee. ,
Myiophasia xnea Wied...........----.- . Ampeloglypter sesostris Lec.

Myiophasia robusta Coq
Hymenoptera.
Proctotrypoidea.

Platygasteride. Platygasterine.

Trichacis rufipes Ashm
Chalcidoidea.

Torymide. Monodontomerine.

Microdontomerus anthonomi Cwid

Eurytomide. Eurytomini.
Eurytoma magdalidis Ashm ............-

Eurytoma tylodermatis Ashm

Bruchophagus herrere Ashm..........---
Cleonymidz. Cleonymine.

Cheiropachys colon L ...

Encyrtide. Eupelmine. Eupel-

mini.
Cerambycobius cyaniceps

IAtghinivees so eee

Pteromalide. Pteromaline. Meta-

ponini.

Bruchobius laticollis Ashm

Pteromalidz. Pteromalinz. Rhaph-

itelini.

Anthonomus grandis Boh.
Balaninus nasicus Say.
Chalcodermus xneus Boh.
Conotrachelus affinis Boh.
Conotrachelus juglandis Lec.
Sphenophorus parvulus Gyll.
Sphenophorus robustus Horn.

Brachytarsus alternatus Say.

Magdalis armicollis Say.
Anthonomus disjunctus Lec.
Anthonomus grandis Boh.
Anthonomus squamosus Lec.
Apion segnipes.@

Lixus musculus Say.

Lixus scrobicollis Boh.
Orthoris crotchu Lec.
Tyloderma foveolatum Say.
Anthonomus grandis Boh.

Magdalis xnescens Lec.

Anthonomus grandis Boh.
Anthonomus albopilosus Dietz.
Bruchus exiguus Horn.@

Lixus musculus Say.
Trichobaris texana Lec.
Tyloderma foveolatum Say.

Bruchus pisorum L.

@Qn the authority of F. H. Chittenden.

.

PARASITES KNOWN TO ATTACK RHYNCHOPHORA.

Parasite. Host.
Chalcidoidea—Continued.
DIAGLUS BDow ied soetis no dee hee ceielaeeeee Magdalis xnescens Lec.
Pteromalide Pteromalinz. Ptero-
malini.
Catolaccus anthonomi Ashm.......------ Anthonomus signatus Say.
Catolaccus celiodis Ashm......--.----- -2 Acanthoscelis acephalus Say.
Catolaccus incertus Ashm ....--.....--- 2 Anthonomus xneolus Dietz.

Anthonomus eugenii Cano.

Anthonomus albopilosus Dietz.

Anthonomus disjunctus Lec.
Anthonomus fulvus Lec.
Anthonomus grandis Boh.
Anthonomus nigrinus Boh.
Anthonomus signatus Say.
Apion decoloratum Sm. @
Apion griseum Sm. 4
Auletes tenuipes Lec.
Bruchus exiqguus Horn.4@
Zyqobaris xanthoxyli Pierce.
Neocatolaccus tyloderme Ashm.......-.... Lizus parcus Lec.
Lixus musculus Say.
Tyloderma foveolatum Say.
Pteromalide. Spalangiine.
Cerocephala pityophthori Ashm ...... . ..- Pityophthorus consimilis Lec.
Cerocephala scolytivora Ashm.........---! A scolytid.
Eulophide. Entedonine. Om-
_ phalini.
Omphale tanda Ashi 3.020555 222220223. Ceutorhynchus rape Gyll.
Eulophide. Entedonine. Ente-
donini.
Asecodes alintarsis Ashm....:..2...:..2.1 Magdalis xnescens Lec.
Entedon lithocolletidis Ashm .......---.-- 2 Anthonomus nigrinus Boh.
Eulophide. Tetrastichine. Te-
trastichin1.
FUE SATE IEG es A aS sh Orthoris crotchii Lec.
Mymaride. Mymarine. Ana-
phini (egg parasites).

Anaphes conotracheli Girault ..........-- Conotrachelus nenuphar Hbst.

Ichneumonoidea.
Ichneumonidz. Ophioninze. Pori-
zonini.

Porizon conotracheli Riley...........--- Conotrachelus nenuphar Herbst.

Braconidz. Blacinz. Calyptini.
Calypinistibiator’ Gressc- 52 aes 5285. 2. 24 Anthonomus signatus Say.
Braconide. Sigalphine.
Sigalphus canadensis Prov.....--.-.-----2 Anthonomus scutellatus Gyll.
Podapion gallicola Riley.?

Sigalphus copturi Riley ms..........-- Cylindrocopturus longulus Lec.

Conotrachelus posticatus Boh.

@On the authority of F. H. Chittenden. b Breed together.

43

44 PARASITES OF THE COTTON BOLL WEEVIL.

Parasite. Host.
Tchneumonoidea—Continued.
Sigalphus curculionis Fitch.....-.------ Anthonomus grandis Boh.

Conotrachelus juglandis? in nuts.
Conotrachelus nenuphar Hbst.
Trichobaris trinotata Say.

ISG OLDRUS Bs Bae Zoic we. alere terme Chalcodermus xneus Boh.
Sigalphus zygobaridis Cwid...-- - oe ee Zygobaris xanthoxyli Pierce.
Urosigalphus anthonomi Owid.....-.---- Anthonomus grandis Boh. (Texas).
Urosigalphus armatus Ashm.....-.-.-.--- Balaninus spp., Conotrachelus spp.
Urosigalphus bruchi Cwid.....-.------- Bruchus prosopis Lec.
Urosigalphus schwarzt Cwid.......------+ Anthonomus grandis Boh. (Guatemala).

Braconide. Chelonine.
Phanerotoma tibialis Hald...........--- Anthonomus nigrinus Boh.

Braconide. Agathidinz. Micro-

dini.

Microdus simillimus Cress.....--------- Lixus scrobicollis Boh.

Braconide. Braconine. Braco-

nini.
Glyptomorpha liai Ashm.....--.-.------ Lizxus scrobicollis Boh.
Glyptomorpha mavaritus Cress...-.--.-- Lixus scrobicollis Boh.
Glyptomorpha novitus Cress....--------- Tixus musculus Say.
Glyptomorpha rugator Say..--.--------- Lixus concavus Say.
Lizus musculus Say.

Vipiobelfrage: Gress: oo. yn e8 eh = oe = Lizxus scrobicollis Boh.
Melanobracon simplex Cress...-------. -- Dendroctonus picexperda Hopk.
Microbracon nwperus Cress.....--------- Orthoris crotchii Lec.
Bracon analcidis Ashm...-.--- - Cotte ee Tyloderma fragarix Riley.
Bracon anthonomi Ashm.......-.------- Anthonomus signatus Say.
Bracom. elUlor Say <2 hos as Se oe Cea Anthonomus eugenti Cano.

Anthonomus albopilosus Dietz.
Anthonomus fulvus Lec.
Anthonomus grandis Boh.
Anthonomus squamosus Lec.
Desmoris scapalis Lec.

Braco DissOdis: NSO e eet os eet a Pissodes strobi Peck.
Bracon rhyssemati Ashm. ms.....-.----- Rhyssematus lineaticollis Say.
Bracon smicronygis Ashm. ms...-------- Smicronyx tychoides Lec.
BT OCOTe Bie a ee cc eee ae eee Tomicus pint Say.
BRACOR ISDS tat = el ee onee eee cee Brachytarsus limbatus Say.
Braton: Spa one Skee tes os eae eee Baris sp.
Braconidze. Rhogadinz. Rhyssa-
lini.
Rhyssalus pityophthori Ashm.....--.---- Pityophthorus sp.
Braconidze. Spathiinse. Spathiini.
Spathius canadensis Ashm...-.---------- Dryocetes autographus Ratz.

(?) Magdalis olyra.
Phleosinus graniger Chap.
Tomicus sp.

BIOLOGIES OF THE WEEVILS CONTRIBUTING PARASITES.

It is of considerable importance in framing means of combating
the boll weevil with parasites that the weevils usually found in the
vicinity of cotton fields should be thoroughly known. In order that

BIOLOGIES OF WEEVILS CONTRIBUTING PARASITES. 45

the conclusions in this regard may be more thoroughly understood,
this subject must be introduced by a series of brief biologies of all +
the weevil species which have contributed parasites to the attack of»
the boll weevil.

CURCULIONID#A. APIONINZE.

1. Apion decoloratum Sm. breeds in Meibomia paniculata. It is
parasitized at Washington, D. C., by Catolaccus incertus.

2. Apion griseum Sm. breeds in Phaseolus perennis, and is para-
sitized by the same species as the preceding.

CURCULIONIN. CLEONINI.

3. Lixus musculus Say makes a gall in the stems of Polygonum
pennsylvanicum. The larve and pupe are found in these galls. The
weevil and its host plant are typical of tne fall, the plant being
found in low moist ground, frequently in close proximity to cotton
fields. The weevil is attacked by two of the important boll-weevil
parasites. .

4. Iixus scrohicollis Say dwells in the stems of Ambrosia trifida.
and psilostachya, two of the commonest roadside and waste-place
weeds throughout the country in summer and fall. The entire stem
is riddled by the weevil larvee, which pupate in cells of frass at the
end of their burrows. Eurytoma tylodermatis and other parasites not
concerned in the discussion have been bred from this weevil.

ERIRHININI.

