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U. S. DEPARTMENT OF AGRICULTURE,
BUREAU OF ENTOMOLOGY-BULLETIN No. 91.
L O. H7WARD, Enomolog amd Chief of Bureau.
HE IMPORTATION INTO THE UNITED STATES
OF THE PARASITES OF THE GIPSY MOTHI
AND THE BROWN-TAIL MOTH:
A REPORT OF PRORESS, WITH SOME CONSIDERATION OF
PREVIOUS AND CONCURRANT EFFORTS OF THIS KD.
L. O. IOWARD,
Chief. Lfurecu ffntomology,
W F. IKE,
In Charge, Gipsjy Oth Parasite Laboratory,
l*t e I J.a Mass.
ISSUED JULY 29, 1911.
:..::CO .-71 1
GOVERNMENT PRINTING OFFICE.
B UREA U OF ENTOMOLOGY.
L. 0. HOWARD, Entomologist and Chief of Bureau.
C L.MARLATT, Entomologist and Acting Chief in Absence of Chief.
R. S. CLIFTON, Executive Assistant.
W. F. TASTET, Chief Clerk.
F. H. CHITTENDEN, in charge of truck crop and stored product insect investigations.
A. D. IOPKINS, in charge of forest insect investigations.
W D. HUNTER, in charge of southern field crop insect investigations.
F. M. WEBSTER, in charge of cereal andforage insect investigations.
A. L. QUAINTANCE, in charge of deciduous fruit insect investigations.
E. F. PHILLIPS, in charge of bee culture.
D. M. ROGERS, in charge of preventing spread of moths, field work.
ROLLA P. CURRIE, in charge of editorial work.
MABEL COLCORD, in charge of library.
PREVENTING SPREAD OF MOTHS.
W. F. FISKE, in charge; A. F. BURGESS, C. W. COLLINS, R. WOOLDRIDGE, J. D.
TOTHILL, C. W. STOCKWELL, H. E. SMITH, W. N. DOVENER, F. H. MOSHER,
D. M. ROGERS, in charge; II. B. DALTON, H. W. VINTON, D. G. MURPHY, I. L.
BAILEY, II. L. MCINTYRE, assistants.
LETTER OF TRANSMITTAL.
BUREAU OF ENToMoLoGY,
Washington, PD I. ., pril 12, 1911.
SIR: I have the honor to transmit herewith the manuscript of a
report of progres on the importation into the United States of the
parasites of the ipsy moth and the brown-tail moth. To this has
been added some con1iderationl of P)reiVols and clnculrrent efforts to
handle the parasites of destructive inec i a practical way. The
work with the foreign parasites of the gipsy moth anl the brown-
tail moth has been going on now for rather more than five years.
It promises excellent results, and the present seems the proper
time to pt-esent to the people interested a somewhat detailed account
of what has been done and of the present condition of the work. I
recommend that this manuscript be published as Bulletin No. 91
of this bureau.
Respectfully, L. 0. HoWARD,
E0t0o1ol00st and Okief of Bureau.
Hon. JAMES WILSON,
Secretary of Agricuture.
*tssCQ ,i ." aL
~Ur; ~ I'~E -,t~gt.2 si"i
ffi..^1111!!^ f'' .^"z r si ss
Digitized by the Internet Archive
Previos work in the practicl handling of natural enemies of injuriou insects.. 16
Early practical work ..................................................... 17
Per ittin the parasites to escape............................ ........ 18
Te transportation f parasites from one art of a diven country to another
part ............................................................... 20
The transfer of beneficial insects from one country to another.............. 23
.............. ........ ........ .............................. 24
Novis in Portugal.......... ......... . . ... ........ ...... 27
c rya in Florida..................... .. ...... ................ 28
Novius in Cape Coony.... .... ..... . .................. 28
Novis in Egpt and the awaiian Islands....... ......... ... .... 28
er a in Italy ............ .......... . . . . ... ....... 29
Icerya in Syria .................... . .. ... ................ 29
The rea s for the d cc s of N viu ........... ... ..... ........ ... 29
Introuction of E don e u Walk into the Unitit States ....... 30
Other intructions by KoOle int Californi .................... 31
Inrnational work with enemies of the black vale. ................ 31
The awaiian rk .. .......................................... 34
An importation of Clerus from Germany ............................... 36
artt's journey for enemies of the San J ale................... 36
The p ts f iaspisa T ........ ........T.... ....... 38
Te work of Mr. orge Cupere................................. 38
rk with e parasite of the elm leaf-beetle .................... 39
Work with pa t of ticks.......................................... 41
Mr. Froatt's jorney t varios parts of the world in 1907-8......... 42
Other work f this kind (by Berlese; by Silvestri in Algeria; in the
Philippi ; by De Bu ; in Per) .............................. 44
Earl ideas on introducin the natral enemies of the y moth............... 47
Cirumstances which broght abou the actal beginning of the work.......... 49
An investigation of the introdction work ................................. 50
Na tie of the p r of the work........................................ 54
Known and recorded p sites of the ipsy moth and of the brown-tai moth.. 84
Establishment and disersion of the newly introduce aaite............... 94
Di e as a fatr in the natral control of the ips moth and the brown-tail
m oth..................................................................... 97
Stdies in the parasit of n n ................. ............. 102
itism as a factor in insect control......................................... 105
The rate of increase of the gipsy moth in New England........................ 109
of additioal control n to check the increase of the gipy moth
.in America ............................................................... 114
The t to which the gipsy moth is controlled through pasitism ab .... 117
tism of the ips mothin Japan.................................... 120
P tism of the gipsy moth in Russia................................... 123
ofthegipsymothin southernFrance----------------...............--.... 129
6 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
Sequence of parasites of the gipsy moth in Europe....-----.....--- ............ 31
The brown-tail moth and its parasites in Europe..---.----......--............. 132
Sequence of parasites of the brown-tail moth in Europe.................... -----135
Parasitism of the gipsy moth in America..................................... 136
Summary of rearing work carried on at the laboratory in 1910 ............. 141
Parasitism of the brown-tail moth in America---.... .......................... 143
Summary of rearing work in 1910 ........................................ 146
Importation and handling of parasite material................................ -152
Egg masses of the gipsy moth.-........................................... 152
Gipsy-moth caterpillars, first stage.---..---........--....................-- 153
Gipsy-moth caterpillars, second to fifth stages..---..... -.................. 154
European importations ..-......-.................................. 154
Japanese importations..................................----...-....- 155
Gipsy-moth caterpillars, full-fed and pupating............................ 156
Gipsy-moth pupae-......------..-----.......---..... .................... 159
Brown-tail moth egg masses.-- ----.............................. ... ... -- 160
Hibernating nests of the brown-tail moth-................................. 161
Immature caterpillars of the brown-tail moth.............................. 161
Full-fed and pupating caterpillars of the brown-tail moth ...............- 162
Brown-tail moth pupe......------......------------------...................... 164
Cocoons of hymenopterous parasites-.......................---........ ...-. 165
Tachinid puparia...........---- ........... ............................. 166
Calosoma and other predaceous beetles.........................----- -.... 167
Quantity of parasite material imported....-...-...... ........................ 167
Localities from which the parasite material has been received.-...--....--- -.. 168
The egg parasites of the gipsy moth........................................... 168
Anastatus b/faseiatus Fonsc............................................... 168
Schedius kuvanx How.................................................... 176
Life of Schedius and its relations to other egg parasites, primary and
secondary...... ..........- --...-.------------...--........----..... 177
Rearing and colonization--....-..............................-..... 184
The parasites of the gipsy-moth caterpillars-----...---.. ----......... ...----.. 188
Apparently unimportant hymenopterous parasites------........... ....... 188
Apanteles solitarius Ratz--....-........................-............. 189
Meteorus versicolor W esm-----...... ................... .... ....-...... 190
Meteorus pulchricornis Wesmr........................................ 190
Meteorus juponicus Ashm.................................---...-- -... 190
Limnerium disparis Vier............................................... 191
Limnerium (Anilastus) tricoloripes Vier- ---....................--....... .. 192
Apanteles fuvipes Hal-........ ...----................................... 193
Secondary parasites attacking Apa- teles fulvipes....................... 198
Tachinid parasites of the gipsy moth.....----......--..-...---....-.......--- 202
The rearing and colonization of tachinid flies; large cages versus small cages. 204
Hyperparasites attacking the Tachinid........--......--..----------.....---.... 207
Perdnlapus cuprinus F6rst ....................................---... 208
Melit/obia acasta Walk-..-.. --.....-.....- -......-.....---...... -----209
Chal s is skei C(rawf ...- .....-- ..-......... ........ ......... ......... 212
Monodotomnerus wxreus Walk.. -...............-............-........ 212
M iscellaneous parasites........ .... ... ................. ........ ...... 213
Blepharipa sm dtellata Desv ........................................... .... 213
Compslurr coidnctnaa Meig-...-......... .................. .............. ... 2128
TachiMn larrar ui I, .. ................. ............................ ..... 225
Tachina japonica Towns ...................... ...... ............... 227
C ONT X T 7
Tachinid parasites of the gipsy moth-Continued. Page.
T icolga gra s Zett.......... ....................................... 228
Paras ge a rgata Rond ..... ..... .......... ....... ............ 229
'a c la r Mcig.... ................... ............................. 231
Z o h nidkcol Tonn..... .. . . . . . . . . 232
Crossorosmia sericiriv Corn.............. ........................ ........ 232
rossoo i a osu l t Schiner ?.......... ....................... 234
Unimportant tachinid panisites of the gip..y moth2: 235
tes of the ............. ...gy n.... .... 23
The gen s Theroniath ............ ... .. .... ...... ........... ..... 2
The enus Pimp la ...... ............ .. .... ... .. . ............ . 237
Ihnu m disp ts Poda ........................................... . .. 239
Thegenus alis....................... . ................. ....... 240
Mon od ntomerus reus W al ....................... ....... ......... 245
The sacophagids ............................................ .. ...... 250
The predaceous beetles.......................................... ...... 251
The parasites of the brwn-tal moth .............................. 256
The genus Trich ram na........ ...............2...................... 256
tlenomun phartiarunm N es............................................. 260
Parnites which hibernate within the webs of the brwn-tail moth ............ 261
diculoides e tricosus Newp................................... ........ 267
P er l egreg i rt .......... .... ........... ........ ........... 268
A pa tees lactiolor .... .. .......................... ........ 278
Apante consper Fiske ............................................. 285
Mleteorus vcrsiwlor W es ................................................ 286
Zygo othria nidicola T n ......................................... .... 289
Parasites attacking the larer caterpillars of the brown-tail oth ............. 295
S. .........enopterous parasit................................... 295
Tachini pa i ... ... ............... ........................ ........ 296
Decodes igrip s Fa .. .. ........ . ..... .......... ........ 296
Pare orista chelon. ......... ... .... ...... ....... 297
Pals pa.ida Meig . . . . . ............. . 300
Ze a lbarix Panz........... . . . .. .. ....... . 302
asi ra s la ia Fal ........... . . . ........ ..... . 303
Eudoro.y.a magn cor is Zet . .. .............. ... . 303
Cyclotophr.s a.ser T n s ........................ ............ . 304
Bleph rd a ulars F ................................. . .... 304
Parasitof the pupw of the bown-til moth ............................... 3C4
Summary and conclusions ................................................ 305
The present status of the intrduced parasites....................... .... 307
The develoments of the year 1910------------..................................... 311
PLATE I. Theb tl ............ .................. Frontispiece.
II. Fi. 1.-View of paraite laboratory at North Saugus, Mass.
Fig. 2.-View of parasite laboratory at Meroe Highlands,
as ................................................... 56
III. Fi. 1- adside oak in Britany, wit leave raged by
ipsy-moh caterpillars. Fig. 2.--M. Ren6 Oberthiir, Dr.
Paul larchal; with roadside oaks ragged by gipsy-moth
caterpillars .. ................ ... ...-......... .....-. 76
IV. Fig. .-Caterpillar huntrs in the south of France, under M.
Dillon, 1909. Fig. 2. Packing parasitized caterpillars at
HFlyres, France, for shipment to the United Statcs, 1909.... 76
V. Fi. 1.-View of interior of one of the laboratry structures,
showing ring caes for brown-tail moth parsites. Fig.
2.-Box ued in shipping immature caterpillas of the gip y
m from Japan ....--- .......------..... ................. 152
VI. The gipsy mot (PPltria dis r.......................... 156
VII. The brn-tail moth (Euprcs c o .................. 160
VIII. Fig. I. -oxes used in 1910 for importation of brown-tail moth
laterpillars, with tubes attached directly to bloxes. Fig.
2-Interior of boxes in which brown-rail moth caterpillar
were imported, showing condition on receipt. Fig. 3.-
ioxs used in shipping caterpillar of the gipsy and brown-
tail m s bye mail ...................................... 164
IX. Fig. 1.--Head r devise by Mr. E. S. I Titu as a protection
against brown-tail rash. Fig. 2.--Show case used when
opening boxes of brown-iail moth caterpillars received from
ab road ................................................. 164
X. Fig. 1.--Lare tube cae firt used for rearing prasiles from
imported brown-tail moth nests and latterly for various pur-
poses. Fig. 2,-Method of packing Caloeoma beetle for
shipm ent ............................. .. . . . . .. .164
XI Fig. 1.-Egg of gipy m~th contaiinig developing caterpillar
of the gi moth. Fig. 2.-Egg of gipsy moth, containing
larva of the parasite A:istaIt bisti tus. Fig. 3.-Egg of
gipsy moth, containing hibernating larva of Anastatus bi'as-
ciatus which in turn is parasitized by three second-stage
arv of S hedis kuran ................................. 172
XII. Fig. 1.-View of cage used for colonization of Awnstatus bifs-
ats in 1910. Fig. 2.-Views of cage prepared for use in
clonization of Anastas bfaciatus in 1911............... 172
XIII. Outdoor parasite cage covered with wire gaze .............. 204
XIV. Outdoor paratecagescovered with cloth ................. 204
XV. View of large cage used in 1908 for tachinid rearing work ...... 204
10 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
PLATE XVI. View of out-of-door insectary used for rearing predaceous
beetles in 1910........................................... 204
XVII. Fig. 1.-Wire-screen cages used in tachinid reproduction work
in 1909. Fig. 2.-Cylindrical wire-screen cages used in
tachinid reproduction work in 1910..................--.... 204
XVIII. Fig. 1.-Blepharipa scutellata: Full-grown larva from gipsy-
moth pupa. Fig. 2.-Blepharipa scutellata: Puparia........ 216
XIX. Fig. 1.-Importation of gipsy-moth caterpillars from France in
1909; en route to laboratory at Melrose Highlands, Mass.
Fig. 2.-Importation of gipsy-moth caterpillars from France
in 1909; receipt at laboratory, Melrose Highlands, Mass..-.. 216
XX. Fig. 1.-Compsilura concinnata: Puparia. Fig. 2.-Tachina
larvarum: Puparia. Fig. 3.-Sarcophaga sp.: Puparia. Fig.
4.-Parexorista chelonie: Puparia......................... 220
XXI. Fig. 1.-View of laboratory interior, showing cages in use for
rearing parasites from hibernating webs of the brown-tail
moth in 1910-11. Fig. 2.-Sifting gipsy-moth egg masses
for examination as to percentage of parasitism................ 244
XXII. Map showing sections of its range in New England from which
Monodontomerus xreus has been collected in hibernating
webs of the brown-tail moth, and subsequently reared...... 248
XXIII. Map showing distribution of Monodontomerus wreus in New
England ..........------- ............ ........... .. ..... 248
XXIV. Map showing dispersion of Calosoma sycophanta in Massachu-
setts from liberated colonies....--..-.................. ... 256
XXV. Map showing distribution of Pteromalus egregius in New
England ...-........................................... 276
XXVI. Fig. 1.-Riley rearing cages as used at the gipsy-moth parasite
laboratory. Fig. 2.-Interior of one of the laboratory struc-
tures, showing trays used in rearing Apanteles lacteicolor in
the spring of 1909........................................ 280
XXVII. View of laboratory interior, showing cages in use for rearing
parasites from hibernating webs of the brown-tail moth in
the spring of 1908... ... -............... ................ 280
XXVIII. Fig. 1.-Cocoons of Apanteles lacteicolor in molting webs of the
brown-tail moth. Fig. 2.-View of laboratory yard, showing
various temporary structures, rearing cages, etc............ 284
FIG. Polygnotus hiemalis, a parasite of the Hessian fly...........-.......... 21
2. Polygnotus hiemalis: Adults which have developed within the "flax-
seed" of the Hessian fly and are ready to emerge ..............----- 21
3. Lysiphlebus tritici attacking a grain aphis .......................... 22
4. The Australian ladybird (Novius cardinalis), an imported enemy of the
fluted scale: Larve, pupa, adult, work against scales................ 25
5. Rhizobius ventralis, an imported enemy of the black scale: Adult, larva. 31
6. Sculellista cyanea, an imported parasite of the black scale........-..... 32
7. Pediculoides ventricosus. ........... ..... ........ ......... ......... 34
8. Erastria scitula, an imported enemy of the black scale: Adult, larvw,
pupa. ...................................................... 34
9. The Asiati ladybird (Chilocorus similis), an imported enemy of the San
Jose scale: Later larval sages, pupa, adults...................... 37
10. Rearing cage for tachinid arasites of the brown-tail moth............. 151
. 11. Map showin various localities in Europe from which parasite material
has been received... .......................... ........... 169
12. Aastau bfascitus: Adult female ................................ 170
13. Anatatu b asatu: Uterine egg................................. 171
14. Aatatu asatus: Hibernating larva ........................... 171
15. Anastaus basatus: Pupa from gipsy-moth gg................... 171
16. Diaram showing two years' dispersion of Aa tus bast from
colony center ...................... .................. 173
17. Shedius ka : Adult female .... ......................... 176
18. Sch dusku a : Eg ............................................ 179
19. Schdius kan: Third-stae larva still retaiing attachment to egg
stalk, and anal shie ....... ................................... 180
20. Schedius kuamn: Pupa................................. 180
21. Shedis k : stalk and anal shield of larva as found in host
Sof i moth from which the adult Shdiu has emered, or in
which the Schedius larva has been attacked by a secondary parasite. 181
22. Sldius kura X: Larval mandibl............................. 181
23. dar na : Larval andibles ............................... 181
24. Pa hy e ro E ...... ................................. 182
25. Pachyneuron g ensis: Larval andibls ....................... 182
26. A tts b t: Larval mandibles............... ............,.. 182
27. Gipsy-moth egg mas showing exit holes of Schedius Lurna ......... 186
8. Apantales soltais: Adult female and cocoon ...................... 189
29. Lim er d par : ocoo ........................................ 191
30. Lim erium disparis: Adult male .................................... 191
31. Apa les fauli pes: Adult ......................................... 193
32. Apantelesufi : leaving gipsy-moth caterpillar............ 194
33. Apantelesfuipes: C(oeons surrounding dead gipy-moth caterpillar. 195
34. Apanteles faripe: Cocoons from which Apantele. and its secondaries
have iue ................................................ 199
35. Blepharipa utela: Adult fem ......... ....................... 213
36. Bepharipa stllata: Egs in siu on fragent of leaf ................ 214
37. ggs of Blephripa telata and Pales pai ia .......... .... ... 214
38. B haripasutellata: Firststa e la ........................... 215
3. Bleharipa selata: Second-stage larva in situ..................... 215
40. Blphara seutellata: Basal portion of tracheal "funnel "............. 216
41. Cop ra codnnata: Adult female and details .................... 219
42. Map showing ditribution of Cormpilura coninnimta in Massachuset. 222
43. Tachina larrarum: Adult female and head in profile................. 225
41. Chalds f pes: Adult ............................................ 241
45. Chalisaipes, female: Hind femur and tibia, showing markings... 212
4B. Chalcis obseurata, female: Hind femur and libia, showing markings.. 242
47. als pe: Full-grown lara from gipsy-moth pupa .............. 243
4 Chalipe: Pupa side view .............................. 243
49. Chasaipes: Pupa ventral view .............................. 243
50. Gipsy-moth pup, sh owingexit holes of Chalis flaripes ............ 243
1. Monodoto rus res: Adult female.............................. 244
52. onodonto erus....................................... 49
53. a onodontomeru..................................... 249
54. Pupa, side viw ..................... ..... 249
5. Ms reus: Pupa, ventral view ........................ 249
56. Gipsy-moth pupa shoing exit hole left by Monodontoers a s.... 250
horamma in act of ovioition in an e of the brow-tail moth. 256
12 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
FIG. 58. Eggs of the brown-tail moth, a portion of which has been parasitized
by Trichogramma sp............................................-- 257
59. Larve of Pteromalus egregius feeding on hibernating caterpillars of
the brown-tail moth..............---.......................... .. 262
60. Portion of brown-tail moth nests, torn open, showing caterpillars
attacked by larve of Pteromalus egregius......................... 263
61. Apanteles lacteicolor: Immature larva from hibernating caterpillar of
the brown-tail moth..........................................---. 263
62. Meteorus versicolor: Immature larva from hibernating caterpillar of
the brown-tail moth........--....-------........-- ...----...-----..... 264
63. Zygobothria nidicola: First-stage larve in situ in walls of crop of hiber-
nating brown-tail moth caterpillar............................... 264
64. Compsilura concinnata: First-stage larva ....-...................... 265
65. Pteromalus egregius: Adult female ....---...----.... .......--....... 269
66. Pteromalus egregius: Female in the act of oviposition through the silken
envelope containing hibernating caterpillars of the brown-tail moth. 274
67. Apanteles lacteicolor: Adult female and cocoon....................... 279
68. Meteorus versicolor: Adult female and cocoons -----.... ...-----------.. 287
69. Zygobothria nidicola: Adult female and details -...--..-- ........-... 290
70. Pales pavida: Adult female and details.............................. 331
71. Pales pavida: Second-stage larva in situ in basal portion of integu-
mental "funnel" .............................................. 302
72. Pales pavida: Integumental "funnel," showing orifice in skin of host
caterpillar..................------...... ......................... 302
73. Eudoromyia magnicornis: Adult female and details---................... 303
74. Eudoromyia magnicornis: First-stage maggot and mouth hook ........ 333
THE IMPORTATION INTO THE UNITED STATES OF THE
PARASITES OF THE GIPSY M1OTH AND THE BROWN-
A REPORT OF PROGRESS,
WIH SOME CONSIDERATION OF PREVIOUS AND CONCURRENT EFFORTS OF THIS KIND.
By L. 0. HOWARD,
hief, Bureau of Entomology.
As il appear from the opening portion of this bulletin, which
gives an accot of previous work in the practical handling of natural
eneies, carried on in various parts of the world, nothing comparable
to the work which is to be described has eer before been und ertaken.
As will appear also, most of the successful work in this direction has
been done with the fixed scale insects. The exceptions to this gen-
eral statement among the measurabli succesful efforts have been the
itroduction of parasits of the sugar-cane leafhopper into Hawaii,
some reported work t in the introduction of South American natural
enemies of fruit flies into Western Australia, an the introduction of
one of the many European enemies of the codling moth from Spain
into Califoria; but it does not appear that practical results of aii
very great value have been achieved by the last two introductions
alth iniformation from Western Australia is scanty. At the
time when the work began nothing practical ha(d been accomplished
with the natural eneies of any lepidopterous insects, and in the whole
history of the practical handling of parasites no work of this character
has ever been attempted upon any thing like the large scale with
which the present work has been caried on. Some studies had already
been made both by the by the ritr Fk on he subject of the
ltensive parasitism of two native species of American moths, and
for ears the bureau had ben keeping records f the rearings of
parasites of lepidopterous insects as well as of other; moreover, the
writer had made a careful stuthe of th rc the rearings of
menopterous parasites from host insects all over the world and
had accumulated an enorous catalogue of such records. Never-
thelss te initial work on such a scale was experimental in its charac-
ter. It smed to the writer that by attempting to reproduce in New
14 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
England as nrearly as possible the entire natural environment of the
gipsy moth and the brown-tail moth in their native homes, similar
conditions of comparative scarcity could surely be reached, and this
view he still holds with enthusiasm. Naturally, in the course of the.
work as it progressed year after year his ideas have been changed as
to methods, and very great improvements have been made upon the
earlier methods, largely through the intelligence and ingenuity of
the junior author of this bulletin. Moreover, the careful, intensive
studies which have been made at the gipsy-moth parasite laboratory
by the junior author and a corps of trained assistants, aided by
abundant material, funds, and supplies, have resulted not only in the
ascertainment of very many facts new to science, but in the accumu-
lation of such facts to such a degree as to enable generalizations of a
novel character and of a sounder basis than could have been had
under other conditions. Many points are brought out in this bulletin
which will doubtless be entirely new to the trained scientific reader.
Mistakes have been made and wrong conclusions have been drawn
from time to time, but these have been corrected, and we are now in
a fair way to see a favorable result from the long and expensive work.
The initial idea was that since a large percentage of gipsy-moth
caterpillars or brown-tail moth caterpillars in Europe contains para-
sites each year, therefore if these caterpillars were brought to America
in large numbers from every possible place we could not fail to rear
from them an abundance of adult foreign parasites. This idea was
sound, and in following it out we have constantly improved the
methods-methods of collection, of packing, of shipment, and of
subsequent rearing. Very large numbers of parasites have been
It was first thought that when parasites had been reared in suffi-
cient numbers they should be widely distributed in small colonies, on
the theory that each colony would remain in substantially the same
general locality and would increase and spread from that point. This
idea was a natural one and was fully justified by previous work which
had been (lone with parasites of other groups of insects, but in this
case it proved to be erroneous, and valuable time and valuable speci-
mens were lost. Eventually it was shown to be of prime importance,
first to establish a given species of parasite in this country, and not
until this has been accomplished to pay any attention to the matter
of dispersion. It seems to be the first instinct of many species that
have been imported to spread widely. Therefore, if the colony put
out be a small one the individuals composing it spread rapidly
beyond all means of meeting and of mating, and thus the colonies in
man y instances were lost. By rearing in the laboratory, however,
until colonies of at least a thousand are to be had, such colonies
while dispersing are much more likely to remain in touch, mate, and
By me thods based upon the first idea, an d by the subsequent modi-
fication of the second idea, some of the most important natural ene
mies of both species have been established in the United States to a
certainty. It has been found with several species that they could not
be recovexed until after three years had elapsed from the time of the
ori~RI colonization; hence it follows with a reasonable certainty
that other species which have not been recovered will ultimately be
recvered as reult of colonization one, two, and three, and even
perhaps four yeairs ago. It is deemed, however, at thi time that
nearly a m uch has been accompli1hed as can be accomplished by the
earlier methods, and subsequent efforts will be devoted to a more
specfic -atteImpt to import the species still lacking several of which
are known in their original homes to be of very great iuportance. As
will be pointed out elsehere, atteml)ts will also be made to import the
peies wichl, wile of lesser illportamre at lhome, nma hee 1fill in
gaps ald rma possibly niultil)ly to an unprecedented extent in the
fce of new conditions la a supertalunlldalce of host ni1erial.
e111 wrk has been going on since 1903. Nothing haos been )pb-
lish (l conlcerin its p rogre exclpt Ithe shrt accounts in the annual
reports of the writer submitted each year to the Secretar of Agri-
cultre, and except a bulletin on the general subject prepare by the
junior author and published by thle State forester of Mssachusetts
It is hoped that the resent account will be deemed a satisfactory
reply to all expressed desire for information as to progress.
The joint authorship of the bulletin is deneed desirable by both
author, bt the writer takes it upon himself to sig this introduction
for the explicit purpose of stating in his own wa': the conditions under
ic it has been prepared. The work from the beginning has been
under the direct supervision of the wrier, and he is therefore to be
held responsible for any failures in the speedy accomplishment of
results, but the greatest credit in bringing about the results which
have been accomplished, he wishes frankl to state, belongs to Mr.
