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ANNALS 


OF 


The Entomological Society of America 


VOEUMEA IT; 19 ke 


EDITORIAL BOARD 


J. H. COMSTOCK, L. O. HOWARD, 


ITHACA, N. Y. 


. W. M. WHEELER, 
Boston, Mass. 


€. j. Ss. BETHUNE, 
GUELPH, ONTARIO, CANADA. 


C. W. JOHNSON, Pak CALV ERE, 


Boston, MASS. 


J. W. FOLSOM, 
URBANA, ILLS. 


V. Ll. KELLOG, 
STANFORD UNIV., CAL. 


HERBERT OSBORN, Managing Editor, 
COLUMBUS, OHIO. 


PUBLISHED QUARTERLY BY THE SOCIETY 


COLUMBUS, OHIO 


PHILADELPHIA, PA. 


WASHINGTON, D. C. 


2\52490 


Ditp. = | Soa-te erereteaive weal > ose See 


CONTENTS OF VOLUME III. 


PAGE 

BRADLEY, J. C:—Minutes of the Boston Meeting.............. 1 
SmitH J. B.—Abstract of Address Insects and Entomologists, 

Rt; cinixelactions to’ the Community atiWargesy) 0.2.5). 12 
WesstTER, F. M.—A Predaceous and Supposedly Beneficial Mite, 

iPedieuloides, Becomes: Noxious to»Many 2.3220 520. 202..5.-. 15 

Banxs, NatHAn.—Myrmeleonidae from Australia.............. 40 


WitiiaMs, Francis X.—The Anatomy of the Larva of Cecidomyia 
INesimicoloidesmwWalliannssescess4 tire OT cid ene ene eye Slee 45 
MitzMaAIn, M. B.—Some New Facts in the Bionomics of the Cali- 
iO Giad Are WO GETIG Mal CAS Us Ain bes Pease 5, 2 ok el oe eet 61 
SEVERIN, H. H. P.—A Study on the Structure of the Egg of the 
Walking Stick Diapheromera femorata Say; and the Biological 
Significance of the Resemblance of Phasmid Eggs to Seeds 83 


ForBes, W. T. M.—A Structural Study of Some Caterpillars..... 94 
Morean, T. H., and Scuutt, A. F.—The Life Cycle of Hormaphis 
MPa DNAS GIS Os Fekete ee cl aM eM is hie enn cere anne, eee ee 144 
HEADLEE, T. J.—An Apparatus for the Determination of Optimums 
Oi hemperatune rand yMoistines 7. ache eae mene, a 147 
SMITH, JOHN B.—Notes on Certain Species of Mamestra........ 154 


GomsTock=|-oh.——The Palpi of MalesSpiderss sso.) ee 161 
WuitmarsH, R. D.—North American Paniscini 
Ewinc, H. E.—The Rediscovery of a Peculiar Genus and Species 


ike) Tele AAAS ears aches tor ts sel cer oaks SANSA oe ce AOE LOG ence CAS 209 
Van Duzer, E. P.—A Revision of the American Species of Platyme- 

{ODORS ARS On NRSC 2h cE RE en) Soke a Me Lp RA 
CHAMBERLIN, R. V.—Diploda from the Western States........ 233 
HaAsEMAN, L.—The Structure and Metamorphosis of the Alimentary 

Canalvor the, Larva or Psychoda alternata Say. ..)23. 0... 277 


Wutson, H. F.—A Key to the Genera of the Sub-family Aphidinae 
anda Notes. om the Synompmiys a.8iie2. et. uae = Ue oe 314 


. oe ge 
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Annala, Vols. I and Ii, complete, cdc 0020s sv decge wetness pu srzes eS 8B .00 pe 
Annols, Vols. Land II, Parts 1, 2 and 4, each. sigh pee wast che at PE IRE Ie, 
Cig EE Part &, each... 5.4; Kine hPa tnd Taegan 2 Fave wel ore 
| REPRINTS GS alice nee 
Fivasdiee of first three phir eis ; Constitution, By-Laws and List of 
Metnberé ie foo occu ney ep teeter hae sevaiiee pein a. weber tte eh tere eeuees 
Wueecter, Wm. M. J poly iiorpblath Of ANB. sos. .ee epee cede ee pede es ps ; F 
‘ Ospoxs, Herpert—The Habits of Insects as a Factor i ‘in Classification. Ne 
Sever, H. H. anv Severn, H. C.—-Anatomical and Histological Studies — 
of the Female Reproductive Organs of the American-Saw fly, Cimbex _ 
Americana, Leach.,...:.-.. Sr bike ti. Gee Meas. CEL cats hice po ee ee ae Oe 
Frur, E. P—Some Problems in Nomenclature.......25... igs Pa a Fee" 
Hammar, A.G.—On the Nervous System of the ate of Corydalis cornuta hi 
Brapiay, J. C.—A) Case of ta eek: Sleeping Habits Seng Aculeate 
(Hymenoptera..c.e. fede. cnet Pa niotehd a2 ale Ge ato the bigs Ee Hye lysine A 
Davis, J. .J.—Notes on the Life History of the Leafy Dimorph ‘of the Box: . 
. elder Aphid, Chaitophorns negandinis Thos. . 2.528.060 ee 20 
Hama.evoy, J. C.—The Genus’ Corizus, with a Review of the North and | cote 
Middle America an Species. PISS. aed Peak cps Meee rages ae tee . 35 
Gmavur, A, A.—Biologieal Notes on Colorado Potato Beetle». <..-.-.... ee 
‘Gixavur, A, AA Monogtaphic Catalogue of the Mymarid Genus “Alaptus - a 
=. Severs, H. H. axp Seven, H. C.—Internal Organs: of Beperanidin, of - 
f Malu Sew-fly...: Soe ce. wid chy ate a eRe seat aves Re ee os 
a Surrn, C. P.—A Preliminary Study of the Araneze ‘Theraphosae of ielitornia’” 
j Davis, J..J,—Studies on Aphis as Nea AO 
Ruuey, W. A.—Monsele Attachment of Insects, /......0...0.05 darter gees Sips | 3 
had | Nexpsam, J. G.—Critichl Notes on the Classification of ‘the. Corduliinae 5 “a 
* ‘¥ (Odonata) .. Peer eben tee eee ee ees Mba egy Phe a ANd wien ti hat “15 
oy Howarp, L. O—A Key to the Ruecien of Prospaltella with ‘Table of Hosts *».. k 
. a Ot and Descriptions of Four New i a Rae ae ee 
Lae Hoop, J, D.—T'wo New Species of Idolothrips. per gigs sclan gd SRSA ey £3 ths 
BONES op ead: MR en pres he vet Gane 
Ae Seaatin Class i a gen 
Sa) ANNALS ENTOMOLOGICAL ‘SOCIETY OF AMERICA, - 
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WM. H. EDWARDS 


Plate 


ANNALS 


OF 


The Entomological Society of America 


Volume [11 NAR CEs hohe Number | 


MINUTES OF THE BOSTON MEETING. 


The Fourth Annual Meeting of the Entomological Society of 
America was called to order by the President at 10:30 A. M., 
December the 30th, 1go9, in the buildings of the Harvard Medical 
School in Brookline. 

The President announced the deaths of the following Fellows 
and Members: 

William Henry Edwards, Honorary Fellow, April 4, 1909. 

Mark Vernon Slingerland, Fellow, March 11, 1909. 

Braxton. Honoré Guilbeau, January 16, 1909. 

William Brodie, August 6, tgo9g. © 

H. M. S. Seib, August 28, 1908. 

The minutes of the last meeting were accepted as printed in 
the ANNALS. 


The President announced the appointment of the following 
committees: 

Commuttee on Resolutions: Messrs. Satterthwait & Brues. 

Commuttee on Nominations: Messrs. Gillette, C. W. Johnson 
and Burgess. 

Auditing Committee: Messrs. Field, Johnson and Sanderson. 

Commuttee to Draft Suttable Resolutions Concerning the Death 
of Mr. Edwards: Messrs. Newcomb, Wheeler and Field. 

Committee to Draft Suitable Resolutions Concerning the Death 
of Professor Slingerland: Messrs. Comstock, MacGillivray and 
Riley. 

The following report from the Executive Committee was read 
by the Secretary-Treasurer, and adopted, the constitutional 
amendments to be brought up at the next annual meeting. 


2 Annals Entomological Soctety of America [Vol. III, 


REPORT OF THE EXECUTIVE COMMITTEE, 


December 30, 1909. 


The following six persons were elected members in June: 


E. W. Berger, 
W. A. Thomas, 
R. A. Cooley, 


E. H. Smart, 
V. I. Safro, 
W. O. Strong. 


Ten more members were elected at the meeting of the com- 


mittee yesterday: 
Miss E. A. Newell, 
Mr. A. C. Lewis, 
Mr. W. W. Chase, 


Mr. S. F. Blumenfeld, 


Mr. P. E. Smith, 


Mr. T. C. Barber, 
Mr. W. V. Reed, 

Dr. C. G. Hewitt, 
Mr. J. W. Hungate, 
Mr. W. H. Shideler. 


The following resignations have been accepted and member- 


ships terminated: 
Rey. G. Birkmann, 
Mr. C. F. Groth, 
Mr. G. H. Chadwick, 
Mr. M. Rothke, 
C. E. Brown, 
A. Ellsworth, 
E. C. Greene, 
T. D. Jarvis, 
J. M. Rankin, 
C. Stevenson, 
A. J. Meidt, 


- 


Mr. C. L. Pollard, 
Miss A. M. Fielde, 
Mr. F. M. Needham, 
J. P. Baumberger, 
J. P. Cockburn, 

E. Gerstenhorn, 

J. C. Huguennin, 
Re ee ea: 

W. D. Richardson, 
W. L. Tower, 

A. F. Winn. 


The Executive Committee proposes to the Society for its con- 
sideration the following amendments to the constitution to be 
brought up at the next annual meeting. 

To amend Section I of Art. IV, by striking out the words ‘‘and 
a Secretary-Treasurer’’ and inserting in their place the words 
‘‘a Secretary and a Treasurer; but these last two offices may be 
held by the same person, so that the section will read: Art. IV. 
Officers. The officers of this Society shall be a President, two 
Vice-Presidents, a Secretary and a Treasurer; but these two last 
offices may be held by the same person.” 


Igo] Minutes of the Boston Meeting 


ics) 


To amend Art. IV., Sec. 3, to read: 

“SEC. 3. Councilor to the American Association. The 
President and preceding Past-President shall represent the 
Society upon the Council of the American Association for the 
Advancement of Science.” 

To amend Art. IV, Sec. 2, by striking out the word ‘‘who”’ and 
inserting after the words ‘‘additional members”’ the words ‘‘five 
of whom”’ and by inserting after the word ‘“‘Society”’ the follow- 
ing: ‘and the sixth shall be, ex officio, the Managing Editor.”’ 

So that the Section will read: 

‘Art. IV, Sec. 2. The business of the Society not otherwise 
provided for shall be in the hands of an Executive Committee, 
consisting of the officers named in Section I, and of six additional 
members, five of whom shall be elected from the Fellows by the 
Society, and the sixth shall be ex officio the Managing Editor. 
Four members of the committee shall constitute a quorum.” 

The committee farther recommends an amendment to the pro- 
posed amendment to Art .V, by inserting at the end thereof the 
following: ‘“‘Their term of office shall commence with the first 
of June following their election.” 

During the year a memorial drawn up by Mr. W. C. Wood 
regarding the tariff on insects and signed by the President and 
Secretary-Treasurer, was sent to the Honorable Sereno E. Payne. 
No action by Congress resulted. 


CLASSIFIED EXPENDITURES 


NonicesmOrorralis, tC. (SOCIELY) ..... a.c0 ocsminen neers $ 10.25 
Sed OUR (OOCICLY 243.4 5c «ai axe ena dian) eras s Saas $ 24.61 
(ASIC TSNGE N00) OSS, enone Rees Pa ere ar 9.25 
Annals, postage and stamps for Annals.......... 13.00 46.86 
Glegke and Stenographers, (Society)... cos... 184 
(Amal Sian cece Sa00he 
—— 16.75 
Printing Annals, 4 issues, 1000 copies each...... .$522.50 } 
mE GOLME IMSELES ch oiic 2-8) 5 Aioyie sla 01g aaah sem ey ain 55.00 
|) ott 00 
Bneravings and) electroty pes... =... s.si0.> 92 4- 61.62 
sxoressand Drayage (Annals)... 0.50 ccna oes 14.74 
NEM IGIMES youre lest ik) eich) 8s co ere sawn amen gaunt species 138.45 
PDO EU rsek ey eyy ch opting sand) shame cs reveals payee Mattenen sear shsrias tess — $866.17 
iBalanceiong hand December k. LOO9K 7: aan. cooeialisealete eienciane ena eie ae 147 .47 


$1013.64 


Of the above total cash expenditures $43.61 were for the 
general expenses of the Society and the remaining $822.56 were 
for expenses connected with the Annas. In addition to the 


4 Annals Entomological Soctety of America [Vol. III, 


total cash expenditures $866.17, there are debits on the ledgers 
to Society $54.00 and ANNALS $12.00, total $66.00, representing 
cancelled subscriptions, and the offsetting of dues of deceased 
and dropped members, etc. 


CasH RECEIPTs. 


Balance on Hand Dec. 15, 1908.................-..-...-- $108.44 
Receipts from members (excluding 10 cents refunded to 
SROEY) ..<. noe oe a nh an ws oe ate Sie eee oe alee 723 .25 
Subscription from non-Members (excluding $3.00 returned). 137.10 
Sale mf Reprints and from Excess Engraving Charges 
Refended..... .. 600505 s% sos cake es eee ee ee 44.75 
—— $1013.54 
ASSETS. 
Balance from 1908. From Managing Editor.........-..-- 10.67 
From Society and Annals...........- 264 .27 
———_— |e 2/5 
Annals: Subscriptions from non-members. WF nates Son Gews = 141.10 
Sabscriptions from members... - 2.5.55 <5 a 20+ estat eee 320.00 
Sale of Reprints 2~ . Refunded Excess Engraving Charges.. 44.75 
se 505 85 
Society. Dues from memibers... . 2... ..2- 2525-3 =25 -56-= 487 .00 
Life membership fees. 2 2.252 2a s'as a we een emne 100 .00 
Gaft from &: Henshaw... )o > 5s bette Beene 6.00 
—_——— 593 .00 
Total Asstts. < oo635 sein veee ins Chae 2s Sees $1373.79 — 


From this sum $66.00 debits representing subscriptions can- 
celled and the offsetting of dues of deceased and dropped mem- 
bers, etc., should be deducted, leaving $1,307.79 as the substan- 
tial assets at the close of the year, and leaving a balance of $339.55 
unpaid assets to carry forward. 

The Society had no liabilities at the end of the year except 
$48.30 paid by members for dues and subscriptions for rgro. 


Nov. 30, 1909. © Unpaid assets:.....3. 4.4.5 k2 (eae $339.55 
Nov. 30, 1909: (Cash on hand... -~..:. 5.43 ee eee $147 .47 
ess Ibaabilities. ..2: 7.0% 5... 5 5. Ae ee eee 48 .30 
Caner. Mice es oie A ee ed 99.17 
Total Assets forward.......... . Aer ek Ce eee eee ee $438 .72 


This shows a healthy increase in the total assets carried for- 
ward of $162.78 over 1908. In comparison of the accounts with 
1908, it must be remembered that that year included the expenses 
of but three issues of the ANNALS, while the present accounts 
include four issues, or one full volume. However, there are about 
75 members who are now two years in arrears, having paid only 


* Foot-note. The Report of the Managing Editor does not show whether 
there are unpaid subscriptions which would increase this sum. 


IgIo] Minutes of the Boston Meeting 5 


the one year’s fee when they were first elected or became Charter 
members. In all probability, many of them will eventually have 
to be dropped for the non-payment of dues, so that somewhere 
up to $150.00 of the present uncollected assets can not be counted 
on as materializing.. 

The appended resolution on the death of Professor Slingerland 
was read and adopted. 

The following papers were read: 

F.M. WesstTer: A Predaceous Mite, Pediculoides ventricosus 
producing a Dermatitis in Man. (Read by Prof. Osborn). 

R. MatTHESON: Remarks on the External Anatomy of the 
Haliphidae. 

Miss A. H. MorGan: Some Correlations of May-fly Structure 
and Habit. 

C. J. TRiGGErRson: The Life-cycle of the Oak Hedgehog Gall- 
fly (Acraspis erinacei). 

H. H. Lyman: An improved Drawer for Insect Cabinets and 
a New Substance for Lining Them. 

A. D. Macciiitivray: The Female Reproductive Organs of 
Corydalis cornuta. 


The Managing Editor gave a brief report for the Editorial 
Board; summarizing the work of the year. Four numbers of 
the ANNALS have been issued with nearly 300 pages of text and 
twenty-nine plates. The list of outside subscriptions, has been 
increased and includes many of the principal libraries. Most of 
these subscriptions are continuous and we may expect further 
additions as the publication becomes known. Members who are 
so located that they can influence librarians in placing the ANNALS 
on their permanent periodical list, can assist the growth of the 
publication by effort in this direction. Moved and carried that 
the report be accepted. 

The Society then adjourned till 2 P. M. 

At that time, the President, in calling the meeting to order, 
spoke of the approaching first International Congress of Ento- 
mology at Brussels, Belgium. It was suggested that delegates 
from this Society be appointed, and there being no objection, 
this was referred to the Executive Committee. 


The following papers were read: 
W. M. WHEELER: ~ On the Effects of Parasitic and. Other 
Kinds of Castration in Insects. 


6 Annals Entomological Society of America [Vol. III, 


W. L. W. Fietp: The Offspring of a Captured Female of 
Basilarchia proserpina. ‘To be published in Psyche. 
F. L. WAsHBURN: A Jumping Seed-gall on the Burr Oak. 


The Society then adjourned to inspect the entomological 
exhibition, which was in conjunction with and under the auspices 
of the Cambridge Entomological Club. 

At 8 p. M., the Annual Public Address was given by Dr. John 
B. Smith, in the lecture hall of the Boston Society of Natural 
History. ‘‘Insects and Entomologists: Their Relations to the 
Community at Large.”’ 

An abstract of this address appears elsewhere in this number 
of the ANNALS. 

At 10 A. M., December 31st, the Society. was again called to 
order bv the President, Dr. Skinner. 

The appended resolutions on the death of Mr. Edwards were 
read by Dr. Wheeler and adopted. 

The President remarked upon the travels of Mr. Edwards on 
the Amazon River, and the high character of the volume descrip- 
tive of that trip, which Mr. Edwards had published, many years 
ago, under the title ‘Voyage up the Amazon.” 

Mr. C. T. Brues read a paper entitled, ‘‘Some Notes on the 
Geological History of the Parasitic Hymenoptera.”’ 

The Nominating Committee made its report and the Secretary 
was directed by vote of the Society to cast a ballot for the entire 
list thus placed in nomination. 


The officers thus elected were as follows: 
President, John B. Smith. 

First Vice-President, Dr. S. A. Forbes. 
Second Vice-President, Prof. V. L. Kellogg. 
Secretary-Treasurer, Mr. C. R. Crosby. 


Additional Members of the Executive Committee: 


Prof. J. H. Comstock, Prof. J. M. Aldrich 
Dr. W. M. Wheeler, Rev. Prof. C. J. S. Bethune, 
Mr. E. A. Schwarz, Prof. Lawrence Bruner. 


Member of Committee on Nomenclature to serve three years: 
Prof. T. D. A. Cockerell (to succeed himself). 


The following report from the Auditing Committee was read 
and accepted. 


TgI0o] Minutes of the Boston Meeting 7 


REPORT OF THE AUDITING COMMITTEE. 


Boston, Mass., Dec. 31, 1909. 
To the Entomological Society of America: 

Your Auditing Committee has examined the books of the 
Treasurer and the Managing Editor of the Annals and find them 
in satisfactory condition. 

Your committee recommends that the manner of keeping the 
financial records of the Society be referred to the Executive Com- 
mittee to determine the future policy. 

Your committee wishes to express the thanks of the Society 
to the Treasurer, J. Chester Bradley, for the painstaking manner 
in which the accounts of the Society have been kept. 

(Signed) E. D. SANDERSON, 
C. W. JOHNSON, 
For the Auditing Committee. 


The following report was read and adopted: 


REPORT OF THE COMMITTEE ON RESOLUTIONS. 


The Resolution Committee begs to submit the following reso- 
lutions: 


Resolved, That we extend thanks to the local committee on 
arrangements, the Cambridge Entomological Club, the Boston 
Society of Natural History, the corporation of Harvard Univer- 
sity and the faculty of the Harvard Medical School for their 
many privileges and courtesies accorded us. 

Resolved, That the editorial management of the ANNALS of 
the Entomological Society of America, especially Professor Her- 
bert Osborn, be highly commended for the excellent standard 
which they have maintained in the publication. 

Resolved, That an expression of thanks be extended to Dr. 
John B. Smith for his admirable public address delivered in the 
hall of the Boston Society of Natural History. 

Resolved, That we also express our appreciation of the untir- 
ing work of our Secretary, Mr. J. Chester Bradley. 

(Signed) -C; I. Bruss, 
A. F. SATTERTHWAIT. 


The other members of the Committee on Nomeclature being 
absent, Dr. Felt stated that owing to the prolonged illness of one 
member of the committee and the somewhat extended absence 


8 Annals Entomological Soctety of America [Vol. III, 


from the Country of another, it had been impossible for the com- 
mittee to formulate any conclusions which could be reported 
upon at the present time. 

The report of the Committee on Nomenclature made to the 
Baltimore Meeting and printed in the ANNALS for 1909 was read 
and adopted, with the provision that the Society express itself as 
standing with the majority of the Committee in Section V. 


Mr. Brues suggested that Professor Felt submit a list of names 
of Gall Insects that he thought could be accepted as standard. 


Mr. Sanderson moved that the request from Dr. Stiles for the 
preparation of a list of one hundred important names to be adopt- 
ed by the Congress of Zoology as standard be referred to the 
Executive Committee. This motion carried. 


The following amendment to the constitution proposed at the 
Baltimore Meeting, was read: 


‘Art. V, Sec. 3, to omit the following: ‘‘All officers, Secre- 
tary-Treasurer excepted, and all additional members of the 
Executive Committee shall be chosen from the list of Fellows. 
Provided, etc.’’ The Secretary, in accordance with the recom- 
mendation of the Executive Committee moved to amend the 
amendment by adding the words at the end of the section: 
‘‘Their term of office shall commence with the first of June follow- 
ing their election.’’ Amendment to the amendment and original 
amendment carried, so that the Section now reads. 


‘‘Article V, Sec. 3. Election of officers. All officers shall be 
elected by ballot at the annual meeting for the term of one year 
and shall be eligible for re-election. Their term of office shall 
commence with the first of June following their election. 


The Secretary was instructed to take a mail vote of all mem- 
bers and Fellows of the Society as to whether the present ar- 
rangement for separate dues and subscriptions to the ANNALS 
should remain in force, or whether they should be combined into 
a single membership fee of two dollars with the provisions that all 
members should receive without further expense the publications 
of the Society. 


Prof. Sanderson suggested the adoption of a uniform style 
of button for both the entomological Societies meeting during 
Convocation Week. Referred to the officers with power. 


Tgto| Minutes of the Boston Meeting 9 


The following papers were read: 

J.C. BrapLey: The Plaiting of the Wings of Hymenoptera. 

T. J. HeapLteE: An Apparatus for the Determination of 
Optimums of Temperature and Moisture for Insects. 

A. D. MacGiiiivray: The Radial Sector in Phlebatrophia 
mathesont. 

W.T.M. Forses: A Structural Study of Some Caterpillars. 

A. G. Hammar: Notes on the Life-History of Fudiobia 
flavipes Ashmead, an Egg Parasite of the Grape Root-Worm 
Fidia viticida Walsh. 

In the absence of the authors the following papers were read 
by title only: 

C. R. Crospy: Some Observations by the late Professor 
Slingerland and the Speaker on the Life-history of Heterocordylus 
malinus Reut. 

M. J. Etrop: The Blackfoot Glacier as an Entomological 
Burying Place. 

J. J. Davis: Chattophorus popultfoliae Fitch versus Chaito- 
phorus populifoliae Oestlund. 

L. HaAsEMAN: ‘The Life-history of a Species of Psychodidae. 


The Society then adjourned, to meet during Convocation 
Weekjof rgro in Minneapolis. 


J. CHESTER BRADLEY, Secretary-Treasurer. 


10 Annals Entomological Society of America  [Vol. III, 


RESOLUTIONS 
ON THE DEATH OF WILLIAM H. EDWARDS. 


WuereEas, By the death of William H. Edwards the Entomo- 
logical Society of America has lost one of the most illustrious of 
its Honorary Fellows, and 

Wuereas, Mr. Edwards, through his magnificent and epoch- 
making contributions to our knowledge of North American Lepi- 
‘doptera was widely known as an investigator and author of the 
first rank; therefore, be it 

Resolved, That this Society express, through these resolutions, 
its grief at this loss to the scientific world, and be it further 

Resolved, That these resolutions be recorded in the Annals 
of the Society, and that the Secretary be instructed to send a 
copy of the same to the bereaved family of the eminent entomol- 


ogist. 
(Signed ) H. H. Newcoms, 


W. M. WHEELER, 
W. L. W. FIELD. 


MARK VERNON SLINGERLAND 


Plate IT. 


Igto] Minutes of the Boston Meeting IT 


RESOLUTIONS 
On THE DEATH OF PRoFEsSOR M. V. SLINGERLAND. 


Mark Vernon Slingerland, naturalist, economic entomologist, 
Fellow of this Society, died at his home in Ithaca, N. Y., March 
II, 1909. His death is a serious loss to the world of natural 
science. As a student, his career was marked by earnestness, 
courage and industry. As a teacher he was direct and forceful. 
As an investigator he was conscientious, unbiased, persevering 
and accurate, and his authority and leadership as an economic 
entomologist received unquestioned recognition. 

His memory is cherished as that of a man whose life, though 
short, stands as a notable example of one who gave his years 
unselfishly and devotedly to the discovery of useful truths in the 
realm of natural history in their relation to the economy of plant 
and animal life. We, his colleagues, give expression to our 
sorrow in the loss of a friend and fellow worker. 

(Signed) J. H. Comstock, 
A. D. MacGILiivray, 
“WILLIAM A. RILEY, 


Committee. 


INSECTS AND ENTOMOLOGISTS: 
Their Relations to the Community at Large. 
By Joun B. Smirn, Sc. D. 
(A bstract)* 


The importance of insects in their relation to the community 
at large has only become recognized during recent years, and the 
work of the entomologist is only now receiving the appreciation 
it merits. Entomologists in the sense used in the address include 
systematists, students of life histories and ecology and collectors; 
but not those studying only anatomy or histology or insects purely 
as hosts for disease-producing orgarirsms. 

Insects are injurious to man directly, as parasites, or as pre- 
datory forms attacking him, e. g., lice in the one case, biting flies 
in the other. Incidentally he may be harmed by urticating 
larvee or such as shed barbed hair, like the brown-tail caterpillars. 

Insects are further injurious as carriers of, and intermediate 
hosts for, disease-producing organisms: two totally different pro- 
cesses for, in the first case, the insect has no necessary relation 
to the disease, e. g., the house-fly to typhoid fever, while in the 
second the insect is a fellow sufferer, and the disease organism re- 
quires both man and insect to complete its life cycle, as in the case 
of the Plasmodia causing malarial diseases. The relation of 
mosquitoes to fevers, of flea to plague, and of Tsetse flies to the 
sleeping sickness was illustrated, and it was stated that if all 
dipterous insects, including fleas, could be at once eliminated, 
mankind would be at once freed from malarial, yellow, dengue, 
jungle, and several other kinds of tropical fevers, the bubonic 
plague, sleeping sickness, filariasis, several forms of eye diseases, 
certain ulcerating sores of tropical countries, and we would reduce 
to a minimum enteric fevers of all kinds, lessen the death rate 
from tuberculosis and pulmonary troubles, and probably modify 
or lessen leprosy and kindred diseases. 

* This address was delivered before the Entomological Society of America 
and its friends on the evening of December 30th, 1909. It was a popular pre- 


sentation of the subject, profusely illustrated by lantern slides, and not suitable 
as a whole for publication in a Scientific Journal. 


I2 


IgIo] Insects and Entomolgoists L3 


All domesticated animals and birds suffer from insect attack 
and parasites. Lice, bots, horse and stable flies, fleas and the 
like, and many animal diseases are carried by insects and their 
allies the ticks. Here also the Diptera contribute the bulk of the 
dangerous and troublesome species, and to our horses, cattle and 
sheep the elimination of all flies would be as great a boon as to 
man himself. 

Insects also live with man as messmates, preying on his stored 
products or acting as scavengers in his dwelling, and all the main 
orders are represented in this heading. They are further injur- 
ious by feeding upon the crops grown by him, the annual losses 
figuring up to enormous sums—estimated at $1,500,000,000 for 
the United States alone. All parts of the plants above and 
below ground, outside and inside are infested, and all kinds of 
plants are attacked. 

Many insects have been introduced from foreign countries 
and some have profoundly influenced our agricultural methods. 
The San José Scale has revolutionized fruit culture in the eastern 
United States and, incidentally, has made more positions for 
entomologists and -stimulated more interest in entomological 
work, than all other species combined:—in which respects it is 
not an unmitigated pest. 

There is, however, another side to this shield and there are 
also insects directly and indirectly beneficial. Bee products are 
of great value, and silk products are enormously so. Some insects 
are used in medicine, a very few for food, and a few also in the 
arts. 

The chief value to man, of insects, 1s as pollenizers to plants, 
and many plants are entirely dependent upon them for their 
continued existence. Pollenizers are found among the Coleoptera, 
Lepidoptera, Diptera, and, pre-eminently, the Hymenoptera. 

Portraits were shown, and brief notes were given of Say, 
Melsheimer, Haldeman, LeConte, Horn, Abbot, Morris, Grote, 
W. H. and Henry Edwards, Hulst, Strecker, Clemens, Hagen, 
Osten-Sacken, Loew, Ashmead, Packard, Scudder, Harris, Glover, 
Fitch, Walsh, LeBaron, Riley, Lintner and Fletcher. <A few col- 
lectors and founders of early societies were also referred to, Ak- 
hurst and Schaupp of Brooklyn, the latter one of the founders of 
the Brooklyn Entomological Society, and Feldman, of Philadel- 
phia, ancestor of three generations of Coleopterists—the Wenzels 
—after whom the Feldman Social was named. Pictures of field 


14 Annals Entomological Society of America  [Vol. III, 


meetings of the New York, Brooklyn, Newark and Philadelphia 
Societies in New Jersey were also shown and the address thus 
summarized. 

‘« > 3 F insects are a factor of very great importance in the 
community, first, because of their injuries, direct and indirect; 
second, because of their benefits, also direct and indirect, and 
millions of dollars annually are involved on both sides of the 
ledger.”’ 

‘‘The entomologist who studies these insects, determines which 
are harmful and which are beneficial, who works out their life 
histories and habits and who determines methods of controlling 
those that are harmful and improving those that are beneficial, 
is a worker of high importance to the community and deserving of 
every possible aid and assistance.’ 


A PREDACEOUS AND SUPPOSEDLY BENEFICIAL MITE, 
PEDICULOIDES, BECOMES NOXIOUS TO MAN. 


By F. M. WEBSTER, 
Bureau of Entomology, Washington, D. C. 


INTRODUCTION. 


Attacks of mites upon humans are by no means new, various 
instances of this character having been observed in Europe and 
recorded in various English and Continental publications both 
entomological and medical. In this country, up to very recently, 
except in the case of the itch mite, Sarcoptides psoriques Megn., 
these have all been grouped under the name “‘chigger,’’ which is 
evidently a corruption of ‘‘chigoe,”’ a tropical species, Sarcopsylla 
penetrans L., which is not a mite at all, but a flea. Thus it has 
come about that people walking during summer in grassy or 
weedy places or in woodlands are very often attacked by “‘chig- 
gers’”’ and suffer serious inconvenience and much pain on account 
of attacks of what are probably the larvae of several species of 
mites; notably the Trombidiums, just which one or how many is 
not at present known. 

The mite involved in the two epidemics of dermatitis in the 
United States, to which this paper especially relates, is quite 
certainly the same as the one discovered by myself in 1882 and 
determined for me by Mons. Jules Lichtenstein of Montpellier, 
France, as Heteropus ventricosus, Plate III, figs. 1, 3, and since 
known in this country as Pediculoides ventricosus Newport. 
Huber has since made this species a synonym of Pediculoides 
tritict Lagreze-Fossat, to which Moniez credits a large number 
of instances of mites attacking man in Europe. 


MITES ATTACKING MAN IN EUROPE AND OTHER COUNTRIES. 


In a publication relating to parasitology, both animal and 
vegetable, by R. Moniez,! quite a number of recorded instances 
have been brought together, some of them it is quite probable, 
involving the species to which this paper relates. Moniez is, 
however, vague and indefinite, and while crediting a large num- 
ber of attacks of mites upon man to Pediculotdes tritici, the fol- 


1. Traite de Parasitologi Animale et Vegetal, Applique a la Medicine, 
Paris, 1896. 


15 


16 Annals Entomological Society of America [Vol. III, 


lowing statements made by him will most assuredly not apply to 
our American species, P. ventricosus, and if these food habits 
exist in P. tritici, this fact alone would separate the two species. 
On page 463 Moniez makes these statements: 

‘It appears certain that the octopod nymphs can only under- 
go evolution if they have at their disposition a liquid nourish- 
ment; they must attach themselves either upon some vegetable, 
or, in default of this, on some animal. In the case of wheat, 
they develop upon the larvae of insects that live at the expense 
of the grain. When the nymphs are famished, they will throw 
themselves upon workmen carrying wheat and attack the skin. 

‘‘Amerling, in Bohemia, did not find the mites in company 
with parasitic insects; they can live on the grain. 

‘‘When the cereals become dry, the mites attack animal life. 
They are forced to quit the vegetable kingdom for the animal. 
In this respect they act as do the Ixodes.” 

From the foregoing one is led to suspect that the obscurities 
surrounding the identity of man-attacking mites is scarcely less 
dense in Europe than it is in this country. 

According to the observations of Lagreze? in 1849, in Espalais, 
France, a number of men engaged in carrying sacks of wheat 
experienced violent itching immediately thereafter. This wheat 
was sent to Bordeaux and Moissac where workmen in unloading 
the cargo were attacked in apparently the same manner. In the 
latter instance the men refused to work on account of the severe 
itching which immediately developed on the chest, arms, face 
and shoulders. In the case of a majority of the workmen this 
irritation of the skin was followed by an eruption of pimples 
more or less inflamed, some of which contained a serum. Later, 
experts who examined this grain reported the presence of numer- 
ous mites in the wheat and after this had been washed and dried 
in the sun, the workmen who handled it were not affected. The 
mite involved in this trouble is now known as Pediculotdes tritict. 

In 1867, Robin’, in the name of M. Rouyer, communicated to 
the Society of Biology some observations on a cutaneous disease 
observed epidemically in a large number of communities of the 
department I’Indre. The peasants engaged in gathering the 


2. (Lagreze Fossat et Montane, ‘‘Sur, la Mite du Ble.’”” Rec. Agronomique 
de la So. de sciences agric. et bellesletters de Tarnet—Garome, t. XX XII, 1851). 

3. Robin, C. R. des seances at memoires de la Soc. de biologie, 4th series, 
t. IV, 1867, page 178.) 


TQIO| Pediculoides Noxious to Man rey 


wheat after the long rains of summer, developed an itching erup- 
tion on all exposed parts of the body. 

May 1, 1872, a baker in the canton of Créon received a num- 
ber of sacks of wheat from Bordeaux. Five men who were 
engaged in carrying these sacks promptly developed severe itch- 
ing on the back, shoulders and arms, followed by an eruption of 
somewhat pointed red pimples. Fear seized the patients and 
their families, who thought themselves poisoned, but experts 
examining this wheat determined the cause of the trouble to be 
what is now known as Pediculotdes tritict. The condition caused 
by this mite has been given the name of “‘grain fever.” 

In 1875, Targioni-Tozzettit reported an eruption produced in 
a laborer who had carried sacks of wheat. 

In 1879, Geber® observed in Lower Hungary an eruptive epi- 
demic coming from barley. It appears that in the first days of 
the month of June, barley which was shipped from Lower Hun- 
gary, in sacks, was being unloaded at a railway station. After 
being engaged in this work for a few minutes, these laborers were 
attacked by a violent itching and burning and to such a degree 
did this become annoying and painful that it was with great 
difficulty that they were induced to continue their work. Geber 
desiring to obtain farther information, visited the railway sta- 
tion in question about ten days after and examined the laborers 
who had been attacked. 

In order to carry out an experiment of his own, Geber em- 
ployed an idiot to carry a sack of barley precisely as the laborers 
had done. The idiot also began unconsciously to scratch and 
immediately an eruption somewhat like nettle rash attacked him. 
It was observed at the time the laborers were handling these 
sacks of barley a yellowish-brown powder of considerable quan- 
tity fell out of the sacks and this circumstance turned the atten- 
tion of the officials to the barley. Upon a small part of this 
powder being brought under a microscope the presence of both 
living and dead mites was revealed. 

The illustrations of Geber’s paper are two figures, figure 2 
representing with reasonable accuracy what might have been a 


4. Targioni-Tozzctti, Relazione intorno ai lavori della Statione di Ento- 
mologie agraria di Firenze per l’anno, 1876, Annali, dell Agricultura, t. I, 1878. 


5. Geber, Entzundliche Prozesse der Haut, durch eine bis jetzt. nicht 
bestimmute Milbe Varursacht; Wiener Med. Presse, Vol. 20, 1879, et V. Ziems- 
sen’s Handbunchd, spec. Pathol. u Therapie, t. XIV, Handbuch, d. Hautkrank, 
2, 1884, page 412. 


18 Annals Entomological Society of America  [Vol. III, 


female of Pediculoides ventricosus, at a stage of her development 
when these mites are most abundant in grain and straw, that is 
to say they had not yet commenced to become gravid. After 
making drawings of the ventral surface, Geber instead of turning 
this same individual over and making a drawing of the reverse 
or dorsal side took for this purpose what he presumed to be anoth- 
er mite of the same species. The facts are that in all probability 
the second mite was a male, as in speaking of the striking agree- 
ment between the two individuals, he says ‘‘it remains to be 
noted only that the individuals shown by figure 2 were very 
rare, only here and there were they to be found and they were 
sasily recognized by their peculiar form.’’ In view of this it 
would seem quite probable that the mite involved in the eruption 
recorded by him might have been what we in America know as 
Pediculoides ventricosus. - 

In July 1882, Koller’ records a case where 36 workmen in 
Budapest were engaged in unloading sacks of barley coming from 
Roumania and were seized within a half hour by an intense 
itching, increasing in intensity during the several succeeding 
days. Vesicles, the largest of which were the size of a millet 
seed, appeared on inflamed bases on the neck, chest and other 
portions of the bodies of these laborers. Koller states in this 
connection that several years previously he had observed a simi- 
lar malady contracted after unloading sacks of wheat from a boat. 

Prof. Howath found a mite in the wheat which completely 
resembled that described by Robin, and several years prior to 
this, similar observations had been made on the banks of the 
Theiss, and in this case there was no other way to avoid the 
trouble except to submerge the boat with its cargo. At the same 
time that Koller observed this case, trouble was observed at 
Cologne with wheat coming from Russia. 

Flemming? in 1884 published the results of similar observa- 
tions upon workmen in Klausenburg, who unloaded wheat 
imported from Russia and were suddenly seized with a skin 
eruption. 

If we compare these records from different parts of Europe 
and involving other adjacent countries as well, we will notice 
the similarity between these epidemics of a dermatitis and those 


6. Koller, G. Ein Getreide-Milbe als Krankheitserregerin. Analysed in 
Biolog. Centralbi, t. ITI, 1884, page 127. 

7. Flemming, J. Ueber eine Geschlechtsreife Form der ala Tarsonemus 
beschrieben Thiere, Zeitschr. f. Naturwissensch B. LVII, 1884, page 472, pl. 2. 


TQIoO| Pediculoides Noxious to Man 19 


occurring in America as recorded in the following pages. It would 
however, be too much to say that in all of these cases Pediculoides 
ventricosus was the species involved; but in view of the fact 
that the Angoumois grain moth, Sztotroga cerealella, Plate III, 
fig. 2, frequently becomes even more of a grain pest in those 
countries than it does here in America, one cannot but suspect 
that either the same or a closely allied species of mite is respon- 
sible for these attacks upon man. Mr. W. D. Hunter tells me 
that in Mexico he was warned not to allow the mites to get upon 
the hands of himself or his associates in his attempts to artifi- 
cially introduce them into cotton fields attacked by the weevil, 
thus showing that its effect on man was not unknown there. 
Quite recently the writer has been informed that precisely similar 
effects have been experienced in New York City by men in hand- 
ling peas from Italy, infested by Bruchus larvae, on which Pedicu- 
loides were observed to be parasitic. 

Pediculoides ventricosus was described in 1853, and P. tritici 
Lag.—Fos was described in 1851. Huber® has made the former 
species a synonym of the latter, which, if sustained would throw 
nearly or quite all of the epidemics of dermatitis recorded to the 
credit of the one species and this would be known as Pediculoides 
tritict Lagreze—Fossat. 

In Zur Morphologie und Ontogenie der Acariden®, Dr. Enzio 
Reuter cites P. ventricosus as a good species but makes no men- 
Tomer P.. tritict. 


THe Mite BENEFICIAL IN AMERICA. 


So far as I have been able to determine, the first published 
record of the occurrence of this mite in America was by myself, 
and was included in a paper printed in the Twelfth Report of the 
State Entomologist of Illinois, for the year 1882, pages 150-151. 
While assistant to State Entomologist Dr. S. A. Forbes, I was 
directed to investigate serious injuries to stored grain by the 
Angoumois grain moth (Sztotroga cerealella) in southern Illinois, 
where Messrs. Halliday Bros., of Cairo, extensive growers and 
shippers of wheat, were at that time experiencing considerable 
trouble from the ravages of this grain moth, not only in their 
grain elevators but also, in barges loaded with wheat to be shipped 
by river to New Orleans and thence exported by steamer. 


8. Bibliographie der klinischen Entomologie (Hexapoden, Acaerinan.) 
von Med-Rath Dr. J. Ch. Huber, Jena, 1899. 

9. Acta Societatis Scientiarum Fennicae. Tom. XXXVI, No. 4, pp. 185 
and 195, 1909. 


20 ~~ Annals Entomological Society of America [Vol. III, 


It was while making these investigations, that this mite was 
discovered attacking the larvae of the grain moth. As the 
original publication containing my observations is becoming 
more and more difficult to obtain, that portion of my paper relat- 
ing to the occurrence of this mite is given herewith in full. 


Pediculoides (Heteropus) ventricosus, Newport. About the 12th 
of October, 1882, a sack of wheat infested with larvae of the grain 
moth was received from Southern Illinois, which, for want of time, 
was put aside for future inspection. On the 13th of November, while 
examining the grains containing larvae, I noticed in a lot of fifty, 
three in which the worms were dead, and on them were numbers of 
globular, yellow objects, which proved to be a species of mite Pedicu- 
loides (Heteropus) ventricosus, Newport. Knowing nothing of the pre- 
daceous habits of these mites, and the limited literature at hand 
throwing little light upon the matter, I did not pay much attention 
to the fact of their occurrence, until the 12th of December, when upon | 
examining one hundred grains with respect to the effect of heat on the 
larva, I found fourteen of the latter infested by these mites. 

In the meantime I had learned that this mite was known to be of 
predaceous habit, in both England and France, (having been first dis- 
covered by Newport, in 1849, in the nests of Anthophora retusa, col- 
lected at Gravesend, England,) and afterwards described by him 
under its present name. It had also been found in France, in 1868, 
by Jules Lichtenstein, of Montpelier, and described by him under the 
name of Physogaster larvarum. This gentleman found it in his breed- 
ing cages, which it so completely overran that, as he informs me, he 
could not for six months breed a single specimen of Hymenoptera, of 
Buprestidae, or Cerambycidae, or of some Lepidoptera. If it has been 
found by any other persons than these, or in any other parts of the 
world, previous to its discovery here by me, I have not been able to 
find the fact recorded. 

On December 31st and January 1st, I examined one hundred infest- 
ed grains of this wheat, which had been continually kept in the labora- 
tory since it was received, and found thirty-two per cent. of the worms 
dead, infested by the mites. 

While making these examinations I frequently threw the grains 
containing infested larvae into a shallow glass dish, where they re- 
mained on my table until the warm weather during the latter part of 
February, when the temperature of the laboratory at night was much 
higher than it had been during the previous cold weather. The effect 
of the change was soon plainly to be seen. The contents of the dish 
began to swarm with newly developed mites, and a larva dropped into 
their midst was immediately attacked, and after that its life was of 
short duration. Larvae placed at some distance from the dish suf- 
fered a like infection. 

To test the matter I placed near the dish some weeds, in the pith of 
which some larvae were hibernating, and in two days the mites had 
found and destroyed them. These young mites when first noticed are 


T9190] Pediculotdes Noxious to Man 21 


very minute, of elongate form, and extremely active, running about in 
search of larvae; and when one is found they immediately puncture the 
skin and suck the juices. 

In a day or two the posterior segments of the abdomen begin to 
enlarge and this process continues until the inflated, bladder-like 
abdomen becomes ten or even twenty times the size of the cephalo- 
uiorax.- Plate Ill, Fis. 3. 

During this time they have gradually lost their ambulatory 
powers, and remain stationary upon their victims. In the meantime 
changes equally wonderful have been going on within the abdomen. 

Eggs are continually forming, and within these the young mites are 

as continually developing, passing through their entire metamor- 
phosis, whtch includes the acquisition of the fourth pair of legs, (an ex- 
ceptional character among mites) within the abdomen of the mother, 
from which they make their way as fast as they reach maturity. 
_ The females are quite prolific. I have counted frequently from 
forty to fifty young and eggs within the abdomen, and believe that 
they produce even. more. The mothers survive the birth of a large 
number, if not a majority of the young. The male I have never found, 
and I am inclined to believe with Mr. Newport, that the species is 
parthenogenous.” The minute size of those young mites admits of 
their free access to the larvae of the moth, through the very small 
opening where this made its entry, and a single mite with its progeny 
would be sufficient to destroy it. 

That this is very often the manner of attack is proved by the 
fact that grains in which the larvae is badly infested frequently have 
no other break in the hull by which even.a young mite could gain ad- 
mission. Like the larvae on which they subsist, their development:is 
retarded or increased by the temperature, they being quite active at 
a temperature of 60° Fah.; but in colder weather able to remain within 
the abdomen of the parent for months in a dormant state, awaiting 
a rising temperature. 


While, as stated, this was probably the first published record 
of the occurrence of this Pediculoides in America, I have since 
had reason to believe that 1t was present many years prior to this 
date, and, indeed, in the light of information that has been ob- 
tained during the present year, 1909, it seems altogether probable 
that it not only occurred but became noxious to man, in Massa- 
chusetts, as early as 1830. 

The particular reference, however, to which attention is called 
may be found in the “‘Prairie Farmer”’ for the year 1845, page 
216. Much is here made of larvae attacking the stems of wheat 
above the upper joint and in connection therewith follows this 
significant sentence: ‘‘In one instance nine eggs were found ina 


10. I have since observed the male though only occasionally, as it is prob- 
able that each Q produces but one G among her numerous progeny. 


to 
to 


Annals Entomological Society of America [Vol. III, 


single straw, one of which had just hatched.” Also, in another 
journal, we are told that specimens of infested straw were for- 
warded to the ‘‘Country Gentleman”’ from Scipioville, New 
York in 1879, which the sender stated contained eggs besides 
larvae and pupae. In both cases, the larvae were almost beyond 
a doubt those of Meromyza americana. According to my own 
observation, these mites attack the larvae of Meromyza americana 
in stems of wheat, and, when observed, one cannot fail to be 
struck with the clearness with which the statements just given 
describe larvae of this species in the stem of grain or grass being 
attacked by these mites, the gravid 2 of which has every appear- 
ance to the unaided eye of being a minuteegg. It therefore seems 
not improbable that this Pediculotides was abroad over the 
country at the earlier date, 1845, which would antedate by sev- 
eral years the description of the species by Newport in England; 
who called attention to the occurrence of this mite as a parasite 
in the nests of Anthophora retusa, in a paper read March 5, 1850, 
before the Linnean Society of London, and with the description 
published in the transactions of this Society, Volume XXI, 2, 
P- 95, 1853. 

In the account given by Dr. T. W. Harris in the second edition 
of his ‘“‘Insects Injurious to Vegetation”’ in connection with his 
discussion of the early occurrences of the barley Isosoma, /sosoma 
hordei, there are two very significant statements that have until 
lately puzzled me very greatly. On page 438, edition of 1852, he 
states that ‘‘in the summer of 1831, myriads of these flies (mean- 
ing the adult /sosoma) were found alive in straw beds in Glouces- 
ter; the straw having been taken from the fields the year before. 
An opinion at that time prevailed, that the troublesome humors, 
wherewith many persons were then afflicted, were occasioned by 
the bites of these flies; and it is stated that the straw beds of 
Lexington, being found to be infested with the same insects, were 
generally burnt.’’ The second reference occurs on page 440 of the 
same volume, in which it is stated that ‘“‘about eight years ago . 
(which would be about 1844) some of these insects (again referring 
to I. hordei) that had come from a straw bed in Cambridge were 
shown to me. They had proven very troublesome to children 
sleeping on the bed; their bites or stings being followed by consid- 
erable inflammation and irritation, which lasted several days. 
So numerous were the insects that it was found necessary to 
empty the bed-tick and burn the straw.”’ 


TQTO| e Pediculotdes Noxious to Man 22 


Now, since first beginning the study of Jsosoma, it has always 
been a puzzle to me, why it was that the adults of /sosoma horde, 
as they were described by Dr. Harris, should have been able to 
bite through bed-ticking and cause the eruption described, and 
yet not be file to gnaw through this cloth and make their escape, 
as every one who has reared these insects in confinement has wit- 
nessed their frantic efforts to escape as soon as they gnaw their 
way out of the straw. It seems to me that, in this mite, now, we 
have as good an explanation as we can expect to secure, after 
a lapse of three-quarters of a century, with no possibility of ob- 
taining actual proof in the case. 

In 1884, I found this same mite Attacking and destroying the 
larvae of [sosoma grande at Oxford, Indiana, and in speaking of 
the occurrence of this larva and its parasites, I made this state- 
ment: ‘‘Curiously enough, during the time it occupies the stub- 
ble in the larval and pupal stages, it sometimes falls a victim to the 
mite Pediculoides (Heteropus) ventricosus, which enters the 
stubble from above after the grain is cut, but whose sense of dis- 
crimination is rather poorly developed, and it is finally victorious 
over the /sosoma larvae, its parasites,.and the predaceous larvae 
of Leptotrachelus dorsalis.’ The same year, and in the same 
locality, I again encountered this mite attacking the larvae of 
Meromyza americana in wheat straw, and again noted the 
remarkable resemblance of the gravid females to minute eggs. 
Since that time, this Pediculoides has been reported by Mr. E. H. 
Ehrhorn attacking the larvae of the peach twig borer, Anarsia 
lineatella Zeller, in California. The same year Mr. Marlatt 
reports it as attacking the eggs of the periodical ¢icada Cicada 
septemdecim.” Still later, in 1904, Messrs. W. D. Hunter and 
W.E. Hinds in Bulletin No. 45, Division of Entomology, page 107, 
called attention to its attack on the larvae of the cotton boll wee- 
vil. In 1908, Mr. W. Dwight Pierce, in Bulletin No. 73, Bureau 
of Entomology, page 30, states that this mite is a common weevil 
parasite in Mexico. Inthe same publication, page 42, he accredits 
it to being parasitic not only on the cotton boll weevil, Anthono- 
mus grandis, but also on an allied species, A. eugeni. Dr. A. D. 
Hopkins informs me that in his studies of forest insects, he has 
encountered it attacking the larvae of wood boring beetles and at 


11. Bulletin No. 10, U. S. Dept. of Agriculture, Div. of Entomology, 
p. 17, 1898. 
12. Bulletin No. 14, Div. of Entomology, n. s. p. 104. 


24 Annals Entomological Society of America [Vol. III, 


one time, in West Virginia, it caused considerable mortality in 


his breeding cages, where he was attempting to rear wood-boring 


Cerambycidae and Scolytidae, precisely as experienced, in France, 
by M. Jules Lichtenstien. 

Mr. W. J. Phillips observed this mite attacking the larve of 
Mordellistina ustulata, in Indiana, October 3, 1905, while inves- 
tigating the occurrence of these larvae in the stems of timothy 
and other grasses. 

In the publication of Messrs. Hunter and Hinds, previously 
referred to, some information is given relative to the attempt to 
use this mite artificially in overcoming the boll weevil. It has 
been experimented with quite extensively by Prof. A. L. Herrera 
and his assistants of the Mexican Commission of Parasitology, and 
upon his return from a trip to Mexico in the fall of rg02, Mr. Hunt- 
er brought with him, through the kindness of Prof. Herrera, a 
supply of the parasites, from which others were reared for experi- 
mental work in Texas. This experiment, however, owing to con- 
ditions beyond the control of man, perhaps happily so, appears 
not to have resulted satisfactorily. One of the principal ob- 
stacles in this case seems to have been that, where the mites 
succeeded in establishing themselves, they were subsequently 
destroyed by the attacks of small ants. 

These references show quite clearly the wide distribution of 
this mite throughout the United States and its great variety of 
host insects. We have, in later years, come to consider it a very 
useful parasite and one that is likely to attack almost any soft- 
bodied larvae, beyond the reach of insecticides, but to which 
it, by reason of its minute size, could gain access and be secure 
from other predaceous insects and adverse meteorological condi- 
tions. 

THe Mite Becomes Noxious To Man. 


While, as stated in the earlier portion of this paper, either 
this or some other closely allied species has long been known to 
occasionally attack man and animals in Europe, when these are 
engaged in handling or come into contact with grain or straw 
infested by their host insects, the first instance of this character 
to be noted in America has been communicated to me by: the 
present presiding officer of this Society, Dr. Henry Skinner, of 
Philadelphia, Pa. It was about the year, 1894 while Dr. Skinner 
was practicing medicine in Philadelphia, that the owner of a 
boarding house in one of the New Jersey suburbs of the city came 


7] 


Igto] Pediculoides Noxious to Man 25 


to him in great distress, stating that the tenant and keeper of 
the boarding house, which accommodated about seventy-five 
persons, would not pay the rent thereon, and further stated that 
the tenant had been threatened with legal proceedings by the 
boarders who had even suggested bodily injury. The cause of 
all of this trouble was an epidemic of a rash like disease, the 
causes of which were suspected to lie in the mattresses of the 
beds occupied by the patrons of the house, because of the occu- 
pants having been attacked by a very mysterious and aggravat- 
ing skin eruption. The owner submitted straw dust and mat- 
tress debris taken from the suspected beds and on examination 
of this Dr. Skinner found specimens of this mite. The house 
was promptly deserted by the boarders, none of whom as it seems 
escaped infection and none of whom were willing to return. The 
matter does not appear to have been further investigated. 

In the Philadelphia Medical Journal for July 6, 1901, Jay F. 
Schamberg, M. D:, of Philadelphia, published a short paper, call- 
ing attention to and describing ‘‘An Epidemic of a Peculiar and 
Unfamiliar Disease of the Skin.” In this paper, Dr. Schamberg, 
who, besides being a practicing physician, is professor of Derma- 
tology, and Infectious Eruptive Diseases, in the Philadelphia 
Polyclinic, described a number of cases that had been treated by 
him a few weeks prior to the publication of this paper. The 
eruption and its effect on the patient were briefly described 
and illustrated, but the causes instrumental in bringing about 
these attacks were still unknown to him; and as several mem- 
bers of the same household were commonly affected the dis- 
ease was considered likely to prove contagious. The dermatitis, 
however, was not lost sight of, and in a paper contributed to the 
Public Health Reports, Volume XXIV, No. 28, July 9, 1909, Dr. 
Joseph Goldberger, Past Assistant Surgeon of the United States 
Public Health and Marine Hospital Service, in cooperation with 
Dr. Schamberg, published the first exact information we have 
relative to the cause of these epidemics and this paper, so far as 
known to me, 1s the first publication in this country in which the 
attack of this mite has been followed up and its dermatological 
effect on humans carefully studied and described. This paper of 
Drs. Goldberger and Schamberg may be briefly summarized as 
follows: 

In the spring and summer of 1909, this peculiar eruptive dis- 
ease became quite prevalent in Philadelphia and neighboring 


26 Annals Entomological Society of America [Vol. III, 


towns. An outbreak among 20 sailors upon a private yacht 
docked in the Delaware River attracted the attention of both 
the city and the Federal Health Authorities. The Surgeon- 
General of the U. S. Public Health and Marine Hospital Service 
delegated Dr. Joseph Goldberger, Passed Assistant Surgeon, to 
proceed to Philadelphia in order to make an investigation of the 
disease. 

After examining the 20 sailors who had been sent to a hospi- 
tal, Drs. Goldberger and Schamberg visited the yacht whence 
they came and made a searching examination of the conditions 
on board. Their attention was directed to the fact that a num- 
ber of new straw mattresses had been received and that the dis- 
ase was confined to those who had slept upon these mattresses’ 
or had placed their clothes upon them. Eleven officers and mem- 
bers of the crew who did not sleep upon the new mattresses 
remained entirely free of the disease. 

At about the same period information was received concerning 
an eruptive disease prevailing among the sailors of four other 
boats, plying along the Delaware River. Investigation disclosed 
the fact that these boats had also received new straw mattresses, 
and, furthermore, that only those were attacked who slept upon 
the mattresses or otherwise came in contact with them. 

In addition to these cases among sailors, Doctors Goldberger 
and Schamberg examined or received authentic information con- 
cerning 70 other cases of this disease occurring in twenty different 
households in Philadelphia and its vicinity. Plate IV. 

In practically every instance they were enabled to determine 
that the patient had either recently slept upon a new straw mat- 
tress or had freely handled the same. Where only one person 
in a household was affected, it was found that he was the only one 
to occupy a bed supplied with a new straw mattress. They were 
able to trace all of the incriminated mattresses to four leading 
mattress manufacturers. | 

Careful investigation warranted them in excluding from con- 
sideration the ticking of the mattresses and the jute or cotton 
topping contained therein. The cause of the disease was, there- 
fore, circumscribed to the straw. Repeated inquiries elicited the 
information that all of the manufacturers had received at the 
time the disease-producing mattresses were made up, wheat- 
straw from a dealer in Salem County, in southern New Jersey. 


TQIO] Pediculoides Noxious to Man 27 


One manufacturer had used straw from this source exclusively 
in the affected mattresses. 

Finding of a Parasite. Drs. Goldberger and Schamberg sifted 
the straw from a mattress through the meshes of a fine flour sieve 
upon a large plate glass covered with white paper. Close scru- 
tiny of the siftings under strong electric illumination soon de- 
tected some slight motion. The moving particles were touched 
with a needle moistened in glycerine and transferred to a glass 
slide. Search with the microscope disclosed the presence of a mite 
of very minute dimensions. The mite was identified for them 
by Mr. Nathan Banks, expert in Acarina of the U. 5. Bureau of 
Entomology, as very close to, if not identical with, the Pedicu- 
loides ventricosus. 

In order to demonstrate experimentally the ethiological rela- 
tionship of the suspected straw mattresses, Dr. Goldberger ex- 
posed his bared left arm and shoulder for one hour between two 
mattresses. At the end of about 16 hours, a number of charac- 
teristic lesions appeared upon the arm, shoulder and chest. Later 
three volunteers slept upon the mattresses and each one devel- 
oped the eruption at the end of about the same period. 

Dr. Goldberger later took some of the sifted straw, divided it 
into two portions and placed it in two clean Petri glass dishes. 
One of these was applied for one hour to the left axilla of a volun- 
teer. At the end of from 16 to 17 hours, the characteristic erup- 
tion was present in the area of the left axilla to which the Petri 
dish of straw siftings had been applied. 

The second portion of the straw siftings in a Petri dish was 
exposed to the vapor of chloroform under a bell jar with a view 
to killing any insect or acarine that might be present. These sift- 
ings were then applied to the right axilla of the same volunteer 
to whose left axilla the untreated siftings had been applied. The 
chloroform evidently destroyed in the siftings the agent that 
was producing the eruption for no lesions appeared after the ap- 
plication of the chloroformized siftings. 

Dr. Goldberger further fished out of some straw siftings five 
minute mites, and, placing them in a clean watch crystal, applied 
the crystal to the axilla of another volunteer. At the end of 
about 16 hours following this application, five of the characteris- 
tic lesions appeared on the area to which the mites had been 
applied. Plate V, fig. 1. 


28 Annals Entomological Society of America [Vol. III, 


INFLUENCES CONTROLLING THE EXCESSIVE ABUNDANCE 
OF PEDICULOIDES. 

It will be noticed that Drs. Goldberger and Schamberg made no 
attempt to discover the underlying causes for the enormous num- 
bers of these mites inhabiting the mattresses involved in their 
investigations, that problem belonging more properly within the 
realm of the entomologist. On my taking up this subject of the 
excessive abundance of the mites, Drs. Goldberger and Scham- 
berg very kindly placed at my disposal everything in their pos- 
session relating to this epidemic, including the mattress which Dr. 
Goldberger had himself used in experiments carried out by him 
at the hygienic laboratory in Washington with this mite. Dr. 
Schamberg was equally kind in placing all of the material, notes 
and photographs in his possession, at my service. 

Almost at the commercement of the investigation, Dr. Wm. 
Royal Stokes of the Maryland State Board of Health informed 
me that a similar but less extensive epidemic had shortly before 
been noted in Baltimore, This he kindly described as follows: 

‘The matter was brought to my attention by several persons 
who came to the Head of the Department and complained of the 
skin eruption described. They stated that a number of people in 
a suburban hotel were similarly affected, but I do not remember 
the number at this late date. These persons volunteered the 
information that they had all been sleeping on some new straw 
mattresses, and that all of the persons similarly affected had used 
these mattresses. 

“T saw Dr. Gilchrist, the Clinical Professor of Dermatology 
at Johns Hopkins University, yesterday, and he gave me the fol- 
lowing description of the one case which he saw at the Health 
Department. I saw two other cases which corresponded with 
these in a general way. 

‘“*The eruption consisted of about 1000 wheals, or erythe- 
mato-withicarial spots, or pauplo-withicarial lesions. As in the 
description in the reprint of Drs. Goldberger and Schamberg of 
the United States Public Health and Marine Hospital Service, 
they varied in size from a lentil seed to a finger nail, and are 
round, oval, or irregular in shape. No vesicles or pustules were 
seen. The eruption was on the neck, chest, abdomen, and back, 
and also on arms and legs. Itching was present, and all lesions 
showed evidences of scratching.’ ”’ 


— aE 


TQIO| Pediculoides Noxious to Man 20 


Besides this, there were several cases reported to me from 
northern Maryland, where farmers in running their wheat 
through a fanning mill had been simultaneously troubled by a 
very similar or identical eruptive disease of the skin. In anoth- 
er instance, a thresherman engaged in feeding the unthreshed 
grain into the cylinder of the threshing machine was also af- 
fected by a disease of the skin, with which the attending physi- 
cian was unfamiliar and who could not classify it with any of the 
urticaroid dermatitis known to him. With my experience of 
previous years, it seemed impossible that this Pediculoides 
should become sufficiently abundant to cause this dermatitis 
without there being an excessive abundance of some host insect 
or insects affecting either the straw or the grain itself. Natur- 
ally, the studies made by me in 1882, led me to suspect that this 
grain moth Sztotroga cerealella might be responsible for the 
abundance of the mites. Then, too, the fact that it attacked 
the larvae of [sosoma grande in wheat straw, led me to suspect 
that, as this particular species is not known to occur in the vicin- 
ity of Philadelphia, while its near relative, the joint worm, 
Isosoma tritict, does occur more or less abundantly over the east- 
ern part of the country, this latter species, too, might perhaps 
be involved. 

With a view of finding out something of the abundance of 
the grain moth in New Jersey, from which State was obtained 
most but not all of the straw entering into the mattresses men- 
tioned by Drs. Goldberger and Schamberg, I applied to Dr. John 
B. Smith, State Entomologist, for information. In reply Dr. 
Smith was kind enough to send me an advance copy of the report 
of his department of the New Jersey Agricultural Experiment 
Station for the year 1908, and from this publication it was 
learned that during the summer of that year, owing to favorable 
weather conditions, this moth developed rapidly in the field 
and there was great damage to wheat among those farmers who 
delayed threshing until September or later. Furthermore, a 
very large percentage of the wheat crop gathered that year 
became useless for milling purposes and so general was the infes- 
tation that grain from some localities was entirely barred at 
mills except when ground for the owner. Some further investi- 
gations carried on in eastern Pennsylvania revealed a very simi- 
lar condition of affairs. It was the straw of 1908, coming mostly 
from New Jersey, but a small part of it from Indiana, that en- 


30 Annals Entomological Society of America  [Vol. III, 


tered into the mattresses, from the use of which came the epi- 
demic in and about Philadelphia. 

In order to settle these points, Mr. V. L. Wildermuth, an as- 
sistant in the Bureau of Entomology, in cereal and forage insect 
investigations, was instructed to examine the straw in the mat- 
tress placed at my disposal by Dr. Goldberger. After a day and 
a half of careful search, only five straws affected by the joint 
worm were found. This seemed to entirely eliminate this species 
from consideration in connection with this particular epidemic. 
There were, however, many wheat heads remaining attached 
to the straw-and these heads contained a great many kernels, the 
contents of which had been eaten out by the larvae of the grain 
moth. Morever, these eaten kernels contained great numbers 
of the dead bodies of Pediculoides. That Sztotroga cerealella 
was the cause of this damage to the wheat was still further proven 
by the emergence of an adult moth from these eaten kernels on 
November 15. The larvae of this moth infested the kernels of 
wheat before the latter were threshed. Many of these infested 
kernels remaining in the straw were included in the material 
going into the manufacture of these mattresses. The greater 
portion of the living larvae of the moth would develop to adults 
during May or early June, thus cutting off the food supply of these 
mites. The mites would therefore very naturally swarm among 
the straw and making their way through the cloth covering of 
these mattresses attack anything that gave promise of furnishing 
food, and preserving them from starvation. It seems that star- 
vation is the final outcome, however, for, as already stated, no 
trouble is experienced in using the mattresses after a certain per- 
iod, which period probably indicates the termination of the life 
of the mites infesting the straw. It therefore did not seem neces- 
sary to seek farther for the primary cause of this eastern epidemic 
of dermatitis, the center of which seems to have been in and about 
Philadelphia. 


A WESTERN EPIDEMIC OF THE DERMATITIS. 


While the problem of the epidemic in the east was apparently 
solved, some of the wheat straw involved therein had come from 
Indiana, and during the last few years an outbreak of the joint 
worm, /sosoma tritici, had been gathering force, throughout Ohio, 
Indiana and southern Illinois, until during the summer of 1908, 
very serious damages occurred. Investigation of the insect dur- 


_— OO —————————— 


TgIoO| Pediculoides Noxious to Man 31 


ing previous years had shown that the outbreak really began in 
the more elevated portions of Virginia, in the upper Shenandoah 
Valley, West Virginia and eastern Ohio, as early as 1904; after- 
wards advancing broadly to the westward. | 

During the summer of 1908, there came to the Bureau, from 
this section of the country, a great number of complaints of serious 
skin trouble among people engaged in threshing grain that had 
been stored for some time in barns; in some localities it having 


. become difficult to secure help to thresh under such conditions. 