5. Desmoris scapalis Lec. breeds in the seed heads of Sideranthus
rubiginosus, a summer weed, which is very abundant in some localities
along roadsides and on prairies. The larve are expelled with the
seed and enter the ground for pupation, maturing in the following
spring. This species is parasitized by Bracon mellitor, which has a
much more rapid development than itshost. (See Pl. III, B, D, F,G.)

ANTHONOMINI.

6. Anthonomus (Anthonomorphus) fulvus Lec. breeds in the buds
of Callirrhoe vnvolucrata, an early spring mallow, which is common
in May and June on moist meadows. The larve feed upon the floral
column of the bud or the imperfectly opened flower and pupate in a
cell of excreta in the capsule or fallen corolla. Bracon mellitor and
Catolaccus incertus have been bred from it. (See Pl. II.)

7. Anthonomus signatus Say, the strawberry weevil, breeds in a
number of rosaceous plants, such as strawberry, blackberry, raspberry,
and wild rose, as also in red-bud. It is an early spring weevil, ovi-
positing in the buds, which soon drop to the ground, where the larva
transforms to the pupastage. It is parasitized by Catolaccus incertus.

46 PARASITES OF THE COTTON BOLL WEEVIL.

8. Anthonomus nigrinus Say is an eastern weevil which breeds in
the buds of Solanum carolinense and various other solanaceous plants,
occurring principally in the spring and early summer, but not con-
fined to one generation. It oviposits in the buds, causing them to
fall. The larva makes its cell in the center of the bud through the
pistil and all of the stamens and pupates in this cell. It is a host of
Catolaccus incertus.

9. Anthonomus albopilosus Dietz is a fall species which breeds in
the seed of Croton capitatus and C. engelmanmi, weeds which are very
abundant in pastures. The larve eat out one seed and then enter a
second and form their pupal cell in this. The weevil serves as a host
to Cerambycobius cyaniceps, Catolaccus incertus, and Bracon mellitor.

10. Anthonomus xneolus Dietz is a spring species which breeds in
the buds of Solanum torreyi, S. rostratum, and S. eleagnifolium. The
larvee feed within one or two anthers, in the latter case forming a cell
which cements the two together. In many cases the flower does not
fall, and is able to fruit. The weevil is a host of Catolaccus incertus.

11. Anthonomus eugeni Cano (xneotinctus Champ.), the pepper
weevil, is a recent introduction from Mexico. It is a fall species,
which breeds in the interior of the cultivated peppers. (See Plate III;
A, C.) Itisa host to Catolaccus incertus, Bracon mellitor, and Pedi-
culoides ventricosus, as determined by Professor Herrera. Anthonomus
mexicanus Boh., another pepper weevil, is very likely identical.

12. Anthonomus squamosus Lec. breeds in the heads of (rindelia

squarrosa, a fall plains plant, which is very abundant in semiarid
Texas. The weevil larva makes a cell among the seed, which is
formed of the hardened. gum of the flower and excreta. It has
probably only one generation. It is very highly parasitized and
principally by Bracon mellitor and Hurytoma tylodermatis.
13. Anthonomus disjunctus Lec. breeds in the heads of Heterotheca
subaxillaris, a fall plant, which is very abundant along roads, waste
places, and on meadows. The weevil larva makes a small cell, formed
of the hardened gum and excreta of the weevil, among the seed.
There is but one generation. Hurytoma tylodermatis and Catolaccus
incertus have both been bred from this weevil.

CRYPTORHYNCHINI.

14. Conotrachelus affinis Boh., the hickory nut weevil, has been
found to be the host of Myiophasia xnea. It is an early. summer
weevil. Pupation is in the ground.

15. Conotrachelus nenuphar Herbst, the plum and peach curculio,
breeds in the fleshy part of the fruit and pupates in the ground. It
is the host of Sigalphus curculionis, which has once been bred from
Anthonomus grandis.

PLATE Il.

Bul. 73, Bureau of Entomology, U. S. Dept. of Agriculture

3
;
a

THE PURPLE MALLOW (CALLIRHOE INVOLUCRATA), A FOOD PLANT OF Two

WEEVILS RELATED TO THE BOLL WEEVIL.

Fig. A.—The buds and flowers, which are subject to attack by Anthonomus fulvus, and

the capsule, subject to attack of Macrorhoptus estriatus. Fig. B.—Bud infested by

Anthonomus fulvus. Fig. C.—Flower injured by adult weevil of Anthonomus fulvus,
(Original. )

Bul. 73, Bureau of Entomology, U. S. Dept. of Agriculture SAGE (Ih

ae

Ait eg
Pie ea

BIOLOGIES OF WEEVILS.

Fig. A.—Pepper, showing egg punctures of Anthonomus eugenii. Fig. B.—Head of Sideranthus
rubiginosus, showing cell of Desmoris scapalis. Fig. C.—Pepper, showing larva of Anthono-
mus eugeniiin situ. Fig. D.—Head of Sideranthus rubiginosus. W ith larva of Desmoris scapalis
initseell. Fig. E.—Section of cotton square, showing pupa of Anthonomus grandis and pupa
of primary parasite Catolaccus incertus. Fig F.—Earthen cell of larva of Desmoris scapalis.
Fig. G.—Desmoris scapalis, adult weevil. (( riginal.)

Hi} atTasd ajutlvongeA to .qs0 .2 .U ,ygolomotnd to usswd ,€1 Wal

EE I TT ST TE

ur: >

i
I
1
1
i
'
|
t

24IIVASW 40 eslacoJtoid

aw MspiwoHse 10 beoH—.d . Vers -ssnosotk to go1donug ego eniwoda 19qqoaI—.A .2it
owoltieh to evtsl eniworle 1eqqod—.) vid 2oqnoa aitosmaal to [loo gitwode aw2osopsdyt
anaes abroad to svisl ctw an-owipsdy asibivobse to DsoH—.C git ita Ar Nssysysyy Bly
ETN DAB sya asssmonotiek to wqsq guivol2 »1BUp2 FOO to MOiv9e—AL wit Ilo9 alt mi
aiincunng eitosiesh 10 6711.1 10 loo foditsI—T vit awhisonh awoonlotn® offemeq yisoing th
(.[eaigitO) Jiveow iinbs inane esvosmesG—.D etl

BIOLOGIES OF WEEVILS CONTRIBUTING PARASITES. 47

16. Chalecodermus «neus Boh. is the cowpea-pod weevil, although
it has once been bred from a cotton square. It is recorded as a host
of Myophasia nea.

17. Tyloderma foveolatum Say breeds in the stems of Onagra
biennis, an early summer evening primrose. Pupation takes place
in the larval burrow. It is parasitized by Eurytoma tylodermatis and
Cerambycobius cyaniceps.

CEUTORHYNCHINI.

18. Auletes tenwipes Lec. This species was found to breed in the
buds of Galpinsia hartwegi at Dallas in the spring. The larvee feed
within a single anther in the bud, leaving it when the flower opens
and entering the ground for pupation. The entire life cycle does not
_xceed 25 days, of which 11 days only is spent in the bud. Catolaccus
incertus has been bred as a parasite.

BARINI.

19. Orthoris crotch Lec. is a very abundant fall weevil in western
Texas, breeding in the seed: pods of Mentzelia nuda, a plains plant.
Pupation is in a cell formed within several seeds. The species is
highly parasitized by several species, the least of which, however, is
Eurytoma tylodermatis.

20. Trichobaris texana Lec. is a stalk weevil, breeding in the stems
of Solanum rostratum, a plant common in waste places in the summer
and fall. The weevil cells are not more than an inch apart through-
out the stem. Cerambycobius cyaniceps has been bred as a parasite.

21. Trichobaris trinotata Say, the potato stalk weevil, is another
host of Sigalphus curculionis.

22. Ampeloglypter sesostris Lec. is a grapevine weevil, and a host
of Myiophasia xnea. It is not recorded from Texas.

23. Zygobaris xanthoryli Pierce. This new species breeds in the
berries of Xanthoxylum at Runge and Beeville, Tex., in the spring.
The larve feed in the interior of the seed until the seed is thrown to
the ground. They then enter the soil and pupate in a tiny earthen
cell, being in,the ground about a month. Catolaccus incertus has been
bred as a parasite.

ANTHRIBID#.
24. Brachytarsus alternatus Say breeds in the stems of Sideranthus

rubiginosus. It is a constant breeder, entirely riddling the stems.
Microdontomerus has been bred from it.

BRUCHID®.

Although not considered Rhynchophora by American writers, it is
very probable that the Bruchide are a connecting link between the
Anthribide and Chrysomelide.

48 PARASITES OF THE COTTON BOLL WEEVIL.

25. Bruchus exiguus Horn breeds in the seed pods of Amorpha
fruticosa in the fall. It is a host of Cerambycobius cyaniceps and
Catolaccus incertus.

It is noticeable from the foregoing that Bracon mellitor attacks
bud, flower, pod, and seed weevils; Catolaccus incertus attacks bud,
pod, and seed weevils; Cerambycobius cyaniceps attacks bud, pod,
and seed, but mainly stem weevils; Hurytoma tylodermatis attacks bud,
flower, pod, and stem weevils; Microdontomerus anthonomi attacks bud,
capsule, and stem weevils; Myiophasia xnea attacks bud, fruit, pod,
nut, and stem weevils, and Sigalphus curculionis attacks bud, fruit,
pod, and stem weevils. Such general habits indicate a generaliza-
tion of habit in the species concerned and give promise of the
possibility of more extensive adaptation to the boll weevil as a host.