Fiske. Followi the breakdown in health of Mr. E. S. G. Titus in
the spring of 1907, as is shown in the bulletin, Mr. Fiske was stationed
at the parasite laborator and has since been given every freedom in
the conduct of its affairs. Nearly every suggestion hich he has
made, while it has been fully discussed by the two of us, has been
adopted. The ingenuity which he has displayed in matters of method
and the broad grasp which he has shown of the whole phenomena of
parasitism i insects, together with his competent and practical
grouping of his ideas, deserve every praise. Such portions of the
bulletin as were dictated by the writer have received the editorial
criticism of the junior author, and the portions preared by the latter
16 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
have received a most careful consideration and editorial pruning of
the writer. Mr. Fiske, by virtue of his practical residence at the field
laboratory and of his intimate charge of all the field notes and labo-
ratorv notes, has prepared all of the matter in this bulletin relating to
the laboratory and field end, subject, of course, to the writer's revision.
The rest has been prepared by the senior author.
Acknowledgements of assistance should be made by the score. The
State authorities of Massachusetts, the admirable corps of laboratory
and field assistants, and above all the very numerous foreign officials,
voluntary assistants, and paid observers have united to make the
undertaking possible. Their individual names are all mentioned in
the following pages in connection with the parts they played, but the
Governments of Austria, France, Germany, Hungary, Italy, Japan,
Portugal, Russia, and Spain should especially be thanked in an official
publication like this for the 'assistance given by the officials of these
PREVIOUS WORK IN THE PRACTICAL HANDLING OF NATURAL
ENEMIES OF INJURIOUS INSECTS.
Two very thorough and careful geneial papers on the subject of the
practical handling of natural enemies of insects, treating the subject
from the different points of view, including the historical side, have
been published in the last few years. The first of these, entitled "The
Utilization of Auxiliary Entomophagous Insects in the Struggle
against Insects Injurious to Agriculture," by Prof. Paul Marchal, of the
National Agronomical Institute of Paris, was published in 1907,1 and
was partly republished in English in the Popular Science Monthly in
1908.2 The other, by Prof. F. Silvestri, of the Royal Agricultural School
at Portici, Italy, entitled "Consideration of the Existing Condition of
Agricultural Entomology in the United States of North America, and
Suggestions which can be Gained from it for the Benefit of Italian
Agriculture," was published in 1909." This paper was in part trans-
lated into English and published in the Hawaiian Forester and Agri-
cult urist for August, 1909. Both of these papers should be consulted
iby lersons wishing to inform themselves thoroughly on this question.
For the present purpose, treatment of the subject must be brief.
The studly of parasitic and predatory insects is old. Silvestri has
lointedl out tIhat Aldrovandi (1602) was the first to observe the exit of
tlte llrv\i of Apant(les glomeratus L. (which he supposed to be eggs)
frromt tlw (m cnon cabbage caterpillar, and that Redi (1668) pub-
lis-lwd t ie same observation and another on insects of different
species Iorn from the same pupa. A later writer, Vallisnieri (1661-
L! m~ a t oI t iNt ionh l 1 groium al I[nst it ul e (Superior School of Agriculture), se.ond seris vo6. f, no.
2, p1 2' I 4, Iri, Ir07.
I' *11: r A I hw MonthlybI, ol. 72, pp. 3T5 .L 4>17 41", \Jpril and May, 1908.
a Ylu11ti. oV t h n* Pe y of[ al i.an Agrultiris*t s oI. 14, no. 8, pp. 3 &367, Apr. 0, 19 .
PR S WORK WITH INSECT PARASITES 17
1730) tly the first to discover the real nature of this phe-
nomenn and to realize the existence of true parasitic insects. Reau-
mur (1831757) and De Geer (1720-1778) each studied the life his-
living ect with great care and among these worked out
te b of number of parasites. Very many descriptive works
ontes were published in the closing years of the eighteenth
and the benning of the nineteenth century, especially by
an (1771828), Nees ab Esenbeck (1776-185), Gravenhorst
(1777-1857), Walker (publishing from 1833 to 1861), Westwood
(pufrom 1827 on through nearly the whole of the century),
(publishing from 1841 on), and Spinola (1780-1857).
Many later writers have contributed to the systematic study of
insects, among them Holmgren and Thomson, of Sweden;
Mayr, of ustria; Motschulsky, of Russia; Ratzeburg, lartig, and
iedeknecht, of Germany; Wesael, of Belgium; aliday,
Marshall, and Cameron, of England; Rondani, of Italy; Brull",
Giraud, Decaux, and others in France; Provancher, of Canada; anld,
in America, Cresson, Riley,ey, oward, Ashea, Crawford, Viereck,
Brues, Girault, and others
The best contribution appearing in Europe and devoted to the
biology of hymenopterous parasites, and especially consideration of
their relations to their hosts, was that by Ratzeburg, whose great
work entitled "Die Ichneumonen der Forstinsekten," was a standard
for many years. Ratzeburg understood the rkle played by parasites
in the control of forest inscts, but did not believe that this control
could in any way be facilitated by man.
EARLY PRACTICAL WORK
Froggatt has pointed out that probably the 'arliest suggestion
made regarding the artificial handling of beneficial insects s as printed
in Kirby and Spence's entomology (1816), were the authors called
attention to the value of the common English ladybird as destroyi
the hop aphis in the south of England. "If we could but discover a
mode of icreasing these insects at will, we migt not only clear our hot-
houses of apides by their means, but render our crops of hops much
more crtain than thy are now." As a matter of fact, gardeners
and florists in England for very many years have reco zed the value
of the ladybirds and have transferred them from one plat to another.
of. A. Trotter, of the Royal School of Viticulture at Avellino,
y, has recently pointed out in an interesting paper entitled "Two
Precursors in the Application of Carnivorous Insects," published in
diin 1908.1 that probably the first person to make a practical
1 liction of the natural enemies of injurious species was Prof.
SRedia, vol. 5, pp. 126-132, Florence, 1908.
18 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
Boisgiraud, of Poitiers, France, in 1840. Prof. Trotter found this
reference in a little-known paper by N. Joly, published in 1842, and
entitled "Notice of the Ravages which Liparis dispar L. has made
around Toulouse, followed by some Reflexions upon a Method of
Destroying Certain Insects." It seems that Boisgiraud, about 1840,
freed the poplars along a road near Poitiers of the gipsy moth by
placing upon them the carabid beetle Calosoma sycophanta L., and
destroyed earwigs in his own garden by placing with them a rove
beetle (Staphylinus olens Mfill). He also experimented against the
same insect with the ground beetle Oarabus auratus L. His experi-
ment must have become rather well known at the time, since Prof.
Trotter points out that in 1843 the technical commission of the
Society for the Encouragement of Arts and Crafts of Milan offered a
gold medal to be given in 1845 to the person who in the meantime
should have undertaken with some success new experiments tending to
promote the artificial development of some species of carnivorous
insects which could be used efficaciously to destroy another species of
insect recognized as injurious to agriculture. This offer drew forth a
memoir from Antonio Villa, a well-known writer on entomology, who
had previously confined himself to the Coleoptera, entitled "The Car-
nivorous Insects used to Destroy the Species Injurious to Agricul-
ture." This memoir was presented December 26, 1844, and he advo-
ca.ted the employment of climbing carabid beetles for tree-inhabiting
forms, rove beetles to destroy the insects found in flowers, and ground
beetles for cutworms and other earth-inhabiting forms.. The paper
of Villa was praised in certain reviews and criticized in others. It
seems to have been entirely lost sight of in later years.
A later Italian writer, Rondani, who devoted himself for the most
part to systematic work, appreciated the practical importance of
parasite work and published tables giving the host relations of differ-
ent species. His work influenced many arguments in the dispute
which sprang up in Italy about 1868 as to the usefulness of insec-
tivorous birds to agriculture, and Silvestri calls attention to the fact
that Dr. T. Bellenghi was referring to Rondani when, in 1872, he spoke
what Silvestri calls "the prophetic words:" "Entomological para-
sitism has a future, and in it more than in anything else Italian agri-
culture must put its faith."
PERMITTING THE PARASITES TO ESCAPE.
The earliest published suggestion as to the practical use of para-
sites of injurious insects, by permitting the parasites to escape while
the host insect is killed, appears to have been made by C. V. Riley
when State entomologist of Missouri. Writing of the rascal leaf-
crumpler (Mineola indiginella Zell.) in his Fourth Report on the
PREVIOUS WORK WrH INSECT PARASITES. 19
Insects of Missouri,' he advocated the collecting of the winter cases of
the destructive insect and placing the cases in small vessels in the
ceter of a meadow or field, away from any fruit trees, with the idea
that the worms would be able to wander only a few yards and would
perish from exhaustion or starvation, while their parasites would
escape and fly back to the fruit trees. It is stated that this method
was put in practice later by D. B. Wier with success.
A French writer, F. Decaux, the following year made practically
the same suggestion with regard to apple buds attacked by Anthono-
mus. He advised that instead of burning these .buds, as was gener-
ally one, they be preserved in boxes covered wit gauze, raising the
latter from time to time during the period of issuing of parasites so as
to permit the to escape. In 1880 lie put this method in practice,
and collected in Picardy buds reddened by the Anthonomius from
800 apple trees, and thus accomplised the destuction of more than
1,000,000 individuals of the Anthonomus, setting at liberty about
250,000 parasites which aided the following year in the destruction of
the weevils. The following year the same p)rocess was repeated, and,
the orchards being isolated in the middle of cultivated fields, all serious
damage from the Athonomus ws stated to have been stopped for
Practically the same suggestion was made later, in 1877, by J. I.
Comstock, in regard to the imported cabbage worm (Pontia rape L.).
Constock deprecated the indiscriminate crushing of the chrysalids
collected under trap boards, on account of the large percentage which
contained parasites. He recom ended instead the collecting of the
chrysalids and placig them in a box covered with a wire screen which
should permit the parasites to escape and at the same time confine the
butterflies so that they could be easily (estroyed. The same author,
in his report upon cotton insects, reco ended a siilar course with
the pupae of the cotton caterpillar (Alabam argillacea iibln.).
Riley later recommended the same l1an for the bagworm ( Thyridop-
teryx ephemereform is Haw.); Berlese in Italy recommended it for the
grapevine Cochylis, and Silvestri for the olive fly (Dacus olew Rossi),
for Prays oleellus Fab., and for Asphonylia lupini Silv.
Writing on the Hessian fly, Marchal has pointed out that the
destruction of the stubble remaining in the field after harest may
have unfortunate consequences, for if this is done a little late there is
a risk that all of the destructive flies will have emerged and aban-
doned the stubble, exposing to destruction only the parasites whose
part would have been to stop the invasion the following year. Mar-
chal also points out that Kieffer has show that one of the measures
SRley, C. Fourth Report on the Insects of Miori, p. 40, 1871.
*An excellent article covering these general questions was published by Decaux in the Journal of the
National Horticultural Society of France, vol. 22, pp. 158-184, 1899.
*Cotton Insects, pp. 230-231, Washinton, 1879.
20 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
advised for 'the destruction of the wheat midge (Coarinia ritic
Kirby), namely, burning the debris after thrashing, has only an
injurious effect, for, while it is true that the pups of the midge are to
be found in this debris, it should be remembered that the healthy
nonparasitized larve of the midge transform in the ground, while
those which remain in the heads are, on the contrary, parasitized.
Still another method of encouraging parasites is pointed out by
Marchal and Silvestri. It is to cultivate in the olive groves various
plants upon which allied insects live which are parasitized by the same
species of parasites as the olive fly. This idea, independently devel-
oped in the United States, has been practically used by Hunter in the
fight against the cotton-boll weevil. Allied insects feeding in certain
weeds along the borders of the cotton fields have parasites capable of
attacking the boll weevil. Careful study of the biology of these allied
weevils and of their parasites resulted in the gaining of the information
that if the weeds are cut at a certain time the parasites are forced to
attack the cotton-boll weevil in order to maintain their existence;
actual experimentation has resulted in the very considerable increas-
ing of the percentage of parasitism of the cotton-boll weevil in this
THE TRANSPORTATION OF PARASITES FROM ONE PART OF A GIVEN
COUNTRY TO ANOTHER PART.
In 1872 attempts were made by Dr. William Le Baron, at that time
State entomologist of Illinois, to transport Aphelinus mali Le Baron,
a parasite of the oyster-shell scale of the apple (Lepidosaphes ulmi L.)
from one part of the State of Illinois to another portion of the same
State where the parasite seemed to be lacking. Some slight success
was reported, and at the end of the year it was stated that the para-
site had become domiciled in the new locality, but, as this parasite
subsequently proved to be one of general American distribution, the
experiment can not- be said to have been worth while except in a very
In France, F. Decaux, above quoted, in 1872, made some experi-
ments in the transportation of parasites from one locality to another.
Riley, in his third report as State entomologist of Missouri (1870),
in considering two parasites of the plum curculio, stated that he
intended the following year, if possible, to rear enough specimens of
Sigalpihus curculionis Fitch to send at least a dozen to every county
seat in the State and have them liberated in someone's peach orchard.
There seems, however, to be no record that this was ever done.
In 1880, in his report on the parasites of the Coccide in the collec-
tion of the Department of Agriculture,' the senior author called atten
ti on to the fuct that with the parasites of scales the matter of trans-
I Annual Report U. S. Department of Agriculture for 1880, p. 351.
P IOUS WORK WITH INSECT 21
portation from one part of the country to another becomes easy, since
all that has to be done is simply to collect tws bearing the scales,
preferably during the
winter months, and
ca them to non-
the parasites being
dormant and pro-
tected each by the
scale of the coccid -
which it had de-
stroyed and it was
mended that the im-
portant parasite of
the black scale (Sais-
set olete Bern.), de-
scribed in the article Y. L-Poly s himai, a parsite of the Hsan fly: Adult.
as Tomocera caifor- Greatly enlarged. ( From Webster.)
nica, could be readily
carried from California and utilized to destroy Lecaniumi scales
in the Southeast.
Excellent ork in this direction 1ha been d(one of
late year by the Bureau of Entomology. li the study
of the Hessian fly (Mayetola destructor Say), under
Prof. F. M. Webster, early-sown plats of wheat at
Lansing, Mich., and Marion, Pa., in 1906, were very
seriously attacked by the essian fly, but when ex-
amined carefully at a later date fully 90 per cent of the
flaxseeds (pupa) were found to have been stung by a
hymenopterous parasite, Polygiot~ s hie(wlis Forbe:;
(figs. 1, 2), and to contain its developing larva. A
field of wheat near Sharpsburg, d., was found to be
infested by the fly, and examination indicated the ab-
sence of the parasite. On Apil 8, 1907, a large num-
.2.-Polygnot ber of the parasitized flaxseeds from Marion, Pa.,
iema: Adults were brought to Sharpsburg and placed in the field.
which hae d On July 8 an examination of the Sharpsburg field
the axseed howed tat the parasites had taken hold to such an
the ean extent that of the large number of flaxseeds taken and
fly and are ready
to e Much brought to the laboratory for investigation not one
s tr.) was found which had not been parasitized. Additional
material secured from Sharpsburg in the spring of 1908
in the same locality showed all of the Hessian flies to be parasitized.
In the same way excellent results have been obtained in the investi-
ation of the cotton-boll eevil under Mr. W. D. Hunter. the
22 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
summer of 1906 a number of parasites were taken from Waco, Tex.,
and liberated in a cotton field near Dallas, Tex., and apparently by
this means the mortality rate due to parasites was raised in a few
weeks about 9 per cent. Later, parasites were introduced from Texas
into Louisiana and increased the niortality of the weevil. Work of
this character is still being carried on by Mr. Hunter, and elaborate,
although as yet unsuccessful, experiments have been made by Web-
ster in the transfer of the hymenopterous parasite Lysiphlebus tritic
Ashm. (fig. 3) from southern points into Kansas wheat fields for the
destruction of the spring grain aphis or so-called "green bug" (Toxop-
tera graminum Rond.), definite results being prevented by the occur-
rence of the parasite throughout the range of the destructive insect,
parasitic, as it is, upon other species of plant lice.
Prof. S. J. Hunter, of the University of Kansas, however, in the
Bulletin of the University (vol. 9, p. 2) states that he was able, in
1908, to hasten the destruction of the Toxoptera in Kansas by the
importation of Lysiphlebus from some other point.
In the last two years
some very interesting
work has been carried
on by the State Horticul-
tural Commission of Cal-
ifornia in.the way of col-
lecting Coccinellidae on a
S- large scale in their hiber-
---- nating quarters, boxing
FIG. 3.-Lysiphlebus tritici attacking a grain aphis. Enlarged. them, and sending them
(From Webster.) to different parts of the
State for use against plant lice upon truck crops. The biennial report
of the commissioner of horticulture for 1907-8, published in Sacra-
mento in 1909, for example, indicates that 50,000 specimens of the
ladybird beetles Hippodamia convergens Gubr. and Coccinella cali-
fornica Mann. had been so collected. This, however, was very small
compared to the scale upon which these insects were collected dur-
ing the winter of 1909-10. Mr. E. K. Carnes, of the commission,
writing to the Bureau of Entomology under date of March 14, 1910,
makes the following statement:
We have quite a sight at the insectary now-over a ton of Hippodamia convergens,
boxed in 60,000 lots each, screened cases, and in our own cold storage. We handle
them in large cages, run them into a chute, and handle like grain. They are for the
melon growers of the Imperial Valley.
This species collects in large numbers late in summer and early in
the autumn at the bases of plants in the mountain valleys and can
easily be collected by the sackful. The actual good accomplished by
the distribution of these ladybirds among the melon growers has not
PREVIOUS WORK WITH INSECT PARAITS. 23
yet been reported upon, but theoretically speaking the experiment
should have excellent results.
THE TRANSFER OF BENEFICIAL INSECTS FROM ONE COUNTRY TO
Dr. Asa Fitch, for many years State entomologist of New York,
was probably the first entomologist in America, or elsewhere for that
matter, to take into serous consideration the question of the transfer
of beneficial insects fro one countr to another. In 1854, following
a disastros attack upon the wheat crop of the eastern United States
by the wheat midge (ontariia fitici Kirby), a species that had been
accidentally introduced from Europe during the early part of that
century, Dr. Fitch, who had made a careful study of the insect both
in this countr and from the European records, was struck with the
fact that in Europe the isect in ordinary season did no damage, and
that when occasionally it became so multiplied as to attract notice it
was but a transitory evil which subsided soon and was not heard of
gain for a number Of years. He was aware that ill Europe certain
parasites of this insect were found, and, comparing the insects taken
from wheat in flower il France wth those taken from wheat in flower
in New York, he found that in France the wheat midge constituted
but 7 pr cent of the isects thus taken, while its parasites constituted
85 per cent; whereas in New York the wheat midge formed 59 per
cent of the isect thus captured, and there were no certain parasites.
He speculated as to the cause for this extraordinary difference and
There must be a case for this markable difference What can that cause be?
I an impute it to only one thing; we here are detitute of natures appointed means
for repressing and subduing this insect. Those other insacts which have been created
for the purpose of quelling this species and keeping it restraied ithin its appropriate
sphere have never yet reached our shore. We have received the evil without the
remedy. And thus the midge is able to multiply ad flourish, to revel and riot, year
after ear, without let or hindrance. Th enly would seem to be the principal if
not thesole cause why the career of this iset here is so very different from what it is
in the Old World.
Quite naturally after this train of reasoning had entered his brain,
Dr. Fitch made an effort to introduce the European parasites of the
wheat midge, and in May, 1855, addressed a letter to John Curtis,
the faous English economic entomoloist, and at that time president
of the Entomological Society of London, informing him of the immense
amount of damage done by the midge in America and suggesting the
manner in which parasitized larve could be secured in England and
transmitted alive to this country. Mr. Curtis was ill and on the point
of starting for the Continent, but laid the letter before the Entomolog-
ical Society of London, which resulted in the adoption of a resolution
24 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
to the effect that if any member of the society should be able to find
parasitized midges he should send them to Dr. Fitch.
Nothing ever came of this effort, but it is of interest on account of
its apparent priority over other experimentation of this kind.
The next international attempt seems to have been made in 1873,
when Planchon and Riley introduced into France an American-
predatory mite (Tyroglyphus phylloxerm Riley) which feeds on the
grapevine Phylloxera in the United States. The mite became estab-
lished, but accomplished no appreciable results in the way of checking
the famous grapevine pest.
In 1874 efforts were made to send certain parasites of plant lice
from England to New Zealand, but without results of value, although
Coccinella undecimpunctata L. is said to have become established.
In 1883 Riley imported the braconid Apanteles glomeratus into the
United States from Europe, where it is an abundant enemy of the
imported cabbage worm (Pontia rapeX L.). This species has since
established itself in the United States and has proved a valuable
addition to the North American fauna.
THE AUSTRALIAN LADYBIRD (NOVIUS CARDINALIS MULS (IN THE UNITED STATES).
But all previous experiments of this nature were completely over-
shadowed by the remarkable success of the importation of (Vedalia)
Novius cardinalis Muls. (fig. 4), a coccinellid beetle, or ladybird,
from Australia into California in 1889. The orange and lemon groves
of California had for some years been threatened with extinction by
the injurious work of the fluted or cottony cushion scale (Icerya pur-
chasi Mask.) a large scale insect which the careful investigations of
Prof. Riley and his force of entomologists at the United States
Department of Agriculture had shown to have been originally
imported, by accident, from Australia or from New Zealand, where it
had originally been described by the New Zealand coccidologist, the
late W. M. Maskell. The Division of Entomology had been for several
years engaged in an active campaign against this insect, and had dis-
covered washes which could be applied at a comparatively slight ex-
pense and which would destroy the scale insect. It had also in the
course of its investigations discovered the applicability of hydrocyanic-
acid gas under tents as a method of fumigating orchards and destroy-
ing the scale. The growers, however, had become so thoroughly dis-
heartened by the ravages of the insect that they were no longer in a
frame of mind to use even the cheap insecticide washes, and many of
them were destroying their groves. In the meantime, through some
correspondence in the search for the original home of the scale insect,
Prof. Riley hald discovered that while the species occurred in parts of
Australia it was not injurious in those regions. In New Zealand it
PREVIOUS WORK WITH INSECT PARASITES. 25
also occurred, but was abundant and injurious. He therefore argued
that the insect was probably introduced from Australia into New Zea-
lad, and that its abundance in the latter countr and its relative
scarcity in Australia were due to the fact that i its native home it
was heId in subjection by some parasite or natural enem, and that in
the introduction into New Zealand the scale insect had been brought
in alone. The same thi, he arued, had occurred in the case of the
introduction into the United States. He therefore, in his annual
report for 1886, recommended that an effort be made to study the
natural enemies of the scale in Australia and to introduce them into
California; and the same year the leain fruit growers of California
in convention assembled petitioned Congress to make appropriations
for the Department of Agriculture to undertake this work. In Feb-
ruary, 1887, the Department
of Agriculture received speci-
mens of an Australian para-
site of Icetra from the late
Frazier S. rawford, of Ade-
was a dipterous insect known
as Lestopkonus iceryx Will.,
and for some time it was cocn-
sidered, both by Prof. Riley
and his correspo ndents and
agents, that the importation
of this particular parasite
offered the best chances for
good resultso. 4.-The Australin ladybirdi ( Novus cardinalis), an
Neither the recom enda- irt ny o the fluted scale: a, Ladybird lane
io ofeeding on adult female and egg sac; b, pupa; c, adult
Ons of Prof. Riley nor of ladybird; d, oringe twig, showing s(ale and lady-
the then commissioner of birds. a-c, Enlarged; d, natural size. (From Mar-
agriculture, Hon. Norman J. lat.)
Colman, nor the petitions of the California horticulturists gained
the needed congressional appropriations, and, since flthere appeared
at that time annually in the bills appropriating to the entomo-
logical service of the Department of Agriculture a clause prevent-
ing travel in foreign parts, it became necessary to gain the funds
for the expense of the trip to Australia from some other source.
A movement was started in California to raise these funds by
private subscription, but it was never carried through. In an
address given by Prof. Riley before the California State Board of
Horticulture at Riverside, Cal., in 1887, he repeated his recomnimenda-
tions. During the summer of 1887 he was absent in Europe, and the
senior author, who was at that time the first assistant entomologist
of the department, by correspondence secured from _Mr. Crawford
numerous specimens of Icerya infested by the Lestophonus above
26 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
mentioned. During the winter of 1887-88 preparations were being
made for an exhibit of the United States at the Melbourne Exposi-
tion, to be held during 1888, and Prof. Riley, after interviewing the
Secretary of State, who had charge of the funds appropriated for the
exposition, was enabled to send an assistant, Mr. Albert Koebele, to
Australia at the expense of this fund. This result was hastened, and
Mr. Koebele's subsequent labors were aided by the fact that the
commissioner general of the United States to the exposition was a
California man, Mr. Frank McCoppin, and his recommendation, joined
to that of Prof. Riley, decided the Secretary of State in favor of the
movement. In order to partially compensate the exposition authori-
ties for this expenditure, another assistant in the Division of Ento-
mology, Prof. F. M. Webster, was sent out to make a special report to
the commission on the agricultural features of the exposition. Mr.
Koebele, who sailed from San Francisco August 25, 1888, was thor-
oughly familiar with all the phases of the investigation of the cottony
cushion scale, and had for some time been stationed in California
working for the Department of Agriculture. His salary was con-
tinued by the department and his expenses only were paid by the
Melbourne Exposition fund. He made several sendings of the Les-
tophonus parasite to the station of the Division of Entomology of the
Department of Agriculture at Los Angeles, where, under the charge
of Mr. D. W. Coquillett, a tent had been erected over a tree abun-
dantly infested with the scale insect; but it was soon found that the
Lestophonus was not an effective parasite.
On October 15 Mr. Koebele found the famous ladybird (Vedalia)
Novius cardinalis in North Adelaide, and at once came to the con-
clusion that this insect would prove effective if introduced into the
United States. His first shipments were small, but others continued
from that date until January, 1889, when he sailed for New Zealand
and made further investigations. Carrying with him large supplies
of Vedalia cardinalis, the effective ladybird enemy, he arrived in San
Francisco on March 18, and on March 20 they were liberated under
the tent at Los Angeles, where previous specimens which had survived
the voyage by mail had also been placed.
The ladybird larve attacked the first scale insect they met upon
being liberated from the packing cages. Twenty-eight specimens had
been received on November 30 by Mr. Coquillett, 44 on December 29,
57 on January 24, and on April 12 the sending out of colonies was
begun, so rapid had been the breeding of the specimens received alive
from Australia. By June 12 nearly 11,000 specimens had been sent
out to 208 different orchardists, and in nearly every case the colonizing
of the insect proved successful. In the original orchard practically
all of the scale insects were killed before August, 1889, and, in his
annual report for that year, submitted December 31, Prof. Riley
PREVIOUS WORK WITH INSECT PARASITES. 27
reported that the cottoy cushion scale was practically no longer a
factor to be considered in the cultivation of orages and lemons in
California. The followig season this statement was fully justified,
and since that time the cottony cushion scale, or white scale, or fluted
scae, as it is called, has no longer been a factor in California horti-
culture. Rarely it begins to increase in numbers at some given point,
but the Australia ladybirds are always kept breeding at the head-
quarters of the State Board of Horticulture at Sacramento, and such
outbreaks are speedily reduced. In fact, it has been difficult for the
State horticultural authorities to keep a sufficient supply of scale
insect food alive for the continued breeding of the ladybirds.
The same insect was introduced direct from California into New
ealand at a later date, and the same good results were brought about.
The Icerya is no longer a feature in horticulture in New Zealand.
NoVIUs iN PORTUGAL.