Also the same disorder was encountered by those who used this 
straw for the purpose of filling bed-ticks, or as a substitute for 
felting under carpets, and in one case, berry pickers had been 
attacked when such straw had been used as a mulch for berry 
plants. This straw came from one of the fields most seriously 
injured by joint worm attacks in 1908. In one instance, a car- 
load of wheat straw was shipped to Pittsburg, Pa., and six men 
engaged in unloading it, were all attacked by some skin eruption, 
and the horses used in hauling this straw after it was unloaded, 
also suffered from what was seemingly the same disorder. Per- 
haps the following from a correspondent of the Bureau of Ento- 
mology, residing in southern Ohio, will give a fair idea of the 
situation on many farms in that section of the country. 

‘“ About four years ago a parasite was found when threshing 
wheat out of barns. They seem to affect the victims almost as 
soon as they get into the mow. The men began to scratch them- 
selves generally on the neck and on the arms (inside) opposite the 
elbow, and on the body back and front. They raised welts as you 
describe and progress about as you describe. They have spread to 
such an extent that farm hands dread and fear them and will not 
change work with neighbors unless they thresh in the field. 
(They are found out of barns.) Here of late they are found in 
wheat straw in the barns, especially if baled. Last week a farmer 
brought me baled wheat straw that seemed to be alive with them. 
They attacked every one that went in the barn and one of my 
horses that was perspiring from effects of a drive was simply 
covered with little knots or swollen places and bit and rubbed him- 
self continually. I had to have the straw hauled out and burned 
and barn disinfected. The farmer stated that they were so thick 
in the shed that contained the straw, that he had to keep all 
stock out of the shed.” 


tN 


Annals Entomological Society of America _[Vol. III, 


ios) 


Many other similar letters from towns in Ohio were received 
by Dr. Schamberg, particularly from Zanesville, Columbus, Vin- 
cent, Springfield, etc., where the affection is popularly believed 
to be due to ‘‘chiggers.’’ A physician from the last named town 
stated that in the fall of 1908, during harvest and threshing time 
he saw in Washington County some 87 cases of the disease in 
question. It affected the harvesters and threshers. ‘This spring 
he observed 38 cases from contact with straw ticks refilled with 
straw of last fall’s crop. The disease is said to have been more 
prevalent last year (1908) than ever before. Information has 
come from Columbus, Ohio, that potters who used straw for 
packing crockery ware have been so badly attacked at times that 
the entire force of packers has been off duty. Many times a 
whole car-load of straw has been so affected that the use of it 
has been abandoned. In Springfield, Ohio, it is said that the 
disease was so bad a year or two ago in the lowlands west of this 
city as seriously to hamper the progress of the construction of a 
large sewer. This, however, might have been due to attack 
by other mites, notably to Trombidium larvae. In Zanesville, 
Ohio, the potters have been obliged to abandon the use of straw 
and employ ‘“‘prairie hay”’ for packing purposes. 

Dr. Schamberg was also informed by a physician of Pittsburg 
that a young woman patient has suffered from an affection closely 
resembling, if not identical, with the one under consideration, 
each time that she has assisted in emptying cases of dishes packed 
in straw. Both the physician and the patient had come to believe 
that something in the straw was the cause of the eruption. 

Indeed, so nearly did the territory from which these com- 
plaints came to us, coincide with that affected by the joint worm, 
that it created the suspicion, not only among those engaged in the 
investigations, but even among farmers themselves, that there 
must be some connection between the two phenomena. Very 
many of these cases were brought to the notice of practicing physi- 
cians, but they were themselves at a loss to account for the preva- 
lence of this dermatitis, many of them supposing it to be some 
species of rash that was more or less contagious, the exact nature 
of which they did not know. 

Among these physicians was Dr. Lyman T. Rawles, of Hunter- 
town, Indiana, who in May, 1909, took up a careful study of a 
number of cases of this dermatitis that had come under his per- 
sonal observation as well as those of some of his associates. Dr. 


- = 


7 iil 


TQIo| Pediculoides Noxious to Man 33 
Rawles’ investigations were very carefully made and the results 
are exceedingly valuable for the reason that, in case of this western 
epidemic, he was able to trace the cause of the dermatitis to 
Pediculoides ventricosus and follow this back to the host insect 
Tsosoma tritici. This paper of Dr. Rawles’, of which a summary 
is given, not only clears up the obscurity with reference to the 
cause of this epidemic in the Middle West, a section throughout 
which the grain moth (Sztotroga cerealella) never occurs in exces- 
sive abundance excepting in grain that is kept in store, and 
then only in the more southern portions of Indiana and Illinois; 
but, furthermore, these studies seem to solve the problem of the 
cause of the skin eruption noted by Dr. Harris to have occurred as 
far back as 1830. 

In May, 1909, Dr. Rawles found in his practice that a very 
strikingly strange skin disease presented itself in his and sur- 
rounding country in epidemic form. Through the press notes it 
seemed to be quite general over the northern part of the United 
States, limiting itself to the wheat growing sections. 

The people generally affected were farmers and those living 
in small villages or towns where straw is used in beds, under car- 
pets and around stables to bed stock. Horses and cattle have 
been seen with a skin disease almost identical to that seen in man. 
The following incident led him to an investigation as to the prob- 
able etiology: 

A family had cleaned house, refilling the straw ticks of their 
beds and placed fresh straw under the carpets, and in about one 
week the family had developed this peculiar skin disease. In the 
beds were found a small, black fly, [sosoma tritict Fitch, about 
the size of an ordinary gnat, which at first it appeared to be, but 
closer observation revealed that it was not of the gnat family. 
Upon examination of the straw it was found that a large number 
of the straws were perforated; these perforations were through 
the wall in the region of the joint, generally about two inches 
from the joint. The perforations were about the size of a small 
pinhole and ranging in number from ten to thirty in a straw. 
Upon examining a section of this straw the small black fly was 
found under many of the openings through the walls. 

Several flies were examined to ascertain if they possessed a 
piercing proboscus, and while observing one which has just been 


taken from under the sheath of the straw, through which there 


13. Journal of the Indiana State Medical Association, August, 1909. 


34 Annals Entomological Society of America  [Vol. III, 


was no perforation over the fly, a small mite was observed crawl- 
ing over the dead body of the fly. 

Placing the bodies of several of these flies under the micro- 
scope and using a }-inch objective and a No. 5 eye-piece, it was 
found that on nearly all flies over which the wall was intact, a 
small mite could be detected, these mites varying in number from 
two to four mites to each fly. Upon furthering the observations 
it was found that the dermatitis lasted after the flies had been 
observed and exterminated. 

The following experiments were tried to prove whether it 
might be the fly or the mite that was the etiologic factor in pro- 
ducing the dermatitis. 

Six live flies were taken, upon which no mites could be found; 
these were placed under a watch glass and bound upon the right 
arm, leaving them in contact with the skin for three hours. Upon 
the left arm four dead flies, on which living mites had been ob- 
served, were placed under a watch glass and left in contact with 
the skin for three hours, after which the glasses were removed 
and results awaited. The right arm showed nothing. Upon 
the left arm there appeared within twelve hours four small 
wheals, the character and evolution of which are later described. 

To further the experiments some fresh lesions of patients were 
scraped and the scrapings examined microscopically and two of 
the mites were found in the scrapings. 

Itching is the most prevalent and first symptom to attract 
the attention of the patient. It is most persistent and intense 
during the after part of the night. At about the time the itch- 
ing was most intense there appeared an urticarial eruption, 
accompanied, in severe cases, with general systemic symptoms, 
such as rise of temperature from gg to 102; in one case the tem- 
perature rose to 103.8; the pulse rate is accelerated to 100, or as 
high as rr0o—in one case to 130. Other symptoms were intense 
headache, anorexia, nausea, in some cases vomiting, and a mild 
form of diarrhoea. In severe cases some complained of general 
joint pains and backache; in these cases the urine was examined 
and albumin in small amount was found, but no casts or blood. 
When the acute symptoms disappeared so did the albumin. 

Many patients who suffered from mild cases complained of 
nothing aside from the intense itching. If all straw was removed 
from the beds and house the symptoms would subside in one or 
two days and completely disappear in a few days more. 


Tgto| Pediculoides Noxious to Man a5 


The lesion which is typical of the disease is the urticaria vest- 
culosa. The urticarial, lesion varies in size from that of a 
split pea to that of a penny; it is surrounded by a pinkish halo, 
varying in intensity of color from a pale pink to a most bright 
pink. The “hive” like lesion is at first blanched, but later 
becomes a rose red color. It is elevated about 1 or 2 mm. above 
the skin surface, and is surmounted by a small vesicle containing 
a whitish fluid marking the place of-inoculation. |The vesicle 
is about 1 or 2 mm. in diameter and elevated about 3 mm. above 
the surface of the urticarial lesion. As the lesion grows old it 
goes through the process of evolution: (1) it 1s blanched and has 
a central vesicle; (2) it is rose red and the vesicle may become a 
pustule; (3) it generally recedes to the skin level with scab for- 
mation, due to the scratching; (4) it leaves a browinsh or green- 
ish-yellow or purple spot on the skin surface. In debilitated 
patients the markings look not unlike faded indelible pencil 
marks. (This was noted in a patient suffering from pulmonary 
tuberculosis.) These discolorations may last for several weeks. 

The anatomical location of the lesions is generally the back, 
sides and abdomen, and less frequently the arms and legs. The 
neck has very few lesions; the face, hands and feet have very 
few or none. 

The number of lesions depends upon the number of mites, 
ranging in number from very few to thousands; in some cases the 
back and abdomen have been almost a solid mass of lesions— 
new lesions on the tops of old lesions, so having lesions in all 
stages of development. 


OBSCURITY SURROUNDING THE OCCURRENCE OF THE 
DERMATITIS. 


The exact nature of this eruptive disease was not at all under- 
stood by the medical profession throughout the country. In 
South-western Virginia, threshermen suffered from the same dis- 
order, but attributed it to- ‘‘chiggers,’ and local physicians, 
though skeptical, were themselves unable to correctly diagnosis 
or to account for the trouble. As the disease is not serious and 
passes away in the course of time without leaving the patient in 
any way permanently injured, it seems to have been passed over 
by medical men without investigation excepting by the physi- 
cians whose publications have just been cited. Among the people 
themselves the eruption was probably more frequently attributed 


36 Annals Entomological Society of America  [Vol. III, 


to attacks of ‘‘chiggers’’ than to any other cause and it is quite 
likely that this common erroneous interpretation of the origin of 
the eruption has prevailed generally throughout the country, 
including the upper Shanandoah Valley in Virginia, where the 
. joint worm was abundant as far back as 1904. It has been con- 
fused with small-pox and more frequently with chicken-pox. It, 
was, consequently, exceedingly unfortunate that, with the begin- 
ning of this disorder, an institution in one of the States involved, 
should publish an unsigned newspaper bulletin, crediting these 
epidemics of this eruption to the attack of ‘‘chiggers,’’ and, further- 
more, at the very time when Drs. Goldberger, Schamberg and 
Rawles, as well as the Bureau of Entomology, were exerting every 
effort to find out the true cause of the difficulty, that a second press 
bulletin, accentuating the first, should have been issued, and, sent 
to every newspaper in the State, and from those copied into other 
newspapers throughout the country. Thus it is that an entirely 
erroneous impression has been magnified and diffused, still con- 
tinuing to prevail throughout the country. 

In order to determine the likelihood of those handling straws 
in the wheat field, being attacked by the small red mites, often 
innocently mistaken for ‘‘chiggers,’’ that abound among the har- 
vested grain at this time, Mr. Wildermuth made a number of 
experiments to determine whether or not these mites, probably 
Tydius sp., were liable to attack men. In no case was he able 
to provoke an attack from them, even when they were confined 
upon the skin of his bare arm. On the other hand, examination 
of straws from various points in Ohio and Indiana have revealed 
the presence of Pediculoides in the cells occupied by the joint 
worm. This seems to entirely eliminate ‘‘chiggers’’ from these 
investigations, because these were probably not present, and there 
does not longer appear to be any doubt but what Pedzculoides 
ventricosus is to be charged with causing these epidemics of this 
. dermatitis, and the cause of its own excessive abundance lies in 
the outbreaks of the Angoumois grain moth among the grain 
itself in the East and the joint worm in the wheat straw in the 
Middle West. 


Licut THROWN Upon OTHER PROBLEMS. 


These investigations have illustrated very nicely the extent 
to which the solution of one entomological problem will at the 
same time also solve other problems more or less closely allied to 


- 
EEE 


Tg1o] Pediculoides Noxious to Man 37 


the original one. The light thrown upon the cases of eruption 
noted by Dr. Harris with reference to Jsosoma hordei has already 
been explained. The present outbreak of the joint worm in the 
Ohio Valley probably originated in the upper Shenandoah Valley 
of Virginia, extending northward and westward throughout West 
Virginia and eastern Ohio. When investigation of the insect was 
taken up in 1904, a parasite, Ditropinotus aureoviridis, was 
also noted in excessive abundance, but for some reason it did 
not overcome the joint worm. This phenomenon has been noted 
continually. Since that time it has been a perpetual enigma to me 
why it was that with such an abundance of its natural enemies 
the joint worm should continue to spread and increase in destruc- 
tiveness. Now, however, that we know that this predaceous mite 
is able to develop thr ouelt a series of years in such immense num- 
bers in connection with the joint worm, the matter comes nearer 
a solution. Dutropinotus, as well as some other parasitic enemies 
of the joint worm, emerge in early July from eggs that were pre- 
viously placed in the cells occupied by the joint worm. As soon 
as these adult parasites emerge they at once oviposit in cells con- 
taining joint worm larvae of the same generation from which they 
themselves developed. The puncturing of these cells by the 
Ovipositor of these parasites, particularly Ditropinotus, opens a 
way for the entrance of this microscopic mite, and, once inside of 
the cell, it will destroy everything therein, whether it be joint 
worm or parasite. Thus the predaceous mite has prevented the 
other parasites from exerting their full influence, because it has 
continually checked the increase of other parasites, thereby pre- 
venting them from increasing and exerting the restraining influ- 
ence upon the joint worm that, but for this mite, they probably 
would have done. 

In the light of the foregoing, it would appear that the only 
way to ee, this disorder among humans, caused by this mite, 
lies the prevention of the occurrence of these two destructive 
grain insects which are responsible for the abundance of the mite 
itself. There is, therefore, a double incentive for the farmer to 
use every effort to prevent the occurrence of these pests in his 
fields. In many fields in Ohio we have found that more than 
one-half of the straws had been attacked by joint worm, and the 
damage resulting from their attacks amounted to a considerable 
percentage of the farmers wheat crop. See Plate V, fig. 2. 
If, in addition to this, his own family and employees are to suffer 


38 Annals Entomological Society of America  [Vol. III, 


annoyance from this dermatitis, and we know that the mite is 
present generally, also those who attempt to use mattresses into 
the construction of which the straws from these fields have entered 
and indeed in even handling the same, the results of his agricul- 
tural methods will be sent wherever, in the entire country, these 
mattresses, or the straw itself, go into practical use within at least 
a year from the time the wheat straw was harvested in the fields, 
and thus people hundreds of miles away, with no possible means 
of knowing of the presence of these mites in mattresses, are caused 
not only great aggravation but intense suffering through their use. 


PREVENTIVE AND PROTECTIVE MEASURES. 


From the foregoing, it will be observed that that public pro- 
tection from this skin disorder is only to be secured through 
revised agricultural methods of the farmer, who, while offering 
this protection, will also materially increase the profits of: his 
business. Throughout the territory involved in the eastern 
epidemic of this dermatitis, which, as has been shown, was due 
to the excessive abundance of the Angoumois grain moth, the 
evidence recently attained by the writer has been overwhelmingly 
to the effect that where wheat was threshed as promptly as pos- 
sible after harvest and directly from the shocks in the field, almost 
no occurrence of this grain moth was observed by millers and 
others handling the threshed grain, and without which there 
would have been no mites. On the other hand, when drawn from 
the field and placed unthreshed in the barn, the damage from this 
pest has varied up to nearly fifty per cent., and has so affected 
the crop as to cause its rejection by millers, except where ground 
on the farmer’s order. Here, then, lies the protection of people 
who use mattresses made of this wheat straw, grown in this sec- 
tion of the country, or otherwise come in contact with the same. 
Reiterating in a condensed statement: Wheat should be 
threshed immediately after harvest and directly from the field. 

In Ohio, Indiana and Illinois, where the mite causing this der- 
matitis has increased enormously on account of the prevalence 
of the joint worm, see Plate V, fig. 3, wheat also placed in barn 
‘ before threshing has been found to be much more. dangerous to 
handle with reference to epidemics of this disorder. At the 
same time, the difference between wheat threshed in the field and 
in the barn is not so striking as where the primary trouble was 
in the abundance of the grain moth. 

A careful study of a large number of wheat fields in central 
Ohio, has shown that the infestation from joint worm, the present 


ANNALS E. S. A. 


VoL. IlI, PLateE III. 


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SCREENS 


Se er ere eet 


Webster on Pediculoides 


III, PLATE IV. 


VOL. 


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ANNALS 


Webster on Pediculoides 


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Tgto| Pedtculoides Noxious to Man 39 


season, has varied from one to ninety-five per cent. Here, too, 
we have found the mite generally in the cells in the straw occupied 
by the joint worm larvae. It has been found that in central Ohio, 
September sown wheat was much more seriously affected by the 
joint worm than that sown in October, and also that the infesta- 
tion 1s worse in both cases on poor soil than on that of an average 
degree of fertility, and still less on good soil. The infestation was 
invariably worse in fields lying adjacent to or where wheat had 
been grown the previous year. Another interesting fact was 
revealed, and that is that fall plowed fields showed the least 
infestation of all. Therefore, it appears that moderately late 
sown wheat on good soil and on land not devoted to wheat the 
previous year, nor lying adjacent to such fields, has escaped with 
the least injury, and that less difficulty with the dermatitis is 
experienced where such wheat has been threshed from the field 
and as soon as possible after the grain was harvested. As the 
joint worm winters over in the stubble, where this can be burned 
during the fall, winter or spring, the destruction of the pest in the 
field will be complete. Where this cannot be done, much good 
may be accomplished by raking over in the spring the last year’s 
stubble fields and burning the stubble thus collected. So impor- 
tant are these measures that practicing physicians might almost 
include them with their prescriptions for this painful skin disorder. 


EXPLANATION OF PLATES. 


lesoyNanioy JB. 

Fig.1. Pediculotdes ventricosus Newport, ungravid 9. Stage in which the 
mite is both migratory and predatory and in which it attacks humans. From 
photograph by Dr. Jay F. Schamberg. Greatly magnified. 

Fig. 2. The Angoumois grain moth, Sztotroga cerealella Oliv., a, eggs en- 
larged; 6b. b., bernels of wheat infested by larvae; c, larva, enlarged; d, pupa; 
e, moth with wings spread; 7, same with wings closed as at rest. Figs. b.b., ori- 
ginal remainder, after Chittenden, Farmers’ Bulletin, U. S. Dept. Agriculture, 
No. 45, p. 6. ; 

Fig. 3. Pediculoides ventricosus, gravid 9. Greatly enlarged. Redrawn 
from Brucker. 

PLATE ENE 

Illustrating various forms of the dermatitis lesions caused by attacks of 

Pediculotdes ventricosus. From Photographs by Doctor Schamberg. 
PLATE V. 

Fig. 1. Lesions caused by bites of Pediculoides ventricosus, in experiment 
of Dr. Joseph Goldberger. From drawing by F. H. Wilder. Courtesy of Public 
Health and Marine Hospital Service. 

Fig. 2. Showing reduction in yield of wheat attacked by jointworm, 
Isosoma tritict Fitch. The tube at left contains yield from 100 heads from unin- 
fested straws: tube at right contains yield from 100 heads from infested straws. 
Photographed by W. J. Phillips, Bureau of Entomology. 

Fig. 3. Showing attack of joint-worm, /sosoma tritict, in field. Note the 
enlarged and distorted stems. From photograph by Geo. I. Reeves, Bureau 
of Entomology. 


MYRMELEONIDZ FROM AUSTRALIA. 
By NATHAN BANKS. 


The ant-lion flies of Australia have been mostly described by 
Walker and Gerstaecker. In collections sent to me by Mr. Dodd, 
from Kuranda and Port Darwin, and by Mr. Perkins, from several 
parts of Queensland, are a number of the described species and 
several that are new. These are described in this paper. 

The Australian Myrmeleondiz are more like the European 
forms than like those of the United States. Two genera, Myrme- 
leon and Acanthaclisis, they have that occur also with us, but these 
genera are generally distributed throughout the world. The new 
general are more allied to the European than to our genera. 
There is practically no similarity to the South African forms. 

The eight genera known to me can be tabulated as follows: 


1. No-spurs, one cross-vein, before radial sector in hind wings; wings broad 
eielibiay 
Spurs present... .- 6... ee ee ee eee eee eee t eens 
2. One cross-vein before radial sector in hind WRUDNES Soyo s0 Soh ks BR ree 
Several cross-veins before radial sector in hind wings; anal ends before fork 
of radial sector:pronotum broader than long.) .y 22, saeau arene 3 
In middle of apical half of each wing is a nearly straight line formed by the 
bent longitudinal veins and very short cross-veins; anal vein of “hind 
wings runs into lower cubitus, legs short and stout. .Acanthaclisis 


5 


ws 


No such line of bent veins and short cross-veins.............0-0e000. + 
4. In both wings the upper cubitus and the median unite long before margin 
of witless “Wines: wery vbnoddis;)-. nner en tek eee Callistoleon 


Upper cubitus and median not united, wings more narrow. .Myrmeleon 

5. Basal joint of tarsi longer than apical joint; anal ends far out on wings, 
beyond origin of first fork of radial sector............. Protoplectron 

Basal joint of tarsi not longer than apical; anal vein ends nearer base of 
wing, at or before origin of first fork of the radial sector.......... 6 

6. Spurs as long as first four tarsal joints: together.s <4. 5,cense Distoleon 
Spurs barely longer than first joint of fATSUS ik. 1 eh he ee eee 7 
Wings broad in stigmal area, in middle of this area is a straight line as in 
Acanthaclisis formed by bent longitudinal veins and very short cross- 
(a er Mn Ah NTC h ety Ein ers | 3 Glenurus 
Wings more narrow; no such line formed by bent veins and short cross-veins 
Formicaleo 


~I 


Glenurus pulchellus Ramb. 
Mid-Queensland (Perkins) also New South Wales (Froggatt). 


Glenurus dissolutus Gerst. 
Port Darwin, 12 May (Dodd). 


Glenurus stigmatus, n. sp. 

Face pale, a large interantennal black spot, a transverse pale band 
above this, and the vertex brown; antenne black; pronotum with a 
broad black median stripe, containing a pale median stripe in fore 
part, sides pale, streaked and dotted with black; rest of thorax black, 
with a few small pale spots on the lobes; abdomen black; legs black, 
tibia I and II with basal and preapical bands pale, sometimes also a 
median spot above; hind tibiz pale behind, except tip, and in front at 


40 


Tgto| Myrmeleomde from Australia Al 


base and tip pale; base of first tarsal joint pale; legs with short black 
hair and longer, erect bristles; spurs rather longer than first tarsal 
joint. Wings hyaline, with many small dots, mostly on forkings of 
veins, and in apical part, larger ones along radius and cubitus, forming 
rather oblique lines at ends of anal veins, and on gradate series in 
fore-wings; stigma a large, black spot in both wings, very prominent; 
wings of usual shape and much like G. dissolutus. 


Expanse 80 mm., abdomen long 20 mm. From Kuranda, 
Queensland, Australia, March, (Dodd). > 


Protoplectron costatus n. sp. 

Face pale, black in antennal region, vertex pale, a transverse 
black line, and behind it a row of spots, a small trapeze in the middle 
and two spots each side toward eye; antenne pale brown, short, with 
large knob; pronotum brown, darkest behind on sides, paler in middle, 
with some long white hair, and also on the pleura; abdomen brown, 
with extremely short white hair; legs pale, femora dark above, tibize 
black at base and tip, tarsi black in middle, pale on most of first and 
fifth joints, with much long white hair, and some black bristles; 
tarsal joint I much longer than IV, one-half as long as tibia, spurs as 
long as first joint. Wings hyaline, narrow, acute at tip, and fore 
wings rather falcate; venation mostly dark, but some patches of cross- 
veins wholly yellow, and long streaks of pale in the longitudinal veins; 
stigma brown at base. In fore wings the lower cubitus runs parallel 
to anal vein for a long distance; but one series of costal cells in either 
wing; eight branches of radial sector before stigma in fore-wing. 


Expanse 48 mm., abdomen 18 mm. 
From Port Darwin, Australia, 10 April (Dodd). 


Protoplectron pallidum n. sp. 

Lower part of face yellowish, upper part and the front black; 
vertex with a broad transverse pale band from eye to eye, divided on 
the median line; antennze yellowish brown, the second joint with a 
darker ring; pronotum pale yellowish; thorax dark brown on sides, 
pale through the middle, and a pale spot above base of each wing; 
pleura mostly pale or hght brown; abdomen black, legs pale, rather 
reddish on femora and tarsi, no bands or marks, clothed with long 
white hair and black bristles. Antenne rather longer than head plus 
thorax; pronotum once and a fourth longer than broad, only slightly 
narrowed in front; abdomen short, not near as long as wings. Legs 
rather short; tarsi very slender, basal joint more than one half as long 
as tibia, the spurs of the same length, joints 2, 3, and 4 very short, 
5th joint more than one-half as long as the first, on its basal part 
beneath are several short, curved spines forming a comb. Wings 
yellowish hyaline, unspotted; stigma scarcely visible; venation 
yellowish, or brownish yellow, not dotted. Wings moderately 
slender, acute at tips, a double costal series of cells in fore wing, single 
series in hind wing; about 7 cross-veins before origin of radial sector 
in fore wing, only one in hind wing; in fore wing the first branch of 


42 Annals Entomological Society of America  [Vol. III, 


the radial sector arises long before end of the anal vein, which is nearly 
one-half way out on the hind margin; apical portions of both wings 
have the longitudinal veins very close together. 


Expanse 58 to 68 mm., abdomen 20 mm. 
Several specimens from Port Darwin, Australia, 8 April to 4 
May, (Dodd). 


Callistoleon n. gen. 


Wings rather broad; anal veins of fore-wings ending long before 
origin of the first branch of the radial sector; posterior branch of 
cubitus running obliquely down toward anal vein, two series of cells 
between it and anal vein; about seven cross-veins before origin of 
radial sector in fore-wings, some crossed; a single series of costal cells; 
the anterior branch of cubitus and the median vein unite before tip, 
and from the anastomosis the veins radiate somewhat, to the margin, 
more strongly so in the hind wings. In the hind wings about four 
cross-veins before radial sector; the -posterior branch of cubitus 
running obliquely down to margin, before origin of radial sector; head 
rather broad; antenne moderately long, pronotum broader than long; 
tarsal joints short, first much shorter than fifth; spurs a little longer 
than basal joint. 


Type—Myrmeleon erythrocephalum Leach. 


Callistoleon erythrocephalum Leach. 
Mid-Queensland (Perkins). 


Distoleon n. gen. 


Wings rather narrow; seven cross-veins before radial sector in 
fore wings, one in hind wings; anal ends before origin of first branch 
of radial sector; a single costal series. In hind wings anal ends soon 
after origin of lower cubitus, not parallel thereto. Spurs as long as 
first four tarsal joints, fifth joint much longer than first; tarsus I about 
as long as tibia. 1; pronotum broad. 


Type—D. verticalis; also includes Formicaleo bistrigatus. 


Distoleon verticalis n. sp. 

Head pale, a black band below and one above base of antenne, 
another band above this, narrowly interrupted in the middle or 
broken into four spots, and on the vertex four spots forming a trapeze 
in the middle and two each side toward the eye; antenne brown, 
annulate with pale; pronotum pale, darker on margin, rest of thorax 
grayish brown, abdomen blackish, a pale spot above in middle of 
each segment, legs pale, some black dots at bases of bristles, black 
and white bristles. Wings hyaline, a dark dot on stigma, one on hind 
margin one-third the way out at end of anal vein, and two at end of 
cubitus and median veins, and other smaller spots in apical part of 
wing, and at forkings of longitudinal veins, these veins black and 
white in long streaks, some cross-veins all black, others all yellow, 
others black and yellow. Hind wings with two dots at end of cubitus 
and less distinctly on apical portion. Antenne long; legs short, spurs. 


Igo] Myrmeleomde from Australia 43 
heavy, curved, as long as first four joints, fifth joint twice as long as 
first. Pronotum broader than long, not narrowed in front. Wings 
rather long acute; one costal series, cells in region of gradate series 
are long and narrow. 


Expanse 68 mm., abdomen long 25 mm. 
Mid-Queensland, Australia, (Perkins). 


Distoleon bistrigatus Ramb. 
(M. striola Walk.) 
Port Darwin 20 April, 28 August; Kuranda, March; Mid- 
Queensland. 
Chrysoleon n. gen. 


No spurs; legs slender, first tarsal joint as long as second plus 
third, hardly as long as the fifth; pronotum slender. Fore wings 
broad, hind wings narrower and longer than fore wings; costals of 
fore wings frequently forked; three cross-veins before radial sector; 
anal ends a httle beyond origin of first branch of medial sector; a 
double series of anal cells; the lower cubitus running obliquely down 
to the hind margin. In hind wings one cross-vein before radial 
sector; anal ending beyond origin of first branch of radial sector; 
lower cubitus running down to margin, upper cubitus uniting with 
median before tip of wing. 


Chrysoleon punctatum n. sp. 

Head yellow, a faint dark mark each side on vertex; antenne 
yellowish, darker at tip; pronotum pale, with some scattered brown 
dots; rest of thorax mostly pale; abdomen pale, a black stripe each 
side; legs pale, minute brown dots at base of bristles, not distinct on 
hind tibie. Wings hyaline, outer apical part of fore wing with small 
brown clouds at forks of veins, and one under the stigma, also in hind 
wing, but fewer in the posterior apical part. Venation pale, frequently 
dotted with brown in fore wings, especially along cubitus. Antenne 
short, not reaching beyond middle of thorax; pronotum slender, 
narrowed in front, this and the rest of thorax with long white hairs, 
abdomen with short white hair; legs with some long white hairs and 
black bristles. Fore wings broad below stigma, barely acute at tips, 
cells of wing small and irregular; radial sector seven branched before 
stigma; costal area broad, the cells narrow. 


Expanse forewings 43 mm., hind wings 47 mm., abdomen 
long 13 mm. 
From Port Darwin, Australia 20 April (Dodd.) 


Formicaleo dispersus n. sp. 

On the same general plan as F. septus and F. bistrigatus, but no 
dark stripes in front wings instead of which are about twenty small 
dark spots in the area of the stripe of F. septus toward the hind 
margin, and in hind wings is a stripe as in F. septus but its apical 
part broken up into small spots. The head is yellow, a large black 
spot on front and upper face; antenne pale on basal part, brown 


44 Annals Entomological Society of America  [Vol. III, 


toward tip; pronotum pale yellowish as also thorax, abdomen rather 
darker on sides and near tip. Legs pale, unmarked, black bristles; 
spurs as long as joint I, 5th joint as long as rest together. Wings of 
general shape of F. septus, and similar in venation, seven cross-veins 
before origin of radial sector in fore wings, one in hind wings; from end 
of anal vein there is no vein bending up and outward as in F. bistrigatus, 
in this respect like Ff. septus. 

Expanse 54 mm.; abdomen long 17 mm. 

From Port Darwin, Australia, 13 March (Dodd). 


Formicaléo marginalis n. sp. 

Pale yellowish; a broad black stripe through antennal region 
from eye to eye; vertex with two black submedian spots, and a median 
spot behind them; antenne pale brown; pronotum with two dark 
spots on front margin and a stripe behind each of them to the hind 
border; thorax striped with dark, a submedian pair in front reaching 
to middle of mesothorax, a lateral one just above wing, and a short 
pair on metathorax, and a dark spot over base of wings; pleura with 
some black spots, mostly on lower parts; abdomen blackish, with 
short white hair; legs pale, with black bristles, and short black hair. 
Wings unmarked, except at posterior margin near tip of both pairs 
where the margin is infuscate for about one-third the way from tip 
to base; venation yellow, subcosta interruptedly black in both wings, 
and some dark dots on cubitus in fore wings. Venation similar to 
F. bistrigatus, a vein bending upward and outward from end of anal 
vein in fore wings, seven cross-veins before radial sector in fore wings, 
one in hind wings. Pronotum rather slender, narrowed in front; legs 
short, tarsal joint I about as long as II plus III, much shorter than V, 
spurs a little longer than first joint. 


Expanse 43 mm., abdomen long 13 mm. 

From Kuranda, Queensland, Australia, March, (Dodd). 
Formicaleo septus Gerst. 

Port Darwin, latter part March through April; also at Kuran- 
da, March. 
Myrmeleon pictifrons Gerst. 

Port Darwin; common, 10 to 21 April; and also 3 September; 
Mid-Queensland (Perkins). 
Myrmeleon uniseriatus Gerst. 

Port Darwin, 9 September; Mid-Queensland. 
Myrmeleon croceicollis Gerst. 

Kuranda, March. 
Acanthaclisis fundatus Walk. 

Kuranda, March (Dodd); Mid-Queensland, (Perkins). 
Acanthaclisis subtendeus Walk. 

From Kuranda, March (Dodd). 


THE ANATOMY OF THE LARVA OF CECIDOMYIA 
RESINICOLOIDES WILLIAMS. 


By Francis X. WILLIAMS. 


In the January number of Entomological News, Vol. XX, 1-8, 
1909, I described this midge in its egg, mature larval, pupal and 
imaginal instars, and gave a brief biological report on it. The 
curious habit of the larva in living in a mass of exuding resin 
might be expected to have brought about some adaptive modi- 
fications of its structure, especially perhaps of its tracheal system. 
It is for this reason particularly that the attempt has been made 
to study the larval structure in detail. As much of the anatomy 
of the larva is described here as could be worked out in a limited 
time, and although over a hundred larvae were examined and 
dissected, the results are far from complete. Considerable diffi- 
culty was encountered in tracing out the various systems of the 
larva, and the writer has deemed it best to omit the doubtful 
things and to declare only those facts which were made manifest 
over and over again by the examination of a large number of 
specimens. 

The work embodied in this paper was done in the entomological 
laboratory of Stanford University. 


EXTERNAL ANATOMY OF THE LARVA: 


ead. PL i, Figs. © and! 2.) The headmise very smalltand 
imperfectly developed. Its brownish-black chitinized portion 
consists of a broad irregular ring from which a pair of prong-like 
rods project well into the supernumerary segment. These rods 
diverge posteriorly, are more heavily chitinized than the ring, 
and have muscles attached to them that run from the body wall. 
The antennae are short, conical, obscurely two-jointed processes 
placed well above the opening of the mouth. Ratzeburg and 
L. Defour believe these processes are palpi, but Osten Sacken as 
well as Laboulbene and Perris, from the position of these organs, 
consider them rudimentary antennae. I incline to the latter 
view since they are situated latero-dorsally on the anterior part 
of the ring and well above the mouth opening. This latter con- 
sists of a soft, fleshy swelling taken by Ratzeburg for the labium. 

Breastbone, Pl. I, Fig. 3. Ventrally on the posterior half 
of segment I, and situated in a sort of fold is the ‘‘breastbone,”’ 


45 


46 Annals Entomological Soctety of America  [Vol. III, 


also known as the ‘“‘sternal spatula”’ or ‘‘anchor process,” an 
organ said to be peculiar to the Cecidomyidae. This is a brownish 
chitinized process varying somewhat in shape, but usually widest 
at the extremities, slightly constricted at or near its middle and 
at the caudal end, and somewhat incised anteriorly where a small 
portion of it projects beyond a transverse fold. The caudal 
extremity is concealed under the posterior fold of segment I. 
The anterior incised portion of the spatula is somewhat scooped 
out, and the more heavily chitinized central axis sends out a 
ridge on either side of this depression. Otherwise the whole piece 
may be quite uniformly chitinized. <A lateral view of this organ 
shows that it follows the curved outline of that part of the seg- 
ment in which it lies. 

The use or the homology of the ‘“‘breastbone’’ has been a sub- 
ject of some speculation. Giard (8) says that some of these 
Cecidomyid larvae have the power.of leaping, using their well 
developed spatula and terminal corneus papillae for this process, 
and in this wise: the larva bends itself almost into a circle, and 
hooking together the breastbone and papillae breaks the con- 
nection thus formed and the spring occurs. In the species re- 
ferred to by Giard however the spatula is well muscled, strongly 
chitinized, projects well externally, and the corneus papillae are 
developed. The advantage gained by this faculty of leaping is, 
according to Giard, the assurance of the dissemination of the 
species, which in turn insures an abundant food-supply and 
therefore the successful perpetuation of the species. The larvae 
of Diplosis loti and jacobeae, cited as examples by him as having 
this leaping power, are gregarious and would not leave enough 
food for their descendants if they were not thus motile. This 
theory does not seem adequate inasmuch as the adults look after 
the food-supply for their progeny, and being active and winged, 
would have no difficulty in finding another suitable plant on 
which to oviposit. The motile habits of the larva brought 
into play just before pupation would indicate a search on the 
part of the insect for a proper place for pupation, the pupa being 
formed underground and probably at no great distance from the 
abandoned food-plant. In C. resinicolotdes the spatula is not 
well developed, projects but slightly anteriorly, and the posterior 
corneus papillae are absent, also as the insect lives both in the 
larval and pupal stages in pitch, the breastbone could hardly be 
used as a locomotor organ. In fact it does not appear to be con- 


rgto| Anatomy of Cecidomyta resinicoloides 47 


nected with any muscles. Owing to the medium in which the 
larva lives, it is possible that it has lost this power of leaping by 
the degeneration of the organs so well developed in the saltatorial 
Cecidomyid larvae. Locomotion in the larva of C. resinicoloides 
is chiefly, if not wholly, accomplished by the extension and con- 
traction of its spiny segments, with the possible assistance of the 
pseudopod-like processes. 

Another theory advanced to explain the use of the breastbone 
is that it is used as a perforator (17). It could thus be used in 
some gall-forming species which pupate outside their gall, to 
force an exit therefrom. L.O. Howard (11) refers to a view pro- 
posed by Enock and sustained by Marchal according to which 
the spatula is used to reverse the position of the larva in the 
puparium. In the case of C. resinicoloides this last theory might 
apply, but it seems more probable that the spatula is used in 
making the cocoon, in pushing out a space in the resin, and form- 
ing the thin, convex cocoon cap. Inasmuch as, according to 
Giard, the spatula is only fully developed in the last moult, it is 
quite evident that this organ is intimately associated with the 
prepupal actions of the larva. Osten Sacken says (14): ‘‘If the 
supplementary (14th) segment be considered as a part of the head, 
this breastbone might be taken for the mentum, in analogy to the 
horny mentum of the larvae of the Tzpulariae.”’ 

Pseudopods. These are rounded, somewhat nipple-like pro- 
cesses, most prominent on the thoracic segments and usually in 
double pairs on the segments. Segment I has a pair on each 
anterior side of the spatula and bordered anteriorly by the 
imaginal leg discs; segments 2 and 3 have each a more prominent 
double pair, transversely arranged in the middle of the segments 
and likewise bordered anteriorly by the leg discs, those of the 
wings being lateral; segment 4 has an inconspicuous pair of pseu- 
dopods; segment ro some indistinct protuberances; while on 
segment 11 are a pair of rather widely separated protuberances 
anterior to the anal slit. The last or 12th segment is terminated 
by two conical projections bearing a bristle at their apices. 

Spiracles. There are nine pairs of spiracles, normally situated, 
one on the first thoracic segment and one each on segments 4-11. 
The last pair is placed apparently on the last segment, but Osten 
Sacken (14) has pointed out that this segment is in reality the 
8th abdominal, the gth being in C. pint and C. pint-inopts, (the 
latter being considered a synonym of C. resinicola) unusually 


48 Annals Entomological Society of America [Vol. III, 


small and concealed under the 8th abdominal segment. The 
spiracles are horny, conical projections placed somewhat above 
the lateral line, especially on segment 1. The anal spiracles are 
far larger than the lateral ones, -and are of most importance to the 
larva. They are truncated conical, with four stout spines. No 
young larvae were examined, but Eckel (5) says of the related 


resimicola: ‘‘The tracheal system undergoes no change with the 
larval growth except the odd development about the anal 
spiracles.’’ In the young resinicola larva each terminal spiracle 


is a long horny tube, whereas in full-grown specimens they 
resemble those of resinicoloides. The 3rd thoracic segment has 
no spiracles although the tracheal tube is present here and well 
developed. Eckel has noted this in the eastern species. 

Hairs and Bristles. These may be divided into two classes: 
(1) longer sparse hairs; (2) shorter numerous bristles. Those of 
the first class are, as far as could be ascertained, present on all 
but the supernumerary segment. On segments 4-10 are two 
larger hairs situated one below the other, the first close to the 
base of the spiracle. On segment 11, laterally and just below the 
spiracle is a large hair. On segment 12 are two apical hairs 
already referred to. These hairs, commonly near the spiracles, 
may assist in keeping the latter clear of the soft resin in which 
the larva lies. 

The bristles* are arranged in transverse rows forming long 
patches on the dorsal and ventral surface of the body and thin- 
ning out laterally. Dorsally these bands are on the anterior 
portion of segments 2, 3, 4, 5 and 6, and on the posterior part of 
segments 8, 9, and 10. The band is heaviest on segment 4 where 
a blank space cuts out a broad mesal notch on its posterior bor- 
der. On segments 2 and 3 the band is broad and almost divided 
by a transverse space, and on segment 5 is further reduced to 
three patches. The patch on segment 6 is almost obsolete. 
These bristles point posteriorly while the simple bands on seg- 
ments 8, 9 and 10 have them pointing anteriorly. 

Ventrally the bristle rows are not broken up but are simple, 
and are present on the anterior part of segments 2, 3, 4, 5, 6, 7, 8 
and 9 where they point posteriorly and on the posterior portion 
of segments 4, 5, 6, 7, 8, 9, 10 and 11, where they point anteriorly. 
These rows pointing forward thin out anteriorly, and on seg- 


* J}lustrated in Ent. News, XX, Jan., 1909, Plate I. 


Tg10] Anatomy of Cecidomyia resinicoloides 49 


ment 4 are quite small and inconspicuous, whereas the sets point- 
ing backward become thicker anteriorly. We see then that these 
rows of bristles are heaviest at either end of the larva and absent 
or imperfectly developed along the middle of the body, there 
being a gradual transition from the posterior—to the forward— 
pointing bristles. The bristles thin out laterally and are more 
numerous on the venter than on the dorsum. 

Although the larva is not an extensive traveller, it is never- 
theless quite active and moves about considerably within the 
limited space of the resinous mass in which it lives. It cannot 
survive very long when completely submerged in the gummy 
exudation, but lives with its anal spiracles at the surface of the 
mass or in a hollow therein, while its mouthparts are in close 
proximity to the abraded cambium. It must also turn itself 
around in its cocoon before pupation. It was observed that the 
posterior portion of the larva is usually more extended and the 
cephalic end often more contracted. When the larva desires to 
advance, its anterior segments are extended, thus exposing the 
strong rows of backward-pointing bristles, while the hinder end 
is somewhat drawn in, thus concealing its bristles, and, by con- 
tracting and expanding the anterior segments the desired move- 
ment can be obtained. It would not seem of so much importance 
that the larva move its whole body backwards, but that it extend 
its caudal spiracles to an opening for air is quite imperative. 
Thus it could firmly anchor its cephalic end to a desired spot, and 
by means of the forward-pointing bristle rows extend its spiracles 
to the surface. By sufficient contraction the rows of bristles at 
one end could be completely concealed and those of the other 
end fully exposed and brought into play. The pseudopods 
may also assist in the movements of the larva. 


INTERNAL ANATOMY. 


Tracheal System. P\. VI, Figs. 4 and 5. This was very care- 
fully worked out, the tracheal arrangement being followed to 
the finest ramifications. As in other insects it consists of longi- 
tudinal trunks connected with the spiracles by lateral branches. 
Figs. 4 and 5 of plate VI shows this system from a dorsal and 
ventral aspect, of a larva submitted to some pressure under a 
coverglass. It consists of a pair of longitudinal dorsal trunks 
beginning at the spiracles of segment 1 and terminating in the 
caudal spiracles of segment 11. At G, Fig. 4, a stout branch is 


50 Annals Entomological Society of America  [Vol. III, 


sent ventrad, which divides almost immediately, the one branch 
proceeding to the head while the other, bending posteriorly, sup- 
plies the brain and sub-cesophageal ganglion. This posterior 
branch is F,, in Plate VI, Fig. 4. 

The two dorsal trunks are connected with the spiracles by 
short branches and are joined to each other in segments 4-9 and 
11 by cross tubes, that of segment 11 being quite stout while the 

rest are slender and send out two re-dividing branches. These 
cross-tubes usually lose their striated appearance at their middle, 
appearing composed of two tubes fused together by a pair of 
sponge-like knobs. In segment 10, there is no fusion, the tubes 
remaining separate. Fig. 6, E,,, Pl. VI, shows a condition observed 
in two larvae in which the left branch is quite swollen basally, 
which perhaps was the normal thickness of both tubes, through- 
out before atrophy occurred and the tubes separated. The fact 
that in C. resinicola there are complete cross tubes in segments 
4-11 would appear to strengthen this view. Furthermore it will 
be noticed that in C. resinicoloides the cross tube in segment 11 1s 


quite stout and may substitute in a measure for the tubes in the: 


above segment. Ramifying tracheae are apparently not so 
necessary in the last few segments. 

The transverse lateral tubes D,—D,,, Fig. 4, terminate in the 
spiracles except in segment 3 where there is no spiracle, in which 
case the tube disappears near the body surface. They are con- 
nected with one another either by the lateral longitudinal system 
through the short tube R,, R,, etc., as in segments 3-8, or they 
may run directly through the former, as in segments g and to. 
This arrangement, however, is frequently asymmetrical. 

The lateral longitudinal trunks arise from the dorsal ones in 
segment 2, Fig. 4, A,, Pl. VI, and terminate as free branches in 
segment 10, C,,, Fig. 4, and apparently correspond to the ventral 
tubes as illustrated by Eckel (5), but differ widely from the latter 
in that they do not join the dorsal trunk posteriorly. They send 
out many branches F,, F,, K,, K;, etc., but the arrangement of 
these is not always constant. The primary function of the 
more anterior of these branches is to aerate and support the ner- 
vous system, that of the more posterior ones to supply the Mal- 
pighian tubules. 

The ventral longitudinal system Fig. 5, Pl. VI, arises from the 
main dorsal one close to the anal spiracles in segment 11, and 
proceeding anteriorly commences ramifying in the middle of 


T9QIO] Anatomy of Cecidomyia resinicoloides om 


segment 8, where it turns dorsally, closely following the Mal- 
pighian tubes to which short branches are sent. 

Circulatory System. This system owing to its transparency 
and delicacy was quite difficult to follow, and so a detailed 
description of it cannot be presented. The dorsal vessel is a 
thin-walled, transparent tube which arises in front of and below 
the brain, and proceeds obliquely upwards until it meets the 
body wall in the posterior part of segment 4 where it is secured 
by the suspensorium. Thence it runs along the dorsum extending 
at least to the posterior transverse tr Ae Aes tule. Sit is diy: ied 
into chambers and the valvular orifices (ostia) on either side 
open and close inwardly. The dorsal vessel pulsates especially 
in its anterior portion where there are several orifices. On each 
side of the heart in the middle of segment 4 is a strip of loose 
pericardial cells which proceed posteriorly, closely following the 
vessel. 

Alimentary Canal. Pl. VII, Fig. rt. The alimentary canal 
consists of the long slender cesophagus D, the larger scarcely 
differentiated fore- and mid-stomachs (proventriculus and ven- 
triculus) G, the slender and much curved ileum or small intestine 
I, the colon K, and the rectum L. A pair of long, recurved 
salivary glands C. F. open into the mouth, and two Malpighian 
tubules H, likewise recurved, are fastened to the fore part of the 
ileum. The whole system is supported chiefly by the trachez. 

Lying above the cesophagus and salivary glands and extending 
well beyond the rods of the head skeleton is a large blind sac, S, 
Figs. 1 and 2, Pl. VI, which probably opens into the mouth. It 
contains in its middle a curious, dark purplish-brown object, A, 
Figs. 1 and 2, Pl. VI, which is lobed anteriorly, and posteriorly 
where it becomes semi-translucent and breaks up into small 
granules. This object is evidently the ‘‘point oculiforme’”’ of 
Giard (8), who noted it also in a Cecidomyia, but what its function 
is he does not state. The large blind sac is apparently not 
muscular, though it is possible that it may serve as a food reser- 
voir. The ‘point oculiforme”’ suggests a strainer of some sort, 
especially if the sac were muscular. The cesophagus extends to 
the end of segment 3 as a very slender and delicate tube, and is 
chitinized within the rods of the head skeleton, C, Fig. 2, Pl. VII, 
anid tio. 6, Pl Vile 

The large, straight, muscular stomach extends from the 4th 
to the 8th segment inclusive, and its cellular coat is made up of 


52 Annals Entomological Society of America [Vol. III, 


more or less regular longitudinal rows of large protruding cells. 
The whole canal, including the salivary glands and the Mal- 
pighian tubules is clearly visible through the larval body wall. 

In the posterior portion of segment 8 the slender ileum arises 
abruptly from the thick ventriculus as a curved and folded tube. 
Its cellular wall consists of several rows of large cells distinct in 
the anterior orange portion, less so in the whitish translucent 
middle part, and disappearing in the slenderer hinder end, which 
opens into the delicate grayish colon. The rectum is a rather 
colourless tube of considerable length terminating in the anus on 
segment It. 

The extreme hind part of the canal probably functions but 
little, except perhaps when the larva is near pupation. It was 
not seen to contain any waste matter, and if the larva was 
submitted to considerable pressure under a cov erglass no excre- 
ment would be discharged. 

Glandular and Excretory A ppendages of the Alamentary Canal. 
The salivary glands are of large size and extend along each side 
of the ventriculus to the middle of segment 6 where they 
recurve dorsally and proceed to the anterior end of segment 5, 
being fastened to the ventriculus at that point. The larger 
posterior portion of the glands are glassy white and of monili- 
forme aspect, with several, rows of large glandular cells containing 
large nuclei. At about the middle of segment 3 the glands 
become narrower, lose their glassy, large-celled appearance, and 
twisting spirally downwards, then upwards, dilate abruptly into 
granular ampullae of a pale yellowish-white color. These are 
three in number, two larger posterior ones and a single incon- 
spicuous anterior swelling. From this latter point the glands 
taper gently, and at about the posterior border of the super- 
numerary segment give place to the tracheoid ducts. These are 
short slender tubes and join to form the common duct, B, Fig. 2, 
Pl. VI, a short distance within the chitinous rods of the head 
skeleton. 

Whether the secretion of the salivary glands of the larva 
produces an irritation in the plant tissue thereby causing a flow 
of resin, or whether ‘‘the twistings of perhaps a dozen spiny- 
skinned larvae smooth out a round cavity’’ (in the resin), ‘‘the 
irritation causing a constant supply of fresh resin,’’ to use Eckel’s 
own words as regards C. resinicola, is a difficult thing to determine. 
Giard (9) thinks that the ‘‘zymase’’ secreted by the salivary 


Tgto] Anatomy of Cecidomyta resinicoloides 53 


glands of the Cecidomyid larva determines the gall. No gall 
is formed in the case of the pitch-inhabiting midges, but the 
salivary secretion in this case might well stimulate and maintain 
the resinous exudation. 

Attached interiorly to the second ampulla of the salivary 
glands and stretching in a curve across the brain which it overlies 
very closely is a curious, loose ill-defined structure of granular 
appearance, F, Fig. VII, Pl. 2, and containing brown patches, 
especially towards the bases. This I believe is Weismann’s 
‘“‘cell chaplet,’’ which he defines thus: ‘“‘It consists of a string of 
large cells closely united which hangs like a garland, free in the 
body cavity. Its two ends are connected with the salivary 
glands,” etc. What its function is remains to be ascertained. 

The Malpighian tubules are of a deep orange color and arise 
from the anterior end of the ileum immediately where the latter 
isejoined to the ventriculus by a delicate; membranes. ihe 
arrangement of these two tubules with regard to the ileum and 
colon is not symmetrical since the left one follows rather closely 
those two divisions of the proctodaum, while the right tubule is 
quite free. Fig. 5, Pl. VII shows the left tubule in its relation to 
the proctodeum. Arising from the ileum each tube proceeds 
dorsally but not quite in symmetry as the right tube bends anter- 
iorly and outwardly while the left first curves somewhat poster- 
iorly, P, Fig. 1, Pl. VII and then follows the common course. Each 
tube is closely appressed to the ventriculus to which it is slightly 
fastened and proceeds anteriorly past the middle of segment 7. 
Here however the left one by reason of its posterior bend does 
not reach quite so far forward as its mate, but terminates slightly 
farther posteriorly and ends somewhat beyond the middle of the 
colon. - Both tubes run along the ventral tracheal trunks in their 
terminal portions. _ 

Giard says (10):—‘‘In all the larvae of the Cecidomyidze 
which I have studied, these tubes, two in number are united into 
an elegantly recurved handle, and open near the anus, the 
proctodeum being excessively short.” In C. resinicoloides the 
tubes end freely, as we have seen, and the proctodzeum, if 
extended would probably at least be as long as the rest of the 
alimentary canal. 

It may be well said here that the alimentary canal (and its 
appendages) of C. resinicoloides much resembles that of Dzplosis 
buxi illustrated by Berlese (2). 


54 Annals Entomological Society of America [Vol. II], 


Organs of Reproduction? At about the termination of the 
Malpighian tubules, and below the same is a pair of elongate, 
fusiform, transparent bodies terminating anteriorly and poster- 
iorly as slender tubes. These bodies were not observed in all 
the larvae examined. They are probably the developing repro- 
ductive organs. 

Nervous System. Plate VIII, Figs. 1 and 2. The nervous 
system is more specialized than in many other dipterous larvae, 
in that there is considerable cephalization of the ganglionic chain. 
This chain is composed of ten distinct ganglia, exclusive of the 
sub-cesophageal one, and extends from the middle of segment 2 
to the middle of segment 4. It is supported chiefly by the con- 
verging branches of the lateral longitudinal tracheae, while the 
brain is kept in place and aerated largely by the stouter pair 
of tracheal branches, F,, which arise from the main dorsal system. 

The brain or supra-cesophageal ganglion consists of two lobes 
of larger size than the rest and broadly joined to the sub-cesopha- 
geal ganglion, the commissure not being apparent. The cesopha- 
gus passes between these two ganglia, as probably also the dorsal 
vessel, the salivary glands running on either side. - At least one 
pair of nerve cords from the brain, and two or more from the sub- 
cesophageal ganglion proceed anteriorly, and each of the gangha 
of the chain sends out several nerves, some of which run along 
the lengths of the supporting tracheae. The last ganglion of the 
chain is more elongate than the others. 

Very little could be made out of the sympathetic nervous 
system, and the paired ganglia, C, Figs. 1 and 2, Pl. VIII, arising 
posteriorly from the lobes of the brain are all that I feel sure of. 

Musculature. The muscles were not studied though it was 
observed that they were numerous and well developed, a condi- 
tion quite necessary in such a restless insect. Several muscles 
are fastened to the chitinous prongs of the head skeleton, and a 
number of large muscles radiate from the caudal cleft, being 
fastened to the body wall. Thus the larva is enabled to retract 
quickly its caudal end into the resinous mass if disturbed. 

Adipose tissue. This is very well developed, so much so in- 
deed that it obscures in a great measure the different visceral 
systems. The fat body is of a dirty yellowish-white color, and 
consists of a loose network of lobes with long meshes, and is con- 
tinuous throughout the body, thickest below the middle, thence 
tapering to either extremity, extending anteriorly to the base of the 


r. 


ie a = 


TQI0] Anatomy of Cecidomyia restnicoloides 55 


first pair of spiracles, and posteriorly to near the anus. It is 
circumscribed and largely supported by the tracheal system. 
Some time before pupation this fat tissue loses much of its com- 
pactness, becoming more or less watery in appearance and 
assumes a reddish color. 

Imaginal Discs of Wings and Legs. Plate VIII, Figs. 6 and 7. 
These are easily seen in large full-fed larvae where they appear 
as more or less circular discs attached to the body wall. 

The wing-buds, Pl. VIII, Fig. 7, A,A,, are larger than those of 
the legs, lie very slightly above the lateral line of the body, and are 
supplied with branches of the lateral and dorsal tracheae. Both 
wing- and balancer-discs arise from about the middle of their 
respective segments, ending posteriorly in a lobe and tapering 
anteriorly into the body wall. 

The three pairs of leg discs, Fig. 6 A,, A,, Az, are situated on 
either side of the ventral line and are more distinctly circular 
in outline than the wing discs and more strongly and completely 
cut by an inner ring. They are supplied by small branches of 
the lateral tracheae. 


BIBLIOGRAPHY. 


1. Aldrich, J. M. (1905). Catalogue of N. A. Diptera, pp. 151-152, 
160. 

2. -Berlese, A. (1906). Sistema Ghiandolare, Ghiandole Confinate, 
(Gli'Tnsetti, I, p. 523). 

3. Comstock, J. H. (1879). Diplosis resinicola, (Rept. Dep. Agr., 
1879, p. 256). 

4. Cannon, W. A. (1900). A Botanical Study of the Gall formed 
by Diplosis pini-radiatae, (Am. Nat., XXXIV, No. 406, pp. 
801-810.) 

5. Eckel, L. S. (1903). The Resin-Gnat Diplosis and Three of Its 
Parasites, (Ent. News, XVI, pp. 279-284). 

6. Felt, E. P. (1903). C. resinicola, (N. Y. State Mus. Memoir 8, 
II, pp. 410-413.) 

7. Felt, E. P. (1903). C. pini-rigidae, (N. Y. State Mus. Memoir 
8, II, pp. 423-425). 

8. Giard, A. (1893). Note sur l’organe appelé Spatula sternalis 
chez certaines larves de Cecidomyes. (Bull. Ent. Soc. France, 
Feb. 22, 1893). 

9. Giard, A. (1893). Sur un genre nouveau et une espece nouvelle 
de Cecidomyide (Drisina glutinosa), (Bull. Soc. Ent. France, 
Dec. 27, 1893). 


Annals Entomological Society of America  [Vol. III, 


Giard, A. (1893). Sur less tubes de Malpighi de les larves de 
Cecidomyes, (Bull. Ent. Soc. France, Feb. 22, 1893). 

Howard, L. O. (1898). The Hessian Fly in U.S., (Bull. 16, Dep. 
Agr., Div. Ent.) 

Kertesz, C. (1902). Catalogus Dipterorum, II, pp. 116-117. 

Mik, Jos. (1897). Einiges uber Gallmucken, (Wein. Ent. Zeit., 
XVI, pp. 290—292.) 

Osten Sacken (1862). On the Habits of the Cecidomyide, (Mon. 
Dip. N. A., Pt. 1, pp. 179-198.) 

Osten Sacken (1871). Biological Notes on Diptera, (Trans. Am. 
Ent. Soc., III, pp. 345-346). OD. resinicola, etc. 

Packard, A. S. Diplosis resinicola and Pini-rigidae, (5th Rep. 
U.S. Ent. Comm., 1886-1890, pp. 797 et seq.) 

Sharp, D. (1901). Use of Spatula in Cecidomyide, (Camb. Nat. 
Hist. Ins., Pt. II, p. 460). 

Snow, W. A. (1900). The Destructive Diplosis of the Monterey 
Pine, (Ent. News, XI, pp. 489-494). 

Townsend, C. H. (1893). Notes on Some Cecidomyiide of the 
Vicinity of Wash., D. C., (Proc. Ent. Soc. Wash., II, pp. 
389-390). D. resinicola, etc. 


1910] Anatomy of Cecidomyia resinicoloides 57 


EXPLANATION OF PLATES. 
All figures greatly enlarged. 
PLATE, VI 


Fic. 1. Dorsal view of head skeleton; S. blind sac; A, point oculiforme; 
O, cesophagus; B, salivary duct; N, nerve. 

Fic. 2. Lateral view of head skeleton; S, blind sac; A, point oculiforme; 
B, tracheoid duct; O, cesophagus; N. nerve. 

Fic. 3. Sternal spatula; A. A, A, dorsal view; B, lateral view. 

Fic. 4. Dorsal view of tracheal system of mature larva; A,—A,,, dorsal 
longitudinal trunk; B,,, ventral longitudinal trunk; C,—C,,, C,,, lateral longi- 
tudinal trunk; D,—D,), lateral transverse tubes to spiracles; E,—Ej,,, E,,, cross 
tubes; F, F,, F,—F,, branches to nervous system; K,, K,, branches to nervous 
system; G, branch from dorsal trunk; H, branch to head; R,, R,, R,, connecting 
branch between C and D; M, head; S, supernumerary segment. 

Fic. 5. Ventral view of tracheal system of mature larva, lettering same as 
in Fig. 4, Bg, dorsal turn of ventral longitudinal trunk. 

Fic. 6. Aborted cross tubes of Seg. 10. 


PraAte Vile 


Fic. 1. . Dorsal view of alimentary canal of mature larva; A, head; B, blind 
sac; C, ampullae of salivary glands; D, cesophagus; E, Weisman’s cell chaplet ; 
F, large-celled portion of salivary glands; G, ventriculus; H. Malpighian tubule; 
I, ileum; J, ventral Jongitudinal trachea; K, colon; L, rectum; M, anus; P, pos- 
terior bend of left Malpighian tubule 

Fic. 2. Head skeleton and portion of salivary glands; A, antenna; B, head 
skeleton; C, cesophagus; D, common duct of salivary glands showing tracheal 
character; E, ampulla of salivary glands; F, Weisman’s cell chaplet. 

Fic. 3. Sagittal section through larva; A, adipose tissue; B, muscle; C, ali- 
mentary canal; O, cesophagus; I, ileum; P, Malpighian tubule; M, M,, salivary 
glands; H, heart; S, blind sac. 

Fic. 4. Cross trachea in segment 11 showing central spongy mass. 

Fic. 5. Lateral view of proctodeum; A, posterior end of ventriculus; B, 
ileum; C, colon; D, rectum; E, anus; F, Malpighian tubule; L, L,, slender portion 
of ileum; O, point of origin of Malpighian tubules. 

Fic. 6. Chitinized portion of anterior end of cesophagus. 

Fic. 7. Dorsal view of anal spiracles, 12—last segment. 


PuaTe VIII. 


Fic. 1. Dorsa] view of nervous system; A, sub-cesophageal ganglion; 
B, brain; C, one of the paired ganglia of the sympathetic system; F, trachea 
feeding brain and sub-cesophageal ganglion. 

Fic. 2. Lateral view of nervous system; lettering as in figure 1. T. 
trachea. 

Fic. 3. Cross section through posterior portion of larva; A, adipose tissue; 
B, muscle; C, ventriculus; H, heart; M, Malphigian tubule; L, L,, slender 
portion of ileum corresponding to L, L.,, of Fig. 5, Plate VII. 

Fic. 4. Cross section through ventriculus showing epithelial cells. 

Fic. 5. Cross section through anterior portion of larva; A, adipose tissue; 
B, muscle; C. ventriculus; M. salivary glands; H, heart; F, pericardial fat cells. 

Fic. 6. Ventral view of anterior portion of larva (portions of segments 1, 
2 and 3) showing imaginal leg discs, A,, A,, A,; B, dorsal longitudinal trunk of 
tracheal system; C, lateral longitudinal trunk; D, tracheal branch feeding brain, 
this is F,, Fig. 4, plate VI. 

Fic. 7. Dorsal view of portion of segments 2 and 3, showing imaginal 
-wing discs, A, and A,; B, dorsal longitudinal trunk; C, lateral longitudinal trunk. 


Vou. II, Pate VI. 


A. 


Ss. 


ANNAIS E, 


Williams. 


F. X. 


eS 


app YEE 


VoL. III, PLATE VII. 


ANNALS E, S. A. 


“som” Gace as 


Williams. 


xX, 


F. 


III, PuatTe VIII. 


VOL. 


S 


ANNALS E. &. 


Williams. 


xX. 


F, 


SOME NEW FACTS ON THE BIONOMICS OF THE 
CALIFORNIA RODENT FLEAS. 


M. Bruin Mirzmatn, B. S., 


University of California. 


The thorough investigations which the United States Public 
Health and Marine-Hospital Service are at present undertaking 
in suppressing bubonic plague in California presented the writer 
with an opportunity to study the parasites associated with the 
transmission of the disease among rodents. 

In this contribution we have not taken into account the fac- 
tors involved in the epidemiology, but have restricted the scope 
of the paper to a discussion of the general habits and transfor- 
mations of the California rodent fleas. Many of the observations 
on the bionomics of the fleas are quite different from those re- 
corded by the investigators of the English Plague Commission in 
India. This is due in a measure to difference in climatic condi- ’ 
tions, to which fleas as a rule are easily influenced; and the differ- 
ence in the species of flea. The English workers confined their 
experiments to one species, namely, the Loemopsylla cheopis 
Roth. We have considered the rodent fleas generally, giving 
most attention to the squirrel flea C. acutus Baker. 


FLEAS IN RELATION TO THEIR ENVIRONMENTS. 


We have made an effort to rear fleas in the laboratory by 
attempting to duplicate conditions found to exist in nature. The 
human flea, P. zrritans Linn, was found to develop very satisfac- 
torily in a medium composed of floor sweepings taken from the 
cracks in the floor. The squirrel fleas and rat fleas were found to 
develop well in material taken from the nests of their respective 
hosts. We have experienced no difficulty in raising these para- 
sites by placing animals covered with fleas in cages with a bed- 
ding of sawdust; and the only provision necessary for complete 
development was the addition of fresh sawdust to prevent the 
accumulation of too much moisture. 

An experiment was made to determine in what media fleas 
away from the host would survive longest. For this purpose 
we tested various materials shown in the following table: 


61 


62 Annals Entomological Society of America  [Vol. III, 


LENGTH OF LIFE IN VARIOUS MEDIA. 


C. acutus. 
‘ = al x l = oe a l 
‘DrySand| Moist B Dae Sawdust | | Moistened 
Days Re- | with Sand i san@) Moistened : | Sawdust 
- hie : rom e: Dry - 
moved from) Squirrel| from | gauirre] with aera aat with 
Host Drop- | Squirrel | "Yeu Horse 2 Wheat 
pings Nest canis Serum | Grains 
EA | 
2 1 ies a pe a Od ev 10 F. 4M. 6F. 4M.6F. 4M.6F 
7 2 Ms5 FE 4F. |Alldead| 1M. 3F. 4F. 1M.5F 
9 ih The 2F. 22. 1M.4F 
10 12 M. 5 F./All dead All dead All dead 1M. 4F. 
14 1M Uk. 1M. 3F. 
16 1M.1F. 
17 All dead 1M.3F 
20 3 FE: 
25 1 Se 
26 7 | “All dead 


*M—Male; + F—Female. 


The controls 5 M. 5 F. were all dead on or before the seventh 
day. 

In the medium of moistened sawdust mixed with a few wheat 
grains it was found that the wheat sprouted in the sawdust and 
held sufficient moisture to provide a suitable condition for larvae 
as well as for adults. The mould which formed in a short time 
did not seem to affect the insect life. 

It is seen that the fleas did not fare well in the medium of 
dry sand. The sand was mixed with clay dust which would rise 
whenever the fleas hopped in the vial. As a consequence death 
resulted presumably by stoppage of the spiracles. 

It will be seen from a survey of this table that fleas taken from 
the natural host may be kept alive without food for a considerable 
time. The medium of moistened sawdust with a few grains of 
wheat seemed to answer the moisture requirements for flea life. 


LOCOMOTION. 


The only literature that has come to our notice on the jump- 
ing powers of fleas appears in the Journal of Hygiene, 1906, Vol. 
6, p. 464. Here we havea note: ‘‘It had previously been found 
that a rat flea could not hop farther than five inches.”’ The 
species in question in these experiments was the L. cheopis, which 
is found to the extent of 99% on the rats in India. 