ROTATION OF HOSTS.

The most important item in the solution of the parasite question
is probably concerned with the treatment of the other hosts. It is
found that the parasites which now attack the weevil are naturally
parasitic on weevils of short season and few generations, and the
most feasible proposition is that of forcing them to attack the boll
weevil only.

Two lines of action are suggested as possible and practicable.
The first is, that such plants as the Crotons, hosts of Anthonomus
albopilosus, might be planted in abundance in the pastures because
they make good feed and are easily destroyed, and that thus the
weevils and their parasites would increase in great numbers and the
latter might perhaps attack the boll weevil.

The counter proposal is that the timely elimination of the summer
series of host plants would cut off the possibility of finding any host
but the boll weevil, which would be in abundance. The cutting of
the weeds around the fields could not result disadvantageously, as
there are numerous pests harbored by these same weeds. At present
the information as to the rotation of hosts is incomplete, and it is
therefore necessary to go very fully into this question 4 as well as to
test both methods which have been suggested.

The known hosts of Bracon mellitor would insure it an uninter-
rupted rotation if they ail occurred in a given locality. They do not,
so there are still other unfound hosts of this species. The season of
its hosts as far as known are as follows:

Anthonomus fulvus breeds from May 3 to July 12.
Desmoris scapalis breeds from July 25 to September 22
Anthonomus squamosus breeds from August 11 to September 19.

Anthonomus albopilosus breeds from August 26 to September 29.
Anthonomus eugenit breeds from October 6 to October 31.

ROTATION OF HOSTS. 49

Important gaps exist in the host rotation of Catolaccus incertus.
Its hosts breed as follows:

Anthonomus signatus breeds from April 11 to June 12.
Anthonomus fulous breeds from May 3 to July 12.

Auletes tenuipes breeds from May 9 to May 25.

Zygobaris xanthoxyli breeds from May 11 to June 29.
Anthonomus xneolus breeds from May 17 to July 12.
Anthonomus nigrinus breeds from June 30 to July 18.
Anthonomus albopilosus breeds from August 26 to September 29.
Anthonomus eugeniwi breeds from October 6 to October 31.
Anthonomus disjunctus was found breeding October 11 to 14.

The known habits of the hosts of Cerambycobius cyaniceps are as
follows:

Tylodernia foveolatum breeds from June 22 to September 23.
Trichobaris texana breeds from June 26 to October 11.

Lixus musculus breeds from August 11 to September 19.
Anthonomus albopilosus breeds from August 26 to September 29.

The following data comprise all that is known of the hosts of
Eurytoma tylodermatis:

Tyloderma foveolatum breeds from June 22 to September 23.
Lixus scrobicollis breeds from July 12 to April 17 (following year).
Anthonomus squamosus breeds from August 11 to September 19.
Orthoris crotchwi breeds from August 11 to September 19.

Lixus musculus breeds from August 11 to September 19.
Anthonomus disjunctus was found breeding October 11 to 14.

In the field at Dallas upon which release experiments were con-
ducted, and in a series of five fields in various directions from Waco,
it was found that the various species of parasites were exceedingly -
localized, indicating that the parasites were derived from the imme-
diate vicinity.

On the Dallas field, by reference to the tables given in Section II,
it will be noticed that Plat B with one small exception was the
only plat in which Catolaccus was found; that Plat B was the only
one from which Cerambycobius was bred; that Eurytoma was not
found in Plat C, but was predominant on Plat E; and, finally, that
Bracon was well distributed.

At Waco on all five fields Bracon mellitor was present in the pro-
portion of 20 to 50 per cent. On the prairie land field, surrounded
only by Ambrosia, Helianthus, and Xanthium, 41.6 per cent of the
parasites were Hurytoma tylodermatis. On the other fields, which
were on the bottoms, only one Eurytoma was taken to a field. Cato-
laccus incertus was present on the prairie land field and one bottom
land field. Cerambycobius was present in all the bottom lands and
very numerous, 58.3 per cent, in a field which had considerable
Solanum rostratum about it; it was entirely absent on the prairie.
The commonest plants on the bottoms were Heterotheca subazillaris,
Croton spp., Solanum spp., and Xanthium.

10292—Bul. 73—08——4

50 PARASITES OF THE COTTON BOLL WEEVIL.

V. CONCLUSIONS AND PROSPECTS.

In conclusion the writer would say that there are a number of very
important points obtained which indicate a possible natural control
of the boll weevil. The actual combinations of conditions which
bring about the instances of high parasitism can not be understood.
It is very evident that the elements in the combination are relative
moisture and light, plant response to. weevil injury, abundance of
other species of weevils in the immediate vicinity, and, finally, the
ability for rapid adjustment to new host relations possessed by the
local parasites.

Very high parasitism has been found, and it may therefore be
expected again, perhaps locally,. perhaps regionally. It may not
necessarily be expected two years in succession at the same place,
for a change in cultivation, an eradication of certain surrounding
plants, or various other conditions might destroy the advantages pre-
viously gained. On the other hand, these same agencies in another
locality might attain the opposite result.

Release of parasites in the open field on a small scale has been
found to increase the rate of parasitization. This will be tried on a
more extensive and better planned outline another year.

‘The abundance of hosts which have parasites in common with the
weevil and the fact that parasitization takes place in the first genera-
tion give promise that the weevil’s advance may be constantly dis-
puted by the parasites.

BIBLIOGRAPHY.
Aupricu, J. M.
1905. Catalogue of North American Diptera. <(Smithsonian Misc. Coll., vol. 46,
no. 1444, p. 427.
ASHMEAD, WitiIAm H.
1896. Descriptions of new parasitic Hymenoptera II. <Trans. Am. Ent. Soc.,
vol. 23, pp. 218, 219.
1902. A new Bruchophagus from Mexico. <Psyche, vol. 9, p. 324.
1902. Pequefia avispa.parasita de picudo. <Boletin de la Comisién de Parasi-
tologfa, vol. 1, no. 9, pp. 403-404.
Banks, NATHAN.
1906. A revision of the Tyroglyphide of the United States. <U. 8. Bur. Ent.,
teeh. bul. .13, p17, pl. 1V-
BARREDA, L. DE LA.
1904. El picudo del algodén. Salvacion de la riqueza de la frontera. <(Comi-
sién de Parasitologia Agricola, cir. 6, pp. 14-27.
CHITTENDEN, F. H.
1893a. The strawberry weevil. <Insect Life, vol. 5, pp. 167-186.
1893b. Observations on some hymenopterous parasites of Coleoptera. <Insect
Life, vol. 5, p. 250.
1895. The potato-bud weevil (Anthonomus nigrinus Boh.). <Insect Life, vol. 7,
pp. 300-352.
1897. The strawberry weevil (Anthonomus signatus Say). <U. 8. Div. Ent.,
cir: 21.
1902. The potato stalk weevil. <U.S. Div. Ent., bul. 33, n. s., pp. 9-18.
Coquitietr, D. W.
1897. Revision of the Tachinide of America north of Mexico, a family of
parasitic two-winged insects. <U.S. Div. Ent., tech. bul. 7.
CRAWFORD, J. C.
1907a. New hymenopterous parasites of Anthonomus grandis Boh. <Can. Ent.,
vol. 39, pp. 133-134.
1907b. New North American Hymenoptera. < Journ. N. Y. Ent. Soc., vol. 15,
no. 4, December, in press.
GILLETTE, C. P.
1890. Insects from Iowa. <Insect Life, vol. 2, pp. 280-281.
Hinps, W. E.
1907. Some factors in the natural control of the Mexican cotton boll weevil.
<U.S. Bur. Ent., bul. 74. In press.

Hopkins, A. D.
1892. Some bred West Virginia Braconide. <Insect Life, vol. 4, pp. 256-259.
Howarp, L. O.
1897. The Mexican cotton boll weevil (Anthonomus grandis Boh.). <(U.S. Div.
Ent., cir. 18.
Hunter, W. D.

1906. Medios para combatir el picudo del algodén. <(Comisién de Parasitolégia
Agricola, cir. 32, pp. 14, 30, 34, 39, fig. 7.
51

59 BIBLIOGRAPHY.

Hunter, W. D., and W. E. Hinps.
1904. The Mexican cotton boll weevil parasites. <U. 8. Bur. Ent., bul. 45, pp.
104-110.
1905. The Mexican cotton boll weevil parasites. <U.S. Bur. Ent., bul. 51, pp.
143-150.
Matty, F. W.
1902. Report on the boll weevil. <Austin, Texas.
Pierce, W. D.
1907a. Notes on the biology of certain weevils related to the cotton boll weevil.
<U8: Bur, Dnt. bul. 63) Ptnie.
1907b. On the biologies of the Rhynchophora of North America. < Ann. Rept.
Nebraska State Bd. Agr., 1906-1907, pp. 249-321.
1907c. Contributions to the knowledge of Rhynchophora. <Ent. News, vol. 18,
pp. 356-363.
1907d. Studies of parasites of the cotton boll weevil. <U.S. Bur. Ent., bul. 73.