Still a tird strikig instance of the value of the Australian ladybird
was seen later in the case of Portugal. Icerya pprchas was robably
introduced into that country in the late eightie or early nineties from
her colonies in the Azores, to which point it was probably introduced
many years previously from Australia. The insect slread rapidly
and threatened the complete estrction of the orange and lemon
groves along the banks of the River Tagus. In September, 1896,
persons in Portugal applied to the senior author for advice as to the
most efficacious eans of fihting the sale insect, and a reply was
made urging them to make an effort to itroduce (Vedalia) Norius
ardinalis and sending information as to the success of the insect in
California. In October, 1897, the chief of the bureau was able to
secure, throuh the kidness of the State Board of Agriculture of
Califora, about 60 specimens of the ladybird, which were sent by
direct mail from Washington packed in moss. But five reached Por-
tugal alive, but these were so successfully cared for that there was a
numerous progeny. Another sending was made on the 22d of Novem-
ber folloing. These were received on the 19th of December and
proved successful. Early in September, 1898, the statement was
published in Lisbon newpapers that already colonies or stocks of the
Vedalia had been established on 487 estates, whence naturally many
others were formed by radiation; gardens and orchards that were
completely infested and nearly ruined were already entirely clean or
well on the way oward becoming so. Since that time the pest has
almost entirely disappeared. The bureau would not have been able
to assist the Portuguese Government to this admirable result had it
not been for the enlightened policy of the State Board of Horticulture
f California coninuing the breeding in confinement of these preda-
28 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
ceous beetles long after the apparent great necessity for such work
had disappeared in California, and had it not been for the courtesy of
the board in promptly placing material at its disposal.
ICERYA IN FLORIDA.
The general effect of the California success on the horticultural
world at large was striking, but not wholly beneficial. Many enthu-
siasts concluded that it was no longer worth while to use insecticidal
mixtures, and that all that was necessary in order to eradicate any
insect pest to horticulture or to agriculture was to send to Australia
for its natural enemy. The fact that the Vedalia preys only upon
Icerya and perhaps some very closely allied forms was disregarded,
and it was supposed by many fruit growers that it would destroy any
scale insect. Therefore the people in Florida whose orange groves
were suffering from the long scale (Lepidosaphes gloveri Pack.) and
the purple scale (Lepidosaphes beckii Newm.) sent to California for
specimens of the Vedalia to rid their trees of these other scale pests.
Their correspondents in California sent them specimens of fhe beetle
in a box with a supply of Iceryas for food. When they arrived in
Florida the entire contents of the box were placed in an orange grove.
The result was that the beneficial insects died, and the Icerya gained
a foothold in Florida, a State in which it had never before been seen.
It bred rapidly and spread to a considerable extent for some years,
and did an appreciable amount of damage before it was finally
Novrus IN CAPE COLONY.
Prior to the introduction of Novius into Portugal, Icerya puschasi
having been established at the Cape of Good Hope, the beneficial
ladybird was, after an unsuccessful attempt, carried from California
to Cape Town by Mr. Thomas Low, member of the Legislative
Assembly of Cape Colony, and on the 29th of January, 1892, living
specimens were placed in perfect condition in the hands of the depart-
ment of agriculture of Cape Colony. These specimens multiplied
and were reenforced late in 1892 by a new sending from Australia
made by Koebele. At the present time the Novius is perfectly
naturalized at the Cape.
NOVIUS IN EGYPT AND THE HAWAIIAN ISLANDS.
At the same time, through the United States Department of Agri-
culture and the courtesy of the State Board of Horticulture of Cali-
fornia, the Novius was sent to Egypt to prey upon an allied scale
insect, Icerya rgyptiaca Dougl., which was doing great damage to
citrus trees and to fig trees in the gardens of Alexandria, Egypt.
Six adult insects and several larve arrived in living condition at
VIOUS WORK WITH INSECT PARASITES. 29
Alexandria These multiplied so rapidly as to cause an almost
com t disappearance of the scales. Later the latter began to
in, but the Novius had not died out and also increased. The
Icea is still held in check in a very perfect-way.
In 1890 the Novus had been introduced into the Hawaiian Islands
for work aainst Icer urc with the same success.
ICERYA IN ITALY.
In1900 Icerya purchs was found also in Italy, in a small garden
at Portici, upon orange trees. By the autumn of 1900 it had mul-
tiplied so abundantly that the owner of the garden tried to stop
the trouble by cutting down the trees most badly ifested, without
botherii himself with the others, so that the infestation continued.
When Prof. Berlese's attention was called to it an attempt was first
made to destroy it by insecticides without success, and then No'ius
cardinis was imported from Portugal and from Aerica. The
following Jue the ladybird in both sexes was distributed in the
garden, prospered wonderfully, and multiplid rapidly. In July the
results were already evident; one could hardly find patches of Icerya
which did not show the work of Novus, and at the end of the month
it was difficult to find adult Iceryas with which to continue the
rearin in the laboratory for food for the reserve su)ply of Novius.
At the present time the multiplication of the scale insect has been
reduced to the point of practically no damage, but the original infes-
tation still persists andthe area of distribution of the scale insect
is slowly enlargi. It is found not only at Portici but in all the
little towns around Vesuvius and in the gardens in Naples; but the
presence of the ladybird allows the culture of oranges and lemons
to go on without interruption.
ICERYA IN SYiRA.
The latest utilization of the beneficial Novius is recorded by Sil-
vestri. It seems that about the year 1905 Icerya made its appearance
in Syia, and in July, 1907, Selim Ali Slam wrote to Prof. Silvestri
that it had spread so greatly about Beirut that it had almost
destroyed the trees Silvestri sent a shipment of Novius in July,
1907, and another one in August. The result was the same in Syria
as it had been in other countries; the Novius multiplied greatly and
produced the desired effect,
THE REASONS FOR THE SUCCESS OF Novis.
It thus appears that in the Novius we have an almost perfect
remedy against Icerya. There have been no failures in its intro-
Sin the above was written (n the autumn of 1909) still another success wit Novius has been by its
from California to osa by Dr. T. Shiraki, the entoologist of the Formosan Government,
wh under ate of Jan28 1910: "To-day it has relieved the region from Icerya and has reduced
Snumber to a racly neligible quantity."
30 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
duction to any one of the different countries to which it has been
carried. Its success has been more perfect than that of any other
beneficial insect that has so far been tried in this international work.
There are good reasons for this-reasons that do not hold in the
relations of many other beneficial insects to their hosts. In the first
place, the Icerya is fixed to the plant; it does not fly, and crawls very
slowly when first hatched, and later not at all. The Novius, however,
is active, crawls rapidly about in the larval state, and flies readily
in the adult. In the second place, the Novius is a rapid breeder, and
has at least two generations during the time in which a single genera-
tion of the host is being developed. In the third place, the Novius
feeds upon the eggs of the Icerya. And in the fourth place, it seems
to have no enemies of its own. This is a very strange fact, since other
ladybirds are destroyed by several species of parasites. For example,
as will be shown later, native American ladybird parasites brought
about a great mortality in the larve of the Chinese ladybird imported
from China into America at a later date by Marlatt. The hymenop-
terous parasites of the widespread genus Homalotylus feed exclu-
sively in ladybird larve, which are frequently also fairly packed with
the minute hymenopterous parasites of the genus Syntomosphyrum,
while the adults are often destroyed by Perilitus, Microctonus, and
The astonishing results of the practical handling of Novius drew
attention more forcibly than ever before to the possibilities of this
kind of warfare against injurious insects, and although its perfect
success as an individual species has never been duplicated, very
many efforts in this direction have been made, some of which have
met with measurable success and some with very positive results of
INTRODUCTION OF ENTEDON EPIGONUS WALK. INTO THE UNTED STATES.
In 1891, with the assistance of Mr. Fred Enock, of London, Riley
introduced puparia of the Hessian fly (Mayetiola destructor Say)
infested with the chalcidid parasite Entedon epigonus Walk. into
America. These were distributed among several entomologists dur-
ing the spring of 1891. One American generation was carefully
followed by Forbes in Illinois, and four years' later (in May, 1895)
the species was recovered by Ashmead at Cecilton, Md., where a
colony had been placed in 1891. Thus the introduction was appar-
ently successful, but if the species still exists in the United States
it must be rare, since extensive rearings of Hessian-fly parasites have
been madle by agents of the Bureau of Entomology in many different
parts of the country during the past few years and not a single speci--'
men of the Entelon has been recognized. The Maryland locality,
however, it should be stated, has not been visited by an entomologist
since Ashecad's trip in May, 1895.
PREVIOUS WORK WITH INSECT PARASITES. 31
OER INTRODUCTIONS BY KOEBELE INTO CALIFORNIA.
Mr. Koebele took a second trip to Australia, New Zealand, and the
Fiji Islands while still an agent of the Department of Agriculture,
but at the expense of the California State Board of Horticulture, and
in 1893 he resigned from the United States Department of Agriculture
and was emp loyed by the State Board of Horticulture of California
for still another trip to Australia and other Pacific islands. He sent
home a large numberof beneficial insects, nearly all of them, however,
coccinellids, Several of these species were established in California,
and are still living in different parts of the State. The overwheling
success of the iprtation of n ofius cardiais
was not repeated, but one of the insects
brought over at that time, namely, the ladybird
beetle zobiu~ vetrals Er. (fig. 5), an enemy
of the so-called black scale o(Saissetia Bern.),
was colonized in various parts of California, and
n districts where the climatic conditions proved
favorable its work was very satisfactory, nota-
bly in the olive plantations of Mr. Ellwood
Cooper, near Santa Barbara. Hundreds of
thousands of the beetles wNere listributed in
California and in some localities kept the black
scale in check. Away from the moist coast re-
ions, however, they proved to be less effective.
INTERNATIONAL NWORK WIT ENEMIES OF THIE BLACK
It will here be convenient to drop the chrono-
logical sequence with which the subject in hand I
has been treated and to refer to the introduction
of a very successful parasite of the black scale, ~'an i.-Rtzed oe:nv teu ,
whose work against this destructive enemy to blackseale: a, Adult lady-
olive and citrus culture in California for a time ird; b, larv. Mc en-
largd. (From M rlatt.)
seeed second only to the success of the Novius
against the Icerya. In 1859 Motchulsky described, under the name
Sctellista cyanea (fig. 6), a very curious little hymenopterous parasite
reared by Nietner from the coffee scale in Ceylon. Subsequently
this parasite became accidentally introduced into Italy and was sent
to the snior author for identification by Dr. Antonio Berlese as a
parasite of the wax scale, Cerop tes rusci L. As there are wax
scales (eropases f eniComst. and C. pediforis Comst.)
which are more or less injurious in Florida and the Gulf States, an
attempt was made, with Berlese's assistance, to introduce this para-
site at a convenient location at Baton Roue, La., with the further
32 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
assistance of Prof. H. A. Morgan at that place. Berlese's sending
arrived in good condition, and the parasites issued at Baton Rouge
and immediately bega to attack the native species. The importa-
tion was successful for a time, but the introduced species was finally
reduced to an insignificant number, presumably through the attacks
In the meantime Prof. C. P. Lounsbury, an American occupying
the position of entomologist of the department of agriculture at the
Cape of Good Hope, on his arrival at the Cape in 1395 and searching
for the usual cosmopolitan scale insects on fruit trees, failed to
find the black scale. He com-
mented on this fact in one of
his first-published papers, and
alluded to the severity of the
scale as a pest in California.
Shortly afterwards he found
the species, and sent the senior
ar thor specimens for identifi-
S/ cation in 1895, together with
parasites which he had reared
from it. Subsequent -cor-
respondence showed other
species, and eventually Scutel-
lista cyanea was forwarded.
Writing to Mr. Lounsbury
SSeptember 14, 1896, the chief
of the bureau made the follow-
ing suggestion: "I think para-
sitized black scales could be
sent to California to advan-
tage. Mr. Alexander Craw
FIG. 6.-Scutellista cyanea, an imported parasite of the would be the proper person to
black scale: Dorsal and lateral views of adult, with to whom to send- them."
enlarged details. Greatly enlarged. (From Howard.) Mr. Lounsbury made further
Mr. Lounsbury made further
studies, and commented in his 1898 report on the existence of
parasites. When this report met the eye of Mr. E. M. Ehrhorn, of
the State horticultural commission, Mr. Ehrhorn wrote Mr. Louns-
bury, under date of December 22, 1899, asking him to send a colony
of the parasite. Mr. Lounsbury had in the meantime, in a letter
to the senior author, suggested that in order to gain authority to
spend time over the matter and incur necessary expense it would be
desirable for the Secretary of Agriculture of the United States to
make a formal req.u:--t for these parasites to the secretary of agri-
culture of Cape Colony. This was done, and in May, 1900, Lounsbury
VIOUS WORK WITH INSECT PARASITES. 33
secured leave of absence and started for America, carring with him
a box of parasitized scales, and landed at New York on June 2.
Hisbox of parasites was at once forwarded to Washington, and the
Burea of Entomology notified Mr. Ehrhorn by telegram, repacked
the box, and sent it to California. Mr. Ehrho succeeded in tem-
porarily establishing the Scutellista indoors and out around his home
at Mountain View, Cal. September 19, 1900, Mr. C. W. Mally,
Lounsbury's assistant, sent two more boxes by post direct to Cali-
fornia, addressing them to S. F. Leib, of San Jose, notifying the
senior author to wire Mr. Ehrhorn to be on the lookout for them. A
third lot was sent October 31 of the sam year. These later sendings
were small, and both failed to yield living parasites. More were
requested, and on Lounsbury's return to South Africa box was
shipped in cool chamber to England and thencedirect to California
by express, Lounsbury's letter of February 28, 1901, to the bureau
stating: "To avoid extra delay in transmission the box goes direct
to California, but will you kindly have a nessage sent to Craw to
advise him of its coming?" Unfortunately the box was detained by
a customs officer at New York, but the bureau secured its lease by
the Government dispatch agent, Mr. I. P. Roosa. A few paraites
emerged after arrival, but failed to propagate. October 1, 1901,
Lounsbury started another sending by letter post to insure quick
transit and noninterference by customs. These boxes were delivered
to Mr. raw n October 31. Only four females of the Scutllista
were reared by Mr. Craw, and probably to these four females are due
all of the Scutellistas subsequently occurr in California. This is
the full stoy of the intrduction of the species, taken from the letter
files of the Bureau of Entomology and the letter files of Mr. Louns-
bury in Cape Town.
Mr. Craw was remarkably successful in his rearings, and during the
following three years constantly distributed colonies in different por-
ions of California. By July, 1902, he had distributed 25 colonies.
It was in the southern part of the State that the parasite did its best
work, and there for a time it surpassed the most sanguine expecta-
tions of everyone. It was established in every county south of Point
Conception and had become very plentiful in Los Angeles, Orange,
and San Diego Counties. In the coloization districts by midsum-
mer, 1903, it was estimated tat over 90 per cent of the black scale
had been destroyed. A year or so later there was great mortality
among these parasites caused by a sudden increase in numbers of
a predatory mite, Pedicloide enicosu New. (fig. 7), which
destroyed the larvo in vast numbers. The Scutellista gradually
recovered from this attack, and is at present to be found in very
many localities in California, keeping the black scale partly in check.
34 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
Another enemy of the black scale was imported in 1901. It is a
*small moth, Erastria scitula Ramb. (fig. 8), the larva of which feeds
in the bodies of mature scales, each larva destroying a number of
scales. An effort had been made by Riley to import this insect from
France in 1892, but without success. In 1901 Berlese sent the senior
author living pupae, which were at once forwarded to Craw and-
Ehrhorn in California. It was reported in 1902 that the insects had
been reared and liberated in Santa Clara, Los An-
geles, and Niles, Cal., but if the species was estab-
lished in the State it has not flourished and has
not recently been found.
A similar lepidopterous insect, Thalpochares coc-
ciphaga Meyrick, was brought over from Australia
in the summer of 1892 by Koebele and left by
him at Haywards, Cal., but the species evidently
THE HAWAIIAN WORK.
In 1893 Koebele resigned from the service of
FIG. 7.-Pedicu.oides the State of California and entered the employ-
enlarged. (From ment of the then newly established Hawaiian Re-
Marlatt.) public for the purpose of traveling in different
countries and collecting beneficial insects to be introduced into
Hawaii for the purpose of destroying injurious insects. Before
leaving California he had introduced a very capable ladybird, Crypto-
lemus montrouzieri Muls., which feeds upon mealy bugs of the genus
FIG. 8.-Erastria scitula, an imported enemy of the black scale: a, Larva from below; b, same, from
above; c, same, in case; d, case of full-grown larva; e, pupa; f, moth. Enlarged. (After Rou-
Pseudococcus. This insect flourished, especially in southern Cali-
fornia, and on arrival in IHawaii he found that coffee plants and
certain other trees were on the point of being totally destroyed by
the allied scale insect known as Pulvinaria psidii Mask. He at
PREVIOUS WORK WITH INSECT PARASITES. 35
once introduced this same Cryptolteus, which is an Australian
inst, with the result that the Pulvinaria was speedily reduced to a
condition of harmlessness.
It may be incidentally stated that within the past year efforts have
been made by the Bureau of Entomology to send the Cryptolsmus to
Malaga, Spain, for the purpose of feeding upon a Dactylopius. The
first attempt was unsuccessful, and the results of the last attempt
have not yet been learned.
Another importation of Koebele's into Hawai was the ladybird
occinela repanda Thunb. from Ceylon, Australia, and China, which
was successful in destroying plant lice upon ugar cane and other
crops. Writing in 1896, Mr. R. C. L. Perkins stated that Koebele
had already introduced eight other species which had become natu-
ralized and were reported as doing good work against certain scale
insects. Among other things he introduced Calcis obscurata Walk.
from China and Japan, which multiplied enormously at the expense
of an injurious lepidopterous larva (Omiodes blackburni Butl.) whiclh
had severely attacked banana and p)alm1 t rees.
Koebele's travels from 1894 to 1896 were through Australia. Cliina,
Ceylon, and Japan. In 1899 he left for Australia and the Fiji Islands,
and sent many ladybirds and parasites to tIawaii, especially to attack
the sle Ceroplastes rtbens Mask. The lawaiian Sugar Planters'
Association, an organization which was responsible for Koebele's
appointment, slubsequently employed M R. I, C. L. Perkins, Mr. G. W.
Kirkaldy, Ir. F. W. Terr, Mr. 0. H. Swezey, and Mr. F. Muir. By
the close of 1902 sugar planters were especially anxious concerning
the damage of an injurious leafhopper on the sugar cane, Pe erkin la
saccharicida Kirk. This insect had been accidentally introduced
from Australia about 1897, had increased rapidly, and by 1902 had
become a serious pest. Koebele had made an effort to introduce
parasites of leafhoppers from the United States into Hawaii, with
unsatisfactry results, and consequently in the sring of 1904 Koe-
bele and Perkins visited Australia and collected all possible parasites
of different leafhoppers. Altogether they suceeded in finding more
than 100 species. Of these the followin hymenopterous parasites
are said to have become acclimated in Hawaii: Amagrms (two species),
Paraag optabilis Perk. and P. perforator Perk. and Ootetrasticus
beatw Perk. These species are all parasitic upon the eggs of the leaf-
hopper. By the end of 1906 observations upon a certain plantation
indicated the destruction of 86.3 per cent of the eggs by these para-
sites. In addition to these egg parsites certain proctotrypid
parasites of hatched leafhoppers have apparently become established,
namely, Haplogonao itiensis Perk., Pseudognatous (two
species), d Echrodep fairchildii Perk. Three predatory
beetles, namely, Veraia f ata Erichs., V. lineola Fab., and Calli-
d testudinari Mls.. were also distributed in large numbers,
36 PARASITES OF GIPSY AND BROWN-TAIL MOTHS. .
The practical results of these importations seem to have been
excellent. There seems to be no doubt that the parasites have been
the controlling factor in the reduction of the leafhoppers.
The good work in Hawaii is still continuing. Koebele is now on a
visit to Europe to import the possible parasites of the horn fly
(HImatobia serrata Rob.-Desv.), Muir is trying to find an enemy to
a sugar-cane borer (Rhabdocnemis obscurus Boisd.), and other similar
work is under way.
AN IMPORTATION OF CLERUS FROM GERMANY.
An early attempt to import beneficial species into the United
States was made in 1892 by Dr. A. D. Hopkins, then entomologist
to the West Virginia Agricultural Experiment Station and now of
the Bureau of Entomology. A destructive barkbeetle, Dendroc-
tonus frontalis Zimm., was extremely injurious in that State in the
years 1889 to 1892, and Hopkins made the effort to import from
Europe another beetle, (Clerus) Thanasimus formicarius L., from
Germany. In Germany he collected more than a thousand specimens
of the Clerus, which he took with him to West Virginia and distrib-
uted in various localities infested by the barkbeetle. The following
year, however, the barkbeetle disappeared almost completely from
other causes, and the Clerus has not since been found.
MARLATT'S JOURNEY FOR ENEMIES OF THE SAN JOSE SCALE.
Another and later expedition was that undertaken by Mr. C. L.
Marlatt, of the Bureau of Entomology, in search of the natural
enemies of the San Jose scale. The question of the original home
of the San Jose scale (Aspidiotus perniciosus Comst.) had been a
mooted point. As is well known, it started in this country in the
vicinity of San Jose, Cal., in the orchard of Mr. James Lick, who had
imported trees and shrubs from many foreign countries. Mr. Lick
died before the investigation started, and no records of his impor-
tations were to be found. The scale was not of European origin,
since it does not occur on the continent. In the course of investiga-
tion it was found that it occurred in the Hawaiian Islands, in Japan,
and in Australia, but in the case of Australia and the Hawaiian
Islands it was shown that it had been carried on nursery stock from
California. In 1897 plants entering the port of San Francisco from
Japan were discovered by Mr. Craw to carry the San Jose scale.
Correspondence, however, seemed to point to the conclusion that it
had also been introduced into Japan from the United States. In
1901-2 Mr. Marlatt made a trip of exploration in Japan, China, anrd
other eastern countries, lasting more than a year. Six months
were spent in Japan, and after a thorough exploration the con-
clusion was reached that the scale is not a native of that country
OU WORK WITH INSECT PARASITES 37
and that wherever it occurs there it has spread from a center of
imported American fruit trees. Finally, as a result of this extended
trip, the native home of the San Jose scale Was found to be in northern
China in a region between the Tientsin-Pekin Road and the Great
Wall, and its original host plant was found to be a little haw apple
which grows wild over the hills. Into that region no foreign intro-
ductions of fruit or fruit trees had ever been made, and the fruits
in the markets were all of the native sorts. Here in China was found
everwhere present a little ladybird, hilocorus similis Rossi (fig. 9),
feeding in all stages upon the San Jose scale. One hundred and fifty
F. .Th Asiatic ladybird (Cilor imili), an mported enemy of the San Jose scale: a, Second
larval stage; sk of same; fulgown larva; d, metod of pupation, the pupa being retained
n the splt larval skin; e, newly emerged adult, not yt colored; f, fully colored and perfect adult.
All lrgedto the same sle. (From Marlatt.)
or two hundred specimens of the beetle were shipped by Mr. Marlatt
to Washinon live, but all but two perished during the winter. One
at least of the two survivors was an impregnated female, and began
laying eggs early in April. From this individual at least 200 eggs
were obtained, the work being done in breeding jars. After some hun-
dred larv had been hatched from these eggs the beetles were placed
on a large plum tree in the experimental orchard and protected by a
wire-screen cage covering the tree. The stock increased very rapidly,
and during August shipmen to various eastern experiment stations
were beun about ,000 secimens bein sent out. At the end of the
38 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
first summer'there remained of the stock at Washington about 1,000
beetles. Among the colonies sent out the best success was obtained
in Georgia. An orchard at Marshallville in that State, containing
some 17,000 peach trees and covering about 85 acres, adjoined a
larger orchard containing about 150,000 trees, all scatteringly in-
fested with the scale. The ladybirds were liberated in August, 1902,
in the smaller orchard, and an examination made 11 months
later indicated that they were rapidly spreading and would soon
cover the orchard. The number of beetles in all stages present was
estimated at nearly 40,000. Colonies established in the Northern
States perished. In the South the almost universal adoption of the
cheap and satisfactory lime-sulphur washes destroyed the possi-
bility of rapid multiplication and destroyed the majority of the bene-
ficial insects. This species has not been found recently, but prob-
ably exists in Georgia. The introduction and establishment of the
species was successful, but it was practically killed out by the cheap
and satisfactory washes in general use. Without the washes the
probabilities are that the ladybird would be found at the present
time occurring in great numbers in southern orchards.
THE PARASITES OF DIASPIS PENTAGONA TARG.
For a number of years the mulberry plantations of Italy had
suffered severely from the attack of the insect known as the West
Indian peach scale (Diaspis pentagona Targ.). This insect occurs
in the United States and is widely distributed in other parts of the
world. In the United States, however, it is not especially injurious.
In 1905," at the request of Berlese, the writer sent parasitized Diaspis
from Washington to Florence, Italy. One of the. parasites which
issued, Prospaltella berlesei How., was artificially reared in Florence
by Berlese and his assistants, and at the time of present writing has
been so thoroughly established in several localities that the ultimate
reduction of the Diaspis to harmless numbers is confidently antic-
ipated by Berlese. Similarly, Silvestri at Portici has introduced
the same species from America, and also certain ladybirds, and is
making the effort to import the parasites of this species from its
THE WORK OF MR. GEORGE COMPERE.
Mr. George Compere, employed jointly by Western Australia and
California as a searcher for beneficial insects, for several years has been
traveling in different parts of the world in search of beneficial insects
which he has either sent or brought to California and Western Aus-
tralia. One of the most interesting of his achievements was sending
living specimens of ('alliephialtes messor Gray., an ichneumon fly, from
Spain to California. This species is a parasite of the codling moth.
PREVIOUS WORK WITH INSECT PARASITES. 39
In California this ichneumon fly has ben reared with great sucess
and has been sent out in large numbers from the headquarters of the
Stte board of horticulture. In the field, however, it is apparently not
succeeding, and there is no evidence that the numbers of the codling
moth have been at all reduced by it. Nor is it, acorin to Froggatt,
effective in Spain.
Mr. Compere has collected many beneficial species attacking many
different injurious insects. He is an indefatigble worker, and his
untiring qualities and s refusal to accept failure are well shown in
his search for the natural enemies of the fruit fly of Western Australia,
eratitis capitat Wied. He visited the Philippine Islands, China,
Japan, California, Spain, returnin to Australia, afterwards visiting
Ceylon and India, and subse uently Brazil. In Brazil he succeeded
in findin an ichneumon fly d a staphylnid beetle feeding upon
fruit-fly lav. IlHe collected some numbers and carried them to
Australia in livin condition, prematurely reporting success. The
fruit fly is a pest in South Africa, and following the announcement of
Compere's importation Claude Fuller and C. P. Lounsbury pro-
ceeded from Afrca to Brazl to get the same parasites. The result of
this journey was discuraging. They did not find the predator
staphylinid, but obtained a braconid parasite, Opiellus trimacilatus
Spin.; they also concluded from inforation gained that the fruit fly
had been introduced into South America more recently than into South
Africa. The material carried home died. Compere left Australia
again about the close of 1904; went to Spain for more coling-moth
parasites, and then went on to Brazil, collecting more fruit-fly para-
sites and carrying them to Australia. The Brazilian natural enemies,
however, did not succeed, and in 1906 he proceeded to India to collect
parasites of a related fly of the genus Dacus, finding several and tak-
ing them to Western Australia. He arrived, however, in the middle
of winter, and the insects perishd. In May, 1907, once more this
indefatigable man returned to India, and in a few months collected
70,000 to 100,000 parasitized pupa, and brought them to Perth,
Western Australia, in good condition on the 7th of December. It is
reported that the parsites iued from this material in great numbers
and in three distinct species. In April, 1908, it was reported that
120,000 parasites had been obtained and distributed, 20,000 of them
having been sent to South Africa. The writer has not seen any
definite reports of success in the control of the fruit fly by these para-
sites, but surely Compere deserves great credit for his efforts.
WORK WITH THE EGG PARA8TE OF THE ELM LEAF-BEETLE.
In 1905 Dr. Paul archal, of Paris, published in the Bulletin of the
Entomological Society of France for February 22 a paper entitled
"Biological observations on a parasite of the elm leaf-beetle," to
40 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
which he gave the name Tetrastichus xanthomelxne. In this very
interesting article Dr. Marchal called attention to the fact that the
elm leaf-beetle had multiplied for several years in a disastrous way
about Paris, skeletonizing the leaves in the parks and along the ave-
nues. In 1904 the ravages apparently stopped, and Marchal's obser-
vations indicated that this was largely due to the work of this egg
parasite. He studied the life history of the parasite carefully during
that year at Fontenay-aux-Roses and published his full account the
Visiting Dr. Marchal in June, 1905, after the publication of this inter-
esting article, the senior author asked him whether he had been able
to make the further observations promised in the article, and he re-
plied that the elm leaf-beetle had so entirely disappeared in the vicin-
ity of Paris that he had not been able to do so. The visitor urged him
to make an effort through his correspondents to secure parasitized
eggs of the beetle for sending to the United States in an effort to intro-
duce and establish this important parasite on this side of the Atlantic.