TQIO| California Rodent Fleas 63 


It seemed desirable to learn the jumping abilities of the com- 
mon California species. | An attempt was made to determine the 
distance upon a horizontal plane, as well as the height that the 
insect could jump. In the broad jump a few experiments were 
conducted with P. zrritans, the most active of our California 
fleas. In this species we have found the jumping distance varies 
considerably with the nature of the container from which the 
insect 1s observed, e. g., a foothold of wood enables the flea to 
jump a greater distance than one of glass. Since the irritans is 
found to predominate on floors of houses, a surface of wood was 
selected as the footing in our tests. In one experiment, five 
specimens were permitted to jump at will and the jumps of each 
were recorded. The mean average of ten jumps of each speci- 
men gave a distance of 7 3-10 inches. The longest jump recorded 
was 11 inches; this was made by a female. In another experi- 
ment, a female, which was starved for five days prior to the test, 
made four jumps of respectively 10 5-10, 11, 12, and 13 inches; 
averaging 11 5-8 inches. Thirteen inches was the longest jump 
recorded on a horizontal plane. A jump of 15 7-8 inches was 
made downward at an angle of thirty degrees. 

This specimen was then permitted to feed fifteen minutes on 
the arm of an attendant, after which it was carefully returned to 
the container, and its jumps for five minutes recorded. The 
longest jump after feeding was 12 inches. 

The jumping powers of squirrel and rat fleas were tested in 
this manner: Three hundred and seventy-five live squirrel fleas 
(C. acutus) and one hundred and fifty live rat fleas (C. fasciatus 
and L. cheopis) were placed in two distinct lots in fifteen open 
specimen vials in a water bath and left undisturbed for two days. 
At the end of the period, the vials were examined, the water 
bath and the surroundings being carefully inspected. No fleas 
could be detected in the water bath or in the vicinity; the fleas 
in the vials were counted, the original number being present. 
They were apparently as active as when removed from their hosts. 
The containers were cylindrical vials 3 3-8 inches in height and 
I 3-16 inches in diameter. The same test was tried in open shell 
vials of the next size smaller (3 1-8 x 1 inch) and the fleas were 
found jumping into the water bath. 

L. cheopis, the rat flea, was tried for its jumping power. It 
had been previously observed that members of this species were 
unable to leap out of open shell vials 3 3-8 inches in height. It 


64 Annals Entomological Society of America  [Vol. III, 


was thought desirable to permit the flea greater latitude making 
the trial fairer and more practical. Twenty fleas (eight males 
and twelve females) were placed in a rectangular porcelain dish, 
the dimensions of which were 11 7—8 inches by 7 5-8 inches by 
2} inches deep. A light sprinkling of moist sand was placed in 
the bottom of the dish to provide a firm footing and the sides of 
the dish were extended to the height of 4} inches, enclosing the 
container entirely by strips of ‘‘tangle-foot’’ paper. At the end 
of twenty-four hours the dish with the fleas was examined, and it 
was: observed that several fleas had leaped high enough to 
become entangled on the adhesive paper. The position of these 
was noted, the distance from the bottom of the pan was meas- 
ured. The entangled insects were pried off, the sticky material 
dissolved with alcohol and the insects examined microscopically. 
Five specimens were collected, two males and three females. 
The locations on the adhesive paper relative to the base of the 
dish were as follows: 

1 F. 24 inches 1 F. 3 1-8 inches 1 F. 2 9-16 inches 
1 M. 2 5-8 inches 1 M. 3 1-16 inches 

A census of the fleas remaining in the bottom of the dish gave 
6 M. and 9 F. showing that none jumped over the surrounding 
paper, and that all jumping over 2} inches were embedded in 
the adhesive paper. 

The jumping powers of P. irritans were further tested, twenty 
fleas of this species being placed in a rectangular museum jar, 
the inside dimensions of which were 2} x 54 x 7} inches depth. 
The glass lid of the jar was coated with ‘‘tangle-foot’’ and the 
sides of the jar within two inches of the top were likewise coated, 
with the same material. The following morning the jar was 
examined and two specimens were seen embedded in the “‘tangle- 
foot’? on the lid. The inside height of the jar being 7? inches 
from the base gives then this height as the perpendicular jump. 

Concerning the fleas ability to walk upwards on glass, we have 
noted that they cannot climb to any considerable distance. A 
great number of counts were made, the greatest distance observed 
being # inches. The climb appears very laborious and in all 
cases the flea dropped to the bottom of the jar after a few efforts. 

We have observed the manner in which these insects can find 
their way about upon their natural hosts. They pilot their paths 
among the dense hairs, walking on the flat of their tarsi, seeming 
to shuffle along. | When on the animal they seldom hop about, 


Igto] California Rodent Fleas 65 


unless disturbed or unless the host snaps at them or scratches 
when unusually annoyed. They hop freely when jumping from 
the animal to the ground or vice versa, or from one host to another. 
When a flea is cornered, that is, when it experiences difficulty in 
passing an obstruction, it proceeds like a swimmer using the side 
stroke. The parasite drops to its side and locomotion ensues by a 
vigorous sweeping movement of the legs, almost entirely by the 
use of the hairs and spines, especially through the medium of the 
powerful spines of the tibia, when it ambles along on the flat 
side in a striking manner. The spines of the leg seem to be 
peculiarly adapted for this side motion: it is in this fashion 
that the nimble parasite manages to become so very elusive. We 
tefer especially to the P. irritans. This may be tested by 
holding a live flea between the thumb and forefinger and unless 
you chance to be a flea trapper of long and painful experience 
the ingenious parasite will surely escape. 


CONSIDERATION OF COLOR ATTRACTION. 


We havea little experimental evidence on the question of attrac- 
tion of these insects towards color in animals. It is the prevailing 
opinion that white animals attract the greatest number of fleas. 
We have to report a few tests of color attraction. The material 
used in the first experiment was six guinea pigs showing extreme 
variations of color: three of these were pure white and three 
pronouncedly dark. These were placed in a large cage, the 
bedding of which was infested with squirrel fleas. After forty- 
eight hours the guinea pigs were removed simultaneously and a 
census of the fleas on the individual animals was made immediate- 
ly. Fleas and animals were anaesthetized at the same time, the 
white guinea pigs yielding respectively ten fleas, five fleas, and 
eleven fleas. The black guinea pigs yielded eleven, seven and 
four fleas; the last number came from a mixed black and white 
guinea pig. A summary of the fleas from the three white guinea 
pigs gave twenty-six fleas, or an average of about nine each; the 
two dark guinea gigs giving eighteen, averaging nine each, and 
the black and white giving four fleas, all of which were of the 
species, C. acutus, the squirrel flea. 

A second experiment was as follows: Four guinea pigs 
were placed in a large container, which was swarming with squir- 
rel fleas. After three days the animals were removed as before, 
and a census of the parasites was taken: one brown and black 


66 Annals Entomological Society of America [Vol. III, 


guinea pig yielded eighty-nine fleas; another brown and black 
guinea pig yielded fifty-seven fleas; a pure white guinea pig 
yielded fifty-seven fleas; and the other white guinea pig yielded 
sixty fleas. These guinea pigs were kept together in the open 
vessel under the same conditions of light and temperature. 

One experiment was conducted by using guinea pigs as traps 
for fleas in an infested house. The results shown are relatively 
meager due to the previous use of adhesive paper for flea trapping. 
In this test six guinea pigs were permitted to wander for twenty- 
four hours through the basement of the house. The fleas col- 
lected from the guinea pigs were P. trritans found on the hosts as 
follows: 


Color of animal Number of fleas 
Black guinea pig 1 
White guinea pig zZ 1 
Black guinea pig 1 
White guinea pig 1 
Black guinea pig 0) 
White guinea pig 0 


It is obvious from these experiments that color does not exert 
the influence generally claimed for it. The white animals are no 
more attractive to the fleas we have used than are the dark colored 
ones. 

TRAPPING OF FLEAS WITH MEAT AS A BAIT. 


An idea prevails that fleas can be attracted and trapped on 
account of their predilection for the odor of fresh meat. This 
idea was put to test in an experimental way. On the 23rd day of 
August, 1909, during a season in the year in which fleas were 
extremely abundant, a vacant house, which was found to be flea 
infested, was chosen for the experiment. To give an idea of 
the abundance of the parasites, two attendants, who had occasion 
to enter the first floor of the dwelling, remained for the period of 
five minutes and emerged covered with fleas. Approximately 
two hundred fleas were taken from their clothing and persons. 
The materials used for the experiment were twelve sheets of 
‘‘tangle-foot”’ fly paper, distributed in pairs among three rooms 
of the house. One sheet of each pair was supplied with a small 
fresh piece of cow’s liver. Care was taken to distribute the sheets 
in such a manner that the influence of light would be the same for 
each pair of sheets in the series. The sticky fly papers were left 
undisturbed for a period of three days, then collected and exam- 
ined. 


IgIo] California Rodent Fleas 67 


A similar experiment was conducted at a later date, December 
5th, 1909, when San Francisco dwellings were still flea infested. 
The sheets of fly paper were left in the basement of the 
house for twenty-four hours. The fleas in both experiments 
were removed by dissolving the “‘tangle-foot”’ in alcohol. The 
parasites which were all identified as Pulex irritans, were dis- 
tributed on the sheets as follows: 


Total number of fleas trapped. 


Experiment 1 | Experiment 2 
‘Adhesive paper) Adhesive paper Adhesive paper | Adhesive paper 
Pair with meat | without meat with meat without meat 
1 0 0) 2 4 
2 2 2 1 3 
Ba 2 1 6 4 
4 | 0 3 47 49 
| 6 12 1 6) 
(oI 4 1 4 1 
Totals) 14 19 61 61 


The table shows that according to our experience, though the 
data is limited, meat used as a bait under the conditions stated 
does not exert any special attraction for fleas. 


COPULATION AND EGG LAYING. 


The most prolific of the rodent fleas, Ceratophyllus acutus 
was set aside for studies in mating. The fleas were observed in 
nature in relation to the host, then microscopically. For the 
purpose of close observation, a live ground squirrel (Citellus 
beecheyi) was placed in a mouse jar, where it was kept for a few 
hours under surveillance. It was noticed that when the animal’s 
body was pressed against the glass that the pelage parted in 
furrows made by some active objects close to the skin. It was 
apparent that the squirrel was harboring a large number of active 
fleas, which occasionally came to view mounted on the hairs of 
thes host: 

The attention of the observer was attracted by a number of 
fleas which appeared gigantic in size. These proved on close 
inspection to be paired. Approximately 1 to every 3 of the total 
number of fleas infesting the squirrel appeared thus in copulation. 
The female of the pair covered almost entirely the body of the 


68 Annals Entomological Society of America  [Vol. III, 


male, which was quite lost to view, only the hind legs projecting 
beneath the abdomen of the female. The pair in copulation was 
observed never to feed, moving about as one insect; and anoma- 
lous as it appears, the member on the ventral side, the male, con- 
tributed no effort in the locomotion during the nuptial wander- 
ings. His fore and middle legs doubled under the sternum, the 
hind pair extending stiffly as though paralyzed beneath the 
abdomen, where they were held securely by the tarsal spurs of 
the female interlocking with the tibial spines of the male. 

The locomotory function was accomplished by the female, the 
front and middle legs of which were used freely in walking; by 
releasing temporarily the entwining spines of the hind legs she 
was able to leap at will. In this manner, the female moved 
about on the body of the host as though unhampered by its 
anchored mate. 7 

Seven pairs of these fleas were collected from the live squirrel 
with the aid of a camel’s hair brush, placed in cell slides and 
covered with glass slips. They were observed under the low 
power microscope, then kept in separate vials, one to several 
hours until the function was completed. 

As observed microscopically, the male adheres to the female 
by the pseudo-joints of the terminal segment of the antennae. 
This attached appendage is extended and pendant, interlocking 
by its hairs the bristles on the ventral side of the second abdomi- 
nal segment of the female. The hind femora of the latter grasps 
the male by the head at a point anterior to the antennal groove. 
The head of the male is held in position between the inner sides 
of the meta-femora of the female. The tarsal spurs of the female 
entangle the spines of the hind tibia of the male. The smaller of 
the pair thus suspended assumes the passive role moving at the 
volition of the female. 

The copulatory act is manifested in the male by the watch- 
spring like action of the spiral, which, coiling and uncoiling inter- 
mittently extends and withdraws the terminal apparatus. 

The male claspers are seen on either side of the vaginal groove 
of the female directly below the anal flap. The male makes a dis- 
tinct concavity in the caudal portion of the back; with the strong 
action of the muscle attachments the plate of the penis is made 
very elastic. The spasmodic pulsations of the curled up abdomen 
are followed by a current of bubbles passing through the genitalia 
of the male upwards to the female. The spermatheca of the latter 


IQIo] California Rodent Fleas 69 


is faintly discernible during the act as it swells and collapses. The 
length of time involving coition in the fleas under observation 
varied a trifle less than two hours to four and a half hours. The 
movements preliminary to the act were not noted. 

The male makes the first effort to disconnect or disentangle the 
pair. In one instance, the male attempted to free itself by press- 
ing vigorously the mid and hind tarsi against the hind tibia 
of its mate. Simultaneously, the antennae were torn from the 
recumbent spines of the abdominal segments of the female. The 
abdomen was straightened, relaxing the clutch of the claspers; 
the spiral contracted releasing the intromittent organ, and a 
final kick of the hind legs gave the male its liberty. The female 
Was seen on ten distinct occasions to approach the male, and the 
latter repelled the advances of its larger companion; at each con- 
tact of the female, 1t would jump in the air, attempting to remain 
on the opposite side of the vial. 

Mammalian blood appears essential for fleas to partake of 
normal functions of copulation and oviposition. In our exper- 
ience insects kept constantly in jars and reared from cocoons never 
having been fed upon a host have not been observed to copulate 
or oviposit. In thirty specimens of P. irritans taken from a house 
which had been vacant for six weeks we found, after three days 
observation, that the fleas were perfectly healthy; and although 
females predominated no eggs were found at the end of this 
period. As a-control, six females of this species collected from 
human hosts when kept in separate vials laid eggs normally, 
depositing from 5—12 in each instance. 

Four experiments with C. acutus, the squirrel flea, have given 
results similar to the foregoing, namely, that this species when 
reared from cocoons and kept starved in jars at room tempera- 
ture has not been observed to mate or lay eggs. 

In twenty-five paired specimens of several species kept under 
observation it appears that the male does not long survive the 
act of mating. It dies even before the female has laid its first 
batch of eggs. | When fertile females are kept under artificial 
conditions the eggs are laid in one laying in a period beginning 
two hours after copulation and extending to a maximum of 
thirty-six hours. When deprived of food the female has never 
been observed to oviposit after this length of time. 

We may note here that when used experimentally the female 
is invariably longer lived. Experiments to determine length of 


70 Annals Entomological Society of America [Vol. III, 


life with human blood diet show that female fleas of all species 
outlive the male by several weeks. This is doubtless true also 
under natural conditions, where we find in collecting fleas from 
the host that the females predominate markedly. 

The eggs require optimum conditions of temperature and mois- 
ture for hatching. They have never been found on the host 
except in one instance. In this case a dog was used in the 
laboratory for supplying fresh fleas; this was done by placing the 
canine on sheets of paper which in a few hours were littered with 
a large number of flea eggs. These were laid loosely on the host 
by the fleas, the eggs falling to the paper, where they were 
collected. Flea eggs have never been found on man, and if 
present, would not hatch under normal conditions of the body 
temperature. We have found that keeping eggs in the incubator 
at blood heat is sufficient to prevent hatching. If the eggs were 
laid on the host we would certainly expect to find them on the 
squirrel, on which animal fleas are most abundant. We have 
taken from two squirrels, respectively, 225 C. acutus and 376 of 
the same species; but in no instance even where fleas are present 
in such large numbers have we been able to find eggs even after 
carefully combing the host. The C. acutus is by all means the 
best criterion in this matter, since we have found that it lays 
more eggs than any other of the rodent fleas. 

The eggs are laid singly in small clusters, and may be viscous 
as in P. irritans, C. fasciatus, L. cheopis, C. acutus; or dry as in 
C. canis and C. musculi. The former adhere to the medium in 
which they are laid, and the eggs of the last two species are laid 
loosely, so that they roll when shaken in the vial containing them. 

Eggs may be laid while the insect is still under the influence 
of an anaesthetic; when covered by a glass slip; and when exposed 
to strong sun light. 

It appears to be the first impulse for the female to lay its 
eggs when removed from the host and placed under artificial 
conditions. The great majority of the eggs obtained were laid on 
the first day, beginning almost immediately after the fleas were 
captured. It is a common observation that many females with 
their abdomens distended lay their eggs as soon as the vial is 
closed over them. The number of eggs laid at one laying by 
different species, varies from 3 to 18; the rat fleas averaging 6 
and the squirrel fleas lay as many as 18. 


TQIO| California Rodent Fleas (a 


THE PROCESS OF HATCHING. 
C. acutus. 


Six eggs laid while the females were kept under observation 
were examined from time to time until hatched seven to nine 
days later. One egg, was observed microscopically during the 
entire process; the other five eggs used as controls were examined 
occasionally. 

On the seventh day of incubation at room temperature the 
premonitory signs of hatching were discerned ina very faint rising 
and falling of the exochorion (outer shell layer) on one side of the 
shell. During the night, seven hours later, the movement grew 
more extensive, the pulsations becoming quite pronounced, caus- 
ing the egg to shift slightly from its position. On the morning 
of the eighth day a deep gash was observed in the side of the egg. 
The gash is made by the egg opener, a wedge-shaped, horny, claw- 
like structure on the dorsal side back of the head of the embryo. 
This is operated so as to hew through the resistant shell by a 
series of slits or gashes. The initial gash increases slowly in 
length, encircling the egg within an hour. 

In the course of a few hours, seven successive gashes are 
made, the location of these is quite constant; two on each side 
of the egg and three at the center, the middle of the latter 
being the most distinct. The young worm produces these slits 
through the chorion (inner skin of shell) by turning with its 
dorsal side against the shell, striking with the egg pick first 
against the base of the egg, rising on its hind prolegs and striking 
rapidly as it climbs upwards. Prior to each stroke the egg pick 
is poised deliberately, the weight of the head accelerating the 
blow; this is augmented occasionally by a lateral shaking of the 
head. 

When the last slit encircles the shell, the embryo is at the 
most active stage, it effects a complete rotation in its shell at 
intervals of ten minutes. The gashes make the shell appear as 
though cut into ribbons. The rotary movement gradually 
subsides until a complete turning consumes twenty minutes, and 
almost imperceptibly there follows a lull. An inactive almost 
comatose condition prevails for a period of about seventeen 
hours. After the resting period, it appears that the egg pick is 
no longer functional; no new gashes are made, the embryo seeming 
contented to concentrate its energies against the middle gash. 


72 Annals Entomological Society of America  [Vol. III, 


This is enlarged by a puffing and dilating of the head; through the 
semitransparent shell is seen a constant bubbling and a subse- 
quent dilatation of the cuticle. 

On the third day of the hatching process, a strong movement 
of the embryo gives decided evidence of the progress of hatching. 
Immediately behind the egg pick, a triangular slit appears. 
through which bubbles emerge to the surface of the exochorion; 
thus the amnion (the embryonic cuticle) makes its initial appear- 
ance through a crack in the exochorion of the shell. 

The amnion splits longitudinally adhering to the chorion 
and bulging out as the insect struggles. It is pushed out of the 
chorion as the abdominal segments are projecting. The emer- 
gence is furthered by the young maggot pressing the head 
against its tail causing the middle of its body to bulge through 
the central gap of the shell. The-amnion becomes noticeably 
darker as it is exposed to the air; it is now a light brownish- 
vellow. 

The pressure of the body against the shell forms nearly a 
round hole through which the larva eventually emerges. The 
opening is enlarged by pressure of the head against the tail, 
raising the body like a hoop, causing an arch or a hump to appear 
with dorsal side outwards. 

The amnion sheds slowly on either side from the middle of 
the arched abdomen ventrally and with. a movement of fluid 
beneath it cracks across the abdomen, peeling and wrinkling as 
the segments telescope. With a final vigorous bubbling and 
wrinkling the amnion sheds off, the moulted skin falling on either 
side, exposing the quite colorless cuticle of the maggot roughly 
wrinkled and bristling with slender hairs. These hairs which at 
first appear transparent turn grayish when exposed to the air. 

When the tail of the larva has been torn loose from the shell, 
the head and thorax are still imprisoned within the egg, necessi- 
tating a maneuvering by waving its tail in the air, twisting and 
squirming while standing on its head. The abdomen doubles up 
ventrally and finally the young larva supported on its tail 
extricates the head by a violent shaking. When the head is torn 
from its fastenings, it is found that the amnion has been holding 
it within the shell. The adhering membrane is cast out when the 
shell is shaken off. The larva has now fully emerged, the colorless 
cuticle has turned grayish and the slender threadlike bristles have 
assumed an iridescent hue. 


TgI0] California Rodent Fleas 73 


The larvae upon hatching busy themselves immediately in 
the quest for food. They experience little difficulty in locating 
it, for at the time of birth a supply of food is found upon the egg 
shell. Here they feed from the first on the tiny blood pellicles 
surrounding the egg shell; this the mother furnishes when the 
egg is laid. When the last of the egg pellicles of blood are con- 
sumed, the insatiable worms look about them for other sustaining 
morsels. The dejecta of the adult flea seems to provide the 
desired ingredients. The young larvae feed ravenously on the 
bloody deposits, apparently satisfied to pass the first few days 
on this unique diet. They can subsist entirely on the bloody 
fragments (flea faeces) when no other food is available, for a 
period of five to six days. 


THE REACTION TO LIGHT. 


The larva is positively heliotropic up to the stage of the initial 
moult. The more advanced sluggish larvae are repelled by the 
light. This is seen when examining the flea breeding cages a 
sight stirring of the nesting material attracts to the surface the 
tiny very active larvae. If the older larvae are desired, it is 
found necessary to nearly invert the container. Prior to the 
final moult when the larva is in readiness to pupate it can be seen 
almost invariably along the edges at the bottom of the box, 
where the greatest number of cocoons are brought to view. 


TROPIC INFLUENCES IN THE ADULT FLEAS. 


Rodent fleas are negatively phototaxic (repelled by light) 
to a very striking degree. The first impulse seems to be to seek 
protection from the light. This is seen in combing a squirrel 
or rat recently killed; the fleas will retreat constantly to the under- 
side, always in the direction away from the light. When shaken 
off, they return to the shadow of the host; in numerous instances 
even when the animal was dead for a period ranging from 24 to 50 
hours the fleas when shaken off would seek the host and bury 
themselves under the hairs away from the light. 

When a number of live squirrel fleas and rat fleas were placed 
in an open test tube and held horizontally with the operator’s 
thumb covering the mouth of the vial and the bottom held 
against the window, the fleas crowded towards the open mouth 
in the direction of the thumb, bounding away from the window 
in an excited manner. When the tube was reversed with the 
open mouth towards the window, it was found unnecessary to 


74 Annals Entomological Soctety of America  [Vol. III, 


plug the mouth of the tube as the fleas did not attempt to jump 
out when given an opportunity to do so. Even when placed 
within a half inch of the open end of the tube with the head of 
the insect turned towards the light, the fleas reversed and 
jumped towards the closed end of the tube. This was repeated 
by tilting the mouth downward to offer an easier exit through the 
open mouth; but even this inducement did not influence the fleas, 
which invariably sought the closed end of the tube in the shadow. 
This was tried with squirrel and rat fleas as well as with human 
fleas and always with the same result. 


LONGEVITY OF THE CALIFORNIA FLEA. 


The great life of our native parasites seems to be anticipated 
at the very incubation. Eggs are laid at all times. We have 
observed oviposition during every month. The broods appear 
to be constant and the number irregular. The incubation stage 
of our Eastern forms taken from the observations of Pergande 
consumes 2—4 days ordinarily. Two days is given as the incuba- 
tion period for the cheopis in India. Our experience with this 
species is a duration varying from 9 to 13 days at room tempera- 
ture. When eggs of this species were subjected to identical con- 
ditions in which wild rats were caged in the laboratory basement 
(at a temperature of 20° to 25° C.) the length of the egg stage was 
1g days. 

Larval life is cited by Eastern and English authorities as a 
minimum of 8 days and a maximum of 24 days. The larval stage 
of our California fleas under laboratory conditions is never less 
than 28 days, often 30 days and sometimes longer. 

Pergande found that the pupal or cocoon stage of the Pulex 
irritans varied from five to seven days in the summer months at 
Washington. Other authors working with this species give 
twelve days during summer months. We have observed several 
instances of cocoon life during the warm days of September. 
Thirty days appear to be spent in this stage. 

The entire life cycle including adult life is given by several 
authors as four to six weeks. The British Indian Plague Com- 
mission gives as the time necessary for the completion of the 
cycle of development (in the case of L. cheopis) that is from the 
egg to the imago, as 21 to 22 days. 

The following is given as a type of the life cycle observed in 
C. acutus. This specimen was kept under observation from the 


Igto| California Rodent Fleas 75 


moment the egg was laid by its parent. An hour after the egg 
was laid it was placed in a small vial with a little sawdust, sand, 
wheat grains and squirrel hair. It was allowed to develop in this 


environment until matured. 


Stage of development Date Age of flea 

Egg laid May 4th, 1909 

Hatched May 12th, 1909 8 days 
Moulted (second stage) May 18th, 1909 14 days 
Moulted (third stage) May 28th, 1909 24 days 
Cocoon June 9th, 1909 36 days 
Adult July 10, 1909 67 days 
Alive (never having been fed) Aug. 11, 1900 99 days 


The English workers in India ascertain the length of time 
which adult fleas live on rats as 41 days. The longest life of this 
species on an exclusive diet of human blood was observed to be 
27 days. We have managed to keep this species (cheopis) alive 
for a period of 36 days by feeding on man. The length of life 
without feeding was also noted. This period varied in the experi- 
ments of the English Commission according to the environments 
in which the fleas were maintained. In the absence of liquid food 
supply, fleas could live in bran for six days, in gunny sacking a 
similar time, and in sand with moist cowdung for 13 days. From 
onr observations we have found the majority of fleas of all species 
to die in five days unless a moist medium was provided. 

As we have stated above rat and squirrel fleas may be kept 
alive for a considerable time when moisture is provided in some 
form. It is interesting to note that fleas which have never 
tasted animal food, having emerged from the cocoon and kept 
under the same conditions in similar material as fleas taken direct- 
ly from the host, will prove longer lived. A number of specimens 
of Ceratophyllus acutus removed from a ground squirrel and kept 
in moistened wheat grains and sawdust lived for 26 days. An 
equal number of fleas of the same species bred from cocoons in the 
laboratory were kept without a host in a similar medium. One 
male lived 38 days and a female lived for 65 days. 

In a series of experiments in which fleas taken from healthy 
rats and squirrels were fed daily on the arm of a man we 
attempted to determine the maximum longevity of these para- 
sites. The fleas were placed individually in open test tubes and 
at feeding time the tubes were inverted over the arm of one of 
the laboratory attendants. Fleas were thus applied daily for a 
period of from 5 to 15 minutes, but only the actual feeding time 
was recorded. The average was about five minutes. 


76 Annals Entomological Society of America  [Vol. III, 


A Loemopsylla cheopis was fed for 35 days, escaping on the 
36th day. One C. acutus died after 58 days and another one at 
the end of 51 days. The common brown rat flea C. fasciatus, 
proved to be the most persistent feeder of them all. Unfortunate- 
ly one of them was permitted to escape after feeding on its 
induced host for 63 days. Another of the group suffered no ill 
effects from its enforced diet for 98 days; and the sole survivor 
had been nourished by its foster host since its removal from the 
rodent host for a period of five months. The experiment was 
discontinued, but the parasite survived a week longer in a 
starved condition. 


LONGEVITY RELATIVE TO SEX. 


We have noted the relative longevity of the sexes under the 
conditions of experiments in which fleas were fed on human 
blood alone. : 

We shall take for consideration the two species common to 
rats, L. cheopis and C. fasciatus and the predominant squirrel flea, 
C. acutus. Four tests with C. fasciatus gave the following data: 


C. fasciatus. 


Six males of this species averaged 8} days, the maximum life 
being 17 days. Fifteen females gave an average of 32 4-5 days 
with a maximum of 125 days. The two unfed controls (male) 
of this series lived for 3 days, and the two control females lived 
for 5 days. 

L. cheopis. 


Two tests with a total of seven males of this species gave 
an average of 10 1-7 days and a maximum period of 15 days. 

Three females lived 28 1-3 days as an average and a maximum 
period of 49 days. 

The two unfed male controls lived 5 days and in four females 
the average was 5} days and the maximum 7 days. 


C. acutus. 


One test with this species furnishes the following data: Three 
males averaged 11 days, and gave a maximum of 11 days. 

Five females averaged 15 1-5 days. The longest life was 53 
days. . 

The unfed males averaged 3 days and showed a maximum of 
4 days. The female control lived 5 days. 


IgIo] California Rodent Fleas Ti 


The greatest length of life of a male of any species is seen to be 
17 days; and the term of life under these conditions for a female 
was 125 days, somewhat over 4 months. 

In these tests for longevity, it should be borne in mind that 
no attempt is made to arrive at the initial age of the fleas, but the 
time is reckoned from the day of removal from the host. 

An effort was made to determine the length of adult life of 
one species, C. acutus, by feeding the insect newly emerged from . 
the cocoon. Ten specimens emerging within a few hours of each 
other were fed on the same day on human blood. One specimen, 
a female, lived for sixty-four days; at which time the experiment 
was discontinued. The unfed controls, as well as the specimens 
given a human blood diet, were kept in ordinary test tubes at 
room temperature. The activities of these fleas may have been 
influenced by changes in the temperature and the absence of 
moisture in the tubes. 


NOTES ON THE FEEDING PROCESS. 


There is a remarkable degree of variation in the feeding habits 
of the different species of fleas. We have not attempted except 
in a superficial way, to study the idiocyncracies of the rodent 
fleas in regard to the biting of their normal hosts. We have, 
however, quite thoroughly observed the manner of biting under 
experimental condition with man as a host. Without taking 
into account the attraction or repulsion which may be exerted 
towards man as a host, we shall consider the more striking features 
of the biting of the parasites. We find that the species do not 
all attack with equal avidity. Pulex irritans the ectoparasite 
of man, is insatiable in its blood craving. It differs in its relation 
to man in being more fastidious in its feeding than the rodent 
fleas. Although its bite is painful, it does not voluntarily feed 
in one spot for any great length of time. The Pulex trritans dif- 
fers from all other species (hundreds of specimens of which were 
tested on human hosts) in that it squirts blood per anum during 
the act of biting. 

The L. cheopis and C. fasciatus, the normal rat parasites, are 
found to bite man with equal readiness and will live about the 
same length of time when fed on human blood. Their biting is 
well defined and effective, but not nearly so painful as that of 
Pulex irritans, nor so prolonged as the common squirrel flea, 
Ceratophyllus acutus. A specimen of C. acutus when starved for 


78 Annals Entomological Society of America [Vol. III, 


several days has been observed to feed on man uninterruptedly 
for a period of nearly one hour (59 minutes) at one insertion of 
its proboscis. 

The bite of the Ctenopsyllus musculi, the blind flea of the 
mouse, is the feeblest we have had inflicted from any flea tested. 
The short weak piercing organs of this species makes a puncture, 
which is scarcely perceptible. In observing the length of time 
this flea bites, it is necessary to depend as a guide on the distention 
of the abdomen with blood rather than the prick of the mandibles. 
The muscult seems not able to adapt itself as an induced parasite 
of man. From a few experiments it appears to live not longer 
than five days on a human host. 

The Ceratopyyllus acutus which proves a very ready parasite 
of man makes its attack even without inducing experimentally. 
Our data compiled from reports and collections of squirrel hunt- 
ers shows that this species, as well as the other common squirrel 
flea, Hoplopsyllus anomalus, will bite man when exposed to their 
attacks. 

The unusually long rostrum in this flea is presumed to be 
the cause of its prolonged feeding at one insertion of the mouth 
parts. This principal is not unprecedented judging from a 
study of numerous parasites in the insect world. 

The following is the description of the method of feeding 
observed in C. acutus. This method is typical. 

The flea when permitted to walk freely on the arm selects 
in a few minutes a suitable hairy space where it ceases abruptly 
in its locomotion, takes a firm hold, with the tarsi, projects its 
proboscis and prepares to puncture the skin. 

A puncture is drilled by the pricking epipharynx, the saw- 
tooth mandibles supplementing the movement by lacerating the 
cavity formed. The two organs of the rostrum work alternately, 
the middle piece boring, while the two lateral elements execute 
asawing movement. The mandibles, owing to their basal attach- 
ments are, as is expressed by the Journal of Hygiene, Vol. 6, No. 4, 
p- 499, ‘capable of independent action, sliding up and down but 
maintaining their relative positions and preserving the lumen of 
the aspiratory channel.’’ The labium doubles back, the V-shaped 
groove of this organ guiding the mandibles on either side. 

The action of the proboscis is executed with a forward move- 
ment of the head and a lateral and downward thrust of the 
entire body. As the mouth parts are sharply inserted, the 


T9To| California Rodent Fleas 79 


abdomen rises simultaneously. The head and middle legs are 
elevated resembling oars. The fore legs are doubled under the 
thorax, the tibia and tarsi resting firmly on the epidermis serve 
as a support for the body during feeding. The maxillary palpi 
are retracted beneath the head and thorax. The labium con- 
tinues to bend, at first acting as a sheath for the sawing mandibles, 
and as these are more deeply inserted, it bends beneath the head 
with the elasticity of a bow, forcing the mandibles into the 
wound until the maxillae are embedded in the skin of the victim. 
When the proboscis is fully inserted, the abdomen ceases for a 
time its lateral swinging. 

The acute pain of biting is first felt when the mandibles have 
not quite penetrated and subsequently, during each distinct 
movement of the abdomen. The swinging of the abdomen 
gradually ceases as it becomes filled with blood. The sting of the 
biting becomes gradually duller and less sensitive as feeding 
progresses. The movements of the elevated abdomen grow 
noticeably feebler as the downward thrusts of the springy bow- 
like labium become less frequent. 

As the feeding process advances, one can discern through 
the translucent walls of the abdomen, a constant flow of blood, 
caudally from the pharynx, accompanied by a peristaltic move- 
ment. 

The end of the meal is signified in an abrupt manner. The 
flea shakes its entire body, gradually withdraws its proboscis by 
lowering the abdomen and legs, and violently twisting the head. 

When starved for several days, the feeding of the rat fleas is 
conducted in a rather vigorous manner. As soon as the proboscis — 
is buried to full length, the abdomen is raised and there ensues a 
gradual lateral swaying motion, increasing the altitude of the 
raised end of the abdomen until it assumes the perpendicular. 
The flea is observed at this point to gain a better foothold by 
advancing the fore tarsi, and, then gradually doubling back the 
abdomen, it turns a back spring, with extreme agility, nearly 
touching with its dorsal side the skin of the hand upon which it 
is feeding. Meanwhile, the hungry parasite feeds ravenously. 

It is interesting to note the peculiar nervous action which the 
rodent fleas exhibit immediately when the feeding process is 
completed, or when disturbed during the biting. Even while 
the rostrum is inserted to the fullest, the parasite shakes its head 
spasmodically, in a twinkling the mouth is withdrawn and the 
flea hops away. 


80 Annals Entomological Soctety of America [Vol. III, 


POSSIBLE VITAL CONSIDERATIONS INVOLVED IN FEEDING HABITS. 


We have previously noted that rodent fleas can live in a 
starved condition, away from the host, during a period of three 
to ten days, when kept in dry test tubes; as long as twenty-eight 
days when a suitable moist medium is furnished. The long 
periods of starvation appear not to affect the vitality of the 
parasite to such an extent that the ability to feed is impaired. 
We have recorded instances in which a specimen of the squirrel 
flea H. anomalus starved for sixteen days, and several specimens 
of C. acutus starved for twenty-seven days had sufficient energy 
to feed 10-15 minutes when an arm was placed in the breeding 
jar. These facts lead us with others to accredit in a measure the 
claims of older authorities on plague, who contended before the 
flea theory was recognized, that clothes and baggage of an 
infected community harbored the germs of plague, which re- 
main viable during long period of time. Modern writers have 
called our attention to the plausibility of infection in clothes and 
baggage due to the agency of fleas, which may be transported 
from infected communities. Bannerman (Journal of Hygiene, 
Vol. 6, 1906, pp. 189-191), writing on the possibility of the spread 
of infection by means of clothes, brings forth several instances in 
which conveyance of infection by clothes seemed the most 
likely means of introduction of plague in some villages in India. 
Plague infected rat fleas secreted in the clothes en route are held 
responsible for the transmission. Bannerman cites an instance 
of plague transmission through clothing where as much as ten 
days elapsed in the transportation of clothing removed from a 
plague infected victim and worn by a relative of the deceased, 
who in turn contracted the disease. 

It has not been our intention to enter into a discussion of the 
flea from the standpoint of plague epidemiology; it is desired 
merely to indicate the possibility that starvation of infected 
fleas may not eliminate the danger of transmission. The presence 
of plague bacilli in rodent fleas as pointed out by the English 
Plague Commission (Journal of Hygiene, Vol. VIII, p. 261) does 
not appear to affect the rate of mortality in these insects. 


Igo] California Rodent Fleas 81 


SUMMARY OF RESULTS. 


rt. The breeding of fleas under laboratory conditions can be 
carried on quite satisfactorily when there is furnished a medium 
simulating the nest of the host. It will be seen that fleas taken 
from the natural host can be kept alive without food for a con- 
siderable time, providing the proper moisture conditions are 
maintained. A medium of moist sawdust with a few grains of 
wheat prolonged the life of these insects to twenty-six days. 
Those without this medium under the same conditions, died on 
the seventh day. 


2. Rat fleas namely, L. cheopis, C. fasciatus can not jump 
higher than 3 1-8 inches. 

P. irritans, the human flea, was found to make a perpendicular 
jump of 7 3-4 inches and as far as 13 inches on a horizontal 
plane. 


3. Experiments with black and white guinea pigs show that 
in relation to color attraction white animals are no more attractive 
to the fleas than are the dark colored ones. 


4. Experiments with adhesive paper baited with meat, 
demonstrate that fleas can not be trapped by the odor of meat. 

5. In the mating of fleas, the female takes the initiative and 
the male assumes the passive role. 

6. Fleas reared from the cocoon kept without a host have 
never been observed to copulate or oviposit. 

7. The eggs are never laid on the host. Oviposition takes 
place within thirty-six hours after the female is removed from the 
host. 


8. The process of hatching consumes a period of 3-4 days. 
The embryo breaks through the shell by means of a special egg 
pick, which is instrumental in the emergence. 

g. The larvae can live on the bloody egg pellicles and the 
dejecta of the parent for a period of 5-6 days. 

to. The larvae are positively heliotropic in the early stages 
and repelled by light in the later stages. 

tr. Adult fleas are negatively phototaxic. 

12. The California rodent fleas have a greater life in all 
stages than fleas of the Eastern United States and India. The 


82 Annals Entomological Soctety of America [Vol. III, 


average length of the stages of development, in C. acutus as a 
type, are as follows: 


Egg stage 8 days 
Larval stage 28 days 
Cocoon 31 days 
Adult without host 32 days and longer 


13. Fleas which have never been fed from the time of 
emergence from the cocoon prove longer lived when starved than 
fleas removed from the host. 

14. Rat fleas may be kept alive on a human host for a 
considerable time, one specimen being kept alive as long as five 
months. 

15. The females of all species of fleas are considerably longer 
lived than the males. 

16. Induced parasitism of rodent fleas on man seems to be 
influenced by the length of mouth parts in the different species. 
One specimen of C. acutus, the species with the longest rostrum, 
fed for a period of nearly one hour at one insertion of the mouth. 

17. It is indicated that starvation of infected fleas, when 
these insects are transported in clothing, may not eliminate the 
danger of transmission of plague. 


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ANNALS 


OF 


The Entomological Society of America 


Volume III JUNE? SEG Number 2 


A STUDY ON THE STRUCTURE OF THE EGG OF THE 
WALKING-STICK, DIAPHEROMERA FEMORATA SAY; 
AND THE BIOLOGICAL SIGNIFCANCE OF THE 
RESEMBLANCE OF PHASMID EGGS TO SEEDS.* 


Henry H. P. SEVERIN, University of Wisconsin. 
(With Plate IX.) 


A detailed description of the eggs of the walking-stick is 
essential because in the systematic work on the Phasmidae, the 
eggs are said to be a very valuable auxiliary means of identifi- 
cation, since their form, “‘ftir die Genera charakteristisch ist.”’ 
(Brunner von Wattenwylt). Kaup (8) even remarks, ‘‘Viel- 
leicht wird man spater die Arten durch die Eier schneller unter- 
scheiden lernen als durch die Thiere selbst!”’ 

Measurements of Eggs: Heymons (6) found with Bacillus 
rossi that the size of the eggs is not constant. ‘‘Es sind mir 
Eier zugekommen, wahrscheinlich die letzten, welche das Weib- 
chen abgesetzt hat, die hochstens die Halfte der tblichen Grésse 
besassen, trotzdem waren sie aber normal gebaut und es sind aus 
ihnen ebenfalls Larven ausgeschlupft.’’ In Diapheromera fem- 
orata a considerable variability in the dimensions of the eggs is 
also present but the smaller eggs were not necessarily the last 
ones which the female lays,—they may appear among the first 
or at any time during the egg-laying. 

The size of the eggs varies between the following dimensions: 


Verity 6 58-2 ye Oe oe 20-29 mm. 
Depth, from dorsal to ventral surface........11-18 mm. 
Width or thickness from side to side.......... 9-14 mm. 


* Submitted as a part fulfillment of the thesis requirements for the 
degree of Ph. D. at the University of Wisconsin. 


+ Paper not accessible. Quoted from v. Brunn’s (2) paper: . 


83 


84 Annals Entomological Society of America [Vol. III, 


The majority of eggs varied between the following dimensions: 


| Voie) 04 7]: A Pn Mme at cm Se. 
WEIR. iss oc SE ERO ee ee 
LG 5 1: rer ren he Pea Ss rh 


Shape and Color of Egg: The eggs resemble very small beans 
with some variation in shape; some are ellipsoid, others ovoid 
and still others nearly spherical in form. Their color is usually 
a glossy black, except on the slightly more convex side which is 
white; instead of the black, however, there may be light shades of 
gray or light chocolate-brown. Out of a thousand eggs, twelve 
showed the light shades of gray and only three, the light chocolate- 
brown color. 

When the operculum is removed the egg, in this region, is 
obliquely truncate and surrounded by an elliptical or oval rim. 
On the inner margin of this rim rested the operculum. The rim 
is provided with a circle of yellow, chitinous, bristle-like projec- 
tions. In most eggs the white color of the slightly more convex 
side is continued around the base of the rim as a white line. 

Operculum: The operculum (Fig. 1, op) fits perfectly within 
the rim of the egg capsule (Fig. 1, 77) and is usually set free when 
the egg is broken. If the operculum is cleared, mounted and 
examined under the microscope a ragged membrane is discern- 
ible at its margin (Fig. 3, vz). This is the torn, so-called ‘‘shell 
membrane”’ to which the operculum was attached. 

Sharp (16) who has described a number of Phasmid eggs, has 
probably overlooked the fact, that the operculum is attached to 
the so-called ‘‘shell membrane.’ He writes, the operculum 
‘is present in all known eggs of the Phasmidae; it is a lid that fits 
very accurately to the truncate anterior extremity of the egg; 
its margin is surrounded by a margin of the capsule, and it is ow- 
ing to the perfect fit between the two that the operculum retains 
its position.”’ Muller (13), however, in the case of Phasma 
ferula claims that, ‘‘Am Rande des Ausschnitts springt die innere 
Schalenhaut etwas vor, ein Rudiment der Verbindung der 
Schale mit dem Deckelchen. Die innere Haut des Deckelchens 
und die innere Haut der Schale sind also an dem unverletzten 
41 ein Continuum.”’ Leuckart (10) who has worked on the struc- 
ture of the eggs of two species of Phasmids belonging to different 
genera also finds that, ‘‘Durch Hilfe dieser Schalenhaut wird der 
Deckel, der sonst vollkommen isolirt ist, in seiner Lage erhalten 
und befestigt.”’ 


Igo] The Egg of the Walking Stick 85 


A number of zoologists who have worked on the structure 
of the eggs of the Phasmidae claim that a “‘shell membrane’ 
exists within the inner surface of the chorion. Muller (13) writes, 
‘Die innere Flache der Schale wird von einer sehr dtnnen hau- 
tigen Lamella, der Schalenhaut, uberkleidet, die sich nur in kurzen 


Stticken von der Schale selbst wegnehmen lasst.’’ Leuckart (10) 
agrees with Muller that a ‘‘shell membrane” lines the inner 
surface of the chorion. ‘‘Dazu kommt als Bekleidung der innern 


Chorion-flache noch eine eigne dtinne ‘Schalenhaut,’ die schon 
von J. Miller aufgefunden ist, also wahrscheinlich unter den 
Phasmoden eine ziemlich allgemeine Verbreitung hat, obgleich 
sie den tbrigen Insekteneiern abgeht.”’ In the eggs of Diaphero- 
mera femorata the so-called ‘shell membrane”’ is the innermost 
layer of the chorion, which peels off in small fragments in the 
eggs that are in the early stages of development, but in the later 
stages, as Leuckart (10) has observed, “‘Bei Cyphocrania lasst 
sich diese Haut ohne grosse Schwierigkeiten in continuo abheben 
—hbis auf die Narbe, an der dieselbe fest mit dem Chorion ver- 
wachsen ist und ein weisses Aussehen hat.’ To Leuckart’s 
observation may be added, in the case of the eggs of Diaphero- 
mera femorata, that a firm attachment also exists at the rim 
between the so-called ‘‘shell membrane’’ and the next outer 
layer of the chorion. 

A microscopical examination of the operculum from the 
outer surface shows a resemblance to the framework of a dome, 
which is shut off at the base by a slightly concave floor of chitin. 
The framework consists of brownish, irregularly-flattened, chi- 
tinous rods. All of these rods arise from an elliptical or oval 
brownish rim of chitin (Fig. 3, br) which is in continuation with 
the similarly colored upper surface of the floor of the dome 
(Fig. 4, e). Some of the rods anastomose, enclosing a large, more 
or less, central space at the top of the dome (Fig. 3, c) and a 
varying number of irregular areas (Fig. 3, d) which are not con- 
stant in number in the different opercula. Those rods which do 
not extend to the top of the dome project free into the irregular 
spaces (Fig. 3, /). 

Various authors have called attention, in different Phas- 
midae, to the resemblance in the histological structure of the 
egg capsule to the structure of certain plant tissue. Murray (14) 
finds, “‘a most striking resemblance to a piece of honey comb”’ 
in the structure of the egg capsule of Phyllium Scythe. Accord- 


86 Annals Entomological Society of America [Vol. III, 


ing to Joly (7) who has worked on the structure of the egg of 
Phyllium crurifolium ‘‘ce tissu présente la plus grande analogie 
avec celui du liége, c’est-a-dire qu’il est formé de cellules irrégu- 
liéres (carrées, pentagonales, sexagonales) trés petites et trés 
serrées. La couche extérieure qui recouvre cette coque, est 
beaucoup plus épaisse et, comme nous l’avons dit, elle ressemble 
a lecorce rugueuse du Chéne-li¢ge, dont elle a la légéreté. Nou- 
velle et curieuse analogie de notre insecte avec le régne végétal: 
analogie qui devient plus complete encore, quand on songe que 
VYoeuf du Phylliaum est muni d’un opercule qui s’ouvre lors .de 
l’éclosion, & la maniére d’une pyxide.’’ Brongniart* who has 
examined the eggs of Phyllium pulchrifolium also compares the 
external envelope to that of cork. According to Henneguy (5) 
who has also worked on the histological structure of the egg- 
capsule of Phyllium crurifolium, ‘‘L’ensemble de la couche 
externe présente une. grande analogie de structure avec la partie 
libérienne d‘une écorce de dicotylédone traversée par les rayons 
médullaires.”’ 

A cross section through the operculum of the egg of Diaphero- 
mera femorata shows that the space between the floor and top of 
the dome is partially filled with chitinous deposits, which, accord- 
ing to a vegetable histologist, resembles somewhat the thin 
walled parenchyma of plant tissue, except that no middle lamella 
was discernible (Fig. 4, a). Muller (13) describes the chitinous 
deposition as ‘‘ein zelliges Gewebe aus deutlich sechseckigen oben 
offenen Zellen bilden. Dies scheint einer besonderen Beachtung 
werth, da die Ercheinung eines regelmassigen zelligen Gewebes in 
der Organisationsstufe der Insecten gewiss die seltenste ist.”’ 
It would be rash to speculate as to the formation of this chitinous 
deposition since the manner in which the operculum is formed 
is obscure. Sharp (16) suggests two methods: ‘“‘first, autotomy 
of the pole of the egg; second, adhesion of the mass of matter 
from the adjacent nutrient chamber, to form as it were a very 
imperfect second egg.”’ 

‘““Hilar area, cicatricula or Narbe” of Egg: On the outer 
surface of the exochorion, of the slightly more convex side of 
the egg is an elliptical region resembling somewhat the hilum 
of a seed. Sharp (16) calls this region the “‘hilar area,’ Miller 
(13) the ‘‘cicatricula’”’ or ‘‘Narbe,”’ Leuckart (10) and Heymons 
(6) also use the term ‘‘Narbe.’’ The margin of this ‘‘hilar area”’ 


* Paper not accessible. Quoted from Henneguy’s (5) paper. 


Igo] The Egg of the Walking Stick 87 


is slightly elevated and buff-colored (Figs. r and 8, m). At the 
posterior end, this margin narrows enclosing a semicircle (Figs. 
t and 8, s) within which lies the micropylar orifice (Figs. 1 and 
8, 0). The margin of this semi-circular area is in continuation 
with a ridge which extends towards the posterior pole of the egg. 
The buff-colored ridge joins a triangular extension of the black 
region of the egg (Fig. 1, 7) and gradually disappears as it passes 
posteriorly into the surrounding black surface. Within the 
slightly elevated buff-colored margin is a white convex region 
which terminates at the posterior end in the microypylar orifice, 
where the white color gives way to black (Figs. 1 and 8, 0). 

Mycropylar Apparatus: Miller (13) did not understand how 
eggs of insects with a hard chorion were fertilized and he takes 
an extreme view as to the way this phenomenon takes place 
in the eggs of the Phasmidae. ‘‘Bei den Phasmen hat die Samen- 
kapsel ausser ihrer einen Oeffnung nach aussen keinen beson- 
dern Ausfthrungsgang. Aber der Eingang der zweihdrnigen 
Samenkapsel liegt gerade ttber dem langlichen Ausgang des Eier- 
gangs. Der Samen tritt also aus der Eingangsoffnung der Kap- 
sel unmittelbar in der Mundung des Eierganges ein, um sofort 
zu den Eierleitern und Trompeten zu gelangen.”’ 

‘Wir haben aber bewiesen, das eine Befruchtung der Phas- 
meneier nicht anders, als vor ihrer Ausbildung und namentlich 
vor der Ausbildung der Schale, méglich sey.”’ 

The micropylar apparatus is very remarkable and differs 
from any of those which Leuckart (10) has described for so many 
insect eggs. As already mentioned the micropylar orifice is 
found at the posterior end of the convex area, just within the 
space enclosed by the semicircular chitinous thickening of the 
margin. This opening leads into a small canal, the micropylar 
canal, which passes a short distance towards the anterior pole of 
the egg. The micropylar canal is elliptical in cross section and 
surrounded by extremely thick chitin (Fig. 5, g). When the 
‘inner surface of the ‘‘hilar area”’ in the region of the micropylar 
apparatus is examined under a microscope, an invagination of 
the inner surface of the chorion is readily seen: at the bottom 
of this inpushing is the opening of the micropylar canal 
(Figs. 6 and 8, 2). If the vitelline membrane in this region is 
now examined under a binocular, one finds that a small, obliquely- 
inclined, membranous tube has been torn away from the opening 
of the canal (Fig. 2, t). A glance at Fig. 7, v, shows the opening 


88 Annals Entomological Society of America [Vol. III, 


of this membranous tube, the vitelline membrane micropyle, 
into the vitelline membrane. The sperm thus enters the micro- 
pyltar orifice, passes through the micropylar canal, then through 
the membranous tube and out of the vitelline membrane 
mycropyle to reach the egg. 

Miuller’s. (13) supposition that the sperm passes from the 
seminal receptacle, through the common oviduct, then into the 
oviducts to the ovarian tubule and fertilizes the egg before the 
chorion is formed is entirely erroneous. Leuckart (10) from 
observations on Gomphocerus found that the micropyles, ‘‘nicht 
von Anfang an dem Chorion zukommen, sondern erst nach der 
Ablagerung desselben durch Resorption ihren Ursprung neh- 
men * * * * _ Eine Bestatigung dieser Beobachtung finde ich 
darin, dass ich nicht selten (wie u. a. bie Borborus, Tetanocera 
und verwandten Fliegen) Eierstockseier antraf, deren Micro- 
pylapparat noch ohen Oeffnung war, sonst aber bereits vollkom- 
men entwickelt schien.”’ 

‘Vor der Ablagerung des Chorions habe ich an der Dotter- 
haut niemals eine Micropyle wahrgenommen * * * * . Wohl aber 
habe ich Falle beobachtet, in denen bei Anwesenheit der Chorion- 
micropyle die Dotterhaut noch ohne Loch zu sein schien.”’ In 
all probability the eggs of the walking-stick are fertilized as they 
pass below the opening of the seminal receptacle. 

The Biological Significance of ihe Resemblance of the Phasmid 
E-ggs to Seeds: A number of naturalists have called attention to 
the striking resemblance of the Phasmid egg to a seed. In 
some cases, the egg resembles the seed of the natural food plant 
of the insect. 

In seventeen species of Phasmids obtained from Lifu and New 
Britan, Sharp (16) has described the eggs of a number of these. 
In regard to the resemblance of the eggs to seeds he writes: ‘“‘The 
climax of the peculiarities is found in the extremely perfect 
structure of their eggs and the resemblance of these eggs to seeds. 
The egg of the Phasmid has not only a general resemblance in 
size, shape, colour, and external texture to a seed, but the ana- 
tomical characters of certain seeds are reproduced on the external 
surface, there being a hilar area, a hilar scar, and a capitulum 
corresponding to the micropylar caruncle of such seeds as those 
of the Castor-oil plant (Ricinus communis). The hilar area on 
the inner surface of the capsule is, in shape, like the embryo of 
a plant. Moreover, naturalists who have examined these eggs 


Tgto] The Egg of the Walking Stick 89 


declare that the minute structure of this curious egg-capsule 
cannot be distinguished histologically from plant-structure.”’ 

Among the leaf-insects the resemblance of the eggs to seeds 
is especially marked. Nab* as early as 1854 has compared the 
egg of one of these leaf-insects to the seed of the ‘‘Belle-de- 
Nuit (Mirabilis Jalapa).”” Henneguy (5) in the case of the egg 
of Phyllium crurifolium claims that, ‘‘Sa forme est celle d’un 
akene d’Ombellifere, et représente, par example, la moitié d’un 
jeune diakéne de Conium maculatum. L’oeuf différe de l’akéne 
d’Ombellifere en ce que son opercule réguliérement conique, est 
situé au centre de sa face supérieure, tandis que le style conoide 
de Vakene est aplati sur la face commissurale.’”’ Morton (12) 
in this same species writes that the ‘‘egg has been confused with 
a seed of Mirabilis and Coniom!”’ 

According to Stockhard (18) the walking-stick, Aplopus 
mayeri, is found only on its food plant, Suriana maritima. ‘‘While 
one may find a close resemblance in size and color between the 
eggs of Aplopus and the seed of Suriana, both of which fall 
from the branches to the ground, where they are obscured among 
the débris” yet the eggs differ from the seeds considerably in 
shape. 

The eggs of Diapheromera femorata were shown to a number 
of botanists, and with one exception, all mistook them for seeds. 
The botanist who did not fall into this error broke the egg 
before giving his opinion. When asked as to what seed the egg 
resembled all failed to recall any particular one. A leguminous 
seed, such as a small bean, was suggested, but none could be 
found in the natural habitat of Diapheromera, that resembled its 
egg in size and color. 

If these botanists are unable to distinguish the egg of a walk- 
ing-stick from a seed, can a grain-eating bird distinguish between 
the two? Ifa bird were to feed on the seeds of the Suriana mari- 
tima which resemble the eggs of Aplopus mayeri in size and color, 
could the bird discriminate between the two, on account of a 
difference in shape? 

Goldi (4) in the case of the eggs of two Brazilian walking- 
sticks raises the question as to how far this imitation is useful 
in the protection of these eggs. Grain-eating birds may eat the 
egg but general insectivorous birds would probably mistake it 
for a seed and leave it untouched. The protective dress of the 


* Quoted from Henneguy’s (5) paper. 


go Annals Entomological Society of America  [Vol. III, 


egg may be only a relative protection in which new dangers are 
involved. He next raises the argument that these eggs on ac- 
count of the resemblance to some Brazilian seeds, deceive, so he 
believes, the egg-parasites. . 

This would imply that through the sense of sight the egg- 
parasite would overlook the eggs on account of their resemblance 
to some Brazilian seeds. It is open to serious question whether 
the egg-parasites are guided to the eggs of their host through 
the sense of sight alone, if at all. One illustration will suffice. 
We (15) have found that Trichogramma pretiosa parasitizes the 
eggs of Cimbex americana, and yet the egg-parasite cannot see 
the eggs of its hosts as Cimbex deposits its eggs in a receptacle 
within a willow leaf. In all probability, the sense of smell plays 
an important role in guiding the egg-parasites to the eggs of their 
hosts. 

A careful survey of the literature shows that the eggs of Phas- 
mids are subjected to the attack of a number of enemies. Accord- 
ing to King (9) the eggs of Anisomorpha buprestoides ‘‘are vic- 
timized in a similar manner by a minute species of Ichneumon 
fly, one of which has fortunately been obtained; it is probably 
one of the Chalcididae: all the transformations take place within 
the egg, and when fully developed the perfect Ichneumon fly 
emerges therefrom.” 

Bates (1) ‘‘observed that the author of the note was probably 
in error in attributing the name of Anisomorpha buprestoides 
to the species in question, which seemed to be a true Phasma.”’ 

Smith (17) ‘‘remarked upon the peculiarity of all the trans- 
formations of the Chalcidite parasite taking place within the 
egg of the Phasma; such a mode of development was novel, if true, 
but he suspected some error of observation.” 

M’Lachlan (11) ‘‘suggested that the cocoon of the Chalcis 
had been mistaken for the egg of the Phasma.” 

von Brunn (2) records the observation of Wolff von Wilfing, 
that the young larvae as well as the eggs have many enemies, 
‘“‘hauptsachlich Springspinnen und Hauseidechsen.”’ 

Girault (3) in his paper on the ‘‘Hosts of the Insect Egg- 
Parasites in North and South America’’ does not record an 
egg-parasite from any Phasmid. 

I think that the resemblance of the eggs to seeds has no bio- 
logical significance as a means of protection against the egg-para- 
sites, if the eggs of the Phasmidae are parasitized. Sharp (16) 


Igo] The Egg of the Walking Stick] gl 


in 


all the species which he has examined believes that these 


resemblances in the eggs have no bionomic importance for the 
species and I am strongly inclined to accept his view in the case 
of the egg of Diapheromera femorata. 


I am indebted to my teacher and friend, Prof. Wm. S. Mar- 


shall, for many valuable suggestions in this work and the more 
than ordinary courtesies extended to me in the use of his excel- 
lent entomological library. 


Zoological Laboratory, University of Wisconsin, Madison, February 1, 1910, 


BIBLIOGRAPHY. 


. Bates, H. W., 1867. [Remarks on King’s (9) paper.] Journ. of Proc. of 
. 80. 


the Ent. Soc. of London. V, p. 8 


. v. Brunn, M., 1898. Parthenogenese bei Phasmiden, beobachtet durch 


einen tiberseeischen Kaufmann. Mittheil. aus dem Naturhist. 
Museum, Hamburg. XV. pp. 147-161. 


. Girault, A., 1907. Hosts of Insect Egg-Parasites in North and South 


America. Psyche XIV, pp. 27-39. 


. Goldi, E. A., 1886. Die Eier zweier brasilianischen Gespenstheuschrecken. 


Zool, Jahrb. Syst. 1, pp. 724-729. 


- Henneguy, L. F., 1890. Note sur la Structure de l’Enveloppe de l’Oeuf 


des Phyllies. Bull. Soc. Philom. Paris II, pp. 18-25. 
Summary in: J. R. Micr. Soc. 1890, p. 596. 


. Heymons, R., 1897. ~ Ueber die Organisation und Entwickelung vou Bacillus 


rossi Fabr. Sitzb. d. Kgl. Akad. d. Wiss. Berlin. XVI, pp. 363-374. 


. Joly, M., 1871. Contribution A Vhistorie naturelle et 4 lanatomie de la 


mouche-feuille des iles Seychelles. Mem. Soc. Toulouse, III, pp. 1-30. 


- Kaup, J. J., 1871. Ueber der Eier der Phasmiden. Berl. Ent. Zeitschr. 


XV, pp. 17-24. 


. King, C. B., 1867. Anisomorpha buprestoides. Journ. of Proc. of the 


Ent. Soc. of London. V, pp. 78-80. 


. Leuckart, R., 1855. Ueber die Micropyle und dem feinern Bau der Schal- 


enhaut bei den Insekteneiern. “Miiller’s Archiv. f. Anat. Physiol. 
pp. 90-264. 


. MWLachan, R., 1867. [Remarks on King’s (9) paper.] Journ. of Proc. of 


the Ent. Soc. of London, V, p. 80. 


. Morton, W., 1903. Notes sur l’elevage des Phyllies. Bull. Soc. Vaud. 


Sc. Nat. XX XIX, pp. 401-407. 


3. Miller, J., 1825. Ueber die Entwickelung der Eier im Eierstock bei den 


Gespenstheuschrecken und eine neuentdeckte Verbindung des Riick- 
engefasses mit den Eierstécken bei den Insecten. Nov. Act. Acad. 
Caes. Leopold, XII, pp. 555-672. 


. Murray, A., 1856. Notice of the Leaf-Insect (Phyllium Scythe). Edin- 


burgh new philosophical Journ. n. s. III, pp. 96-111. 


5. Severin, H. H. P. and Severin, H. C. M., 1908. Habits of the American 


Saw-fly, Cimbex americana Leach, with Observations on its Egg- 
Parasite, Trichogramma pretiosa Riley. Trans. Wis. Acad. Sci. Arts 
and Letters. XVI, pt. I, pp. 61-76. 


. Sharp, D., 1898. Account of the Phasmidae with Notes on the Eggs. 


Willey, Zool. Results, Cambridge. pp. 75-94. 


. Smith, F., 1867. [Remarks on Ki g’s (9) paper.] Journ. of Proc. of the 


Ent. Soc. of London, V, p. 80. 


. Stockard, C. R., 1908. Habits, Reactions, and Mating Instincts of the 


Walking-Stick, Aplopus mayeri. Publ. No, 103. Carnegie Inst., 
Washington. pp. 43-59. 


92 Annals Entomological Soctety of America [Vol. III, 


EXPLANATION OF PLATE IX. 


All figures were drawn with a camera lucida. 


Fig. 1. ‘‘Hilar area” on the slightly more convex side of the egg: op, 
operculum; rt, rim in which the operculum fits perfectly; m, margin of the 
**hilar area’’; s, semicircular margin; r. buff-colored ridge which passes over 
into the triangular extension of the black region of the egg; 0, micropylar orifice. 

Fic. 2. Vitelline membrane of egg, after the chorion of the “‘hilar area”’ 
has been removed: ¢, membranous tube, which connects with the opening of 
the micropylar canal on the inner surface of the chorion. 

Fic. 3. Operculum from the outer surface: vt, torn so-called ‘‘shell mem- 
brane” to which the operculum was attached; br. brownish rim of chitin from 
which the irregularly-flattened chitinous rods arise; c, large, more or less, central 
space at the top of the operculum enclosed by the anastomosing rods; d, irregu- 
lar areas which are not constant in number in the different opercula: p, rods 
which project free into these irregular spaces. 

Fic. 4. Cross section of a part of the operculum: e, brownish upper surface 
of the floor of the operculum; a, chitinous deposits, between the floor and top 
of the operculum, resembling somewhat the thin-walled parenchyma of plant 
tissue except that no middle lamella is discernible. 

Fic. 5. Cross section of the micropylar canal g, which is surrounded by 
extremely thick chitin. This canal opens to the exterior by the micropylar 
orifice. 

Fic. 6. Posterior region of the inner surface of the “‘hilar area”’ of the 
chorion: 4, opening of the micropylar canal in an invagination of the inner 
surface of the chorion; 0, micropylar orifice. 

Fic. 7. Vitelline membrane in the region of the micropylar apparatus: 
v, opening of the membranous tube, the vitelline membrane micropyle, into the 
vitelline membrane. 

Fic. 8. ‘‘Hilar area” on the outer surface of the exochorion; lettering as 
in Figs. 1 and 6. 


ANNALS E. S. A. 


VoL. III, PLatr 1X 


I~‘ e, 
Ped te 


3. ase 
= Ree | 
ee eae | 
ARTA | 

5 Aree { 
PILES AT ETD } 
GEA oe 


H. H. P. Severin. 


A STRUCTURAL STUDY OF SOME CATERPILLARS.* 


By Wi.i1AMmM T. M. Forses, 


Clark University, Worcester, Mass. 


Although the Lepidoptera have been studied for a long time 
little attention has been paid as yet to the minute descrip- 
tion of their caterpillars. In describing a caterpillar most 
authors have contented themselves with the colors and markings, 
adding an occasional note on the structure if especially striking. 

My problem, therefore, has been to discover external struc- 
tures characteristic of the groups of caterpillars, especially such 
as have been less carefully studied by others. This paper then 
deals mainly with the parts and setae of the head. I have also 
summarized my studies on the prolegs, and have included some 
of Dyar’s characters derived from the body-setae. I have tried 
so far as possible, to lay emphasis also on such characters as could 
be determined from the cast skin. It appears that, with a little 
care since they are brittle, the characters of the head can be made 
out quite as easily from cast skins as from killed material. The 
body can not be studied quite so well, but at least the arrange- 
ment of the proleg hooks, and also the type of vestiture, whether 
primary, secondary or tufted, can be made out without trouble. 


MATERIAL AND METHODS. 


My material is made up of specimens representing about 125 
genera, preserved in various ways. The major part were col- 
lected personally in the course of the past summer and preserved 
in alcohol. There were also a few cast skins, and a few caterpil- 
lars dried whole without any preparation. To this I added a num- 
ber of species from the American Entomological Company, which 
were preserved in formalin, and a series of inflations, mostly of 
Microlepidoptera, from Staudinger and Bang-Haas. 

Most of my alcoholic material was cleared by boiling in 10 per 
cent. caustic soda, cutting the skin down one side, and separating 
the maxillae and labium from the rest of the head. The dried 
caterpillars were treated in the same way. Often one mandible 
was also removed to give a clearer view of the labrum. The 
prepared skins were then preserved in alcohol, except a few that 


_._* A Dissertation submitted to the Faculty of Clark University, Worcester, Mass., in partial 
fulfillment of the ig heist for the degree of Doctor of Philosophy, and accepted on the recom- 
mendation of C. F. Hodge. 


94 


1910] A Structural Study of Some Caterpillars 95 


were mounted in Farrant’s gum-glycerine. The same method 
was tried on formaldehyde specimens, but with much less success, 
as the muscle would not dissolve easily and had to be picked out 
piecemeal. 