QUAINTANCE, A. L.
1906. The principal insect enemies of the peach. <(U. 8. Dept. Agr., yearbook
1905, p. 327.

RanGEt, A. F..

1901a. Segundo informe acerca del picudo del algodén (Insanthonomus grandis
I. C. Cu.).. <Boletin de la Comisién de Parasitologia Agricola, vol. 1,
no. 5, pp. 171-176.

1901b. Tercer informe acerca del picudo del algodon. < Boletin de la Comision
de Parasitologia Agricola, vol. 1, no. 6, pp. 197-206.

1901c. Cuarto informe acerca del picudo del algodén (Insanthonomus grandis
I. ©. Cu.). <Boletin de la Comisién de Parasitologia Agricola, vol. 1,
no. 7, pp. 245-261.

1901d. Quinto informe acerca del picudo del algodén. <Boletin de la Comisién
de Parasitologia Agricola, vol. 1, no. 8, pp. 302-317.

Ritey, C. V., and L. O. Howarp.
1890-91. Some of the bred parasitic Hymenoptera in the National Museum.
<Insect Life, vol. 2, pp. 348-350; vol. 3, pp. 57-61, 151-158, 460-464;
vol. 4, pp. 122-126.
ToOwNnsEND, C. H. Tyumr.
1895. Report on the Mexican cotton boll weevil in Texas (Anthonomus grandis
Boh.). <Insect Life, vol. 7. pp. 295-309.

END-EX;

Page.

Acanthoscelis acephalus, host of Catolaccus celiodis.............-.....---+++--- 43
Ambrosia, occurrence with reference to that of Hurytoma tylodermatis ......... 49
psilostachya, food plant of Iixus scrobicollis................+.+---+--- 35, 45

aa toud. plant Of Dass scrovreoules.. 2. Sos ns os ees 25 ot ie ee 25, 35, 45

Amor pha fruticosa, food plant of Bruchus exiguus...........--..------2++---+--- 48
PA DELO LI MLCrisesOstriss DIOLOP ICAL MOLES). <- ./05 xl - 2 oiols 2 ec ees gee bee ee 47
Ave Ole 210 PNOSUO NCO. «22 oes eos Bacon sd - HS a Soe 40, 42

Anaphes conotracheli, parasite of Conotrachelus nenuphar.............--------- 43
I CONE DIONE DOS sOly BPACOR-MELULOT ncn 2s ete os oa nc a ie taco oe es phew 38
Anthonomor phus fuluus=Anthonomus fulvus.........------------+-+++--++- ine 45
PAT THONOMLUS LNCOMLSs DIO OPICAl NOLESs- -.2- 2 s<co- = hase5 ss ons coos cate ce ee cee 46
Ree dlmaeNenOdise Menjae son te ade nos occ e so oe Sige wee aa 3 49

MOSt Ol C MLOlACCIO INCU saa) os Se ede 43

Ee COLITIS CUMEM UD eit oie es ie ons ea Se pee oe 2 46

GIO PILOSi 84) DIOIOMLCH NOLES: a-eeeia tes ao as 2 eS . . See wes 46

breeding period..............-- ic Mp Be Sees ne 48, 49

host of Bracon mellitor..... - PD ee ee on Sa Se, a ee 39, 44

Catolaccusmncentisesss ete sec ee See 37, 43

CorambyCobmustCyOnice ps sees as eee = a eae 36, 42

int Or Otol CU PIAL Seto nee aa teins ieee eae 3/,.09

CHGELIMATURE serene A> Cats core hes ae 36-37

GOrotonsee crac: sate nin es ee ee Ae ets Soe ee 4-5, 48

GES ERELUSA SOMO | OMLCALOLES ars ae tee SS Sank ths oe tae ee 46

PTE CCITEy PCHIO Gear rarer wee eat sa nT SN, 22 ha oy Pe 49

hast Gr Carolccns: tech! is eee.) 5 Fae =e eo ss 37, 43

LER EO MOM LOGEN VANS ans, tars ot eee ee aes 34, 42

IMME CLEROULECUNS EL UOLULONISa= =e ee ee ee ee 34, 37

UG CHI Ac, ENCOUNCEUS: &, BYNOU YIM 25) oo oe ie a) aalas a en ee a 46

DiOlOpT CHEMO es =m aoe ty eae yi aero ae nana ee 46

DLEeECINE Pp Ellodh eos tots has tes a ose Gt 3 oc id 48, 49

ORDLOL CES TACOTS TILL LELOI tee eee Ge Sea teats AS 44

LOTIMN COT SAL Oia eae ere a 39

Catolaccis incense hee ee oe ee 37, 43

PEGE GTILOLACS INET COSTUS = ee aa aoe 42

TOG UY DILUROREDICE DE™ Pests tase paces Ae ee 42

|NGu 18° 210) O(CIr™ SESE Ene capa ke a pe EE ee CERIN et T= 37, 39

EELS a LO Ope al an Ohedee se a SPOR 5 fe eels Sat Sc eae pee oes 45
LCCHINOENEMOU ae: Cerise. acres a's 6S Sa te woe ee 48, 49

HOStHOLM ES ACOTMINEULILOT ams she eee re eee ee 39, 44

LOUIE CLOT SCLC Tee eee ee 39

Malolaccusancenttsecn = ate van Be: oa ae eae 43

TC AUMENOC INDO ONG) Je ie =e eee = a ele te ee ys 39

grandis (see also Boll weevil).

54 PARASITES OF THE COTTON BOLL WEEVIL.

Page
Anthonomius grandis; host’ot Aspergillus Sp. ..22 2 Sc= secs esse ee ee 41
BraconidS .: 5. 22. So eee ee 29, 40
Brovon mellitor so eee Lee eee 44
and tts iorme. 22's ee ee ee ay)
Bruchophagus herrete 22. oS ee ee 42
CRiOlACeis Tn Cerise ee ws Slee eee ee 43
Cerambycobius cyaniceps -...:...-.2--2-- 2052s. 0s. 42
Cordijceps Sp 2a... e ts eek eae ee ee 41
Burytome WilOdemans = <2 ee ee eee 42
Microdontomerus anthonomi .........------------ 28, 42
MUO DROS, NCO eho ccc (RC. ee ee 42
Pediculoides: Veni icons Sop ape ee ae 42
Sigal phusicurculionisc.. ¢2-s022 25 are eee 44, 46
PyrogliphusDtewice pin. 2 a2 Oe tee Pe eee 42
Uroswpalphirs anthonomis - 202) ee 38, 44
SCLAWONEU so eee ee a ee 38, 44
possibly a host of Catolaccus anthonomi.........-...----- 31
MERICONIES IODA DIY = 9A. (CUPENT sees ence oe ee es 46
negrinus, bibliographic relerenee! 2222225 20. es eee ee 51
DiOlorical MObes oe es Gee ee he ee oe ee 46
breeding period? 2227.20 s. Sete ee oe 49
hostot :Calolactiusinecrius—. = 21s... Soo eo ee 43
Emiedon lithocolletidts 3. = 085.3 .0025-c22es open 43
Phanerotoma thivahs 4. . st a2. = eee ee ee 44
scutellatus, host of Sigalphus canadensis...-.......---:------+--22- 43
signatus, pibliographic reterence:.. .--2ce oc se oe eee oe 51
biological notes.<-+ Beso pepe ee ee ee eee 45
breeding period -.- te ee ae ee ee ee 49
hostiol Braconanthonoml. 2 Stara ces ee ee ee 44
Calapius: tebvator. Ree ee ae 43
Catolaccusonthononitas= ace eee ee 31, 43
ATVCETEUS fists Oat rs Sepa ee, SE 37, 43
squamosius, biological motés.-- 52.25. - 55. een See oer oe ee 46
breeging Pernod. 2. hc cos PN a eee ee ae 48, 49
host qt Braconmellitot secs oe: oo sete Sete eee 39, 44
form: anthostignia.. 2) eee 39
Hurytomea tyulodermans.-- 222 ncn se eee 34, 42
in -Grindelia: squarrose qiudas. +. 6225.55. 2 see ee 34, 39
Ant. (See Solenopsis geminata.)
Apion decoloratum, biological motes.< !: 2. 50: 2 ee eee ee eee 45
hiost of ‘Catolaccus incertits:.—- 2. + 2-0 eee ee eee 37, 43
in.Mevbomia paniculata: 2.0 2: ae fee eee 37
gnseum, biolopical notes... 20. ho. Soran eee 5 es eee 45
host.of Catolaecusaneertus.: \onos.. voe gases Goce eee eee 37, 43
in Phaseolus penenniseac:.. 5.2. -s8 sos o2e Heh eee 37
seqnipes, host ot Hurytoma.tylodermatise....02- --eeas* oe ae ee ee 42
Asecodes albitarsis, parasite of Magdalis xnescens...........-.----------+00+2--- 43
Aspergillus sp., fungous enemy of boll weevil (Anthonomus grandis) ..-.--- 11, 28, 41
Auletes‘tenurpes; biological notes-:.. 222525: So scee cee soe. eee eee 47
breeding period. +: :22. 22: g226 Steet eee 49
host of (Cotolaceus miceriuss.. 2222.2 nee ee ee eet ete teS -.. 43
Balaninus, host of -Urosigalphus armaius..- 02-5 -* aoe owe eee 38