It was considered hopeless to attempt the introduction that summer,
as the time was so late and it was not then known in what part of
France the elm leaf-beetle could be found abundantly. During 1906
practically the same conditions existed. A locality was found, but
the parasites did not seem to be present. In 1907, reaching Paris
about the 1st of May, the visitor again reminded Dr. Marchal of his
desire to import the parasite into the United States, and meeting M.
Charles Debreuil, of Melun, the subject was again brought up and M.
Debreuil later in the season forwarded eggs of the beetle to the United
States, which were promptly sent to the parasite laboratory at North
Saugus, Mass., but the time was too late, and the parasites had
emerged and died.
In April, 1908, the Entomological Society of France published in its
bulletin (No. 7, p. 86) a request from the senior author that eggs of the
elm leaf-beetle should be sent to the United States for the purpose of
rearing parasites. This notice brought a speedy and effective response.
About the 20th of May Prof. Valery Mayet, of Montpellier, France, a
personal friend, secured a number of leaves of the European elm car-
rying egg masses of the beetle, placed them in a tight tin box, and
mailed them to Washington. They were received May 28, and at
once forwarded to the junior author at the parasite laboratory at
Melrose Highlands. On opening the box the junior author found a
considerable number of active adults of the parasite. Most of them
were placed in a large jar containing leaves of elm upon which were
newly deposited masses of the elm leaf-beetle eggs. Probable ovipo-
sition was noticed within an hour after the receipt of the sending.-
There were probably somewhat more than 100 adults received in the
shipment and very few emerged from the imported egg masses after
PREVIOUS WORK WITH INSECT P ITS. 41
the t day. The adults lived certainly for 35 days. Reproduction
curred in the experimental jars, and the adults secured by this
borator reproduction were liberated in two localities near Boston
and tized eggs were sent to Prof. J. B. Smith at New Brunswick,
N. Prof. V. Slinerland at Ithaca, N. Y., and others to Washin
ton. The first of theMassachusetts colonies consisted of about 600
parasites inclosed in an open tube tied to a tree in the Harvard yard,
Cambridge, Mass., on June 22. Mr. Fiske thinks that more than 100
found their freedom on the same da, and almost certainly all of the
rest within a week. A little mor tha a month later Mr. Fiske
found parasitized es one-fourth of a mile away from this colony.
At Meose Highlands more than 1,200 were liberated on the 21st of
June and the 8th of July; and on the 27th of July fresh native eggs in
the neigborhood produced parasites, indicating the development of
a generation nmerican soil. In the summer of 1909 none of the
parasites was found, but this by no means indicates that the species
has not become established. Both the eggs and the parasites are
verv small, and the writer expects that even from this first experiment
good results will follow, Arrangements had been made for a repeti-
tion of the sending in May, 1909. from Montpelier, this southern
locality allowing such an early sending as to insure the arrival of the
parasitized eggs in the United States at the proper time of the year.
Relying upon Prof. Mayet's promises and his great experience as an
entomologist, no other arrangements were made. Most unfortu-
nately, however, just before the time arrived Prof. Mayet died, and
the introduction was not made. It should be stated that in the death
of this admirable man France lost one of its most enlightened and able
economic zoologists. It is hoped to repeat the introduction, through
the kindness of Dr. Marchal in France and Prof. Silvestri in Italy.
Silvestri has proised aso to send other natural enemies of the elm
leaf-beetle from Italy.
WORK WITH PARASITES OF TICKS.
S1907 the senior author described the first species of a hymenop-
terous parasite ever recorded as having been reared from a tick.
The name given to it was odipagus teanus, and it had been
reared from the nymphs of Hwmaphysalis leporis-palustris Pack.
colected on a cotton-tail rabbit in Jackson County, Tex., by Mr.
J. D. Mitchell. In 1908 he described another, Hnterells hooki,
rear by Mr. W.A. Hooker at Dallas, Tex., from Rhipicephalus
SBanks taken from a Mexican dog at Corpus Christi, Tex.,
by H. P. Wood. Inasmuch as a closely allied if not identical
tick, Ripicepals sanins Latr., is supposed to be a transmitter
of a trypanosome disease in South Africa, sendings of the Hunterellus
wee made in the autumn of 1908 to Prof. Lounsbury at Cape of
42 PARASITES OF GIPSY AND BROWN-TAIL MIOTHS.
Good Hope and to Mr. C. W. Howard, entomologist to the govern-
ment of Lourenco Marques, Portuguese East Africa. In June, 1909,
Mr. C. W. Howard reared parasites from engorged nymphs of Rhipi-
cephalus sanguineus taken from dogs, with which transmission
experiments with trypanosomiasis were being made. Examination
showed them to be Hunterellus hookeri. Mr. C. W. Howard is of
the opinion that these 1909 reared specimens could not have been
the offspring of those sent over in the autumn of 1908, since, as he
writes under date of September 3, 1909, the latter arrived while he
was absent in the Zambesi country, and, as he was gone nearly three
months, they remained on his desk unopened. When he returned
they were all dead. He kept the ticks some time, however, in a
sealed jar to see if any more parasites might emerge, but none did so.
In his opinion there is absolutely no possibility that the 1909 speci-
mens are the descendants of those sent from Texas. Of course Mr.
C. W. Howard is probably correct in his surmise, but a most interest-
ing question arises as to the original home of the parasite. Could it
have been carried accidentally from Texas to Africa at an earlier
date ? As a matter of fact, during the Boer War thousands of horses
and mules were shipped from southern Texas to Cape Town, and
much of this stock came from the very region in which the Texas
Rhipicephalus occurs. Banks, in his revision of the ticks,1 records
this species from horses as well as from dogs, the horse record coming
from New Mexico. The suggestion regarding the importation' of
horses and mules from Texas to Cape Town during the Boer War
was made to the writer by Mr. W. D. Hunter, who also suggests that
as Rhipicephalus sanguineus occurs throughout Africa and Mediter-
ranean Europe, and that as in 1853 several shipments of camels
were brought to Texas from Tunis, being turned loose at Indianola
and roaming wild throughout the territory around Corpus Christi
for some years, it is possible that the Rhipicephalus was brought to
Texas on these camels, and the parasite as well. This seems unlikely,
however, since the parasite had never been found in Africa or Europe
until the specimens referred to were reared by Mr. C. W. Howard in
MR. FROGGATT'S JOURNEY TO VARIOUS PARTS OF THE WORLD IN 1907-8.
As a result of a conference of Australian Government entomologists,
held in Sydney, July 9, 1906, and of a conference of State premiers,
held in Brisbane, June, 1907, it iYas agreed that Mr. W. W. Froggatt,
entomologist to the Department of Agriculture of the State of New
South Wales, should be dispatched to America, Europe, and India,
to inquire into the best methods of dealing with fruit-flies and other'
pests, the expenses of the journey to be shared by Queensland, South
SU. S. DIepartment of Agriculture, Bureau of Entomology, Technical Series No. 15, p. 35, 1908.
PREVIOUS WORK WITH INSECT PARASITES. 43
Australia, New South Wales, and Victoria. As a result of the trip
following this authorization, Mr. Froggatt has published a report on
prasitic and injurious insects, issued in 1909, in which he considers,
(1) the commercial value of introduced paraites to deal with insects
that are pests; (2) the range and spread of fruit-flies, and the methods
adopted in other countries to check them; (3) the value of parasites
in exterminating frit-flies; (4) the habits of cosmopolitan insect
pests. On his journey, which began the end of June, 1907, Mr.
Froggatt visited Hawaii, the United States, Mexico, Cuba, Jamaica,
Barbdos, England, France, Spain, Italy, Austria, Hungary, Turkey,
Cyprus (spending a day in Smyrna and two days at Beirut on the
way), Egypt, India, Ceylon, and thence to Australia, stopping in
Western Australia before his return to Sydney. In the course of
this trip Mr. Froggatt not only studied the question of parasites and
of economic entomolog in general, but looked into a large number
of matters of agricltural interest, and has given a report which
can not fail to be interestirg to every one occupied with any branch
With regard to the practical handling of parasites, and especially
international work, he is inclined to be rigidly critical. Ilis motive
obviously was to look everwhere for accomplished results and
where he could not find these to distinctly state the fact. Ile depre-
cates all claims that are not or have not been justified by practical
results of value. Thus, while admitting the good work of the intro-
duced parasites of the sugar-cane leafhopper in Hawaii, he states
that the tdvocates of the partsite system do not take into account
the alteration of methods of cultivation which occurred about the
same time, namely, the burning of the refuse (probably containing
many eggs and larv) ilnstead of buying it as formerly, and the
introduction of new varieties of cane more resistant to the leaf-
hoppers. In California, he admits the value of the introduction of
the Australian ladybird, but states that his observations show that
no good has followed the introduction of the codling-moth parasite
from Spain, although it had been claimed previously that this parasite
would prove a perfect remedy for the apple pest, and pointing out
that when he visited Spai he found that a very large percentage of
the apple crop is always infested by the codling moth. He states
that the promises of the advocates of the parasite method in C(ali-
fornia have not been fulfilled; that Western Australian claims that
staphylinid beetles destroy the majority of the fruit-fly maggots in
Brazil, and that nature's forces in that country control the destruc-
tive fruit flies are to be contrasted with the statement of the South
African entomologists that only a few months after the visit of the
West Australian entomologist to Brazil they found that "all along
the Brazilian coast it was difficult to obtain a fruit that had not been
44 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
punctured by a fly." The statement that nature controls the
destructive fruit flies in India he opposes, as a result of his own
observations in India. He does not contend that this work has not
a great practical value, but insists-that it should be done by trained
entomologists, and that full information of the habits and life his-
tories of both the pests and their parasites should be understood
before liberation is attempted. As already stated, he especially
deprecates premature claims, and points out that in New South
Wales the passage of the very necessary vegetation diseases bill was
delayed for some years by the outcry "Why should we be made to
clean up our orchards and spend money, when the department can
send out to other countries and get us parasites that will do all that
is needed?" In conclusion he states:
Let the whole question be judged on its results. Allow that one or two experiments
have shown perfect results; yet because mealy bugs or scale insects in a restricted
locality have once or twice been destroyed by parasites, that can be no reason why
the parasite cure alone should be forced upon anyone. Its admirers should be
perfectly honest; and if a friendly introduced insect from which, rightly or wrongly,
great things had been expected turns out upon further trial to be a failure, they should
say so; and they should never proclaim results for a parasite till those results have
actually been proved in its adopted country, for the wisest can never be sure of the
results of any experiment. Economic entomology is a great commercial science,
and those at work for its far-reaching interests could do it no greater harm than by
misleading or unproved statements.
OTHER WORK OF THIS KIND.
Reference has already been made to the importations of Prospal-
tella berlesei into Italy to attack the destructive mulberry scale,
Diaspis pentagona, through cooperative arrangements between the
senior author and Prof. Berlese, of Florence. Prof. Berlese has been
successful in establishing the species, and believes that it is best to rely
upon this species only, and not to attempt to introduce the predatory
enemies of the scale, his idea being that coccinellids will feed indis-
criminately upon parasitized and unparasitized scales and that thus
the Prospaltella will not have a chance to multiply to its limit. The
contrary view is taken by Prof. Silvestri, at Portici, in the south of
Italy, and he has been making every effort to introduce from all parts
of the world all of the enemies, whether parasitic or predatory, of the
mulberry scale. He has brought over and has had breeding in his
laboratory at Portici, as well as in an experimental olive orchard
southeast of Naples, a number of species of Coccinellidaz brought from
different parts of the world. At his request, in May, 1910, the senior
author carried from Washington a box containing possibly 200 living
specimens of Microweisia misella Lec. and a few specimens of Ch(lo-W
corus bivulnerus Muls. These were carefully packed with plenty of
food in a small paper-covered wooden box, approximating a 10-inch
cube. He sailed from New York direct to Naples and, through the
PREVIOUS WORK WITH INSECT PARASITES. 45
ki eof the officers of the Royal Italian Line steamship Duca di
G was enabled to suspend the box by a cord from a crossbeam in
the ordiary ld room of the steamer. After an eleven days' passage,
the box was opened in Prof. Silvestri's labratory in Portici, and prac-
tically very coccinellid was found to be alive and in apparently good
Efforts have been made by the Bureau of Entomology, in coopera-
tion with the Pasteur Institute in Paris, to introduce a large bembecid
wasp (Monedua carolina Fab.) from New Orleans into Algeria to prey
upon the tabanid flies concerned in the carriage of a trypanosome
disease of drom aries. The wasps were sent in -their cocoons in re-
frigerating baskets from New Orleans by direct steamer to Havre and
from New York by direct steamer to Havre. There they were met by
agents of the Pasteur Institute, arried to Marseilles by rail and thence
byboat to Algeria, and were planted under conditions as closely as
possible resembling those under which they were found in Louisiana,
care being taken to simulate not only the character of the soil but the
exposure to light, the prevailing wind irections, and the moisture
condition. Adults issued, but the species has not since been re-
covered, although it is quite possibly established.
In the same way an attempt was made to introduce the common
bumblebee Bombs pe nnslvanicus De Geer of the United States into
the Philippine Islands for the purpose of fertilizing red clover. These
were sent in refrigerating baskets, carried by hand by Filipino stu-
dents returning from the United States to the Philippines, and for the
mst part in the pupal stage. These were properly planted upon
arrival and reared, and a few specimens have been recovered.
In the summer of 1910 Dr. L. P. De Bussy, biologist of the Tobacco
Planters' Association of Deli, Sumatra, visited the United States for
the purpose of investigating damage to the tobacco crop by inects
and disease and to make an effort to import into Sumatra the parasites
of the destructive tobacco wo known as Heliothi obsoleta Fab.
Aleady shipments of an egg parasite, Tricogra a petsa Riley,
have been made to Sumatra a sterdam, but information as to
the results of these reliminary shipments has not yet reached this
Prof. C. H. T. Townsend, an assistant in the Bureau of Entomology,
receiving a temporary appointment as entomologist to the Depart-
ment of Agriculture of Peru, pecially to study the injurious work
done by the scale insect icioapis mior Mask. on cotton, has
during the past year, with the assistance of the bureau, imported a
number of shipments of Pro ltella berleei from Washington into
Peru. It is too early to announce results.
In July, 1910, Mr. R. S. Woglum, agent of the Bureau of Ento-
Swas sent abrad to find the oriinal home of the white fly of
46 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
the orange, Aleyrodes citri R. & H., and to attempt to find parasites
or satisfactory predatory enemies. In November, 1910, he found the
white fly at Saharampur, India, and discovered that it was killed by a
fungous disease (lately determined as a species already occurring in the
United States- Egerita webberi-by Prof. H. S. Fawcett, of the Florida
Agricultural Experiment Station). He also found that it was at-
tacked by two species of Coccinellidm (Verania cardoni Weise and
Cryptognatha flavescens Motsch.). A preliminary shipment of the
ladybirds by mail was apparently unsuccessful. Later shipments by
direct steamer from Calcutta to Boston were also unsuccessful.
At Lahore, India, Mr. Woglum found his first evidence of parasit-
ism by hymenopterous parasites. A certain proportion of Aleyrodes
citri was found to contain the exit holes of a true parasite. The
specimens on leaves sent in by Mr. Woglum were examined with
great care. None of the full-grown larvae or nymphs contained para-
sites, but five specimens of a very minute aphelinine of the genus
Prospaltella were found dead and attached to the orange leaves in
close vicinity to the perforated aleyrodids. The size of the specimens
was such as to justify the conclusion that they had issued from the
aleyrodids, and their juxtaposition and the known habits of the genus
confirm this conclusion. The species was described by the senior
author as Prospaltella lahorensis in the Journal of Economic Ento-
mology for February, 1911, pages 130-132. Efforts will be made to
import this parasite into Florida.
The occurrence of a European weevil, Phiytonomus murinus Fab.,
in the alfalfa fields of Utah in alarming numbers and the difficulty of
fighting the pest by mechanical or cultural means has started an in-
vestigation as to its parasites in its original home. Mr. W. F. Fiske,
of the Bureau of Entomology, sent from Naples, Italy, on March 17,
1911, a large lot of stems of alfalfa containing eggs of an allied weevil
parasitized by a minute mymarid, which at the time of this writing
are on their way to Utah.
In the meantime the State board of horticulture of California has
been continuing its efforts to import beneficial insects of different
kinds. Mr. George Compere returned from a lengthy trip during the
summer of 1910, bringingg with him a number of interesting species,
among them a new coccinellid enemy of mealy bugs in which he has
great faith and which promises to be a valuable addition to the insect
fauna of the United States.
Entomologists and horticulturists all over the world have become
greatly interested in this aspect of economic entomology and for the
ilnlle(liate future a great deal of experimental work has been planned
by the officials o(f different countries.
EARLY IDEAS ON INTRODUCTION 47
EARLY IDEAS ON INTRODUCING THE NATURAL ENEMIES OF THE
Promptly with the discove that the gipsy moth had become
acclimatized in Massachusetts, in 1889 there was published by Prof.
C. H. Fernald a special buletin of the Massachusetts Agricultural
College Hatch Experiment Station, in. which he gave popular descrip-
tions of the different stages of the inect and recommended spraying
with Paris green. He stated that the insect is generally held in
check by its natural enemies in Europe, but occasionally becomes
very destructive, and stated that 11 species of hymenopterous para-
sites and several of dipterous paraites had been noticed in Europe.
This bulletin was published in November, 1889. In Janua-y, 1890,
an illustrated article on the i psy moth, by Riley n Howard, wa
published in Isect Life,' and a list of 24 Europeanl hymenopterous
parasites compiled by Howard wa published.
Immediately followin this lpublication, there was received at the
Department of Agriculture, from Rev. II. Loomis, of Yokohama,
Japan, a letter in which he stated that he had seen reports of the
ravages of the gipsy moth in Massachusetts and had taken consid-
erable interest in the matter. He al stated tha ehad seen the
gipsy moth caterpillar on a wistaria vine near his house in Yokohama,
and that it had ben attacked and killed by a parasite. Several of
the parasites were sent in an accompanying box, and proved to be
Apanteles. Subsequent attempts were made by Mr. Loomis to send
this parasite in living condition both to the Department of Agricul-
ture ad to the State of Massachusetts, but all arrived dead, for the
most part having been killed by secondary parasites.
In March, 1891, a conference was held in the rooms of the com-
mittee on agriculture at Boston, at which were present Prof. N. S.
Shaler, Gen. F. H. Appleton, and Mr. William R. Sessions, of the
State board of agriculture; Prof. C. V. Riley, entomologist of the
United States Department of Agriculture; Prof. C. 11. Ferald, ento-
moloQist of the State Experiment Station; Mr. S. H. Scudder, a well-
known entomologist; the mayors of Medford, Melrose, Arlinton, and
Malden, and others In the course of the conference, which was held
for the purpose of discussing the best measures to be taken against
the gipsy moth by the State, Prof. Riley advocated an attempt at
extermination b spraying. M. Scudder advocated the destruction
of the eg, and in the course of the discussion Prof. Riley made the
I would make one other on, and that is, that as an auxiliary method it would
be well to spend $500 or00 in sending one or two persons abroa next summer with
no other object than to go to some section of northern Europe to collect and transmit
to authorized persons here a certain number of the primary parasites of this species,
I Insect Life Division of Entomology, U. S. Deartment of Agriculture, vol. 2, pp. 208-211, 1890.
48 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
which are known'to check its ravages over there. The insect was undoubtedly brought
over by Trouvelot without any of its natural checks. In my judgment it would be
well worth trying to import its parasites from abroad. The advantage would be this:
If you failed to exterminate it by spraying, its parasites, seeking for this particular
host, would be more apt to find the overlooked or escaped specimens than man would.
No action was taken upon this suggestion, and the State author-
ities, believing that such an attempt would be useless owing to the
fact that their effort for some years was consistently devoted to the
aim of absolute extermination of the gipsy moth, perhaps wisely
saved the expense of a mission abroad for this purpose. Then, also,
there was some hope that the native parasites, particularly the ich-
neumon flies and the native species of Apanteles, as well as tachina
flies and some of the carabid beetles, might gradually accommodate
themselves to the imported pest and prove prominent factors in the
fight against it.
This last faint hope, however, was not justified. In the course of
the careful work done by the State during the next seven or eight
years, the better part of which is summarized in the admirable Report
on the Gipsy Moth, by Forbush and Fernald, published in 1896, sev-
eral native parasites and predatory insects were observed to attack
the gipsy moth in its different stages, but at no time was the per-
centage of parasitism sufficiently great to have any value as a factor
in the suppression of the pest. At no time was there a greater per-
centage of parasitism by native parasites than 10, whereas the con-
dition in Europe is such that the percentage reaches frequently well
above 80. It may be worth mentioning that parasitism by native
species has never exceeded 5 per cent in any collections made since the
present laboratory was established. It is nearer 2 per cent on the
In discussions among the Washington entomologists it was repeat-
edly pointed out by E. A. Schwarz and by B. E. Fernow (at that time
Chief of the Division of Forestry of the United States Department of
Agriculture) that one of the most important of European enemies of
the gipsy moth, and the nun moth as well, is one of the tree-climbing
ground beetles known as Calosoma sycophanta L. There exist a num-
ber of species of this same genus Calosoma in the United States, but
none of them has the tree-climbing habit developed to the same ex-
tent as have Calosoma sycophanta of Europe and its relative Calosoma
inquisitor L. Prof. Fernald, writing to the famous German authority
on forest insects, Dr. Bernard Altum, early in 1895, asked his opinion
as to the advisability of importing these tree-inhabiting ground beetles,
but received the reply that such an importation would not give good
results. Prof. Altum considered the services of the hymenopterous.
parasites of the old genus Microgaster as of much more importance.
In the report just cited Fernald disposed of the question of import-
ing parasites in the following words:
CIRCUMSTANCES OCCASIONING WORK. 49
o tempt has been ade to import parasites thus far for the reason that the law
requires the work to be conducted with direct reference to the extermination of the
ipy moth, and, therefore, the general destruction of the insect would also destroy
the paites. There is no reasn why our native hymenopterous parasites may not
prove to be quite as effective as those of any other country, since there is no parasite
known which confines itself exclusively to the gipsy moth, and, as has been shown,
we have several species which attack it as readily any in it native country.
This position with regard to the nonimportation so long as exter-
mination of the gipsy moth was the end, held until the State of
Massachusetts ceased its appropriations, in the year 1900.
CIRCUMSTANCES WHICH BROUGHT ABOUT THE ACTUAL BEGIN-
NING OF THE WORK.
During the five year that elapsed before the State again began to
appropriate money for the suppression of the gipsy moth and the
brown-tail moth, as iswell knon, the gipsy moth spread from a
restricted territory of 359 square miles throughout an extended range
of 2,224 square miles and even more. As son as the effort to exter-
minate it was abandoned, owing to the lapse of the appropriations
for the year 1900, the project of importing parasites was taken up
by the Chief of the Bureau of Entomology, who began correspond-
ence with a number of European entomologists with this end in view.
Especial efforts were made to import the Calosomas, but failed,
partly owing to a lack of interest in the matter on the part of the
Europeans. In 1902 Mr. B. Alwood, entomologist of the Vir-
ginia Agricultural Experiment Station, went abroad for a series of
months and was requested by the chief of the entomological service
of the United States Department of Agriculture to endeavor to find,
in some well-placed situation in Europe, one or more competent col-
lectors of insects who would undertake systematically to send gipsy-
moth parasites to America. This effort also failed, and Mr. Alwood
was unable to find the proper persons. Finally, in December, 1904,
Congress was asked to make a small appopriation for the distinct
purpose of attempting the importation of these parasites, and the
sum of $2,500 was appr iated for this purpose in the session of the
winter of 1904-5. During te corresponding sessin of the Massa-
chusetts State Legislature, State appropriations began once more. In
1904 it was apparent to everyone that the old areas had become
reinfested and that the insect had spread widely. Private estates
and woodlands in June and July of that year were almost completely
defoliated. Kirkland wrote:
From Belmont to Saugus and Lynn a continuous chain of woodland colonies pre-
sented a sight at once disgusting and pitiful. The hungry caterpillars of both species
of moths swarmed everywhere; they dropped on persons, carriages, cars, and auto-
mobiles, and were thus widely scattered. They invaded houses, swarmed into living
95677 -Bull. 91-11---4
50 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
and sleeping rooms, and even made homes uninhabitable * *. Real estate in
the worst-infested districts underwent a notable depreciation in value. Worst of all,
pines and other conifers-altogether too scarce in eastern Massachusetts-were killed
outright by the gipsy-moth caterpillars, while shade trees and orchards were swept
bare of foliage.
There was a general demand upon the State legislature and an
excellent bill was prepared and passed with the appropriation of
$300,000, $75,000 to be expended during 1905, $150,000 and any
unexpended balance during 1906, and $75,000 and any unexpended
balance during 1907, up to May 1, 1907, inclusive. And to this
appropriation there was added the clause "for the purpose of experi-
menting with natural enemies for destroying the moths, $10,000
is additionally appropriated for each of the years 1905, 1906, and
1907." There was then available in the spring of 1905 the appro-
priation of $2,500 by the General Government and that of $10,000
by the State of Massachusetts for work with the natural enemies.
Mr. A. H. Kirkland was appointed superintendent for suppressing
the gipsy and brown-tail moths, by Gov. Douglas, and immediately
following his appointment, and with the approval of his excellency
the governor, went to Washington, and by arrangement with the
honorable the Secretary of Agriculture, Mr. James Wilson, arranged a
cooperation between the State and the Department of Agriculture
whereby the Chief of the Bureau of Entomology of the department
was practically placed in charge of the details of the attempt to import
parasites from abroad, in consultation with Mr. Kirkland.
The reasons which influenced Mr. Kirkland in entering into this
cooperation between the State and the United States Department of
Agriculture were expressed in his first annual report (p. 117).
At this time for more than 25 years the chief of the bureau had
been devoting his especial efforts to the study of the parasitic Hymen-
optera, and had especially interested himself in the subject of their
biology and host relations. He had accumulated a card catalogue
of more than 20,000 entries of records of the specific relations of
parasites to specific insects, the great majority of these being Euro-
pean records and covering all of the published information regarding
the parasites of the gipsy moth and the brown-tail moth. He also
had the advantage of the personal acquaintance of most of the
European entomologists interested in this kind of work. These
facts were known to Mr. Kirkland and caused his action.
AN INVESTIGATION OF THE INTRODUCTION WORK.
From the beginning of the work, and even before, certain citizens
of Boston, irnliressed1 by the claims of the State Board of Horticulture
of California as to the results said to have been achieved by the
agents of the board in the introduction of beneficial insects, urged
INVESTIGATION OF INTRODUCTION WORK. 51
the employment of thse agents in the work of introducing the para-
sits of the gipsy moth and brown-tail moth. The arguments in
favor of this proposal were duly considered by the superintendent
of the Massachusetts ork, who decided for many reaso to con-
duct the introduction experiments along the lines just described
and not to call in the assistance of the California people. In his
third report, submitted January 1, 1908, _Mr. Kirkland expressed
the situation as follows:
In spite of all the thoht, energy, and skill that have been brought to bear on this
mst important problem of introdcing the natural enemi of the moths-a problem
entirely novel in the field of entomol oy-it was apparent during the winter of 1906-7
that sveral of our influential citizens had expecte immediate results from the
importation of the psites, and were beginning to get restive becsrae such results
had not been obtained. Several expred a doubt if everything poible was being
dne to secure the succesful introduction of the parats. Other became enthusi-
atic 4over the specious prositin put forward by a certain western horticulturist
(not an entomol ogst), who offe to su the gipsy moth in Measchusetts by
means of parasits for the sum of $, ",0(K) no cure, no pay." This state of affairs
ws no doubt a natural outcome of te desire to avoid a repetition of the great damage
to property caused by the moth in past years. Aain, men without any technical
knowl ige f entomology or of the life histories of the arasites not realizing the difli-
culties in securing, -hipping. breeding, and diseminag these beneticial insects.
and equally igno ant of how long it takes n imported inset to become established
ven under the most favorable conditions, might well be pardoned for expecting
alst immediate relts from the itrocto o te relatively shmall number of
parasitesmall indeed in comparin with the tremendous numbers of the moths
Coming before the legislature during the session of 1906-7, this
group of Boston citizens stated that it was their opinion that the
work pa es not p with parsites as no(t p swit h1 sufiicient rapi(dity, and
asked the legislatre to appropriate funds and to instruct the super-
intedent to secure dditional counsel and advice in the matter to
determine whether the work was going on in the right way. The
legislature agreed and appropriated the a(ddIitional sun of $15,000 to
enable the superintendent to secure such ad-vice.