The remainder of the specimens were examined entire, as 
opaque objects. By placing them in full sunlight, or even con- 
centrating it on them with a lens, they could be examined suc- 
cessfully, even with rather high powers of the microscope. A 
binocular microscope was very useful, especially in getting a cor- 
rect idea of the relations of parts, in dissecting, and in hunting 
for setae. 

Inflations of the smaller species were sometimes immersed 
in xylol, or in absolute alcohol, when they could be examined by 
transmitted light. They did not collapse on being taken out and 
dried again. 

Most of the drawings were sketched with the camera lucida, 
the details being put in free-hand under a higher power. They 
were not drawn to a single scale, as they differ a great deal in 
size, and the size is of little classificatory value. 

The purchased specimens were received with names. In the 
case of the others, which were named by the writer, there was 
often some uncertainty, as indicated by question marks in the 
text. This was especially the case in the Noctuidae and Geome- 
tridae, where I have listed less than half of my specimens. 


** * *K K * * * ok 


I wish to express my thanks to the many who have made this 
study possible by their help in supplying specimens, in allowing 
the use of instruments and books and by advice and information. 

Wo. T. M. Fores. 


STRUCTURE. 


A caterpillar is an insect larva in which the thorax and abdo- 
men are similar in general appearance, the head alone is heavily 
chitinized; there is no sclerite between the antennae and the 
mandibles; the mouthparts are small and largely retractile, with 
lacinia not recognizeable as such and no glosse or paraglossz. 
The whole structure of the maxilla and labium is peculiar and 
not closely paralleled in other orders. 

Most of the head as seen from in front, is composed of the 
two epicrania (compare Fig. 1), each of which usually bears eleven 
primary setae and the eyes. There are also almost always 


96 Annals Entomological Society of America  [Vol. III, 


present additional setae or punctures, mostly in the neigh- 
borhood of setae i and ii. (One pair of punctures is shown in 
the figure) .* 

Lying above the mouth and separating the epicrania is a 
triangular sclerite, the front. It has two sete near its lower outer 
angles, and usually between them a pair of punctures. In 
Hepialus, however, the sete lie between the punctures. The 
front and the epicrania are separated by a pair of narrow adfrontal 
sclerites, each of which has two setz and a puncture between 
them.+ 

Bordering the lower side of the front is the clypeus, with a 
pair of setae at each end; and hanging from that, the movable 
labrum, which belongs functionally to the mouth. Its structure 
will be mentioned with that of the mouthparts. 

On each side, between the two articulations of the mandible 
is the antenna. (Fig. 29.) It is four-jointed, but all but the 
second of these are minute. There is a large membranous cone 
at the base, which Scudder treats as a true joint. It is inverted 
when the antenna is retracted, and would seem to be no different 
from the membrane which separates all the joints of the antenna. 
The antenna is surprisingly constant in its structure, the arrange- 
ment of setae shown in the figure occurs with little change in all 
the caterpillars studied. The Lasiocampidae alone have a few 
secondary sete. 

The mouth parts are the labrum, or upper lip (Fig. 12), the 
two mandibles (Fig. 28), and the lower lip (Fig. 2), which in 
caterpillars is formed of the maxillae as well as the labium. The 
labrum is oblong, wider than high, with the free edge rounded, and 
with a notch, through which the food is guided into the mouth. 
It bears (1) a row of four sete across the top of the notch (1 and 11), 
(2) two pair of sete on the lateral edge (iii and iv), (3) two pair 
of setae on the tip of its two lobes (v and vi). There are several 
punctures, the most noticeable of which is the one indicated in the 
drawing and marked za. On the inner side there are a large 
number of sensory cones, and near the outer edge three larger 
similar cones. The latter are indicated in outline on the figure. 
The labrum has been especially useful in furnishing characters- 


+ The front, as the term used in this article, has been usually called the clypeus, the adfrontals 
being known as paracylpeals, and the clypeus as anteclypeus or epistoma. I believe that the set 
of names used here agrees better with their homologies in other orders. 

* Dyar’s numbering of the setae, which has been copied in Fig. 1, is published in Journ. of the 
N. Y. Entomological Society IV, 93, with a figure. 


Tgto} A Structural Study of Some Caterpillars Q7 


It varies considerably in the position of its seta, and is flat, not 
easily distorted, and easily studied under the microscope. The 
setae which appear on the margin in figure 12 have often migrated 
inward a distance on the surface of the labrum, and in different 
groups it is different setae that have migrated. As 1s often the 
case these differences define smaller groups in the butterflies, 
than in the moths. For example, vi has migrated up toward u 
in the Sphingide, in Apatelodes, which shows other resemblances 
to the Sphingidae also; and in a part only of the Nymphalinae 
among the butterflies. (Figs. 14 to 18). 11 migrates inward in 
the Pierids. Another striking arrangement occurs in the Skip- 
pers, as shown by Figs. 23 and 24. 

The mandibles are the heavy jaws, and are the only ones used 
for biting. They bear two setz on the outer side and the edge is 
more or less notched, the notches of the two sides fitting into 
each other. Taxonomically they seem more characteristic of 
genera than of larger groups. The position of the distal seta 
far out on the scrobe, as shown in the figure, is typical of the 
Sphingide. The Arctideze show interesting variations in struc- 
ture of the mandibles. 

The labium forms the middle of the hind margin of the 
mouth. It is roughly cylindrical. The basal part is formed by 
a long, usually lightly-chitinized joint, the mentum, which bears. 
two setae near its middle. Resting on that is the heavy ring- 
shaped sclerite, a, whose setz are shown in the figure as free 
in the membrane distad of the sclerite. This is often the case. 
The tip of the labium is retractile, and therefore largely formed 
of membrane (see Fig. 3). Projecting from its center is the 
cylindrical or flattened spinneret, which is of membrane, strength- 
ened by three chitinous bands. Surrounding the base of the 
spinneret is the ring, c, composed of an inner and an outer semi- 
circular sclerite. It bears a puncture at each side, and may 
be either very wide and heavy as in the Psychidae, or reduced 
to a narrow ring about the puncture, as in typical Noctuidae. 
The labial palpi spring from an area of membrane on each side 
of c. There is one large and one minute joint, each bearing a 
seta. In Catocala there is also a rudimentary basal joint. The 
remaining sclerite, b, of the labium forms a semicircle about the 
base of the palp. It bears two punctures at its ventral end. 

The submentum is divided into a pair of triangular sclerites 
at the base of the labium. These are usually separated by the 
base of the mentum. (Compare Figs. 2 and 25). 


98 Annals Entomological Society of America [Vol. III, 


All the parts of the labium and maxillae are described as if 
looking at the exposed surface from below or from behind accord- 
ing to the position in which the caterpillar holds its head. This 
surface morphologically consists of the caudal aspect of the 
labium, and the caudo-lateral aspect of the maxillae. 

The maxillae are fused at the base with the labium, but are 
free at the tip. The main part is made up of a usually lightly 
chitinized joint, the stipes (plus the palpifer) which bears two 
setae at its distal end. The cardo forms a small triangle between 
the base of the stipes and the submentum. At the tip of the 
stipes there are two very incomplete rings, which would seem 
to be the basal joints of the palpus, but which are completely 
fused with the maxilla proper. In Hepialus the more distal of 
these is shaped like a normal joint of the palpus, and in Microp- 
teryx, according to Packard, it is free. From the end of these 
arises the two-jointed free part of the palpus, and mesad of this 
springs the large basal joint of the galea, which forms an incom- 
plete ring. The tip of this joint bears the two maxillary lobes 
(which are the distal joints of the galea and lacinia?’), and is 
also armed with two large cones (anterior to the maxillary lobes 
in Frenate, and mesad to them in Hepialus), two small cones 
(posterior to the maxillary lobes) and a step-cone between them. 
The latter is composed of a larger chitinous ring, bearing a thin- 
walled cone at its tip. Each of the maxillary lobes, also, bears 
a sense-cone at its tip. 

Chitinous sclerites are but little developed as a rule on the 
body. The greatest development that I have seen on the abdo- 
men occurs in Incurvaria. In this caterpillar each segment has 
two dorsal and two ventral plates (see Fig. 34). The anterior 
ventral plate, or sternum, extends between the prolegs. 

The thorax has more extensive sclerites in Adela, as the 
sketch of the pro- and meso-thorax (Fig. 35) shows. Not many 
of the sutures are traceable, so that they can be only roughly 
homologized with those of other insects. The praesternum and 
sternellum of the prothorax are more distinct. 

In higher caterpillars only the coxz, which are divided by a 
clear suture into coxa and meron, are preserved. Besides the 
coxa, the legs have very short femurs and tibiz and a single- 
jointed tarsus, which bears one claw. 

One of the chief characteristics of the caterpillars is the 
definite arrangement of setae. These are either primary, which 


T9I10] A Structural Study of Some Caterpillars 99 


are common to all the caterpillars, and many of which can be 
homologized with setae occurring in the Trichoptera and Panor- 
pata; or they are secondary, in larger numbers, and not definitely 
arranged. The setz are more or less completely carried over into 
the pupa, with comparatively slight changes in their arrangement. 
Often in place of the primary hairs there are tufts of hair spring- 
ing from the same wart. This is the condition spoken of as 
‘“‘with tufted hair.’’ In such caterpillars as Melalopha, this is 
due to the occurrence of secondary hair on the tubercle of the 
primary hair, which remains quite distinct, but in the Arctiidae 
it seems rather to be a reduplication of the primary hair. 

Where the secondary hairs are very few, like the primaries, 
they take definite positions, and are then known as subprimaries. 

The arrangement of the primary setz on the meso- and meta- 
thorax of the typical Frenatz is shown in figure 4, while different 
arrangements for the abdomen are shown in Figures 5, 33 and 34. 
The small primaries numbered wa, 7x and x are usually over- 
looked. The numbering of the others is after Dyar, except that 
figure 33 1s changed to make it agree with the others. 

The prothorax has its arrangement modified by the persist- 
ence of three sclerites, the cervical shield, with many sete and 
some punctures, the prespiracular wart, with two or three setze 
(sometimes fused with the cervical shield) and the subventral 
wart, with two or three setae. 

Tufted or secondary hair on the body is usually accompanied 
by secondary hair on the true legs and head. It reaches its great- 
est development in the Lasiocampidae, where it occurs even on 
the antennae and palpi, and on the sclerites, a, b, and c of the 
labium. 

PROLEGS. 

One may take as a typical proleg that of the Tortricidae or 
higher Tinerdae. It is a fleshy, more or less conical projection, 
the tip of which, the planta, is more or less retractile, and rounded 
or flat. Around the edge of the planta is a series of hooks or 
crotchets, each imbedded about two-thirds its length in the 
skin, but with the tip free and hooked toward the center of the 
circle. To the center of the planta is attached a muscle, by which 
it can be completely inverted, even the hooks disappearing from 
sight. Just above the hooks on the outer side there may be a 
horny edge, and above that, a larger plate bearing three setae 
(vii). On the front of the base is a minute seta (sometimes 
entirely in front of the leg,) and on the inner side another (viii). 


100 Annals Entomological Society of America [Vol. III, 


The possibly primitive state of the proleg can perhaps be 
reconstructed from Adela and Incurvaria. Each segment bore 
on the ventral side two chitinous sclerites, between which was a 
fold of membrane, which was rough with minute conical granu- 
lations like the rest of the skin. In this membrane there may 
have developed a concave fold, which lost its granulations, while 
those at each edge of it were enlarged, and became regularly 
imbricated, gradually becoming like the unmodified granules as 
you go away from the fold toward the two sclerites. This is the 
condition of Adela, (Fig. 6), except that the sclerites still pre- 
served in Incurvaria, are lost. On segment A6 the front side 
of the fold only develops hooks, and the folds of the remaining 
abdominal segments (1, 2, 7, 8, and g) are traceable, but entirely 
without hooks. There are no anal legs. 

In Incurvaria (Fig. 7) the posterior rows of hooks have disap- 
peared on all the segments, so that A6 is no longer different from 
A3 to 5, and the hooks are reduced to a single row. 

Hepialus has kept two rows of hooks, and in part, three, but 
the terminal row is much more highly developed, while the others 
are quite rudimentary. The two bands have fused on the inner 
side, and approach each other on the outer side, thus enclosing 
the fold, which becomes the planta. In many higher micros the 
outer ends have also fused, making the circle (ellipse) complete. 
In all above the Tineidae there have developed hooks on the 
last segment, but there is never a posterior series, or a complete 
circle; in some there is a straight band, which becomes a loop, 
but is always open posteriorly. It is nearest a circle in the Psy- 
chidae and lower butterflies. In the higher forms (butterflies and 
Noctuidae for instance), it has again become straight, but longi- 
tudinal, by the disappearance of the outer hooks. 

The ventral legs have a similar evolution. The circle which 
serves as a type for all the higher species may be broken on the 
outer side (Pyralididae), on the inner side (Psychidae), or on both 
(4Egertid@). It may be broken in front and back (Papilionidae, 
Thyatiridae, Libytheidae,) and in that case the outer half becomes 
weaker, and then disappears, leaving an inner band only (Noctu- 
na, Bombycina, higher butterflies, etc.) 

In the Tineina the last ventral legs may entirely disappear 
(as has been mentioned they are only half developed in the very 
primitive type Adela) while in several higher families the first 
ventrals are lost (Noctuidae, Geometridae, Nolidae). In the 


1910] A Structural Study of Some Caterpillars TOI 


Thyatiridae, Drepanidae and Notodontidae the last legs either 
disappear or are modified, while in the lowest Tineids they may 
have never: developed (Adela, Incurvaria). In the Euclezdae 
and Nepticula, as well as some other leaf-miners there are no 
hooks at all on the prolegs, but the area of roughened skin in 
Nepticula suggests an even more primitive condition than occurs 
in Adela. 

The Lycaenidae, as well shown in Scudder’s figure, have 
developed an outer fleshy lobe, apparently from the planta. The 
planta is apt to be unrecognizable in those which have lost the 
outer part of the circle of hooks, but its retractor muscle serves 
as in the lower ones to withdraw the hooks that remain. (Com- 
pare Jasoniades with Papilio). 

In Lagoa the line of hooks is sharply angulate at the middle 
and there the hooks are shortened. 

Another variation is in the arrangement of the hooks in the 
band. In the most primitive forms all those in a single line are of 
equal length gradually decreasing to the end, but a little higher 
in the scale, there develops an alternation of length by which the 
hooked tips are thrown into two or more ranks. In many of the 
highest families the single length again prevails, apparently as a 
secondary modification. In the lower forms (Micros) these two 
arrangements intergrade, but in the Macros they separate 
families or even superfamilies very sharply. 

The proleg typically bears four setae, three on the outer side 
(vii) and one on the inner or anterior side (viii). The minute 
primary ix may migrate up on to the anterior side. 


HOMOLOGIES AND NAMES OF THE PARTS OF THE HEAD. 


About the identity of the front and clypeus; and of course of 
the mandibles, labrum, maxillae and labium as a whole, there 
can be little or no doubt. The epicrania, as the name is used in 
this paper, include also a number of other sclerites which are 
fused with it so completely as to show no suture. A large part 
of the epicrania becomes the compound eyes in the imago. The 
postgenz will not be discussed to any extent, but they are quite 
distinct. 

The naming of the parts of the maxilla and labium on the 
other hand has been done mainly in order to refer to them intelli- 
gibly. Such grounds as I have for this tentative homologizing 
may be largely drawn from the figure of the Elaterid beetle larva, 


102 Annals Entomological Soctety of America  [Vol. III, 


figure 25. The arrangement is about the same in Phryganea 
(one of the herbivorous caddis-flies), but in a sawfly, as shown in 
figure 32, there is no close relation to the others, the modifications 
having taken quite a different direction. In all three of these 
non-lepidopterous types the lacinia of the maxilla is well devel- 
oped, but in the Elater the proximal joint of the galea is open on 
the inner side, and one can see how the lacinia might easily fuse 
to it in much the same way as the galea is fused to the two basal 
joints of the palpus in the higher caterpillars. 

Especially striking is the complete disappearance of the 
glossee and paraglossee, not only in the larve figured, but in the 
adult. Elater also. The setae marked ai and the more veutral 
of the two sclerites which form the ring c, come closest to the 
normal position of the glosse and paraglosse. 


ARTIFICIAL KEY TO THE FAMILIES OF FRENATS. 
(Incomplete in the Microlepidoptera. ) 
1. Prolegs each of one or two transverse bands of rudimentary hooks, no 


ADA) MEO SS as. sew FSF ats eas Sea ae eas te eee aa Tineidae 
1. Hooks of prolegs in curved or longitudinal bands, or in a circle, anal pro- 


legs wanting only./in-Drepanidaes.. << fem. des OA aa es ee 3 
1. No prolegs, or proless without bhaols./f .25..0.t: che ee ee 2 
2.) Shig-catetpillass, “exposed Feeders so... 26. a7 ods hse eae Eucleidae 
2. “uplender leat=cosmens yar en... stands ayes, oe are Nepticula and other Tineina. 
3. Hooks of prolegs in two curved transverse bands.......... ASemnae 
3. Hooks of prolegs in a complete circle or horseshoe. . : 4 
3. Hooks of prolegs in a longitudinal band, on inner side of leg, with or without 

a rudimentary outer Series. - ou... dct G's awe ve Sen ee ee 8 
A, Wath much fine secondary. hair iW1 he Sa oad ns ceeeane ees ee ee 
4. With a single subprimary on the proleg................... Taceaaaanee 
4.) ‘Without\secondary Rainn i7 oA) + -) th hic & api eee ee 6 
5. Head: much larger than’ second ‘sesment...-.. 5.1252 05.8 Hesperiidae 
5. Head smaller than second sepment:....:..:.:.:....... Megathymidae 
6. Circle of hooks interrupted caudo-mesally, and as complete on anals as 

On the ofher Orolegss:. : joes tact ie ere ee Psychidae 7 


6. Circle of hooks complete, or interrupted outwardly, anal prolegs with a 
shorter, straight or curved band. 
The ty pical Micros, including Cossidae, Thyrididae and Orneodidae. 


7. Setaiof abdomen dorsal of ii, front much higher than wide. . . .Solenobiinae 
7. Seta ii of abdomen dorsal of i, front as wide as high............Psychinae 


8. Hooks of prolegs alternately of three lengths, at least with some short 
hooks: on: anal: PaOlege «s+ 5: asia: 91) ee nas Fk ie a ae Papilionina 9 
8. Hooks of prolegs alternately of two lengths......................4:. 21 
8. Hooks of prolegs of the same length, or gradually growing shorter to the 
end: of ‘the. Lows: wy. sooo reo sone Peane Gate nat tet eee 12 


9. Maxillary lobes minute; labrum with 18 or more secondaries. . Papilionidae 
9. Maxillary lobes at least half as long as terminal joint of palpus; labrum 
witb: less than 18isecondariés...% chcigt 2. sos cee ie ets een ee 10 


10, ‘Slugeaterpilars with ‘small heads. wc jc .26 usnianier aes Lycenidae 
10.. Normal. cylindrical caterpillars: ...3,7.'0. S306 oo emus nek ee ee 11 


TgIo] A Structural Study of Some Caterpillars 103 


Seta iii of labrum as near to ii as to iv, far from margin........ Pieridae 
Seta oMmlabrum On) Marcin open ey Trek rah rae ike soe calc: Nymphalidae 
Hooks of prolegs in an angulated band, shortest at the angle; with addi- 

tional prolegs (two pair) without hooks. Bier .Megalopygidae. 


Hooks of prolegs in a straight or curved band; no additional prolegs. ..13 


Anal prolegs reduced or modified, with fewer or no hooks. . Notodontidae 
Anal’ proless larger than: ventralomes: tw tcety amie eso i.tosh ect cay 14 


With dense fine secondary hair on head, body with tufted hair........ LS 
With some sparse secondary hair on head, body with tufted hair. . Arctiidae 
Without secondary hair on head, body often without secondary or tufted 


1A eae heen ee eer eer eee PML Nase nwt ote bro Reones eet a coms. ol 17 
Eversible dorsal glands on posterior abdominal segments; a tuft of second- 

ary hair on the adirontals; five warts on mesothorax....... Liparidae 
No eversible dorsal glands; no such tuft of secondary hair........... 16 
Three warts on mesothorax, the upper very large.......... Syntomidae 
Four nearly equal warts on mesothorax...........Noctuidae (Pantheinae) 
Bodypwith: primary hair only... i... 0. 81h). at eee OCtuidae 
Body, with: tuited: orpsecondary hair’ cr).' 5005, «x. <,-.sp) vache ee os a eee 18 
Rounteentlegs (inst prolegswamntiney))/ amet. 5 cise ee eR eee Nolidae 
SUR beCM METS ac. Sey.) Perse dere ee ee ee he nayts © ss ciclaice dae eee aa IK) 


Two warts above spiracle on each segment of abdomen.......... 
Zygaenidae (Pyromorphide) 
Three warts above spiracle on each segment of abdomen, or with second- 
ARYA es Ey ces, 28 os de eee ae ero oh eer eee oe ce ee 20 


Adfrontal puncture three times as far from lower as from upper seta. . 
Arctiidae 
Adfrontal puncture not more than twice as far from lower as from upper 
SOU Inne Goo att ke acl nes POP nes Le Gari de cian Noctuidae (Acronyctinae) 


Anal legs wanting; a series of small outer hooks on prolegs. . Drepanidae 
Anal legs reduced; a few outer hooks:on prolegs.......... Thyatiride 
Anal legs large; ventral prolegs reduced or wanting; no outer hooks on 

PEO CaS ese A sete eyed bys are oma yee cca are ects Mice arene at Geometridae 


Secondary hair on antennae, palpi, and sclerites a, b and ¢c of labium 
; Lasiocampidae 
No secondary hair on antennae, palpi, or tip of labium............... 24 


Seta vi of labrum distant from margin, migrated toward ii*........... 25 
Seta.vi' Ol labrum marsinalvor mean SO. eee ees ee ee 26 


Hain! smMicroscopics Ss Withoubsnbubbedy icin severest ate oe ee Sphingidae 
Hair conspicuous; with tufted as well as secondary hair....Apatelodes} 


An unpaired dorsal spine or tubercle on ninth abdominal segment 
Citheroniidae 
Usually with an unpaired dorsal spine on eighth abdominal segment, 
but never with one on the ninth.......... Bankes .Saturniidae 


* This separation does not entirely hold, see the descriptions of the individual families. 
t+ Probably does not belong to the Notodontidae. 


104 Annals Entomological Soctety of America [Vol. III, 


SPECIAL PART. 
THE SUBORDERS. 


JucaTaE: Unfortunately I have only the highest family, 
Hepialidae, of this suborder for examination. Probably the 
head characters will hold in the other families, so far as they are 
primitive in their nature, such as the palpi, ocelli and galea, but 
the others will be very likely to fail. 


Frontal punctures more widely separated than frontal setae; 
the four anterior eyes arranged in two pairs; galea with the area of 
membrane on the mesal aspect, one of the large cones being shifted far 
proximad on that side; second joint of palpi more distinct than in the 
Frenatae and shaped like the free distal ones; an anterior subdorsal 
seta present on the meso- and metathorax, besides the minute pri- 
maries; iiia of abdomen large and dorsal to i111; ix of abdomen large, on 
leg-base. 

The abdominal setae iv, v and vi of Hepialus hectus are ar- 
ranged very much as in Incurvaria. The hooks of the prolegs 
form a circle broken shortly on the inner side; they are all of 
one length, but there is a double series of rudimentary hooks at 
their base. 

Fipures 27, 30, 33- 

FRENATAE (including Rhopalocera): Frontal punctures closer 
together than frontal sete; the four anterior eyes forming an arc of a 
circle; base of galea with the area of membrane on the anterior side, 
and with the large cones nearly opposite the maxillary lobes; first two 
joints of palpi quite similar, very short, and completely fused into 
the maxillae; no anterior subdorsal seta on thorax (except in Litho- 
siidae, which are otherwise very highly developed Frenatae); iiia of 
abdomen minute, usually lying between iii and the spiracle; ix of 
abdomen minute. 


The prolegs of Adela are probably more primitive than those 
of Hepialus. 


SUPERFAMILY PAPILIONINA (including the Hesperiina) : * 


Prolegs with hooks alternately of three, very rarely of two, lengths, 
either in a complete circle or a straight line. Secondary hair present 
(on body, epicrania, front, mandibles and stipes, and sometimes 
everywhere except on the tips of the mouthparts and the antennae.) 
Labium narrower than the maxillae, and narrower at the base than 
oné of the submenta. 


* See Scudder, Butterflies of Eastern North America; for a good many additional characters, 
and for descriptions of all the known eastern species. 

Of the Hesperiina I have seen no caterpillars of the Megathymide. The characters I have 
italicized should be expected to apply to them as well as the Hesperiidw, while the small, normal 
head will separate them. 


TgI0| A Structural Study of Some Caterpillars 105 


1. Labrum very shallowly notched, all six setae near the outer edge; cervical 
region chitinized in two pair of sclerites, which fuse more or less to the 
corresponding sclerites of the head; postgenae very wide. Large cones 
with a tuft of branches (unique). Prolegs with hooks in a complete 
circle, or shortly broken on the outer side; part of primaries modified into 
flat plates, the others inconspicuous. Labial setae at close together. 

Hespertina, Hespertinae 
* Feet with 125 hooks, alternately of 3 lengths. Head square; 
postgenae with setae; adfrontals abruptly widened at top of front; 
gula as wide at base as it is long. Example: Epargyreus tityrus (Fig. 23). 
* Feet with 50 hooks, alternately of two lengths; head heart- 
shaped; postgenae fused with gula and without setae; adfrontals very 
wide and long; gula twice as wide. Example: Pholtzsora catullus. 
(Fig. 24). 

1. Labrum usually with setae normal (about as in Noctuidae) not all on the 
terminal margin, usually moderately notched; cervical region not 
chitinized, postgenae meeting, or else very narrow. Large cones 
simple, normal. Proleg with hooks never in a complete circle or horse- 
shoe (the outer band preserved only in Jasoniades, Euphoeades) 
primaries unmodified, usually similar to secondaries; labial setae az 
widely separated. Papilionina go to 2. 


bo 


Labrum with about 18 secondaries. Maxillary lobes very small, front 
wider than high, not reaching half way to vertex; clypeus as broad as 
1-3 height of front, with many scattered secondaries, primaries sep- 
arated by half the distance that separates the two primaries i from each 


other. Lower end of adf. about as broad as clypeus. Paptlionidae 
* Feet with an outer row of 45 small hooks and an inner row of 
80 large ones. Example: Jasontades glaucus (Figs. 11 and 22.) 


Euphoeades trotlus. 
* Feet with a single row only of about 50 hooks. 
Example: Paptlio polyxenes (Figs. 10 and 21.) 
Laértias philenor. 

2. Labrum with less, or usually without secondaries. Maxillary lobes at least 
half as long as last joint of palpus. Front higher than wide (except 
in Anosia) usually reaching more than half way to vertex; clypeus 
usually much narrower, and with less or no secondaries, when widest 
the adfrontals are only about half as wide as it at the bottom. go to 3. 


3. Head small and body stout; prolegs with fleshy outer lobes curving down 
over the row of hooks; clypeal setae very far apart; about as in Papilio, 
but on a narrow clypeus. Seta iv of labrum some distance from the 
margin. Adfrontals apparently very narrow. Lycaenidae 

Example: Thecla ilicts. (Fig. 38.) 
3. Head at least half diameter of body, which is approximately cylindrical; 
- prolegs normal; seta iv of labrum variable. Claws of true legs deeply 
lobed. go to 4. 


4. Primaries on body mostly distinct, in P. brassicae conspicuous; adfrontals 
very narrow; clypeal setae far apart; ii of labrum distant from margin 
nearly as far as from seta ii. Pieridae 

Example: P. rapae, (Fig. 20), P. brassicae. 

4. Primaries on body not distinguishable from secondaries, at most a little 
larger; adfrontals about 1-5 as wide as height of front; clypeal setae 
separated by a distance not more than 4 that between the two setae 
i; 11 of labrum close to margin. Nymphalidae 


5. Prolegs with outer hooks (4 scattered ones); segments of abdomen divided 
into four equal annulets, each bearing a row of larger black setae. 
Libytheinae 

Example: L. celtis. (Fig. 37.) 
5. Prolegs with the inner row of hooks only; one annulet of each segment 
broad; the larger black setae, if present, irregularly arranged, or on 
spines. go to 6. 


106 Annals Entomological Society of America [Vol. III, 


6. Front broad and epicrania high (as in Papilio) labrum with secondary 

setae; mentum only 1-4 as wide as stipes. Euploeinae 
Example: Anosta plextppus (Fig. 12). 

6. Front much higher than broad, and usually reaching more than half way 

to vertex; labrum with very few or no secondary setae; mentum wider. 

go to 7. 


7. Labrum with i and ii low down near margin, and notch very shallow; 
mentum without secondaries; two minute points projecting back 
from rear of body, but otherwise unarmed. Satyrinae 

Example: Cercyonts alope. (Fig. 19). 

Labrum normal, with moderate notch, setae all about equal and i and ii 
far above the level of v and vi; mentum with secondaries (onlyjone 
odd one in my specimen of Euphydryas phaeton). Body more gener- 


ally spined, (or else entirely unarmed, in Anaea). Nymphalinae 
Examples: Argynnts cybele. Argynnint (Fig. 14). 
Euphydryas phaeton. Melttaetni (Fig. 15). 
Polygonia tnterrogationts, Vanesiint (Fig. 16). 
Euvanessa antiopa, Vanessim (Fig. 17). 


Basilarchia distppus. Nymphalini (Figs. 18 & 36). 
Spines very unequal in Basilarchia. - 
No unpaired spines in Basilarchia, Argynnis. 
Two unpaired spines on last segment in Euphydryas. 
Two spines to a segment in subventral row in Euphydryas. 
Hooks of prolegs less regularly arranged in Basilarchia. 
Ventral legs with hooks of two lengths only in Euphydryas, of 

four lengths in Polygonia. 

30 hooks in Euphydryas, 50 in Vasessini, 60 in Basilarchia. 
Clypeus and adf without secondary setae in Basilarchia. 
Mentum with most setae in Euvanessa. 
Postgena wider in Euvanessa. 
Labrum with secondary setae in Argynnis. 


THE SPECIALIZED MACROFRENAT. 


Following the Butterflies there comes a series of families 
which include the larger and better-known moths. These may 
be divided for convenience into two groups, which perhaps 
represent very early-separated lines of descent. 

The first group, which seems to be especially associated with 
a tree habitat, may be roughly defined by the presence of four 
setae on the outer side of the legplate (the upper one of which will 
be spoken of in this article as vib), by the usually rather small 
front and large epicrania, and especially by the hooks of the 
prolegs, which are alternately of two lengths. The caterpillar 
very frequently has secondary hair, obscuring the arrangement 
of the primaries; and the imago shows a very strong tendency to 
lose the frenulum. (It is minute or wanting in the Saturniidae, 
Ceratocampidae, Lasiocampidae, Endromidae, Bombycidae, Dre- 
panidae and Lacosomidae, also in a few Sphingidae and Geome- 
tridae). Most of the families are small, and the type, except as 


1910] A Structural Study of Some Caterpillars 107 


represented by the Geometridae, seems to be geologically past 
its prime. . 

On the other hand the group typified by the Noctuidae is the 
dominant modern type of moths. Though they are much more 
uniform in adult structure, there is a vast number of species, and 
a surprising variety of larval types. In these families the outer 
side of the leg-base never has more than the three typical setae 
of viz unless tufted hair is present. There is never but one 
length of hooks on the prolegs; the front is usually somewhat 
larger in proportion. Secondary hair is very rare, occurring only 
in. Thaumetopcea, Panthea and Apatela, and only in. ‘some 
species of the latter. A decided majority of the species live on 
low plants. 

Group I. (with vib, and usually with secondary hatr). 

This group again can be divided for convenience into two sec- 
tions, those with and those without secondary hair. The first 
includes the Sphingidae, Saturniidae, Bombycidae, Lasiocampi- 
dae and related families: the second, the Notodontidae, Thya- 
tiridae, Drepanidae, Geometridae, and Lacosomidae, doubtless 
also the Epiplemidae and Dioptidae. 

The Notodontidae have been an especially puzzling group. 
Apatelodes shows no distinct affinity to the others, but is in every 
traceable way more like the Bombycidae and Saturniudae. In 
hairiness it surpasses both, and approaches the Lasiocampidae. 
The single-haired Notodontidae (such as Cerura and Hetero- 
campa) show seta vib unmistakably. Datana has secondary 
hair on the body, and in Melalopha it has invaded the head and 
the tubercles, producing a kind of tufted hair in which the pri- 
maries still remain dominant. The distribution of hair on the 
head is quite different from that of Apatelodes, with which the 
moth has been associated. In most ways it would seem 
more natural to derive the series from the Thyatiridae, and con- 
sider such genera as Gluphisia and Nadata, primitive. Still the 
hairy prolegs of Gluphisia* and Nadata would point toward an 
origin from the series with secondary hair. In the latter case 
the primitive Notodontan must have had very nearly the structure 
of Melalopha, with a recessive tendency to have a seta vib. 

. Lacosoma is a synthetic form between the rest of the series 
and the Microlepidoptera, with positive relations to both, as 
will be discussed under the heading of its family. 


* See Packard's figure, Monog. Bombyc. I. Pl. Fig. 


108 Annals Entomological Society of America  [Vol. III, 


P SPHINGIDAE. 


Minute secondary hair on epicrania, front, adfrontals, maxillae, 
labium and body; none on clypeus, labrum, mandibles, or distal parts 
of maxilla and labium; primary hairs 717 and iv of abdomen usually 
distinct (directly above and below the spiracle), 2, 71 and zt of epi- 
crania also sometimes distinct, never with tufted hair or branching 
spines. Epicrania large, extending more than the height of the front 
above its top, usually much more. 7 of adfrontals about at their 
middle (they are usually the lowest setae the adfrontals bear) ; labrum 
with vz decidedly more distant from the margin than v, 777 and iv 
not very close together; mandible with one seta far out near tip of 
scrobe; mentum very wide at base, without distinct submenta; prolegs 
with a single band of hooks alternately of two lengths, the anal prolegs 
the largest; caudal horn present except in Ellema. 

SESIINAE: Primaries i, ii and 111 of epicrania easily recognized by 
their enlarged punctures; lower ocellus rather farther than usual from 
the others; front wider than high, its width nearly half as great as 
height of head; adfrontals very narrow and with very irregular 
outer margin; clypeus very narrow, the setae close together, separated 
by only about 1-7 the distance between the two setae 7; head setae 
black and conspicuous under a lens; labrum with setae 7 and 7 in a 
straight line, 277 farther from the margin than iv, vi distant from the 
margin only about 1-6 of height of labrum, notch shallow; two conical 
spines on anal plate.* 

Example Pseudosphinx tetrio, Figs. 47 and 50. 


(The remaining subfamilies have primaries of epicrania rarely 
distinct, front usually higher than wide, adfrontals wider, clypeus 
usually wider, especially at the two ends, with the setae of each pair 
farther apart, head setz colorless, labrum with sete 7 and 7 out of 
line, 777 often as near the margin as iv, vt usually much higher, and the 
notch much deeper, anal plate unarmed). 

ACHERONTIINAE (Sphingicae only were examined). Head higher 
than wide, rather regularly rounded, or moderately tapering toward 
the vertex, a large number of very minute sete on the epicrania, aver- 
aging eight or ten in a portion of the epicrania the size of the front, 
arising from depressions on the surface, front with very sinuous mar- 
gins, labrum deeply notched; skin usually smooth in last stage; horn 
always normal. 


Body rough and granular, four short soft horns on thorax; two 
oblique lines of granules on each segment besides the mid-dorsal line; 
labrum with v distant from margin; mandible with teeth less distinct 
than usual in the family; posterior ocellus rather high, front only 1-3 
height of head and with exceptionally sinuous outer margin; head dis- 
tinctly tapering toward vertex, widest near base. 

Example Ceratomia amyntor Figs. 45 and 49. 

Widest point of head about halfway up; front proportionately very 
wide toward the apex. Horn at the middle about as thick as at base. 
Posterior ocellus a little lower; front about two-fifths height of head. 

Example Sphinx gordius. 


* Hemaris is less aberrant. 


Tg10] A Structural Study of Some Caterpillars 109 


Head decidedly tapering upward, upper part of front narrow, horn 
tapering regularly; otherwise about as in Sphinx. 
Example Dolba hylaeus Figs. 39-41. 


Labrum much less deeply notched, head intermediate between that 
of gordius and hylaeus; front hardly more sinuous than in Deilephila. 
Horn strongly curved and regularly tapering. 

Examples Phlegethontius celeus. 
PB. sexta, Big. 151% 

AMBULICINAE: Head much higher than wide and tapering much 
to the vertex; epicrania with only three or four setae in an area the 
size of the front, set on decided tubercles; front with margins sinuous 
as in the preceding subfamily, quite small proportionately; Labrum 
very deeply notched. Skin rough, horn always conical, variable in 
Size. 

Example Paonias myops Figs. 42-44. 

PHILAMPELINAE: Head squarish; front small in proportion to 
head, and higher than wide; with much straighter margins than in the 
preceding groups; posterior ocellus low, as near to lower ocellus as to 
the fourth one; labrum deeply notched; a subspiracular ridge on the 
anterior segments, disturbing the incisure between thorax and abdo- 
men; skin always smooth, horn variable. 


Head wider than high, third ocellus much enlarged; caudal horn 
shorter than height of head; supraanal plate nearly an equilateral triangle, 
and acute at tip. 

Example Amphion nessus. Figs. 48 and 53. 


Head higher than wide, with the sides nearly parallel; third ocellus 
much enlarged, caudal horn nearly normal; supraanal plate as narrow 
as in the last, but with the tip rounded off. 

Example Darapsa myron. 


Head higher than wide, the sides converging somewhat toward the 
vertex, caudal horn replaced by a low button; supraanal plate broad 
and rounded. 

Example Pholus pandorus. 

CH@ROCAMPIN®: Head decidedly wider than high, the upper 
part nearly spherical; epicranial seta 2 distinct; posterior ocellus 
high, half farther from lower than from fourth ocellus; front wider 
than high, its width equal to about half height of head, its margins 
nearly straight, as in the last subfamily; clypeus quite narrow with 
the setae close together as in the Sesiinae; labrum with notch shallow; 
7 and a nearly in a straight line; zzz farther from the margin than iv, 
vt not very far from the margin. This comes nearer than any of the 
others to Pseudosphinx. 

Example Deilephila gallii Figs. 46 and 52. 


APATELODES. 


Secondary hair on epicrania, front, adfrontals, mandibles, maxillae, 
labium and body, but not on distal parts of maxillae and labium or on 
clypeus or labrum; tufted hair also on body. Front rather large, 
the epicrania extending about its height above it, labrum with setae 
vt quite distant from margin, 1277 and v also not marginal. Secondary 
setae of labium arranged in two diverging rows.  Setae az of labium 
close together. Prolegs with hooks in a single band, alternately of 


110 Annals Entomological Society of America [Vol. III, 


two lengths. No caudal horn. Frontal punctures close together, 
but the setae 7 are also close together. Mentum somewhat wider at 
the base than one of the submenta. 

It may be distinguished from the Saturniidae by the much denser 
hair on the maxillae, the larger front and the fact that setae az of the 
labium are much closer together. This genus is entirely isolated in 
the North American fauna, it is perhaps most often placed with the 
exotic family Eupterotidae. 

Examples A. torrefacta Figs. 54, 55 and 56. 
A. angelica. 


LASIOCAMPIDAE. 

Very dense secondary hair on all parts except the tips of the palpi, 
even on the antennae; tufts also present (7, 771, 1v and viz), but reduced; 
front rather small, the punctures close together, and the primary setae 
rather close together; adfrontals enlarged opposite top of front, with 
the setae close together. Prolegs with hooks alternately of two lengths 
in a single row. ; 

Second joint of antenna three times as long as wide. Labrum with 
most of the secondary hair toward. the tip of the lobes; prolegs spread 
laterally; a low tubercle in place of caudal horn; lappets on legless seg- 


ments simulating the legs, also on first segment each side of the head. 
Example Artace punctistriga ’. 


Second joint of antenna twice as long as wide, labrum with most of 
the secondary hair in the neighborhood of setae 1 and ii; no lappets or 
tubercles, prolegs normally placed. 

Examples Malacosoma americana. 
M. disstria (Figs. 59 and 60). 


SATURNIINA. 


Secondary hair on epicrania, front and body, and sometimes on 
adfrontals, mandibles, maxillae, and labium; tubercles, at least in part 
with tufted hair; tubercles 27 of either eighth or ninth abdominal seg- 
ments fused across the middle line (that of the eighth would be the 
caudal horn), except in Saturnia; mentum wide, its base wider than 
one of the submenta; prolegs with hooks alternately of two lengths, 
in a single band; setae az of labium far apart; punctures of front far 
apart, with reference to the sete, but close together in comparison to 
the width of the front. 


CERATOCAMPIDAE: Tubercles ii of ninth abdominal segment fused. 

Adfrontals, maxilla and labium without secondary hair; front as wide 
as high and head wide; wt of adfrontals about 2-5 up the front; clypeal 
sete close together; secondary hair of body minute; armed with many 
long horns. 

Example Citheronia. Fig. 63. 


Maxille with secondary hair but none on adfrontals; front wider 
than high, zw of adfrontals only 1-3 way up it; clypeal setae close together; 
head as wide as high; secondary hair of body long. 

Example Basilona. Figs. 61 and 65. 

Maxillew, adfrontals and labium with secondary hair; head and front 
both higher than wide; adfrontals wider; both v and vi of labrum distant 
from margin; secondary hair of body minute. 

Example Anisota. Figs. 62 and 64.» 


Tgto| A Structural Study of Some Caterpillars IIE 


SATURNUDAE: Tubercles of ninth abdominal segment separate. 

Labrum notched about half its depth; adfrontals without secondary 
hair, the puncture very close to the upper seta; secondary hair of labium 
in two parallel rows, (as in Apatelodes) secondary hair also on maxilla, 
but not on labrum or mandible; maxillary lobes well developed, decidedly 
longer than broad. 

Hyperchiria io. Figs. 66 to 68. 

Labium with not more than one or two secondaries; mandible with 
secondary hair; maxillary lobes minute; labrum cleft two thirds its width 
or more; adfrontals with several secondary hairs. 

Labrum with secondary hairs; warts of body larger,. several 
haired. 
Example Telea polyphemus. Fig. 69 and 70. 


Labrum and warts with primary hair, warts smaller. 
Example Tropaea luna. Figs. 71 and 72. 


(The labrum in this figure is aberrant). 
GYNANISA ISIS agrees structurally with this family. 


ENDROMIDAE. 


With secondary hair on epicrania, front, maxillae and body, 
but none on adfrontals, clypeus, labrum, mandible, labium, etc.;. 
hair minute. Epicrania large and front small, higher than wide, 
punctures of front far apart and lower than setae; labrum deeply 
notched, seta vz a short distance from the margin, 777, 1v and v close 
to the margin; mentum not very wide, about as wide at the base as one 
of the submenta; upper ocellus minute; prolegs with hooks in a single 
row, alternately of two lengths, anal legs larger than the ventrals. 
There are no North American species of this family. 

Example Endromis versicolor. Fig. 57. 


BOMBYCIDAE. 


With secondary hair on epicrania, front, and body, and with one 
each (in the specimen before me) on adfrontals and maxillae, hair 
minute, but denser and longer than in Endromis. Tufted hair repre- 
sented in adult by a single rudimentary tuft in the subventral region. 
Punctures, etc., about as in the Sphingidae and Saturnoidea; setae az 
of labium moderately far apart, but not so far as in Saturniina. Basal 
joint (the one fused to the maxilla) of maxillary palp unusually broad. 
Prolegs with a single row of hooks, alternately of two lengths. Caudal 
horn present. There are no North American Species of this family. 

Example Bombyx mori. Fig. 58. 


NOTODONTIDAE, 


Head with secondary hair, only in.Melalopha; body with or without 
general secondary hair, but always with it on the prolegs; seta vib- 
distinct unless covered by secondary hair; epicrania with setae 7 well 
above the top of the front; submenta large and nearly meeting; frontal 
punctures farther apart than in most Noctuidae, often close to the 
sete, front small, the head extending twice its height above its top; 
prolegs with hooks in a single band, not alternately of two lengths; anal 
prolegs reduced or modified, with fewer hooks; primary setz distinct, 
arranged as in the Noctuidae, often on enlarged tubercles. For super- 
ficial structural characters in this family see Packard’s Monograph of 
the Bombycine Moths, Part I. 


112 Annals Entomological Society of America [Vol. III, 


MELALOPHINAE: With dense secondary hair on epicrania, front, 
adfrontals and body, also on the tubercles, but not obscuring the 
primary hair; notch of labrum shallow, the setae 7 and 7 not far out 
of a straight line; anal legs moderately reduced. 

Example Melalopha apicalis. Figs. 76 and 83. 


PyG#RINAE: Head without secondary hair; dense secondary 
hair on body, usually not obscuring the primary hair; epicranial setae 
u three times as far apart as 7; anal legs are rudimentary stemapods, 
with one or two retracted hooks, labrum small proportionately. 

Examples Datana ministra. Fig. 78. 


= 


D. integerrima. Fig. 73. 


NoTopONTINAE: Without secondary hair except on prolegs; anal 
legs well developed in one form or another (in the species studied fairly 
normal), with hair about as on the other legs; epicranial setae 77 very 
high up; and little if any farther apart than 7; anal dungforks not 
developed. ; 


Labrum half as high as wide, cleft about half its height. Frontal 
punctures close to set. 5 
Examples Schizura concinna. Fig. 74. 
S. badia. 
S. unicornis. 


Labrum very high and deeply cleft; frontal punctures about trisecting 
the space between the setae; anal legs rather more reduced. 
Example Nadata gibbosa. 
Labrum and trontal punctures similar; anal legs very little reduced. 
Example Lophodonta. 
Frontal punctures close to setae: anal legs a little longer and more 
conical. 
Example Heterocampa guttivitta. Figs. 80 to 82. 
CERURINAE: Anal legs modified into stemapoda without hooks 
and with a retractile lash, with many setae; no secondary hair except 
on the stemapoda and prolegs; epicranial setae 72 intermediate in 
position; anal dung-forks strongly developed. Labrum moderately 


notched. 
Example Cerura. Figs. 75 and 77. 


Family THYATIRIDAE. 


Prolegs with a band of inner hooks, alternately of two lengths, and 
with a few minute outer hooks; anal legs slightly reduced; vib present 
but without other subprimaries; epicrania decidedly wider than high, 
front small; adfrontals wide, with 7 low down, frontal punctures 
close; labrum with vw slightly up from margin, and very deeply notched. 
Mentum rather short, but submenta nearly meet in the middle line. 


Head about a fourth wider than high; a slight hump on segment A8; 
body with a regular pattern of marks. 
example Habrosyne derasa. 


Head about half wider than high; body entirely lacking any hump, 
and entirely without marks, setae less distinct than in Habrosyne. 
Example Bombycia or. Fig. 88. 
Family DREPANIDAE. 


Prolegs with a band of inner hooks, alternately of two lengths, and 
with a short band of rudimentary outer hooks; no anal legs; a subpri- 


1910] A Structural Study of Some Caterpillars 113 


mary hair near 727, and from two to four in a longitudinal line between 
qv and v, and vit (vib): head sometimes with subprimaries, but without 
secondary hair; epicrania high, and head higher than wide; supraanal 
plate with a median horn bearing two setae at its tip; adfrontals high 
with 71 near top of front; mentum short and broad, but apparently 
with the submenta nearly meeting; labrum with vw a little distance 
from the margin. 


Three or four subprimaries on head, four setae represent mb; granu- 
lations long and setitorm, giving an appearance of fine secondary hair; 
subdorsal tubercles large, and present on meso- and metathorax (7b), 
and on segment 2 of abdomen (i); labral notch somewhat shallower. 

Examples Drepana falcataria. 
D. arcuata. Figs. 89 to 92. 


Less subprimaries on head, two setae represent vib on abdomen; 
granulations minute and conical; subdorsal enlarged tubercles smaller 
on thorax, and wanting on abdomen. Labral notch very deep. 

Example Cilix glaucata. 


GEOMETRIDAE. 


No secondary hair, except occasionally on prolegs; at least one sub- 
primary subventrally, often several; prolegs with hooks usually alter- 
nately of two lengths, the series often interrupted in the middle; 
(see Fig. 87). Epicrania full and rounded, or with the region of seta 
2 produced into an angle; front large, the punctures rather closer 
together than the distance between a puncture and the corresponding 
seta; adfrontais narrow over the top of the front; clypeus and labrum 
normal; mandibles usually rather thin, with sharp teeth and setae 
close together; maxillae and labium normal, of about equal width, 
the mentum at the base decidedly narrower than one of the submenta. 

The number of secondary setae on the proleg varies; in Cosymbia 
only the subprimary v7zb is present; in Alsophila and an unidentified 
species there is a large number; in Brephos, Hydria, Aplodes, Zerene, 
etc., the number is intermediate, usually one besides vib. Alsophila 
and Brephos, which have additional rudimentary prolegs, have no 
additional setae on them. 

The planta of the Geometridae seems to be developed as a sucker 
on both ventral and anal prolegs and in several species the hooks in 
the center of the line are rudimentary to give it room. Such are 
Alsophila, Zerene, etc. 

The ventral proleg varies in position with reference to its segments. 
In Brephos it is directly below its segment, in Alsophila below the 
incisure between its segment and the next one behind; in Aplodes, 
Synchlora, Zerene, Ennomos, etc., it has shifted back almost beneath 
the following segment. 

Examples Brephinae Brephos nothum 


Hydriomeninae Alsophila pometaria 
Hydria undulata 


Sterrhinae Cosymbia lumenaria 

Geometrinae Aplodes sp. 
Synchlora aerata 

Ennominae Fidonia truncataria ? 


Zerene catenaria. Fig. 87. 
- Lycia cognataria? Figs. 84 to 86. 
and several other species unidentified. 


114 Annals Entomological Society of America [Vol. III, 


Family LACOSOMIDAE. 


Prolegs with a complete circle of hooks, alternately of two lengths; 
body with four setae representing vz (the fourth one being probably 
vib) but otherwise without subprimaries; 7v and v approximate below 
the spiracle, and on a level; all the setae clubbed; head Macro in its 
type of appearance; higher than wide; front rather high and extend- 
ing half way to vertex; anal legs with a practically complete circle of 
hooks; labium with high narrow mentum, and rather small submenta, 


not meeting. 
Example Lacosoma chiridota, halfgrown. Figs. 93, 94 and 95 


THAUMETOPOEIDAE. 


Secondary hair on epicrania, front, adfrontals, mandibles, maxil- 
lae and labium, but not on clypeus or labrum. Front small; setae 
moderate, the punctures trisecting the space between the primaries; 
as wide as high, adfrontals narrow; labrum with a very shallow cleft, 
111 and iv distant, vz a very short distance from the margin. Proiegs 
with hooks all of the same length, in a-single band, the anal prolegs 
with the same number of hooks (18) as the ventrals. Body with 
tufted hair, the tufts 72, 72, 7v, and v1, as well as viz and vii of the 
legless segments, being distinct, the secondary hair is in two transverse 
bands. Tuft zis of short dense hair, making the caterpillar look like 
a Lymantriid. There are no eversible glands. True legs with very 
deeply and curiously cleft claws. The family is not American. 

These characters suggest very strongly the Lymantrudae, especially 
Euproctis, without allowing one to deny that they may be due to 
divergence from a more primitive Notodontan origin. Staudinger and 
Rebel place it between the two families, which come together in their 
arrangement. It is usually considered Notodontid. 

Example Thaumetopoea (of Europe). Figs. 96 and 97. 


LYMANTRIIDAE. 


With secondary hair on epicrania, adfrontals, maxille and labium, 
and sometimes on front, none on labrum, mandibles, etc.; front rather 
large, somewhat higher than wide, the punctures close together, the 
setae, when not obscured by secondaries, very high up and far apart; 
labrum quite variable; maxilla and labium much as in the Noctuidae 
in form. Prolegs with a single band of hooks of a single length, the 
ventrals and anals equal. Body with tufted, but no secondary hair; 
wart 7 quite variable and furnishing generic characters, vitt somewhat 
diffuse, as if there were a little secondary hair associated with it. 
Claws of true legs moderately cleft as in most Lepidoptera (similar to 
Fig. 101). Dorsal glands on 6th and 7th abdominal segments. 

1. Labrum more deeply notched, one third its height or more, seta 
vt not on the margin, adfrontals enlarged at the top, with a tuft 
of secondary sete; front and upper part of epicrania without second- 


aries. 
2. Labrum cleft about half its height, sete i and ii far out of line 
with each other; body with tufts i small, and ii moderate, both normal. 
Example Porthetria dispar. 


1gto] A Structural Study of Some Caterpillars I15 


Labrum cleft about a third its height; setae ii only moderately 
out of line with 7; body with tufts ¢ and a of four abdominal segments 
fused into large square masses. 

Example Hemerocampa leucostigma. Fig. 98. 

1. Labrum less deeply notched, vz nearly marginal; adfrontals 
slender, their secondary sete inconspicuous; front with secondaries, 
especially in the lower part; epicrania with a good deal of hair above 
setae 11° warts 7 and 27 of the first abdominal segments fused to each 
other, but not fused across the median line. 

Example Euproctis chrysorrhea. Fig. 99. 


NOCTUIDAE. 


Prolegs with hooks in a single band, not alternately of two lengths. 
Usually without secondary or tufted hair, but both are present on the 
body in Pantheinae and Acronyctini; head with secondary hair on 
the epicranta, maxillae and labium in the Pantheinaeonly. Front large, 
more than a third of the height of the head above the clypeus, the 
frontal punctures usually close together, never nearer to the setae 
than to each other; labrum with puncture za considerably nearer to ¢ 
then to 72, not very high up; adfrontal puncture rarely more than twice 
as close to upper as to lower seta; maxillae and labium of about equal 
width; the submentum at least as wide as the base of the mentum, the 
labial setae az close together. Seta of abdomen all well separated, 
t somewhat higher than 72, iv higher than v, vz always single and viz 
with three setae except in forms with tufted hair; no caudal horn. In 
those with tufted hair there are four warts on thorax above the legs, 
on abdomen three above and two below the spiracle, besides viz and 
vit. Never with eversible dorsal glands; anal legs with more hooks 
than ventrals, one or two pairs of ventral legs often reduced or absent, 
but never three. 

PANTHEINAE: Epicrania, maxillae and labium with considerable 
secondary hair; adfrontals wide above the top of the front, the punc- 
ture about half way between the setae, and above the top of the front. 
Body with tufts and pencils, with secondary hair also, in Panthea. 

Example Demas coryli. Figs. 100 and 101. 


NoctuiNaE: Head without secondary hair, body occasionally with 
tufted and secondary hair, or tufted hair only. Adfrontal puncture 
below the top of the front, and adfrontals rarely much widened at the 
top (Apatela interrupta) ; anal legs directed downwards and not length- 
ened ; setae usually short except in hairy forms, distance between upper 
adfrontal setae less than half height of front;-distance between 
frontal punctures not less than a third that between seta and puncture. 

In this group of Noctuinae (or Trifidae) there is a good deal of 
minor variation between different genera, and even species of the 
same genus, but my series of forms, though larger than of any other 
family so poorly represents this enormous group that I shall only 
characterize the two tribes Acronyctint and Cuculliint (Cucullianae of 
Hampson) and mention a few peculiarities of some other genera.* 

* J have done nothing whatever with the body setae. Dyar discusses seta iv in Proc. Ent. Soc. 


Wash. IV, 370 and Pird, in Can. Ent. Vols. 32-34, 39, 40 refers often to variations in the tubercle 
of the genus Papaipema. See also mv ‘Field Tables,’’ page 140. 


116 Annals Entomological Society of America [Vol. III, 


Acronyctini: with several-haired warts and often with secondary 
hair; with fine setiform granulations, with which there may be mixed 
larger conical ones. Epicrania high, extending over 1 1-3 times the 
height of the front above its top; adfrontals very wide, the upper 
setae far apart and high above the top of the front; clypeal setae closer 
together than the frontal seta and puncture; 7 and za of labrum sep- 
arated by a distance less than the width of a setigerous puncture; 
last joint of maxillary pulpus short. 

With secondary hair; labrum more deeply notched, wii and iv of labrum 
well separated, head dark, front about as wide as high, setae i and i of 
labrum form an angle of about 60 degrees with the horizontal. Adfrontals 
not especially wide at the top of the front. 


Examples A. (Hyboma) hasta. Fig. 103. 
A. (Triaena) hastulifera. Fig. 102. 


Similar; adfrontals with an enlargement at the top of the front, 

which contains the setae and puncture; labral notch not so deep. 
Example A. (Triaena) occidentalis (interrupta). 
Figs. 1, 12 and 26. 

Similar; head pale, labral notch shallower; front narrow; adfrontals 
unusually high and yet narrow; clypeal setae closer together; setae 7 
and wi in a line only 30 degrees from the horizontal; ta distant from 1. 

Example A. (Acronycta) leporina. Fig. 105. 


No secondary hair. labrum deeply notched; head black; front narrow ; 
wi and iv of labrum obliquely placed, separated by only half the vertical 
distance that separates i and w horizontally. 

Example A. (Eulonche) oblinita. Fig. 104. 


No secondary hair; labrum notched only 1-4 its height; front wide. 
Example Simyra (Arsilonche) henrici. Fig. 106. 

In the remaining forms there isno tufted or secondary hair; there are 
no fine setiform granulations, though there may be coarse ones; the 
front is larger in proportion to the epicrania and to the adfrontals; the 
adfrontal setae are closer to the front and to each other; 7 and za of 
labrum are more widely separated. 

Cuculliini Clypeal setae closer together than the distance between 
frontal seta and puncture, epicrania not reduced, labral setae about 
evenly spaced. 


Adfrontals i level with top of front; clypeal setae much closer together 
than frontal seta and puncture; labral setae i and wi quite evenly spaced; 
front as wide as high; adfrontals narrower; sete of labial palpi long. 

Example Cucullia sp. (undescribed). Fig. 110. 

Adfrontal setae i higher, clypeal setae farther apart; labral setae dis- 
tinctly arranged in pairs; front narrow, adfrontals wider; setae of labial 
palpi minute. 

Example Scopelosoma sp. 

Adfrontals 7 high, clypeal setae much closer together than frontal seta 
and puncture; labral setae i and w distinctly arranged in pairs; front dis- 
tinctly higher than wide; setae of palp intermediate. 

Example Calocampa curvimacula. 


Earias chlorana (of Europe) is made the type of a subfamily or 
placed with Nycteola. Claw very deeply notched; second joint of 
antenna short, tubercles 71 of 8th abdominal segment enlarged, and 
also 7b of thorax; 11 of epicrania directly below 7 and close to edge of 
adfrontals (as in some Micros) front a nearly equilateral triangle, 
adfrontals quite broad, and rather ill-defined; upper ocellus behind 
second instead of being above it. Fig. 112. 


1910] A Structural Study of Some Caterpillars 117 


Rhodophoia. Skin coarse and granular (Fig. 138), the coarser 
granules much more prominent than in Feltia; frontal punctures tri- 
secting the space between the setae; punctures of adfrontals half way 
between the setae; labrum hardly notched, with setae 7 and a in a 
straight line. The granular skin occurs also in Heliothis. 

Nectua Labrum with setae 777 and iv on a level, prolegs with 40 
hooks. (c-nigrum and other unidentified species.) 

Feltia. Coarse nodular granulations; first prolegs reduced to half 
their normal number of hooks and second prolegs slightly reduced; 
epicrania very low, in the last stage the adfrontals reach the vertex 
in a way similar to Zygaena. (Three stages). Fig. 108. 

Prolegs with only 10 crotchets; epicrania very short, adfrontal 
punctures half way between the setae. An unidentified species, agree- 
ing with Slingerland’s description of Euxoa scandens. Fig. 109. 

Hadena (Trachea) turbulenta. Outer margin of front very sinuous. 
Fig. 107. 

In Pyrophila pyramidoides, Ceramica picta and two species of 
Leucania, there were no decided characters. The epicrania were a 
little larger than in the preceding genera. 

NycTEOLINAE: Considered the type of a distinct family by some. 
Epicrania extending above top of front 1 1-2 times its height; ep. 72 
nearly twice as close together as z, yet distant from the adfrontals; 
labial palpi long and slender; adfrontals broad and ill defined out- 
wardly ; ocelli small, the distance between two ocelli being much more 
than the width of an ocellus. Body-hair long and fine. 

Example Nycteola revayana. Fig. 111. 


CATOCALINAE: Like the Noctuinae, but with the anal legs length- 
ened, or produced backward; head sometimes held horizontally ; never 
with tufted or secondary hair. 


Hind legs not lengthened ; prolegs with about 25 hooks; setae i and i 
of labrum at an angle of 30 degrees to the horizontal, iii and iv obliquely 
placed and close; epicranial setae ii low opposite adj. i and distant from 
ep. (Ne 


Example Ingura sp. 


Anal legs somewhat lengthened, prolegs with about 25 hooks; setae 
i and w of labrum nearly on a level; tv higher than ii; the two upper ocelli 
in contact; ep. i and w close together near vertex ; epicrania extending less 
than its height above its top. 
Example Drasteria erechtea. 
D. crassiuscula. 


Anal legs very long and produced backward, head held horizontally ; 

45 hooks on prolegs; labrum with setae i and ii nearly on a level, iii and iv 

close, wi directly above iv; frontal punctures less than twice as close to 

each other as to the setae; notch of labrum rather shallow; clypeal setae 

much farther apart than frontal seta and puncture; ep. i and ii close 

together on the face. (The conical hump of C. cara lines between them.) 
Example Catocala cara. Figs. 2 and 3. 


Anal legs very long; prolegs with over 50 hooks; labrum with setae 
iand wat an angle of over 45 degrees; iii and iv distant, frontal punctures 
decidedly closer together, frontal setae high up; labrum deeply notched; 
ep. t and w close together near vertex. 

Example Panapoda rufimargo. 


118 Annals Entomological Society of America [Vol. III, 


HypENINAE (Deltoides); Setae long and stout, both on head and 
body ; distance between adfrontal setae 7 more than half height of front, 
and they are also high up; distance between frontal punctures hardly 
more than 1-4 that between puncture and seta; front small, its setae 
usually high. 

Slender, green, a semilooper with only 14 legs, anal legs produced 
backward as in Catocala. 
Example Hypena humuli. 


ARCTIIDAE. 


Head usually with sparse secondary hair on epicrania, front and 
maxillae, often also on labium; none on clypeus, labrum, mandibles 
antennae, etc., or body, rarely on adfrontals (Apantesis). Epicrania 
rounded, front large, with the punctures about as far apart as the 
distance between a puncture and a seta; labrum normal, vw nearly 
marginal, puncture usually very high up, and more nearly over 1 
than 1; adfrontals usually very narrow and irregular, with the punc- 
ture close to the upper seta. Maxillae usually somewhat narrower 
than labium, the base of the mentum about twice as wide as one of the 
submenta. Body without secondary, but with tufted hair; four warts 
above legs on thorax, and three above spiracle on abdomen,—all the 
tubercles below the spiracles developed as separate warts. 

LitHosiiNAE: Head without secondary hair, adfrontal puncture 
close to upper seta; hair not feathered; no pencils or dense dorsal tufts; 
adfrontals very narrow and wavy-edged; thorax with the two upper 
warts side by side as on the abdomen. Frontal punctures twice as 
close together as distance between puncture and seta. 

Example Lithosia complana, 


ArctuNAE: Head with ‘secondary hair (except Euchaetias, in 
which the adfrontals are quite wide) ; hair serrate or feathered on body; 
often with dense pencils and dense dorsal tufts; warts of thorax all in 
a vertical line. Punctures of front nearly trisecting the distance 
between the setae. 


Adfrontal punctures about a third way down from upper seta, adfront- 
als broader; warts 7 and ii of abdomen transversely elongate and side by 
side, bearing dense tufts; with pencils of hair, hair very feathery. Labrum 
with puncture rather near to setae as in the Noctuidae (When young the 
puncture is a little nearer to the normal Arctiid position). Frontal 
punctures decidedly helow.the level of the setae. 

Examples Halesidota caryae. Figs. 119 and 120. 
H. maculata. 
H, tessellaris. 


Adfrontal puncture near upper seta; adfrontals, warts and hair sug- 
gesting Halesidota; head without secondaries (when very young the warts 
are as in normal Arctiidae, rather than as in Halesidota). Frontal setae 
and punctures on a level. 

Example Euchaetias egle. Figs. 117 and 118. 


In the remaining genera the hair is serrate rather than feathery, with- 
out dense tufts or pencils; puncture za of the labrum at least as far from 
i as wis; epicrania and maxillae, at least, with secondary hair. Frontal 
setae and punctures nearly on a'‘level. 


1910] A Structural Study of Some Caterpillars 119 


Front, labium and maxillae each with several secondaries; an anal 
tuft of long hair. 
Example Eubaphe nigricans. 
Front with several secondaries: maxillae with about ten secondaries, 
labium with a single pair; some scattered long hair. 
Example Hyphantria textor. 


Front, adfrontals, and maxillae each with several secondaries, labium 
without any; adfrontal punctures about a third way down from the 
setae; no long hair. 

Example Apantesis parthenice (?) young. Figs. 113 and 116. 


Front without secondaries, labium with a single pair, maxillae with 
about six. Frontal puncture and seta closer together than clypeal 
setae. 

Example Diacrisia virginica. 

Front with or without secondaries (at most a single pair); maxillae 
with five secondaries, labium with none; clypeal setae as in the last; no 
scattered longer hairs. 

Example Isia isabella. Figs. 114 and 121. 


Front without secondaries, stipes with six or seven; clypeal setae 
nearer together than frontal seta and puncture. 
Example Estigmene acraea. 
SYNTOMIDAE. 


Prolegs with hooks in a single band, all of the same length. Head 
with secondary hair on epicrania, front, maxillae and labium, but not 
on adfrontals, clypeus, labrum, mandibles, etc.; body with tufted hair. 
Front quite small, the epicrania extending fully twice its height above 
its top; frontal setae rather close together; adfrontal setae and punc- 
ture close together, the puncture almost as near the lower as the upper 
seta. Labrum with puncture za decidedly nearer 72 than 1; labium 
short and broad, the submenta separated by more than their width 
at the base. Thorax with only three tufts above the legs, the upper 
one twice as large as normal and elongate; abdomen with tufts as in 
the Arctiidae, 2 forming pencils. 

Example Ctenucha virginica. Fig. 122. 


ZYGAENINA. 


Head retracted within the first segment of the body; epicrania 
with setae reduced; the vertex cleft nearly to the top of the 
front, or with the cervical skin attached to the epicrania along a 
similar line; front triangular, about as high as wide, the adfrontals 
narrow and not extending much above it; frontal punctures about 
trisecting the distance between the setae, a little farther apart in 
Zygaenidae; clypeus with setae rather far apart, labrum with 7 
no higher than 7; mandible with setae close together; maxilla 
with seta zv distinctly arising from the galea, submenta about as 
wide as base of mentum, or membranous and indistinct; sclerite 
b of labium broad and massive as in the butterflies. Body with 
w and v approximated, or forming the same wart, 7 and 7 sep- 
arate or forming the same wart; with tufted or secondary hair, 


120 Annals Entomological Soctety of America [Vol. III, 


or with primary hair only, when there is tufted hair there are only 
two warts on the abdomen above the spiracle. Prolegs with 
hooks not alternately of the two lengths, or without prolegs. 