nasieus, host of Mysophasia sneas.:2- 2. S522 ee ee '. 40, 42

INDEX. 55

‘ ‘ Page.
Ealeninns NAsiciistcc). NOt. Ole MICNACIS PUsLPES..-.-- =. .22 sends Sioe<%.- see Fes. 42
Spp a lostewon Urosigalphus armas. 22.2... 2 a2 gee enc. -'-- 22 2a 44
1ST Cag) Oe aL st 0a 370 (07 OU 0) Se a ie ceca Pe Ad
birds -tactoram conirol of boll weevils... 33... ost. 22 ages - =~ eRe Ee Sen 3
Blackberry, food plant of Anthonomus signatus..............------- Re AOE * 45
Bolls, fallen, percentage of parasitism of boll weevil...-............---- 14, 15, 16, 21
hanging, percentage of parasitism of boll weevil. .....--.....-.-- 14, 15, 16, 21
percentage of parasitism of boll weevil in 1905.................--..----- 13
Boll weevil (see also Anthonomus grandis).
CHOC RS! OT LNG KOANC Men eS hak fen 2p as ote sot Enis iow 3
chronology and: parasihismls tons. -s.4 tans oi sj. i's se ee 21
Gonitvol tne wlde SPACING setae Soa sabe minyn Sachs ss oes tie 5, 22
ACOs LUNCOTETO Erm ee eens ON. Les = meh SS te Nace eee 3
Ii penPenanikes list ey aa. hen eetey Sere SE eS aN A ag tte Bos 28
parasites. (See Parasites of boll weevil.)
tee areOus SMeMmMICHMste: cee os ol sds Shes soci oe eee 28
jDUTTO NAN Aloe OB Nero he/she Slee eee ee are 27-28
stages in fallen forms, parasitism.........-- Se ioe ae aes cee 22
NGUPIMP MOTUS PAFASIMISM 3.004222 5025022556 o~ a toa 22
Biigiep ames PAPABI SMe tek Aen 2 le oe le ce Pe wit ets ake 19-20
BACLT OLLerRGaLUS.) DIOLOPICAL NOLES! = 5 <2. oe pe ne oe ge 2 eee cee ieee 47
host of Microdontomerus anthonomi..........--------- OonAD
TOS UMC ANINUS MU OTOIMOSIUG. = 22 bo 2 oP ue 32
TRIES DOS Ole AnCGM SP estas one oe Ie oe. Mio. ss heen els welar en 44
Bracon.cnacdts. parasite of Tyloderma fragame....< <2 222-222. -6- 28 bashes 44
anthonomi, parasite of Anthonomus signatus............-..--.-------- 44
dOnSOtOTL=—NORMnOl es MEUtOh mace eee ee ee a Sas ee 11, 29, 38
OCCURTEN COS easier gee yee senses Se Sey oat L en Ole ena 29
PALASLFO NOL, DOLE OCW A 46 aks Soe eI 2 le ot SE 11, 28
RETCRpuOCEUILO NCO ale ns Oy pti pS ada a ales ea lode ee Paes 49
mellitor, accidental host of Cerambycobius cyaniceps, Eurytoma tyloderma-
tis, and Microdontomerus anthonomi.........-.- 28, 33-34, 34-35, 37
artificial propagation as parasite of boll weevil..........-..---- 22
Ibiolomicalnatess ates ei ee ee a oS oe a tee 38-40
Biracon GOTsGLOn aS YMOMVAN <0 2 Fe Sale oe. athe wa ies oS aid oe 11, 29
LUM UIOSUGIEG 2 SY MOWUVIN: 2S 2S hot Oe te 29
center of activity as boll weevil parasite..............--.------ 9
Peneriocelmiemcemat Amat es tee beet, 2 Sete 49
geographical and seasonal distribution................-...-...- 31-32
ORG Shs lS tees panne ergs eR Pe a koi eee Be 44
OG CUTE COMES cain sae septs ay ss ee a. Saas deca ee 29, 30
parasite of Anthonomus albopilosus.......-.-...-....-..-+----- 46
CUR Mae ee 5) D3, Sac Sa a 46
ULI SS a pt RE A eee mae eee 45
SOUANUOSUSE 2 = eae =e eget Cc er eroes Se: 46
olileyee vile ek seen teee eee ot i te 2) on es Eee 11, 28
bud, flower, pod, and seed weevils.........-....--- 48
ED CST VOMSESCO:D GUUS REE ee alone aa ees eee 45
esis MSS) Coheg Oey rl oes Salt ees ee ee eae ae eee ea 25, 26
ERRNO MO Tee Se ars eh a Ee Oe eo Mes oo oe ete 48
Pissudis.- Patasite Or Misa0Ges SOUT 2 c5 hos pis oa Sie oi <2 stwvaie cid So a's 2 DS Ad
rhyssemati, parasite of Rhyssematus lineaticollis..............-..-------- 4

snucronygis, parasite of Smicronyg, tychovacs........0..-..2--+----225-: 44

56 PARASITES OF THE COTTON BOLL WEEVIL.

Page.

Bracon 8p-; parasite of Barisispsassree o> 02 ao eee oa o eee eee ie eee ee 44
BTOCHY LAPS US UVIY OGLE ore. ore ea ee Ee 44

POMAUCUSH DIY EAEE EA SS SD ea ee it ee eee ere 44
manthostignia== -B-melltopan -o.. eee ete eee oe ae er eee 29, 38
OCCUTTEMIC CURE See ee ER CE ices tte) emo nite ener 29
Braconid:- parasite of boll weewilat tee cs.25. 22 see aae Serle Sens Serene 29, 40
Breeding records of parasites of boll weevil, 1906............-.---..---------- 13-22
work on ‘parasites of boll" weevils... ass ree oe eee eee 11, 27-41
Bruchidez, probably a connecting link between Anthribide and Chrysomelide. - 47
Bruchobius laticollis, parasite of Bruchus pisorum.....<-....------+-------+-+-- 42
Bruchophogus herrere, OCCurremee. 22s Ae le = oe a 2 se er ie Fee ee 29
parasite of boll weevil (Anthonomus grandis) ..- - - 11, 27, 36, 42

new, bibliographic reference. sss et eB te see etree ae 51
Bruchus exiguus,, bidlogical motes ht: 3002 20 ee Bei aera eis tera 48
NOSt Of GALolaccustinicentits seer rear ee eee 37, 43
Corambycobtus CYOntCOp3s et. nat vee ee eee 36, 42
pisorum,. hostiol Brovchobias 1avicol lise oes ee nee oe oe are eee 42
PPOSO PS, ADOSt Ol TOSLGGL PUG UMUC a= 8 see are see ne cre Sete yee es eat ie ae 38, 44
IMP FOSO PIS QLENOULOSA= =a an ete CEE eae ee nn et 38
Callirrhoe involucrata, food plant of Anthonomus fulvuus...........---/-------- 39, 45
Calyptus tibiator, parasite of Anthonomus signatus...........--------+++------ 43
Cathartius cassie, enemy of bollsweevil 22225222. 0 = - on sie eal ae eee 28-41
OCCUTTENGCO:.2 SEIS ee Sse ae tava thse nya = ate Se lS/epel are oa ieyata Sear 30
Catolacciistanthonpmi-.0 CCUnreNnICensse: see nese see eee ene ee eee eee 30-31
parasite of Anthonomus signatus=.--.. 5. Jess ea ee 31, 43
possible primary parasite of boll weevil..........------ 28
celiodis, parasite of Acanthoscelis acephalus...:......-----.--------- 43
ancertus: biological Motes ss =~. sh Te Sete eta © tera ae eee 37-38
center of activity as boll-weevil parasite............-..---- 9
geographical and seasonal distribution........-.-.-------- 31-32
INOSOS, LIEBE .oeeaase tere Soe Soe ae oat he tee 43
localization(: =. soa Neen sete te eres fat seen ene eee 49
OCCURPEN CES. se ee eee ete e oe ee a eee ae oer eens 29, 30
parasite of Anthonomus albopilosus.......--.-------------- 46)
GISHUNGUIS: sar veto a oe ee 46
CUGET SE OLR i oS. ce eae 46
PUPS STE ee ee een ee 45
RAOTUMUS=2 oD ae ee eehe oe ee 46
SUQNGLUS 2-222 se ew ertne ae ee eee 45
Apion decoloratum....-.- WP se a een eas tel 45
OMISCUM Sais soge tess eee eee 45

Aniletes tenwipesscn0 2 oe ate Ce ee 47
boll weevil’. = cee re ee tere ee See Toi
BT UC US OVUG MAS Se Des ce tae Ses eae eee ie 48
bud; ‘pod, and seed weevils... J. .-.2-<:.<.---226 48