It was first sugested that he consult only with certain California
men who had had experience in importing parasites of scale insects.
He, however, considered that consultation with men whose experi-
ence had been confined to a single group of insects, not to the same
group as the gipsy moth and the bron-tail moth, while possibly
helpful, would not be broad enough to throw any great light on
the Massachusetts problem. To use his own words-
It seemed much wiser and certainly more thoroughgoing, since this entire work
iht be called in question at any time, and in view of the lage aount of money
husett was expding in securing parasite, to consult not with the trained
entomologists of a single State, but with as many entomologists of national or even
world-wide reputation as possible. In other words, that a large number of entomolo-
gists of the hihest possible scientific standing, and particularly those having practical
riee in dealin with arasitic insects, ould be invited to visit Massachusetts,
52 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
learn of our difficult problems on the spot, examine into the methods of importing,
rearing, and distributing parasites, and then give us the benefit of their criticism and
counsel, based on a full knowledge of the facts at hand. He also suggested that,
since by some this movement might be taken as a criticism on his management and
on his judgment in placing the direction of the work in the hands of Dr. Howard, it
would be well to have some outside board or commission take charge of the matter,
so that it should be entirely an ex parte affair, free from any suggestion of influence
by the present administration of the work. The suggestion to authorize the super-
intendent to invite the entomologists was heartily indorsed by the legislative com-
mittee which had the matter under consideration, while the arrangement of the entire
affair was left in his hands.
In his selection of experts, Mr. Kirkland was aided by Prof. C. H.
Fernald, of the Massachusetts Agricultural Experiment Station,
one of the oldest and best posted entomologists in the country; and
Mr. Kirkland himself, it must be remembered, had been engaged in
active entomological work for 15 years and had held official posi-
tions in the Association of Economic Entomologists, thus having a
very broad personal acquaintance with the best workers. The list
selected, as quoted from Mr. Kirkland's report, was as follows:
Prof. Edward M. Ehrhorn, deputy commissioner of horticulture, State of California,
a man of large practical experience in importing, breeding, and disseminating insect
parasites, particularly those of scale insects, and also a man well trained in applied
Prof. Herbert Osborn, Ohio State University, one of the country's best known
teachers of entomology, and of large experience in investigation and laboratory work.
Dr. John B. Smith, entomologist, New Jersey Agricultural Experiment Station,
an investigator of the highest order, a successful teacher, and the author of numerous
standard works on insects.
Prof. S. A. Forbes, State entomologist, Illinois, a most successful teacher and investi-
gator, and one of the most prominent entomologists of the Middle West.
Prof. E. P. Felt, State entomologist of New York, a well-known writer on and investi-
gator of insect pests, and particularly ingenious in devising laboratory methods.
Prof. H. A. Morgan, director of the Tennessee Agricultural Experiment Station,
of large experience, and one of the best-known entomologists of the Southern States.
Prof. M. V. Slingerland, Cornell University, New York, an investigator with hardly
an equal, and one who has had great success in studying life histories of beneficial
and injurious insects.
In addition to these, the following well-known foreign entomolo-
gists, visiting Boston, were asked to investigate the situation care-
fully, to study the laboratory and field methods, and to report:
Prof. ('harles P. Lounsbury, entomologist, Cape Town, South Africa, one who has
had great. experience as well as great success in importing beneficial insects.
Prof. W\alter W. Froggatt, government entomologist, New South Wales, and also
investigator for Victoria and Queensland. Prof. Froggatt's work has been practi-
cally along the same line as that of Prof. Lounsbury, and has met with a large measure
of suc(c'ss.o .
Dr. James FIlether, lominion entomologist, Canada, well known for his success in
working out difficult points in the life histories of insects, and more particularly in
dealing with a wide rang(e of injurious species.
Prof. R. Blanchard, Universit.y of Paris, and member of the Academy of Medicine.
Dr G. orvath, director of zoological section, National Hungarian Museum, mem-
INVESTIGATION OF INTRODUCTION WORK. 53
ber of the Academy of Science of unar and formerly director of the entomological
station of Hunar. The last two gentlemen are entirely familiar th the two moths
an their parasites.
Dr. Richard Heymons, extraordinary honorary professor and custodian at the Zoo-
logical Museum of the Royal Institute of Berlin. Dr. Heymons has made a large
study of the injurious insects of central Europe, and particularly of their natural
Prof. A. Severin, conservator at the Royal Museum of Natural History of Belgium,
and member of the Superior Council of Forests. Prof. Severin's position is naturally
that of one of the best psted entomolgists, particularly with reference to dangerous
In ad(litio to these foreign entoologists, Prof. Filippo Silvestri,
of the Royal Agricultural School of Portici, Italy, visiting America
on an official mission in the summer of 1908, visited Boston, and
was asked to give his professional opinion of the work, his report
being printed in the fourt annual report of the superintendent,
ssued( Januar, 1909, by L. Worthley, acting superitendent.
It is worthy of note that Prof. Silvestri had been comrnssioned by
the R. Accademia dei Lincei and by the royal minister of agriculture
of Italy to investigate the work in economic entomologv )being done
in the Unite States, and had visited all portions of the country,
including California and Hawaii, studying with especial care all the
work being done with parasites. It should be pointed out also that
the California claim were prfectly well understood by all of the
American experts, Mr. Ehrhorn himself being the second ranking
olicer in the Caliornia service, and the others having either visited
California partly for the purpose of investiating this work, or being
perfectly familiar with the situation bystudy of the publications and
by correspondence. Moreover, of the forein experts, Mr. Froggatt
had just come from California on an investigating trip for the gov-
ernment of the Federated Colonies of Australia which subsequently
carried him around the world, Mr. Lounsbury had visited California
for the purpose of studying this work, and Dr. Fletcher had repeat-
edly visited that State.
The reports of all of these experts, with the exception of that of
Prof. Silvestri, are published in the third annual report of the Massa-
chusetts superintendent, Boston, 1908, Prof. Silvestri's report being
published, as above stated, in the fourth annual report of the super-
It will be entirely unnecessary to quote from these reports, since
they may be found in full in the State documents mentioned. It
will suffice to state that the work was commended, it is safe to say,
with enthusiasm by every individual. Specific consideration was
given to the California suggestion by Mr. Lounsbury, by Ar. Frog-
gatt, and by Prof. Slingerland. Suggestions were made by several
of them that the study of the fungous, bacterial, and protozoan
dis s of the larv~e should be taken up. Dr. Felt and Dr. Smith
54 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
recommended' the importance of the introduction of the Japanese
parasites, and Dr. Felt suggested the importance of careful biolog-
ical studies of the parasites, not only in America but in Europe. All
of these suggestions coincided with plans already made which were
about to be entered upon, as indicated in following pages.
The subject of the study of the diseases of the caterpillars does not
come under the range of the present bulletin, but since it has been
mentioned, it should be stated that the State superintendent has for
the past two years been having this subject investigated and that it
is now going on under the expert supervision of Dr. Roland Thaxter,
of Harvard University, and Dr. Theobald Smith, of the Harvard
Mr. Kirkland's summary seems fully justified. It is as follows:
It will be seen from the foregoing that the work of importing parasites of the gipsy
and brown-tail moths in Massachusetts has been thoroughly examined by practically
a congress of the world's leading entomological experts. And it is believed that their
consensus of opinion, which is, in the main, that everything possible to secure the
successful importation of these insects is being done, will be taken as authoritative
and final. It would seem that the last word has been said on this matter, and that
there should be no further occasion for that kind of adverse criticism, whose sole
effect is to harass those who are giving their best thought and most sincere effort to the
accomplishment of the desired result. Destructive criticism of scientific work, by
the amateur or dilettante, is absolutely valueless. Constructive criticism, such as
these reports make on certain minor details of this important work, is helpful and a
NARRATIVE OF THE PROGRESS OF THE WORK.
Down to the time when this work was begun, all attempts at the
international handling of beneficial insects had been done either by
correspondence or by the sending of an individual collector to search
for such insects and to forward them by mail or express or to bring
them back himself in comparatively small numbers, the beneficial
species being either at once liberated in the field or reared for a time
in confinement and then liberated. In planning the present work
the normal geographic ranges of both the gipsy moth and the brown-
tail moth were well known and most of their parasites had been listed,
so that the problem seemed to be a comparatively simple one. Owing-
to the fact that the most abundant of the Japanese gipsy moths (four
of them are listed) liresents rather marked differences from the Euro-
Wpea aLnd New England form--so much so, in fact, as almost to justify
the opinion tlat it is a distinct species-and as the ancestors of the
New England gipsy moth came from Europe, it was decided to con-
centrate the effort, for a time at least and in the main, upon Euro-
pean parasites and natural enemies. From the outset the idea was'
to secure as many parasites belonging to as many different species as
possible from all parts of Europe, in the hope of establishing in New
England a)pproximately the natural environment of the gipsy moth
NARRAIVE OF PROGRESS OF WORK. 55
and the brown-tail moth in so far as their natural checks are con-
cerned. It was the aim to establish, not one or half a dozen of its
natural enemies, but all of them, aiming at the same time to avoid
the introduction of hyperparasites-that is, those species that prey
upon the true parasites of the injurious forms-thus, if possible, bring-
ing about an even more favorable situation for the primary parasites
in New England than exists in Europe.
On account of the enormous numbers in which both gipsy and
brown-tail moths existed in Massachusetts, it was considered that
the simplest way to secure the true European parasites was to
collect caterpillars and chrysalids wherever they could be found
in Europe, box them, and ship them directly to Boston; this always
with the certainty that a certain percentage, high or low, would
contain living parasites which would probably issue in the adult
condition on the journey or after arrival in America, in which event
they could be cared for, reared until sufficiently multiplied, and then
A temporary laboratory for the receipt and care of specimens
was immediately established by Mr. Kirkland at Malden, Mass.,
and a careful search ws begun for a suitable location for a perma-
nent laboratory for the care of parasites. It was considered desirable
that this laboratory should be placed in a region in which both the
gipsy moth and the brown-tail moth occurred in abundance, so
that there might be plenty of material for food for the parasites
at all ties; and it was also considered of importance that a con-
siderable area of land should be secured which could be controlled
for outdoor experiments.. r. Kirklad finally found a small farm
with buildings in North Saugus, the location easily accessible by
electric cars and sufficiently isolated. (See P1. II, fig. 1.) The
house was large enough to give ample room for laboratory use,
and at the same tie furnished dwelling rooms for the state official
in charge. In the iniediate vicinity there was a chain of large
woodland colonies of the gipsy moth and numerous orchards infested
by the brown-tail oth, as well as a large area of scrub-oak land
where the brown-tail moth occurred very abundantly. A portion
of the building occupied as a laboratory was fitted up by the State
with shelves, tables, rearing cages, and all necessary apparatus
and upplies, and the State employed Mr. F. H. Mosher, with Mr.
E. A. Back and Mr. O. L. Clark as assistants, to help care for the
While, as just stated and for the reasons given, the main effort
was made with Europe, correspondencewas begun with the Imperial
Agricultural Experiment Station at Nishigahara, Tokyo, Japan,
and the Imperial Agricultural College at Sapporo, in order to secure;
if possible, the services of exert Jaanese entomologists in sending
56 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
Japanese parasites, and Prof. S. I. Kuwana immediately prepared
an important sending, which, however, was not productive, through
accidents in transportation. The method tried by Prof. Kuwana
was interesting. A small tree carrying a number of infested gipsy-
moth caterpillars was packed in a large wooden case with wire-
gauze sides; another case of small elms was shipped with the insects,
and they were thus supplied with fresh food from time to time as
far as Hawaii. The case, however, shrunk in transit, making open-
ings through which the parasites for the most part escaped.
In May, 1905, the Chief of the Bureau of Entomology visited
Boston for conference with Mr. Kirkland, and on June 3 sailed
from Boston to Naples. Landing in Naples on June 13, he at once
proceeded to the Royal Agricultural School at Portici, some miles
away, and held a conference with Prof. F. Silvestri, the entomologist
of the college, and his principal assistant, Dr. G. Leonardi. By
good fortune, Prof. Silvestri was able to point out a locality in
Sardinia where, during 1904, there had been a severe outbreak of
the gipsy moth and where, therefore, during 1905 parasites could
with almost absolute certainty be predicted to occur in numbers.
With true scientific enthusiasm, both Prof. Silvestri and Dr. Leonardi
volunteered their assistance, and Dr. Leonardi was at once com-
missioned by his chief to proceed to Sardinia and to collect such
caterpillars as he could find and forward them in tight wooden
boxes, with a supply of food, to Boston. His expedition was a
success, and there were received from him at Boston, on the 15th
of July, 7 boxes, on the 26th of July 24 boxes, and on the 1st of
August 7 boxes, all containing valuable material, the most important
being a large series of living puparia of certain parasitic tachina
This extremely cordial and profitable reception at Portici by
Prof. Silvestri and Dr. Leonardi, both personally known to the
chief of the bureau from former visits, was but a foretaste of the
encouragement which was to be met at all points, and it may very
properly be said in advance that throughout the whole of the work
many European and Japanese entomologists, both officials and
private individuals, have shown an extreme liberality in their offers
of assistance in this great piece of experimental work, and the State
of Massachusetts and the United States Government are under
great obligations to them for their help and encouragement. For
the work done by Dr. Leonardi, just described, and for similar
work done in ensuing years, with Prof. Silvestri's permission, no
compensation would be accepted, and the State of Massachusetts
has paid simply for the expenses, such as packing, postage, small
traveling expenses, and items of that general character.
Bul. 91, Bureau of Entormoloy, U. S. Dept, of Agriculture. PLATE II.
FIG. 1.-VIEW OF PARASITE LABORATORY AT NORTH SAUGUS, MASS. (ORIGINAL.)
FIG. 2.-VIEW OF PARASITE LABORATORY AT MELROSE HIGHLANDS, MASS. (ORIGINAL.)
NARRATIVE OF PROGRESS OF WORK. 57
Ater Portici, Florence was visited, where a conference was held
with ProfA. Berlese, of the Royal Station of A cultural Ento-
mology, nd his assistants, Drs. Del Guercio and Ribaga. It seemed
that no occurrences of either the gipsy moth or the brown-tal moth
were known that season in Tuscany or adjoining portions of Italy.
Prof. Berlese spoke of the destruction of an outbreak of the gipsy
moth in southeirn Italy some years lreviously by a disease which
he considered to be identical with the pebrine of the domestic silk-
worm. He promised to keep up a watch for occurrences of the
pests and wherever possible to assist in the introduction of larasites.
A few days ere then spent in Lombardy, earching for tlhe larv
of either of the injurious species, but without success. Then, pro-
ceeding to Vienna, thle celebrated Natural Iistory Museum was
visited and the well-known curator of Lepidoptera, Dr. hans
Rebel, was interviewed. Dr. Rebel stated that both the gipsy
oth and the broAw-tail moth were to be found rather commonly
in parts of Austria, and it was decided to employ a professional
collector to assist in the work of shipping larvw to Boston. Upon
Dr. Rebel's recommendnation, Mr. Fritz W agner was employed.
Mr. Wagner was and is a resident of Vienna, is well versed in the
subject of European butterflies and moths, and p)erfectly familiar
with all the best collecting placess for many miles about Vienna.
Mr. Wagner accompanied the writer on several expeditions. The
frt trip was taken to tthe suburbs of Vienna, and there the first
European specimen of the ipsy-moth larva was found. It was
esting on the trunk of a locust tree by the side of. the street, and
further examiationl showed that there were a hundred or more
caterpillars on the trunk an limbs of the same trre, er was
some evidence of parasitism, and the white cocoons of a microgaster
parasite (Apahteles fulaipes Hlal.) were fountl here atnd there in
the crevices of the bark. This particular tree and nother one,
to be mentioned later, indicate very well the condition of the gipsy
moth in Europe. A hundred nearly full-grown larva were present,
but there was hardly ay evidlence of defliation. A trained ento-
mologist walking by the tree would not have noticed that insects
had been feeding upon it to any serious extent. On the other
hand, a similar tree in any of the small towns about Boston would
have carried not 100 larva, but probably some thousands, and at
that time of the year would hardly have had a whole leaf. These
specimens were collected and sent to Boston.
Later a trip was taken into the country to the battlefield of Wa-
gram, and here on two roadside poplars was found another colony of
the caterpillars raning in size from the second stage to full-grown
arv There was here more extensive evidence of parasitism by
58 PARASITES OF GIPSY AND BROWN-TAIL MIOTHS.
microgaster parasites. Their white cocoons were found abundantly,
and here again, although there must have been 250 or more larva on
the trees, the evidences of defoliation were very slight-so much so
that at a rather short distance the trees appeared in full leaf. Dur-
ing the remainder of June and July Mr. Wagner continued the search
and sent considerable material to Mr. Kirkland, at Boston.
After Vienna, the city of Budapest was visited. At the Natural
History Museum in that city Dr. G. Horvath, the well-known director,
and Prof. Alexander Mocsary were consulted, Prof. Mocsary being one
of the first authorities in Europe on the subject of parasitic Hymen-
optera. Neither of these gentlemen, however, was able to give any
new points in connection with the parasites of the gipsy moth and
the brown-tail moth. The agricultural experiment station in the sub-
urbs of Pesth was then visited, and Prof. Josef Jablonowski, the
entomologist of the station, was consulted.* By this time it was
the 4th of July, and already the season in Hungary was far advanced,
being about two weeks or more earlier there than at Vienna. Prof.
Jablonowski stated that gipsy moths had been found in certain
localities in Transylvania, but that the adults were already issuing
and that the brown-tail moths had been flying for some time. He
exhibited, however, a large box full of the previous winter's nests
of brown-tail larvae, and stated that in the early spring he had
reared from these nests many hundreds of parasitic insects. This at
once seemed to indicate a very easy way of importing such parasites,
since these nests could be readily collected in the winter in large
numbers and sent to Boston in great packages-a bushel or more in
each package-in the late fall or winter season, and Prof. Jablonowski
volunteered to make every effort the following winter to send over
a large quantity. Taking into consideration the small size of the
brown-tail moth caterpillars during hibernation, it seemed very
strange that they should be so extensively parasitized as indicated
by Jablonowski. The larger caterpillars in the late spring and early
summer would seem to be much more likely to be extensively infested.
These winter nests, remaining alone on the trees after the leaves have
fallen, would seem to be an attractive place for small Hymenoptera
of various kinds, in which they might seek shelter for hibernation, and,
while of course there was a chance that some of the true parasites of
later stages might thus be sheltered, it was with considerable doubts
as to the ultimate result that the writer arranged for the importation
of these nests in large quantity. Even if unsuccessful, however, it
seeted thathe experiment must be tried.
From Budapest, Dresden was reached, and, as in Vienna and
Budapest, the principal museum (the Zoological Ethnological
Museum) was at once visited. Dr. K. l Heller, at that time acting
direcu(tor of the museumn was asked to reconnmend a good man who
NARRATIVE OF PROGRESS OF WORK. 59
ight be eIployed as a professiol collector to undertake work in.
ths manner as that done by Fritz Wagner in Vienna. Dr.
Heller recommended Mr. Edward hShopfer, who was at once enaged.
Although at the date of the first visit to him the season was already
considerably advanced (July 7), Mr. Schopfer had rearing caes in
operation i his rooms, and in these ces were a number of nearly
full-grown larva of the gipsy moth. He knew the localities about
Dresden whre these inects were to be found, and at once began
ending specimens to Boston. The well-known Forest Academy at
Tharandt near Dresden, was visited, and Prof. Arnold Jacobi and his
asistant. Mr. W. Baer, were interested anrd promiied assistance,
espeially in the matter of sending specimens of (Closeioz &ycaphanta
(see PI frontispice) and C. inquisitor. Other trips were made
in the icinity of Dresden, and then the journey wa resumed to
Zurich, where, through the kindness of Dr. Ierbert Ilaviland Field,
director of the Concilium Bibluioraplicum Zoologicum, the writer
met Miss Marie RHil, aeditor of the Societas EntoIologica, a very
well-posted entomologist, especially on matters relating to Lepidop-
teran, wlo had aund b as a COr espolnCe 1t nc hroughout nortthern
Germany. She was eng agd as the official aent of the investigation
forthat part of German an able, through her own work and that
of er correspondents, to send a large amount of material to Bsato n
befoe the clse of the son of 1905, ad has since continued the
From Zurich th trip w rest.umed to Pari, where some time was
spent in ilterviewing Dr. Paul Mrchal, the entomologist of the
agricultural school conducted under the ministry of ariculture, and
other ntmoloists, and in visiting the scientific societies for the pur-
poe of interesting naturalists in the work. Many trils were taken
to towns aroud Pari in searc of the )pulp of the gi ly moth and to
visit local clectrs in search of information, after which the return
jourey was nade to Aerica.
The result of this initial trip was to demonstrate that it is an easy
matte and a comparatively inexpnsive one to iml>ort certain of the
parasites of both the gipv moth and the brown-tail moth in living
condiion into the United States. The most important part of the
Europen rage of th t two spies was visited, and the entomologists
were orgaized into an active body of ass nts.
Mention has alread been made of the number of boxes sent in by
Dr. eonardi from Sardinia. Ten boxes were shipped by Fritz Wag-
ner from Vienna, 47 boxs from Schopfer in Dresden, and 36 from
Miss Rill in Zurich, all of these containing pfaasitized larvl or pupat
of the gipsy moth or bron-tail moth.
Acting upon Prof. Jabloowski's observation concering the
stence of arasites in the interin nests of the brown-tail moth,
60 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
arrangements were made with Miss Riihl, Mr. Schopfer, Prof. A. J.
Cook, who was then in Berlin, and several volunteer collectors to send
in numbers of the winter nests. During his visit to Paris in July, the
chief of the bureau had addressed a meeting of the Entomological
Society of France on the subject of his mission and asked the members
of the society to assist in the work. The most remarkable response to
this request came from Mr. Rene Oberthiir, of Rennes, who, although
not present at the meeting, read the account in the bulletin of the
society, and placed himself and his services entirely at the disposal
of the United States authorities. During the autumn of 1905 and
the winter of 1905-6 he sent to Boston more than 10,000 winter
nests of the brown-tail moth. In all, 117,000 nests were received
and cared for during that winter.
In the autumn the laboratory house (P1. II, fig. 1, p. 56) at North
Saugus was taken possession of by Mr. Kirkland, fitted up as pre-
viously described, and occupied by Mr. Mosher; the parasite material
from Malden was brought over and installed, and arrangements were
made for the receipt of the brown-tail winter nests. Very many
large boxes were constiucted, somewhat on the plan of the Cali-
fornia parasite-rearing cage, each one large enough to contain from 500
to 1,000 nests of the brown-tail moth, the front being pierced with
auger holes in which'were inserted round-bottom glass tubes into
which the emerging parasites would come in search of light and
through which they might be examined to differentiate between the
primaries and the hyperparasites. Much carpenter work was done
during the autumn and winter months and on into the spring.
Double windows and double doors were provided, and every crack
in the laboratory rooms was sealed. Realizing that many different
kinds of insects might emerge from this large supply of silken nests,
including possibly species injurious to agriculture not previously
introduced into the United States, as well as dangerous parasites of
beneficial insects, every possible effort was made to prevent the escape
of any insect whatever from the laboratory rooms.
On account of the importance of a speedy detection of injurious
forms coming from these rearing cages, and on account of the
necessity for the most expert supervision of the laboratory end of the
experiment, Mr. E. S. G. Titus, an especially well trained expert
from the Bureau of Entomology, was assigned in the spring of 1906
to the charge of the laboratory end of the introduction.
In March, 1906, Mr. Titus, with the chief of the bureau and with
Mr. Kirkland and Mr. Mosher, visited the parasite laboratory, and
for the first time examined the contents of the imported nests.
There were in the different cages, well separated as to localities,
winter nests from almost the whole of the European range of the
brown-tail moth, from Transylvania on the southeast to Brittany
NARRATIVE OF PROGRESS OF WORK. 61
on the northwest, and from the Pyrenees on the southwest to the
shores of the Baltic on the northeast. In spite of the voluntary
assistance of such men as Rene Oberthiir and Josef Jablonowski, the
expense of getting these nests to Boston had been very considerable,
and the moment when this examination was begun was considered
to be rather a critical one. No published record of the rearing of
parasites from these winter nests was recalled by the senior author
or by any of his European correspondents, and the expensive experi-
ment rested solely on the unpublished observation of Jablonowski,
and h himself had simply seen parasites emerge from nests in the
spring. Would they prove useless ? Iad the parasitic insects, even
if useful, simply crawled into the nests for hibernation ? Or were they,
soe of them, true parasitesof the young larva ? Representative nests
were examined from a number of different localities, and the relief
and joy wee great when parasitic arva. were founll ill co1nsiderable
lnumbers il each of the nests examined, feedig within thle nest pockets
externally upon the brown-tail larva. This particular explerimnent
was a success, and the expenditure of money and trouble was justified.
About April 25 these parasites began to issue from the nests. The
nests had been gathered in all from 33 different localities, and from
some of them only a small number of )arasites was reared. In all,
about 70,000 issued, of which about s per cent were hyperptrasites.
In the earig rcaes above mentioled it was a comparatively easy
matter for Mr. Titus to selarat te the yperlrasites from tlie true
parasites and to destroy the former. Of the species issuing in that
spring-tand they continued to issue until about June 15-there were
two species which appeared to be important, namely Pteroabuw
egregius First. and IIabrobracon brevicrnis Wesm,. The latter species
proved later to have entered the nests for hibernation only.
With the cooperation of Mr. Kirkland, several lcalits were found
in which there as slight danger of forest fire and in which no work
against the moths would apparently be unlertaken for at least some
months to cole, and cololies of various sizes the three principal
ones including, respectively, 10,000, 15,000, and 25,000 parasites-
were liberated in the open. Outdoor cages had been built over trees,
and some smaller colonies of the prasites were placed in these cages.
Both the outdoor experients and the open experiments were
seriously hampered, however, by the fact that the season proved to
be one of extraordinary humidity, which caused the appearance of a
fungous disease which destroyed a large proportion of the brown-tail
moth larve in the vicinity of Boston.
Concident ith the issuing of these parasites from the nests, as the
season grew warm the young larvr swarmed from the nests and
filled the glass tubes in the breeding cages and were constantly being
destroyed by the asistants in the laboratory, and when the parasites
62 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
ceased to issue the remaining nests and larvae were burned. But
later observations showed this destruction to have been a mistake.
It. was not considered likely that other parasites could be reared
from these imported larvae if they were fed and reared as far as pos-
sible, but such proved to be the case, as will be shown later.
During the winter of 1905-6 efforts were made to import in winter-
ing conditions the two large European ground-beetles, Calosoma
sycophanta (see P1. I, frontispiece) and C. inquisitor. No success in
importing living specimens was gained until March, 1906, but from
that time on until July small consignments of living adult beetles
were received, and in all 690 living specimens of Calosoma sycophanta
and 172 of (. inquisitor arrived at Boston alive, some of them dying
soon after arrival. Colonies were started in various localities about
Boston. Consideration of the history of these two species will be
given in Bulletin 101.
After visiting the parasite laboratory in March and determining
the success of the importation of the brown-tail nests, the senior
author sailed from New York on the 17th of the month for Europe,
returning to America May 17.