HETEROGYN!DAE: With primary hair only; prolegs with a single 
normal band of hooks; head black and heavily chitinized; epicrania 
with a deep cleft at the vertex; front with the punctures decidedly 
nearer the corresponding setae, than to each other; labrum with 71 
strong, and no higher than 7; antenna normal; body with 7 and 77 well- 
separated, 7v and v approximated, two setae on a level, in the position 
of vz, but one on the outer side of the proleg, and two on the inner side. 
It seems clear that the anterior seta on the inner side of the proleg 
belongs to vi1, but more doubtful whether the additional seta in the 
position of vz is the other missing one. 

Example Heterogynis paradoxa. 


ZYGAENIDAE: Head as in the preceding, seta 7 of the labrum 
decidedly higher than 7, but fully developed; body with somewhat 
diffuse tufts of hair, representing, 7+ 71, 212, 1v+-v, v1, and two tufts 
representing viz; vii1 single-haired; hair serrate, but not feathery. 

Example Zygaena trifolii. 


MeEGALopyGIDAE: Head pale, and lightly chitinized, submenta 
membranous; epicrania with the cleft in the vertex filled up, apparently 
by the growing together of its edges; their setae rudimentary ; labrum 
with 7a higher than in Zygaenidae, much smaller than 7, none of the 
setae marginal; frontal punctures rather nearer to each other than to 
the corresponding setae; antenna with first joint about as long as sec- 
ond, second less than twice as long as wide and without any long seta. 
Body with tufts as in Zygaenidae, but in addition with two isolated 
setae on a hump, near the tip of the prolegs. The row of hooks on the 
ventral prolegs is angulate in the middle and the shortest hooks come 
next to the angle. viz is opposite the apex of the angle. Second and 
seventh abdominal segments with rudimentary prolegs, on which the 
setae are arranged as on the normal ones, but without any hooks. 

Example Lagoa crispata. Figs. 123, 124, 125, 126 and 135. 


EucLeipAE: Mostly like the preceding family. Hair more or less 
reduced, diffuse, modified into branching spines, or absent ;* without 
prolegs, and without setae on the ventral part of the body; with a row 
of ventral suckers. 

Examples Cnidocampa flavescens. 


Empretia stimulea. 
Euclea delphinii. 


NOLIDAE. 


With tufted, but no secondary hair. Epicrania, front and 
clypeus as in the Eucleidae, but with normally developed setae. 
Labrum with setae normal, 7 lying between setae 77, and nearly on 


* See Dyar, Journ. N. Y. Ent. Soc., vols. iii to vii for details of structure of the various species. 


Tgto] A Structural Study of Some Caterpillars T21 


a level with them;. basal joint of antenna nearly as long as second 
joint, which is short, submenta heavily chitinized and well sep- 
arated. Body with tufts representing 7+17, m1, w+uv, vt, vu, 
and vii, with only two, therefore, above the spiracles, while 
there are three in the Arctiid types. Prolegs with a single unin- 
terrupted row of hooks, all of the same length; no prolegs on third 
segment of abdomen. 
Example Nola cucullatella. 


PSYCHIDAE. 


Prolegs with hooks all the same length, in a circle broken postero- 
mesally; anal legs similar to the others. Adfrontals massive, their 
setae well separated, not reaching far above the top of front; 7 of epi- 
crania close to them; frontal setae far apart, punctures close together 
and somewhat lower; antennae normal with short second joint; pro- 
thoracic spiracle piercing the cervical shield; labrum with vz not on 
the margin. Mera of true legs much enlarged and nearly or quite 
meeting in the middle line; all segments of the thorax with dorsal 
plates. Body with setae 7 and variable, 72v and v close together, 
vw variable, 22 of ninth abdominal segment distant from each other. 

SOLENOBIINAE: ‘True legs very long, and slender; front twice as 
high as wide; abdomen with 7 lower than 7, normal; prolegs with about 
15 hooks. 

Example Solenobia pineti. 

PsycHINAE: True legs short and very stout; front shorter; abdo- 

men with zz higher than 7; prolegs with over 20 hooks. 


Front nearly as wide as high; 7% of abdomen almost directly over 7; 
Example Thyridopteryx ephemeraeformis. T[igs. 128 and 134. 


Front half higher than wide; w of abdomen on the next annulet 
behind 7. 
Example Psyche zelleri. Fig. 127. 


COSSIDAE. 


Prolegs with hooks in an uninterrupted circle, alternately of three 
lengths, but with no great difference between the three lengths; the 
anal legs with hooks in a curved band. No secondary or tufted hair. 
Epicrania separated by a membranous area at the vertex, a slender 
prolongation of the adfrontals reaching the vertex; head not retractile. 
Front higher than wide; the setae far apart and-the punctures close 
together, level with the setae. Adfrontals large, reaching vertex, 
the setae rather close together, opposite the upper half of the front. 
Labrum with a shallow notch, 17, 1, a, and iv nearly on a level, 
none of the setae quite on the margin; mandibles extend forward with 
the cutting edge turned upward for gnawing wood. Mavxillary palpi 
with second joint very wide and massive, bearing the large cones, which 
turn wnwaid as in the Hepialidae; the galea arises from its apex as in 
other Frenatae, and is quite small. Labium normal with sclerites b 
massive, and ¢ slender; palpi rather short; submenta not in contact. 
Prothoracic spiracle distant from cervical shield; body with setae as in 


122 Annals Entomological Soctety of America [Vol. III, 


the Tortricidae; but on a9 the tubercles 77 are not fused across the 
middle line. 
In the structure of its lower lip this family is very different from 
the Tortricidae, and may be the most primitive of the Frenatae. 
Example Cossus cossus (three stages). Figs. 129, 130 and 151 


AEGERIIDAE (SESIIDAE). 


No secondary hair; prolegs with hooks in two curved transverse 
lines, all of the same length, anals rudimentary, with few hooks. 
Epicrania low, with 7 rather near the adfrontals, adfrontals practically 
reaching vertex, their setae close together near the middle, puncture 
opposite upper seta; front high and lanceolate, setae high, punctures 
low and not very close together; clypeal setae well separated. Labrum 
normal; maxillae with large cones part way down the inner side as 
in Hepialus, but galea arising from second joint of palpus as in Frena- 
tae. Basal joint of antennae massive. Body with zv and v fused, wv 
is a vertical row of three setae, the middle one the longest. Last 
spiracles dorsal and higher than 171; setae 21 of A9 separate. 

Example Melittia cucurbitae. Fig. 152. 


MICROLEPIDOPTERA. 


As my series of Microlepidoptera is quite short, it does not 
seem best to discuss the families separately. 

Setae 77 of the epicrania are usually rather close to the adfront- 
als, as in Argyresthia, but often somewhat more distant. They 
are never as far away as in the Bombycid series. The front is 
usually much higher than wide, often twice as high, but in 
Endrosis it is not, and in the Pyralide there are various inter- 
mediate conditions. When the front is high, the adfrontals 
nearly or quite reach the vertex, but when it is lower the adfront- 
als may merely border it, as in most Macrolepidoptera. The 
frontal setae are often far apart, close to the outer edge, as in 
Depressaria and Homoeosoma, or they may be closer together, 
but are never very close, as in the Bombycid series and Zygaenina. 
In the lower forms the punctures trisect the distance between 
them, but in the Tortricidae, Pyralidae, Depressaria, Simaethis, 
etc., they are much closer together, and lower, as in the Noc- 
tuidae. The adfrontal setae are very close together in Gracilaria, 
but often are not so. The puncture is apt to be about half way 
between the setae. The labrum is usually not very deeply notched, 
its setae 7 and 27 nearly on a level. 

The antennae are normal, but often with a seta on the side 
of the second joint, as in Yponomeuta and Cacoecia, figured. 
Yponomeuta is very aberrant in the proportions of the joints. 
The maxillae and labium are rather long, with the sclerites tending 


1910] A Structural Study of Some Caterpillars 123 


to be well-developed, the maxilla with stipes, palpifer and sub- 
galea sometimes separately chitinized, the submenta not widely 
separated, or even in contact, (Fig. 137). Setae of mentum rather 
nearer the base than is typical of the Macrolepidoptera. 

There is never secondary hair on the head, and secondary and 
tufted hair on the body only in the Pterophoridae. 

Thorax with cervical shield well developed—in the lower forms 
with additional sclerites ventrally and laterally, which reach their 
greatest development in Adela (Fig. 6). The true legs are 
absent in Nepticula, replaced by patches of enlarged granulations 
similar to those representing the prolegs. 

Abdomen usually with anal plate only, but with two dorsal 
and two ventral plates on each segment in Jucurvaria. Setae 
za and 2 well separated except in the Pterophoridae, iv and v 
approximated, except in Y ponomeuta, where they are distant and 
wv is higher than v, they are usually on a single tubercle, and zv 
may be much reduced. vz 1s quite variable, most often in the 
Tortricidae the setae form an oblique line, while in the Tineina 
the middle one is anterior. In the Pterophoridae, Thyrts, and 
Simaethts they lie, not on the leg, but on a plate at its base. 
Adela and Incurvaria are somewhat different (Figs. 34 and 35). 
vi lies on the anterior side of the leg in Adela and Incurvaria. 
On the ninth abdominal segment setae 27 are both on a single 
median tubercle in the Tortricidae, but in the others they are 
distinct, and often distant. 

The hooks of the prolegs in the Pyralididae and Tortricidae 
are oftenest in a complete circle, alternately of two or three 
lengths. They are similar in Thyris, Endrosis and Depressaria. 
Most Tineidae, as well as Phalonia and Orneodes, have but a single 
length. In Gracilaria there is half of a second band (Fig. 7). 
Adela has two areas of minute hooks, grading into the granula- 
tions, which in Incurvaria are reduced to a single transverse row. 
Y ponomeuta has three or four complete circles of hooks. Nepticu- 
la would appear to have the hooks replaced by a vague area of 
enlarged conical granulations. In Panorpa, there is such an area 
of enlarged, but setiform granulations, on the posterior side of 
each of the slender prolegs.* 

Tineola resembles Solenobia in its head and ventral prolegs, 
but its anal prolegs are normal. 


*Iam indebted to Dr. E. P. Felt and Prof. J. H. Comstock for the loan of this specimen from 
the Cornell University collection. 


124 Annals Entomological Society of America [Vol. III, 


The anal. prolegs are wanting in Adela and Incurvaria, those 
of the sixth abdominal segment are absent in Gracilaria and 
Coleophora, and reduced in Adela. 


Examples: 
Thyris vitrina. Simaethis oxyacantha 
Oxyptilus hieractii. (Figs. 139 and 140.) 
Botys polygonalis. Argyresthia goedartella 
Hydrocampa nymphaeata. (Fig. 136.) 
Homoesoma nebulella. Sitotroga cerealella. 
Crambus falsellus. Depressaria putridella 
Galleria mellonella. (Fig. 142). 
Orneodes hexadactyla. Cosmopteryx scribaiella. 
Caccecia cerasivorana. Coleophora. ~ 
(Figs. 133 and 137). Endrosis lacteélla (Fig. 143.) 
Carpocapsa pomonella Gracilaria alchimiella. (Fig. 7.) 
Several other unidentified Tor- Nepticula pomivorella. 
tricidae. Tineola bisselliella. 
Phalonia alcella. Incurvaria koerneriella. (Fig. 34.) 
Yponomeuta cagnagellus. Adela degeerella (Figs. 6 and 35.) 
(Fig. 141). 


SUMMARY. 

1. Useful classificatory characters may be found in the 
structure of the sclerites of the caterpillar head, and the arrange- 
ment of their setae. 

2. The Sphingidae, Saturniina, Bombycidae, Notodontidae 
and, perhaps the Lacosomidae, with their related families, show 
positive points of resemblance, aside from the mere presence of sec- 
ondary hair in most of them. This is found in the prolegs, sub- 
primary setae, frontal setae and proportions of front and head, 
and in their habits. 

3. The genus A patelodes is a synthetic form with suggestions 
of Lasiocampidae, Saturniidae, Bombycidae and perhaps Sphingi- 
dae. It is not near the Notodontidae. Melalopha is a fairly 
typical Notodontid. 

4. Lacosoma is a synthetic form between the Microlepidop- 
tera and Bombyx-Notodontid series, nearer (at least when young) 
to the Microlepidoptera. 

5. Some Papilios have the proleg structure of the skippers 
and Microlepidoptera. It is correlated with a nestbuilding habit. 

6. Thyris is a typical Microlepidopter. 

7. Cast skins and specimens dried without preparation make 
fairly satisfactory material for study, thus making it possible to 
found complete descriptions of larvae on the identical specimens 
that are bred through and accurately named. 


1910] A Structural Study of Some Caterpillars 125 


BIBLIOGRAPHY. 


Epwarps, Henry, Bibliographical Catalogue of the Described Transformations 
of North American Lepidoptera. Bulletin of the United States 
National Museum, No. 35; Washington, 1889. 

A list of references to descriptions of North American Cater- 
pillars, complete to its date. 

Dyar,H.G. A List of North American Lepidoptera, and Key to the Literature 
of this Order of Insects. Bulletin No. 52 of the United States National 
Museum, Washington, 1902. 

Refers to the Bibliographies of American Lepidoptera. 


DESCRIPTIONS OF CATERPILLARS. 
(Only a few of the most important titles are given, see Edwards, cited above) 


Lepidoptera. 
Buck ter, W., The Larvae of the British Butterflies and Moths; edited by 
George T. Porritt. London. Completed in 1901. 

Contains 154 plates, illustrating very fully the British cater- 
pillars, but with comparatively little descriptive matter. 
Horan, Ernst, Die Raupen der Grossschmetterlinge Europas; Stutt- 

gard, 1893. 
Describes and figures the known European Macrolepidop- 
terous caterpillars. 50 plates. 
Forses, Wo. T. M., Field Tables of Lepidoptera, Worcester, 1906. 
Contains an analytical key to the better known macrolepidop- 
terous caterpillars of Eastern North America, arranged artificially 
for convenience in identifying specimens; also their foodplants; 
and the dates of appearance of a number of the more generally 
interesting ones. 


Rhopalocera. 
ScuppER, S. H., The Butterflies of the Eastern United States and Canada, 
Cambridge, 1889. 
A mongoraph, as complete as it could be made. Colored 
plates. 
Epwarps, W. H., Butterflies of North America, Boston and New York. 
Completed in 1987. 
Sphingidae. 
FERNALD, C. H., The Sphingidae of New England. Orono, 1886. 
Describes the species then known, and gives a few figures. 
Exror, Ipa M. and Souteg, C. J., Caterpillars and Their Moths, New York, 
1902. 
Although popularly written, this is the authority to go to for 
the sphinx caterpillars of Eastern North America. Photographs. 
Roruscuitp, W., and Jorpan, K., A Revision of the Lepidopterous 
Family Sphingidae. Novitates Zoologicae, Supplement to vol. ix., 
Tring, 1903. : 
Describes the known caterpillars of the world, and makes a 
slight attempt to classify them. Bibliography. 
Ceratocampidae. 
PACKARD, ALPHEUS S., Monograph of the Bombycine Moths, Part II. 
Memoirs of the National Academy of Sciences, vol. ix, Washington, 
1905. 
A monograph of the family for North America; with colored 
plates. 
Notodontidae. 
PACKARD, ALPHEUS §S., Part I of the same. Memoirs of the National 
Academy of Sciences. Vol. vii, Washington, 1895. 
A monograph of the family in North America north of 
Mexico. Colored Plates. 


126 Annals Entomological Society of America [Vol. III, 


Syntomidae, Arctiidae and Noctuidae in part. 

Hampson, GeorGeE F., Catalogue of the Lepidoptera Phalaenae in the 
British Museum. London, nine volumes published so far. 

Long descriptions by Dyar of the North American caterpillars, 
and briefer ones by the author, of those of other parts of the world. 

Noctuidae. 

Smitu, J. B., and Dyar, H. G., Contributions toward a Monograph of the 
Lepidopterous Family Noctuidae of Boreal North America; A Revision 
of Acronycta (Ochsenheimer) and of Certain Allied Genera. Proceed- 
ings of the United States National Museum xxi, 1; Washington, 1898. 

Describes and gives a key to all the species, and figures many 
of them. There is also a discussion of the setae as a means of 
classification. 

Dyar, H. G., Descriptions of the Larvae of Fifty North American Noctu- 
idae. Proceedings of the Entomological Society of Washington, iv, 
315; Washington, 1899. 

An artificial key to the species described. 

Geometridae. 

PackKaRpb, A. $., A Monograph of the Geometrid Moths, or Phalawnide, 
of the United States. United States Geological and Geographical 
Survey of the Territories, Report, vol. x, Washington, 1876. 

Describes the species then-known, and figures many: now 
more or less out of date. 

Dyar, H. G., Life-Histories of North American Geometridae. Psyche, 
Cambridge, vols. ix to xiv, 1900 to 1907. 

Gives descriptions of all the stages of the caterpillars studied, 
with their superficial structural characters. There is little or 
nothing about their life history proper. 

Eucleidae. 

Dyar, H. G., and Morron, E. L., The Life-Histories of the New York 
Slug-Caterpillars. Journal of the New York Entomological Society, 
vols. iii to vii, New York, 1895 to 1899. 

Monographs, with colored figures, of the eighteen New York 
species. Important discussions on setae and skin granulations. 
In the same place Dyar describes several exotic slug-caterpillars 
also. 

Dyar, H. G., Annals of the New York Aeademy of Science, viii, 193. 

Cited under ‘‘Setae,’’ below. It summarizes the structure 
of the Eucleid larvae, species by species, paying especial attention 
to the setae. 

Pyralididae, Crambinae. 

Feit, E. P., On Some Grass-eating Insects. Ithaca. 

Descriptions and Life Histories. 

FERNALD, C. H., The Crambidae of North America, Boston, 1896. 

Contains about the same material. 

Se olan : 
yar, H.G., The North American Nymphulinae and Scopariinae. Journal 
of the New York Entomological Society, xiv, 77; New York, 1906. 
Sesiidae. 

BEUTENMULLER, WM., Monograph of the Sesiidae of North America north 
of Mexico. Memoirs of the American Museum of Natural History. 
Vol. I, part vi, 217, New York, 1900. 

The descriptions of the caterpillars by Dyar are unusually 
complete. Keys arranged by structure and by food-plant. 


STRUCTURE, 
General Anatomy. 
LYONNET, Pierre. Traite Anatomique de la Chenille qui Ronge le Bois 
de Saule. La Haye, 1760. 

This contains the only accurate figures I have seen of cater- 
pillar mouth-parts. Also a full description of the external and 
internal anatomy of Cossus cossus. A couple of figures from it are 
copied on plate 10. 


1gto}] A Structural Study of Some Caterpillars 127 


Prolegs. 
Samney it Car J., Om Msekternas Extremiteter samt deras Hufvud och 
Mundelar. Till K. Vet. Akad. Inlemnad d. 11 Januari, 1860. 
Figures caterpillar mouth-parts and legs. 
GoossENs, TH., Les Pattes des Chenilles. Annales de la Societe Entomo- 
logique de France, vii, 385, Paris, 1887. 
He has a pretty series of figures of the true legs as well as the 
prolegs, but unfortunately it is a very superficial study. 
CuHapMAN, T. D., On Some Neglected Points in the Structure of the Pupae 
of Heterocerous Lepidoptera and Their Probable Value in Classifica- 
tion; with some Associated Observations on Larval Prolegs. Transac- 
tions of the Entomologica! Society of London, 1895. 97 to 119, Lon- 
don. 
Refers more especially to the presence of a complete circle of 
crotchets in the lower forms. 
Setae. 
Dyar, Harrison G., A Classification of Lepidopterous Larvae. Annals of 
the New York Academy of Science. viii, 194 to 232, New York, 1895. 
Although he has led up to it in a couple of descriptions in 
Entomologica Americana, this paper of Dyar’s is really the pioneer 
work on caterpillar setae as a means of classification. A large 
number of caterpillars are discussed from this point of view. 
Figures. 
Dyar, Additional Notes on the Classification of Lepidopterous Larvae. 
Transactions of the New York Academy of Science xiv, 49 to 62, New 
York, 1895. 
Additions and corrections to the last, with more attention 
to the first stage. 
PacKarp, A. S., Monograph of the Bombycine Moths, I. See above 
under ‘“‘ Notodontidae.”’ 
PackKarp, A. S. Ona New Classification of the Lepidoptera. American 
Naturalist, xxix, 636, Philadelphia, 1895. 
Gives figures of the mouthparts of Eriocephala (Jugatae). 
Dyar. A Classification of the Lepidoptera on Larval Characters. Ameri- 
can Naturalist. xxix, 1066 to 1072, Philadelphia, 1898. 
Discusses especially the Jugatae, with figures of Hepialus, 
Micropteryx and Eriocephala. 
Dyar. On the Larvae of the Higher Bombyces (Agrotides Grote). Pro- 
ceedings of the Boston Society of Natural History. xxvii, 227, Boston, 
1895. Ks 
Gives the structural characters of a large number of forms, 
of the following families.. Drepanidae, Apatelidae (Noctuidae 
in part), Arctiidae, Nolidae, Lithosidae, Euchromiidae, Euptero- 
tidae (no typical species), Lemontidae, Lymantriidae, Lasiocam- 
pidae, etc. There is also a genealogy and a key to the families. 
Dyar. The number of Stages in Apatelodes torrefacta. Psyche, vii, 316, 
1895. Cambridge. 
With a description and figures of the seta-plan of the first 
stage. 
Dyar. Note on the Head-Setae of Perophora melsheimerii. Journal of 
the New York Entomological Society, iv, 92, New York, 1896. 
Figures the head, numbering the epicranial setae. 
Tutt, J. W. A Natural History of the British Lepidoptera. London, 1900. 
A chapter on caterpillar anatomy with a long discussion of 
the setae. 
Most of Dyar’s descriptions of caterpillars contain more or less in reference 
to their setae. In particular the following may be mentioned: 
Journal of the New York Entomological Society iii, 68 
(Eudeilinea) i111, 17 (Thaumetopoea), iii, 130; iv, 68 (Pericopidae). 
Canadian Entomologist xxvii, 325; xxvii, 159 (Apatelodes); 
Foam, Boxeayiii, IO}. 
Entomological News vi, 38 to 40 and 100. (Pterophoridae). 
Psyche vi, 259 (Clisiocampa). 


128 
Exc, .4 
HyrG. 2. 
(See 
Pyc. 9. 
Fic: 4. 
Fro. ‘5. 
ric. 6. 
BxG. 7. 
ric. 8: 
ric, 9%, 
Fic. 10. 
Fic. 14. 
Fic. 12. 


Annals Entomological Society of America  [Vol. III, 


EXPLANATION OF THE FIGURES. 


PLATE X. 


Front view of head of caterpillar (Apatela), with setae numbered. 
adf., adfrontal sclerite; adj. 1, adj. 7, its setae. 

adj. o., adfrontal puncture. 

jr. 2, frontal setae. fr. o., frontal puncture. 

cl. t., and cl. 1., clypeal setae. 

lbr., labrum. 

ant., antenna. 

md., mandible. md. 1. and md. 11. its setae. 

1 to xt, setae of epicrania. 

Under lip of caterpillar (Catocala). 

S.m., Submentum. 

Ment., Mentum. 

m. t., mental seta. 

c., Cardo. 

St., Stipes. 

7 and 2, Setae of stipes. 

11, Seta of first joint of maxillary palpus. 

mx. p., Free part of maxillary palpus. 

also Figures 3 and 26.) 

Tip of labium of Catocala. 

Seta of sclerite a. 

a. t. b., Sclerite b, perhaps the palpifer. 

b. o., Punctures of sclerite b 

c. o., Puncture of sclerite c. 

seg. 1, Lb. p, seg. 3, The three joints of the labial palpus. 

Sp. Spinneret. 

Seta plan of a typical (Noctuid) caterpillar; metathorax. In these 
diagrams a single segment is represented as if cut on the mid-dorsal 
and mid-ventral lines, and laid flat. The anterior edge is to the left, 
and the mid-dorsal line at the upper edge. 

Seta plan of a middle abdominal segment of the same type. 

Proleg of Adela, with the setae numbered, ventral view. 

Proleg of Gracilaria, lateroventral view. 

Proleg of Noctuid caterpillar, lateral view; seta vili is shown in dotted 
lines as if seen by transparency, and the roots of the hooks are repre- 
sented in the same way. 

Arrangement of hooks on the proleg of a Sphingid or Bombycid 
caterpillar. 


PLATE XI. 


Proleg of Papilio polyxenes, somewhat flattened; seen from the 
median side. 

Proleg of Jasoniades glaucus, split down the side to the planta and 
flattened out. 

Labrum of Acronycta, showing typical arrangement of setae, and one 
of the punctures. 


Fics. 13 to 24. Labra of butterflies: drawn to the same scale. 


Fre. 25. 


3. Anosia plexippus. 19. Cercyonis alope. 
14. Argynnis cybele. 20. Pieris rape. 
15. Euphydryas phaeton. 21. Papilio eae a 
16. Polygonia interrogationis 22. indies 5 Ss glaucus. 
17. Euvanessa antiopa. 23. argyreus tityrus. 


18. Basilarchia disippus. 24. Pholisora catullus. 
Labium of an Elater beetle larva (compare with Figs. 2 and 32). 


Fic. 2 


Fic. 


Fre. 
Fic. 


RIG. 
Exc: 
Fic. 
Fic. 
Fic. 
Fic. 


Fic. 
Fic. 
Fic. 
Fic. 
Fic. 
Fic. 


Fre. 


Fic. 


36. 


37 
38 


39. 
. Sketch of head of Paonias myops, to show the form. 


42 


A Structural Study of Some Caterpillars 12Q: 


PLATE “XIE. 


Tip of left maxilla of a Noctuid caterpillar, caudal aspect. 

qv and v, setae of second segment of palpus. 

palp. 1, First free segment of palpus. 

2, Second free segment of palpus. 

base, Basal segment of galea. 

mx. l., Maxillary lobes. 

l. c., Inner large cone, the outer one is behind the outer maxillary 
lobe. 

SiG, ,Step.GOne: 

s.c¢., Smallcone. The inner one is in front of the inner maxillary lobe. 

Tip of maxilla of Hepialus (Jugatae), caudal aspect. 

A typical mandible (Phlegethontius quinquemaculatus, side view of 
right mandible). 

A typical antenna (Diacrisia virginica), side view of right antenna. 
(The long seta is posterior). 

Arrangement of eyes of right side in the Jugatae (Hepialus hectus). 

Arrangement of eyes of right side in the Frenatz (Demas coryli). 


. Maxilla and labium of a sawfly. The right maxilla isnot shown. The 


part of the maxilla covered by the labium is indicated in dotted 
lines. 
Seta plan of middle abdominal segment of Hepialus hectus; compare- 
Figs. 4 and 5. 
Same of Incurvaria, showing also the dorsal and ventral sclerites. 
Ventral view of pro- and mesothorax of Adela. 
St., Sternum. 


G32 (Comxar 
F., Femur. 
1h, Abillsvey 


tr., Trochanter. 


PLATE XIII. 


Labium and maxillae of Basilarchia disippus. The drawings of lower 
lips were made with the camera lucida, and they are drawn as they 
appeared in the prepared specimens. They are usually somewhat. 
retracted, on one or both sides. In life the labium is usually folded 
so that the spinneret lies at right angles to the rest, and the base 
of the spinneret and the setae az at least, are concealed. 


. Labrum of Libythea celtis, on the same scale as the other butterfly labra- 
. Labrum of Thella ilicis. 


SPHINGIDAE. 
40 and 41. Head, labrum and lower lip of Dolba hylaeus (?). 


43 and 44. Front and labrum more enlarged. 


45 
46 


27 


48 
49 
50 
51 
52 


. Head of Ceratomia amyntor. 
. Head of Deilephila gallii. 
. Head of Pseudosphinx tetrio. 


PLATE XIV. 


and 53. Head and labrum of Amphion nessus. 
. Labrum of Ceratomia amyntor. 

. Labrum of Pseudosphinx tetrio. 

. Labrum of Phlegethontius carolina. 

. Labrum of Deilephila gallii. 


54, 55 and 56. Head labrum and lower lip of Apatelodes torrefacta. 


ENDROMIDAE. 


57. Labrum of Endromis versicolor. 


BOMBYCIDAE. 


58. Labrum of Bombyx mori. 


. OL: 
Ra a7 
>. 63. 


Annals Entomological Society of America [Vol. III, 


LASIOCAMPIDAE, 


_59 and 60. Antenna, and front and labrum of Malacosoma disstria. 


CERATOCAMPIDAE. 


Head of Basilona imperialis. 
Labrum of Anisota senatoria. 
Labrum of Citheronia regalis. 


PLATE XV. 


. Head of Anisota senatoria. 


Labrum of Basilona imperalis. 
SATURNIIDAE. 


}. Head of Hyperchiria io. 
. Maxillae and labrum of a half grown larva, supposed to be H. io. 
s. Maxillae and labium of adult larve of H. io. 


Labrum of Telea polyphemus. That of normal Tropaea luna is similar, 
but lacks the secondary hair. 


70. Maxillae and labium of T. polyphemus. 


Labrum of Tropaea luna. An aberration, apparently due to injury 
and imperfect regeneration. The form is changed, and setae 11t, 
iv and vt are lost on the right side. 


2. Head of Tropaea luna. That of polyphemus is similar. 


NOTODONTIDAE, 


. Front and labrum of Datana integerrima. 
. Front and labrum of Schizura concinna. That of S. badia is quite 


similar. 


. Front and labrum of Cerura, penultimate stage. 


PLATE XVI. 
Head of Melalopha. 


. Ventral proleg of Cerura, extended. 


Anal leg of Datana ministra, penultimate stage, seen from the ventro- 
lateral point of view. 


. Lateral view of stemapod, or anal proleg of Cerura, with the tip of the 


body. 


. Nearly lateral view of anal proleg of Heterocampa guttivitta; about half 


of the row of crotchets is shown. 


. Ventral proleg of the same, half retracted; extended it would resemble. 


Figure 77 quite closely. 


2. Labrum of H. guttivitta. Nadata is quite similar. 
. Labrum of Melalopha. 


GEOMETRIDAE. 


4. Head of Lycia cognataria (7). ; 


85. Lateral view of sixth abdominal segment of the same, showing normal, 


Ennomid position of the proleg, and seta vib. 


). Labrum of the same. 
7. Ventral view of proleg of Zerene catenaria, opened on the outer side 


and flattened, to show the sucker, interrupting the row of hooks. 
THYATIRIDAE. 


8. Heax of Cymatophora (Bombycia) or. Sketch to show form of 


epicrania. 
DREPANIDAE 


. Front view of head of Drepana arcuata. 

. Seta plan of the same. The leg is indicated very diagrammatically. 

. ‘Labrum of the same. 

. Lateral view of ventral proleg, showing the three true setae vit, the. 


outer row of rudimentary hooks, and the two ends of the developed 
inner row. 


Igo] 


. 100 


‘1G, 101 


li02 


LOG, 
ANON 
. 108. 
. 109. 
aD: 
Lael: 
2. 


A Structural Study of Some Caterpillars es 


PLATE XVII. 
LACOSOMIDAE. 


. 93. Ventral view of proleg of half-grown Lacosoma chiridota. 
. 94 and 95. Lower lip and head of L. chiridota. 


THAUMETOPOEIDAE. 


x 96 and 97. Labrum and claw of true leg of Thaumetopoea (Cnethocampa) 


pityocampa. 
* LYMANTRIIDAE. 


. 98. Front and labrum of Hemerocampa leucostigma. 
. 99. 


Labrum of Euproctis chrysorrhea. 


NOCTUIDAE. 

Head of Demas coryli. 

Tip of true leg of D. coryli. The moderately notched claw, and the 
three spatulate setae are typical, but not universal in the Macro- 
lepidoptera. 

to 105. Labra of various species of Acronycta, to show variation within 
the genus. 

Labrum of Arsilonche henrici. 

Head of Hadena (Trachea) turbulenta. 

Head of Feltia sp. 

Part of head of Euxoa sp. 

Front of Cucullia sp. 

The ocelli of Nycteola revayana. (Right side). 

The ocelli of Earias chlorana. 


PEATE, Vil, 
ARCTIIDAE, 


3. Head of Apantesis parthenice (?); the setae are somewhat shorter 


than in life, but not so much so as in most of the figures of heads - 
in this paper. 


. Head of Isia isabella. 

5. Mandible of the same, seen from the inner aspect. 
. Mandible of Apantesis. (Of the opposite side). 
. Front and labrum of Euchaetias egle. 

. Lower lip of E. egle. 


and 120. Front and labrum more enlarged of Halesidota caryae. 


. Labrum of Isia isabella. 


SYNTOMIDAE, 


. Labrum of Ctenucha virginica. 


MEGALOPYGIDAE, 


23. Lower lip of Lagoa crispata. 


Antenna of L. crispata (seen from the ventral side). 


PEATE x 


5. Labrum of Lagoa crispata. 
. Half of the same, more enlarged. 


MICROLEPIDOPTERA. 


. Front view of head of Psyche zelleri. 
. Ventral view of proleg of Thyridopteryx ephemeraeformis. 
. Sketch of lower lip of Cossus cossus, showing the general arrangement 


and proportions of parts. 


132 


Annals Entomological Society of America [Vol. III, 


Fic. 130. Tips of maxillae and labium of Cossus cossus; copied from Lyonnet. 


Fic. 


Fic. 
Fic. 


Fic. 


Fic. 


Fic. 
Fic. 
Fic. 


Fic. 


Fic. 
Fic. 
Fic. 


131. 


132. 
133. 


133. 


135. 
136. 
137. 
138. 


139 


141 
142 
143 


Twice the size of the original engraving. Only a small part of 
Lyonnet’s figure is shown. 

g. Subgalea. 

d. f. Maxillae. 

e. Mentum. 

H. Maxillary palpus. The dotted line runs to the enlarged 

second joint characteristic of Cossus. 

K. Labial palpi. 

I. Sclerite c. at the base of the spinneret. 

T. Large cones. 

Skin of Cossus cossus. Opened from the dorsal side, and with the 
larger muscles removed to show the retractor muscles of the proleg 
(2). The proleg itself is represented by an indistinct ring at the 
right end of the muscle. The midventral line runs just to the left 
of the muscle marked P. This is also a copy of a small part of one 
of Lyonnet’s figures, enlarged about twice. 

Antenna of Melittia cucurbitae. (Sesiidae). 

Antenna of Cacoecia cerasivorana (Tortricidae). 


PLATE XX, 


Various FAMILIES. 


Partly lateral view of spinneret and neighboring parts of Thyridop- 
teryx ephemeraeformis (Psychidae) to show an unusual amount 
of development of the sclerites. 

Ventral view of proleg of Lagoa crispata (Megalopygidae). 

Head of Argyresthia. (Tineina). 

Maxillae and labium of Cacoecia cerasivorana. (Tortricidae). 
Portion of skin of Rhodophora florida (Noctuidae) to show the type 
of granulation. 
and 140. Head and antenna more enlarged of Simaethis oxyacantha 

(Tineina). 

Antenna of Yponomeuta cagnagellus. (Tineina). 

Head of Depressaria putridella. (Tineina). 

Head of Endrosis lacteella.  (Tineina). 


ANNALS FE. S. A. 


Vou. III, PuatEe X. 


a 
y 


= 
o,. 


Hn 09 
ae 


f 
Hh 


H99900 
Wi) i nn 


! 


99, 
a) 


W. T. M. Forbes. 


ANNALS E. S. A. 


VoL. II], PLATE XI. 


Submentu 


W. T. M. Forbes. 


ANNALS E. S. A. VoL. III, PLATE XII. 


W. T. M. Forbes. 


VoL. II, PLATE XIII. 


W. T. M. Forbes. 


Vou. III, PLATE XIV. 


ANNALS E. S. A. 


W. T. M. Forbes. 


ANNALS E. S. A. Vox. III, PLaTe XY. 


W. 7. M. Forbes. 


ANNALS E. S. A. VoL. III, PLATE XVI, 


W. T. M. Forbes. 


ANNALS E. S. A. VoL. III, PLhate XVII. 


-— 


: Awe 
vii . — 
MINS 


hastulifera 


106 


W. 1h M. Forhe &. 


ANNALS E. S. A. Vou. III, PLATE XVIII. 


Apantesis farticeiec? 


W. T M. Forbes. 


ANNALS E. S. A Vou, IIT, PLATE XIX. 


W. T. M. Forbes. 


ANNALS E. S. A. Vor Lil. PuArE xox 


W. T. M. Forbes. 


THE LIFE CYCLE OF HORMAPHIS HAMAMELIDIS. 


By T. H. MorGan and A. F. SHULL. 


Pergande* has described the life cycle of this species as con- 
sisting of seven generations, the first two and the sexual genera- 
tions living on the witch hazel, and the other four on the black 
birch (Betula nigra), the latter four generations consisting of three 
aleurodiforms and one winged migrant. 

Failure on our part to discover the aleurodiform generations 
in localities where the witch hazel was abundant, and the discov- 
ery that the winged migrants continued to emerge from the galls 
from the end of July until October, thus leaving no time for the 
intercalation of the four birch generations, led us to examine the 
life cycle of this species in the vicinity of New York. 

Leaves of the witch hazel bearing galls of Hormaphis were 
enclosed in bags of paraffin paper about the first of August, when 
the migrants had begun to leave the gall. The bags were opened 
at intervals of about a week, and the under surface of the leaves 
examined. Nothing was found until about the first week in Sep- 
tember, when the sexual forms were discovered on the leaves 
which had been in the bags. At the same time the sexual forms 
were also found on the leaves outside. The results show at least 
that the birch is not a necessary link in the life cycle of this species; 
and since no aleurodiform individuals were seen at any time on 
the witch hazel, their necessary occurrence in the life history 
seems, in this locality at least, to be excluded. 

The final and conclusive evidence was obtained from a micro- 
scopic examination of the migrants within the witch hazel galls, 
and of the young stages, pupal and prepupal. Serial sections 
show that all these forms contain embryos that are males or fe- 
males. The determination of the male embryos can be made 
owing to the fact that the testes early develop, and in the embryos 
in the winged stages of the migrant the characteristic spermat- 
coyte divisions can be found. In all of these, as in other aphids, 
a lagging chromosome is present, and two classes of sperm, func- 
tional and rudimentary, result. The determination of the sexual 
female can be made owing to the presence of a sypegesis stage 
in the eggs in the ovary. 


* Pe ‘rgande, Theo, ‘‘The Life History of Two Species of Plant-lice Inhab- 
iting Both the Witch- ae and Birch."’ U.S. Dept. of Agr., Div. of Entomol- 
ogy, Technical Series No. 9, 1901. 


144 


rgto| Life Cycle of Hormaphts hamamelidts T45 


At Cold Spring Harbor, where these observations were made, 
three birches are present: Betula lenta is common, B. populifolia 
is uncommon, and B. luteais rare. B. nigra is said to occur, but 
was not found. Winged migrants were placed on the leaves of 
the first three of these species in the evening; those on leaves of 
trees outside had left by 9 A. mM. the next day; some of those on 
branches in the laboratory remained during the day, and were 
seen as late as 4 Pp. M., but had left by 8 a. mM. on the following 
morning. They appear not to have deposited any young, since 
none appeared on the leaves during the following two weeks. 

A branch of witch hazel with leaves bearing galls, and one 
of a birch (B. lenta), growing near together, were enclosed in the 
same bag. After about three weeks the bag was opened, when 
sexual individuals were found on the witch hazel leaves, but none 
on the birch. 

The observations show that in the vicinity of New York, 
Hormaphis has a much simpler life history than that ascribed to 
this species by Pergande for the vicinity of Washington. It 
would seem to follow, either that further south there is a longer 
life cycle including an alternate host, or that Pergande has inter- 
calated in the life cycle of this species several generations of some 
other (aleurodiform) species. Fortunately the question can be 
very simply decided by making a few serial sections of the winged 
migrants in the galls found in the District of Columbia. 

Pergande’s statement is so definite and detailed that it scarcely 
seems possible that he could be mistaken in regard to the life 
history of the Washington form. For example, he states on page 
17, regarding the third generation; ‘‘Toward the middle or end 
of June the insects cast their third or final skin and assume a 
most remarkable mimicry; in fact, mimic now so closely certain 
Aleurodids that for some time I was completely deceived as to 
their true nature, which only after close examination of numerous 
specimens, in connection with its earlier stages, was disclosed. 
When seen on the leaves they are to all appearances true Aleu- 
rodids, both in shape and size, resembling to some extent the scale- 
like form of Aleurodes corni or related species.’’ The fourth and 
fifth generations are also aleurodiform. ‘‘With the appearance 
of the sixth generation a new cycle of forms begins to make its 
appearance, in which the aspect of the insects has changed com- 
pletely, so much so that the casual observer would fail to trace 
a connection between them and the Aleurodiform generations. 


146 Annals Entomological Society of America  [Vol. III, 


Continued observations, both in the woods and on small potted 
birches to which the insects were transferred, removed, however, 
all doubt as to the close relationship of these aberrant forms. 
This generation develops in time into the return migratory gene- 
ration.”’ 

The return of the migrants to the witch hazel is described by 
Pergande as follows (p. 21): ‘‘Having cast their fourth or final 
skin, they acquire wings and, after feeding for some time to com- 
plete maturity, forsake the birch and migrate back to witch hazel 
to deliver themselves of the ultimate or sexual generation. Each 
migrant, according to size, contains from seven to fifteen or per- 
haps more larvae. Migration continues for about a month and 
a half, according to conditions of the season and other natural 
causes, and commences usually toward the end of August and 
terminates during the early part of-October. In general appear- 
ance they are essentially the same as those of the spring migrant 
from the witch hazel, though they are uniformly smaller * * * .”’ 
From these statements it appears that Pergande has observed 
not only all the intermediate stages between the young of the 
spring migrants from the witch hazcl and the aleurodiform genera- 
tions on the birch, but also the development of the return migrant 
from the aleurodiform individuals on small potted birches in 
confinement. 


AN APPARATUS FOR THE DETERMINATION OF OPTIMUMS 
OF TEMPERATURE AND MOISTURE. 


By Tuomas J. HEADLEE, 


Manhattan, Kansas. 


Pioneer work in economic entomology consisted in the 
determination of the insect’s life history and habits without more 
than casual regard to the environment in which it lived. From 
time to time, however, various workers have called attention 
to the fatal effects of the extremes of temperature and moisture, 
and a few persons have pursued systematic inquiry into the rela- 
tions existing between temperature and insect life. Others have 
used various arthropods in determining the response of proto- 
plasm to various stimuli. The last two types of investigation 
have proceeded far enough to show that insects in common with 
other organisms have minimum, optimum and maximum rela- 
tions to each important stimulus to which they are subjected. 
More than enough work has been done to show that the life 
economy of the insect depends to a very large extent directly 
and indirectly upon the physical, chemical, and animate environ- 
ment in which it lives, and that no fundamental understanding 
of its life economy can be reached until the effect of its environ- 
ment is understood. 

While the study necessary to the accumulation of sufficient 
data to arrive at such an understanding is one requiring much 
time and expense, certain insects are of such transcendent eco- 
nomic importance that the expenditure of enough time and money 
to make the most exhaustive study is entirely justifiable. Such 
insects have as a rule already received pioneer study and a few, 
owing to their especially marked response to environmental fac- 
tors, have received more fundamental attention. The writer 
was first led to see the necessity for making a more fundamental 
study of highly injurious species by the observation that for cer- 
tain of the insects most injurious to staple crop production— 
insects that exact a yearly toll of millions from the state in which 
he is now located—only inadequate measures of control have 
been devised, although they have been subjects of study for many 
years. 

In making'‘a study of the relation of environmental factors to 
the life economy of insects, either the investigator must deal 
with a sufficiently large number of individuals and instances to 


147 


ANNALS E. S. A. VoL. III, Puate XXI. 
FIG. | 


T. J. Headlee. 


ANNALS E. S. A. Vou. III, PLhaLrE XXII. 


wiAGRAMATIC VIEW OF CONS TANT TEMPERATURE AND MOISTURE INC UBAT OR, 

Fi@ét = TOP VIEW 

FiG2 =FRONT view 

FiG.3 = SIDE viEW 

A =EXHAUST FAN AND PASSAGE WAY TO LEAD BOX 
B =PASSAGE FROM LEAD BOx 

c SPIRAL LIFT 
DB = THERMOSTAT 
—E =EXPERIMENT CHAMBER 
F =LEAD BOX CONTAINING. CALCIUM CHLORIDE 
G =ICE CHAMBER 
H =HYGROSTAT 
| =HEATING COIL 
J =GLASS FRONT OF CHAMBER 
K 
ts 
M 
N 
i?) 
P 
Q 
R 


=>WATER JACKET 

=WEROSENE HEATER 

= VALVE 

=TEMPERATURE-MAGNETIC CIRCUIT 
= " -MOTOR 7) 

= MOISTURE-MAGNE TIC "” 

= " -MOTOR ” 


= MOTORS 
S =ELECTRO-MAGNETS 
T =LEVER U~ ELECTRODE 


FIG.3. 


8) 


_———— 


RRNA |] AAO ALTE 
Q ULL 


T. J. Headlee. 


150 Annals Entomological Society of America [Vol. III, 


reduce the error of the average to a negligible quantity, or he 
must deal with smaller numbers under conditions in which the 
important variables are reduced to constants. 

In planning a study of the life economy of certain insects most 
injurious to staple crop production, the writer has adopted the 
plan of using the smaller number of individuals and of reducing 
the number of variables to a minimum. Of course it has been 
easy to eliminate natural enemies and to prevent large variation 
in the quantity and quality of food supply, but of the physical 
factors he has thus far been able to reduce only temperature and 
moisture to constants. This has been accomplished through the 
construction of an incubator, in which, within limits, desired 
degrees of each can be maintained. 


CONSTRUCTION OF THE APPARATUS. PLATES XXI-X XI. 
TEMPERATURE PHASE. 


Essentially the incubator consists of a water-jacketed chamber 
E (figs. 2 and 3) with special provision for heating and cooling the 
water within the jacket, the whole being surrounded by a box 
fled with non-conducting. packing.. The packing used in this 
‘nstance consisted of wood shavings. The 12” x 12” x 18” cham- 
ber is jacketed on three sides only, the fourth being closed by a 
double glass door for the purpose of admitting light. The water 
is cooled by the inflow of ice water from tank G (Fig. 3). This 
exchange is automatically controlled by means of mercurial 
thermostat D (fig. 2), which projects far into the jacket. 

The platinum-tipped electrode U (fig. 2) has been so adjusted 
that when the temperature of the water within the jacket rises 
higher than is necessary to bring the air in chamber E (fig. 3) 
to the desired point, the rising mercury column in D (fig. 2) makes 
contact with it and completes magnetic circuit N (figs. 2 and 3), 
magnetizing electromagnet 5 (fig. 3), pulling lever T (fig. 3) down 
upon it, thus pulling valve M out of its seat and allowing ice water 
to flow by gravity into the jacket. The pulling of the lever down 
on the electromagnet S (fig. 3) completes motor circuit O (figs. 
1 and 3), and sets spiral lift C (fig. 3) in motion. This interchange 
impelled by gravity is thus hastened by pumping. This inter- 
change continues until enough cold water has been introduced 
into the jacket to cause the mercury column in -D (fig. 2) to with- 
draw from the electrode U (fig. 2). So soon as this happens valve 
M (fig. 3) falls back into its seat and spiral lift C (fig. 3) stops. 


1910] Determining Optimums of Temperatrue and Moisture 151 


The water within the jacket is heated in coil I (fig. 2) by means 
of kerosene burner L (fig. 2). The method of heating could be 
sreatly improved where constant electric current is available by 
the installation of electric heaters under chamber E (fig. 3) and 
the controlling of the amount of current delivered by some form 
of thermostat. This portion of the incubator was devised and 
constructed by the ‘‘International Instrument Company,” and 
later so modified by the writer as to fit it for his use. 


MoIsTURE PHASE. 


While certain companies would undertake the construction 
of constant low temperature incubators, we were unable to obtain 
a combination constant low temperature and moisture incubator. 
On the arrival of the constant low temperature incubator, we set 
about devising a means of bringing the relative humidity under 
control. After trying many things the writer adopted the method 
of placing enough plants or water vessels in chamber E (fig. 3) to 
bring the relative humidity to roo°, then when the relative humid- 
ity reached the desired point to prevent its further rise by passing 
the air over calcium chloride. 

An exhaust fan A (figs. 2 and 3) was placed in the rear wall 
of the chamber E and the air led through a 24” passageway into 
a leaden box F (fig. 3) partly filled with calcium chloride, and 
from there through a similar passageway B (figs. 2 and 3) back 
into the chamber. The fan, passageways, and leaden box are all 
included in the packing space of the incubator wall. <A strand of 
human hair, after having the oil removed from it with sulphuric 
ether, was stretched in an adjustable brass frame. A lever was 
attached by its short arm to this hair in such a way that variation 
in the length of the hair strand would cause the distal end of the 
long arm to move through a considerable space. To this end of 
this arm was attached a platinum electrode which descended into 
a mercury-filled tube. Of course, the contact of the plati- 
num point and the mercury was designed to close magnetic 
circuit P (figs. 2 and 3) and this to close motor circuit Q (fig. 3) 
which would set the exhaust fan in motion. So long as the plati- 
num point remains in the mercury, the fan continues to change 
the air. Less than a minute after the connection is made is 
usually sufficient to dry the air to a point at which the contraction 
of the hair breaks the connection. 


152 Annals Entomological Society of America [Vol. III, 


OPERATION. 


In operating the incubator, it is necessary to set the automatic 
apparatus for both temperature and moisture. The former is 
accomplished by introducing a standard thermometer into 
chamber E (fig. 3) and when the rising temperature reaches the 
desired point, adjusting the electrode so that it almost touches 
the rising mercury column in D (fig. 2). The latter is accom- 
plished by introducing a standardized hygrograph into chamber 
E. (Fig. 3) and when the rising relative humidity reaches the 
desired point, setting the tension of the hair strand so that any 
further loosening and lengthening will allow the platinum point 
to make contact with the mercury. 

So long, then, as the heat maintains a constantly rising tem- 
perature and the ice in the tank is replenished, constant tempera- 
ture will be maintained in chamber E (fig. 3) and so long as a con- 
stantly rising relative humidity is maintained in chamber E 
(fig. 3), and the calcium chloride is replenished, the constant rela- 
tive humidity will be maintained. 


Its Uss. ILLUSTRATED.* 


Inasmuch as the data derived from use of the incubator in the 
study of Toxoptera graminum Rondaniare most available, they will 
be used for illustration. 

The determination of the optimum temperature was first at- 
tempted. The incubators were so placed that they would get 
the same quantity and quality of light. The moisture apparatus 
was set at 75° relative humidity and the temperature apparatus 
at different temperatures: A considerable number of individual 
viviparous Toxoptera graminum were allowed to complete their 
life. cycles. The temperature was then changed and the experi- 
ment repeated. This process was continued until the effect 
of temperatures of 50° F., 70° F., 80° F., and go° F., under con- 
stant relative humidity of 75° had been tested. 

It is generally accepted that the optimum temperature of 
an organism is that temperature under which the organism’s vital 
processes are most active. 

»@ The temperature under which these insects experienced the 
lowest mortality and produced the largest number of healthy 
young in a given time has been considered the optimum tempera- 

5 a The writer desires hereby to acknowledge the aid afforded him by his student assistant, 
Mr. Francis B. Milliken, who under his immediate direction has tested different plans for the 


control of moisture and collected data from the insects under observation while the incubators 
were in operation, 


1910] Determining Optimums of Temperature and Moisture 153 


ture. Chart No. 1, curve No. 4 shows that in our tests Toxoptera 
graminum experiences the lowest mortality at 80° F., curve 
No. 3 shows that the highest daily reproduction is at 80° F., 
curve No. 1 shows that the shortest time from birth to maturity 
occurs at 80° F., and curve No. 2 shows that the average period 


of reproduction is only a little less at 80° F., than at 70° F. 


CHART NO. | 


NO.|= NO.OF DAYS FROM BIRTH TO MATURITY 
NO.2= LENGTH OF REPRODUCTIVE PERIOD 
NO.3= DAILY RATE OF REPRODUCTION 

NO.4= MORTALITY 

NO.5= NO. OF YOUNG 


4 


> 


Be 


ican asc 


OUNG NO.5 


—t 
ew 


a 
i) 
aT 


NO. OF INDIVIDUALS, NO.3 


w 


uw 
i} 
ii: 


AND NO.OF Y 


oe 


ES 
fiers 
DAYS!NO.1,2, 


wn 


I 


60°F. 70°F. 


TEMPERATURE 


Cuart No. 1.—Plotted data showing the relation of Toxoptera graminum 
Rondani to temperature under constant relative humidity of 75°. In curves 
No. 1 and No. 2, point at 50° F. represent the average of 6 individuals, at 70° 
F. 27, and at 80° F. 28. In curves No. 3 and No. 5 point at 50° F. represents 
the average of 6 individuals, at 70° F. 27, at 80° F..28, and at 90° F. 201. In 
curve No. 4 point at 50° F. represents the average of 54 individuals, at 70° F. 
108, at 80° F. 57, and at 90° F. 201. 


90°F, 


Clearly, taking into consideration the effect of higher daily rate 
and shorter period of immaturity on the geometric rate of increase, 
T. graminum will under constant relative humidity of 75° produce 
the maximum number of progeny in a given time at 80° F. It 1s, 
therefore, reasonable to conclude that the optimum temperature 
for T. graminum under 75° relative humidity is about 80° F., 
possibly a little above or a little below. 


NOTES ON CERTAIN SPECIES OF MAMESTRA. 
By Joun B. Smirn, Sc. D. 


In 1852, Guenée described Hecatera laudabilis in Vol. II, p. 
30 of his Noctuelites, and figured it very recognizably on Pl. VIII, 
figure 4. The locality given was “‘Amerique septentrionale,”’ 
Coll. Doubleday, and the larva was described from a figure of 
Abbot. The type is now in the British Museum. 

In 1856, Walker described Hapalia tndicans in Vol. X, p. 359 
of the British Museum Catalogue, and records two specimens, 
and ° from E. Florida, presented by Doubleday. In 1857, in 
Vol. XI, p. 511, of the same publication, Walker refers to Hecatera 
laudabilis Gn., gives a brief latin diagnosis and records 4 examples: 
2 from East Florida out of the Doubleday collection, and 2 with- 
out locality out of the Milne Collection. He apparently had no 
idea that this was the same species that he had described in a 
previous volume. 

In 1868, Grote and Robinson referred tnmdicans as a synonym 
of laudabilis, in the Trans. Am. Ent. Soc. II, p. 78, after examin- 
ing the types, and that reference was accepted by me and more 
recently by Hampson. 

In 1860, Wallengren described Hecatera strigicollis in the Wien. 
Ent. Monatschr., IV, 170, and gave the locality as California. 
That species remained unidentified in our lists until 1891 when, 
in my revision of Mamestra, I re-published the description with- 
out, at that time, suggesting its identity with any other described 
species. In 1893, after seeing the British Museum collections, I 
referred the species, in my Catalogue of Noctuidz to laudabilts, 
and in this reference Hampson has also followed me. 

In 1875, Grote described Mamestra tallaudabilis in Vol. VII, 
p. 27, of the Canadian Entomologist, differentiating it very briefly 
from laudabilis. It is recorded from California and from Van- 
couver Island, out of the Henry Edwards Collection, and both 
sexes were present. In 1881, Mr. Grote lists tllaudabilis as a 
variety of laudabilis, and again points out certain color differences 
between the eastern and western specimens: differences which, 
unfortunately, are not constant nor, as the distribution given 
proves, very useful in separating the species. In my revision of 
1891, I accepted Mr. Grote’s ranking of the species, but was in 
error as to the form to which the name dllaudabilis should be 
applied. In my figure of the genitalia I obviously got hold of 


154 


Igto] Notes on Certain Species of Manestra rss 


an imperfect or broken structure and, while it is reasonably accu- 
rate so far as it goes, it is altogether misleading as representing 
the real structure of the zllaudabilis form which, obviously, I used 
for the dissection. In 1905, Hampson, having before him the 
types of Guenée, Walker and Grote, lists all names under /auda- 
bilis, but as ‘‘Ab. 2, tllaudabilis,’ he designates those forms in 
which the green tinge is almost entirely replaced by white. 

During the two or three years last past it has been my fortune 
to handle very large series of these forms from all parts of the 
country, and the more of them I handled, the less satisfied I be- 
came with the association. Recently, in re-arranging the species 
in this series, I gathered in all my material for comparison, and 
demonstrated to my own satisfaction two very good species; the 
one extending throughout the eastern and southern States and 
into Texas, the other through the Rocky Mountain region into 
Arizona and west to the Pacific Coast. The true laudabilis 1s a 
chunky, heavily built species with comparatively short, broad, 
obtuse primaries. Strigicollis, which must be used for the other 
species, is slighter throughout, the primaries narrower, more 
trigonate, with apices more obvious. In color, laudabilis when 
fresh, is always greenish, fading out to whitish, with the median 
space ranging all the way from reddish to black, often greenish 
below the sub-median vein. The space also tends strongly to 
narrow inferiorly. In both sexes the secondaries may range from 
blackish to almost pure white, and the general impression is that 
of a stout, heavily built insect. Strigicollis, on the other hand, 
never has that delicate green tinge in even the freshest examples, 
and many of them are almost clear white. In others there is a 
mossy Olivaceous tinge which often darkens the normally pale 
portions of the wing. I have never seen a specimen with a red- 
dish median area, but this may range anywhere from olivaceous 
brown to black. The median space while it tends to narrow in- 
feriorly, never approximates the median lines so closely, and does 
not often tend to give a wedge-like impression. The secondaries 
are more uniformly pale in both sexes, and the impression, as 
already stated is of a slighter species than laudabults. 

Finally, as there was plenty of material available, I tested the 
male genitalic structures once more, and demonstrated the dis- 
tinctness of the two series beyond peradventure. A comparison 
of figure 1 with figures 2, 3 and 4, will show that it is not a matter 
of slight differences, but of quite a radical change. In laudabulis, 


156 Annals Entomological Society of America  [Vol. III, 


of which specimens from New York to Texas were examined, 
there was not enough variation to warrant more than a single 
figure. The harpes are very broad at base, with a very narrow 
rather short extension, and a small extension at tip. There are 
two distinct claspers and one of them is spatulate. The corneous 
sheath of the penis is very long in all the examples. 

Seven examples of strzgicollis were studied, coming from Colo- 
rado, Arizona and Utah, and three figures represent all the varia- 
tions found. These are practically all in the size of the penis 
sheath, though none is in the least like that of laudabilis. The 
differences in the uncus are due to differences of position, the 
drawings having been made with a camera lucida. Attention is 
especially directed to the uniformity of the outer angle of clasper 
at point of constriction. ; 

Two examples a co and & from Kaslo, B. C., do not fit into 
the series of strigicollis and, in my opinion, represent a good 
species. 


Mamestra restora n. sp. 


Head and collar pale greenish over white; palpi black at sides, the 
small terminal joint pale; vertex with an admixture of black scales. 
Collar with a distinct black band crossing above the middle. Thoracic 
disc mottled with black, white and pale green. Primaries, the pale 
areas light greenish, basal and median lines white-filled, ornamenta- 
tion otherwise black. All the lines geminate and broken. An irreg- 
ular black spot at the termination of the basal line inferiorly. Median 
space mostly black, marked with mossy green below the sub-median 
vein. The outer part of wing is black powdered, forming a black 
patch at anal angle, more conspicuous than in its allies. The fringes 
are deeply marked with black and narrowly cut with white. Claviform 
deeper black, extending almost across the median space. Orbicular 
small, round, with a blackish central dot and a greenish annulus. 
Reniform large, superiorly dilated, annulate with white, with a mossy 
greenish filling. Secondaries smoky in both sexes; uniformly so in 

paler at base in oc’. Beneath, primaries blackish; secondaries with 
blackish powderings along costa and outer margin, a dusky discal 
spot and an extra-median line. 


Expands 1.08—1.12 inches equals 27-28 mm. 

Habitat; Kaslo; BGs Vln 26, ater. 

One © and one & in good condition. Type of maculation like 
strigicollis, but darker throughout, the paler areas shaded with 
green, the anal angle of primaries with a dark blotch, secondaries. 
dark in both sexes. 


1910] Notes on Certain Species of Mamestra is 


It is doubtful whether I would have dared to describe this as 
a distinct species in spite of its different appearance, were it not 
for the difference in the genitalia of the male. Comparison of 
figure 5, with figures 2, 3 and 4 will show to what I refer. In 
restora the harpes curve evenly to the narrowly extended tip, 
and form no obvious angle, while the extension of tip itself is 
shorter and broader. The claspers are also more slender and 
more separated than in the allied forms, and for the present I 
believe that we have a very good species to deal with, although 
undoubtedly a recent off-shoot from sétrzgicollis. 

Mamestra marinitincta Harvey, was described in 1875, in the 
Bull. Buf. Soc. Nat. Sci., II, 273, and is evidently a local offshoot 
from strigicollis, in a different direction from restora. Here 
the angle of harpes is intensified instead of lessened, and the 
extension of the tip is carried further. The penis sheath is very 
much elongated and more like laudabilis; but with a very long 
irregular series of short spinules running longitudinally. The 
small clasper has been reduced to a mere vestige, while the larger 
has not changed materially. 

Superficially the wing form of strzgtcollis is held, while the 
median area is an exaggeration of the tendency to narrow it in 
laudabilis. Accompanying this is the evening of the median lines 
and the elimination of the second element, so that they are scarce- 
ly geminate except on costal area. Thus far the species is 
recorded from Texas only, and most of the examples are from the 
Belfrage collections in Bastrop Co. In my collection is one 2 
from Kerrville; and one 9 labelled “Tenn.,”’ out of the Kemp 
collection. These two examples are very similar to each other; 
but may possibly be distinct from marinitincta. I do not think 
they are, at present, and await further material for closer study. 

Mamestra spiculosa Grote, is a species that has always been 
rare in collections and the two pairs now before me I owe to the 
kindness of Mr. Doll. The figure of the & genitalia in my 
revision lacks detail, and a better drawing is presented here as an 
illustration of an intermediate form between the olivacea and 
laudabilis types;—the harpes of zllaudabilis with the tip of oliva- 
cea indicated by the drawn out point. The clasper is long, slender 
and single. 

In Mamestra stricta Wlk., and its variety ferrea, the structures 
become more compact except for the clasper, which is longer and 
more slender. I have twenty examples before me illustrating 


158 Annals Entomological Soctety of America  [Vol. III, 


the gradations from the deep red brown to the yellowish brown 
type, and the species as a whole seems quite recognizable and, 
except for this variation in ground, very constant. 

Mamestra circumcincta Smith, was described from two Cali- 
fornian examples representing the two sexes, which I placed with 
stricta on genitalic characters, while comparing it with olivacea 
in fascies. Theo’ type is in my collection, and another co exam- 
ple recently received from San Francisco, California, makes re- 
examination possible, as well as a new figure of the & genitalia 
available. Hampson in his Vol. V, p. 176, makes this a synonym 
of stricta; but in my opinion quite without warrant. The total 
habitus and ground color are different, while the differences 
pointed out in my original description are intensified in the fresh 
example which is darker and more smoky throughout. Neither 
example has in the secondaries any of that yellowish tinge that 
is in all specimens of stricta ever seen by me. As for the genitalia, 
I can claim very little for circumcincta as against stricta. The 
two are very much alike, and such differences as exist might 
easily be within range of variation. A comparison of figures 
8 and g will make this clear. 

Mamestra tenisca, recently described by me in the Proc. N. Y. 
Ent. Soc., is an intensified and enlarged stricta, and I have little 
doubt is mixed with the older species in collections; but I believe 
it to be well distinguished and take this opportunity to offer a 
figure of the male genitalia which, while preserving their close 
resemblance to those of s/ricta, depart noticeably from the type. 
Especial attention is directed on this point to the curved series 
of spinules on the penis sheath, as compared with those in 
stricta and circumcincta. 


EXPLANATION OF FIGURES ON PLATE XXIII. 


Fic. 1. Mamestra laudabilis, from N. Y., Ga., and Fla. specimens. 
Fic. 2. Mamestra illaudabilis, from Ariz. and Denver, Colo., specimens. 
Fic. 3. Mamestra illaudabilis, from Arizona, desert specimens, 

Fic. 4. Mamestra illaudabilis, from Utah specimen. 

Fic. 5, Mamestra restora: from type @. 

Fic. 6, Mamestra maritinitincta: Texas example. 

Fic. 7. Mamestra spiculosa: Arizona. 

Fic. 8. Mamestra stricta. 

Fic. 9. Mamestra circumcincta. 

Fic. 10. Mamestra tenisca: from one of the para-types. 


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"CONTENTS oF THIS NUMBER, ee 


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Volume II] Syetig LEAVE Bsr. bose) Number 3 


THE PALPI OF MALE SPIDERS. 


By JoHN HENRY COMSTOCK. 


INTRODUCTION. 


The remarkable modification of the palpi of the males of 
spiders into organs for the transference of the seminal fluid to 
the female at the time of pairing of the sexes attracted the 
attention of naturalists at a very early date; and the great 
variety of forms presented by these organs has led systematists 
to make much use of them in taxonomic work. In practically 
all of the more important works on the classification of spiders 
there are figures and descriptions of the palpi of males. 

Notwithstanding the general recognition of the value of 
these organs for taxonomic purposes our knowledge of their 
structure 1s very inadequate. Several important contributions 
to this subject have been published and are well known, notably 
those of Westring (’61), Menge (’66), Bertkau (75 and ’78), 
Wagner (’87), Van Hasselt (’89), and Chamberlin (’04 and 
08). Still we find, even in the more recent publications, 
figures of palpi given with almost no effort to identify their 
parts; and even when some of the parts arenamed we find differ- 
ent terms appled to homologous parts in the descriptions of 
different genera. 

The necessity of selecting from the many terms that have 
been proposed for parts of the palpi, a set to be used in a hand- 
book of North American Spiders that the writer has in prepara- 
tion, and the need of terms for parts that have not been des- 
cribed, has led to the preparation of this paper. It is hoped that 
the publication of it may tend to bring about a greater uniform- 
ity in nomenclature and an increased use in systematic works of 
the extremely valuable characters presented by these organs. 


161 


162 Annals Entomological Society of America — [Vol. III, 


THE MORE GENERALIZED TYPES OF PALPI. 


In all spiders the external opening of the reproductive organs 
of the male is on the lower side of the adbomen near its base, 
in the epigastric furrow. Some time before pairing the seminal 
fluid is emitted from this opening and is stored in a tubular 
cavity in an appendage of the last segment of the palpus, where 
it is retained until the pairing of the sexes, and from which it 
then passes to the spermathece of the female. As the object 
of this paper is purely morphological the details of this trans- 
ference of the sperm will not be discussed here; the reader is 
referred to the recently published papers by Montgomery 
(03 and ’10) for a review of the subject and for an account of 
original observations. 


Fic. 1. Tarsus of Filistata hibernalis; Fic. 2. Diagram of the receptaculum 
1, lateral aspect; 2, cblique view; seminis. 
3, mesal aspect. 


The genital appendage of the palpus of the male is exceed- 
ingly complicated in structure in the more specialized spiders, 
as in the Argiopid; but it is comparatively simple in some of 
the more generalized families. A few illustrations of the sim- 
pler forms will be given here. 

Tue FiristatA Type oF PALrus.—lIn Filistata hibernalis, 
which is a very common house spider in the South, is found the 
most simple type of male palpus that I have seen among spi- 


1910} The Palpi of Male Spiders 163 


ders. In the males of this species, the distal end of the last 
segment of the palpus, the tarsus, contains a coiled tube (Fig. 1); 
this 1s the receptaculum seminis (Wagner ’87). The proximal 
portion of this tube is slightly enlarged and ends blindly; the 
distal part is slender and extends through a slender, twisted 
prolongation of the tarsus ending at its tip by an open mouth. 
The modified terminal portion of the tarsus, which contains the 
receptaculum seminis, 1s the genital bulb. By looking directly 
at the tip of the palpus, instead of at one side of it, it can be 
seen that the base of the bulb is situated in a cavity in the end 
of the main part of the palpus (Fig. 1, a.). This cavity is the 
alveolus (Menge ’66). The slender prolongation of the bulb, 
which contains the terminal portion of the receptaculum 
seminis is the embolus (Menge '66; style, Simon, '92). Except 
ing the specialization of the distal end of the tarsus, the segments 
of the palpus of Filistata resemble quite closely the correspond- 
ing segments of a leg, the relative length of the femur, patella, 
and tibia being quite similar; there is not the shortening of the 
tibia, which is so marked in many of the specialized forms, as 
in Aranea for example. 