Zyqovaris Lanthorylt.. 2 eee a lee eee 47
release experiments... 22.222 :o5 250 e oe eee es ae ee ee 25, 26
TOLAtION; OLHOSIS NSS ssa oe aoe ee ee een ee eee 49
localizationtearsc ee ae ee tore ee onc ee eee 49
Cerambycobius cyaniceps, accidental parasite of Bracon mellitor......------- 28, 34, 37
biological notes'*: <r. cue eae ee ee ee 36-37

center of activity as boll weevil parasite..........---- 9

geographical and seasonal distribution..........----- . 31-32

INDEX. ie 1

Page

Cerambycobius. cyanieeps, hosts, Mist= 22:2 4ie225- i == 123+ 2 eacls Plaine ciers slot 42

hyperparasite of boll weevil........-..........-. 28, 34, 37

COCURFEN CELE tn ete 2, Se see se nis os se a 29, 30

parasite of Anthonomus albopilosus..........------- 46

Loo teonial eee ee ees emer sy ane ML, Bz

PSTLCIMUSKERUG ULLLS 4 aa ne See ore ne 48

bud, pod, seed, and stem weevils......-.- 48

tosis, misculuss sabes sages es st aes 25

DT IENGVOTRS LOCGTUM <= a ees ahaa asl Sale 47

tiimotatapee saee Sacer 25

Tyloderma foveolatwmscs.. a2 oh So 47

TOLCASCLEXPCHIMENESS: 22,048 saci see oe ee eee es QON2iT

MOM MOMN Ol NOSUS Suen woaeoR hoot e wooden: Cee ae 49

' Localization Mere se yoe re atta ige 22 sepa ctt4 teeta a oleh ele ev 49

Cerocephala pityophthori, parasite of Pityophthorus consimilis............------ 43

SCOLUM VOTO PArasibexOlescOly tl deere asserts ees ae = eee 43

Reuconuynchus rape. nost.oL Onuphale tanta: 2.2202 22 25.2 Sse - 2s. ce eee eet 43

MMRIPOUCTITUS LONGUS a DLOLOCLEA AOR 434-242 U sts vie sel eae nee eps ES SS 47

INGO AVON PENNA Moan cceascaeoe bd AaeeeoreBeeGeococ 42

SEGAL DIU SIR Mere epam che peas Joa 8 acs tase Ad ae 44

Sp pWOntiv Oi Mito pNasta ene a 52 ok = Saat ese. oe 5 Seas Se se in 40

Cheiropachys colon, parasite of Magdalis enescens........-...-------------+----- 42

Chronology of boll weevil as related to parasitism.........---.-.-.----------- 21

Panormachelis: ayints. DlOlLOPICAlLMOUCS=.n 2 22-5. 5-20.22 sete o's eae eet 46

INGE OH LN MOpN PSIG) CANA 3 sccdanca ~ ~sbocsaneeeadeues cae 40, 42

MMT E KOT eT en eases tata a i eS Se aera 40

“uglandis nost or -Metaderia basalts: =... <b... <.-002-24-¥--2 >< 42

IM DOPOLOISIOR CUA a's ae onccern 5 Eeuebeodegses sooane 40, 42

juglandis (?), host of Sigalphus curculionis.......:---.---------- 44

nen phan, biological motes: eas ote = ated Saas. ok Seer 46

hostiol, Ana phes conotrachelts-y<04 as aie ss festa 43

EP OFRZOTACOROUGENCLUSS pnts ot fe inc ee 43

Sigalphus cuncwulionis=. 29-0 .. o8-o tac 4 wins ae 38, 44

posticnius. host Gt Sigal phys copert: +.) 20.55. teeta a2 asin eee 43

spp eehostsoieU rosigal pus! armatus 24a at ee oe et Se ee . 44

Cordyceps sp., parasite of boll weevil (Anthonomus grandis).........-.------ 10, 28, 41

Cotton, food plant of Chalcodermus xneus......2.-222222 522202 eee eee eee 47

forms; tallen’, parasitismion boll: weewil-2.4s-2: 32 $226 yo 8e ot see eee 22

hanging sparasitismeof boll. weevilt.ce- 2s. se-sese) 3.2 sss 22

vanlenestayorablestoubolle weevilucontrolesassee sees oss oe Sees 5, 22

Wide spacine ior control of bolliaweewils< S20 azesc 22 2S cee s 32S 22 5, 22
Cowpea-pod weevil. (See Chalcodermus xneus.)

Croton capitatus, food plant of Anthonomus albopilosus.........-.---------- 37, 39, 46

engelmannt, food plant of Anthonomus albopilosus.....-...--------- 36, 37, 46

spp., food plants of Anthonomus albopilosus.........-.--------------- 4-5, 48

occurrence with reference to that of Cerambycobius..........---- 49

Cylindrocopturus longulus, host of Sigalphus copturi.......-.-..----.-+------- 43

Dendroctonus picexperda, host of Melanobracon simplex..........-.----.---+-+- 44

Dexmones scapula Dior end NOLES S crm. ac tees ee See eee ee a ea 45

kee Wingy Demo Gs tae 32ers fh ae eee sk. Si ole heme 48

host of Bracon mellitor...... - Lage Melber! Perched ey best ae 39, 44

Orman Gar salons le 3 Sel bey Vil tya es ey 39

IME SLOCRANIEILS ET LLOCO UNO S1LS ae eee a en ee 39

58 PARASITES OF THE COTTON BOLL WEEVIL.

Page.
Determinate growth, factor in control of boll weevil.............!-..-....--.-- : 3
Dinotus'sp.; parasite of Magdalistenescens!: 3. Aer ee ol oe eee ee 43
Dryness and sunlight favorable to parasitism of boll weevil! See ee 22
Dryocetes autographus, host of Spathius canadensis............----.------+----- 44
Empusa (Entomophthora) spherosperma, fungous disease of Phytonomus punc-

LAEUS s DIRe ee SER aR SASS terete apse ee ole S sister wa ee ve )s wiewicke oe wiele ae ben seers 4]
Entedon lithocolletidis, parasite of Anthonomus nigrinus......-....----+-+------ 43
Entomophthora phytonomi, fungous enemy of Phytonomus punctatus..........-- 41

sphxrosperma. (See Empusa [Entomophthora] sphxrosperma.)
Epicerus imbricatus, host of Sporotrichum globuliferum.....-....-..-.-------+-- 41
Eupelmus cyaniceps— Cerambycobius\eyaniceps: 2 3-22-25 cee eis se ee ee 11
Bury ton ;localivationver’ co.cc cee roars 2 eee ee eet hee nee Oa eee ee 49
magdalidis, parasite of Magdalis armicollis............------------- 42
species parasitic on-bollweeval 22-4. oocth ee eee ee eee oe ee 11
tylodermatis, accidental parasite of Bracon mellitor.........--- 28, 34, 35-36
biological motests< 25 Aatee Se ee ee eee eee 34
center of activity as boll weevil parasite............-- 9
geographical and seasonal distribution...........-.-- . o1-32
hosts, Waste s-seee Jers Se SE ee ee De ee ees 42
hyperparasite of boll weevil..........-..-...---- 28, 34, 35-36
localizations: tia seer esse Leese eee ee Cee ee 49
@CCUETENCE as pe oe Boa ee Se Re ee 29, 30
parasite of ‘Anthonomus disjunetis:.<.- 22 222 e sewer 46
SGUGMLOSUSE = teen eee eee 46
A pron scynipes ae Te ok, eae ee 42
boll weewnle scl SIE B= See a ee eee 127
bud, flower, pod, and stem weevils. ......- 48
isGUSNSCrODUCOLLISe seer eee 25, 26, 27, 45
Orthoris \Giotchtt. 6s 222 een 47
Tyloderma foveolatumocs:. 229. 220s A a see 47
rotation f+ hostesses Re Ee ah epee 49
Examination work-on ‘parasites of boll’ weewil (25212 22 222.22 3.) ee ee 11-22
Formica fusca subpolita perpilosa, enemy of boll weevil................------- 28
Fungous enemies of boll weevil. (See Aspergillus sp. and Cordyceps sp.)
Epicexrus imbricatus. (See Sporotrichum globuliferum.)
Phytonomus punctatus. (See Empusa [Entomophthora]
sphxrosperma and Entomophthora Reese x
Galpinsia hartwegi, food plant of Auletes tenwipes........-..-.-.--------------- 47
Geographical considerations governing parasitism of boll weevil........--.----- 18-19
Glyptomorpha lai, parasite of Inxzus scrobicollis.........-....------------------ 44
mavaritus, parasite of Kigus scrobicollis. 2232-52 s2-2- = 22 = 22 ee 44
novitus, parasite iol Ligus musculis: sess. Poss se oe ee 44
rugator, parasite of Lixus concavus and L. musculus......-..----- 44
Grapevine, food plant of Ampeloglypter sésostris.........---..--+--------- ae 47
Grindelia squarrosa, food plant of Anthonomus syuamosus.....-.--------------- 46
nuda, food plant of Anthonomus squamosus.......--------- 34-39
Heat and dryness, factor in ‘control of boll weevil. >2 - 220. -422 52222 52-2 eee 3
Helianthus, occurrence with reference to that of Eurytoma tylodermatis.........- 49
Heterotheca subaxillaris, food plant of Anthonomus disjunctus........-.------ 34, 37, 46
occurrence with reference to that of Cerambycobius. - - 49
Hibernation of boll weevil parasites in bolls...............------------------ 12
Eickory, food plant. of ‘Conotrachelus ayinisai--co-2-)-s-- 2 22s a 40

nut weevil. (See Conotrachelus affinis.)