Proceeding directly to Paris, Mr. Rene Oberthiir was met by appoint-
ment, and the whole subject of the summer work was carefully con-
sidered. Mr. Oberthiir is a man of affairs, proprietor of a large
printing business, a learned amateur entomologist, and the possessor
of one of the largest insect collections in the world. His advice and
assistance throughout the whole work has been most important, and
he assures the American representatives that he has highly appre-
ciated the opportunity of being of assistance and of taking part in
such an interesting piece of work. At his advice the writer proceeded
to the south of France, after interviewing correspondents and agents
in Paris, and visited Prof. Valery Mayet at the agricultural school
at Montpellier, Dr. P. Siepi, of the Zoological Gardens in Marseilles,
and Mr. Harold Powell, of Hyeres. Both Prof. Mayet and Dr. Siepi
stated that both of the injurious species of insects were rare in their
vicinity, but both promised to assist in the importation of the Calosoma
beetles. Mr. Powell proved to be a lepidopterist who had been
employed professionally by Mr. Oberthiir as a collector, and he was
engaged to collect parasitized larvae in IHyres and in the Enghadine
district. Ile sent in much good material, and later, as will be shown
in subsequent pages, organized a very efficient service in the summer
of 1909. The visit to Prof. Mayet at Montpellier, moreover, was by
no means devoid of results, since at a later date he was able to send
a few specimens of carabid beetles, and in 1908, as a result of this
personal interview, he was able to send to America the first living
specimens of the European egg parasite of the imported elm leaf-
beetle, Tetrastichus xanthomelemnx Marchal, which, as a result of this
NARRATIVE OF PROGRESS OF WORK. 63
sedin, i now possibly established in New England, although it
was. not recovered diring the summers of 1909 and 1910.
While at Marseilles interviewing Dr. Siepi, April 10, the news was
received of the eruption of Vesuius and the partial destruction
by lava flow of Bo3scatrecase and other villages on the slope of
v Ha g to interview Prof. Silvestri and Dr. Leonardi at
Portici, and fearing for their safety, the visitor proceeded at once to
Naples, arrving there the day of the great market-hous accident in
which the roof fell in from the weight of volcanic ash and a number
of rsons were killed. Eve rthing in Naples was in a state of con-
fusion; the streets were filled with volcanic ash alnost knee-deep,
and it was with great difliculty that a conveyance could be secured
to drive to Portici. Portici is almost on a direct line between Naples
and Mount esuvius, anld the agrcultural college was found to be in
bad condition; the gardens were utterly destroyed by ashes, and the
roof of the old buildin was deeply covered. The accident haplened
the week before Easter, an the majority of the faculty and students
had, on account of the catastrophe, anticipated their Ester vacations
and had depad departed for their homes, Silvestri and Leonardi among the
rest. Letters were forwarded to them, however, iving detailed
suggestions s to methods of packing ald shipment of parasites.
As in 1905,, Flornce, Vienna, Buda1)st, Dresden, Tharandt,
ad Zurich were ited. Efforts were made to learn of localities
where either the gipsy moth or the brown-til moth might reasonably
be expcted to be abundant during the sumner of 1906, and a number
of such loc alties were learled and the informatiof given to ageits.
All of the arents Id correspondentIs ere give full instructions
regading the work for the summer of 1906 and the winter of 1907.
The experience of 1905 with regard to the best methods of packing
and shipment and the best kinds of boxes used was related to all,
and these points were fully discussed, with the result that the
material received during the summer of 1906 was not only greater
in quantity but better in condition than that received during the
In Vienna the visitor ha( the good fortune to find Dr. )Gustav
Mayr, hom lhe had issed i the sumer of 1905. Dr. Mayr (since
deceased) was the European authority on several of the groups of
parasites most intimately comnected with the work in hand, and the
writer had a long consultation with him concerning the systematic
osition of some of the forms already imported and concerning the
practical possibilities of the whole series of Mcrohmenoptera.
Through him was learned the probable importance of certain egg
parasites of the brown-tail moth, which he himself had reared in
Europe a had described. As a result of this information the
ents visited later were instructed to send over egg masses of the
64 PARASITES OF GIPSY AND BROAWN-TAIL MOTHS.
brown-tail moth to Massachusetts in midsummer, and later to send
over egg masses of the gipsy moth. From the brown-tail moth eg
masses parasites were reared by Mr. Titus at North Saugus and
were observed to oviposit in native eggs. Mr. Titus reared not
only the species referred to by Dr. Mayr, namely, Telenomus phals-
narum Nees, which came from eggs forwarded by Miss Riihl and
collected in Croatia, but he also reared an interesting parasite of the
genus Trichogramma from egg masses received from Wiirtemberg,
Dalmatia, and Rhenish Prussia.
At Budapest the visitor was especially glad to be able to announce
to Prof. Jablonowski the success of the rearings of parasites from the
winter nests of the brown-tail moth, so many of which had been
brought over from Europe the previous winter on the basis of Jablo-
nowski's unpublished observations. At the time of this visit Prof.
Jablonowski was too busy completing his important work upon the
migratory grasshoppers invading Hungary to be able to promise
much assistance beyond that of corresponding with foresters and
other persons well located in Hungary in order to obtain information
as to good places to secure material.
Returning to America about the end of May, the laboratory at
North Saugus was again visited, with Mr. Kirkland and Mr. Titus,
and the work of preparing indoor cages and field cages was pushed.
In the course of the summer a number of outdoor houses were con-
structed, and in these houses it was hoped to study the breeding
habits of the imported insects.
During the summer the number of shipments received from Europe
was so large that Mr. Kirkland made no attempt to list them in his
Second Annual Report published January 1, 1907. In June, in ad-
dition to egg masses previously mentioned, larve and pupe of both the
gipsy moth and the brown-tail moth were received in number from
many different European localities, and from these a large number
of parasites of several different species were reared, the most abund-
ant having been tachina flies. In one lot received from Holland more
taclhinilds were reared than there were gipsy moth caterpillars orig-
inally. Nearly 40,000 gipsy-moth larve and pupa were received
and more than 35,000 brown-tail moth larvae and pupme. The receipt
of predatory beetles is recorded in a previous paragraph.
It will be noticed that in the work conducted so far the effort to
import parasites was confined to the continent of Europe west of
Russia, whereas the well-known occurrence at intervals in large
numbers of the gipsy moth in parts (f Russia, and especially in
southern Russia (a very good account of which will be found in the
Third Report on the Gipsy Moth, by Forbush and Fernald), seemed"
to render it desirable that search should be made in those region
for parasites. The fact, however, that during these two years
NARRATIVE OF PROGRESS OF ORK. 65
the writer ad been unable to secure answers to letters addressed
to correspondents in Rusia and the reported unsettled condition
of affair in that country deterred him during the 1905 and 1906
trips from visiting the Russian southern Provinces. In the late
summer of 1906, however, advices were received from Prof. J. Por-
chinsk, of the ministry of riculture at St. Petersbug, with the
information that in the southern part of Russia both the gipsy moth
and the brown-tal moth were at that time occurring in suficiently
great numbers to enable the collection of parasites and commending
the writer to certain officials, trained entomologists, n Simferopol
(Crimea), Kishenef (Bessaabia), and Kief. Prof. Porchinsky wrote
that he had apprised these officials of the intended visit, and plans
were therefore made to include southern Russia in the itinerary for
the spring of 1907.
During the autumn of 1906 egg masses of the gipsy moth con-
tinued to be received from parts of Europe, and during the winter
hibernating nests of the brow-tail moth were sent in. More than
111,000 nests were reived from-diferent portions of the European
range of the species. These were placed in the especially constructed
cages, and from many of them large numbers of parasites were
reared, issuing mainly during the month of May, 1907. As it hap-
pened, the month of May in New England, as well as in other parts
of the United States, was phenomenally cold and wet. As a result
of this unlooked-for condition very man of the parasites refused
to leave the nests until they were so weakened as to be unable to
survive the close confinement and careful scrutiny to which they
were necessarily subjected in order to eliminate the danger of intro-
ducing secondary parasites. As a.result, a smaller number of Ptero-
mau egreu was colonized in the sumer of 1906, but 40,000 speci-
mens were put out in several localities, the pfincipal colonies consist-
ing, respectively, of 13,000, 11,000, and 7,000 individuals. At this
time, as well as in the summer of 1906, although this fact has not as
yet been stated, a number of important parsites of the genus Mono-
dontomerus issued from the winter nests and were allowed to escape.
As will be shown subsequently, this parasite has proved to be more
important than the Pteromalus and has made a phenomenal spread.
In this iportant work wth the introduced hibernation nests
of the brown-tail moth it was early found most difficult to preserve
the health of the labratory assistants The irritating and poisonous
hairs of the browntail moth larv, of which the nests are full, soon
penetrated the skin of the assistants handling them, entered their
es and throats, and the atmoshere of the laboratory became
almost filled with them. It was necessa that the rooms should
be kept thoroughly closed; double windows and screens were used,
66 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
and the doois of the rooms were doubled, in order that a possible
secondary parasite, if accidentally liberated, should have no chance
of escape. This made the rooms very warm and increased the
irritating effect of the larval hairs. Some of the assistants employed
could not stand the work and resigned. One of the best and most
experienced helpers was induced to continue the second year only
upon the promise that he would be relieved from this especial class of
work. Spectacles, gloves, masks, and even headpieces were invented
to avoid this difficulty, but these, while greatly increasing the suffer-
ing from the heat, were not entirely effective. The most serious
result of this trouble was the breaking down in health of Mr. E. S. G.
Titus of the bureau, in charge of the laboratory at Saugus, who was
obliged to resign in May, 1907, on his physician's advice, in order
to save his life. The difficulty in Mr. Titus's case was the intense
irritation to his lungs from the entrance of the barbed hairs. Mr.
Titus was soon after appointed entomologist of the Utah Agricul-
tural Experiment Station, and the change of work and climate
fortunately brought about a speedy recovery. His necessitated
departure in the midst of important work, however, threw us into
what appeared to be a serious dilemma, but fortunately it so hap-
pened that the services of the junior author, then occupying another
position in the Bureau of Entomology in Washington, could be
spared from the other work upon which he had been engaged, and,
since he had made especial studies of the parasitic Hymenoptera
and had done a large amount of rearing of parasites in the course of his
other work, he was sent on from Washington to replace Mr. Titus
in the parasite laboratory and has since had charge of the laboratory.
One of the early points to which the junior author devoted his
attention was the invention of new methods of handling the brown-
tail nests in order to avoid the serious effect upon the work of the
breaking out of the rash on himself and his assistants. He soon
devised an apparatus like the ordinary show cases that are seen in
shops, the glass on one side being replaced by cloth with armholes,
through which the gloved hands of the worker could be thrust and
the brown-tail nests handled in full sight through the top glass.
Most of the work with these nests, it has been found, can be done in
these cases with a minimum escape of the barbed hairs. There still
continued, however, considerable trouble from the rash, since much
rearing of brown-tail larvae must be carried on under conditions in
-which such cases can not be used, and this difficulty still exists.
Miss Rilh, of Zurich, in handling and repacking the large number of
nests sent to her by her European correspondents and forwarded.by
her to Boston, has been a great sufferer from the rash. She has made'
for herself a complete costume of an especially finely woven cloth,
and has made a large light helmet covered with cloth and provided
NARRATIVE OF PROGRESS OF WORK. 67
with a cape, the spae opposite the eyes being fitted with a sheet of
very transparent celluloid. Of course this costume would be very
uncomfortable in the sumer time on account of the heat, but since
she handles her nests for the most part in the autumn and winter,
he has been able to reduce the discomfort of the brown-tail rash to
Sailin aain for Europe on April 20, 1907, the senior author landed
at Cherbourg and proceeded directly to Paris, and from Paris to
Budapest by the Oriental Express. At Budapest, by prearrange-
ment, he met Mr. Aexander Pichler, whom he had engaged as a
guide and courier for the Russian trip. After a conference at Buda-
pest with Dr. Ilorvath and Prof. Moesary, of the Natural History
Museum, and Prof. Jablonowski, of the agricultural station, he pro-
ceeed to Kief, via Lemburg. Prof. Porchiky, of the ministry of
agriculture, had arraned with Prof. Wahldemar Pospielow, of the
University of Kief, to consult with the Chief of the Bureau of Ento-
mology about future arrangements, and a conerence with Prof.
Pospielow was held, in the course of which it was agreed that one of
Pospielow's assistants, engaged especially for the purpose, at 34
rubles per month, should occupy himself throughout the summer,
under Pospielow's directions, in collecting lanrv of the gipsy moth
and brown-tail moth, forwarding material to Boston, rearing and
stuying the parasites, and conducting observations in an orchard
in the suburbs of Kief, rented by the writer for the State of Massa-
chusetts for the summer at the rate of 20 rubles per month. This
procedure ~as novel in the work, but was later tried in another
locality, as will be showi in subsequent pages.
From Kief, Pichier and the visitor proceeded to Odessa and from
Odessa to Kishenef, at which point he had been recomnnended to Dr.
Isaak Krassilstschik by Prof. Porchinsky. Through some mis-
understanding as to dates, owing to the difference between the
Russian calendar and the one in use in other parts of the world, Prof.
Krassilstschik ad mistaken the date of arrival announced in the
letter sent in advance, and was absent from Kishenef on a brief visit
to Germny. Full written instructions, however, were left for him
at Kishenef, and the visitor returned to Odessa and thence by boat
to Sebastopol, and by train to Simferopol. At Simferopol he was
expected by Prof. Sigismond okshetsky, the director of the Museum
of Natural History at that place and an enthusistic economic ento-
mologist, through whose efforts American methods in the warfare
against insects had been introduced into southern Russia. Prof.
Mokshetsky had done some rearing of the Russian parasites of both
the gipsy moth and the brown-tail moth, and was able to furnish
much valuable information. His hospitality and cordiality were of
the most encouraing nature and after consultation as to the best
68 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
methods, he promised his hearty support to the work, refusing, how-
ever, to accept any compensation from the State of Massachusetts or
from the United States Government.
The visitor then proceeded by boat from Sebastopol to Con-
stantinople, but was unable to learn of any person in Turkey having
any information on the subject of insect pests, nor was he able in the
country about Constantinople to find any indication of the occurrence
of either gipsy moth or brown-tail moth.
Leaving Constantinople, the expedition proceeded to Vienna, drop-
ping Mr. Pichler at Budapest. At Vienna the Seventh International
Congress of Agriculture was held, beginning May 22, 1907. The
visitor met there a number of delegates from the different countries
in Europe, with whom he discussed the question of parasite importa-
tion, receiving warm assurances of support, especially from Prof. Dr.
Max Hollrung, of the Agricultural Department of the University of
Halle, Prof. Dr. Karl Eckstein, of the Forest Academy at Ebers-
walde, and Prof. Dr. J. Ritzema Bos, director of the Phytopatho-
logical Station at Wageningen, Holland. While in Vienna arrange-
ments were made with Mr. Fritz Wagner for continuance of the
work, and a further consultation on the subject of parasites was held
with Dr. Gustav Mayr.
After Vienna, Mr. Schopfer was visited in Dresden, Dr. Hollrung
at Halle, Dr. R. Heymons in Berlin, Dr. Eckstein in Eberswalde,
Miss Riihl at Zurich, and Prof. G. Severin at Brussels. Prof. Severin
is connected with the Royal Natural History Museum at Brussels,
is an admirably well-posted entomologist, and is connected with the
Forest Conservation Commission of Belgium. He was able to give
good advice in the parasite work and promised assistance.
Returning to France, an important conference was held with Mr.
Rene Oberthiir, and it was arranged to establish during the summer
of 1908 a field station at Rennes, to be placed in charge of a special
expert, Mr. A. Vuillet, chosen by Prof. Houlbert, of the University
of Rennes. Through Mr. Oberthiir's courtesy it was arranged to
establish field rearing cages at a convenient point near the University
of Rennes and to carry on the work in much the same way as it had
been arranged for the present summer at Kief. The University of
Rennes having a certain connection with the University of Paris, it
was considered desirable that the cooperation of the scientific faculty
of the University of Paris be gained by direct application. This was
readily arranged, through the cordial and sympathetic cooperation
of Prof. Alfred Giard, of the faculty of science of the University of
Paris (since deceased).
In dealing with the European parasites reared at North Saugus,
considerable difficulty was experienced in ascertaining their names.
It was very desirable, of course, to have a definite name by which to
NARRATIVE OF PROGRESS OF ORK. 69
designate each species, and by which to correlate it with published
accounts of observations already made. With the assistance of Dr.
0. Schmiedeknecht, of Cassel, Germany, a number of these forms had
been named, but with others it seemed practically impossible to bring
this about by correspondence. As a result, on the trip in question
the writer made an effort, by studying the collections i some of the
principal European museums, to determine a few of the unnamed
forms reared in America from European matrial. The difficulty of
this search was surprising. The Pteroalus, for example, which had
been reared in Boston by scores of thousands and which, there-
fore, must be a very common European insect, was found to be
absolutely uprereented in the lare natural history museums of
Vienna, Dresden, Berlin, Brussels, and London; nor did it occur in
the tpe collections of Ratzebur crefully preserved by Dr. Eck-
stein at the Forest Academy at Eberswalde, where, on acount of
Ratzeburg's important work on the paraites of European forest
insects, one would naturally expect to find it. At last, in a small
special collection in the Museum of Natural Histor in the Jardin
des Plantes at Paris, Mr. I. du Buysson of the Iuseum found in the
laboratory a box contaiing parasites reared many years ago by the
French entojologist, Sichel, which had been named for him by the
eminet autholrity on parasitic HImenop)tera, Arnold Firster, of
ermany In this box were spimen of the Pteron)alus labeled
"Pt. egregis" in the handiting of F rster hiself.
Especial efforts were made on the trip to arrange for the importa-
tion of large number of the egg parasites of both species and to
mtroduce in living condition the important parasi4es of the genus
Apanteles, which, according to the visitor's field observations, are
among the most important of the European enemies of the gipsy
mot. Previous imprtations of these parasites had failed, owing
to the fact that they emerged and died on the journey. On this
trip, however, specific directions were given to agents to send in
young larNe of the second stage, and by this means living specimens
in considerable numbers were later reared in the laboratory at North
Saugus. These on issuing laid their egg in the gipsy-moth larvae
of the first stae, and from these caterpillars were secured the
cocoons of adults of a second generation which was reared through
all of its stages on American soil.
From Kief there were received two species hitherto unknown as
parasites of the gipsy moth, and one of these, being a rapid breeder,
promised to be of much assistance. This species, beloning to the
genus Meteors, seemed to produce cocoons in about 10 days after
gg laying, and will be considered later in this bulletn. -
We have previously referred to the destruction in 1906 of the
great bulk of brown-tail cateriars imorted from Europe after the
70 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
early appearance of adult parasites. Mr. Titus, in 1906, tried the
experiment of rearing a very few of these imported larve, and found
that in their later growth they gave out a second lot of parasites
entirely different from those reared in May from the very young
hibernating larvae, indicating a delayed development of eggs which
must have been laid by adult parasites the previous autumn. Among
these were at least two species, one belonging to the genus Apanteles
and the other a Meteorus. Before his resignation in 1907 he started
an extensive series of rearing experiments with the end in view of
securing these parasites in large numbers. Partly on account of his
enforced absence from the laboratory during a critical period, and
partly through the unsuitable character of the rearing cages which
were employed, the project did not meet with entire success. Only
about 1,000 of the parasites were reared, of which all but a small
percentage were the Apanteles.
The importations of the summer following the trip above described
were very large, and reasonably successful, and during June alone
872 boxes were received, many others following during July and into
August, shipments of brown-tail eggs and gipsy-moth eggs following,
and of brown-tail winter nests in the late autumn and during the
winter. As in 1906, tachinids made up the great bulk of the para-
sites secured through the importation of pupae and active caterpillars.
Notwithstanding the. improvement in methods of shipment over
previous years, Apanteles invariably hatched en route, and only dead
adults or secondary parasites were received.
Before the close of the summer it had become obvious that better
quarters for the Massachusetts laboratory were necessary. The
heating and lighting arrangements at North Saugus were insuffi-
cient; the building was not sufficiently commodious, and the location
was not convenient. Therefore, after considerable search, Mr.
Kirkland found and leased for a term of years a commodious house
at Melrose Highlands (No. 17 East Highland Avenue) (see P1. II,
fig. 2, p. 56.) The building was remodeled so far as necessary to fit
it for the work. The grounds back of the house were sufficiently
ample to enable the building of several outdoor laboratories, properly
screened and ventilated, which were planned and erected under the
direction of the junior author. The building is well warmed, lighted
with electricity, and, being close to fire protection, possesses many
advantages over the old laboratory. Moreover, it is much nearer
the central office in Boston, enabling an important saving of time in
sending to the laboratory shipments of parasites received from abroad.
The rental and the expense of construction were all borne by the
State of Massachusetts. The new quarters are also within a stone's
throw of a large area of waste land covered with scrub oak.
SARRATIE OF PROGRESS OF WORK. 1
In planning the work for the season of 1908, several new features
were introduced. The parasites constantly sent over by agents
belong to three main groups, namely, those of the order Hymenop-
tera including the ichneumon flies, the chalcis flies, and others;
those of the Diptera, including the tachina flies, and those of the
order Coleoptera, icluding the predaceous ground beetles. The
amount of material reeived had been so great, and the character of
the different life histories of the insects involved had been so diverse,
that no one expert was able to do the fullest justice to the situation.
Thereore, while the junior author was left in general charge of the
whole mass of importations and retained his expert supervision of
the work on the biology of the parasitic Ilymenoptera, Mr. C. H. T.
Townsend, of the Bureau of Entomology, was assigned to the work
on the biology of the dipterous parasites, and Mr. A. F. Burgess, also
of the Bureau of Entomoloy, was ssigned to the expert charge of
the ground beetles.
Owi to the fact that the condition of European sendings by mail
and express during the sumer of 1907 had been by no means uni-
fornly good-those fro eastern Europe, subjected to long railway
journes in addition to the sea voyage, frequently arriing in bad
condition-the second innovation was made by establishing at
Rennes, France, a general laboratory depot in addition to the field
cages and rearig station mentioned in a previous paragraph. The
expert assistant designated by Prof. Houlbert, of the University of
Renes, was Mr. A. Vuillet, who was placed in specific charge of the
general laboratory depot under the general supervision of r. Rene
Oberthir. Ir. Vuillet placed himself in relations withl the steamship
company agents at Cherbour and Havre and was kept informed as
to the dates of the sailings of steamers. Nearly all of the European
sendings were shipped to Rennes, examined, repacked, and carried
persnally by Mr. uillet to Cherbourg or Havre on the known days
of sailing of certain steamers and then placed in the hands of chief
stewards of the vessels and carried in the cold rooms to New York,
whence they were sent to Boston. Early in the course of the work
the honorable the Secretary of the Treasury, upon request of the
honorable the Secretar of Ariculture, had issued orders to the col-
lector of the port of ew York to admit all such packages without
examination and to hasten their departure for Boston through the
United States dispatch agent. The steamship oflicials showed them-
selves uniforml courteous, and as a result of thi new arrangement
the average condition of the material received proved to be much
With the installation of the new laboratory at Melrose Highlands,
and with the added space afforded by the new structures in the gar-
72 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
den, the junior author was able to carry out some new ideas with
admirable results. The first of these was the carrying on of active
winter work with parasites, especially those secured from the imported
nests of the brown-tail moth, which began to come in from Europe
in December. It was found quite possible to rear these parasites
in artificially heated rooms, feeding them upon hibernating native
brown-tail larvae brought in in their nests from out of doors, feeding
the latter upon lettuce and other hothouse foliage, and in the early
spring securing more normal food for them by sending it up in boxes
by mail from Washington and points south. In this way the rear-
ing of the parasites of the genus Pteromalus was carried forward
uninterruptedly throughout the winter, and, as during the rearing of
successive generations they multiplied exceedingly, it was possible
later in the year to liberate a vastly greater number of individuals
than had the imported species been allowed to hibernate normally in
the nests. In the course of this work the junior author invented a
rearing tray which was of the utmost advantage and which has since
greatly facilitated parasite rearing work. This tray will be described
With the importation of brown-tail moth eggs it often happened
that they hatched too soon to be of use in America; or too late,
arriving after the American eggs had all hatched. It was ascertained
by the junior author during the summer and autumn that native eggs
can be kept in cold storage until the arrival of the European egg
parasites, which were found to lay their eggs and breed in these cold-
storage eggs as freely as in those which they attack in the state of
nature. It was found that this process can be carried on for a long
time, and that successive generations of these egg parasites may be
reared from eggs retarded in their development by cold storage. It
was thus shown that it is easy to rear and liberate an almost
infinitely greater number of these egg parasites, and under favorable
conditions, than would have been possible from a simple importation
of European parasitized eggs which would have to arrive in America
at a specific time.
In the same way great advance was made in the rearing of the
tachinid parasites in Mr. Townsend's charge. This expert devised
methods and made observations that greatly added to our knowledge
of the biology of these insects and resulted in the accumulation of a
store of information of the greatest practical value, not only in the
prosecution of the present undertaking but in any problem of parasite
introduction or control that may arise later. Extraordinary and
almost revolutionary discoveries were made in the life histories of
certain of these flies, and without this knowledge the greatest success
in handling them practically could not have been reached. Certain
of these facts regarding the most important of these parasites are
NARRATIVE OF PROGRESS OF WORK. 73
related in a later part of this bulletin, and many of them have been
described in some detail in Technical Series No. 12, Part VI, Bureau
of Entomology, United States Department of Agriculture (1908),
by Mr. Townsend.
Similarly r. Burgess, in charge of the Coleoptera, succeeded in
a very perfect way in rearing and liberating the important European
predatory beetle, Calosoma sycophanta, as well as some other insects
of the family CarabideI.
While these extensive importattions from Europe were going on,
Japan had by no means been lost sight of. While it seemed probable
that the European parasites in themselves would succeed in reestab-
lising the balance of nature in New England, and in spite of the
~mewhat dangerous nature pf Japanese importations on the ground
that the Japanese gipsy moth is probably a different species and
might prove in New England even more voracious and destructive
than the European moth, there was at no tine any intentin to neglect
Japanl in the search for effctive parasites. Contllluous correspond-
ence had been carried on with Japanese entomologists, and some
hipments had been made by correspondents which resulted unsuc-
cessfully. For some tile the Apanteles previosly mentIioned was
the only gips-moth parasite known to occur in Japan. Later
inforation was received from Prof. Nawa, of Gifu, Japan, to the
effect that there exists in Japan an important egg l)arasite of the
gipsy moth. Durn the preious annual trs of tip the Chief of the
Bureau of Entomology to Europe the Eurolean service of collectors,
agents, 1and advisers had been well organized 1and istructed, and the
work duri 1908 was reasonably sure to be well continued without
further personal consultation; it was therefore decided to interrupt
the European trip for 1908 and to send a skilled agent to Japan. In
consid ring the appointment of suchan agent, Prof. Trevor Kincaid,
of the University of Washington at Seattle, was at once suggested
to the mind of the writer, primarily on account of his extraordinary
skill a collector, as indicnted in the remarkable results of his work
on the Harriman expedition to Alaska in 1S99, and also on account
of his comparative proximity to Japan and the fact that he was per-
sonally acquainted with man persons in Japan. iHe was therefore
recommended to the State ofials of Massachusetts for appointment,
and was commissioned by the State to undertake the expedition.
At the same time he was formally appointed a collaborator of the
Bureau of Entomology of the United States Department of Agricul-
ture, and the Japanese Government was forall notified by the
honorable the Secretary of Agriculture, through the Department of
State, of the intended visit, the writer having also notified by per-
sonal correspondence some of the well-known Japanese entomologists.