A study of the palpus of Filistata gives a clue to the probable 
course of the evolution of the genital bulb. It is evident that 
the bulb is a specialization of the tip of the tarsus, and its most 
striking feature is the presence within it of the coiled recepta- 
culum seminis. Regarding the origin of the receptaculum 
seminis, the fact that it is furnished with a transversely striated 
intima, like the intima of a trachea, indicates that it is merely 
an invagination of the body-wall. In its primitive form, it 
was probably a cuplike depression in the tip of the tarsus. 

In its most perfect form, as seen in the more specialized 
spiders, the receptaculum seminis consists of three quite dis- 
tinct parts: first, the proximal end of it, the fundus, is enlarged 
so as to form a pouch, the wall of which is more delicate than that 
of the other parts (Fig. 2, fu.); I have not been able to see 
teenidia in the intima of this part, and infer that it serves as a 
compressible bulb; second, the intermediate portion, the reser- 
voir, is a large coiled tube occupying the middle division of the 
genital bulb (Fig. 2, res.), in this part the tanidia of the intima 
are well-developed and are sometimes very prominent; third, 
the terminal portion constitutes the ejaculatory duct; this is the 
slender tube traversing the apical division of the bulb (Fig. 2, 


164 Annals Entomological Society of America  |Vol. III, 


ej. d.); the wall of this duct is often darkin color, which renders 
it easy to trace the course of the duct in an expanded bulb. 

The tracing of the course of the ejaculatory duct is often the 
only method by which the embolus can be recognized in a com- 
plicated palpus; for when the embolus is: small or when it is 
lamelliform a slender apophysis may be mistaken for it. Even 
Menge ('66), who proposed the term embolus for this part 
labe!s the terminal apophysis as embolus in several of his 
figures of Epeira. 


\ 


Fic. 3. Palpus cf Hypochilus thorelli. Fic. 4. Palpus of Lonxosceles rufescens. 


There is no reason to believe that the lumen of the recepta- 
culum seminis communicates with the body cavity; the meati 
sanguinis described by Wagner (’87) do not exist. 

After the stage represented by Filistata had been reached, 
a shifting in the position of the bulb occurred in most spiders. 
Instead of occupying a terminal position, at the tip of the tarsus, 
it has moved to one side of the tarsus in all spiders known to 
me except Filistata. In the tarantulas and in Hypochilus thor- 
elli, the most generalized in many respects of the true spiders, 


1910] _ Ihe Palpi of Male Spiders 165 


the genital bulb is nearly terminal but 1s, nevertheless, distinctly 
on one side of the tarsus (Fig. 3). In other spiders it has moved 
to a greater or less extent towards the base of the tarsus, which 
it has nearly reached in many, as for example in Lowxosceles 
rufescens (Fig. 4). It has been suggested by Nelson (’09) that 
this shifting of the position of the bulb is for the protection of 
it from mechanical injury. 

In Hypochilus (Fig. 3) and in Loxosceles (Fig. 4), the alveolus 
is comparatively small; but in many spiders it is large, resulting 
in the tarsus being more or less cuplike in form; this is shown in 
some of the figures of the more specialized palpi given later. 
This cuplike form of the tarsus as distinguished from its appen- 
dage, the genital bulb, suggested for it the name cymbium 
(Menge ’66), which is the classical name of a small drinking 
vessel. The term /amina, proposed by Westring (’61) antedates 
cymbium; but I have adopted the later term, as it is the one 
in general use. 

The well-known fact that tarsal claws do not exist on the 
palpi of male spiders is easily understood if we regard the 
genital bulb as a specialization of the tip of the palpus, as is 
indicated by the structure of the palpus of /ilistata described 
above. Sometimes, as in Lycosa, the tip of the cymbium bears 
one, two, or three stout spines; these have been regarded as 
‘transformed claws’? (Chamberlin ’08); it seems more probable 
for the reason given above, that these are secondarily devel- 
oped structures instead of vestigial claws; in fact there are fre- 
quently strong spines distributed over the surface of the cym- 
bium. 

The genital bulb in Flzstata is helicoid; this is due, so far as 
the larger basal part is concerned, to the fact that the wall of 
it 1s molded over the coiled receptaculum seminis; but the 
twisting of the bulb is continued to the tip of the embolus, 
although in this part, the receptaculum seminis is not coiled 
but extends in a nearly direct line. I know of no other case 
where the helicoid form of the genital bulb is so well-marked 
as here; but there is always a more or less spiral arrangement of 
parts. 

THE TARANTULA TYPE OF PALPUS.—In those spiders that 
are commonly known in this country as tarantulas, and which 
represent the more generalized of the two principal divisions of 
the order Araneida, there exists a comparatively simple type of 


166 Annals Entomological Society of America  [Vo.1 III, 


palpus; but in none of them that I have seen, or of which I 
have seen figures, is it as generalized as is the palpus of Filistata. 

In the palpi of the tarantulas, the genital bulb has migrated 
to one side of the tarsus; but it is still near the tip of this seg- 
ment of the palpus (Fig. 5). A striking feature of the bulb is 
that it is divided into two distinct segments. The smaller 
basal segment may be termed the basal division of the bulb 
(Fig. 5, b.d.). The larger segment consists of two parts: a large 
stout part, which may be termed the middle division of the bulb 
(Fig. 5, m. d.), and a slender terminal portion, which may be 
termed the apical division of the bulb (Fig. 5, a. d.); there 1s, 
however, no distinct line between the middle and the apical 
divisions, the one gradually merges into the other; but in the 
more specialized palpi these two divisions are distinctly sep- 
arated. 


Fic. 5. Tarsus of Eurypelma. Fic. 6. Genital bulb of Eurypelma 
californicum. 

In the articulating membrane which joins the bulb to the 
tarsus, there is on one side a distinct sclerite, which can be seen 
by removing the bulb from the alveolus (Fig. 6, pet.); this is 
doubtless homologous with what has been termed the petiole 
(Chamberlin '04) in more specialized palpi. 

The greater part of the wall of the bulb in the tarantula type 
of palpus is very densely chitinized but there is a longitudinal 
area on the concave side of the middle and apical divisions 
which is comparatively soft (Fig. 6, p. p.); it may be that this 
part is distended by blood pressure at the time of pairing, as 
is the hamatodocha in the more specialized palpi; but upon this 
point I have no data. This soft strip may correspond to that 
portion of the spiral type of embolus, described later, that I have 
designated the pars pendula. 


1910] The Palpi of Male Spiders 167 


THE PALpus OF LOXOSCELES.—In certain genera of the true 
spiders, the palpi are as simple as in the tarantula type. In 
Loxosceles of the family Scytodide, for example (Fig. 4), 
although the bulb has migrated nearly to the base of the tarsus; 
the bulb: itself is very simple in structure. The basal division 
of the bulb is. inconspicuous; the middle division is nearly 
spherical, and the apical division is long and slender. Here 
the receptaculum seminis is differentiated into the three parts 
described above; the reservoir is large, while the eee duct 
is very slender. dye 


THE PaLpus oF DyspDERA.—In the 
family Dysderide two quite distinct 
types of palpi occur. In Arzadna the 
palpus resembles very closely that of — 
Loxosceles; but in Dysdera it is of a \ 
very different form (Fig. 7); thisis due \ 
to the fact that the apical division of 
the bulb is not slender, and is sharply 
differentiated from the middle division, 
its wall being much less densely chitin- 
ized. But there is on each margin a 
distinct sclerite; and this part of the 
bulb bears distinct apophyses. At the 
tip of the apical division there appears 
to be the beginning of a separation 
into embolus and conductor. 

A summary of the parts of the tarsus in the more generalized 
types of palpi of males is shown by the following table: 


Fic. 7. Palpus cf Dysdera interrita 


Body of tarsus or cymbium, containing the alveolus. 
Genital bulb. 
Internal parts. 
Receptaculum seminis. 
Fundus. 
Reservoir. 
Ejaculatory duct. 


External parts. 
Petiole. 
Basal division. 
Middle division. 
Apical division or embolus. 


16S Annals Entomological Society of America [Vol. ITI, 


THE INTERMEDIATE TYPES OF PALPI. 


There are palpi which hold an intermediate position as re- 
gards complexity of structure between the comparatively simple 
tarantula type and the exceedingly complex forms to be des- 
cribed later. These intermediate types occur in widely sep- 
arated portions of the araneid series; but agree in their more 
essential characteristics; for sake of brevity, I will discuss only 
a few examples of the intermediate types; and will then pass to a 
description of forms in which the maximum number of parts is 
found. 


emb.--- 


Fic. 8. Tarsus of Atypus bicolor. Fic. 9. Genital bulb cf Pachygnatha, 
extended. 


The most important characteristic of these intermediate 
types is that the apical division of the bulb is separated into 
two, more or less nearly, parallel parts. One of these parts 
contains the ejaculatory duct of the receptaculum seminis, this 
is the embolus (Menge ’66); the other is intimately associated 
with the embolus and is known as the conductor of the embolus, 
or the conductor of the style, or, simply, as the conductor. 

A comparatively simple example of this group of palpi is 
that of Atypus bicolor. Here the terminal part of the con- 


1910] The Palpi of Male Spiders 169 


ductor is a broad concave plate (Fig. 8, con.), in which the ter- 
minal portion of the embolus rests. 

A more complicated form of the apical division of the bulb 
exists in Hypochilus thorelli (Fig. 3). Here the embolus is 
coiled about the conductor, the terminal part of which is con- 
cave so as to support the terminal portion of the embolus; the 
tip of the conductor bears a delicate membranous flap. 

In Hypochilus the tarsus bears a branch which supports 
a prominent bunch of bristles (Fig. 3, p. c. )}; this may be a rudi- 
mentary form of paracymbium, a part that is well developed in 
Pachygnatha. 

A somewhat similar condition exists in Pachygnatha (Fig. 9). 
Here the proximal part of the embolus is coiled about the con- 
ductor, which is a broad twisted plate; and the terminal portion 
of the embolus is supported by the corresponding part of the 
eonductor.2 When at, rest the apical «division of) the bulb 
rests in the concave tip of the cymbium; but in the specimen 
figured the bulb has been extended so to show the parts better; 
and the embolus and conductor have been separated at the tip. 

In Pachygnatha the tarsus is divided into two distinct 
parts, which are joined by a movable articulation at the base. 
The larger part is the cymbium (Fig. 9, cym.) the smaller part, 
the paracymbium (Menge ’66) or the accessory branch of the 
tarsus (Simon ’92) (Fig. 9, p. c.). The cymbium and the para- 
cymbium resemble the other segments of the palpus in the nature 
of their cuticula and in the fact that they are clothed with 
hairs. 

The term conductor is in general use and was substituted 
for the term spermophorum of Menge, which was suggested by a 
misconception of the function of this part. As to the particular 
part to which the term should be applied there is no doubt. 
Menge (66, Plate 15) clearly indicates, in his figures of the 
palpus of Tetragnatha extensa, the part to which he applied the 
term spermophorum; and the term conductor must be applied 
to the homologous part whenever it is used. This, however, 
has not been done; in many descriptions an entirely different 
part has been termed the conductor, merely because it is more 
or less nearly parallel with the embolus. <A discussion of the 
function of the conductor is given later. 


170 Annals Entomological Society of America  [Vol. III, 


THE MORE SPECIALIZED TYPES OF PALPI, 


In the development of the bulb of the male palpus in the 
more specialized families of spiders there has been evolved an 
exceedingly complicated organ, which is difficult to understand, 
on account of its small size and the fact that when at rest it 
is compactly folded. Fortunately when such a palpus is boiled 
in a solution of caustic potash (10%) the bulb expands so that 
its parts can be seen; and if preserved in glycerine, it remains 
flexible, so that it can be easily manipulated. The expanded 
bulbs figured below were prepared in this way. Even with the 
best of preparations, it is sometimes difficult to make out the 
relation of parts; this can be most easily accomplished by the 
use of a stereoscopic binocular microscope. 

The extreme specialization of the palpi of males is marked 
chiefly by the development of hamatodocha, to be described 
later, and by an increase in the number of distinct parts and 
appendages of the bulb. The maximum degree of specialization is 
to be found in the Araneine, of which the palpi of several species 
of Aranea are described later. The understanding of the rela- 
tion of the parts of the bulb in this genus will be facilitated by 
a study first of a more simple form, such as is found in the 
Linyphiide. 

THE LINYPHIA TYPE OF PALPUS.—The very common Liny- 
phia phrygiana will serve as an example of the Linyphiide. 

As in Pachygnatha, just described, the body of the tarsus of 
Linyphia consists of two parts; the cymbium (Fig. 10, cym.), 
and the paracymbium (Fig. 10, p. c.). The alveolus is a circular 
cavity near the base of the cymbium. 

When the bulb is expanded, the three divisions of it are dis- 
tinctly separated, there being a slender neck between the basal 
division (Fig. 10, b. d.) and the middle division (Fig. 10, m. d.), 
and also a similar slender neck between the middle division and 
the apical division (Fig. 10, a. d.). 

The wall of the basal division of the bulb consists of two 
parts; the basal heematodocha, and the subtegulum. 

The basal hematodocha.—The genital bulb is attached to the 
cymbium, within the alveolus, by means of a saclike structure, 
which, ordinarily, is inconspicuous or completely concealed 
by other parts of the bulb, but which is very conspicuous in the 
expanded bulb (Fig. 10, 6. h.). This has been named the 


1910] The Palpi of Male Spiders ils 


hematodocha from the fact that at the time of pairing it is 
distended with blood (Wagner ’97). The wall of the hematodo- 
cha appears to consist of elastic connective tissue; hence the name 
spiral muscle applied to it by Menge is inappropriate. In fact 
no muscle tissue has been found within the genital bulb. As 
similar extensible blood-sacs are present in more distal parts of 
the bulb of many spiders, I suggest that this one be termed the 
basal hematodocha. 


6. d. 


ae ae 
Fic. 10. Expanded bulb cf Linyphia Fie. 11. Palpus cf Linyphia phrygiana 


phrygiana. 


The subtegulum.—The proximal end of the basal hemato- 
docha is attached to the cymbium, the distal end, to a ringlike 
sclerite, for which I propose the term subtegulum (Fig. 10, 
s. teg.). The existence of a sclerite in this position was indi- 
cated by Wagner, and it is lettered in his figures s. teg., but its 
ringlike form has not been described; in fact, Wagner states 
that the hematodocha ends in the tegulum. 

The middle division of the bulb.—The middle division of the 
bulb (Fig. 10, m. d.) is that part which contains the chief portion 
of the receptaculum seminis, the reservoir; its wall is the tegu- 
lum, and it bears an appendage, the median apophysis. 

The tegulum.—The term tegulum was applied by Wagner to 
all of the more densely chitinized parts of the wall of the genital 
bulb; but as it is desirable that the different sclerites should 


172 Annals Entomological Society of America  [Vol. III, 


bear distinctive names, I propose that this term be restricted 
to the sclerite that forms the wall of the middle division of the 
bulb. In Linyphia, the tegulum, in this restricted sense is a 
ringlike sclerite (Fig. 10, teg.). 

The median apophysis.—Arising within the distal margin 
of the tegulum there is an appendage, only the tip of which is 
shown in the view of the bulb figured here (Fig. 10, m. a.); 
this is the median apophysis. In many spiders this appendage 
is very conspicuous; and to it have been applied several names. 
In fact in several instances a writer has applied different names 
to this part in his descriptions of different genera. Among the 
names that have been applied to it are /amella characteristica and 
apophysis mediana (Chyzer et Kulezynski '91), clavis and unca 
(F. O. Pickard—Cambridge '97-’05), and scopus (Chamberlin 
04). The term median apophysis. occurs frequently in descrip- 
tions, and is the older name for this part. 

The median apophysis is articulated to the middle division 
of the bulb near the point from which the apical division arises; 
and in some cases, as in Aranea, it appears to be more closely 
articulated with a basal segment of the apical division, the radix, 
than it is with the tegulum. 

The apical division of the bulb—This division includes that 
portion of the bulb which les distad of the middle division; it 
consists of two subdivisions: the conductor and the embolic 
subdivision. The embolic subdivision is traversed by the ejacu- 
latory duct and is composed of several distinct parts. In fact 
the multiplication of parts of the embolic subdivision is the 
most characteristic feature of the more specialized types of 
palpi as contrasted with the intermediate types described above. 

The conductor.—The conductor (Fig. 10, con.) is easily 
recognized by its relation to the embolus, which rests upon it, 
and by its membranous texture. Its attachment to the middle 
division of the bulb is by means of an exceedingly delicate 
membrane. 

In Linyphia the embolus rests upon the conductor through- 
out its length; but in many genera the palpi of some of which are 
described later, the function of the conductor is evidently to 
protect the tip of the embolus in the unexpanded bulb. In many 
cases the embolus is very long while the conductor is short; 
but in every case the embolus in the unexpanded bulb occupies 
such a position that its tip is protected by the conductor. 


1910] The Palpt of Male Spiders 173 


In most cases the conductor can be recognized at a glance by 
its peculiar texture; sometimes it is chitinized to a considerable 
extent, but even then it usually has a membranous margin; 
and in any case it can be recognized by its relation to the tip 
of the embolus in the unexpanded bulb. 

The embolic subdivision.—Closely connected with the mem- 
branous base of the conductor is the base of a separate subdi- 
vision of the apical division of the bulb; as this portion bears 
the embolus it may be termed the embolic subdivision. 

The radix and the stipes —Immediately following the mem- 
branous neck that connects the middle and the apical divisions 
of the bulb and parallel with the membranous base of the con- 
ductor, there are two segments of the embolic subdivision; to 
the basal one of these I apply the term radix (Fig. 10, va.); and 
maeriessecond, the term. sizpes (Figy 10. sh). * For a, more sdis= 
tinctly segmented condition of the base of the embolic sub- 
division see the figures of Avanea circulata given later (Fig. 18 
and 19), where the corresponding parts bear the same letters. 

The embolus.—The organ through which the ejaculatory 
duct opens, the embolus, is comparatively simple in Linyphia, 
being a short spinelike part (Fig. 10, emd.). 

The lateral subterminal apophysis.—In Linyphia phrygiana 
there is developed a remarkable platelike apophysis, which 
serves to protect the exposed face of the unexpanded bulb. 
In Figure 10 (/. st. a.), only the edge of this apophysis is shown; 
but in Figure 11, the broader face of it is represented. I desig- 
nate this the lateral subterminal apophysis as it occurs on the 
lateral aspect of the unexpanded bulb, and also to distinguish 
it from an apophysis developed on the opposite face of the bulb 
in a subterminal position, which occurs in certain other genera. 

The terminal apophysis.—The embolic subdivision ends in 
a strongly chitinized lobe, which may be designated the terminal 
apophysis (Fig. 10, ¢. a.). To apophyses of this kind Menge 
applied the term retinacula; but as this term predicates their 
function, which in some cases is obviously not that implied by 
the name, I prefer apophysis with a modifying term indicating 
the position of the particular apophysis described. 

THE ARANEA TYPE OF PALpus.—I have selected the palpus 
of Aranea frondosa (Epeira strix) as an example of an extremely 
specialized palpus. In Figure 12 the entire palpus, with the 
bulb unexpanded, is represented slightly twisted so as to show 


174 Annals Entomological Society of America  [Vol. III, 


Fic. 12. Palpus cf Aranea frondosa. Fic. 13. Lateral aspect cf an ex- 
panded bulb cf Aranea frondosa. 


Mic. 14. Mesal aspect cf an expanded Fic. 15. Lateral aspect cf an ex- 
bulb ef Aranea frondosa. panded bulb cf Aranea ocellaia. 


Or 


1910} The Palpi of Male Spiders sr 


the ventral aspect of the proximal segments and the lateral 
aspect of the bulb. | 

The proximal segments of the palpus.—This account of the 
palpi of male spiders is devoted almost entirely to a discussion 
of the parts of the tarsus, the proximal segments being well- 
understood; there are, however, a few features of these segments 
in the aranea type that merit attention here. 

Upon the coxa there is a prominent spur, the coxal spur 
Mare i2e.s.); and wpon the inner side-of the femur mear its 
base, there is a groove, the femoral groove, into which the coxal 
spur fits when the palpusis extended forward. The presence or 
absence of this spur and groove is an important generic char- 
acteristic in the Araneine. 

The patella bears two prominent spines at its apex (Fig. 12, 
p.). This is also true in the males of several other genera; 
but in the greater number of genera of the Araneinze there is 
only a single spine in this position. 

The most striking feature of the tibia is its shortness, 
being of about the same length as the patella. 

The tarsus.—As in Linyphia, the tarsus of Aranea consists 
of two parts; the cymbium and the paracymbium. But in 
Aranea the paracymbium (Fig. 13, p. c.) is merely a prominent 
apophysis arising from the base of the cymbium and is not 
articulated with the cymbium by a movable joint as in Linyphia 
and in Pachygnatha. The alveolus is much more extended than 
itis in Linyphia; here it occupies nearly the whole length of the 
Cymibium (Fig. 13, a.). 

The unexpanded bu!lb.—In the Sea AGS bulb of Aranea 
frondosa, the subtegulum (Fig. 12, s. t2g.), tegulum (Fig. 12, 
ies.), and a terminallobe-of the ee division of the bulb, 
bearing a long and slender terminal apophysis (Fig. 12, ¢. a.) are 
visible. Two prominent appendages can also be seen; the 
median apophysis (Fig. 12, m. a.) and the conductor (Fig. 12, 
con.). 

The expanded bulb.—Two figures of the expanded bulb are 
given here; Figure 13 represents the lateral aspect of the bulb, 
the aspect that 1s exposed when the bulb is not expanded; and 
Figure 14, the mesal aspect, the one that is next the cymbium 
in the unexpanded bulb. 

The basal hematodocha is essentially the same as in Liny- 
pria (Pig; 13-and 14; b. h.). 


176 Annals Entomological Society of America  [Vol. III, 


The subtegulum is a ringlike sclerite but its form is like that 
of a seal-ring being narrow on the mesal aspect of the bulb 
and wide on the lateral aspect (Fig. 14, s. teg.). This wider part 
of the subtegulum 1s all of it that is commonly observed and 
has been termed the lunate plate (Chamberlin ‘()4). 

The specimen represented in Figure 14 was more fully 
expanded than that used for Figure 13. In the more expanded 
specimen there is evident a large heamatodocha between the 
subtegulum and the tegulum; this I designate the muddle 
hematodocha (Fig. 14, m. h.). The dark axial object seen 
through the wall of the middle hematodocha is the fundus of 
the receptaculum seminis (Fig. 14, fw.). 

The tegulum is also a ringlike sclerite, which is broad on the 
lateral aspect of the bulb (Fig. 13, teg.), and is narrow on the 
mesal aspect (Fig. 14, ¢eg.). : 

The median apophysis (Figs. 12 and 13, m. a.) is a conspic- 
uous appendage, which projects from the ventral side of the 
bulb. Although the position of this appendage in Linyphia, 
in which the middle and apical divisions of the bulb are dis- 
tinctly separated, shows that the median apophysis is an appen- 
dage of the middle division, in Aranea it appears to be articu- 
lated with the base of a proximal segment of the apical division, 
the radix. 

The conductor (Fig. 14, cov.) arises at the base of the apical 
division and is closely connected with the tegulum. 

The radix (Fig. 14, va.) is much larger than in Linyphia. 
Here it forms the wall of one side of the basal segment of the 
embolic subdivision of the apical division. That this is the case 
is more clearly shown in the bulb of Aranea circulata (Fig. 18 
and 19, ra.), where the segmentation of the embolic subdivision 
is much more marked. 

The stipes (Fig. 14, st.) is also much larger than in Linyphia; 
it is articulated with the distal end of the radix. Like the 
radix, the stipes forms the walls of one side of a segment of the 
embolic subdivision of the bulb, a fact which is also well shown 
in the bulb of Aranea circulata (Fig. 18 and 19, st.). 

The embolus is borne by the embolic subdivision distad of 
the stipes; it projects ventrad between the aistal end of the stipes 
which is mesad of it, and the conductor, which is laterad of it 
in the unexpanded bulb. In the specimen represented in 
Figure 14, the distal end of the stipes and the embolus have been 
pushed away from the conductor in the expanding of the bulb. 


1910] The Palpi of Male Spiders PEG 


The distal hematodocha.—The most striking feature of the 
embolic subdivision in the aranea-type is the presence of a 
large hematodocha, which when expanded over-shadows all 
other parts. This hamatodocha I designate the distal hema- 
todocha (Fig. 13 and 14, d.h.). It is doubtless due to the devel- 
opment of this hematodocha that the radix and the stipes are 
restricted to one face of their respective segments of the apical 
division in Aranea frondosa, the remaining parts of the wall 
of these segments forming a part of the distal heematodocha. 

The mesal subterminal apophysis.—On the mesal aspect of 
the bulb, there arises from the distal haamatodocha a prominent 
apophysis (Fig. 14, m. st. a.); this may be termed the mesal sub- 
terminal apophysts. 

The lateral subterminal apophysis——On the lateral aspect 
there is also an apophysis borne by the distal hamatodocha 
(Fig. 13, 1. st. a.); this may be termed the lateral subterminal 
apophysts. 

The terminal apophysis.—In Aranea frondosa, the tip of the 
embolic subdivision of the bulb ends 1n aspearshaped apophysis 
(Fig. 13, f. a.); this may be termed the terminal apophysis. 

THE PALPUS OF ARANEA OCELLATA.—A glance at the palpus 
of Aranea ocellata (Epetra patigiata) will show that it is of essen- 
tially the same type as that of Aranea frondosa but is different 
in some details. It is figured here to illustrate the kind of 
variations in form that serve to distinguish closely allied 
species (Fig. 15.) The median apophysis differs markedly in 
form from that of A. frondosa; the tegulum bears a small but 
distinct apophysis; the lateral subterminal apophysis bears two 
prominent teeth; and the terminal apophysis is lacking, the 
embolic subdivision ending in a blunt lobe. 

THE PALPUS OF ARANEA CIRCULATA.—The most striking 
modification of the aranea type of palpus, taking the palpus of 
Aranea frondosa as typical, is that of Aranea circulata, which 
is the most complex palpus that I have studied. *. In the unex- 
panded bulb, there appears to be no resemblance to the bulb of 
Aranea frondosa. In Aranea circulata (Fig. 16 and 17), the bulb 
is very large and the cymbium comparatively small and narrow 
Gauge iy, cym.)s.-Thexbasaly hematodocham (Hic telGs04 jes)" 1S 
conspicuous, which is the result of the other parts of the bulb 
being twisted into unusual positions. The median apophysis 
is large and projects beyond the tip of the bulb (Fig. 16, m. a.). 


178 Annals Entomological Soctety of America 


(Vol. ITI, 


Fic. 16. Unexpanded bulb of 
Aranea circulaia. 


p.m. wf 
vw 
4y 


Fic. 18. Expanded bulb of Aranea 
circulata. 


b, he 


cym.- 


Fic. 17. Unexpanded bulb cf Aranea 


circulata. 


—— -$f, 
r-2.7.a. 


“s 7a. 


Fic. 19. Expanded bulb .of Aranea 
circulata, 


1910} The Palpi of Male Spiders 179 


But the most remarkable feature is an elbowed structure on the 
mesal aspect at the base of the bulb (Fig. 17). The fact that 
the ejaculatory duct can be traced throughout the length of this 
elbowed structure gave the first definite clew to the relations 
of the parts of the bulb. The part containing the ejaculatory 
duct evidently pertains’ to the apical division of the bulb, 
although it appears to arise from the base of the bulb. 

When the bulb of Aranea circulata is expanded and untwisted, 
as occurs in the process of expansion, the relation of the parts 
is more easily seen. Figures 18 and 19 represent two views of 
a preparation of this kind. If Figure 19 be studied it will be 
seen that the relations of parts are essentially the same as in 
Aranea frondosa (Fig. 14); the basal hamatodocha, subtegulum, 
and tegulum follow in the same sequence; the median apophysis 
and the conductor project from beneath the tegulum in the cor- 
responding positions, and the elbowed structure, which in the 
unexpanded bulb appears to arise at the base of the bulb is 
here clearly seen to be the embolic subdivision of the bulb. 
The most remarkable differences are the lack of a prominent 
distal haamatodocha and the fact that the radix (Fig. 19, ra.) and 
stipes (Fig. 19, st.) are each a complete cylinder, instead of 
merely forming one face of the wall of a segment of the apical 
division, as in Aranea frondosa. At the distal end of the stipes, 
between this part and the embolus and the terminal apophysis, 
there is a vestigial distal heamatodocha (Fig. 18. d. h.). 

In this species there is an apophysis which like the median 
apophysis is joined by a flexible articulation to the tegulum 
within the cuplike cavity formed by the distal margin of the 
tegulum (Fig. 17 and 19,°p. m. a.); this may be termed the. 
paramedian apophysis. As I have not found this apophysis in 
other palpi, I do not consider it a fundamental part. 

THE PALPUS OF ARANEA GIGAS.—The preceding species, 
Aranea circulata, and several others, have been separated from 
Aranea by Pickard—Cambridge and placed in the resurrected 
genus Eriophora of Simon. The peculiar form of the genital 
bulb in Aranea circulata appears to sustain this separation. But 
in the palpus of Aranea gigas (Fig. 20 and 21) we find a form in- 
termediate between the aranea type and what may be termed 
the eriophora type. 

In the unexpanded bulb of Aranea gigas (Fig. 20) the parts 
are twisted so as to render the basal hamatodocha conspicuous 


180 Anna.s Entomological Society of America [Vol. III, 


m. St. a. 


\ . 
Fic. 20. Unexpanded bulb of Aranea Fic.§ 21. Expanded bulb of Avanea 


WM civas. gigas. 


emb,-- 


Vil. 


PaGhe Wliaese fh 


pay 
emb--7* vad 4 [7S 


-- (02. 


Fic. 22. Expanded bulb of Dolomedes Fic. 23. Unexpanded bulb of Dolo- 
fontanus. _w medes fontanus. 


1910] The Palpi of Male Spiders 181 


as in Aranea circulata; but otherwise there is little similarity 
in appearance to either this species or to Aranea frondosa. 

In the expanded bulb (Fig. 21) it can be seen that the 
embolic subdivision, is intermediate in form between the two 
types, resembling the aranea type in having a large distal 
hematodocha and a well-developed median subterminal apo- 
physis; and resembling the eriophora type in the form of the 
embolus, which is lamelliform (Fig. 21, 2). In the general view 
(Fig. 21) the embolus is covered by the tip of the stipes. 

THE PISAURID TYPE OF PALPUS.—In the family Pisauride 
there is a type of palpus which, while it resembles the aranea 
type in its more general features, differs from that type in several 
important particulars. The palpus of Dolomedes fontanus 
(Fig. 22 and 23) may be taken as an example of this type. 

A study of an expanded bulb of this species (Fig. 22) 
reveals the following characteristics: There is a well developed 
petiole of the bulb (Fig. 22, pet.), which, in this species, consists 
of two nodes with an unchitinized internode. The subtegulum 
bears very prominent anelli (Fig. 22, an.), which are described 
in a later paragraph. The median apophysis is prominent 
tiie. 22, m..a.).. The conductor (Pig. 22) ‘con:) is extremely, 
membranous. The radix and the stzpes are not developed as 
distinct segments. The embolus is of the spiral type (Fig. 22, 
emb.) The terminal apophysts is modified into an organ for the 
support of the embolus (Fig. 22, ful.), which may be termed the 
fulcrum of the embolus. This type of terminal apophysis has 
been termed, incorrectly, the conductor. The true conductor 
in this species, as in all others studied, is an organ whose func- 
tion is to protect the tip of the embolus in the unexpanded bulb. 
At the base of the terminal apophysis, at the point where the 
embolus arises, there is a lamelliform lateral subterminal apo- 
wrysis. (Bis. 22.1 st. a.) 

In the unexpanded bulb (Fig. 23), the long embolus makes a 
curve in the distal end of the alveolus beyond the end of the 
bulb. The fulcrum is applied against the embolus on its con- 
cave side, and has a furrow on its distal face within which the 
embolus rests. The distal part of the conductor is wrapped 
about the tip of the combined embolus and fulcrum, serving, 
as in all other cases observed, as a protection to the tip of the 
embolus. 


182 Annals Entomological Society of America  [Vol. III, 

THE ANELLI OF THE SUBTEGULUM.—In Aranea, the chiti- 
nized part of the wall of the basal division of the bulb, the sub- 
tegulum, is reduced to a ringlike sclerite (Fig. 24, 1.); but in 
certain other genera, the subtegulum is cup-shaped or basket- 
like. In Agelena nevia (Fig. 24, 2), one side of the subtegulum 
is greatly thickened; at the proximal end of this thickening, 
which corresponds to the lunate plate, there is a condyle, which 
articulates with the petiole; and at the distal end, there is a 
condyle, which articulates with the tegulum. The other side 
of the cuplike subtegulum contains in its wall several parallel, 
incompletely ringlike sclerites; these may be termed the anelli 
of the subtegulum. In Dolomedes fontanus (Fig. 24, 3), the 
anelli of the subtegulum are greatly thickened and form prom- 
inent projecting ridges. 

It is probable that the presence of the anelli of the subte- 
gulum, and their nature when present will afford characters of 
use for taxonomic purposes. 


Fic. 24. Three kinds of subtegulum. 


I. 


THE DIFFERENT TyPEsS OF EmMBOLUS.—The form of the 
embolus varies greatly in different species of spiders. Two prin- 
cipal types can be recognized, the connate and the free; and the 
free type includes three subtypes. 

The connate type of embolus.—In the connate type, the 
embolus is not separate from the middle division of the bulb 
but is merely a more slender continuation of it, as in the taran- 
tulas, Loxosceles (Fig. 4), and Arizadna. 

The free type of embolus.—In the free type of embolus, there 
are one or more movable articulations between the embolus and 
the middle division of the bulb. In the free type, the embolus 
varies greatly in form; but the different forms can be grouped 
under three heads: coniform, lamelliform, and spiral. 

A contform embolus.—In this type, there may be a broadly 
expanded base; but the projecting part of the embolus is.a 
straight or slightly curved cone. The embolus of Aranea 
frondosa (Fig. 25, 1) is an example of this type. 


1910} The Palpi of Male Spiders 183 


A lamelliform embolus.—In this type the embolus 1s flattened, 
and may bear a greater or less number of apophyses; an example 
of this type is found in Lepthyphantes minuta (Fig. 25, 2.) 

A spiral embolus.—In the spiral type, as seen in A gelena, for 
example, the embolus is long, slender and coiled; and, in a well- 
expanded specimen, it is seen to be composed of three distinct 
parts: first, the wall of the convex side is densely chitinized, 
forming a gutterlike sclerite, which may be termed the trunk 
of the embolus (truncus) (Fig. 25, 3, t. e.); second, the greater 
part of the wall of the embolus is membranous, and forms a 
loose flap along the 
concave side of the 
organ, which contains 
the ejaculatory duct; 
Baas tap: (Pic. +25, 3; 
p. p.) may be desig- * 
nated the pars pendula 
of the embolus; third, 
at the distal end of the 
pars pendula, there is 
a triangular, chitinized 
area, through which Fic. 25. Three types of embolus. 
the ejaculatory duct 
opens (Figs. 25, 3, a. s.), this may be termed the apical sclerite 
of the embolus. 

The pars pendula and the apical sclerite may be completely 
withdrawn into the trunk of the embolus, so that only the latter 
is visible; the embolus then appears to be merely a strongly 
chitinized style; it is in this condition that it is usually seen and 
described. 


CONCLUSION. 


In the preparation of this paper many palpi other than 
those figured here have been studied; and it is believed that the 
series examined has been sufficiently large to warrant the con- 
clusions given regarding the fundamental parts of the genital 
bulb. There remains to be determined the manner in which the 
different types of palpi have been specialized in other families of 
the order, and the details of the modifications characteristic of 
genera. This, however, is too great an undertaking to be at-. 
tempted at this time, and must be left for those who monograph 
the different families. 


LS4 Annals Entomological Society of America  [Vol. III, 


I wish, however, to urge the importance of describing palpi 
from expanded specimens. <A large proportion of the figures 
of palpi that have been published, being of unexpanded examples 
show comparatively little of the structure of this organ. The 
labor involved in expanding the bulb of a palpus is very little; 
a preparation can be made in five minutes; and in no other way 
can so much be done to make possible a description that will 
describe. 

The following tabular statement shows the relations of the 
fundamental parts of the tarsus in the more specialized types 
of palpi; not all of these parts are invariably present, and fre- 
quently subordinate apophyses are developed. 


Body of the tarsus. 
Cymbium, containing the alveolus. 
Paracymbium. 
Genital bulb: 
Internal parts. 
Receptaculum semuinis. 
Fundus. 
Reservoir. 
Ejaculatory duct. 
External parts. 
Basal division of the bulb. 
Basal heematodocha. 
Petiole. 
Subtegulum. 
Lunate plate. 
Anelli of the subtegulum. 
Middle division of the bulb. 
Middle haematodocha. 
Tegulum. 
Median apophysis. 
Paramedian apophysis. 
Apical division of the bulb. 
Conductor. 
Embolic subdivision. 
Radix. 
Stipes. 
Embolus. 
Body of embolus. 
Pars pendula. 
Apical sclerite of the embolus. 
Distal hamatodocha. 
Lateral subterminal apophysis. 
Mesal subterminal apophysis. 
Terminal apophysis, sometimes developed into a 
fulcrum. 


1910] 


The Palpi of Male Spiders 185 


In conclusion I wish to acknowledge the very efficient assist- 
ance in the preparation of this paper, of Miss Anna C. Stryke, 
who prepared the illustrations for it. 


NAMES OF THE PARTS OF THE PALPUS AND ABBREVIATIONS USED 
FOR THEM IN THE ILLUSTRATIONS. 


Accessory branch=paracymbium. 
Alveclus, a. 
Anelli cf the subtegulum, an. 


Mesal subterminal apcphysis, m. sf. a. 
Middle divisicn cf the bulb, m. d. 
Middle hamatodccha, m. h. 


Paracymbium, p. ¢. 
Paramedian apcphysis, p. m. a. 
Pars pendula cf the embolus, p. p. 
Patella, p 

Peticle cf the bulb, pet. 

Radix, ra. 

Receptaculum seminis, 7. s. 
Reservcir, res. 

Secpus=median apcphysis. 
Spiral muscle=hematodccha. 
Stipes, st. 

Style=embolus. 

Subtegulum, s. feg. 

Tegulum, teg. 

Terminal apcphysis, f. a. 

Wibia. ¢. 

Trechanter, tr. 

Trunk cf the embclus, f. e. 


Apical divisicn cf the bulb, a. d. 
Apical sclerite cf the embclus, 2. s. 
Basal divisicn cf the bulb, 0. d. 

Basal heematodccha, b. h. 
Clavis=median apcphysis. 

Cenductcr cf the embclus, con. 
Coxayc: 

Cexal spur, c. s. 

Cymbium, cvm. 

Distal hematedccha, d. h. 
Ejaculatcry duct, ej. d. 

Embclic subdivisicn cf the bulb, e. s. 
Embclus, emb. 

Femur, f. 

Fulcrum, ful. 

Fundus cf the receptaculum seminis, fu. 
Lateral subterminal apcphysis, /. st. a. 
Lunate plate=subtegulum in part. 
Median apophysis. m. a. 


BIBLIOGRAPHY. 


Westring, Nicola (1861). Araneae Svecicae Descriptae. Gothoburgi. 

Menge, A. (1866). Preussische Spinnen. Danzig. 

Bertkau, Ph. (1875). Ueber den Generaticnsapparat der Araneiden. 
Naturgeschichte. Jahrgang 41, Band 1. 
Bertkau, Ph. (1878). Versuch einer naturlichen Ancrdnung der Spinnen. 
fiir Naturgeschichte. Jahrgang 44, Band 1. 
Wagner, W. (1887). Copulationsorgane des Mannchens als Criterium ftir die 
Systematik der Spinnen. Horae Societatis Entomolcgicae Rosicae. 22. 

Van Hasselt, A. W. M. (1889). Le Muscle Spiral et la Vésicule du Palpe des 
Araignées Males. Tijdschrift vocr Entcmologie. Deel 32. 

Simon, Eugene (1892). Historie Naturelle des Araignées, deuxieme edition, tome 1. 

Pickard-Cambridge, F. O. (1897-1905). Biclogia Centrali-Americana, Arachnida, 
Viole2 

Montgomery, T. H. (1903). Studies cn the Habits cf Spiders, particularly those of 
the Mating Period. Proc. Acad. Nat. Sci. Philadelphia 1903, p. 59. 

Chamberlin, Ralph V. (1904). Notes cn Generic Characters in the Lycoside. 
The Canadian Entomologist, Vol. 36, p. 145-148, 173-178. 

Chamberlin, Ralph V. (1908). Revision cf North American Spiders of the Family 
Lycoside. Proc. Acad. Nat. Sci. Philadelphia, 1908, pp. 157-318. 

Nelson, Jas. A. (1909). Evolution and Adaptation in the Palpus of Male Spiders. 
Annals of the Entomclogical Scciety cf America. Vol. 2, p. 60. 


Montgomery, T. H. (1910). Further Studies cn the Activities of Araneads, IT. 
Proc. Acad. Nat. Sci. Philadelphia, 1909 p. 548. 


Archiv ftir 


Archiv 


NORTH AMERICAN PANISCINI. 


By RayMonpd DEAN WuitMarsH, B.S 
Massachusetts Agricultural College, Amherst, Mass 


INTRODUCTION. 

The tribe Paniscini is rather a small one in this country, con- 
taining but two genera, and eleven species so far described. 
The external anatomy here given was prepared from our com- 
monest species—Paniscus geminatus (Say)—added to by com- 
parison in some respects with the corresponding structures in 
Opheltes glaucopterus (Linn). It was my intention when I 
commenced this paper to include a key to the species of Paniscus, 
but I have been unable to see specimens of all the species, and 
the entirely inadequate descriptions of several has made it 
impossible to prepare a key that would be strictly reliable. I 
have therefore included the original descriptions of the different 
species here, leaving it to others with better opportunities to 
prepare the key. 

This paper has been prepared under the supervision of Dr. 
H. T. Fernald of the Massachusetts Agricultural College, and 
to him I wish to extend my heartiest thanks for his suggestions, 
verification of observations, and other assistance in the course 
of the work. 


EXTERNAL ANATOMY. 
HEAD. 

The head is of medium size, and of the hypognathous type. 
It is shghtly higher than broad, and when viewed from the front 
appears nearly circular in outline. From above, it appears 
slightly bi-concave; the anterior concavity being lost at the 
level of the insertion of the antennae and below them. 

The compound eyes are large, extending from the top of the 
head to a point almost touching the base of the mandibles. 
Looking from the front these organs occupy half or slightly 
more than half of the front of the head; from the side view the 
eye takes up about two-thirds the width. The anterior margin 
of the eye, about one-third the distance from the top, is sharply 
excavated which is very noticeable. The antennz are inserted 
at the line of these two indentations or very nearit. The lateral 
margins appear nearly straight. Above the point of concavity, 


186 


1910] North American Paniscint 187 


the eye which is elliptical in form extends toward the middle of 
the head, somewhat, thus making the distance between these 
organs at the vertex, a little less than at any other point. 

The cheek, which lies behind the eye, is nearly uniform in 
width from the top to the base, and is only slightly narrower 
below where it extends slightly beneath the eye. The face in 
general is clothed with a short thick mat of sericeous pubescence. 

CLYPEUS. 

The clypeus, which is more or less irregular in outline, being 
rounded above (convex) and nearly straight along its free margin 
occupies about a third of the distance from the antenne to the 
mandibles. On either side there is a projection, which extends 
just below the compound eyes, and about half way between the 
extremities of these projections and the dorsal edge of the cly- 
peus are situated a pair of oval depressions, one on either side 
known as clypeal foveze. The dorsal edge of the clypeus ‘is 
marked by a transverse suture (or rather what was once appar- 
ently a suture, as there are no distinct sutures separating the 
clypeus from the frons) some little distance below the insertion 
of the antenne. Between these there is quite a noticeable 
depression, the lower side of which 1s most pronounced. 


FRONS. 


The frons may be regarded as extending upward from the 
clypeus to the ocelli, with a downward extension on each side 
of the clypeus, where it appears to meet the cheek just below 
the eye. About two-thirds of the whole distance from the 
dorsal line of the clypeus to the ocelli, is the place of insertion 
of the antennze. That portion of the frons, from the antennze 
extending downward, projects in front of the level of the eye, 
while the portion above the antennz is slightly sunken behind 
this level. 

OCELLI. 

The ocelli, which are of nearly equal size, are three in num- 
ber, and are arranged to mark the corners of an equilateral 
triangle. The two lateral (posterior) ocelli nearly touch the 
compound eyes. An imaginary line passing through the pos- 
terior ocelli marks the vertex as treated in this paper. No 
sutures separating the vertex from the posterior portion of the 
head, above, of from the cheeks, behind the eyes, are present, 
and the limits of these parts are therefore somewhat indefinite. 


ISS Annals Entomological Society of America  [Vol. III, 


The portion of the head, showing behind the compound eye, 
is termed the cheek, in this paper. The width of the cheek 
does not vary a great deal. The central portion, however, 
(that portion in the region of the attachment of the head to the 
thorax) is slightly wider than other parts. 


LABRUM. 


The labrum is attached to the lower, inner edge of the cly- 
peus, leaving the outer edge of the clypeus well defined. In 
preserved specimens, the labrum is usually bent backward, at 
nearly right angles to the clypeus, with the mandibles closed 
over it, so that it is not accessible for study. For this reason, 
it has not seemed advisable to use it for analytical purposes. 


MOUTH PARTS. 


None of the mouth parts seem to be useful for the determina- 
tion of the species and their description therefore is not included 
here. I might say, however, that possibly the mandibles, 
labial palpi, and maxillary palpi would be of use in the determi- 
nation as in most of the insects examined, the palpi show to the 
full extent. Maxillary palpi five jointed, labial palpi four 
jointed. 

The mandibles are somewhat curved, having two teeth, 
which are blackish to their bases. When the jaws are closed 
they overlap. 

ANTENN. 

The antenne are long. The basal portion of the bulb of the 
proximal segment or scape is rather small and articulates with 
the head in a socket. Its diameter, at this end, is about equal 
to its length to where it unites with the enlarged portion of the 
scape, but it is somewhat narrower toward the place where it 
enlarges into the scape proper. The bulb appears to be an 
entire segment, but this is not the generally accepted view. 
The scape which is the stoutest segment of the antenne is some- 
what smaller at its base, gradually increasing to the base of the 
next segment, the pedicel. This seems to extend down into the 
scape for some little distance. There is also a third segment 
before we come to the filament proper which is nearly twice as 
broad as long, and is called the ring joint. This seems to be 
rather unusual. The filament which consists of about fifty- 
four segments has those near the base a little longer than the 
others, gradually decreasing in length as we go towards its 
extremity. 


1910] North American Paniscini 189 


THORAX. 


The thorax in general is strongly compressed laterally, 
very finely pubescent, and is of a yellowish russet color. 


PROTHORAX. 


The prothorax, as seen from the side, is triangular in outline, 
marked by fine, distinct, oblique, transverse striations which are 
most noticeable near the lower anterior portion of the plate, while 
viewed from above it is a very narrow transverse band. Its 
anterior margin is straight and elevated, forming a flange, 
which projects slightly forward. Looking down from above, 
there can be easily detected with the aid of a microscope, a 
semicircular indentation or groove, which leads to the query, 
“Was this portion once a distinct plate representing a much 
reduced pronotum while the large triangular plate at the sides 
represented a much enlarged pro-pleuron?”’ It is set out very 
prominently from the mesethoracic plates by quite conspicuous 
sutures. The facial portion of the prothorax, lying just behind 
the head (episternum?) is somewhat hoof-shaped having a 
deeply impressed line extending from the point where the head 
joins the prothorax to a point near the insertion of the fore 
coxa, thus dividing it into right and left halves. 


MESOTHORAX. 


The mesonotum is a broad plate distinctly convex, lying 
between the fore-wings and extending forward to the dorsal 
portion of the prothorax. Starting near the anterior margin 
and at equal distances from the middle of the plates, are two 
lateral grooves known as the parapsidal furrows which curve 
Slightly inward and extend nearly toy the scutellum:, | dihese 
grooves become weak posteriorly, being only slightly noticeable 
after they have passed the middle of the plate. The anterior 
margin of this plate is bent slightly upward or reflexed, and this 
is continued around the sides as far back as the tegula, where it 
leaves the margin and becomes a distinct ridge which passes 
obliquely backward across the plate. 

Directly behind the mesonotum already described, is the 
scutellum, the median portion of which is trapezoidal in outline, 
longer than broad, the longer base anterior, the shorter base 
being posterior and slightly over half the length of the former. 
Between the mesonotum and median portion of the scutellum 


190 Annals Entomological Society of America  {Vol. III, 


is a deep trough-like depression, whose ends are closed by a con- 
tinuation backward onto the scutellum of the ridges, spoken 
of previously, which may be designated as the dorso-lateral 
ridges. On the mesonotum, just in front of the trough and 
branching off from the dorso-lateral ridge is a short oblique ridge 
which extends outward and forward to the insertion of the fore- 
wing. The sides of the trapezoid are continuations backward 
of the dorsal-lateral ridges. Near the rear margin of the scu- 
tellum these become lost. Along this margin there is a reflexed 
edge or flange which extends obliquely towards the insertion 
of the fore-wings. 

The surface of the lateral portions of the scutellum lies at 
a sharp angle with that of the median portion (trapezoidal) 
and extends from the dorsal lateral ridge downward and for- 
ward to the base of the forewing. As already stated the hinder 
margin of this portion is reflexed nearly to the wing attachment. 
The mesothoracic pleuron is a large plate which lies below the 
wings, its posterior margin being approximately indicated by 
an oblique groove extending downward and backward from 
beneath the fore-wing to just beneath the hind wing where it 
bends a little more downward, and takes a nearly straight path 
to the meso-coxa. From here it passes forward, curving some- 
what until about opposite the base of the fore coxa. It then 
turns sharply upward, and passing by the base of the fore-coxa 
follows along the edge of the prothorax to the base of the fore- 
wing. : 

There are several ridges and grooves on this plate, but noth- 
ing which can be termed a distinct suture, that is, dividing the 
pleuron into subordinate plates. The meso-coxa is attached 
near the extreme posterior edge of this plate, occupying about 
one-third of its width, which is somewhat greater ventrally 
than dorsally. 

METATHORAX. 


Just to the rear of the scutellum and below it is a narrow 
plate which extends from one hind wing to the other and is 
known as the post scutellum. The rear margin of this plate 
is reflexed as in the case of the rear margin of the scutellum, 
and extends forward nearly to the base of the hind wings. 

The part of the metapleuron nearest the hind wing (epi- 
sternum) is triangular in outline. The apex of the triangle 
extends ventrally and comes to a point just between the meso- 


1910} North American Paniscint 191 


thoracic pleuron, and the metathoracic epimeron, while the base 
of the triangle lies dorsally below the furrow of the post scutel- 
lum. 

The metathoracic epimeron is a large plate bordered on its 
dorsal side by the median segment, anteriorly by the meta- 
thoracic episternum above, and the mesothoracic pleuron below, 
while posteriorly it bears the coxa of the hind leg. 

The median segment is a large arched plate distinctly 
striate above covering the area between the post scutellum, in 
front, and the petiole behind, and extending transversely from 
the metathoracic pleuron of one side to that of the other. A 
short distance behind the angle where the median segment, 
epimeron and episternum of the metathorax meet is an oval 
stigma on each side. Near the hinder margin of the plate, on 
each side, is a crescentic elevation, immediately behind which is 
a depression of the surface. Extending forward from this cres- 
cent to a point near which the median segment meets, the pos- 
terior dorsal! angle of the metathoracic episternum is an elevated 
area, just below which is situated the dorsal edge of the stigma. 
This elevated area may be used for convenience as indicating the 
division between the dorsum and the sides of the median seg- 
ment, though there is no evidence of a separation of these 
portions into separate plates. 

At the extreme posterior end of the median segment there 
is a semi-circular carina which is quite prominent. The median 
segment is marked by fine, distinct, transverse striations and 
the metathorax, as a whole, is pubescent, and very finely 
punctured. 

ABDOMEN. 


This portion of the body has more or less of a russet brown 
color varying from a light yellowish brown, in some specimens, 
to a reddish brown, in others. The abdomen is composed of 
eight segments in both male and female, and is compressed 
laterally, more so ventrally than dorsally. The segments, as a 
general thing, decrease somewhat in length after the second one 
which is only about half as long as the first, but rapidly increas- 
ing in height up to the sixth inclusive, but the seventh and eighth 
are smaller. 

The median segment has been included with the description 
of the metathorax. While it is undoubtedly the first segment of 
the abdomen, for convenience of description, we may consider 


192 Annals Entomological Society of America {Voll III, 


the segment, behind the thoracic mass, as a whole, as being the 
first segment. The first segment of the abdomen, counted in 
this way, is attached at the ventral side of the posterior end of 
the median segment. It is straight and nearly twice as long as 
the second segment. Just above its attachment to the median 
segment and below the dorsal portion of the posterior carina 
is a small but prominent muscle or funiculus, which is attached 
to the anterior dorsal portion of the ventral plate. A little in 
front of the middle of the segment, on each side, is an oval 
stigma, and situated about two-thirds the distance from the 
stigma to the posterior end of the thorax is an elliptical opening 
which, in some cases, apparently extends clear through the seg- 
ment from side to side. On closer examination one will readily 
detect a thin membrane, stretched across this opening about 
in the middle of the body, separating it completely into right 
and left halves. 

This opening is formed where the dorsal plate bends upward, 
and apparently leaves the ventral plate, only the thin membrane, 
already referred to, forming a connection between the two. 
It then turns downward and becomes closely connected with the 
ventral plate, again extending as narrow bands along the upper 
margins of the same plate nearly to the median segment. 
Viewed from above, the dorsal plate appears to divide just 
behind the funiculus, and extends as arms to the median seg- 
ment, forming the lateral portions already described. 

The ventral plate of the sixth segment is most prominent. 
From this portion of the sixth segment is protruded the ovi- 
positor, and its palpi or feelers, which are hairy, except their 
basal portions which are concealed by the remaining segments. 
The ovipositor and feelers extend quite a distance beyond the 
extremity of the abdomen. On the eighth segment, a pair of 
cerci are situated at its extreme posterior margin, and approxi- 
mately half way between the ventral and dorsal portions of this 
plate. 

In the female the stigmata of the second and third segments 
are well in front of the middle, while the remainder are closer 
to the anterior margin, and none are perceptible on the last 
two segments. In the male the stigmata approach the anterior 
margins of the segments more and more passing backward, 
but they do not come close to it. On the seventh and eighth 
they appear to be absent. 


1910} North American Pantiscini 193 


The genitalia of the male protrude prominently between and 
beyond the last dorsal and ventral plates of the abdomen. The 
last segment in the males being noticeably shorter than the 
others, while in the females the last two are noticeably shorter. 

All of the ventral segments of the abdomen are rather soft 
and membranous. The claspers are large and broad. 

WINGS. 

The wings are quite large and colorless, with the exception 
of a yellowish tinge, produced by the presence of reddish brown 
nerves and numerous short hairs, generally distributed over 
both the upper and under surfaces. Cresson’s nomenclature 
of wings is the one used in this paper. 


FORE WING. 

Separating the marginal, or radial cell, from the median, 
or externo-medial cell, is a well-developed stigma, which is 
long, narrow and yellowish brown in color. It is considerably 
wider beyond its middle than before. The marginal or radial 
cell is lanceolate. The greater part of its anterior and longer 
side is formed by the costa and its remaining basal portion by 
the stigma. Posteriorly it is separated from the disco-cubital 
cell by the first abscessa of the radial nervure which runs from 
the stigma to the apex of the small triangular areolet. It is 
separated from the third submarginal cell by the second abscessa 
of the radial nervure, which is longer than the first, and is quite 
strongly bent at its origin where it forms approximately a right 
angle with the radial nervure. The median cell which is one of 
the largest of the wings is about four times as long as its greatest 
width. This latter dimension, occurring about three-fourths 
of the distance from the base of the wing to the outer end of the 
cell. It is separated from the sub-median cell posteriorly by 
the externo medial nervure and from the disco-cubital beyond 
by the basal nervure. The third side is formed by the costal 
and sub-costal nervures fused. 

The costal cell is absent due to the fusing of the costal and 
sub-costal nervures, the sub-costal portion being however, more 
prominent than the costal. The disco-cubital cell which is 
probably the largest cell of the wing is irregular in outline, is 
bounded by five different nervures, namely, first abscessa of the 
radial, first transverse cubital, disco-cubital, discoidal and basal 
nervures, and a portion of the stigma. 


194 Annals Entomological Society of America  [Vol. III, 


Of the above, not mentioned before, the first transverse 
cubital separates this cell from the areolet. The disco-cubital 
nervure which separates it from the third discoidal cell has quite 
a prominent stump of a vein near its middle known as an abbre- 
viated cubital nervure which extends into the cell. About half- 
way from this stump, to the triangular areolet is a short bulla. 
This cell is separated from the second discoidal cell by the dis- 
coidal nervure and ‘receives but one recurrent nervure at the 
point where the disco-cubital and discoidal nervures meet. 
There is but one completely enclosed sub-marginal cell which is 
regarded as the second or areolet. It is small, approximately 
triangular and in all the specimens I have examined, the second 
transverse cubital nervure which separates this cell from the 
third sub-marginal cell has a bulla extending half way from the 
base to the apex of the triangle, beyond which the vein bends 
slightly, taking a straight line to the apex of the triangle. 

The third sub-marginal cell is separated posteriorly from the 
second apical cell by the cubital nervure, and extends to the 
margin of the wing. Its other bounding lines have already been 
described. Submedian or internomedial cell is quite long and 
narrow, the externo-medial nervure separating it from the 
median in front, and the first abscessa of the anal nervure from 
the anal cell behind. Outwardly it is separated from the second 
discoidal cell by the transverse medial nervure which joins the 
externo-medial nervure just beyond the origin of the basal 
nervure, thus making it post fercal. The second discoidal cell 
is separated posteriorly from the anal cell by the second abscessa 
of the anal nervure which is about one-half the length of the 
first abscessa. Outwardly it is separated from the third dis- 
coidal cell anteriorly and first apical cell posteriorly by the first 
recurrent nervure, that portion of this nervure however sep- 
arating it from the former is much shorter than the latter. The 
first recurrent nervure has a small bulla patch where it joins the 
margin of the wing. The third discoidal cell is separated pos- 
teriorly from the first apical cell by the first abscessa of- the 
subdiscoidal nervure and outwardly from the second apical 
cell by the second recurrent nervure, at the center of which 
is a distinct bulla patch, while at its anterior extremity just 
below the areolet is another small bulla. The second apical 
cell is separated posteriorly from the first apical cell by the 
second abscessa of the sub-discoidal nervure which is much 


1910] North American Paniscint 195 


shorter than the first abscessa, and extends to the margin of the 
wing. The anal cell is the longest cell of the wing. It is bound- 
ed, anteriorly, by the anal nervure and, posteriorly, by the 
margin of the wing which has a small fold near the extremity of 
the cell (frenal fold.) This cell extends to a point slightly 
beyond the outer end of the second discoidal cell. The first 
apical cell has already been described with the exception of 
mentioning that its posterior and outward boundary is formed 
by the margin of the wing. 
HIND WING. 


The costal cell is quite long extending somewhat over half 
the entire length of the wing, and it is, by far, the narrowest cell. 
It is separated, posteriorly, from the median cell, by the sub- 
costal nervure. Anteriorly it is bordered by the costal nervure 
which extends somewhat less than half the length of the cell, 
and by the margin of the wing, which is a continuation of the 
latter. The marginal or radial cell is separated posteriorly 
from the median cell by the first abscessa of the marginal or 
radial nervure, and from the submarginal or cubital cell by the 
second abscessa of the same nervure. Its anterior and outward 
boundary 1s formed by the margin of the wing. 

Six frenal hooks are found near the middle of the anterior 
margin of this cell. Submarginal or cubital cell is separated 
posteriorly from the discoidal cell by the second abscessa of the 
cubital nervure, inwardly from the median cell by the transverse 
cubital nervure, and extends outward to the margin of the wing. 
The median cell is the longest and largest cell of the wing. It 
is separated posteriorly from the submedian cell by the externo 
medial nervure and from the discoidal by the first abscessa of 
the cubital nervure. The submedian cell is separated poster- 
iorly from the anal cell by the anal nervure and from the sec- 
ond discoidal cell by the first abscessa of the transverse medial 
nervure which is somewhat longer than the second abscessa, 
and forms with it approximately a right angle. Outwardly it is 
separated from the first discoidal cell by the second abscessa 
of the transverse medial nervure. The discoidal cell is separated 
posteriorly from the second discoidal by the discoidal nervure, 
and extends to the margin of the wing. Second discoidal 
cell is separated posteriorly and inwardly from the anal cell 
by the second abscessa of the anal nervure and extends to the 
margin of the wing. 


L9G Annals Entomological Society of America  [Vol. III, 


TEGULA. 
The tegula is a small chitinous plate, more or less triangular, 
with rounded corners, lying over the base of the fore wing. 
It is slightly sericeous, and to some degree arched. 


LEGS. 


The legs are quite long, rather slender, hairy, and all these 
segments beyond the femora are armed with spines to a greater 
or less degree. Coxa more or less bulb shaped. The two tro- 
chanters taken together are as long or a little longer than the 
coxa, and they are of about equal length in the hind leg, while 
in the middle and fore legs the outer segment is only about 
half the length of the other. Femur is second in length to the tibia, 
the longest segment, and in stoutness to the coxa which 1s the stout- 
est segment. The tibiais largest atits distal end. It is armed with 
spines above, below, and on the sides, these showing a partial 
arrangement into longitudinal rows. At its outer end where it 
joins the first tarsal segment is a circlet of spines, two of which 
are much longer and stouter than the others in the posterior 
and middle legs, which are called tibial spurs. The inner spur 
is nearly twice as long as the outer one. These spines are 
clothed with hairs. The fore leg instead of bearing two tibial 
spurs has one long spur-like appendage nearly half as long as the | 
segment, curved for about one-third of its length from the base 
to the extremity, bearing hairs on its inner surface which in 
connection with a corresponding modification at the base of 
the first tarsal segment acts as a cleaning apparatus. 

There are five tarsal segments. The first is by far the long- 
est, being about twice as long as the second, and three times as 
long as the third which is about as long as the fourth and fifth 
taken together. The fourth segment is slightly shorter than the 
fifth. The first three segments are armed with spines beneath, 
on the sides and at their outer ends. The fourth segment is 
the shortest tarsal segment, and is provided with spines at its 
end where it joins the base of the fifth and last segment. The 
fifth segment is not armed with spines except for a few long 
spine-like hairs at its extremity where the pair of well developed 
claws and a pulvillus between them occur. On the under side 
‘of the claw are borne about twenty teeth. 


1910] North American Paniscint . 197 


SEXUAL DISTINCTIONS. 


Aside from the presence of a sting in the females and of 
more or less copulatory organs in the male, other differences 
may be noticed in the sexes. Ocelli seem to be closer together 
in the male, and also to the compound eyes than in the females. 
The male has fifty-seven segments in the antenne, and the 
female fifty-eight, but it would seem probable, considering the 
large number of segments that this might vary somewhat. 


TABLE OF GENERA. 


Terminal tocth of mandibles longer than the inner; cheeks and temples not 
broad; scutellum more or less margined laterally; transverse median nervure dis- 
tinctly postfurcal to the origin of the basal nervure..... Pantscus Gravenhorst. 

Teeth cf mandibles equal in length; cheeks and temples broad; scutellum not 
margined laterally; transverse medial nervure interstitual with the origin of the 
Dasalonervure, rarely <slichtly postiurcals=. .,.24...5 0-000: OPpHELTES Holmgren. 


DESCRIPTION OF SPECIES. 


Opheltes glaucopterus (Linn). Description taken from Provancher. 
Faune Entcmclogique Du Canada, II, 1888, p. 359. 

Female—Length, 68 inch. Black variegated with red. Head varied 
with russet, except a black spot covering all the vertex and the back 
side of the head. Antennae russet, brown at the extremity, almost as 
long as the body. Thorax black; scales clear, a line before and another 
below, two lines on the back of the mesothorax, and its exterior bor- 
ders, also sometimes the scutellum, all the legs with trochanters, the 
abdomen except the last three segments, of a yellowish russet color. 
Wings with the costa and the stigma yellow, the nervures brown; 
areola small, not petiolated, subtriangular; median nervure not appen- 
diculate, arched. Metathorax bearing a little elevated canal in the mid- 
dle, and a carinae on each side with another transversal at the apex, 
these carinaes being elevated in the form of acute tubicles in certain 
spots. Hind part (hind coxae) russet black at the base. 


Paniscus alaskensis, Ashm. 
Proceedings of the Washington Academy of Sciences, IV, 1902, pp. 237-8. 
Male—Length 10 mm. Head yellow, with purplish-brown eyes, and 
very much as in PANISCUS GEMINATUS Say, except that the ocelli are 
not placed on a black spot, and the lateral ocelli do not quite touch the 
eye margin, as in that species. The thorax and abdomen are pale 
honey-yellow; a stripe on sides of prosternum, the lateral margins of the 
mesonotum, and a stripe on the parapsidal furrows behind, are yellow- 
ish-white; the apical transverse carina is indicated only laterally, being 
wholly obliterated medially, the surface of the metanotum before it 
being very finely, transversely aciculate, behind it polished and impunc- 
tate. Wings hyaline, the venation as in P. GEminatus, the costal vein 
and the stigma yellow, the subcostal vein and the internal veins being 
brown or brown black. External claspers similar to those in P. cEm- 
INATUS but slightly narrower. 


LOS Annals Entomological Society of America  [Vol. III, 


Type—Cat. No. 5688, U. S. Nat. Museum. From Kukak 
Bay, July 4. Five specimens. 


Paniscus albotarsatus Prov. 
Faune Entomologique Du Canada, II, 1883, p. 361. 

Male—Length .40 inch. Beautiful yellow, face of a yellowish 
white, the ocelli black, contiguous with each other. Posterior tarsi 
white except at the base of the first segment; their claws (hooks) brown. 
Abdomen elongated, compressed; slightly arched, slim, scarcely tinged 
with brown at its extremity. Wings colorless, nerves brown; stigma 
yellow. Median nervure arched, without a rudiment of a nervure at its 
middle; areola subtriangular, very oblique, very small. G. 

The color almost uniform, color of the species with its posterior 
tarsi which distinguishes it at first sight. In the female the abdomen is 
a little obscure towards the extremity, the ovipositor longer than the 
length of the abdomen. 


Paniscus albovarigatus, Prov. 
Faune Entomologique Du Canada, II. 1883, p. 361. 

Female—Length .38 inch. Uniform, honey-yellow; the face below 
and above the antennae, the orbites all around the eyes, a line on the 
edges of the mesothorax prolonged to the tip of the scutellum, two lines 
upon the back of the mesothorax, a line below the anterior wings, a 
large spot lower upon the flanks (sides) with another smaller one below 
the posterior wings, of a beautiful white. Ocelli not touching the eyes, 
and also distant from each other. Metathorax uniform (simple, even) 
obscurely transversely acieulate; areola of wings small, triangular, 
pedunculated, slightly oblique, abdomen convex, compressed from the 
third segment, brownish in its posterior half. AC. 