Page.
iHydnocera pubescens, biological motes! 1255-22 F202 Le rae tee oe ee 41
enemy or bollhweewile fe 2.2 LS eee ae 28
OCCUITEMCER she aa eae Es 5 Se EP cine Se «oslo 30

Insanthonomus grandis. (See Anthonomus grandis and Boll weevil.)
Heal=wormenactorin: controlot bolliweevilit: sa. = soe ee ee Bee ee 3
Taxus concavus, host of Glyptomor pha rugator. 2.222. bose eset ee en 44
TURES IOlOOICA NOLES! ance qe ees so nee done re bab aac oeeeaes 45
IncedmneyperTigd’: a5 sie cess hae ers 2 ee Se ee Leas See actos 49
host of Cerambycobvus cyaniceps >. 8.2 ele ise Le Bore. 25, 36, 42
Bruryionma tylodermals:<(ivecest. Se Ske Pee Se 34, 42
GI PLOMmOn PRO NOWLUS a 0228 dos sek eee a Be eee sans 44
MUG QUOT = eon eee eee none eM ctavanavars A Sas 44
Neocatolaceus tylodertns £2 J P22. 2a ee ee 43
in: Polygonum pennsylvanmicum 2.0.2... i222 ee eas 25, 34, 36
pareus, nost ot WVeocatolaceus tylodermm. 229.2222. 2. 2b eee ote eee 43
BEraDTcOl iss WIOlOMed | MOLEN-=- = 8 Sisecee Sess Sta ee ce ete ede ee ree EAS 45
DReC@aMeePETIOG = yeast 2.2 Perea on es ae ee oie 49
host of Hurytomatylodermatis: .... 0-2. 2+---------+++'- 25, 34, 35, 42
GUY DLOMOnDRGiiien 22 sss. ee Pe ot, on noes see 44
PVUBAT IVS Tae SERN 5a Me ws 5b wig cals 44
ISG ELEVA SRC HTH HE] Os eRe a Le ¢ At
apiouelmagevess 4058 Swaseree tae) 2 8 sete cae s,s 44
AMPA TAD LOSUMST OMe oe eee ae sata ee ne ota wa 2 isles aise eed 25, 34
AT LOSLACHUO TSS oe OG OES ole a. 2 ciate poke 34-35
Magdalis xnescens, host of Asecodes albitarsis.........--.-.----------+---++++--- 43
Cheivopmcny sealants an eee te oa hacia shears 42
DUMOUUS Spy eer Aa norte eo eee Sato oe 43
armicollis, host of Eurytoma magdalidis..........----------------+++- 42
ola. (%)s hostor. S paths canaidengs <2. 922. 2225222. 2 52 Reese: 44
Meibomia paniculata, food plant of Apion decoloratum......------------------ 37, 45
Melanobracon simplex, parasite of Dendroctonus picexperda...........---------- 44
Menizelia nuda, food plant of Orthoris crotchii.............--------------++--+- 34, 47
Metadexva basalis, parasite of Conotrachelus juglandis.........-.-------+---+----- 42
Microbracon nuperus, parasite of Orthoris crotchii .......-..--------------+---- 44
Microdontomerus anthonomi, accidental parasite of Bracon mellitor....------ 28, 33-34
bialoptealmones- 24. 2. assent oes sss eas Sale 32-34
hyperparasite of boll weevil.........-..-..----- 28, 33-34
OCCURICN COm-e eee ta teeta ose nisi ee 28
parasite of boll weevil (Anthonomus grandis)... -. - 27, 42
Brachytarsus: alternatus. 2.2... 2-- 42
bud, capsule, and stem weevils .........-.------- 48
parasite of Brachytarsus alternatus.......------ ae Se aes AT
Microdus simillimus, parasite of Lixus scrobicollis.............-.--------------- 44
Min nkastdcaed.npiolopacal NOLES: 145-4- <2 ese ee Oe =e ee re a nln ete Pa 40
OSES SOLIS Gop eee eee ay fey Se Son i ae ere al ofc rae Ses a mecishes 42
OE CURED CO sae er ee ee ee ate areca msl erate ae 30
parasite of Ampeloglypter sesostris...-....-.--.--------+-+-- 47
pO arseit aerate atta SS pee Ol ok oc ee aa 28
bud, fruit, pod, nut, and stem weevils........----- 48
Chalcodennustenews 2 2 5-5-3552 2 ae. seh 47
CORGITACRELUS EE UNIS Week Played oa nage He =A 46
robusta, parasite of Sphenophorus robustus.......----------+------- 42
Neocaiolatcus tylodermice Wests, LSts clan eoseh ne ot wines Sites cht 3 Sane 43

60 PARASITES OF THE COTTON BOLL WEEVIL.

; Page.
Omphale livida, parasite of Ceutorhynchus rapx.....-..:..-..----+------+----- 3
Onagra biennis, food plant of Tyloderma foveolatum.........-....-------+---- 34, 36, 47
Opthahis crotchz; biological motes’. sas sot yo ie eee eae eee eee 47
breeding pemodasts 2. wchiceaeie ices ca ehe 2 ee eee cece 49
host.of Hunjtoma tulodermatiss-2. -eesaetese eee eee ee 34-42
Microbrocon 1aiperus s25258-S. ese eee oe eee 44
Tetvasiichiisisp >. sew sce SS eee eee ee oe ee 43
an, Monizelia, Wud =| on ee ee Se ec a ae ee ee eee 34
Parasites» factonumucontrol of bollsweevilesssss2 eee ete ee eee gah
known tovattack-Rhynchophoras:2< 253". te -ceseece oe acre eee 41-44
of boll weevil, artificial propagation:<-4-.:cetes Soe ase ee eee 4, 22
bibliography... “ise tes 2¢ hee eee eee 51-52
biological notessotxctte aig Ee bet ee ee oe 32-41
biologies of weevils contributing them.........-.---- 44-48
breeding'records of 1906+ 302 fro 53 oe eR ee 13-22
. brecditia wobec 22 Seo eee ee Se eee ee 11, 27-41
CONCLUSIONS Ss! jo asec oe) Bee es Soto Bee eee 50
Tromewouke ond G0 G5=-eer eee eee 21-22
elimination of natural hosts [other weevils]. ....-.-.--- 4-5
examimatrombworks ese ee cat le ee ee 11-22
field works 7s: soscatpcetc lone tees =e ce eee 24-27
geographical and seasonal distribution...........-.--- 31-32
hibernation anybolll sees a sea = See ee eee ee eee 12
WistOry 2722.0 Bol skeet Aes see eee eee ee eee 1O=11
intensive Centersie.cs4-52% oes os be ee eee 9
OCCUIFENCELOLSDECLES see ee eer ne oe eaeee eee eee 28-31
percentage in bolls versus squares in 1905........----- 13
propagation workeonsscelt fee. ce igseee eee 11, 22-27
PIOspects: tse Sse Sse Se ee ee 50
records priorto 19067225. 2 ee Lee = Sh oe eee ee 12-13
release €xperimentsn.<... inh <ctee- 22 ee eee 24-27
rotation of hostsié-t2q<h<cG=0: is JU eee ee 48-49
schemes for increasing their work. .... Pe Me SS Ps acs 48
SOUrCE Work vee ee se eee be ee te eee or 11, 41-50

transfer or artificial propagation, apparatus and diffi-
culties. oie ett goss ig c Eee ke eee ee 22-24
workon90G ¢conclusions=eessee eee eee eee ae 11, 50
Parasitism.and status of oll weevil-cc- a. 2 tence eee’ ope eee eee eee 19-20
of boll weevaliandychronologyetascb tae ee RS eee 2 ee 21,
bynmlonthis, 1906.2 e832 soe ek ot ece a eee eee 14
fieldiconditionsss< 32 Gahaks hee aos oc e eeee ee 17-18
geographical. considerations;.<2% ---pie<<-as2----<s-0 18-19
im itallen* boll sine ase Sasa eee eae 14, 15, 16, 21

FORMS 22 fo58 Be. Se eee eee eee Repdes

in shade versus sun.......-------- 17, 18, 22
SQURTeR Meee asec 3 ae ee ns 14, 15, 16, 21
forms in prairie versus woodland........-..---- 17, 18,22
ihanein os balllseee=eeeeeseras eh OE Oy 9 2a 14, 15, 16, 21
fOTIMS wats AOS sels Se wee ee eee 22
SQMares He sssese cele tase eee 14, 15, 16, 21
percentage by years, prior to 1906.........----------- 11
plant conditions most favorable..........----------- 15-17

Peach curculio. (See Conotrachelus nenuphar.)