Prof. Kincaid sailed from Seattle on March 2, and the results of his
74 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
expedition far more than justified the expense involved. A very
large amount of parasite material was received from him in good
condition at Boston, and very many parasites from Japan were
colonized in the woodlands in New England. Prof. Kincaid was
received with the most extreme courtesy and cordiality by the
Japanese Government and by .official and private entomologists
everywhere. His work was facilitated in every possible way; assist-
ants were placed at his disposal and in this way a large number of
individuals occupied themselves in the collection of parasitized.
material. After consultation with the Japanese entomologists,
whose great cleverness in manipulation and ingenuity in devising
methods are well known, Prof. Kincaid was able to pack his shipments
in such a way as to bring about a minimum of mortality on the jour-
ney. The steamship companies showed him every courtesy, and
much of his material arrived at Melrose Highlands in better condition
than corresponding sendings received from Europe. A single indi-
cation of the value of Prof. Kincaid's work may be mentioned: From
one shipment of cocoons between 40,000 and 50,000 adults of the
Japanese Apanteles were reared and were liberated directly in the
open in Massachusetts, and this. is the species which, although
repeatedly sent by correspondents, had never arrived in New England
in such condition that a single living adult could be reared.
The European importations in the meantime continued to arrive
in numbers, and at the close of the summer it was found that the
actual number of beneficial insects liberated had been far in excess
of that for 1906 or 1907, and that the list included several species
of apparently great importance and promise that had never before
been received at the laboratory in living condition.
The successful European importations all came from western
Europe, and unfortunately the few shipments sent from Russia
arrived in very bad condition. This is considered to have been most
unfortunate, since several of the Russian parasites were very promis-
ing, and the subject of improving the Russian service was taken into
With the great success of the summer's Japanese work, and the
question of the great desirability of similar work in Russia in his mind,
the senior author, visiting the Pacific coast in the autumn of that
year (1908) on a tour of inspection of the field laboratories of the
Bureau of Entomology, called on Prof. Kincaid at Seattle and dis-
cussed with him at length the plans for 1909. Although Kincaid
expressed himself as charmed with Japan and anxious to repeat his
visit to that most interesting country, his innate honesty compelled
him to state that he considered the expense of the trip unnecessary;
that he had found the Japanese entomologists, officials, and others
so intelligent and so thoroughly competent, and at the same time
NARRATIVE OF PROGRESS OF WORK. 75
so heartily interested in the experiment, that he considered them
not ony perfectly able, but perfectly willing to carry on the work by
thenselves. Ater this authoritative expression of opinion from one
who knew the ground so well, the visitor asked Mr. Kincaid whether
he would care to spend the early summer months of 1909 in Russia,
and, upon his affrmative reply, later recomended his reappointment
to the Massachusetts State authorities for that purpose.
During the autumn and winter shipments of eggs of the gipsy
moth were received from Japan, principally from Prof. Kuwana.
From these eggs were reared numerous specimens of Anasttus bIas-
clats Fonsc., a previously known European parasite of these eggs,
a1l of another parasite belongi to a genus and species new to
science (sinc1 named by the senior author SlcheWdis kuvrans) which
has tufred out to be an important primary parasite and which is
considered in later pages. Durin the winter, also, Prof. Jablonowski,
of Budapest, sent over several thousand egg masses of the gipsy moth
collectedin various localities in I ungary. After they arrived in
Massachusetts there were reared from them and liberated under the
most favorable conditions more than 75,000 adult individuals of
Anastatub. This was a surprising thing to the laboratory
workers, since less than 1,000 parasites of this species had been
reeived from all localities, the earlier ones having come from southern
Ruissia 'd froml Japalln.
The winter of 1908-9 was spent at the laboratory, in additional
rearig operations, some of them on a large scale, amd in studying the
parasites already reared, and planning for the coming sununer.
As it happened, during the winter the brown-tail moth was intro-
duced into the United States upon nursery stock from France in
large numbers. ShiFpments of nursery stock bearing winter nests
of this insect were sent to many States of the Union. Fortunately
this was discovered early in the winter, and through prompt action
and the cooperation of the customs oficials and the railroads prob-
ably every sending was traced to its ultimate destination, and was
there inspected and the nests destroyed either by State oflicials or by
persons appointed for this purpose by the United States Department
In the sprin of 1909 it seeed necessary for the chief of the
bureau to proceed to Europe for the purpose of making an investi-
gation of the European methods of growing nursery stock, with a
view to the prevention of similar introductio in the future either
by general legislation by the United States Government or in some
other way. On this trip .he utilized the opportunity to consult
further with European agents in the importation of the parasites
and to arrange for the summer's work.
In the meantime Prof. Kincaid, whose apointment had been made
bythe State of Massachusetts and who had again been made an
76 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
official, collaborator of the Bureau of Entomology of the United
States Department of Agriculture, securing leave of absence from
the University of Washington, proceeded to Russia, and stationed
himself in Bessarabia for the purpose of collecting and sending para-
sitized material from that country to the United States. It had been
noticed by Mr. Vuillet at Rennes during the preceding summer that all
material coming from Russia had been opened on the journey and had
deteriorated in consequence. Before Prof. Kincaid's departure from
America, Russian officials had been communicated with through
correspondence between the chief of the Bureau of Entomology and
Prof. Porchinsky, of the ministry of agriculture, and also directly
between the United States Department of State and the American
ambassador at St. Petersburg through the instigation of the honor-
able the Secretary of Agriculture. The United States Government
was assured that the Russian Government would welcome the expe-
dition and would facilitate the sending of material in every way
The chief of the bureau landed at Cherbourg May 12. He pro-
ceeded immediately to Paris, where a conference had been arranged
in advance with M. Oberthiir, M. Vuillet, and Mr. Henry Brown, the
latter an English entomologist resident in Paris. At this conference
it was decided to abandon the forwarding laboratory at Rennes and
to station Mr. Vuillet, during the forwarding season, at Cherbourg.
He was instructed to engage quarters at that seaport and to arrange
for cold-storage facilities, with the intention that shipments from
France, Switzerland, and Italy should be forwarded to him to be
kept in cold storage until the date of sailing of vessels, and then
should be transferred to the cold room of the next steamer, thus
practically keeping all living specimens dormant from the time of
arrival in Cherbourg until the time of arrival in New York, making
the exposure to summer temperature practically only 24 hours or less
in Europe and 24 hours or less in the United States. In the mean-
time Mr. Oberthiir was authorized to arrange for an extensive service
in the south of France, through Mr. H. Powell, of Hyeres, one of
the agents for the year 1906. The preparation of the requisite boxes
was intrusted, as in previous years, to the superintendence of Mr.
Oberthiir, and Mr. Powell was authorized to engage as many col-
lectors as the material would seem to need, with full instructions as
to packing and shipping to Cherbourg.
The visitor then proceeded to Wageningen, Holland, where he
arranged for further assistance from Prof. Dr. J. Ritzema Bos. From
there he went to Hamburg, where he arranged with the American
Express Co. to care for shipments coming from Germany, Ru a,
and Austria-Hungary, arrangements being made to keep the material
on ice until the next steamer should sail, and in case of the breakage
Bul. 91, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE III.
FIG. 1.-ROADSIDE OAK IN BRITTANY, WITH LEAVES RAGGED BY GIPSY-MOTH CATER-
PILLARS. (L O. HOWARD, JUNE, 1909.) (ORIGINAL)
FIG. 2.-M. RENE OBERTHOR (IN CENTER), DR. PAUL MARCHAL (AT RIGHT), WITH ROAD-
SIDE OAKS (BEHIND) RAGGED BY GIPSY-MOTH CATERPILLARS. (L O. HOWARD, JUNE,
Bul. 91, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE IV.
FIG. 1.-CATERPILLAR HUNTERS IN THE SOUTH OF FRANCE, UNDER M. DILLON, 1909.
FIG. 2.-PACKING PARASITIZED CATERPILLARS AT HYERES, FRANCE, FOR SHIPMENT TO
THE UNITED STATES, 1909. (ORIGINAL.)
NARRATIVE OF PROGRESS OF WORK. 77
or other ba condition of packages arrangements were made with
Dr. Re, of the Hamburg Museum, to act as expert adviser of the
From Hamburg he proceeded to Berlin for a short consultation
ith Dr. R. emons, and thence to St. Petersburg. At St. Peters-
burg ie was assured by Mr. Montgomery Schuyler, the secretary of
the embassy, that all arrangements had been made ith the Russian
Gverme, and the sa*e assurance was given" by Prof. Porchinsky.
The Russian officials insisted that none of the 1908 packages going
out of Russia had been opened by the Russian postal authorities, and
stated that in their opinion the opening must have been done at the
German frontier by German officials. A strong letter was then writ-
ten to the IHon. David J. i1, United States ambassador to Germany,
reciting the facts, dwelling upon the importance to America of these
importations, and urging him to secure from the German Govern-
ment orders to postal officials to pass ithout opening boxes of these
parasites addressed to the American Express Co. in Hamburg. Later,
in Dresden, a reply was received from Abassador iill, stating that
the German Government consented to issue the necessary instruc-
tions, but still later, ill Paris, a aldditional colunication from the
abassador requested detailed information as to the points on the
German frontier here these sendings would enter the Empire. By
telegraphic co umm ication with Prof. Kincaid, in southlern Russia,
and the Austria agents, this information was furnished, but there
seems still to have been some opening of the Russian boxes with
resulting damage to their contents.
After Russia, Dresden, Tetschen, Vienna, Budapest, Innsbruck,
Zurich, and Paris were consecutively visited, and agents were
instructed concerning the new arrangements for shipping material.
At nsbruck the iitr tor e r the first time Prof. K. W. von Dalla
Torre, the author of the great catalogue of the hymenoptera of the
world, and got his views on the subject of the larasitic Hymenoptera
and their practical handling.
From Paris he took a trip into Normandy and Brittany with Dr.
Paul Marchal, of the ministry of agriculture of France, and Mr. Ren6
Oberthiir, for the pupose of examining into the export nursery indus-
try, and at the same time with a view of observing gipsy-moth and
brown-tail moth conditions in that part of France. (Se P1. III,
fig. 2.) It transpired that both of the injurious insects were unusu-
ally abundant in portions of this territory, and by good fortune a
small oak forest covering some hundreds of acres was found not far
from Nantes, in wich there had been an outbreak of the gipsy-moth
more serious than either Dr. larchal or Mr. Oberthiir had ever seen
or had ever heard of in France. Practically every tree was defoli-
ated (see Pl III, fig. 1), and at the time of the visit, the last week
78 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
in June, the larvae were about full grown and making ready to spin.
The natural enemies of the gipsy-moth were not abundant in this
forest, although a few were seen on trees along the highway in this
general region. Nevertheless the invariable experience in Europe is
that following such an outbreak as this parasites congregate in the
region the following year and multiply in enormous numbers. The
finding of this area, therefore, seemed fortunate, since during the
season of 1910 it seemed probable that parasites would be abundant
at that point. This hope was not fulfilled, however, and in 1910
practically no gipsy-moth larvwe were to be found in that general
In the meantime the honorable minister of agriculture for Japan
had at the request of the honorable the Secretary of Agriculture of
the United States designated Prof. S. I. Kuwana, of the Imperial
Agricultural Station at Tokyo, to be the official representative of the
Japanese Government in the parasite work to be carried on during
the spring and summer of 1909, and to conduct his operations in
cooperation with and in correspondence with the chief of the Bureau
of Entomology of the United States Department. Prof. Kuwana
has shown himself in this, as in his previous work, a man of extra-
ordinary intelligence and activity, and has sent in a number of inter-
esting and valuable lots of parasitic material which were received at
Melrose Highlands in uniformly good condition. This was due to
the great care and intelligence shown by Prof. Kuwana in its collec-
tion and in his methods of packing and shipping.
The most nearly perfect European service during the summer of
1909 was secured in France, owing to The arrangement made at the
May conference in Paris. In the south of France very many people
were employed under Mr. Powell, and several thousand boxes of good
material were received at the parasite laboratory from this region.
(See P1. IV, fig. 2.) In quantity it exceeded the total of all the
importations of a similar character made since the inception of the
work, and from it have been reared a greater number of important
tachinid parasites than have been reared from all other importations
of similar character taken together. The size of the French shipments
is largely due to the intelligent energy of AIr. M. Dillon (see P1. IV,
fig. 1), with whom the bureau was placed in relations by Mr. Powell.
Quantities of miscellaneous material were also received, as for-
merly, from numerous collectors in Germany, Austria, Italy, Hol-
land, Belgium, and Switzerland.
Prof. Kincaid's account of his Russian observations is as follows:
At the request of Dr. L. O. Howard, Chief of the Bureau of En-
tomology, United States Department of Agriculture, the writer
visited the provinces of Russia bordering upon the Black Sea during
the summer of 1909 with a view to the introduction into America
NARRATIVE OF PROGRESS OF WORK. 79
of the parasites of the gipsy moth reported to exist in that part of
Europ. Proceeding to St. Petersburg via New York and Paris, an
interview was had with Prof. Porchinsky of the Russian Bureau of
Entomology, who supplied valuable information and suggestions for
the furtherance of the investigation. Leaving the Russian capital
on April 28, a journey of 48 hour brought the writer to the city of
Kishenef and after making a survey it was decided to establish
a base of operations in the forest of Gauchesty, an area of wooded
hills adjacent to a villae of that name about 30 versts northwest
from ishenef. Since the accommodations in the village of Gau-
cesty were of an unsatisfactory character, Mr. Artemy Nazaroff,
the manager of the estate of Prince Manook Bey, on the lands of
which the more important infested areas existed, invited the writer
and his interprter to become his guests during the progress of the
investigation. A suite of roms in the guest house of Gauchesty
castle was placed at our disposal, and AMr. Nazaroff did all in his
power to forward our interests and to make agreeable our stay in
that part of Russia. An outbuilding upon the farm of the estate
ws trainsforJled into a laboratory in which was erected a set of
reariing frames for the rearing of the parasites. During the first
week of April systematic exploration of the adjacent wooded areas
was begun. The forest cover was found to consist almost exclusively
of young oaks, with a few scattering trees of other species. The
ground beneath the trees was fairly ee f rom underbrush and was
carpeted wit a rich profusion of shrubs and flowers. At a distance
of 7 v rsts from Gauche~ty w an area covered with trees of con-
siderable age aRiong Which the unlldPrbrush was Comiparatively d(ense.
Fro the forester in charge of the timbered areas upon the estate
it was learned that the gipsy moth had done great ( mage to the
forest during the previous season, lar e areas having beein completely
defoliated Thi statement was borne out by the iunense nuiber
of eg masses attached to the trees. At the time we commenced
our ivestigations the caterpillars had emerged from the eggI but
were still resting upon the hark. Few signs of previous parasitic
activity were observed beyond the discovery of a number of empty
cocoons of Apa ntele solitariu Ratz. attafrhedt to the brk of the
trees. In the ancient forest mentioned above the egg masses were
very numeros, but the number of larva upon the bark was remark-
ably small. From the abnormal appearance of most of these egg
masses, and from the fact that several Microhymenoptera were dis-
covered in them, it seemed probable that a considerable number of
the eggs had been destroyed through this agency. In other parts of
the forest no evidence was secured indicating the presence of egg
The brown-tail mnoth seemed to be practically absent from the
forested areas, but in the open rolling country between .ishenef and
Gauclesty many wild pear growing in cultivated fields were found
to be completely defiated. A large number of the larvz were
placed in rearing fraes but yielded no parasits, not even Meteorus
making its appearance.
By June 1 the caterillars of the gipsy moth had passed into the
second stae and the trees were showg obvious signs of damage,
but up to this date there was no indcation of the emergence of
SRussian of =3 Enh feet 6 versts=approxtey 4 English miles.
80 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
hymenopterous parasites either in the field or from the thousands of
larvar reared in rearing frames. It became apparent that the con-
ditions were unfavorable for the purposes in mind of assembling
parasites for export, and it was decided to shift our headquarters to
a more promising locality.
On June 5 a new base of operations was established at the town
of Bendery on the Dniester River. Quarters were selected in the
principal hotel, the Petersburgia, and in a remote corner of the exten-
sive grounds of the hostelry a temporary laboratory was constructed
in which several tiers of rearing frames were erected. The forest
conditions in this district were much more diversified than at Gau-
chesty. To the northeast of the town at a distance of 7 versts was
the' forest of Gerbofsky, occupying a dry elevated area of about
5,000 acres and consisting almost exclusively of mature oak trees.
To the southward, on the banks of the river, was the forest of Kitz-
kany, composed largely of black poplar, maple, and willow. In both
of these forests the caterpillars of the gipsy moth were found in
immense numbers, and evidence of attack by both hymenopterous
and dipterous parasites was readily obtained, although nowhere in
the abundance hoped for. For two weeks the two forests, as well as
the extensive orchards in the vicinity of Bendery and the neighboring
town of Tiraspol, were scoured for parasites. A number of Russian
boys were pressed into service and trained to assist in making collec-
tions, at which they became quite expert. Except for a few clusters
of cocoons derived from Apanteles fulvipes Hal., the only hymenop-
terous parasite to appear in considerable abundance was Apanteles
solitarius. Caterpillars of the gipsy moth attacked by this species
crawl down to the trunk or lower branches of the tree and collect in
colonies on the lower side of the branches, under bark, in cavities
and other sheltered places. Here the larva of the parasite emerges
and spins its cocoon beneath the body of its host. The task of col-
lecting these scattered cocoons was a tedious one, since it was neces-
sary to remove each one carefully from the bark without undue
pressure and also to disentangle it from the hairy body of its host.
In the forest of Kitzkany, where the conditions were favorable for
bacterial infection owing to excessive dampness, the caterpillars of
the gipsy moth were swept away in vast numbers by a bacterial
disease before any extensive defoliation took place.. The search for
hymenopterous parasites in this district soon become a vain one,
since very few of the caterpillars appeared to have escaped the
The forest of Gerbofsky, owing to its being elevated, open, and
well drained, was not favorable for bacterial infection and no trace
of disease was observed. This forest was therefore almost com-
pletely defoliated by the caterpillars, and multitudes of the insects,
failing to find any further nourishment upon the oaks, descended to
the ground, where they died in great numbers, apparently from
starvation. IIynmenopterous parasites seemed to play a relatively
small part in the destruction of the caterpillars, since the attacks of
Apanteles solitarius were of the most scattering character. In the
shrubbery growths adjacent to the main forest, where new planta'-
tions had been recently established by the forester in charge, a con-
siderable number of Calosoma were found at work destroying the
caterpillars, but their operations did not appear to extend into the
NARRATIVE OF PROGRESS OF WORK. 81
main forest, where the open grass-covered ground id not offer suf-
ficient concealment for th beetles.
Th principal check to the depredations of the caterpillars of the
psy moth in this forest came with the advent of the tachinds, the
atter apearing upon the scene after the trees had been almost or
entirely defoliated. Chalcid.flies also appeared at this time, but not
in considerable numbers. The species of Limnerium, a few speci-
mens of which had been previously received from Russia, and of
which it had been hoped to secure a supply for transfer to America,
pved to be exceedingly rare, only three s iens being found.
he larva of this paraite on emrgg fro its ost spins an elongated
silken thread, at the end of which it spins a cocoon and transforms
to the pupal state.
Considerable numbers of the cocoons of Apanteles solitaius were
collected from the forest, from the extensive orchards of the neigh-
borhood, and from clups of willow bushes conmonly found at the
edges of fields. For several weeks shipments were made almost
daily to Hamburg, from which port the packages were sllhipped in
cold storage to New York. Many difculties arose in attempting to
make rapid shipments. The postal connections were very unsatis-
factory and caused annoying delays, while at the German frontier
another cause for loss of time developed through the formalities of
the customs authorities of the German Government.
The brown-tail moth seemed to be quite uncommon in the region
about Bendery and no parasites were obsered upon the small num-
ber of larv ollected at this point.
Since it seemed desirable to cover as extensive a territory as pos-
sble during the season the writer, leaving an assistant in charge of
the laboratory and collecting organization at Bendery, journeyed
northward on June 17 and established a new center of exploration at
the city of Kief, in the province of the same name. Through the
courtesy of Prof. Waldemar Pospielow the writer was furnished with
much valuable information in regard to the forests of this portion of
Russia and concerning the areas in which the gipsy moth was known
to exist Severa immene forested areas were traversed, but as the
were for the most part purely conierous in character the gipsy motl
appeared to be quite a rare insect. Through information supplied
by Prof. Pospielow it was ascertained that at Mechnigori, a monas-
terial institution on the banks of the Dnieper, several hours by
steamer from Kief, an area of woodland existed which was infested
to a moerate extent by caterpillars of the gipsy moth, among which
the Farasite were reported to be much in evidence. A visit to the
locality showed an interesting condition. The monastery was sur-
rounded by beautiful groves of elm and oak trees in which the gipsy
moth had made considerable inroads, but the parasites had developed
to a sufficient extent to practically clear the foliage of caterpillars.
Almost the sole agency in bringing about this condition was Apan-
teles ruipe, which attacksthe larve of the gipsy moth in a manner
closely resembn Apantele ponicus, as observed during the pre-
ceding season in Japan, but in the case of the latter the caterpillars
usually die upon the leaves of the trees, whereas in the former the
caterpillars descend to the trunk and lower branches to form colonies.
On emerging from the caterpillars the parasites spin cocoons beneath
82 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
the host, which are also attached ventrally to the bark f the tree,
and as numerous caterpillars die in a restricted area a mass of Apan-
teles cocoons, often of considerable thickness, is formed. Such
masses standing out as white patches against the dark tree trunks on
which they rest may be seen for considerable distances. Cocoons of
Apantales solitarius were also observed in the forest of Mechnigori,
but were comparatively rare, so this species evidently did not repre-
sent a very important element in the control of the gipsy moth.
In the forested areas about Kief the caterpillars of the brown-tail
moth were rarely met with, but in several of the parks on the out-
skirts of the city they were found in abundance. In the grounds of
the military school a large number of magnificent oak trees were
almost denuded of foliage, and some of the other deciduous trees and
shrubs, such as poplars, rose bushes, and Crataegus, were severely
damaged. The usual brown-tail parasites were found at work, the
most effective being Meteorus. Almost every branch of the injured
trees bore the suspended cocoons of this parasite. Tachinids were
also active, so it was obvious that very few of the caterpillars would
On departing from Kief on July 9 the season was practically over,
and gipsy moths were in flight.
Returning to Bendery, it was found that the season was over so far
as Apantales solitarius was concerned, but large numbers of tachinid
puparia were in evidence. As many as possible of these were assem-
bled and shipped to America. Thie chrysalides of the gipsy moth
were also forwarded in considerable numbers in the hope of securing
These lines of work were continued till July 16, by which time the
season was so advanced that the moths were beginning to deposit their
eggs for the succeeding season. From the abundance of moths in
flight it was obvious that unless the natural parasites multiplied suf-
ficiently to control the situation the region would experience another
visitation of the same character during the following year.
Leaving Bendery on July 16, the writer returned to Paris via
Odessa, Constantinople, and Naples, arriving in New York August 28.
Owing to various unforeseen conditions, and principally owing to
the deficient transportation facilities, the material received as the
result of Prof. Kincaid's expedition proved to be unsatisfactory on
In May and June, 1910, the senior author went to Europe once
more, visited agents and officials in Italy and France, and, through
the courtesy of the Spanish and Portuguese Governments, was able
to start new official services in each of these countries for the collec-
tion and sending of parasitized gipsy-moth larvae to the United
States. In Italy Prof. Silvestri at Portici and Dr. Berlese at Flor-
ence were visited and informed as to the latest ideas of the laboratory
regarding methods of shipment. In Spain Prof. Leandro Navarro,
of the Phytopathological Station at Madrid, volunteered his services
with the approval of the minister of agriculture. In Portugal Senhor
Alfredo Carlos Lecocq, director of agriculture, placed the visitor in
relation with Prof. A. F. de Seabra, of the Phytopathological Station
NARRATIVE OF PROGRESS OF WORK. 83
at Lisbon, and the latter gldly consented to act as the agent of the
bureau in this work in Portugal. In France arrangements were made
with M. Dillon as durig the previous year in the south of France,
and arrangements were renewed with Miss R l in Zurich and Mr.
Shopfer in Dresden. The distributing aency in Hamburg was con-
ued, and a new distributin aency was started at Havre, France,
on account of its convenient proximity to the American line steamers
starting from Southampton. In order to insure the best results, Mr.
Dillon accompanied certain large shipments from Hyeres to Havre,
and personally saw that they were placed upon the channel steamer
the night before the sailing of an American line steamer from
Sendings from Japan were continued in the same manner as dur-
ing the previous year. The minister of agriculture for Japan, at the
request of the Secretary of Agriculture of the United States, again
designated Prof. S. I Kuwana, of the Imperial Agricultural Experi-
ment Station at Tokyo, to be its official representative in this work,
and he continued hisextremely valuable sendigs.
The amount received durig the sunner was larger than ever
efore, but the results obtained, owing partly to the condition of the
material on reeipt ad owing to curious seasonal fluctuations and
differences in the countries of origin and in the infested territory in
America, the results by no mean corresponded with the increased
mate.al. The work carried on in the laboratory during the season
and the resul obtained are entioned later.
In the aumn the junior author visited France and Russia for the
purpose of studing certain important points regarding the question
of alternate hosts of the parasites and methods of hibernation. The
results of his observations will be given in detail in the later section
headed "The extent to which the gipsy moth is controlled through
At the close of the season of 1910, and in part owing to the prepara-
tion of the present bulletin, a general review of the whole work was
und en, and a summing up of present conditions seemed to indi-
cate that nearly as much had already been accomplished by present
methods as could be expected. The great need at this time seemed
to be a careful study in the countries of origin of the species of appar-
ent importance wich have been sent over but have not become
established, in order to ascertain the reasons for the apparent failure;
and, further, to see on the spot what can be done wth regard to the
importation of parasites of apparently lesser importance, but which,
through the fat that they may fill in gaps in the parasitic chain and
may at the same time increase beyond their native wont when con-
fronted with American conditions may be very desirable. Accord-
84 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
ingly the junior author was commissioned to visit France, Italy, and
Russia in the winter and early spring of 1911, and subsequently to
spend the breeding season- if found desirable, in Japan. He was
given authority to employ the necessary agents in each of these coun-
tries. He sailed January 5, 1911.
KNOWN AND RECORDED PARASITES OF THE GIPSY MOTH AND OF
THE BROWN-TAIL MOTH.
When the work of introducing the parasites of the gipsy moth and
of the brown-tail moth was begun in 1905, the available assets con-
sisted of generous appropriations by the State of Massachusetts -and
the Federal Government, an abundant faith in the validity of the
theory which was to be put to test, and a long bibliographical list
of the parasites which were recorded as attacking these insects in
Europe and Japan. Of these, the appropriations have withstood
most effectively the ordeal of the years which have since passed.
Our faith in the validity of the principle at stake has also stood out
wonderfully well, when the numerous trials to which it has been
subjected are taken into consideration. It is not too much to say
that at the present time it is stronger than ever, notwithstanding
that a good many facts have come to light in this period which are
more or less flatly in contradiction to the theory of parasite control
as generally accepted at the beginning. It has more than once been
necessary to modify beliefs and ideas as previously held, in order to
make them conform to the actuAl facts. To take a pertinent exam-
ple, it was necessary to place an entirely different value upon the
bibliographical list above mentioned than that which was placed upon
it when the work was begun, and when the policies of the laboratory
were first determined.
Nearly thirty years ago the present head of the Bureau. of Ento-
mology undertook the compilation of a card catalogue of references
to the host relations of the parasitic Hymenoptera of the world.
For more than twenty years the work was continued until some
30,000 such references were accumulated. From among them. those
in which the gipsy moth was mentioned as the host were collected
and a list of gipsy-moth parasites was published in Insect Life.1
With the exception of a comparatively few recent additions this list
forms the basis of that which follows. That of the parasites which
have been recorded as attacking the brown-tail moth is largely from
the same source.
1 U. S. Department of Agriculture, Division of Entomology, Insect Life, vol. 2, pp. 210-211, 1890.
KNOWN AND RECORDED PARASITES. 85
HYMENNOPTEROUS PARASITES OF THE GIPSY MOTH (Portheria dispar L.).
Reared at laboratory. Recorded as parasite
Apantelesful s (Hal.). Apantekefuvipes (Hal.). 1
el solitarus (Rtz.). Apanteles soltariti (Ratz.).' 2
Microgaster caleata Hal.1 2
Apanteles tenebrosus (Wesm.).'