Paniscus geminatus (Say). 
Faune Entcmologique Du Canada, IT, 1883. p. 360. 

Female—Length .65 inch with a uniform yellowish russet tinge. 
Face a little clearer; ocelli contiguous with the eyes, and also with each 
other. Wings colorless, nerves black, stigma yellow. Median nervure 
strongly arched, with a rudiment of a nervure very apparent towards 
its middle. Metathorax pubescent, very finely punctured. Legs and 
posterior tarsi a little darker than the rest. Abdomen compressed, 
slightly convex, brownish at-the extremity; ovipositor longer than the 
length of the abdomen. AC. 


Paniscus medius, Ashm. 
Proceediings of the California Academy of Sciences, [V, 1894, p. 128. 

Female—Length 12 mm.; ovip. 2 mm. _ Brownish-yellow; face and 
clypeus yellowish white; ocelli large, prominent, the lateral touching 
the eye; eyes distinctly emarginate within; thorax smooth, trilobed; 
metanotum smooth, without a vestige of carinae; wings hyaline, the 
costa and stigma brownish-yellow, the other nervures black, the dis- 
coidal nervure with a distinct stump of a vein; abdomen twice as long as 
the head and thorax united, strongly compressed, viewed from the side, 


1910] North American Paniscint 199 


not much broadened towards apex; the petiole is a little more than 
one-third longer than the second segment, with the spiracles placed at two- 
fifths its length, the second segment is about two and one-half times as 
long as wide at apex. 

Described from one specimen, from San Esteban (Haines). 

This species resembles somewhat P. texanus, but is larger, 
differently colored, with a stump of a vein in the’ discoidal 
nervure, and with the segments of the abdomen relatively 
different. 


Paniscus melanostigma, Cam. 
Biclogia Centrali Americana; Hymencptera, I, 1886, p. 303. 

Rufo-testaceous, mesonoto nigro; capite flavo; flagello antennarum 
fusco; alis hyalinis, stagmate nigro. Female. 

Long. 15 millim. 

Hab. Panama, Volcan de Chiriqui (Champion). 

Face closely punctured; metathorax closely transversely striated. 
Areolet moderately large, straight, the recurrent nervure received a 
little beyond the middle; there is no branch on the cubital nervure. 


Easily known from its allies by the black stigma and ner- 
vures. The petiole is longer compared to the second segment 
(the relative length of the two being as in P. geminatus) than in 
P. tinctipennis. 


Paniscus nigripectus, Ashm. 
Preceedimgs cf the United States Naticnal Museum, XII, 1889, p. 425. 

Female—Length 16 mm. This species is much more closely allied 
to P. geminatus than is P. texanus, and structurally it is almost iden- 
tical, but the mesonotum, mesopectus, stigma, and veins are black; the 
lateral ocelli touch the eye; the stump of the vein at the middle of the 
cubitus is wanting, and the submedian cell is only one-third the length 
of the transverse median nervure longer than the median cell; the 
second abdominal segment is two-thirds the length of the petiole, the 
spiracles situated at about one-third its length. 


Plabivat.——Pexas: 
Described from one specimen in Belfrage Collection. 


Paniscus ocellatus, Vk. 
Proceedings Entcmclogical Scciety of Washington, Vcl. XI, 1909, p. 211. 
Female—16 mm.; tegument pale castaneous to castaneous, orbital 
margin yellowish, tips of mandibles and edge of ocelli more or less 
blackish; wings faintly cloudy, transparent, stigma and costa pale, 
translucent, other veins mostly brownish or almost black; anterior 
ocellus elliptical, transverse, the shortest distance between the anterior 
ocellus and the nearest point on the eye; antennae 59-jointed, cylindri- 
cal, the joints well defined, the fifth to penultimate subequal and longer 
than wide at base, the apical joint rounded, subconical, a little longer 


200 Annals Entomological Society of America [Vol. III, 


than the penultimate; discocubital vein with a trace of a stump, recur- 
rent vein interstitial with the second transverse cubitus, aerolet quad- 
rangular, metathorax transversely wrinkled above and with a paren- 
thesis-shaped carina on each side; abdomen shining and with a bloom- 
like pubescence that is conspicuous only in certain lights, the rest of 
the body very similarly pubescent, the pubescence pale, seemingly 
tinged with golden. 

Male—Very like the female, but with a yellow face and 56-jointed 
antennae; male paratopotype with a distinct stump of a vein. 


Type.—Female and male and paratopotype, No. 12320, 
U.S. National Museum. 

Type locality.— Wellington, Kans., E. G. Kelley, collector, 
Webster, No. 5431, U.S. Department of Agriculture, Bureau of - 
Entomology. 


Paniscus subfuscus, Cress. ; 
Proceedings cf the Entcmclegical Scciety cf Philadelphia, IV, 1865, p. 57. 

Dull rufo-fuscous, legs and antennae paler; basal segments of 
abdomen piceous; wings hyaline. 

Female—Dull rufo-fuscous, opaque; clypeus pale, pubescent; 
antennae as long as the body, slender, dull fulvous, slightly dusky at 
tips. Thorax with the dorsal lines deeply impressed; scutellum prom- 
inent, carinated on each side; metathorax minutely sculptured, incised 
at base, with a small acute tubercle on each side behind. Wings 
hyaline; nervures fuscous, stigma testaceous; areolet small, subpetio- 
lated, oblique. Legs pale fuscous, the femora slightly dusky. Abdomen 
stout, arcuated, subcompressed, clothed with a very short, appressed, 
pale pubescence; Ist segment elongate, slightly broader at tip, with a 
faint tubercle on each side before the middle; 2nd segment about one- 
third shorter than the Ist; the three basal segments above, except the 
extreme base of the Ist, stained with piceous; apical margins of the 
remaining segments pale; ovipositor about as long as the Ist segment 
of the abdomen. Length 9 lines; expanse of wings 15 lines. 


Collection.—Ent. Soc. Philad. One specimen. 


This does not agree with the description of P. Rufus Brulle, 
which I have not seen, and which is said to inhabit Cuba. 


Paniscus texanus, Ashm. 
Proceedings of the United States Naticnal Museum, XII, 1889, p. 425. 

Female—Length 9 mm. This species has probably been confused 
in our collections with P. geminatus Say; but besides its much smaller 
size 1t can be readily separated by the following differences: 

The third joint of the antennae is only slightly longer than the 
fourth; the mesonotum is smooth without parapsidal grooves; the lateral 
keels of the scutellum become delicate posteriorly; the spiracles of the 
petiole are placed just anterior to the middle; the second abdominal 
segment viewed from above is trapezoidal but slightly longer than wide 


1910] North American Pantscini 201 


and only half the length of the petiole; the basal joint of hind tarsi only 
twice the length of the second; the submedian cell is as long or slightly 
longer than the median cell, the length of the transverse median ner- 
vure; while whereas, in p. geminatus there is always a distinct stump 
of a vein at the middle of the cubitus, in P. texanus it is entirely wanting 
or only the base remains. 


Habitat.—Texas. 
Described from one specimen in Belfrage Collection. 


Paniscus tinctipennis, Cam. 
Biologia Centrali Americana; Hymencptera, I, 1886, p. 308. 

Fuscous, testaceo variegatus; capite flave; pedibus pallide testaceis, 
cox1ix posterioribus femoribusque posticis fuscis; alis fumatis, stigmate 
testaceo. Female, Long. 15 millim. 

Hab. Panama. Volcan de Chiriqui 2000 to 3000 feet (Champion). 

Antennae a little longer than the body, microscopically pilose; the 
flagellum dark fuscous, the scape testaceous. Head yellow, the occiput 
dark fuscous; the ocellar region black; the face obscurely punctured, 
projecting 1n the middle; mandibles testaceous, the tips black; palpi 
yellow. Thorax dark fuscous; a line on the pronotum, the sutures of 
the mesonotum, a mark below the tegulae, the sides of the scutellum, 
and the lower part of the metapleura, pale testaceous. Metathorax closely 
transversely striated. Abdomen fuscous, the base of the first three 
segments and their ventral surface pale testaceous. Wings with a 
fuscous: tinge; areolet minute, not oblique; the recurrent nervure 
interstitial. 


Differs from P. geminatus in the fuscous colour of the body 
in the wings having a decided smoky tinge, in the expansions 
on each side of the metathorax being much more distinct, in 
the abdomen being broader and not so compressed, and in the 
petiole being thicker and shorter compared to the second seg- 
ment. 


INDEX TO NoTEs oF INTEREST ON Paniscus Not INCLUDED IN THIS 
PAPER. 


Nuttal: Insects and Disease, p. 41, 1899. 

Packard: Text book of Entomology, p. 517. 

Proceedings of the Entomological Society of Washington, IV, 
pp. 45-47. 

4. The Canadian Entomologist, XIX, p. 80. 


CON 


202 Annals Entomological Society of America  [Vol. III, 


INDEX TO LETTERING PLATES. 


a, anal cell. 

ai, ae, anal nervure (First and Second abscissas). 
ab, abdomen. 

ac, abbreviated cubital nervure. 

ap, first apical cell. 

api, second apical cell. 

aw, anterior wing. 

b, bulb. 

bn, basal nervure. 

c, costal nervure. 

C1, costal cell. 

cc, Cerci. 

cf, clypeal foveae. 

cl, clypeus. 

cla, claspers. 

c + se, costal and subcostal nervures, blended. 
cu, cubital or submarginal cell. 

cu, Cue, cubital nervure (First and Second abscissas). 
cus, cubital nervure. 

cx, coxa. 

d, disccidal nervure. 

d;, discoidal cell. 

ds, second discoidal cell. 

ds, third discoidal cell 

de, disco-cubital nervure. 

dc;, disco-cubital cell. 

e, eye. 

em, externo-medial nervure. 

em, externo-medial or median cell. 

ep, triangular plate, Epimeron? 

epm, metathoracic epimeron. 

eps, metathoracic episternum. 

epsi, hoof-shaped plate, Episternum? 

f, funiculus. 

ff, frenal fold. 

im, interno-medial or submedian cell. 

m, marginal or radial cell. 

mj}, Me, marginal or radial nervure. (First and Second abscissas). 
mc, median cell. 

md, mandible. 

mn, mesonotum. 

mp, mesothoracic pleuren. 

ms, median segment. 

0, Ovipositor. 

oc, ocellus. 

p, parapsides. 

ped, pedicle. 

pn, pronotum. 

po, palpi of the ovipositor with tactile hairs. 
psct, postscutellum. 

pt, antennal pit. 

pw, po sterior wing. 

r, ring joint. 

re;, first recurrent nervure. 

res, second recurrent nervure. 

s, spiracle of median segment. 

sc, subcostal nervure. 

sd, sdg, subdiscoidal nervure (First and Second abscissas). 
scp, scape. 


1910] North American Paniscini 203 


sect, scutellum. 

sm, submedian cell. 

sms, second submarginal cell or areolet. 
sm;, third submarginal cell. 

sp, spiracle of abdomen. 

st, stigma. 

t, tegula. 

tc, transverse cubital nervure. 

tc,, first transverse cubital nervure. 
tes, secend transverse cubital nervure. 
tm, transverse medial nervure. 

tm, tms, transverse medial nervure (First and Second abscissas). 


EXPLANATION OF PLATES. 
The following figures were drawn with the Camera Lucida. 


PLATE XXIV. 


Fie. 1. Dersal view of the thorax of Paniscus geminatus. 

Fic. 2. Side view of the thorax of Paniscus geminatus. 

Fic. 3. Anterior tibial comb cf Opheltes glaucopterus. 

Fic. 4. Anterior tibial comb of Paniscus geminatus. 

Fic. 5. Side view of the anterior end of the first segment of the abdomen of 


Paniscus geminatus. 
Fic. 6. Dorsal view of the anterior end of the first segment of the abdomen of 
Paniscus geminatus. 


PLATE XXV. 


Fic. 7. Anterior wing of Paniscus geminatus. 
Fic. 8. Posterior wing of Paniscus geminatus. 
Fic. 9. Anterior wing of Opheltes glaucopterus. 
Pic. 10. Posterior wing of Opheltes glaucopterus. 


PLATE XXXVI. 


Fic. 11. Posterior leg of Paniscus geminatus. 

Prec. 12. Clypeus and labrum of Paniscus geminatus. 
Fic. 13. Mandible of Opheltes glaucopterus. : 
Pic. 14. Mandible cf Paniscus geminatus. 


PLATE XXVII. 


Fic. 15. Front view of the head of Paniscus geminatus. 
Fic. 16. Side view cf the head of Paniscus geminatus. 
Pic. 17. Maxillae and labium of Paniscus geminatus. 


PLATE XXVIII. 


Fic. 18. Abdomen of Paniscus geminatus (female). 

Fic. 19. Abdomen of Paniscus geminatus (male). 

Fic. 20. Basal joints of the antenna of Pariscus geminatus. 
Fic. 21. Claw of Opheltes glaucopterus. 

Fic. 22. Claw of Paniscus geminatus. 


VoL. III, PLATE XXIV. 


ANNALS E. S. A. Vor. TL, PEATE Xe. 


R. D. Whitmarsh. 


Vou. II], PLATE XXVI 


R. D. Whitmarsh. 


ANNALS E. S: A VOL. III, PLATE XX VIL 


Fig. 15 


Fig. 16 


Fig. 17 


R. D. Whitmarsh. 


Vou. III, Puate XXVIII. 


Fig. 2 We 


Fig.22 


R. D. Whitmarsh. 


THE REDISCOVERY OF A PECULIAR GENUS AND SPECIES 
OF ORIBATIDAE. 


By H. E. Ewine, 


Ames, Lowa. 


Years ago Mr. Theo. Pergande made a drawing of a very 
peculiar species of the order Acarina. The specimens of this 
species were collected by Mr. E. W. Allis, at Adrian, Mich., 
in October, 1885. This acarid, with many others, it appears, 
was found in connection with a study of the eggs and larve 
of the 17-year locust. ‘Some time later when Mr. C. L. Marlatt 
was ready to publish his bulletin upon the 17-year locust, he 
asked Mr. Nathan Banks to determine the Acarina for him. 
Mr. Banks looked over the mite figures (these were very accu- 
rately drawn, and represented beautifully the characters of the 
different species), and named what he could: It was Mr. 
Banks’ intention to describe these species when he had examined 
the specimens. Later he did describe some of them, but the 
slide of this species he could not find! 

Notwithstanding the fact that this slide was never found, 
the figure and name of this interesting species were published in 
Mr. Banks’ treatise on the Acarina.* Whether or not Mr. 
Banks thought at the time his paper was published, that this 
genus was good, it 1s hard to tell. No comment whatever, is 
made upon this genus or species, and in his key to the genera of 
the Oribatide given on the opposite page from the picture, this 
genus is omitted. 

While looking over some slides in the cabinet of the Depart- 
ment of Zoology, lowa State College, I came across a slide of 
this species which had the following data upon it: ‘‘In galls on 
elm twigs. Ames, Iowa, April 16, 1894.’ The slide contained 
about a dozen individuals well preserved but poorly mounted. 
I dissolved off the cover-glass and obtained the specimens with- 
out injury for my work upon the species. I found that this 
species represents in almost exact detail all the peculiar charac- 
ters shown in Mr. Pergande’s figure just referred to. These 
characters are given in the following pages. 

*A Treatise on the Acarina, or Mites. Proc.-U. S. Nat. Mus., Vol. XXVIII, 
p. 70, Fig. 136. 


209 


210 Annals Entomological Society of America  [Vol. III, 


Now in regard to the question of who is to be given credit 
for the genus Oripoda for which this species stands as a type. 
Mr. Banks writes that he never has seen any of the individuals 
of this species, and that the species never has been described. 
Mr. Pergande was not a special student of the Acarina, and did 
not give a name to the species. While the author himself 
is the only one who has ever actually studied specimens of this 
species with the idea of finding out the systematic value of their 
characters, yet I am not responsible for the name or figure in 
any way. In the International Code, I can find nothing which 
states that the person who first describes, or in any way makes 
recognizable a new species or genus, must have actually seen the 
species he is naming or describing. Morally it does not appear 
proper to describe species upon secondhand information, still, 
some of the greatest advances in ‘science are made in the inter- 
pretation of facts observed by others. Under these conditions, 
it appears to me that to give both Mr. Banks and Mr. Pergande 
credit for the genus and species is the best way out of the diffi- 
culty: in reality the conditions are very similar to a joint author- 
ship. 

The genus Oripoda I count as a very distinct and peculiar 
genus. The form and shape of the cephalothorax are entirely 
different from that of any other species of the Oribatidae; the 
queer shapes and forms of the segments of the second pair of 
legs represent a great exaggeration of the variations from the 
common type, as is shown in a few of the known species; while 
the whole body is drawn out to such an extent as to make the 
creature look out of all proportion when compared with the 
other members of the group. The genus may be described as 
follows: 

Gen. OripopA Banks and Pergande. 

Pteromorphe very long and narrow, extending almost the 
,entire length of the body, and attached to the cephalothorax 
as wellas tothe abdomen. Cephalothorax very long and narrow 
with the sides almost parallel; covered above by a single, 
chitinous plate which projects slightly, hke a hood, over the 
mouth-parts (Fig. 1). Cephalothorax completely bare above 
except fora single pair cf bristles. Pseudostigmata entirely cov- 
ered above by the pteromorphz, and occupying an extreme 
postero-lateral position on the cephalothorax. Tarsi of the first 
two pairs of legs very short and truncate anteriorly (Figs. 5 and 


1910] A Peculiar Genus and Species of Oribatide Daal 


6). Femora of second legs large, flattened and with a lateral 
blade-like expansion (Fig. 5). 

The genus for which Oripoda elongata stands as a type will 
occupy a place next to the genus Gymnobates Banks, a genus 
represented by only two species, both of whitch are American. 
It shares with this genus the elongate form of the body and the 
truncate nature of the tarsi of the first two pairs of legs. The 
pteromorphe represent an exaggeration of the form shown in 
Gymnobates, being extended farther forward and united by a 
transverse ridge at about the middle of the cephalothorax! It 
differs from this genus in having an elongate, cylindrical ceph- 
alothorax, entirely devoid of either the true or the lateral 
lamelle; and in the structure of the second pair of legs. The 
species may be described as follows: 


Oripoda elongata Banks and Pergande. 


Light chestnut brown. 

Cephalothorax very long, cylindrical, fully twice as long as 
broad; sides of cephalothorax almost parallel. No lamelle, 
translamella or lamellar hairs present. A superior pair of 
bristles present, each pectinate, almost straight and about 
as long as the tibia of leg I. There is a slight, chitinous ridge 
connecting the two pteromorphe, which passes transversely 


across the middle of’ the cephalothorax. Pseudostigmatic 
organs, short, capitate and entirely concealed by the ptero- 
morphe. 


Abdomen cylindrical, about twice as long as broad. Ptero- 
morphe very long and narrow, extending the entire length of 
the abodmen and for about one-half the length of the cephalo- 
thorax. The abdomen bears dorsally about twenty rather small, 
slightly curved, pectinate bristles. There is an anterior lateral 
marginal pair; a dorsal pair; a posterior terminal pair; and about 
seven pairs situated upon or near the base of the pteromorphe. 
Genital covers (Fig. 3) each about one-half as broad as long and 
semi-disc-shaped. They are situated over twice their length in 
front of the somewhat larger anal covers. Anal covers (Fig. 3) 
subrectangular in shape and almost approximate to the posterior 
margin of the ventral plate. 

Anterior pair of legs extending about one-half their length 
in front of the tip of the cephalothorax; tarsus of leg I two-thirds 
as broad as long, truncate anteriorly; tibia twice as long as the 


212 Annals Entomological Society of America  [Vol. III, 


tarsus; genual slightly over one-third as long as the tibia; femur 
as long as tibia; coxa almost completely hidden in its socket 
(Fig. +), where it has a double hinge upon which it rotates. 
Second pair of legs (Fig. 5) shorter than the first and normally 
bent at the genual; tarsus of leg II almost as broad as long; tibia 
much broader distally than proximally; genual slightly longer 
than broad; femur of a very peculiar shape, being much flat- 
tened, with an expanded latero-ventral margin; coxa as in leg I. 
The last two pairs of legs are of the common type; the last ex- 
tending slightly beyond the tip of the abdomen. Ungues tri- 
dactyle; dactyles equally developed. Length 0.42 mm.; breadth 
0.22 mm. 

I have examined the mouth-parts of this species in hope that 
they would show some peculiarities, especially since this species 
has a rather extraordinary habitat, but they present the typical 
acarid form (Fig. 1); however, they are somewhat weaker than 
usual... The mandibles (Fig. 2) appear to be normal. The 
internal anatomy presents no special peculiarities. The organs 


are less compact than in the common species, as would be ex-, 


pected, because of the drawn-out form of the body. |The ven- 
triculus, or stomach, and its ceca appear to be rather small, 
but this is probably due to the shrinking of the tissues. In the 
females often there can be seen several large developing ova. 
The ovipositor is rather longer than usual. 


EXPLANATION OF PLATE XXIX. 


Fic. 1. Mouth-parts as seen from below. 

Fic. 2. Side view of one of the mandibles. 

Fic. 3. Ventral surface of the body. 

Fic. 4. The coxe of legs I and II as seen from below, so as to show their dcuble- 


hinged arrangement. 
FIG. Inside of leg IT. 
Fic. 6. Tarsus and tibia of leg I. 


Crt 


ANNALS IE. S. A. Vols Tile “Pisce! XOX: 


Maya I) 


f 


H. E. Ewing. 


A REVISION OF THE AMERICAN SPECIES OF 
PLATYMETOPIUS. 


By E. P. VAN DUZEE, 
Buffalo, N. Y, 


The Hemipterous genus Platymetopius was founded by Bur- 
meister in 1838 in his Genera Insectorum for the European 
rostratus H. S. and vittatus Fabr ( =undatus DeGeer,) the former 
of which should doubtless be taken as the type. It is widely 
distributed in the palsarctic and nearctic regions. Twelve 
species have been recorded from the former and from the latter 
twenty-three are enumerated in the present paper. Three 
additional species have been described from tropical America, 
where the genus may be well represented. 

This genus is easily distinguished from the allied Jassid 
genera by the long pointed vertex, the narrow front and the 
strong elytral venation. There are three anteapical and five 
apical areoles in the elytra of which the outer is broadly triangu- 
lar; the second sector is connected with the branches of.the first 
by two transverse veinlets and there is usually a series of eight 
or ten oblique veinlets crossing the costal areole. The elytra 
in the more typical species are more or less closely inscribed with 
fine brown or fuscous pigment lines and ornamented with a few 
oval or rounded whitish spots placed near the ends of the areoles 
of which those occupying the apex of the anteapical and the 
base of the apical are most regular and persistent. The angled 
base of the front usually has a concentric pale or white line a 
little below the sharfi edge of the head where there is frequently 
a second line; these lines being accentuated by darker borders. 
In the paler faced species these lines may become obscure or 
entirely obliterated. In fully marked species there are usually 
five longitudinal pale vittce on the pronotum and about five 
white calloused spots on the margins of the scutellum of which 
three are on the angles of the posterior field. The characters 
of the genitalia are quite distinctive in some of the species but 
in most they run so close that they are of little value for diag- 
nostic purposes. 

I do not now like to express any very decided views on the 
phylogeny of this genus but in a linear arrangement of the 
genera I would certainly place it between Jfesamia and Delto- 


214 


1910} American Species of Platvmetopius 20 


cephalus. Through the former it is related to the Dorydini 
and through the latter with the Athysanint, while with this last 
tribe it is independently connected by genus Scaphoideus. I 
still adhere to the views expressed by me many years ago as to 
the value of the second cross nervure between the first. and 
second sectors of ‘the elytra as a’ diagnostic character in the 
Jassid genera. It'is but one character: among several which 
unite a fairly homogeneous group of genera with MWesamia 
at one extreme and Deltocephalus at the other. ‘The form of the 
vertex alone can never be used for.separating the Jassid genera 
into groups nor do I know of any other single character so avail-. 
able as is the elytral venation. That certain species of Athysanus 
or Eutetttx show an occasional second cross nervure only proves 
to me that those genera have but recently developed from their 
parent stem and are still plastic... The whole group of genera 
included, by me in the Deltocephalinit and Athysanini seems now 
to, be in their ascendancy and rapidly developing in both species 
and genera, many of which have not yet become well differ- 
entiated. 

In the preparation of this paper I have had before me repre- 
sentatives of ali but two of the described American species, 
modestus and elegans. The latter however is a distinct species 
well known to me and it has been included in the synoptical 
table. Of the others I have examined the types of all but 
acutus Say, madgalensis Prov. and cinereus O. & B., the two 
former of which certainly refer to our most abundant eastern 
species. 

I wish here to extend my thanks to those who have assisted 
me with the loan of material. To Prof. Herbert Osborn I] am 
indebted for a number of his types and the opportunity of exam- 
ining all the material in his collection and in that of the Ohio 
State University at Columbus, Ohio; Dr. Ball has sent me his 
very valuable material in the genus including the types of the 
species recently described by him; and at the suggestion of 
Prof. Baker, Dr. Howard has kindly sent me the types of the 
three Baker species described in 1900 and with them examples of 
ten other species from the National Museum collection. In 
addition to this my own collection contains representatives 
of twenty species mostly in good series. From this material 
I have found it necessary to describe as new four species and 
one variety. 


216 Annals Entomological Society of America  [Vol. III, 


In preparing the following dichotomous table of the species 
I have used the most available characters I could discover. 
Many of these characters are obscure or difficult to express in 
words which is especially true of the markings of the vertex 
although these markings seem to be good and constant charac- 
ters. It must be borne in mind that the vertex in the female 
is always longer and more acute at apex than in the male. 
Unfortunately the type of tenuifrons Baker was received too 
late to allow of its being included in the key to the species but 
its long vertex with a peculiarly broad and rounded apex and 
the brown face will readily separate it from verecundus to which 
it is most closely related. 


KEY TO THE SPECIES OF PLATYMETOPIUS. 


Elytra uniformly whitish-hyaline crossed by three narrow but usually well distin- 
guished fuscous vitte; oblique costal veinlets wanting; vertex rather short, 
1. hyalinus Osb. 


Elytra more or less hyaline, usually pigmented and veined or inscribed with fus- 
cous with the costal areole hyaline and crossed, at least at apex, by ob- 
lique fuscous veinlets; never uniformly hyaline with narrow transverse 
LEGS stad ese dota yh Sap Son an eae Det | ite eR CRUD, RnR rae ae Nh ee il 

1. Elytra quite uniformly colored cr subhyaline with at most but slight veining 
toward the apex; oblique veinlets of the costa wanting antericr to the 
anteapical, arecles. <1 en cs aise eg bates eros eae is Rade eh ett eet eae 
~. Elytra white or whitish hyaline more or less clouded with fulvcus cr cinereous 
and inscribed with fulvous-brown or fuscous, at times almcst entirely 
fulvous or black; oblique veinlets at least indicated in the costal arecle 


Color a polished sub-hyaline brown; vertex large, parabolic, dull fulvous; 
clavus and a broad band across the anteapical areoles which extends 
back along the costa, whitish; face deep black with the edge and an angu- 
late line white, conspicuous; length 4 mm.; inhabits California. 

3. majestus Ball. 

~. Color greenish or creamy white; vertex of the male short, about right-angled, 
of the female strongly produced, lineated at apex with brown and pale; 

elytra slightly infuscated at apex with about three oblique veinlets against 

the anteapical areoles; length3's-4’ mm............ 2. nigriviridis Bail. 

3. Elytra but partially clouded and inscribed, marked with a conspicuous yel- 

low dorsal stripe along the closed elytra; inhabits California. 
4. elegans VanD. 
~, Elytra variously clouded and inscribed but without a conspicuous pale dorsal 
DAT: ae Re Aaa Se Sr A Ne Ama Gres te O pee Oo oS 4. 
4. Vertex flat, Thamnotettix-like, not depressed nor channeled toward its apex, 
anterior margin rounded to the front; pronotum and elytra recalling 
Mesamia vitellina, wanting the white areolar spots of Platymetopius. 
13. dorsalis Ball. 
~, Vertex more or less channeled toward the apex which may be depressed or 


to 


produced and a little elevated; anterior edge thin, acute or subacute... .5. 
5. Face pale or yellow, usually more or less infuscated at base and sides, the for- 
mer frequently marked with an angulate pale line.................+:: 6. 
=, Face entirely brown or fuscous. .... 0. 205. sce cee etc s meste e e ee es 18. 


6. Markings of the vertex in the form of lineations, usually a pale median line 
more distinct anteriorly and edged with brown, and a faint slightly 
divergent line either Sides sc. Si sv. ese wets cv leis ooo eR ee i? 


1910] American Species of Platymetopius 217 


10. 


Pale markings on the vertex taking the form cf a broken transverse vitta 
before the eyes, more cr less complete, the median line shert and apical. 15. 
Elytra subhyaline, either cinereous or fulvous, usually clouded and reticulated 
anterior to the anteapical areoles with the white areolar spots apparent; 
angular white line on the base of the front distinct (except in /atus) ....8. 
Elytra fulvous and cpake anterior to the anteapical arecles, the white areolar 
Spots'obscure in the female, obsoleteinthemales jae sa. see eon 14. 
Genera! color cinereous with the elytral markings and areolar spots distinct; 
beneath mostly black; face distinctly infuscated at base and sides...... 9. 
General color fulvcus with the elytral markings usually indistinct and the 
lower surface pale; face but slightly infuscated on the sides, the angular 
Dasalmanknearlv@OSOleter: sere a cy eee re nen oor ee ier an 10. 
Length 5 mm.; vertex of the male little lenger than the width between the 
eyes, cf the female somewhat longer; inhabits from Canada to Georgia. 

5. acutus Say. 


Length 4 mm.; vertex very Ilcng and pointed, at least twice as long as wide 


between the eyes; inhabits Florida...2.2..........-: 9. slossoni VanD. 
Face obscurely infuscated on the sides, the angular white line at base distinct. 
tke 


Face uniformly yellowish, the angular white line at base obsolete; vertex at 
least twice as long as wide between the eyes; elvtral reticulations and 
areolar spots obsolete at least in the female............ 7. latus Baker. 


Kengthratsleastio mim. ceneral colomtulouse ase seer oe eee nee er 12. 


Length 314-4 mm.; color cinereous, usually tinged with greenish. 
10. cinereus O. & B. 


Male plates short and broad, hardly passing the middle cf the pygofers; 
areolar spcts practically obsolete anterior to the anteapical areoles; 
general color darker, the males clearly marked........ 8. cuprescens Osb. 


Male plates long triangular, but moderately exceeded by the pvgofers....13. 
Last ventral segment of the female scarcely longer than one half its basal 
width; male plates narrow; pronotal vitte usually distinct. 
5. acutus var. dubius VanD. 
Last ventral segment of the female long and narrowed to the nearly truncated 
apex; much longer than one half its basal width; male plates large and 
broad; pronoetal watte obsolete.< 92 0.\..c68 ase oe 6. oregonensis Baker. 
Larger, 444-5 mm.; valve of the male distinctly angled at apex. 
12. fulvus Osb. 
Smaller, 3-4 mm.; valve of the male scarcely angled at apex. 
11. angustatus Osb. 


Hace, palesor-yellow,nintuscated:at, the sidesipy scm sai eee) ee ert eee 16. 
Face entirely yellow, sometimes showing obsolete irrorations quite uniformly 
AIS irDU LEC SA ah Sipe Tues 3 at Fat ean nec Oe Ue BUC at NRL POM NN ile 


Vertex more obtuse; sides of the face distinctly and abruptly darker; the 
angulate pale line at base obvious across nearly the whole width of the 
GYC@s Aierrner< SaNOshy olla eee GPs boebeuceuuconese. 14. frontalis VanD. 

Vertex more produced; face showing traces of darker irrorations especially 
at the sides; angulate basal line reduced to an angled transverse spot; 
vertex mostly white, the markings of the whole upper surface more strong- 
ENC OTMLTAS Cima Oran Ae a5, SN Sood 33 Porting ce eee ae a ar 15. nasutus VanD. 

Vertex longer, about as in nasutus; pleuree and legs mostly fuscous brown; 
inhabits Florida to Colorado and Mexico.......... 16. loricatus VanD. 

Vertex shorter, about as in frontalis; pleurzee and legs whitish or mostly so; 
iInnabibs* famaiea eel a4 oh ewes wc s esa eee 17. nanus VanD. 

Male plates triangular, their apex acute or subacute; last ventral segment 
of the female long with its apical margin oblique either side cf the middle 
which is distinctly produced and sometimes abruptly angled.......... 19: 

Male plates short, transverse and rounded at apex, little exceeding the valve; 
last ventral segment of the female short, truncate, its hind margin a little 
SINUATSD AD. PE Ries aoe hat ee A cee Fe 24. fuscifrons VanD. 


218: Annals Entomological Society of Ameriea  [Vol. III, 


19. Larger, 5 mm.; white markings cf the vertex scarcely ferming a transverse 
vitta before the eyes, consisting cf the apical line, a curved mark either 
side, and sometimes two dashes behind these, and four approximate marks 
on the base; inhabits eastern states and Canada........ 21. obscurus Osh. 

Smaller species, 34-4 mm.; markings of the vertex forming a transverse vitta 
before ‘the’ ‘696s io. 72 i nen seks es dis te ee a LT ore nate roa 20. 

20. Vertex forming at least a right angle befcre, the apex subacute; ,....5.:- 21. 
. Vertex‘short,, rounded’ at lapert on, ..\ aa ee kte Pe aban ke it ee ee 23. 
21. Vertex strongly produced, almost acute at ‘apex, its length nearly cr quite 
twice the width between the eyes; pale markings complete but not strongly 
contrasted; transverse vitta interrupted at the middle and scmetimes on 
either Side: inhabits Plorida,...$525. “Aes se es eee 19. verecundus VanD. 

Vertex shorter, scarcely mcre than right angled at-apex......... » see 

Vertex paler at base shading to fusccus antericrly; transverse v itta ef four 

subequal white dashes; apical dash short, isolated; areclar white spots few 
but. well distributed over the elytra; inhabits California and Mexico. 
18. irroratus VanD. 

-, . Perm of frontalis nearly; general color darker or blackish-fusccus; areclar 
white spots clustered abe ut the apical transverse nervures; white markings 
cn the vertex consisting cf the apical line, a marginal line either side, two 
larger spots representing the transv erse V itta, and two cthers at the base; 
inhabits: Kansas: . s).24.62 esc Se ACR Se ke 20. scriptus Ball. 

23. Anterier edge cf the head marked with five uniform contigue us W rhite spo tS; 


to 
to B 


tinct. transverse spot; inhabits Jamaica ..........,.. 23. pene VanD. 

-. Anterior margin cf the head marked with a white apical point and an cbscure 

pale patch either side about the ocelli; white mark on the base cf the frent 
represented by a shert slender arcuated line; inhabits Guatemala. 

22. osborni VanD. 


1. Platymetopius hyalinus Osborn. 
Ent. News, xi, p. 501, 1900. 

This insect is. quite aberrant‘in its genus by its uniformly 
whitish hyaline elytra crossed by three narrow fuscous bands. 
Prof. Osborn described it from a series taken from an imported 
tree at Washington, D. C., and strongly suspected that it might 
be an exotic form introduced with material added to the Botan- 
ical Gardens ‘there. In that case it probably has become accli- 
mated as Mr. C. W. Johnson has sent me a specimen he took 
near Philadelphia. 

2. Platymetopius nigriviridis Ball. 
Ent. News, xx. p. 163, 1909. 

Another very distinct little species from California. It is 
of a pale straw color tinged with green especially on the head, 
anterior margin of the pronotum and abdomen. On the elytra 
there is a longitudinal line on the clavus, some of the discal 
nervures near the apex, about four of the oblique costal veinlets 
posteriorly, and a spot at the apex of the inner apical areole 
blackish. Face immaculate. Vertex of the male short, about 
right-angled at apex; of the female strongly produced but little 
shorter than in s/ossont. The variety dixianus Ball is still 


1910] American Species of Plaiymetopius 219 


paler with the vertex of the female shorter and similarly lineated 
with fuscous at apex. This variety, of which I have seen only 
the types, is from Utah. 
3. Platymetopius majestus Ball. 
Ent. News, xx. p. 164, 1909. 

A small but very distinct species thus far reported only 
from California. Its deep black face marked with a’ white 
basal edge and concentric line, the large pale yellowish parabolic 
vertex and the brownish iridescent and uninscribed elytra make 
its identification easy .. I have seen only the types which were 
kindly loaned to me by Dr. Ball. 


a 


4. Platymetopius elegans Van Duzee. a ae 
Ent. Americana, vi, p. 94,’ 1890. 

In this species there is a broad yellowish dorsal stripe from 
the middle of the vertex to the tip of the clavus. The corium 
is brownish-cinereous obscurely banded with whitish hyaline, 
the cinereous areas are minutely inscribed with fuscous, . the 
apical submargin and some of the adjacent nervures are heavily 
embrowned and the clavus and disk of the corium are sprinkled 
with obscure rufous dots. The lower surface is yellowish with 
the base of the face cinereous and the angled pale line distinct. 
The vertex of the female is about one half longer than the width 
between the eyes. Length 5 mm. I have seen only the unique 
female type which was from California but among my Florida 
captures was one apparently scarcely mature specimen showing 
a similar yellow dorsal vitta, which I cannot identify with this 
species. It may represent a still undescribed form. 


: 5. Platymetopius acutus Say. 
jl. Acad. Nat. Sci.,, Phila., vi, p. 306,.1831: Compl. Writings, 11, p. 382. 
Platymetcpius magdalensis Prov. Pet. Faun. Ent. Can. 1i1, p. 275, 1889. 


This, our most widely distributed Platymetopius, seems to be 
the stem form from which have branched off most of the closely 
allied species found in this country, to all of which it is connected 
by certain variable characters. Its typical form, agreeing per- 
fectly with Say’s description, 1s found throughout the middle 
and nerthern states but in one form or another it occurs from 
the Atlantic to the Pacific coasts and from northern Quebec and 
British Columbia to, or nearly to the gulf coast.* The typical 
form may be characterized as follows: 


Vertex in the male a very little longer than broad between the eyes, 
in the female distinctly longer. Color: Vertex fulvous-brown finely 


220 Annals Entomological Society of America  [Vol. III, 


irrorate with testaceous forming a longitudinal line next each eve and 
another midway between that and the median sulcus; a line at apex, 
an oblique mark either side of this and four marks on the basal margin 
white bordered with piceous black. Pronotum fulvous-brown, closely, 
confluently irrorate with testaceous-white and showing five whitish 
longitudinal vitta more or less conspicuous. Scutellum fulvous with 
four spots on the anterior field and the three angles of the posterior 
field whitish, outlined in black, the disk of the anterior field sometimes 
variegated with pale. Elytra white, rather closely inscribed with fulvous 
brown; nervures darker brown; costal area white crossed by nine or ten 
heavy oblique veinlets; round areolar white spots most conspicuous in the 
apex of the anteapical areoles, the base of the apical, and near the tip 
of the claval. Face light yellow, abruptly shading to fuscous on the 
base of the front and outer half of the cheeks, the former marked with a 
white angled line and a more slender one follows close under the anterior 
edge of the vertex; the lower continued as a longitudinal line behind the 
eyes. Beneath and legs deep black, more or less varied with whitish, 
the tibize white dotted with black at the base of the white spines. Geni- 
tal characters. Male: Last ventral segment short, broadly and very 
obtusely excavated. Valve large obtusely triangular, apex rounded. 
Plates triangular, exceeding the valve by about its own length, their 
sides rectilinear or more generally slightly convexly arcuated to an obtuse 
apex, sometimes a little concave with their apex subacute; closely 
fringed with stiff bristles. Pygofers narrow, distinctly exceeding the 
plates. Female: Last ventral segment scarcely longer than one half 
its basal width, moderately compressed suggesting a median carina; 
viewed from directly below the hind margin is broadly rounded; ovi- 
duct scarcely exceeding the pygofers. Length about 5 mm. 


Toward the north and west this species varies to a lighter 
almost fulvous color with little if any black beneath, the face is 
paler, scarcely shading to darker on the sides of the cheeks, and 
the vertex is more strongly produced, at least one half longer 
than broad between the eyes. I call this variety dubius to 
distinguish it from the typical form with which it is connected 
by almost insensible gradations. It in turn forms a connect- 
ing link with oregonensis, which however I believe to be a dis- 
tinct species. I have listed this species as Jatus in Canadian 
Entomologist, xl, p. 157, 1908. 


6. Platymetopius oregonensis Baker. 
Can. Ent. xxxii, p. 49, 1900. 

This species was described from two males from Oregon but 
Dr. Howard has sent to me from the Baker collection one male 
and one female marked as types. These agree in every par- 
ticular with material in Dr. Ball’s collection and my own from 
California, Utah, Colorado and New Mexico. In the male the 


1910} American Species of Platymetopius PAPA 


vertex 1s shaped almost exactly as in the male of acutus, but it is 
shorter than in the female acutus with which sex I assume 
Prof. Baker may have compared his specimens. The female 
has a much longer vertex, its length being somewhat more than 
twice its width between the eyes. In color this species is still 
paler than in the variety dubius of the preceding species. It is 
of a light fulvous, paler and almost immaculate beneath; the 
face 1s pale with but very faint infuscation on the base of the 
front and sides of the cheeks but with the angular white lines 
on the former distinct. 

This species 1s best distinguished by the form of the genital 
pieces: The valve of the male is more angled and subacute at 
apex; the plates are larger with their outer margins distinctly 
concavely arcuated and their apices a little divergent and sub- 
acute. The last ventral segment or the female is conspicuously 
longer and narrower than in acutus with its apex slightly 
subangularly produced. 

7. Platymetopius latus Baker. 
Cane Pnte xcxxitee py oO snl G00) 

I have one female of this species taken by Mr. W. J. Palmer 
of this city at Quinze Lake, Quebec, which agrees perfectly with 
Prof. Baker’s Colorado type received from Dr. Howard. It 
may be distinguished by its fusiform shape, narrow pointed head 
with a vertex twice as long as its width between the eyes, the 
broad flaring elytra, the almost uniform coloring which is ful- 
vous-brown above, without irrorations on the vertex, pronotum 
and scutellum, and almost without elytral reticulations or white 
areolar spots, the nearly concolorous nervures, those of the 
costal areole being a little more distinct, and the uniformly pale 
vellowish-testaceous coloring beneath with the base of the front 
darkened only at the extreme tip and not showing the angular 
white mark. Genital characters nearest to those of acutus; 
the last ventral segment a little more angled and the pygofers 
smaller. This is a rare species and probably is confined to 
northern Canada and the higher regions of the Rocky Moun- 
tains. 

S. Platymetopius cuprescens Osborn. 
20th Rept. N. Y. State Ent. p. 517, 1905. 

Another closely allied species which comes nearest to acutus 
but has a longer and sharper vertex, about as in dubdbius and 
oregonensis; the elytra are more infuscated with but slight traces 


222 Annals Entomological Society of America  [Vol. III, 


of the white areolar spots. The face is pale yellow, but little 
darkened on the base of the front and sides of the cheeks, with 
the pale angular line distinct. The genital characters of the fe- 
male are very close to those of acutus, but the last ventral seg- 
ment is more carinated with its apex distinctly notched when 
viewed from below. In the male these characters are very dis- 
tinctive. Last ventral segment longer, not shorter, than the 
preceding; valve shorter; plates much shorter, rounded, about 
two thirds the length of the valve; pygofers broad, exceeding the 
plates by about the length of the latter and similar to them in 
color and texture. Prof. Osborn’s type was taken by me at 
Phoenicia, N. Y., I have taken another female at Niagara Falls 
and a male at Boulder, Colorado, and another at Ogden, Utah, 
and Mr. Palmer took one at Quinze Lake, Quebec. 


9. Platymetopius slossoni n. sp. 


Very close to acutus but smaller with a longer and more 
acute vertex and shorter male pygofers. Length 4 mm. 


Vertex longand acute at apex, stilllonger thanin oregonensis and latus 
rather more than twice as long as wide between the eyes and obviously 
longer than the pronotum and scutellum together; marked with three 
pale longitudinal lines as in aculus; the lateral curved, becoming approx- 
imate and parallel for a short space at base. Face light yellow with the 
angled base of the front and sides of cheeks broadly distinctly infus- 
cated, the former with a very distinct acutely angled concentric white 
line which is projected to the apex of the head and is strongly repro- 
duced behind the eyes. Pronotum with the longitudinal vittze distinct 
as in acutus. Beneath and abdomen dark fuscous; in the female ob- 
scurely marked or irrorate with pale in places; last ventral segment 
shaped as in aculus, whitish at base with about four black ligulate marks 
on the basal margin. In the male the pectus and abdomen are deep 
black, obscurely marked with pale on the venter and at the tip of the 
valve; last ventral segment angularly but not deeply emarginate; valve 
triangular, subacute at apex; plates rather broad-triangular, on their 
sutural margin as long as the valve, their outer edges moderately sin- 
uated and fringed with bristles, their apex a little narrower than in 
acutus; pygofers slightly but distinctly shorter than the plates which 
they scarcely exceed at the sides. 

Described from one pair taken at Jacksonville, Fla., in 
May, 1910, by Mrs. Annie Trumbull Slosson whose enthusiasm 
and industry have added so much to our knowledge of the insect 
fauna of Florida and other parts of the eastern United States. 
I took a good series of what is certainly the same species at 
Crescent City, Sanford, Sevenoaks and Ft. Myers, Florida. 


1910] American Species of Platymetopius 223 


They differ only in being slightly larger and paler. This species 
is very close to acutus with which the typical form is almost 
identical in color and markings but its small size, remarkably 
long and acute vertex and the short male pygofers preclude its 
reference to that species. 


10. Platymetopius cinereus Osborn & Ball. 
Proc. Towa “Acadi* Sei im ps) Lost elso7- 

This is a very small and somewhat greenish species with a 
vertex formed about as in acutus; face greenish-white, very 
slightly infuscated on the base of the front where the angular 
white line is distinct; markings on the vertex and pronotum 
nearly obsolete but the five pale vittz of the latter are generally 
discernable; abdomen mostly black in the male, rarely so in the 
female; elytra closely but lightly reticulated. Last ventral 
segment angularly produced in the female; valve of the male 
obtusely triangular; plates nearly as long as the valve on their 
suture, triangular, blunt at apex, considerably exceeded by the 
pygofers. 

This, one of our smallest species, is found throughout the 
warmer portions of the central and southern states from Iowa 
to Florida. 


11. Platymetopius angustatus Osborn. 
20th Rept. N. Y. State Ent., p. 518, 1905. 


Prot. Osborn has very kindly sent me one of his male types 
and I have examined the female in ‘his-collection. This is the 
smallest species known to me and has been reported from the 
type locality only: Long-Island, N. Y.' The female is distinctly 
larger with a longer vertex than the male. It has a light green- 
ish color with coppery reflections; the round areolar spots of the 
elytra are confined to the apical and anteapical areoles; the costa 
is hyaline with the oblique veinlets dark and the abdomen is 
marked with black. The male is smaller with a much shorter 
vertex; its color is a bright fulvous becoming yellowish on the 
head; the elytra are subopake, almost unicolorous on their base 
but becoming recticulated with brown on their apical and ante- 
apical areoles where the round white spots are fairly distinct; 
oblique veinlets of the costal areole almost obsolete. In both 
sexes the base of the front is feebly infuscated with the pale 
angular line discernible. 


224 Annals Entomological Society of America [Vol. III, 


Last ventral segment of the female rounded. Valve of the 
male large scarcely angulate at apex; plates triangular, on their 
suture shorter than the valve; terminated by the long sete; 
the brown apex of the pygofers much surpassing the plates. 


\2. Platymetopius fulvus Osborn. 
20th Rept. N. Y. State Ent., p. 519, 1905. 


This insect has about the form and size of frontalis but it is 
of a bright fulvous color with the elytra mostly opake and the 
markings almost or quite obliterated from the base nearly to 
the apex of the ante-apical areoles; vertex and scutellum more 
yellowish with indistinct pale vermiculate lines; oblique costal 
veinlets brown on a concolorous ground; face almost uniformly 
pale yellow with the basal angular pale line rarely indicated. 

Last ventral segment of the female rounded behind with a 
blackish arcuate mark either side which gives it the appearance 
of being strongly produced on the middle. Valve of the male 
ovate-triangular, its margins arcuated to the obtuse apex; 
plates large, triangular, exceeding the valve by its own length; 
strongly ciliate on their outer margins; pygofers slender, longer 
than the plates. 

Prof. Osborn’s types were from Cold Spring Harbor, Long 
Island, N. Y. About Buffalo it is not uncommon among the 
hills at Colden and Gowanda and it has been reported from 
Pennsylvania and Ohio. 


13. Platymetopius dorsalis Ball. 
Platymetepius frontalis var. dorsalis Ball. Ent. News, xx, p. 164, 1909. 

I cannot agree with Dr. Ball in placing this as a variety of 
frontalis. It seems to me very distinct and withall a difficult 
species to place. It is fulvous and pale yellow in color and has 
much the aspect of a small Mesamza vitellina but it has the ver- 
tex of a Thamnotettix and the front of a Platymetopius, to which 
genus I think it is most closely allied by its venation and struc- 
tural characters. The anterior edge of the head is obtusely 
rounded as compared with the allied species. The lower sur- 
face, vertex and scutellum are pale yellow and immaculate. 
The pronotum and elytra are bright fulvous with coppery 
reflections, with the sides of the pronotum, costa at base, apical 
areoles and a vague saddle across the apical half of the clavus 
yellowish. I have seen only the unique type from Kansas 
kindly loaned to me by Dr. Ball. 


L910] American Species of Platymetopius 225 


14. Platymetopius frontalis Van Duzee. 
Can. Ent. xxii, p. 112, 1890. 


Common and widely distributed throughout a great part of 
the United States and Canada. This species may be distin- 
guished by its stout form, short vertex and blackish color both 
above and below, with a clear yellow face which becomes infus- 
cated at each side but scarcely at the pointed base where the 
angled lines are but poorly distinguished. The vertex has a 
white median dash at apex and a transverse vitta of similar 
marks across the disk before the eyes, more or less conspicuous. 
The pronotum sometimes shows faint traces of the five pale 
vittz and there may be some white marks on the scutellum 
forming two broken longitudinal vitta. Elytral areoles with pale 
reticulations about their margins, sometimes nearly obsolete 
in the males; round areolar spots well developed and conspic- 
uous; costa white, the oblique veinlets heavy; apical areoles 
generally infuscated beyond the white basal dots, the extreme 
edge clear white. Male genital characters about as in fulvus; 
the valve broader and more rounded and the plates a little 
shorter; last ventral segment of the female short and broadly 
rounded on the apical margin. Dr. Ball has sent me for 
inspection a typical example of frontalis taken by Prof. Gillette 
in Mexico. 


15. Platymetopius nasutus Van Duzee. 
Bul. Buf. Sec. Nat. Sci. viii, No. 5, p. 64, 1907. 


While very close to frontalis I believe this a sufficiently dis- 
tinct species. The vertex is obviously longer and more pointed 
and there 1s much more white in the coloring of the upper sur- 
face. The vertex is mostly white with the apex and a trans- 
verse eroded vitta black, the former with a median line and two 
oblique dashes white; the pale basal portion is veined with 
fuscous and the white median area is crossed by a black longi- 
tudinal line either side of the middle; the pronotum is irrorated 
with white and fuscous with the anterior margin and depressed 
sides mostly white and the longitudinal vittz scarcely indi- 
cated. The elytra are white with a few fulvous-brown areas in 
the areoles and with fuscous reticulations more or less extended, 
the white areolar spots being much obscured. Beneath black 
with the tibia and tarsi white, dotted with black; the femora 
sometines largely invaded with white. 


226 Anna.s Entomological Society of America  |Vol. III, 


In this species the face while mostly yellow shows a strong 
tendency to a uniform fuscous irroration which becomes more 
conspicuous laterally (thus forming a connecting link with the 
group of brown-faced species), and the angled base may also be 
darkened, with a transverse crescentic white mark. The male 
plates are proportionately a little larger and more obtuse at apex 
than in frontalis and the last ventral segment of the female is 
shorter and more angled. The types of this species were from 
Jamaica and Colorado and I have seen others from California. 


16. Platymetopius loricatus Van Duzee. 
Bul. Buf: Soc. Nat. Sci., v., p. 205, 1894. 


Allied to nasutus but sometimes smaller and showing strong 
indications of the frontal irrorations found in fuscifrons. Face 
in typical examples entirely pale yellow. Vertex shorter than in 
nasutus but longer than in frontalis; its markings substantially 
as in frontalis but here the white vitta consists in great part of 
two longer approximate and dislocated white dashes and a few 
minute irrorations, the base with two median white dashes. 
Pronotum with the five pale vitte fairly distinct. Elytra 
mostly blackish as in frontalis but with the white areolar spots 
smaller and fewer anteriorly. Face frequently showing strong 
indications of fuscous irrorations, especially toward the base 
of the front where there is then a short angled white mark. 
Last ventral segment of the female short, its hind margin 
hardly angled at the middle, the lateral angles rounded. Valve 
of the male short and rounded or very feebly angled; the plates 
broadly but acutely triangular, scarcely longer on their suture 
than the valve; slightly exceeded by the pygofers. Described 
from California but I have seen others from Colorado, Florida, 
Orizaba, Mexico, and Gualan, Guatemala. 


17. Platymetopius nanus Van Duzee. 
Bul. Buf. Soc. Nat. Sci., viii, No. 5, p. 65, 1907. 


Very close to Joricatus but the colors are paler with the white 
elytral spots less distinct, the pleura are yellow with a brown 
cloud anteriorly, not black as in /oricatus, and the vertex is a 
little shorter but similarly marked. The last ventral segment of 
the female is more produced. In the male the last ventral seg- 
ment is subangularly excavated behind; the plates also are larger 
and white, surpassing the apex of the pygofers which is not the 
case in loricatus. 


1910] American Species of Platymetopius DT 


The aspect of these two species and the next shows a greater 
difference than is sustained by their structural characters and 
it is possible, but I hardly think likely, that material from inter- 
mediate localities may necessitate merging them in one. The 
smaller manus has thus far been found only in the island of 
Jamaica. 


18. Platymetopius irroratus n. sp. 


Differs from Joricatus in being paler in color and generally 
smaller and in having the hind edge of the last ventral segment of 
the female distinctly angled either side and the plates of the male 
proportionately shorter but equally acute at apex leaving the 
margins more deeply arcuated. In this form the face is always 
more or less irrorated with brown or fuscous or at times entirely 
pale brown irrorated with pale. In the paler individuals the 
facial irrorations are always more dense toward its angled base 
where the pale line 1s distinct. The vertex is somwehat shorter 
than in loricatus and the elytra are cinereous instead of almost 
black as in that species. It is close to brevis but may be dis- 
tinguished by the longer vertex which is darkened at apex and 
marked with four white dashes as in J/oricatus. The genital 
characters are also quite different from brevis. Length 4 mm. 

Dr. Ball has sent me a long series of this species taken at 
Chino, Riverside, Pasadena and San Diego, California, Phoenix, 
Arizona, and Ti Juana, Mexico. 


19. Platymetopius verecundus n. sp. 


Allied to trroratus and obscurus but with a longer vertex 
than in either. Length 3 1-2 to 4 mm. 


Vertex about as in oregonensis, much produced and acute but dis- 
tinctly shorter than in s/ossoni; in the female about twice as long as wide 
between the eyes. Clypeus much constricted near the base with its 
apex produced and rounded as in the allied species. Hind margin of 
the last ventral segment of the female distinctly angled and slightly 
produced at the middle and more obscurely angled on either side. 
Last ventral segment of the male subangularly emarginate; valve large, 
scarcely angled at apex; plates as in Joricatus, a little longer than in 
irroratus and smaller than in scriptus and obscurus. 

Color fulvous-brown with rather strong ornamentation. Vertex 
marked as in the allied species, the pale apical line more developed; 
transverse pale vitta sometimes almost entire or broken only by the 
fulvous median stripe and a brown line near it on either side, or there 
may be a second interrupting line exteriorly; base with a white median 
vitta bisected by a fuscous longitudinal line and sometimes terminated 
outwardly by a similar line. Pale vitta of the pronotum distinct. 


228 Annals Entomological Society of America  |Vol. III, 


Scutellum distinctly fulvous with the extreme angles and two vitt« 
irrorate with white. Elytra strongly inscribed; the areolar white spots 
distinct; costal areole whitish with the oblique veinlets strong; apical 
areoles narrowly fuscous next to the slender white apical margin. 
Front fulvous-brown, obscurely irrorate with pale, the white angular 
mark on the base of the front well developed and reaching about one 
half way to the eyes. There are two white lines beyond the eyes and an 
obsolete oblique one at the upper: angle of the lorze; pectoral pieces 
fulvous-brown, conspicuously paler beneath. Abdomen fuscous or 
brown, varied with pale. Legs pale dotted with fuscous. 

Described from numerous examples taken by me in Florida 
and listed as ‘‘obscurus Osb.?”’ in my report on the Hemiptera 
taken at that time. The exact localities represented are Cres- 
cent City, Sanford, Sevenoaks, Clearwater and Estero. This 
species with manus from Jamaica, trroraltus from Mexico and 
scriptus from Kansas forma puzzling group of very closely allied 
forms which run into Joricatus on one hand and obscurus on the 
other. Their genital characters vary slightly but are not suffi- 
cient for separating them. Students with ‘‘lumping’’ pro- 
clivities will doubtless place them as varieties of a single species. 

20. Platymetopius scriptus Ball. 
Ent. News, xx, p. 165, 1909. 

Male: A little larger and darker with a shorter vertex than in 
the preceding two species. Closely allied to frontalis but with a 
brown front and differently marked vertex. Last ventral seg- 
ment nearly as long as the preceding, broadly angularly exca- 
vated behind and distinctly carinate on the median line; valve 
large, produced but scarcely angled at apex; plates large, sub- 
acute at apex, sides nearly straight; on their suture much shorter 
than the valve; considerably surpassed by the pygofers. Color 
blackish fuscous, nearly as dark as in frontalis, paler on the scu- 
tellum; white areolar spots present in the apex of the ante- 
apical and base of the apical areoles, elsewhere usually wanting. 
Face brown, coarsely irrorate with pale, the angular basal mark 
distinct. Pectus, abdomen and legs black; tibia mostly white 
dotted with black; median apical line and anterior edge of the 
vertex white; the latter forked about the ocelli and connected 
with a squarish white spot indicating the transverse vitta of 
loricatus; basal white dashes usually connecting with another 
exteriorly forming an irregular pale patch. Scutellar angles 
more or less conspicuously fulvous. Length 4 mm. 

Known to me only from the five type specimens kindly 
loaned to me by Dr. Ball. 


L910} American Species of Platymetopius 229 


21. Platymetopius obscurus Osborn. 
Ohio Nat. v. p. 274, 1905. 


This species very closely resembles acutus in size, color, 
markings and shape of the vertex. The face however is ful- 
vous-brown, coarsely irrorate with paler; the valve of the male 
is more angled, the plates are smaller and the hind edge of the 
last ventral segment of the female is more transverse and angled 
on either side. Length 5 mm. 

Described from New York, Pennsylvania and Ohio. Mr. 
Palmer ‘found it abundant at Lake Temagami, Ont., and 
Quinze Lake, Quebec. I took it at Sharon and Ipswich, Mass., 
and have examined others from Georgia. 


22. Platymetopius osborni n. sp. 


Near brevis but larger with a shorter vertex. Length 4 to 4 1-2 mm. 
Vertex scarcely longer than its basal width; anterior margin rounded, 
the angle at apex hardly indicated. Front broader than in brevis, 
its sides parallel or nearly so to the superior tip of the lore, then 
abruptly narrowed to the clypeus. Genital characters substantially 
the same as in the allied species except that the last ventral segment of 
the male is shorter and the pygofers are longer, surpassing the plates. 
Last ventral segment of the female distinctly angled at about the middle 
of the latero-posterior margins where they are practically rectilinear in 
brevis. Color about as in brevis becoming paler in some females. 
Vertex fulvous-brown varied with darker, base with four pale marks; 
a line adjoining the eye and the broad apex ivory-white, the latter inter- 
rupted by a median vitta forked at apex and a small broken spot half 
way between this and the black ocelli. Face irrorate with fuscous points 
which become confluent on the front and base of the cheeks and omit 
the sides of the clypeus, a vitta on the lores and another on the cheeks 
above its suture; pale angled line on the base of the front reduced to a 
small transverse spot less conspicuous than in brevis. Pronotum rather 
dark with the five pale vitte distinct. Scutellum tinged with fulvous 
with a deeper fulvous mark within the lateral angles, the margin with 
five white spots. Elytra with strong nervures and recticulations, the 
latter forming a longitudinal median vitta in the anteapical areoles; 
apex infuscated with the extreme edge white as far as the outer areole; 
costa crossed by about ten oblique veinlets; the white areolar spots 
distinct. Beneath and legs varied with pale and fuscous; the disk of 
the pleural pieces and base of the venter infuscated. The female is 
paler but with all the markings indicated. 


Described from one female and two males from Prof. 
Herbert Osborn and taken at Los Amates, Guatemala, January 
17th and February 28th, 1905, by Prof. J.S. Hine. This can 
be confused with no species but brevis from which it may be 


230 Annals Entomological Society of America — |Vol. III, 


distinguished by its larger size, broader form, shorter and more 
rounded vertex, the brown vitta in the anteapical areoles, the 
longer male pygofers, and the angle on the margin of the last 
ventral segment of the female. It has somewhat the aspect of 
scriptus but is very distinct in having a much shorter vertex, 
in the form of the last ventral segment of the male and in the 
more angled valve. 
23. Platymetopius brevis Van Duzee. 

: Bul. Buf. Soc. Nat. Sci. viii, No. 5, p. 66, 1907. 

This tiny species is very close to osborni but it averages a 
little smaller, the vertex is a little more angled, the transverse 
vitta is crowded close to the anterior margin of the vertex and 
consists of a pale cloud against each ocellus which may be ob- 
scurely connected to a pale basal mark either side of the middle. 
The anterior edge is white and calloused and is nearly or quite 
twice interrupted on either side forming five conspicuous white 
marginal spots. The general colors in this species are pale, 
largely obliterating the white areolar spots on the elytra. It 
has been found only on the island of Jamaica, unless a series 
from California in the Ball collection belong here. 


24. Platymetopius fuscifrons Van Duzee. 
Bul. Buf. Soc. Nat. Sci. v., p. 206, 1894. 
Platymetopius abruptus Ball, Ent. News, xx., p. 165, 1909. 

This is very distinct from any other known dark-faced 
species by the short transverse male plates and the short trun- 
cated last ventral segment of the female. In the male the 
front is almost black; the vertex is a little more than right- 
angled at apex and is obscurely marked, with the white apical 
line distinct. The male is deeply colored as in frontalis while the 
female seems to be paler than the female of that species, but 
in both sexes the white areolar spots are well developed. I have 
been unable to detect any characters by which to distinguish 
abruptus, the type of which Dr. Ball has kindly sent me for 
examination, from fuscifrons. This species inhabits Arizona 
and California. ; 

25. Platymetopius modestus Stal. 
Of. Vet. Akad. Forh., xi, p. 255, 1864. 
This species has not been recognized by later students and 


from the short description I am unable to place it in my synop- 
tical table. I append a copy of Stal’s description: 


1910] American Species of Platymetoptus Pea | 


“Vertice thoraceque flavis, illo apicem versus concavo, longitudi- 
naliter anguste nigrolineato, hoc basi virescente; hemelytris dilute 
virescentibus, nervis flavescentibus, margine costali albido-subhyalino ; 
subtus dilute flavotestaceus. Long. 8, lat. 2 millim.—American septen- 
trionalis.”’ 

26. Platymetopius tenuifrons Baker. 
Can. Ent. xxx, p: 50; 1900: 


The type of this species was received too late for its inclusion 
in the synoptical table. It belongs to the group of brown- 
faced species and really comes nearest to verecundus from which 
it differs at once by its much larger size and the still longer 
vertex. It has somewhat the aspect of the female oregonensis 
but the vertex is longer, broader, and more rounded at tip. 
The elytral reticulations and areolar spots are finer but dis- 
tinct; the general color is more cinereous than fulvous; the front 
is fuscous-brown irrorated with paler and becomes darker or 
almost piceous at the pointed base below the short but distinct 
and acutely angled white line, and the lower surface of the body 
is varied with pale and brown. The last ventral segment of the 
male is about as long as the third and is broadly subangularly 
excavated; valve rather short and broadly rounded; plates 
broad-triangular and shorter than the valve. The pale pro- 
notal vitte are obvious but not conspicuous and the vertex 
is blackened on the immediate apex with a white point at tip 
and a pale marginal mark on either side and another half way 
to the eye. The unique male type was taken in November at 
Chapada, Brazil. 


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ANNALS. 


OF 


The Entomological Society of America 


Volume III DECEMBER: 19:10 Number 4 


DIPLOPODA FROM THE WESTERN STATES. 


By Ratpu V,. CHAMBERLIN. 
Brigham Young Univ., Provo, Utah. 


FAMILY CALLIPODIDAE. 
Lysiopetalum mutans sp. nov. 


General color of body above dark brown to almost black; a narrow 
median line of yellow extending from the anterior margin of the second 
dorsal scutum to the end of the anal scutum; a broader pale stripe of 
darker caste than that of the median line along each side from the 
anterior margin of the first segment to the end of the body; lower por- 
tion of sides and the entire venter yellowish. Legs yellow on inner 
or ventral surface proximally, the distal articles commonly darker; 
dorsally brownish, likewise darker distally. General color of head 
brown; vertex areolate with dark-brown over a lighter ground; a 
smoky or blackish brown area between eye and antenna of each side 
and those of the other, the area narrowing to a median point down the 
front, the region each side of this elongation areolate like the vertex; 
clypeus and labrum light brown to yellow. Antennae deep brown 
or blackish excepting the two distal articles which are whitish. 

Vertex of head crossed by a narrow median longitudinal furrow 
which is subdivided by a fine line-like elevation; head depressed trans- 
versely a little ventrad of bases: of antennae; subdensely hirsute with 
short stiff hairs or setae which are somewhat longer in the clypeal 
region. 

Gnathochilarium broad; greatest width of mentum 3) times as 
great as the median length; lingual lamellae, exclusive of its processes, 
twice as long as the greatest width. 

Eyes well developed, black, subtriangular; the ocelli numerous, 
compactly and seriately arranged. Below eye area proper on each 
side is what appears to be an organ of sense; it is much larger than an 
ocellus, convexly elevated and transparent, the convex portion appar- 
ently covering a pit in which is a conical body like a sense-cone of the 
antennae. 

Antennae long and moderately clavate; clothed with mostly short 
hair intermixed with some longer ones; length 2 mm. 


233 


234 Annals Entomological Society of America  [Vol. III, 


Body gradually attenuated from near anterior end of caudal third 
of the length cephalad, more abruptly narrowed or constricted over 
several segments immediately caudad of the head. Dorsum depressed 
or sub-complanate, rather more convex cephalad than caudad. Sides 
sub-vertical. A fine median dorsal keel extending from the anterior 
margin of the second segment to the caudal end of body. Each side 
of the median keel upon the dorsum are three conspicuously elevated, 
sharp edged carinae which extend entirely across the segment and each 
of which bears at its caudal end a short clavate seta projecting dorso- 
caudad; laterad of the outermost of these dorsal keels is a much thicker 
keel with wide flat surface which bears a similar seta adjacent to caudal 
end; between each two of these major carinae is an intermediate lower 
carina which bears no seta; on the side ventrad of the thickened carina 
above mentioned are two additional, large setigerous carinae and be- 
tween the thickened carina and the first and between the first and 
second of these a lower non-setigerous carina; while ventrad of the sec- 
ond are other lower carinae which become obscure ventrad. Segments 
anteriorly transversely furrowed, the carinae over and cephalad of the 
furrow lower and weaker but continuous. On the second, third and 
fourth segments the setae are not borne at caudal ends of the carinae 
but spring from tubercles contiguous with the carinae at or cephalad 
of their middles and extend vertically from the surface or nearly so; 
on the fifth segment are two series of setae, one consisting of setae along 
caudal margin as on more posterior segments, and the second more 
cephalad as on the preceding segments, the setae of the two series 
alternating. 