INDEX. 61

Page.
Pediculoides ventricosus, introduction into Texas an unsuccessful attempt... --- 40-41
OCCUITEN CEs: Meenas. ee pide a Le 7 See eR 30
parasite of Anthonomus eugenii..............-.-------- 42-46
boll weevil (Anthonomus grandis). 10-11, 28, 40, 42
Peppers, food plants of Anthonomus eugenit...:.-..-.----..-2--22-22+--+- 37, 39, 46
Phanerotoma tibialis, parasite of Anthonomus nigrinus..........-.------------ 44
Phaseolus perennis, food plant of Apion grisewm........-..-.-----------2------ 37-45
Phleosinus graniger, host of Spathius canadensis..............-..----------+--- 44
ROROPICEISICONUPLONA— ANCY ILS! COMPLANGe ease aoe ae ee aoe ee ee ee 38
Phytonomus punctatus, host of Empusa (Entomophthora) sphxrosperma and
LY LEC SOLD ACRE O OOO Dee SOS ES Hoe AeA OB So Boone nee aol cle aoe 41
Teoma sO00s NOSt Ol BraCOl PISSOdIs2= = 22 22 ates oo. 02 Sake | Pee is 44
Pityophthorus consimilis, host of Cerocephala pityophthorv....-. yet Kepat ee te 43
Bp: host of Rhyssalus pityophilont i... sore >. 22g Sela ee es 44
Plant conditions most favorable for parasitism of boll weevil...........--:-.-- 15-17
Plum curculio. (See Conotrachelus nenuphar.)
Podapion gallicola, host of Sigalphus copturt...-...-.-..---22-/22--2-+2+--+- 43
Polygonum pennsylvanicum, food plant of Lixus musculus.....-..-.----- 25, 34, 36, 45
Porizon conotracheli, parasite of Conotrachelus nenuphar.....-.......-.--------- 43
Potato-bud weevil. (See Anthonomus nigrinus.)
stalk weevil (see also T’richobaris trinotata).
biplogeap tie relerences se 2s eS Sse ee eS Le 51
eae: Parasitiem: Ob boll weevil opi <p see mer Sea eS Ve eR Se ieelisa oe
Eroliterationeeractorin: control of bollisweevales= sass eee eer he eee 3
Propagation of boll-weevil parasites artificially, apparatus used, and difficulties. 22-24
work on) parasites. of boll weevil: : 7.3. -20e 04. We eee 11, 22-27
Prosopis glandulosa, food plant of Bruchus prosopis..........-..--.--------- ae 38
Raspberry, food plant of Anthonomus signatus.....-- - PAN FRO Ti IS Sey eae 45
Red-bud, icod plant of Anthonomus signatus: --2 222022. 22. 2.2 I 45
Rhyssalus pityophthori, parasite of Pityophthorus sp........-----.------------- 44
Rhyssematus lineaticollis, host of Bracon rhyssemati...............---+---------- 44
Taynehopnora patasites, lstec2. fess sc 5 2set 8 Poe SE ee cece 41-44
Rose, wild, food plant of Anthonomus signatus.....-..-.-.-.------+-+++----+--- 45
peolytid,, osteo Cerocephala scolytwvorass 022 SOS. Lees 2 foe. Fe - Ey eas 43
Scyinnus in cotton squares...2: 2.22225. 20.22 REL FUEL Ne tesa e ais et 10
Siades parasitisma of, bolliweevil =: 32552 PO EE ae on ees aM sy
Sideranthus rubiginosus, food plant of Brachytarsus alternatus........--.-------- 32, 47
DeEsMOristsca palis aes v8 2 Saas ee ee 39, 45
Sigalphus canadensis, parasite of Anthonomus scutellatus............---------- 43
COphU ry MOstSs MISt ise: Gwe eso 2 Aue NS BR SASS ORS s 0 De ae 43
CUncHlionis * ple lopical nObes 62 <3 59 essere ass D> pecs Lys cae 38
DGstse higher ere ye et enero stra lee SS ee 44
Fa
OCCUIFEN CEM. S42 Apr eycenmetny meter tleo nein s:./ mebseeat (2) 23, 29
parasite of boll weevil (Anthonomus grandis) ....... -- 11, 28, 46
bud, fruit, pod, and stem weevils. ........... 48
Soret actin TEMUNPNGT 2 SS sah = Sees ONG 222d oe 46
Trichobvaris: tnnotara sa ersye 82.25 ES. 47
Sen Patasite. Oli CRALCOGEIVUS PNLUSss 22 852s See CAS. 2 2 Se Se 44
zygobaridis, parasite of Zygobaris xanthoxyli.....-...-..-.------+----- 44
Smicronue tychoides, host: of Braconismicronygig?.< 22.2. Fo) 22225. 22 sn Lee 44
Solanum carolinense, food plant of Anthonomus nigrinus.......-.-------- Sk Ne 46
eleagnifolium, food plant of Anthonomus xneolus...........-.-.----- 46

rostratum, food plant of Anthonomus xneolus........--.------+-+---+- 46

62 PARASITES OF THE COTTON BOLL WEEVIL.

Page.

Solanum rostratum, food plant of Trichobaris texana.............------- 25, 36, 38-47

occurrence with reference to that of Cerambycobius...... - 49

spp., occurrence with reference to that of Cerambycobius..........-- 49

torreyt, food plant of Anthonomrs aneolus.—.. 222-5). 5-52- eee 46

Solenopsis geminata, enemy of boll weevil...........--...--.----..----------- 3, 28

Source work on. parasites of boll weevils: =-2ocoaeee aee ene e eee : oe 11, 41-50

Spathius canadensis, hosts, list. -22-22,--eie es 2 eee eee nee 44

Sphenophorus parvulus, host of Myiophasia xnea.........-.-...----------1--.- 40,42

rovustus, host of Mirophast robusta: ss... sees ee oe Se ee 42

Sporotrichum globuliferum, fungous enemy of Epicxrus imbricatus........-..-- 41

Squares, fallen, percentage of parasitism of boll weevil...............-- 14, 15, 16, 21

hanging, percentage of parasitism of boll weevil ............---- 14, 15, 16, 21

percentage of parasitism of boll weevil in 1905............-...------- 13

Strawberry; food plant.ot Anthonomus signatus. 222 522s.) eh eee ene ee 45

leaf-roller. (See Ancytis comptana.)
weevil (see also Anthonomus signatus).

bibliographic Felerence.. < 32 ocietgha pve oa ee ae eee 51

Sunlight and dryness, favorable to parasitism of boll weevil ........------------ 22

Sin, jparasitismoi pall myeeyileet ree Sr es oe fl eer ee eae Welle), BP.

Tachinids, conditions favorable to parasitism of boll weevil .......-..-------- 22

Tetrastichius sp, pakasiteot OFiOris ChOlehtt. 2 22 =. one 2 ees Doe Se eee eee 43

TOMmscnsspige ROM Oleb TAL SP tn. 2 2 Jie ae aac el Rote ne reese eee eee 44

Spr uhost Gkos pals canadens?s.<30. 2... 22. . paee een Sade eee eee 44

Trichacis rujipes, parasite of Balaninus nasicus (?)....2-2:--.---2- 206+ 0= --50-e 42
Tichobaris ecana Mpielogical motes:- 2.6 5.) 252k decease tee ER ee Bie

lpreedine period: . 22. ois poe Lin: Getiscew eee oe eee 49

host of Cerambycobius cyaniceps... . - aE eee nS 25, 36, 42

in Solanum rostratum.........---- iageit UAE C Secs lie ace eee 25, 36

trinotata, host of Sigel phus curcwlionss.-22 22S. 22 es a oe ee 38, 44, 47

inl /S DLO NIL, FORPFOLUMs.2 = 3. 2eths els aes er eee cease ee 38

Tyloderma foveolatum, biological notes. 2iee=s0-sesdk 222 ot oe) see hee ae eee 47

breeding period: 225) S82 csc. foe eee ee ee 49

host of Cerambycobius cyaniceps ..........----------+-<- 36, 42

Buryioma tylodermatiss oo). 22- o2c eee eee 34, 42

Neocatolaccus tyloderme.. 2%. 2-22. ee eects 43

in OnNagrd DIENWIS: 2 os ence oe eee eee ee eee 34, 36

fragarie, host:ol Braconanaledis2.. 6.22.5 Se cle as enews 44

Tyroglyphus breviceps, parasite of Anthonomus eugenii.....-...-.------------- 42

boll weevil (Anthonomus grandis).. 11, 28, 30, 41, 42

Urosigalphus anthonomi:, biological motes ease oo. cawe nels yee ae eee eee 38

MOVIN eA ae eG one ae Ooh oancsneatedcks pasos 29

parasite of boll weevil (Anthonomus grandis)... . - - 11, 28, 44

armatus, parasite ol (Balanintts) 20.22 Spiess asin eee ee 88, 44

Conotrachelus, 8pp2 <.-e rece naen oa ose eee 44

brucht, paxasite,of bruchus prosopiss 23. = ones AB eee 38, 44

(robustus). (See Urosigalphus anthonomi.)

SCHUATZ1. (OCCUITEN CO nate Suekee aa ae ee ee one ee ness eee ot oe 29

parasite of boll weevil (Anthonomus grandis)... .--.. 28, 38, 44

Vipio belfraget, parasite of Inrus scrobicollis tse 2 © ee nee eee: Sede 44
Weevils contributing parasites of boll weevil, biologies............---.--------- 44-48 |

Winter temperatures, factor in control of boll weevil.......-....-------------- 3

Woodland, parasitism of boll weevalo>-. o-oo ee eee ee 17, 18, 22

INDEX. 63

Page

Xanthium, occurrence with reference to that of Cerambycobius. ...........-.-- 49
Eurytoma tylodermatis ....... - 49
Xanthoxylum, food plant of Zygobaris xanthoxyli.......-..------- Poss ere 47
Buacuars ranthoaiy, biological notes: 52 2.02.05... 222. $222k e--. oe 47
ree uMi peri Odum emtcad ec os at eee mere ss age! 49
host on Catolaceuaecenlise 82 so Foo eee oles eee 43

SLAG DNNIS CUGOUAILOISS: 5 = til nate Se ee eid Lee 44

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