Microgaster tibialis 'ees.z
( Microgaster) Apanteles fiulpes lipari is
Apanteles glomeratus (L.).
Apantels solitarius var. mlanoscelu
Apanteles solilarius? ocmnri.e Svanov.
Meteorus rersicolor (Wesm.). 1eteorus sculellator (Nees).,
tors puchricornis (Wesm.).
Meteorus japonicus Ashm.
Pimpla (Pimpla) instigator (Fab.). Pimpa (Pmpa) instiato (Fab.a).' 2
Pimpa (Pimpla) porthetri Vier.
Pimpl (Pimpla) eminaor (Fa.). Pimpla eaminator (Fab.).1
Pimpla (Pimpla) plto Ashm.n
Pimpla (Apechths) brassicarir (Poda).
Pimpl (Pmpla) disparis Vier.
Theronia atalan (Poda). Theroi atalan oda).1
Limnerum (Hyposoter) disparis Vier. Campopler coicus Ratz.'
Linerim (Anilas ) tolorip Vier. siara t intris (Gray.).
Ihneumon disparis ( Poda. Ichumon disparis (Poda).' 2
khntumon pictus (Gmel.).1 2
Amblyteles varips dw.2
Trogusflaitorius [sic.] lutoris (Fab.)? 12
( Cryptus) Artrans am u (Gra.).
ptus cyanator Gray.
PROBABLY SECONDARY BUT RECORDED AS PRIMARY.
Mesochorus pectoralis Ratz.' 2
Msochorus graclis Brischke.1 2
Mesochor splndidlus Grav.' 2
Mesochorus confusus Holmgr.1
Mesochorus semirufus Holmgr.1
(emitdes) Astomaspisfuipes (Grav.).1 2
=A. nanus (Gray.) according to Pfan-
Hemiteles bicolorius Gravy.
Pezomachue hortensis Gray.2
Pezomachusfasciatus (Fab.) '=Pezomachus
SRecorded by the senior author in a card catalogue of parasites kept in the Bureau of Entomology.
Recorded by Dalla Torre in Catogus Hymenopterorum.
3 Japanese species.
86 PARASITES OF GTSY AND BROWN-TAIL MOTHS.
Reared at laboratory. Recorded as parasites.
Pteromalus halidayanus Ratz.1
Pteromalus pini Hartig.1
Dibracitys boucheanus Ratz.1 (Second-
Eurytoma abrotani Panzer 1 2=appendi-
gaster Swed. (Secondary.)
Eupelus basatus Fonsc. Eupelmus bsciat ifasciatus Fonsc.1 2
Monodontomerus wreus Walk.
Chalcis flaipes Panz. Chalcis callipus Kby.3
Chalcis obscurata Walk.4
Schedius kuvanx How.4
HYMENOPTEROUS PARASITES OF THE BROWN-TAIL MOTH (Euproctis
Reared at laboratory. Recorded as parasites.
Meteorus versicolor (Wesm.). Meteorus versicolor (Wesm.).5
Meteorus ictericus (Nees).5
Apanteles lacteicolor Vier. Apanteles inclusus (Ratz.) 2 5
Apanteles ultor Reinh.1 2 3
Apanteles difficilis (Nees).5
Apanteles liparidis (Bouche).5
Apanteles vitripennis (Hal.).5
Apanteles solitarius (Ratz.).5
Microgaster consulars (Hal.) -= Microgas-
ter connexa Nees.
Microgaster calceata Hal.'
Rogas geniculator Nees.2 s
Rogas testaceus (Spin.).'
Rogas pulchrnpes (Wesm.).1
Pimpla (Pimpla) examinator (Fab.). Pimpla (Pimpla) examinator (Fab.).2 "
Pimpla (Pimpla) instigator (Fab.). Pimpla (Pimpla) instigator (Fab.).1 2 3
Plmpla (Apechthis) brassicarix (Poda).
Theronia atalantx (Poda). Theronia atalantas (Poda).12 3
Campoplex conicus Ratz.5
(Campoplex) Omorgus difformis (Gmel.).6
Cryptus moschator (Fab.).'
(Cryptus) Idiolispa atripes (Grav.).'
Ichneumon disparis (Poda).6
Ichneumon sutellator (Grav.).2
I Recorded by the senior author in a card catalogue of parasites kept in the Bureau of Entomology.
2 Recorded by Dalla Torre in Catalogus Hymenopterorum.
3 Reared by Dr. S. I. Kuwana.
s Recorded by Emelyanoff.
KNOWN AND RECORDED PARASITES. 87
PROBABLY SECONDARY, BUT RECORDED AS PRIMARY.
Reared at laboratory. Recorded a parasites.
Mesocu pectoralis Ratz. 22
Mesochorus dilutws Ratz.2 3
Hemitles socialis Ratz.3
Pteromalus sp. Pteroml s rotundatus Ratz.3=Pt. chry-
sorrhra D. T.X 2
Pera nidlns Thorns. =Pt. egregius Ptermals procone Ratz.' 2
Digochis omiora Walk. Pteromai ndians Thoms.
Pteroumls Ipparm L.3
Dibrachs bouchanus (Ratz.).3 (Second-
Chalcis sdrropoda F6rst.,
Monodontomerus ae s Walk. Torus onephs Rat z. = oo o-
rus ;r#ls Walk.1 2
n ntoiurs d ntips WB1oh.1 2
AIruits oiorus 1a Rondni.2
Trichograma p. I.
Trichogramma sp. II.
Tenou ph nam Nes (?). Te nos phabl nm Nes. 2 3
DIPTEROUS PARASITES OF THE GIPSY MOTH (Portheoria dispar L.).
The followin are lists of the dipterous parasites reared and
recorded from Portetria dpar L. and Eupr s chrysorrha L. Each
list is supplemented by a list of recorded hosts for each species
These lists have been compiled from various sources, the principal
being the "Katalog der Palliarktischen Dipteren," Brauer & Bergen-
stami's "Die Zweifllgler des Kaierlichen Museums zu Wien," Fer-
nald and Forbush's "The Gipsy Moth," and the senior author's "List
of parasites bred from imported materi durin the year 1907"
(3d annual report of the superintendent for suppressing the gipsy
and brown-tail moths).
In the choice of names of the forein tachinids the Katalog der
Paliarktischen Dipteren has been followed with the exception of a
few cases in which other names have ben in use at the Gipsy Moth
Parasite Laboratory; in these few case, to avoid confusion, no
change has been made.
I by Emelyanoff.
SRecorded by Dalla Torre n Catalogus Hymenopterorum.
Recorded by the senior author in a card catalogue of parasites kept in the Bureau of Entomology.
88 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
FOREIGN TACHINID PARASITES ON PORTHETRIA DISPAR.
Blepharipa scutellata R. D. Argyrophylax atropivora R. D.
Carcelia gnava Meig. Carcelia excisa Fall.
Compsilura concinnata Meig. Compsilura concinnata Meig.
Crossocosmia sericarix Corn. Echinomyia fera L.
Dexodes nigripes Fall. Epicampocera crassiseta Rond.
Parasetigena segregata Rond. Ernestia consobrina Meig.
Tachina larvarum L. Eudoromyia magnicornis Zett.
Tachina japonica Towns. Exorista affinis Fall.
Tricholyga grandis Zett. Histoch.eta marmorata Fab.
Zygobothria gilva Hartig. Lydella pinivore Ratz.
Meigenia bisignata Schin.
Parasetigena segregata Rond.
Phryxe erythrostoma Hartig.
Ptilotachina larvincola Ratz.
Ptilotachina monacha Ratz.
Tachina larvarum L.
Tachina noctuarum Rond.
Zenillia libatrix Panz.
Zygobothria gilva Hartig.
Zygobothria bimaculata Hartig.
N. B.-It is interesting to note that only four species are common to both lists.
RECORDED HOSTS OF FOREIGN TACHINID PARASITES OF PORTHETRIA DISPAR REARED
AT THE GIPSY MOTH PARASITE LABORATORY.
BLEPHARIPA SCUTELLATA R. D.:
Acherontia atropos L.; Vanessa antiopa L.
CARCELIA GNAVA Meig.:
Malacosoma neustria L.; Orgyia antiqua L.; Stilpnotia salicis L.
('OMPSILURA CONCINNATA Meig.:
See list of recorded parasites of P. dispar
CROSSOCOSMIA SERICARI2E Corn.:
Antherxa yamamai Guer.; A. mylitta Moore; Sericaria mori L.
DEXODES NIGRIPES Fall.:
Ascometia caliginosa Hb.; Agrotis candelarum Stgr.; Bupalus piniarius L.;
Cucullia asteris Schiff.; Deilephila euphorbix L.; Eurrhypara urtica L.; Heliothis
scutosa Schiff.; Hybernia sp.; Mamestra pisi L.; Miana literosa Hw.; Ortholitha
cervinata Schiff.; Phragmatobia fuliginosa L.; Plusia gamma L.; Porthesia
similis Fussl.; Tapinostola elymi Tr.; Tephroclystia virgauriata Dbld.; Thau-
metopaea winivora Tr.; Vanessa io L.; V. polychlorus L.; V. urticx L.; Lophyrud
sp.; Nematus ribesii Scop.
PARASETIGENA SEGREGATA Rond.:
(See list of recorded parasites of P. dispar.)
TACHINA LARVARUM L.:
(See list of recorded parasites of P. dispar.)
TACHINA JAPONICA Towns.:
Porthetria dispar L.
TRICIHOLYGA GRANDIS Zett.
Arcti caja L.; Mamestra oleracea L.; M. pisi L.; Saturnia pavonia L.; S. pyri
Schiff.; Sphinx ligustri L.; Thaumetopoea pityocampa Schiff.; Vanessa io L.
KNOWN AND RECORDED PARASITES. 89
RECORDED HOSTS OF FOREIGN TACHINID PARASITES RECORDED ON PORTHETRIA
ARtGYOPHYLAX ATROPr A R. D.:
P. dpar L.; Acherontia atropos L.; Notodonta trepda Ep.; anessa io L.
CARCELIA EXCISA Fall.:
Abrotola tripartita Hufn.; A. triplasia L.; Arctia caja L.; A. hebe L.; A. illica
L.; Bupalu pnaru L; Callimorpha ominula L.; Cucullia scrophulariT
Cap.; Dasychira pudibunda L.; Endromis versicolora L.; Hyloicus pinastri L.;
P. dar L.; P. monacha L.; alacooa castresis L.; nutra L.; Orgyia
antua L.; Phragmatobia fuliginosa L.; Pterostoma palpina L.; Pygaera
urtula L.; aturnia pyri Schiff.; Sphinx gustr L.; Stilnotia salis L.;
po re pannonia Frr.; Thumetoa procsionea L.
COMPSILURA CONCINNATA Meig.:
Abraas grossularita L.; Aronycta aers L.; A. alni L.; A. uspis Ib.; A.
egarephala F.; A. rumics L.; A. triden Schiff.; Araschinia lerana L.; A.
prorsa L.; Arcti caja L.; Attarus cynthia L.; Catocala proissa Esp.; ranio-
phora liustri Fab.; ullia lactu Esp.; C. verbasci L.; Dasychira udibunda
L.; Dlina tilae L.; Diobia carucephala L.; Dipterygia sabriuscula L.;
Drymonia honia IHb.; Euprotis chrysoha L.; Hyloius pinastr L.; Liby-
tha eltis Laich.; Porthtria dispar L.; P. moncha L.; Macrothylarca rubi L.;
Mamestra brassice L.; M. oleracea L.; 3M. pericari L.; Malacosoma neustria L.;
eonistiqudra L.; Phera bephala L.; Pieri brassi L.; P. rapr L.; Plusia
fettu L. ; P. gamma L PLelocampa populi L.; Porthes siilis FiI,.;
Pygra anaora Fab.; Pyrame atalanta L.; Smernths populi L.; Spilo-
soma lubrcped L.; menthstr Esp.; Stauropu fag L.; Stilpnotia salii L.;
Tniocam pa stabilis View.; Taumetopa processionea L.; T. pityocampa
Schiff.; Tmandraamaa L.; Traheaatrplis L.; Van antpa L.; io L.;
V. urtiEC L; xanthomelas E1p.; ponometa padeila L.; Cimbex humeralis
Fourer.; Tr*icioam pus i~m inalis Fall.
EINOMYIA FERA L.:
Agroti greosa Esp.; Arctia aulica L.; Leuania oboleta Sb.; Portetra dispar L.;
P. mnaha L.; ametra pi L.; eonits quadra L.; Panolis griovariegata
EIrAMPOCEA CRASSI ETA Rond.:
Porthria dipar L.; Tumetopw proessionea L.
ERNESTIA CONSOBRINA Meig.:
Cuullia artemisii Hu1 .; Portria dispar L.
EUDOROMYT.A MtIAGNICORNIS Zett.:
Art sp. ind.; adena aduta Ep.; Porthetra dspar L.
EXORISA AFFNIS Fall.:
Acronyta alni L.; Arctia caja L.; Portetria dispar L.; Pachytelia villosella 0.;
Saturnia paronia L.; S. pyri Schiff.
HISTOCHUETA MARMORATA Fab.;
Arctia aja L.; A. queselii Payk.; A. villca L.; Cucullia vrbani L.; Malacosoma
neutria L.; Porthetria dispar L.; Goniarcta rufipes Payk.
LYDELLA PINIVOR~E Ratz.:
Porthetria dispar L.:
MEIGENIA BISIONATA Schin.:
Porthetria dispar L.:
PARASETIGENA SEGREGATA Rond.:
Portheria dipar L.; P. monacha L.; Lophyrus pini L.
PHRYXE ERYTHROSTOMA Hartig:
Dendrolimus pini L.; Haloicus pinastri L.:
90 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
PTILOTACHINA LARVINCOLA Ratz.:
Porthetria dispar L.:
PTILOTACHINA MONACHA Ratz.:
Porthetria dispar L.:
TACHINA LARVARUM L.:
Acronycta rumicis L.; Agrotis praecox L.; Arctia caja L.; A. villica L.; Catocola
fraxini L.; Cosmotriche potatoria L.; Cucullia prenanthis B.; Dasychirafascellina
L.; Deilephila gallii Rott.; D. euphorbia3 L.; Dendrolimus pini L.; Gastropacha
quercifolia L.; Lasiocampa quercus L.; Porthetria dispar L.; P. monacha L.;
Macroglossa stellatarum L.; Macrothylacia rubi L.; Malacosoma castrensis L.;
M. neustria L.; Mamestra brassicme L.; Melitaea didyma 0.; Melopsilus porcel-
lus L.; Ocneria detrita Esp.; Olethreutes hercyniana Tr.; Orgyia ericx Germ.;
0. gonostigma F.; Orthosia humilis F.; Panolis griseovariegata Goeze; Papilio
machaon L.; Plusia iota L.; Saturnia pyri Schiff.; Stilpnotia salicis L.; Vanessa
antiopa L.; Vanessa io L.; V. polychloros L.; V. urticxe L.; Yponomeuta
evonymella L.; Lophyrus pini L.; Pamphilius stellatus Christ.
TACHINA NOCTUARUM Rond.:
Cosmotriche potatoria L.; Porthetria dispar L.:
ZENILLIA LIBATRIX Panz.:
Abrostola asclepiadis Schiff.; Brephos nothum Hb.; Dasychira pudibunda L.; Laren-
tia autumnalis Strom.; Porthetria dispar L.; Malacosoma neustria L.; Pygwra
pigra Hufn.; Thaumetopcaa processionea L.; Yponomeuta evonymella L.; Y.
ZYGOBOTHRIA GILVA Hartig:
Porthetria dispar L.; Stauropus fagi L.; Lophyrus laricis Jur.; L. pallidus Klug.;
L. pini L.; L. rufus Latr.; L. variegatus Hartig.
ZYGOBOTHRIA BIMACULATA Hartig:
Lymantria monacha L.; Lophyrus pallidus Klug.; L. pini L.; L. rufus Latr.; L.
socius Klug.; L. variegatus Hart.; L. virens Klug.
NATIVE DIPTERA REARED FROM PORTHETRIA DISPAR.
Exorista blanda 0. S. Exoristafernaldi Will.
Exorista pyste Walk. Tachina mella Walk.
Other than Tachinidae: 2
Aphiochxta setacea Aldr. Phora incisuralis Loew
Aphiochata scalaris Loew. Sarcophaga sp..
Gaurax anchora Loew.
1 These have only been reared very occasionally at the Gipsy Moth Parasite Laboratory.
2 At the Gipsy Moth Parasite Laboratory these have been recorded only as scavengers and not as para-
Ii : ":-
KNOWN AND RECORDED PARASITES.91
DIEROUS PARASITES OF THE BROWN-TAIL MOT (Euproctis chrysor-
FOREIGN TACHNID PARASITES OF EUPROCTIS CHRYSORRH(EA.
.Complura concinnata Meig.
Compilura concinnata Meig. Echinomyia praeceps Meig.
Cycloophry aner Town. yciaferrugiea Meig.
Dexodes nigripes Fall. Pales pavida Meig.
Digoni ta setipeni Fall. Taclina laros Rond.
Donichmta spinipeniis Meig. Zenillia fauna Meig.
Eudoromya mag i Zett. Zenillia libar Panz.
Masicera ylvatca Fall.
Nemorilla notabilis Meig.
Pales parida Meig.
Parexorista cheloniB Rond.
Tachina larrarum L.
Tricholyga grandis Zett.
Zenillia libatrix Paunz.
Zygobothria nidicola Towns.
N. B.-It is interesting to note that only three spies are common to both lists.
RECORDED HO~ orF FOREIGN TACHINIDS REARED FROM EUPROCTIS CHRYSORRH(EA
AT ~HE GISY MOT PARASITE LABORATORY.
BLEPHARIDEA (PHIYRXE) VULOARIS Fall.:
Abraxas grossularata L.; Adopa lineola 0.; Aporia ragi L.; Araschina levamn
L.; A. prorsa L.; Arynnis lathoia L.; Arctia hebe L.; Boarma lariiria Dbld.;
rotoloia tiulosa L.; alymna trapezna L.; Cootrhe potatoria L.;
Cucullia anthemidis Gn.; C. asteris Schiff.; C. verbasci L.; Dendrolimus pini L.;
Ephyralinearia Hb.; Epineuronia cespitis F.; Euchloe cardamines L.; Euple.ia
lucipara b.; yberna dfolara Cl.; Ilylo pnatri L.; Hylophila prai-
aa L.; Leuania alipca F.; L. lythargyria Esp.; 1aestra d a F.;
M. per ri L.; M. retiiata Vill.; Mita athala Rott.; Metopsils porcelus
L.;; nia typia .; Parasem plantaginis L.; Piers brac L.; P. dapli-
dice L.; P. rap L.; Plusia Lgamma ; Thamnnna avar L.; Thumtopa
pityoampa chiff.; T. proceion L.; Too pa painum Tr.; nessa
antiopa L.; V. o L.; urt L.; anthomelas Esp.; Zygna achille
Esp., ab. janthina; Z. filipendul L. (?); Prorues cria L.
CoMPSruURA CONCINNATA Meig.:
See host list of tachinid paraite of P. dipar.
CYCLOTOPHRY8 ANSER ,Towns.:
No record other than at the Gi Moth Parasite Laboratory.
DEXODES (LYDELLA) NIORIPES Fall.:
A etia caliginosa Hb.; Agrots candelarum Sr.; Bupalu piiarius L.;
ullia ater Shiff.; Deilephila euphrbi L.; Eurrhypra urtcata L.; Heli-
ot utoa Schiff.; yberia sp.; Mamestra pii L.; Miana literosa w.;
Ortholitha cervinata Schiff.; Phragmatobia fuliginosa L.; Plusia gamma L.;
Porthea imil F .; Tapinostola elymi Tr.; Tephroclystia virgaureata Dbld.;
Thmeopa piniora Schiff.; anessa io L.; V. polychloros L.; V. urticx L.;
Lophyrus sp.; Nematus ribesii Scop.
DIGONICE TA SETIPENNIs Fall.:
Grapholitha strobilella L.; Notodonta trepida Esp.; Pheosia tremula Cl.'(?); Forf!-
cula auricularia L.
92 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
DIGONICHAETA SPINIPENNIS Meig.:
Lasiocampa quercus L.; Panolis griseovariegata Goeze.
EUDOROMYIA MAGNICORNIS Zett.:
Agrotis sp.; Hadena adusta Esp.; Porthetria dispar L.
MASICERA SYLVATICA Fall.:
Apopestes spectrum Esp.; Cucullia verbasci L.; Deilephila euphorbias L.; D. gallii
Rott.; D. vespertilio Esp.; Dilina tilix L.; Gastropacha quercifolia L.; Lasio-
campa quercus L.; Nonagria typhlize Thbg.; Pieris brassicse L.; Satrnia pav-
onia L.; S. pyri Schiff.; S. spini Schiff.; Sphinx ligustri L.
NEMORILLA NOTABILIS Meig.:
Notocalia uddmanniana L.; Plusia festuce L.; Sylepta ruralis Scop.; Tachyptylia
PALES PAVIDA Meig.:
Acronycta tridens Schiff.; Agrotis stigmatica Hb.; A. xanthographa F.; Attacus
cynthia L.; A. lunula Fab.; Eriogaster catex L.; Emphytus cingillum Klug;
Euproctis chrysorrhcea L.; Orgyia erics Germ.; Panolis griseovariegata Goeze;
Plusia gamma L.; Thaumetopoea processionea L.
PAREXORISTA (EXORISTA) CHELONIIE Rond.:
Ammoconia cacimacula Fab.; Arctia caja L.; A. hebe L.; A. villica L.; Hadena
secalis L.; Macrothylacia rubi L.; Orthosia pistacina Fab.; Phragmatobia fuli-
ginosa L.; Rhyparia purpurata L.; Spilosoma lubricipeda L.; Stilpnotia salicis L.;
Cimbexfemorata L.; Pamphilius stellatus Christ.
TACHINA LARVARUM L.:
See host list of foreign tachinids recorded from P. dispar.
TRICHOLYGA GRANDIS Zett.:
Arctia eaja L.; Mamestra oleracea L.; M. pisi L.; Saturnia pavonia L.; S. pyri
Schiff.; Sphinx ligustri L.; Thaumetopoea pityocampa Schiff.; Vanessa io L.
ZENILLIA LIBATRIX Panz.:
See host list of foreign tachinids recorded as parasites of P. dispar.
ZYGOBOTHRIA NIDICOLA Tn.:
No record other than at the Gipsy Moth Parasite Laboratory.
RECORDED HOSTS OF FOREIGN TACHINIDS RECORDED AS PARASITIC ON EUPROCTIS
COMPSILURA CONCINNATA Meig.:
See list of recorded hosts of foreign tachinids recorded as parasitic on P. dispar.
ECHINOMYIA PRMECEPS Meig.:
Ilemaris fuciformis L.; Euproctis chrysorrhcea L
ERYCIA FERRUGINEA Meig.:
Euproctis chrysorrhcea L.; Melitxa athalia Rott.; M. aurinia Rott.; Porthesia
similis Fiissl.; Vanessa io L.
PALES PAVIDA Meig.:
See list of recorded hosts of foreign tachinids reared from E. chrysorrhoea L.
TACHINA LATIFRONS Rond.:
Euproctis chrysorrhoea L.; Zygxna filipendulae L.
ZENILLIA FAUNA Meig.:
Acronycta rumicis L.; Cossus cossus L.; Euproctis chrysorrhoea L.: Smerinthus
ZENILLIA LIBATRIX Panz.:
See list of recorded hosts of foreign tachinids reared from E. chrysorrhea L.
KNOWN AND RECORDED PARASITES. 93
NATIVE (AMERICAN) TAC IDS RARED FROM EUPROCTIS CHRYSORKH(EA L., AT THE
GsarY MT PARASITE LABORATORY.
B lWied. ? Phorocera leucani Coq.
Euphorocra claripenni Macq. Sturmia discas Coq.
Sgan V. de Wulp. Tachina ell Walk.
N. B-Th above spe have only been reared very occasionally. The spies, however, doubtfull
to P i Coq. has been reared through to the pupal stage in coiderale numbers.
The up have always been imperfect anrm and at the time of writing none as been reared
through to the adult.
The compilation of the catalogue of parasites was originally under-
taken in the expectation that it would prove of great servce upon
exactly such occasions as the present, when the application of the
theory of control by parasites should be put to the test. Its value
naturally depended upon the accuracy of the original records, and it
was only right to suppose that in the majority of instances these
could be depended upon. It wa equally natural to suppose that
the parasitic fauna of such common, conspicuous, and widely dis-
tributed insects as the gipsy moth and the brown-tail moth would
be well represented in these lits, which were based upon a thorough
overhauling of European literature, and it was not expected that any
parasites of particular importance would be found which were not
thus recorded, unless, indeed, they were confined to Continental Asia
or to Japan.
In the fal of 1907, as soon as the turoil of his first summer's
work permitted, the junior author attempted to make use of the
numerous bibliographical references for the purpose of learning as
much as possible of the insects with which he was to deal. One
after another, various species were taken up, until he was in possession
of practically all of the published inforation concerning perhaps
hal of the Hyenoptera listed. Then he stopped, because the
information thus gained was obviously not worth the labor. It was
not so much that recorded information was scanty, or lacking in
interest, but it was because ia great many instances it was contra-
dictory to the results of the actual rearing work which had been
carried on in the laboratory throughout the sumer. It was obviously
impossible to accept everything at its face value, and apparently
next to impossible to choose between the true and the false. But
one thing reained to be done, and that was to determine at first
hand everything which it was necessary to know concerning the
numerous spcies of parasites which it was desired to introduce into
If the list of parasites which have been reared at the laboratory
from imported eggs, caterpillars, and pupe of the gipsy moth and
the brown-tail moth be compared with the lists which have already
beengiven, the numerous and obvious differences which are immedi-
ately apparent will serve better than words to illustrate the situation
which confronted us at the close of the season of 1907.
94 PARASITES OF GIPSY AND BROWN-TAIL MOTHS.
ESTABLISHMENT AND DISPERSION OF THE NEWLY INTRODUCED
In the beginning we were very far from accrediting to that phase
of the project which has to do with the establishment and dispersion
of the newly introduced parasites the importance which it deserved.
Many widely diverse species of insects were known to have been
introduced from the Old World and firmly established in America.
Presumably they were accidentally imported, as was the case with
the gipsy moth and the brown-tail moth; presumably, also, they had
spread and increased from a small beginning, at first very gradually
and later more rapidly, until they had become component parts of the
American fauna over a wide territory. The circumstances under
which the gipsy moth was imported were well known, and a good
guess had been made as to those which resulted in the introduction
of the brown-tail moth. But these were and are rare exceptions in
this respect, and for the most part the preliminary chapters in the
story of each of the insect immigrants never have been and probably
never will be written.
Because the two very conspicuous instances of the gipsy moth and
the brown-tail moth were constantly and automatically recurring
whenever the probable future of the intentionally introduced para-
sites was considered, it was, perhaps, taken a little too much for
granted, that they were to be considered as typical and significant
of what to expect. In each instance the invasion started from a
small beginning, and while the subsequent histories were different,
the more rapid spread of the brown-tail moth was directly due to the
fact that the females were capable of flight, and the relatively slow
advance of the gipsy moth into new territory to the reverse. Even
the brown-tail moth was for some years confined to a comparatively
limited area, and it was rather expected that the parasites, if they
established themselves at all, would remain for a similar period in the
immediate vicinity of the localities where they were first given their
Accordingly, in accepting this theory without submitting it to a
test, attempts were made to encompass the rapid dissemination of the
parasites coincidently with their introduction. In 1906 and 1907
the parasites which were reared from the imported material were
mostly liberated in small and scattered colonies. In a few instances
this procedure was the best which could have been adopted; in others
the worst. Small colonies of Calosoma, for example, remained for
several years in the immediate vicinity of the point where the parent
beetles were first liberated before any material dispersion was appar-
ent (see P1. XXIV), and the small colony was thus justified.
The gipsy-moth egg parasite Anastatus, as was later determined,