Collum or first segment in outline half-moon shaped, the caudal 
margin being straight and the lateral and anterior margins together 
evenly convexly rounded or semicircular. Carinae extending only 
across the posterior half of the segment, the setae being borne at their 
cephalic ends. A second series of setae toward the cephalic margin. 

Anal plate triangularly narrowed caudad, the posterior margin 
truncate or weakly widely rounded and bearing the usual two long 
setigerous papillae; anteriorly, longitudinally carinate; bearing trans- 
verse rows of setae of which the most caudal become longer and con- 
spicuously and finely drawn out apically. 

Anal valves smooth; an obliquely transverse, somewhat curved 
impression over caudal portion on each side; each valve bearing two 
pointed setae, one at the caudal edge and one at middle toward the 
mesal margin. 

Anal scale transversely somewhat diamond shaped, the meso- 
caudal and meso-cephalic angles a little rounded; a longitudinal, some- 
what incurved, impression crossing each side; bearing two pairs of 
finely pointed setae, one median and between the longitudinal impres- 
sions and the other near the caudal margin with each seta laterad of 
the impression of the corresponding side. 

Distal article of first and second legs in the female with a comb- 
or calamistrum-like series of setae along the ventral surface, the setae 
being very closely and regularly set. 


1910] Diplopoda from the Western States 235 


In the female the coxae of the second to eighteenth legs inclusive 
and of the twentieth or, as appeared in one specimen, of the second 
to nineteenth legs inclusive, bear on their caudo-ventral faces con- 
spicuous, distally inflated, sub-fungiform outgrowths; the processes 
of the third legs, however, reduced to smaller, low convex elevations. 

Oviducts of female protruding as greatly elongated appendages or 
ovipositors which when extended caudad reach the eighth segment 
of the body; distal portion of processes enormously enlarged, bearing 
very long stiff bristles; proximally glabrous; showing a tendency to 
segment into articles; color white. 

Number of segments 49 to 53. 

Length 26 mm. Width 1.5 mm. 


Locality—Stanford, Cal. (W. M. Mann). 


FAMILY CRASPEDOSOMIDAE. 


Conotyla deseretae sp. nov. 


Dorsum brown; a dark band of black or bluish black color along 
each side immediately ventrad of the carinae, the band commonly 
extending dorsad on the prozonites and sometimes, especially on the 
posterior segments, forming thus a band entirely across the dorsum; 
a median longitudinal dark band along dorsum, and this typically 
geminated throughout its entire length by a narrow light line. Venter 
and legs paler, light brown to yellowish. Head commonly a consider- 
ably darker, more reddish, brown than that of the body. Antennae 
deep brown to blackish. 

Head widely and shallowly depressed or furrowed transversely 
between the eye patches. Evenly hirsute with short hairs. 

Lingual lamellae of gnathochilarium with sides sub-parallel for 
most of length; their length, inclusive of processes, three times the 
greatest width. Lingual stili conspicuous, their lateral teeth distinct. 
Spatula large, distally broadly rounded. Stipites a little less than five 
times as long as the greatest width. Mentum with greatest diameter 
somewhat more than twice the greatest length; its anterior margin 
widely rounded; as a whole sub-semicircular in shape. 

Eye patches triangular, the apex directed forward; ocelli 25, more 
or less, arranged mostly in seven series (6, 5, 5, 4, - 2), 

Antennae long, of typical form and proportions: length 3.0 mm, 
wee Pl. XXXI, fig. 8. 

Body strongly narrowed caudad and cephalad, the first segment 
constricted in the usual way; a sharp, low, median keel extending 
along dorsum for its entire length, a narrow furrow impressed each side 
of the keel. Second segment short, the inner bristles relatively farther 
laterad than on the first, and these bristles located more and more dis- 
tantly from the median line on subsequent segments back to the mid- 
dle region of body. The lateral bristles, as usual, becoming directed 
more and more strongly caudad in proceeding from anterior segments 
posteriorly. Inter-segments or prozonal divisions in part encircled 
by a series of fine transverse impressed lines, the elevations between 
these crossed with fine striations. 


236 Annals Entomological Society of America  [Vol. III, 


First segment somewhat semi-circular or sub-reniform, the anterior 
median region extending into broad excavation of head. Carinae 
very weak, continuous with the finely raised line of cephalic margin. 
The inner bristle on each side more than one-third the distance from the 
distinct longitudinal median keel to the outer bristles. The two outer 
bristles of each side upon carina, extending dorso-laterad and the 
caudal one in a transverse line with the inner ones. 

Anal scutum truncate caudad, but the caudo-lateral angles extended, 
the intervening margin widely set in. 

Anal valves equalling the scutum; the caudal margin of each nearly 
straight or but little curved, distinctly raised; an impressed longitudinal 
line toward dorsal margin and a similar one toward the mesal; three 
bristles borne a little cephalad of the caudal margin, the uppermost 
of these being close to the anal scutum, and the lowermost near the 
middle line of the valve. 

Anal scale trapezoidal, the caudal margin wide, widely weakly 
incurved; a long bristle borne near each caudo-lateral angle. 

First two pairs of legs of male slender, six-jointed as usual; the 
claws large; without special protuberances, and with no roughenings 
on the distal articles. 

Third legs stout and, like the subsequent ones, seven-jointed; the 
ultimate joint roughened over the ventral and ventro-lateral surfaces, 
except proximally, with rows of short transparent processes; femur 
swollen into a wide enlargement on the dorsal side. 

Fourth and fifth legs of male similar to the third but with the dor- 
sal face of femora not bulging. 

Sixth legs of male like the fifth but the coxae bulging ventrad. 

Seventh legs of male with coxae strongly bulging ventrad into 
prominent rounded processes as shown in the figure (Pl. XXXII, fig. 6). 

Anterior pair of processes of male gonopods broad plate-like struc- 
tures with teeth along mesal portion of distal edge; posterior pair 
large, strongly inclined caudad, a secondary, somewhat flattened, 
accessory division along proximal portion of caudo-lateral surface; 
a conical process springing from the base of each (see Pl. XXXII, figs. 
4, 5, and 6.) 

Eighth legs or accessory gonopods with the distal article strongly 
enlarged as usual, the basal article more than commonly elongate, and 
extended ventro-mesad into a conical process which bears distad a 
number of very long bristles; below the conical process the proximal joint 
is shallowly excavated for the reception of a rounded protuberance 
of the posterior gonopod of the corresponding side. 

Length 23 mm.; width 2.9 mm. 


Locality—City Creek, Mill Creek, Provo and other canyons 
of the Wahsatch Mountains, Utah. 


1910} Diplopoda from the Western States Zod 


Conotyla coloradensis sp. nov. 


Dorsum and sides brown, the prozonal divisions of segments some- 
what lighter than rest; sometimes finely areolated with paler brown. 
Lower region of sides and the venter paler, light brown to yellowish. 
No black bands or spots. Legs yellowish proximally, the distal joints 
brown or marked with brown in spots and streaks. Antennae brown. 

Head hirsute, the vertex more sparsely so than the frontal region. 

Lingual lamellae of gnathochilarium widest anteriorly, incurved 
posteriorly, posterior arms a little divergent, three times as long as 
greatest width. Stipites five and a half or more times as long as the 
greatest width. Spatula conspicuous, distally subtriangular. Mentum 
one-half as long as greatest width. 

Eye areas subtriangular, but with the sides convex; ocelli about 22, 
in five curved series, counting from outermost mesad as follows: 5,5,5,4,3. 

Antennae very long, proportions as represented in Pl. XX XIII, fig. 
2. Length 4.2 mm. 

A fine median longitudinal keel over all segments excepting the last. 
Carinae increasing in size from the first caudad. Inner setae becoming 
relatively farther and farther from middle line in going caudad, these 
setae inserted in a wide longitudinal depression extending along each 
side, the dorsum between the depression a little convexly elevated. 
Setae, as usual, becoming directed more and more strongly caudad in 
proceeding from anterior to posterior segments. 

First segment of the usual form, the anterior margin broadly extended 
cephalad mesally and fitting into excavation of head, the lateral por- 
tions of margin a little incurved. Carinae small, continuous with 
anterior margin of plate. Innermost bristle about one-third the dis- 
tance from median to the edge of the carina of the corresponding side, 
in transverse line with the more anterior of the lateral bristles which is 
situated cephalad and considerably mesad from the posterior one, the 
latter close to the caudal margin. Segment considerably depressed or 
furrowed transversely, the inner setae located in this depression. Setae 
erect. 

Anal scutum truncate caudad, the posterior margin weakly incurved. 

Anal valves not equalling the scutum, widely rounded and broadly 
margined. The three bristles nearly equidistant, the most ventral 
three or more times as far from the ventral scale as the most dorsal 
is from the scutum. 

Anal scale sub-trapezoidal, the lateral and cephalic margins weakly 
convexly rounded, the caudal distinctly incurved between the two 
long bristles. 

Appendages of second segment of female conspicuous, strongly 
enlarged distad, the distal end with long bristles. A sub-lobe sheath- 
ing the cephalic surface of main appendage, this bearing long bristles 
apically. See Pl. XX XMlT, figs 7: 


Locality—Colorado (T. D. A. Cockerell). 


238 Annals Entomological Society of America [Voll III, 


Tingupa gen. nov. 


Antennae moderate, strongly clavate distally, the fifth article 
longest and much thickest, the third article second in length. 

Eyes well developed, consisting of numerous ocelli (8-20) arranged 
subseriately. 

Gnathochilarium with promentum well developed, triangular and 
moderately low. 

Segments convexly arched dorsally and with well-developed carinae; 
dorsum covered with densely arranged, laterally compressed and in 
part conically pointed granules which extend to edges of carinae and 
on caudal segments may project as spinous processes from carinae and 
caudal margin of segments; setae moderate to long, clavate. 

Segments in adult 30. 


Type—Tingupa utahensis sp. nov. 

Distribution—Two species known, the type from Utah anda 
second form, 7ingupa monterea, from California. 

The genus seems to have closest affinities with Pseudotremia. 


Tingupa utahensis sp. nov. 


General color brown, like that of the dead leaves among which it 
lives; in some with finer mottlings of paler color on sides. Venter pale. 
Legs from yellow or almost white to pale brown. Antennae brown, 
the articles often paler at distal ends. 

Segments with major subdivision strongly granulate, the granules 
or scales mostly somewhat compressed laterally or elongate antero- 
posteriorly and closely arranged; similar but narrower elevations closely 
covering the dorsal portion of the minor subsegments; scales larger on 
posterior segments, series of larger ones with conical apices directed 
caudad as acute spinous projections. Segments mesally well arched, 
depressed immediately mesad of the carinae; entire dorsum weakly 
longitudinally furrowed, the furrow divided by a fine, at times almost 
obsolete keel. Segments bent cephalad laterally, the latero-cephalic 
margin widely rounded, the acute anterior angles situated close to body. 
Edges of carinae often appearing finely crenate or dentate from the 
presence of the projecting tubercles. All setae clavate, in proceeding 
from first to posterior segments the two lateral setae on each side 
become more extended in these directions, more strongly directed 
caudad and cephalad respectively, in some being horizontally extended 
in these directions. 

Vertex of head crossed by a low, rounded, median longitudinal 
ridge; head also elevated along a line connecting the angles of the eyes; 
transversely depressed between bases of antennae; conspicuously and 
rather densely hirsute with moderately long stiff and erect hairs or setae. 

Gnathochilarium with lingual lamellae inclusive of processes three 
and a fourth times as long as the greatest width. Stipites in length 
about 4.7 or a little less times the greatest width. Promentum moderate, 
triangular in general outline, the basal margin broadly angularly 


1910] Diplopoda from the Western States 239 


excised. Mentum with distal border broadly angular, fitting into exci- 
sion of promentum; greatest width apparently about three and a half 
times the mesal length, but the position of the sclerite and some 
difficulty in determining the proximal limit leaves room for some uncer- 
tainty. 

Ocelli arranged in a narrowly sub-triangular patch, the apex directed 
mesad and toward that of the opposite eye; caudal margin shortest, 
a little convex, the dorsal margin convex near caudal margin and con- 
cave near apex or mesal angle, the ventral margin concave. Ocelli in 
four series, counting from ventral row dorsad as follows: 7, 6, 4, 3, 
giving all together 20 or thereabouts. 

Antennae strongly clavate, the fifth article longest and thickest, 
the first three especially much more slender. 

First segment clearly longer than the second, the cephalic margin 
mesally protruding, the lateral portions straight or but little concave. 
Caudal margin nearly straight, distinctly excised immediately mesad 
of carinal corners, the latter angularly projecting caudad or latero- 
caudad, the posterior of the two lateral bristles projecting caudo- 
dorsad and somewhat laterad from bottom of this excision. Depressed 
or widely furrowed longitudinally on each side between median line 
and the lateral margin, the anterior of the lateral bristles springing 
from the cephalic portion of its depression 1n a dorso-cephalic direction. 
The innermost bristle near cephalic margin and well toward the median, 
extending dorso-cephalad. 

Anal scutum in outline as viewed from above somewhat trapezoidal, 
the lateral margins not strongly converging; caudad widely sub- 
truncate, the lateral angles somewhat rounded. Papilla on each side 
nearly midway between the median line and the lateral margin, bear- 
ing a long curved bristle. A pair of long bristles borne on weak eleva- 
tions each side of the median line and half-way between caudal and 
cephalic limits; and each lateral margin with a long bristle arising 
from near anterior angle and a weak tooth or acute obtuse bristle 
midway between anterior and posterior ends. 

Anal valves with caudo-dorsal angle sub-acute, apically but weakly 
rounded, the caudo-ventral angle obtuse. Each valve bearing three 
long bristles, a little within the ventro-caudal margin, these bristles 
projecting caudad, the most ventrad being at the obtuse caudo-ventral 
angle or extension and closer to dorsal scutum than to anterior end of 
free margin, the other two equidistant from this one, with the upper 
one closer to scutum than to median bristle. 

Ventral scale bearing two long, caudally directed bristles. 

Anterior pairs of legs, in female at least, with distal joint bearing 
beneath a conspicuous comb-like series of stout bristles. See Pl. XX XIII, 
fig. 8. 

Appendages of second segment large and distinct. Two sub- 
cylindrical processes springing from a main plate or fold and extending 
ventro-cephalad, these processes translucent or transparent distally. 
see Pl. XXXIII, fig. 7. 

Length 7-8 mm. 


240 Annals Entomological Society of America  [Vol. III, 


Locality—Mill Creek Canyon, Utah. 

Found among dead leaves along the margin of a deeply 
shaded ditch on south side of canyon, the accumulation of 
leaves remaining very damp. 


Tingupa monterea sp. nov. 


Dorsum dark brown, a row of light spots along each lateral margin, 
a spot being on edge of each carina, and two other rows, more or less 
evident, between this row and a pale median longitudinal line, the 
inmost row being close to the latter. Venter pale. Feet white or yel- 
lowish. Vertex of head dark brown, the other portions paler, yellowish. 
Antennae brown, the articles pale distally. 

Segments bent first forward laterally and then again somewhat 
_ caudad distally, especially in case of the posterior segments, the lateral 
margins lying parallel with axis of body and not widely rounded or 
curved in mesad, the cephalo-lateral angles being in fact farther from 
the body than the caudal in most segments; a little incurved mesad 
of each caudo-lateral angle, but less conspicuously so than in utahensis. 
Segments, excepting first and last, with a fine longitudinal keel along 
mesal line. Carinae strongly developed and conspicuously extended 
laterad and meeting together rather closely; finely denticulate. Gran- 
ules or scales covering the segments densely in compact rows, their 
conical apices projecting like minute spines, at times giving the appear- 
ance of hirsuteness when viewed parallel to surface of segment, the whole 
surface thus appearing finely spinescent. Setae long and clavate, 
situated nearly as in utahensis. 

Anterior margin of first segment bow-shaped, being mesally moder- 
ately curved cephalad or convex, then each side concave, and lateraly 
again convex. Not mesally furrowed or keeled. 

Anal segment as viewed from above with the sides strongly converg- 
ing caudad, incurved; widely truncate caudad; a large mesal tubercle 
or elevation a little cephalad of caudal margin. 

Anal valves appearing a little to exceed the scutum. Dorso-caudal 
angle widely rounded, the free margin first curving evenly ventrad 
and cephalad and then running straight cephalad parallel with the 
dorsal margin. Dorsal and ventral edges elevate, the caudal not so. 

Head with vertex elevate, somewhat longitudinally rugose along 
caudal border. Conspicuously depressed on each side from base of 
antenna caudad, the antenna being inserted in anterior end of the 
furrow. Head densely hirsute with short setose hairs over entire sur- 
face, sometimes appearing a little sparser on vertex. 

Eye area small; the ocelli numbering eight or thereabouts, arranged 
in three series, counting from above toward antenna, 3, 3, 2; the side 
of patch toward median and that toward dorsal surface subequal, each 
consisting of three ocelli, and the two lying at right angles to each other 
or nearly so. 

Antennae short, strongly clavate, the fifth article decidedly longer 
and very much thicker than the third. 


1910] Diplopoda from the Western States 241 


Gnathochilarium with lingual lamellae, inclusive of processes, 
about 3.8 times as long as the greatest width which 1s anteriorly. Sti- 
pites inclusive of processes, about 4.25 times as long as the greatest 
width. Greatest width of mentum apparently three times the length, 
though the caudal margin was made out with some uncertainty. Pro- 
mentum distinct and well developed. Spathula evident. Stili not 
detected. 

Size approximately that of T. utahensis. 


Locality—Pacific Grove, California (June, 1902.) 


Caseya irritans sp. nov. 


General color in alcohol light brown; indications of pale, whitish 
transverse bands, one across each segment, and of a row of whitish 
spots laterally. Venter and the feet yellowish. Head with the vertex 
pale, finely areolated in brown; frontal region brown, a distinct pale 
spot ventro-mesad of each antenna, the spot transversely elongate, 
and a median pale spot immediately dorsad of the inner ends of these 
two spots; clypeal region pale; labrum yellow. 

Head widely excavated caudad. Vertex transversely elevated, 
smooth and shining in the specimen as at present, crossed by a faintly 
developed fine median longitudinal line. Frontal and clypeal region 
of head sub-complanate; a conspicuous transverse impression above 
clypeus, indented or bent dorsad mesally. 

Eye patch elongated in a meso-ventral direction. Ocelli strongly 
developed, 13 in number and arranged in three series, which, counting 
from above toward antennae are as follows: 7, 5, 1; on one side the series 
appearing rather as four, 1, 6, 5, 1. 

Antennae long and slender, scarcely clavate. The third article 
much longer than the fifth; the seventh much smaller than the sixth, 
being but little more than half its length. 

Promentum moderately large, triangular; mentum a little less than 
three times as wide as the median length. Length of lingual lamellae, 
inclusive of processes, a little less than three times as large as the great- 
est width; which is caudad of the middle. Stipites, inclusive of pro- 
cesses, about 4.4 times as long as the greatest width. Sensory cones 
of outer process of stipes 5 or 6 in number, of inner process apparently 
6. Stili not detected. 

First dorsal scutum sub-reniform, the anterior median portion 
rather narrowly protruding. Caudal margin with lateral angles evenly 
rounded, the median portion almost straight, slightly obtusely indented 
at mesal line. A fine median longitudinal line impressed across caudal 
portion of plate. Lateral regions of segment marked with numerous 
fine oblique striae, there being similar striae on the anterior rounded 
median portion but these extending but little caudad. 

Body fusiform, attenuated both caudad and cephalad. 

Segments caudad of the first with numerous striae over sides be- 
neath; on the second there are three striae above the obsolete shoulder- 
like bulging occurring where carina would usually be developed, on the 


242 Annals Entomological Society of America [Vol. III, 


third four or five, the striae existing farther and farther dorsad on sub- 
sequent segments, from the fifth caudad extending completely across 
the dorsum. From the fifth segment to the ultimate a band-like area 
across dorsum marked apparently with coarser striae and distinctly 
delimited laterally; on the more caudal segments the band extending 
caudad a little beyond the usual limit of segments and presenting weak 
crenations or dentations, the band farther cephalad on segment in 
median and anterior region of body. 

Anal scutum in outline as seen from above with sides converging 
caudad, nearly straight or but little incurved mesally; caudal margin 
mesally emarginate. Two impressions extending from anterior margin 
on each side of the middle and converging to meet in an acute angle at 
a mesal point on the caudal emargination of plate. Papillae slender, 
gradually acuminate. 

Each anal valve crossed longitudinally over dorsal portion by two 
or three striations. The caudo-ventral edge nearly straight, margined. 
Length, 10 mm. Greatest width 1.5 mm. 


Locality—Portland, Oregon (Aug., 1902). One female. 


FAMILY STRIARIIDAE. 
Striaria nazinta sp. nov. 


Body uniform light brown in color. Legs yellow. Antennae 
white. 

Body cylindrical, gradually attenuated cephalad as well as caudad. 

First segment large, expanded on each side in a plate which extends 
cephalad over lateral portion of head. Mesally the front margin is 
straight or nearly so. Anterior border with a ridge-like elevation con- 
tinuous laterally with edges of the lateral plates, the two ridges meet- 
ing in a mesal elevation and at an angle open cephalad, the surface of 
plate in front of this elevation obliquely descending to head, closely 
tuberculate. On main area of plate each side of the median line are 
six strongly elevated and relatively thin carinae of which the one nearest 
the meson is lower and begins farther cephalad than the next laterad; 
the farthest laterad of these carinae is much lower and shorter than any 
other and is the only one not attaining or nearly attaining the caudal 
margin. Laterad and cephalad of the carinae and to the very edges of 
the lateral plates, the segment is densely tuberculate. Between the 
carinae the granules are irregularly scattered, and relatively few, not in 
definite rows, though they may usually be regarded as representing two 
broken series in each interspace. 

Second segment with twelve dorsal and dorso-lateral carinae in 
addition to the major lateral carinae, but crossing the entire width of 
the major sub-segment and, as on all subsequent segments, more strongly 
elevated at caudal ends. Lateral carinae or plates less strongly ex- 
tended horizontally than the lateral plates of first but in general similar, 
the lateral portions less strongly granulate. 

Third and subsequent segments with twelve dorsal and lateral 
carinae in addition to the lowermost pair, the latter not at all carried 


1910} Diplopoda from the Western States 243 


out on horizontal plates or extensions like those of first two segments 
but reduced and comparable in size to the one immediately dorsad 
of it, and projecting cephalad instead of caudad as do the other carinae. 
These segments with only scattered and weakly developed granules, 
some appearing almost smooth. All marked with the usual fine median 
keel. 

Anal scutum conspicuously granulate with strongly developed 
tubercles. 

Anal valves not equalling the last dorsal scutum, narrow and elon- 
gate. Granulate with strongly developed tubercles similarly to the 
anal scutum. 

Caudal margin of head straight, not at all excavated. Vertex 
of head transversely elevated, the lateral portions ridge-like, the ends 
abrupt and conspicuous, while the mesal portion of the elevation 
extends ventrad and narrows to a mesal point attaining the imaginary 
line connecting the bases of the antennae, in front of which extension 
the level is depressed. Vertex crossed by a narrow median longitudinal 
furrow. Entire head granulate, the lateral portions more coarsely so. 
Hirsute, more strongly so over clypeus and adjacent region. 

Labrum set off by a transverse depression or furrow; lateral por- 
tions rounded; mesally with two conical teeth. 

At each lateral end of transverse elevation of the vertex a small 
darkened area apparently representing an eye; but in the type speci- 
men ocelli indistinct, three or four obscurely indicated areas apparently 
representing them, the granular character of the general surface making 
their certain identification difficult. 

Antennae short; strongly clavate; appressed close to head and in- 
conspicuous. Fifth segment longer and much stouter than the third; 
the seventh smaller than the sixth. Subdensely hirsute with short stiff 
hairs, the proximal articles rather more sparsely clothed. 

First legs bearing along the cephalo-ventral surface a calamistrum- 
like row of stout setae, elsewhere with fine hairs which are more sparse 
on proximal joints. A moderate tuberculate elevation toward distal 
end of femur on ventral surface like those present in males of some 
craspedosomatids. 

Other legs mostly but sparsely provided with hair; all tuberculate. 

Length, 13mm. Width, 1.2 mm., nearly. 


. Locality—Portland, Oregon (Aug. 1902). One female spec- 
imen. 


FAMILY XYSTODESMIDAE. 
Fontaria tuobita sp. nov. 


Black, the carinal and lateral margins narrowly bordered with 
pale. Anterior portion of prozonites yellowish. Sometimes the black 
of the dorsum is in varying degrees supplanted with brown which 
encroaches in irregular streaks and spots. Sides paler, smoky yellow, 
the dark color more pronounced adjacent to bases of legs and on caudal 
segments or forming distinct bands down each segment in alternation 


244 Annals Entomological Society of America  [Vol. III, 


with the bands of yellow of the prozonites. Venter pale. Legs yellow. 
Vertex and frontal region of head dark, a background of yellow being 
closely covered with a network of black; clypeal and labial region 
yellow. 

Head with median furrow of vertex sharply impressed, extending 
to an imaginary line tangent to the dorsal edges of the bases of antennae 
where it touches the mesal angle of an impression extending between 
antennae. Finely hirsute. 

Antennae slender, rather short. 

First dorsal plate a little narrower than the second. The lateral 
and cephalic margins together forming an even semicircular curve; 
caudal margins of carinae oblique, the intervening caudal margin of 
segment nearly straight. Mostly smooth and shining, finely rugose 
along cephalo-lateral border. 

Dorsum well arched, the carinae somewhat depressed below the 
horizontal. Second segment with cephalo-lateral and caudo-lateral 
angles a little larger than rectangular, narrowly rounded. In subse- 
quent segments the cephalic border adjacent to cephalo-lateral angles 
becomes more and more protruded cephalad, and the caudo-lateral 
angles become more and more produced caudad in an acute angle or 
process. 

Anal scutum terminating in a cylindrical process which is curved 
ventrad and is truncate distally. 

Anal valves strongly raised along free mesal margins. 

Anal scale triangular, the lateral margins convex proximally, else- 
where nearly straight; caudal angle narrowly extended a little and 
rounded. 

Ventral scuta transversely impressed, more strongly so on caudal 
segments; a little extended at a small elevation or process at base of 
each leg. 

Gonopods of male with the coxae relatively large and stout, distad 
of this extending ventro-cephalad; narrowed distad and terminating 
in a blunt pointed process which curves mesad and a little ventrad and 
then toward apex bending back proximad.. On the ventro-mesal sur- 
face at one-third the length from coxae, an acutely-pointed, blade-like 
process, which curving meso-ventrad and then distad crosses its fellow 
in the middle line. See further, Pl. XXXV, figs. 7 and 8. 

Length, 19 mm. Width 4.5 mm. 


Locality—Cloudcroft, Sacramento Mountains, New Mexi- 
co (Sept. T. D. A. Cockerell). 


FAMILY POLYDESMIDAE. 


Scytonotus piger sp. nov. 

Dorsum dark, sometimes of reddish caste. Venter but slightly or 
not at all paler than the dorsum. Legs uniform brown or yellow. 
Head and antennae dark brown. 

Head roughened. Vertex crossed by a short median longitudinal 
impressed line. Hirsute with short stiff bristles which are rather 
uniformly arranged. 


1910} Diplopoda from the Western States 245 


Antennae clavate. Moderately to sub-sparsely clothed with 
mostly short hairs. 

First dorsal plate less in width than either head or second dorsal 
scutum but relatively wider than in granulatus. Somewhat semicir- 
cular in outline but at each side extended into a narrow carina which is 
dentate, there being commonly two major teeth of which the caudal 
has a minor or secondary denticle on its posterior edge and in front of 
the anterior of which are several small denticles. Front margin mesally 
sub-straight, laterally convexly rounded. 

Depressed along the cephalic margin, the depression extending 
caudad mesally and narrowing to a furrow which again expands trian- 
gularly adjacent to the caudal margin. The segment elsewhere ele- 
vated. Densely covered with tubercles, the setae much as in gran- 
ulatus. 

Second segment with the lateral ends curved cephalad and embrac- 
ing the first. Segment longer laterally than mesally. Bearing three 
transverse rows of large tubercles. Carinae with cephalic angles acute 
but caudally widely rounded; laterally showing eight or nine rather 
small crenulations, the incision between them shallow; no translucent 
margin such as is present in granulatus. 

Subsequent segments gradually increasing in length and showing 
from three to five or six transverse rows of tubercles. Posterior angles 
of carinae becoming less widely rounded and extending farther and 
more acutely caudad, in the last few segments being a little produced. 
Lateral margins of carinae with small crenulations or teeth which for 
the most part are well rounded and not at all acute; teeth or crenula- 
tions numbering from eight to ten or twelve, the lesser number occur- 
ring on the spiraculiferous segments. Prozonites very finely and densely 
granulate. 

Anal plate convexly elevated proximally, bearing separated rows 
of conically pointed tubercles; two tubercles projecting laterad at 
base or proximal end of the cylindrical apical portion of scutum, this 
portion being slightly depressed ventrad and apically rounded and bear- 
ing on caudal surface four setigerous tubercles. 

Anal valves with mesal and caudal margins elevated and smooth. 
In some an irregularly oblong area over posterior portion of each valve 
roughened with sharply separated depressed areolae. A setigerous 
tubercle in furrow near caudo-mesal angle and a second similar one 
near middle of plate nearer mesal than dorsal margin. 

Anal scale with caudal extension convexly rounded and bearing 
two setigerous tubercles. Lateral portions in outline something like 
that of the fore part of a shoe. 

Length 12-13 mm. Width ad 2 mm. 


Locality—Upper branches of Mill Creek Canyon, Utah. 
A number of specimens were obtained about springs in 
upper side branches of the canyon. It seems to be close to 


S. granulatus though clearly distinguishable in numerous» 
details. 


246 Annals Entomological Society of America  [Vol. III, 


Polydesmus (Kepolydesmus subgen. nov.) anderisus sp. nov. 


Brown to dark reddish brown; carinae along lateral edges and above 
of lighter color, often yellowish, the pale bands commonly more dis- 
tinct caudad and frequently extending mesad along the caudal margin 
of each segment; often some of the anterior segments are dorsally dis- 
tinctly areolated with black, which may form a more solid transverse 
band along the cephalic margin excepting its lateral ends. Venter pale. 
Legs yellow or light brown. Head brown, a dark blackish band between 
the bases of the antennae, this band concavely excised on ventral side; 
labral region yellowish. Antennae darker than the legs, being concolor- 
ous with the dorsum of body. 

Vertex of head crossed mesally with a sharply impressed, distinct 
longitudinal furrow. Vertex and frontal region sub-sparsely clothed 
with short setose hairs, the clypeal region more sparsely provided with 
longer hairs. 

Antennae long; a little clavately enlarged distad. 

Body gradually narrowed cephalad from about the twelfth segment, 
less strongly so caudad. 

First dorsal plate scarcely narrower than the head inclusive of 
genae, its greatest width about 2.3 times the mesal length. Anterior 
margin mesally straight or nearly so, the lateral angles widely rounded. 
Caudal margin slightly and widely mesally incurved, laterally extending 
gradually cephalad to meet the lateral margin at a rounded angle. 
Entire border of margins depressed. A transverse impression caudad 
of the cephalic border, a broad median longitudinal depression extending 
from this to the depressed border along the caudal margin. The rows 
of minute setigerous granules distinct, somewhat uneven; but the 
main tubercles very obscure or not at all indicated. Segment finely 
roughened laterally. Each lateral margin with a single minute denticu- 
lation. 

Second segment bent cephalad on each side. Antero-lateral cor- 
ner sub-rectangular as is also the cauda-lateral, but the latter well 
rounded. Sharply margined laterally, caudally and also along lateral 
portions cephalically. Lateral margin of carina with one distinct 
denticle at antero-lateral angle and one or two faint or obsolete crenu- 
lations or denticles indicated farther caudad. 

Third segment nearly as the second; but the anterior margin imme- 
diately mesad of the cephalo-lateral angle more rounded and with a 
‘small indentation. 

From the second segment caudad the cephalo-lateral angle becomes 
more and more rounded and the caudo-lateral less and less so, the latter 
finally becoming somewhat produced though not acute. The produced 
caudo-lateral angle of the penult segment somewhat indented caudally. 
Segments dorsally depressed mesad of the carinae, the depression 
anteriorly bending obliquely cephalo-mesad. Carinae bent a little 
dorsad. Transverse furrow deep and distinct, extending across dorsum 
about half way from meson to each lateral margin. Tubercles not 
sharply delimited, but rather more distinct on caudal segments. 


1910] Diplopoda from the Western States 247 


Anal scutum with four denticulations along each lateral margin, 
each of these teeth bearing a stout seta. Dorsal surface with trans- 
verse rows of setae. Caudal process of the scutum straight, bearing 
four long, finely pointed setae. 

Ventral plates impressed with the usual transverse furrow and with 
a less distinct longitudinal one. Smooth except for a moderate cloth- 
ing of hair. 

Legs rather long and slender, the second article inflated. 

In the male the first two pairs of legs are conspicuously smaller 
and the ultimate joint is clothed within with stout setae subseriately 
arranged, the row being more even and well marked on the second. 
Other legs on inner surface with rows of setigerous tubercles, the setae 
being short, otherwise with sparse short hair only, except on the proxi- 
mal articles. Second legs showing a cone-shaped, apically truncate 
process on the ventral surface of coxa. 

For structure of copulatory appendages of the male see Pl. XX XVII, 
figs. 2, 3, and 4. 

Length of female, .26 mm. Width, 3 mm. Length of antennae 
ad 3.8 mm. 


Locality 


Kendrick, Idaho (W. M. Mann). 


Polydesmus (Kepolydesmus subgen. nov.) sontus sp. nov. 


Dorsum deep brown to almost black, the prozonites mostly paler; 
carinae yellow. Pleural region between carinae and the bases of the 
legs brown. First segment commonly darker than the adjacent ones, 
often black, the caudal and carinal borders and the minute setigerous 
granules yellowish. Venter and legs yellow. Head with the vertex 
dark brown to blackish, the dark area extending forward between the 
antennae as a narrow, median tongue-like band, the color commonly 
deeper at the base or proximal end of this band; sides of head dark, a 
yellow band between dark area of vertex and the sides extending obli- 
quely forward over insertions of antennae to the yellow lower frontal 
and clypeal area. Antennae yellow to brown, the sixth and seventh 
articles commonly conspicuously darker, the seventh being black and the 
sixth dark brown to black. 

Body gradually attenuated cephalad; over posterior portion with 
the sides subparallel excepting for the few last segments which are 
narrowed in the usual way. 

Vertex of the head crossed longitudinally by a fine median 1m- 
pressed line which extends almost to an imaginary line tangent to the 
upper edges of the antennal sockets. Frontal and clypeal region clothed 
. with setose hairs of moderate length, the vertex appearing glabrous or 
nearly ‘so. 

Antennae short, a little less than or at most equalling the width 
of the body. Scarcely clavate. Sub-sparsely hirsute, the hairs more 
dense on the flexed side. 

First dorsal plate in outline semi-circular or half-moon shaped, 
but the caudal margin a little incurved. Wider than the head inclu- 


248 Annals Entomological Society of America  [Vol. III, 


sive of the genae. Anterior border margined. Carinae narrow; bent 
dorsad; lateral margin with a single emargination but with no real 
denticulation. Crossed by a fine median longitudinal line. 

Second segment bent forward at the sides. The anterior angles 
of the carinae sub-rectangular, somewhat rounded, overlapped by the 
first plate. A.small denticulation on outer side of the cephalo-lateral 
angle and two weak or obsolescent emarginations indicated farther 
caudad. Caudo-lateral angles not rounded, a little obtuse. Caudal 
margin of carina excavated or abruptly bending meso-cephalad. Cari- 
nae rather narrow, bent somewhat dorsad. Inter-carinal region of 
dorsum strongly elevated and convex as it is on subsequent segments. 

Lateral margin of carina of third segment shorter than that of the 
second. Anterior angle rounded, the emargination caudad of it as on 
the second segment, the denticle in front of the emargination likewise 
low and rounded. Posterior angle rectangular or acute. Caudal 
margin of carina straight, extending cephalo-mesad without showing 
any curvature. 

Fourth segment similar to the third but the lateral carinal margin 
longer, sub-equal to that of the first, and the caudal angle rectangular 
or slightly obtuse. Only the single anterior emargination indicated. 

Fifth segment abruptly longer. Lateral carinal margin bulging 
laterad, being mesally very obtusely angular. Caudal angle obtuse 
and rounded. The anterior emargination very weak. 

Sixth segment a little longer than the fifth, the lateral carinal 
margin but weakly and evenly convex. The emargination very faint. 

Subsequent segments sub-similar but the emarginations not at all 
manifest. The caudal angles in proceeding caudad become more and 
more acute and finally extended or produced sharply. All but a few 
of the segments with the dorsal transverse sulcus sharply impressed 
and extending well toward the carinae where it becomes more indis- 
tinct. A median longitudinal impression which is distinct in front of 
the transverse sulcus which is mostly vague or absent caudad of it. 

Anal scutum with the process truncate and apically the caudal 
end shallowly emarginate. 

Anal valves with the caudal margin a little incurved, meeting the 
mesal margin at an acute but rounded angle. Mesal margin elevate. A 
short curved impression near the mesal margin of each valve and about 
the middle of its length, the concave side being mesad. A longer and 
also slightly curved impression toward dorsal margin, the concavity 
being dorsad or ectad. 

Anal scale relatively long, the caudal border emarginate. Later- 
ally weakly emarginate at proximal ends of sides. 

Ventral plates with a longitudinal median impression which is 
distinct on the posterior segments but is indistinct or not at all evident 
cephalad. Transverse impression mostly very weak. All rather dense- 
ly clothed with hair, some of which is conspicuously long. 

Legs sparsely clothed with short stiff hairs which are most abun- 
ant on distal joints and on the ventral or inner side. 


1910] Diplopoda from the Western States 249 


In the male the first legs are, as usual, small, but the second are 
not conspicuously reduced. The bristles on the ventral surface of the 
distal joint of first legs more densely and sub-seriately arranged than 
on the others. 

Copulatory appendages of the male moderately small, strongly 
bent cephalad and parallel with the body. The main outer process 
of each side bearing distally three chitinous appendages. The most 
proximal of these is short and blade-like and extends dorso-mesad and 
a little cephalad, crossing with its mate at the middle line. The sec- 
ond takes its origin cephalad and dorsad of the first and as a long, con- 
spicuous, flattened and somewhat twisted blade extends first cephalad 
and then curves ventro-mesad. The third takes its origin a little dorsad 
from the base of the second and curves first dorso-mesad and then 
ventro-mesad, the distal portion curving finally ectad again and ending 
in a slender acute point. From the ventral concavity of this appendage 
springs a slender process which bears distally a stout curved seta which 
extends caudad to end in the neighborhood of the first process. See 
buctner Pl: XOXCOXVIlI figs. 2 and 3. 

Length 19-20 mm. Width 2.5 to 2.7mm. Antennae ad 2.5 to 
At Tam. 


Locality—Los Angeles, Cal. (June, 1909.) 


Polydesmus amandus sp. nov. 


Segments with the main dorsal divisions whitish, a tendency for the 
median area contiguous to the cephalic margin to be brown, the ulti- 
mate segments and a few of the most anterior commonly more or less 
completely brown; the prozonites clear brown. First dorsal plate clear 
brown. Venter white, excepting a few of the most caudal and of the 
most anterior segments which are brown. Legs white but with the 
coxae all brown and with a decided tendency for the distal articles like- 
wise to be brown or brownish. Head uniform brown, the color being 
of a lighter shade than that of the first segment. Antennae uniform 
brown or with some of the median articles whitish in whole or in part. 

Head nearly smooth, the vertex crossed longitudinally by a short 
median impressed line. Clothed rather densely over nearly the entire 
surface with very short hairs. 

Antennae a little longer than the body is wide. Clavate, the sixth 
article being considerably the thickest. 

First dorsal plate narrower than the head inclusive of genae and 
than the second segment. Transversely sub-elliptical. Lateral cari- 
nal margin with three teeth; a fourth tooth projecting from edge on the 
cephalo-lateral curve. _Tuberclesin numerous rows; strongly developed; 
apically acuminate and setigerous. The setae curved or frequently 
hooked distally. A transverse impression or sulcus cephalad of the 
row of tubercles along caudal border. 

Second segment strongly bent cephalad at the sides. On the cari- 
nal margin immediately caudad of the acute denticle of the cephalo- 
lateral angle is a more obtuse one and caudad of this five larger, low 


250 Annals Entomological Society of America  [Vol. III, 


crenations which are continuous as a series with a row of crenations or 
projecting tubercles along the caudal margin; all are setigerous. Tuber- 
cles all strongly developed. Carinae sub-horizontal. 

Carinal margin of the third segment rather longer than that of the 
second but similarly sculptured. 

Subsequent segments becoming longer, the rows of tubercles they 
bear more numerous, the carinal crenulations weaker, and the caudo- 
lateral angles more rounded excepting in the last few where they tend 
toward rectangular. In the antepenult segment alone are the caudo- 
lateral angles a little produced. A transverse sulcus between the sec- 
ond and third or the third and fourth rows of tubercles from the caudal 
margin. Two longitudinal impressions, one a little each side of the 
median, extending from cephalic margin a short distance caudad and 
then curving out laterad. Wide spaces intervening between the carinae 
of adjacent segments. 

Anal scutum bearing transverse rows of strongly developed, coni- 
cal setigerous tubercles like those of other segments, these rows well 
separated from each other. Apical process of scutum decurved, coni- 
cal, not truncate, and bearing long setae as usual. 

Anal valves with the dorsal and mesal margins sub-parallel, the 
caudal margin meeting the mesal in a rounded but as a whole somewhat 
acute angle. A little cephalad of the caudo-mesal angles is a transverse 
impression crossing the median line from valve to valve. Mesal mar- 
gins strongly elevated. A conspicuous conical and setigerous elevation 
near the middle of each valve, this elevation closer to the mesal than 
to the dorsal margin. 

Anal scale with each half in outline like that of shoe with high 
instep. Caudal margin crenately incised a little mesad of each caudo- 
lateral angle, the crenulation adjacent to each angle bearing a long 
seta. 

Ventral plates pilose. The transverse impression in most more 
strongly developed than the longitudinal. 

Legs rather long, very sparsely hirsute proximally but more dense- 
ly so distally. Second and third articles inflated dorsally. 

The first legs reduced as usual, the second intermediate in size. 

Gonopods of male of moderate size. Each one presenting two main 
chitionous processes of which the cephalo-mesal one is apically biden- 
tate and presents near its base an acute conical tooth projecting in a 
cephalo-mesal direction. The second process lies against the first for 
most of its length but distally diverges from it, extending more directly 
ventrad and ending in an acute point. See further Pl. XXXVIII, 
fig. 6 and Pl. XXXIX, fig. 1. 

Length of male 18 mm. Width 1.9 mm. Length of antennae 
2+ mm. 


Locality—Mill Creek Canyon, Salt Lake Co., Utah. 
(Sept., 1900). 


bo 
Or 
— 


1910] Diplopoda from the Western States 


Polydesmus sastianus sp. nov. 


Dorsum brown; the carinae paler, yellowish. Sides pale brown. 
Venter and legs yellow. Head light brown. Antennae light proxim- 
ally, the distal joints brown. 

Head finely roughened. The vertex crossed with the usual im- 
pressed line. Clothed rather densely with very fine short hairs. 

Antennae short. Clavately thickened. Distal articles sub-densely 
clothed with moderately short hair; but with only a few hairs on the first 
and especially on the second articles. The four sense cones conspicuous. 

First dorsal scutum narrower than the head and also distinctly 
narrower than the second segment. Transversely sub-elliptical with 
the caudal margin, however, but little convex. Tubercles strongly 
developed, conical, in crowded series. The setae conspicuous, those of 
the rows of tubercles along cephalic and caudal margins very long and, 
as usual, curved or hooked distally. 

Second segment not strongly bent forward laterally. The conical 
tubercles strongly developed and densely arranged as on the first one. 
Carinal margin with the anterior and the posterior angles about equally 
rounded; laterally with six, large, angular, setigerous teeth, of which 
the most cephalic, which is located on mesal side of cephalo-lateral 
angle is smallest and most acute, the fourth and fifth with a secondary 
indentation on their cephalic edge and three similar teeth on caudal 
side. The series of teeth continued along the caudal edge of plate by 
a row of large projecting tubercles. 

Third and fourth segments with their lateral margin shorter than 
that of the second; the caudo-lateral angle of carina more rounded; 
lateral margin with same number of teeth as the second. 

On the subsequent segments the caudo-lateral corners of the 
carinae become more angular, first becoming sub-rectangular and then 
on the last two preceding the anal plate a little produced caudad. 
Entire dorsum with a longitudinal median keel-like elevation which is 
better developed on the prozonites. Transverse sulci rather weak. 

Anal scutum strongly tuberculate, the tubercles conical and 
setigerous like those of the preceding segments. Caudal process nar- 
rowly rounded. ; 

Anal valves somewhat as in the preceding form. 

Anal scale sub-triangular. Caudal angle a little rounded. Seven 
pairs of setigerous tubercles along the lateral margins. 

Most of the ventral plates are very distinctly marked both with 
the transverse and with the longitudinal impressions; of these impres- 
sions the longitudinal one is narrower and usually appears more sharply 
impressed. 

Legs moderately clothed with rather short hair, the first two joints 
more sparsely so. 

Length: 10-11 mm. Width 1.5 mm. Length of antenna ad 
1.5-1.6 mm. 


Locality—Shasta Springs, Cal. (Aug., 1909.) 
Three specimens were secured, one adult female and two 
not fully grown males. 


252 Annals Entomological Society of America  [Vol. II], 


Polydesmus socarnius sp. nov. 


Dorsum brown; the carinae commonly paler; a median longitu- 
dinal dark line indicated in some, this more distinct caudad. Venter 
scarcely paler than dorsum. Legs yellow or light brown. Head brown, 
the genae and the clypeal and labial region paler, yellowish. Antennae 
brown, darker than the legs, but light distally. 

Vertex of head crossed by the usual median longitudinal impressed 
line, this line extending to the angle between two furrow-like impres- 
sions which extend from the median ventro-laterad to the bases of the 
antennae. Head conspicuously longitudinally furrowed along each side 
in line with the antenna, the latter being inserted in the furrow. Some- 
what finely rugose over vertex. Moderately pubescent with rather 
short hairs. 

Antennae a little longer than the width of the body, clavate to the 
sixth joint, seventh article comparatively large, showing the usual 
sense cones distinctly. Clothed densely with mostly short hairs. 

First segment wider than the head without the genae but dis- 
tinctly narrower than the head inclusive of the genae and than the 
second segment. Transversely somewhat elliptical; the cephalic mar- 
gin widely and but weakly convex; cephalo-lateral angles strongly 
rounded, the margin evenly curving to the middle of the lateral carinal 
edge where it ends on each side in the single obtusely angular tooth. 
Caudo-lateral corners of the carina sub-rectangular, the caudal margin 
of the carina being nearly straight, untoothed. A row of four contig- 
uous tubercles adjacent to the caudal margin along its median portion. 
The median convex area of segment with tubercles weakly or but 
obscurely indicated. 

Second segment bent cephalad laterally as usual; overlapped by the 
first plate. Lateral carinal margin with three strongly developed teeth 
of which the cephalic is most acute and of which the most posterior is 
at or but little caudad of the middle of the margin, the latter caudad of 
this tooth being straight or but little curved to the sub-rectangular 
caudo-lateral angle. Posterior edge of carina straight and untoothed. 
Three rows of tubercles which are not closely placed, the two first rows 
extending from carina to carina; the row along the caudal border con- 
sisting of six elongate and roundly conical tubercles, which project a 
little beyond the caudal margin and are widely separated from each other, 
the two more median ones being closer to each other than are any other 
two. 

In the following segments a fourth tooth, at first ventral in posi- 
tion, occurs between the caudal angle and the median tooth of the 
carinal margin. The caudo-lateral angle becomes gradually more and 
more acute and extended caudad, on most of the segments being conspic- 
uously produced. Most of the segments typically with a narrow longi- 
tudinal furrow adjacent to the carinal margin, a little mesad of this 
with a wider and deeper furrow and again a little farther mesad a sec- 
ond narrow furrow, there being, between the large and more evident fur- 
row and the narrower one each side of it a carina-like elevation which 


1910] Diplopoda from the Western States 253 


is commonly rather conspicuous. Dorsum as a whole complanate. 
Carinae a little elevate dorsad. 

Anal plate with distal process apically decurved. 

Anal valves with the mesal edges strongly margined and elevated. 
The usual setae present. 

Anal scale sub-triangular. 

Ventral plates with the usual cruciform impression, the transverse 
furrow being the wider and deeper in most plates. 

Legs clothed with short stiff hairs, this rather longer and more 
abundant on distal joints. 

Length, 10-11 mm. Width, 1.3-1.4 mm. Length of antenna ad 
1.6 mm. 


Locality 


Salt Lake City, Utah (June 23, 1900). 


FAMILY PARAIULIDAE. 
Paraiulus zakiwanus sp. nov. 


Color brownish red. The first segment clear reddish without 
darker markings; but most of the body subsequent to this with a row 
of black dots along each side, one to each segment, which on the more 
posterior segments is supplemented with an area of smoky or blackish 
which tends to extend or diffuse about the segment and thus much to 
darken it. No distinct median dorsal line and no light spots. Vertex 
of the head dark, a band between the eyes darkest, black; the head else- 
where reddish, the red intruding into lower border of the dark area in 
three semicircular incisions. Legs brown. 

A sharply impressed furrow extending from the upper margin of the 
eye area mesad curving evenly mesad and then bending sharply back 
dorsad to meet its fellow at an angle on the mesal line; the median longi- 
tudinal line extending distinctly to this angle. A second impressed 
line a little cephalad of the caudal margin of the head, this line being 
bent forward at a sharp angle mesally. Head very finely rugose. 

Eyes are large, sub-triangular, the ventro-lateral side and the caudal 
side convex, the mesal being concave. Ocelli 63-65, arranged in nine 
transverse series which, counting from above ventrad are as follows: 
11 1059,-9) (8), 7 6, 5, ey. 

First dorsal plate with the lateral margins not angularly produced 
beneath but nearly straight and subparallel with the axis of the body. 
Entire border of plate distinctly margined. Finely rugose or nearly 
smooth, not at all striate. 

Second segment with four deep striae on each side beneath and on 
caudal division, the lowermost margining the lower edge. 

Subsequent segments also deeply striate beneath, the striae well 
spaced apart. 

Spiracles rather small; touching the transverse suture which is bent 
at their level. 

In the male the sixth and seventh segments are moderately extended 
beneath the lower margin of the latter evenly convex in outline, not at all 
angular. 


254 Annals Entomological Society of America  [Vol. III, 


First legs of male strongly enlarged as usual. Uncinate. Inner 
surface covered with rounded tubercles which are arranged in transverse 
rows as shown in Pl: XL, fig. 4. 

Second legs of the male with the coxae much enlarged in the usual 
manner; produced mesally into a long tongue-shaped process which dis- 
tally is enlarged and sub-truncate and which extends ventro-cephalad 
between the legs of the first pair. Claw of legs small, concealed with 
hair which clothes the distal end of legs sub-densely. 

First pair of gonopods in the male with the anterior division much 
shorter than the posterior, strongly and subquadrately enlarged distad 
and clothed densely with long hairs. The anterior branch with the 
more slender distal portion bent first mesad almost at right angles and 
then caudad parallel with the axis of the body, drawn out distally to a 
narrow, slightly rounded point, the lower margin of the mesally extend- 
ing division irregularly toothed. See Pl. XL, figs. 1, 2 and 3. 

Second pair of gonopods inserted a considerable space from the base 
of the anterior pair, above base curving forward parallel to the body 
and then curving ventrad in engagement with the inner branch of the 
first pair. From about the end of the first third of the length arises a 
slender process which in lateral view appears to terminate in a spear- 
shaped head, this process crossing the main branch in a caudo-dorsal 
direction. Main process terminating in two branches, a broad plate- 
like branch through which the duct opens and a more slender acute pro- 
cess springing from the ventral aspect of the former. See further 
Pl. XX XIX, figs. 6 and 7. 

Width of body 2.6 mm. Length uncertain, the caudal end of the 
body missing. 


Locality—Sacramento Mountains, New Mexico. (T. D. 
A. Cockerell.) 


Paraiulus tiganus sp. nov. 


Deep brown to almost black. A yellow to white stripe encircling 
the body at junction of adjacent segments and a narrower line-like ring 
of same color encircling each segment near transverse suture. On the 
dorsum of each segment each side of the median and on the anterior 
portion of segment is a curved light line, the concavity of which is 
caudad. <A dorsal median longitudinal line of black which is thickest 
on each segment between the mesal ends of the two curved light lines. 
A row of black dots along each side, there being a dot on each segment 
cephalad of the narrow light ring. Back of each black dot is a white 
one which encloses spiracle. First dorsal plate deep brown or blackish, 
a solid black band along the cephalic border which is extended caudad 
mesally, narrowly edged with pale. Vertex of head dark brown; clypeus 
and an area on each side embracing insertion of antenna and extending 
ventrad from it pale; eye patches and a triangular area between them and 
extending ventrad black, the median area including three light dots. 
Antennae blackish. Legs brown to yellow. Last plate like the first 
one, deep brown or blackish, narrowly edged with pale. 


1910] Diplopoda from the Western States 255 


Head with a furrow-like depression between the two eyes, the fine 
median longitudinal line extending from this across the vertex. 

Eye areas oval but with the inner ends obliquely truncate. Ocelli 
convex, arranged in eight transverse rows as follows, counting from 
eye entrade 4.8, 9,9, 10, 11, 9, 5, making a total of 65, in ae speci- 
men described. 

Antennae of medium length, slender. 

Cervical plate large. A weak median longitudinal impression 
which is obsolete on anterior portion. Lower lateral portion impressed 
with a longer striation near and subparallel with margin and dorsad 
of this and “extending from the caudal margin a second, short stria, one 
or two other similar ones sometimes indicated. 

Other segments sharply striate beneath. Spiracles small, not 
touching the transverse suture which is sinuate at its level. 

Anal scutum smooth. Distal portion triangular, the apex sharply 
angular, straight, not at all decurved. 

Anal valves with the mesal margin elevated, setigerous. 

Anal scale roughly triangular; each lateral side mesally convex; 
incurved adjacent to proximal and distal angles; cephalic margin widely 
convex. 

Male: Gnathochilarium with stipes not produced mesally. The 
very large promentum obovate, a little crenately incised each side of the 
middle on distal margin and also proximad of each disto-lateral corner. 
Hypostoma free except laterally. See Pl. XL, fig. 6. 

First legs of male strongly enlarged, uncinate. The inner surface 
tuberculate as usual, and bearing long bristles as shown in the figure. 

Second legs of male reduced excepting for greatly enlarged coxae, 
as usual; the claws small. See Pl. XL, fig. 7 

Gonopods of male large and conspicuously exposed. The anterior 
pair with the mesal branches a little the longer, attenuated from the 
base distad but produced caudo-laterally near or a little below its mid- 
dle height. Outer branch clavately enlarged distad, embracing the 
first on “the antero-lateral surface; pilose with long hairs over the distal 
end and proximally along the disto-mesal face for part of the length. 

Posterior gonopods with the outer distal branch somewhat clavate 
apically and densely pubescent with fine short hairs. Inner branch 
almost straight, blade-like, acutely pointed. From the inner edge of the 
outer branch springs a third, spine-like, process which extends distad 
ee the two main ones. See Pl. XL, fig. 8, and Pl. XLI, figs. 1, 
2 ancy 3 

Length of a male 55 mm. Width 3 mm. Length of antenna ad 
3mm. Number of segments 51-57. 


Locality—Canyons of the Wahsatch Mountains, Utah. 

The most common diplopod in Utah. It is evidently 
closely allied to P. furcifer, which it resembles much in general 
appearance and in the general type of gonopods in the male. 
The latter are readily to be distinguished, however, both from 
characters in the anterior pair and especially those of the pos- 


256 _ Annals Entomological Society of America  [Vol. III, 


terior pair which differ in ending in three processes instead of 
in two, a slender spinous process appearing between the two 
major ones which are alone present in P. furcifer, etc. 


Paraiulus furcifer Harger var. sinampus, new. 


A row of small dark, inconspicuous dots along each side toward 
dorsum, the dorsum above and between the two rows of dots light yel- 
lowish brown, the sides below the dots distinctly darker, brown; each 
segment with a dark blackish brown to black annulus which is broadest 
across dorsum and narrows ventrad down each side; a narrower and rath- 
er less deeply colored dark line along junctions of adjacent segments 
which below disappears in the brown of the sides. A fine median dorsal 
dark line, on each side of which on each segment is a yellowish white 
line which does not extend laterad as far as the line of dark dots. First 
dorsal plate dark brown or chestnut, especially the caudal border pale. 
Penult and antepenult segments and the proximal portion of anal scu- 
tum pale. Venter pale. Legs medium brown. Vertex of head black, 
the dark area extending ventrad between the antennae; clypeal and later- 
al regions of head pale. Antennae black or blackish. 

Body in the not fully adult male being described somewhat con- 
stricted caudad of head, the first segment narrower than the second and 
subsequent ones. 

First segment moderate; the lateral margins running obliquely 
from caudal edge of head ventro-caudad to junction with posterior mar- 
gin which curves forward to angle. Lower lateral portion of plates 
marked with two distinct striae which extend forward a short distance 
from the caudal margin, in addition to the line margining the lateral 
edge. Caudal border not margined, the anterior one obscurely so. 

Second segment with the lower lateral border extended ventrad 
mesally in an acute angle of which the cephalic side is much the longer. 
Deeply striate beneath, above somewhat roughened, as on subsequent 
segments. 

Spiracles more or less elongate longitudinally, moderate, separated 
from the transverse suture which is somewhat weakly quadrately emar- 
ginate opposite them. 

Anal plate not roughened. Distal portion subtriangular, straight, 
apically somewhat rounded. Extending a little beyond the anal valves. 

Anal valves with mesal edges narrowly margined, not elevate. 
Each valve with a longitudinal impression ectad of its middle. 

Anal scale subtriangular, the caudal angle acute, not at all rounded. 

Legs rather long. Dorsally glabrous. Inner surface with longi- 
tudinal series of stout, spinescent setae. 

Head roughened with areolations formed by a network of impressed 
lines. A bow-shaped line impressed between inner angles of the two 
eye areas, the line angularly bent dorsad at middle where it is joined 
by the median longitudinal line of the vertex. A wide, shallow, median 
depression from frontal region to clypeus which is deepest ventrad. 
Head furrowed below eye area on each side from base of antenna toward 
gena and then dorsad along the border of the head. 


1910] Diplopoda from the Western States 257 


Eye area large, subtriangular, the dorso-mesal and ventro-lateral 
sides convex and the ventro-mesal nearly straight. Ocelli arranged in 
eight straight transverse rows which, counting from above ventrad, are 
as follows: 8, 10, 9, 8, 7, 6, 5, 3, making a total of 56. 

Antennae slender, a little enlarged distally, the sixth article thick- 
est. Clothed with short hairs intermixed with longer ones, the hair 
more dense distally. 

First legs of male thickened, but as yet straight and not distally 
hamate. Coxae fused in middle line. See Pl. XLI, fig. 8, and PI. 
CM figs 2. 

Second legs not as yet reduced, as large as the third but the four 
distal articles forming a hook which seems firm and is directed cephalad. 
Claw large. 

Lower margin of seventh segment but slightly angularly extended. 
Gonopods enclosed in a single sheath, the body being very long and pre- 
senting indications of the separate pieces only distally. See Pl. XLI, 
figs. 5 and 6. Upon removal of the envelope the copulatory organs 
are revealed in their usual character and relations, but the posterior 
pair not yet engaged distally with the inner branches of the anterior 
pair as in the adult furcifer, its two prongs being straight and closely 
appressed together and the outer one distally pilose. Outer branch 
of anterior pair provided with long hairs as in the adult furcifer. 

Number of segments 58. 

Locality—Portland, Oregon (1902). 

The description above is of a male which as indicated pre- 
viously, is not wholly mature, although apparently or probably 
within one moult of being so. While the copulatory organs 
when released from the peculiar sheath are seen to be fully 
formed and in most respects in close agreement with those of 
typical furcifer, the coloration is markedly different from that 
usually found in P. furcifer of the corresponding age. Further- 
more, an adult female collected at the same time has precisely 
the same coloration and differs from furcifer. It has seemed 
best to designate the form for the present at least, therefore, 
as a distinct variety. Because of the interesting points pre- 
sented the young male is described in full and illustrations of a 
number of its features presented. 


FAMILY PAEROMOPIDAE. 
Paeromopus lysiopetalinus Karsch. 

Figures of this interesting form, so long obscure as to its 
affinities because of incompleteness of the original description 
and lack of specimens in collections, are presentedin Plate XLII, 
figs. 3to 5,andin Plate XLIII, figs. 1 to3. The author has speci- 
mens of this form from Portland, Oregon, and from Pacific 
Grove and Stanford, California. 


258 Annals Entomological Society of America  [Vol. III, 


FAMILY CAMBALIDAE. 


Paiteya, gen. nov. 


Eyes fairly well developed, consisting of a moderate number of 
ocelli arranged in two or more rarely in three transverse series. 

Antennae short, a little clavately thickened distad. Second article 
longest, the sixth next; the others much shorter, the third, fourth and 
fifth not much differing in length. The four sensory cones slender and 
moderately long. 

Gnathochilarium narrowest proximally. Promentum elongate, 
triangular completely separating the laminae linguales which are strong- 
ly narrowed proximad. 

Body iulus-like in general form. All segments striate beneath; 
dorsum of segments from the fourth to the antepenult, not inclusive, 
each with a series of strongly developed keels of which the most ventrad 
on each side is thickened and bears on the enlarged caudal portion of 
edge the spiracle. ‘ 

Legs definitely armed with spiniscent bristles. 

First legs of male reduced; with six articles; normally armed. 

Second legs of male likewise reduced but otherwise normal in 
structure. 


Type—FPatteya errans sp. nov. 
Distribution—Southern California. 


Paiteya errans sp. nov. 


Brown in transverse encircling bands which, except on anterior 
segments, alternate with paler yellow bands which are broadest on dorsal 
surface and increase in extent on posterior segments; fifteen or so of the 
segments immediately following the fourth with an oval pale spot 
enclosed in the dark band laterally, these spots on posterior segments 
becoming confluent on the caudal side with the yellow of the adjacent 
yellow stripe; first four segments usually unmarked laterally with any 
such pale spots. A row of black spots on each side in line with the 
spiracles and extending caudad as far as the antepenult segment, not 
inclusive; these dark spots sharply defined excepting on the most anter- 
ior segments where they tend to be more diffuse and commonly enclose 
a paler area. First four segments commonly with a pale transverse 
band on dorsum, this band narrow at mesal line and expanded laterally, 
commonly areolated with dark. All dorsal carinae dark along their 
edges. Anal scutum brown excepting the caudal portion. Venter, 
legs and antennae yellow. Vertex crossed by a transverse yellow band 
which expands about the black eye area on each side. Between eye 
and antenna of each side and those of the opposite is a broad brown 
area which is often areolated with pale. Head below this latter dark 
band yellow. 

Body constricted caudad of head, most strongly so over from about 
fourth to ninth segments. 


1910] Diplopoda from the Western States 259 


Eyes in adults with ocelli in two rows which run a little oblique to 
the lateral portion of front margin of the first segment, which covers 
several of the more caudal ocelli. Ocelli commonly about nine arranged 
with the greater number in the dorsal series, thus: 6, 3; occasionally a 
single eye in a third row. . Eye area extending almost transversely. 

Antennae short. A little clavately thickened. The second and 
sixth articles longest; others distinctly longer than broad. Proximal 
articles sparsely provided with hairs of moderate length, especially 
toward the distal ends, the distal articles more densely clothed. 

Gnathochilarium conspicuously broader distally than proximally. 
Lingual lamellae completely separated by the long, triangular promen- 
tum. Laminae strongly narrowed proximally, 2.75 times as long as the 
greatest width. Mentum toward proximal border with a sharply out- 
lined semi-circular impression which has its concavity distad; wider at 
base than the median length in the ratio 11:9. Stipites nearly 3.5 times 
as long as the greatest width, the processes being included. In one 
specimen the processes of stipes ioe apparently completely fused togeth- 
er as shown in Pl. XLIII, fig. 

First segment nearly as long eee as the second and third taken 
together. Extending over the caudal portion of head. Posterior mar- 
gin straight. Lateral margins widely rounded from caudo-lateral angles 
cephalo-mesad. The cephalic margin substraight mesally. Within 
each lateral margin several longitudinal striae extend cephalad from 
the caudal margin a mostly short distance. 

Second segment with deep striae below, the striae on subsequent 
segments extending farther dorsad, on all but the most anterior and the 
most posterior reaching the lateral carina which is borne on all but the 
first segments. Spiracles beginning on the fifth segment. Dorsum of 
segments from the fourth to the antepenult (not inclusive) each with 
six strongly developed carinae, of which the lowermost on each side is 
conspicuously thickened. A weak, fine median keel along dorsum. 

Anal scutum large, smooth, caudally rounded. 

Anal valves exceeding the anal plate, the free margin thickened. 

Legs without true hairs; but armed with definitely arranged spines 
or spinescent bristles. Claws very large, each armed with a well- 
developed spine springing from its base on the ventral side. 

In the male the first two pairs of legs are reduced, but otherwise 
unmodified, having six articles and being normally armed. 

Gonopods of male concealed excepting for the distal end of slender 
curved process which projects caudad on each side from beneath a 
covering plate. 

Number of segments 47-49. 

Length 19mm. Width 1.4 mm. 


Locality—Southern California. 


260 Annals Entomological Society of America  [Vol. III, 


EXPLANATION OF PLATES. 


PLATE XXX. 
Lystopetalum mutans sp. nov. 


Fic. 1. Gnathochilarium. X 70. 

Fic. 2. Antenna. X 70. 

Fic. 3. Eye. X 70. 

Fic. 4. ‘‘Ovipositor’’, lateral aspect. X 39.5. 

Fic. 5. The same, cephalic aspect. X 39.5. 
Fic. 6. First legs, caudal aspect. X 47. 

Fic. 7. Second legs, cephalic aspect. X 39.5. 

Fic. 8. Third legs, caudal aspect. X 39.5. 

Fic. 9. Fourth legs, caudal aspect. X 39.5. 

Fic. 10. Fifth legs. X 39.5. 


PLATE XXXI, 
Lysiopetalum mutans sp. nov. (cont.) 


Fic. 1. Ninth leg, caudal aspect. X 39.5. 

Fic. 2. Thirteenth leg, caudal aspect. X 39.5. 
Conotyla deseretae sp. nov. 

Fic. 3. First legs of male. 

Fic. 4. Gonopods of male, caudal aspect. 

Fic. 5. Gonopods of male, cephalic aspect. 

Fic. 6. Gonopods of male, lateral aspect. 

Fic. 7. Eighth legs. 

Fic. 8. Antenna. 


PLATE XXXII. 
Conotyla deseretae sp. nov. (cont.) 


Fic. 1. Second legs. 
Fic. 2. Third legs. 
Fic. 3. Fourth legs. 
Fic. 4. Fifth legs. 
Fic. 5. Sixth legs. 
Fic. 6. Seventh legs. 
Fic. 7. Gnathochilarium. 
Conotyla coloradensis sp. nov. 
Fic. 8. Appendages of second segment of the female. 
Fic. 9. Eye. 


PLATE XXXITI. 
Conotyla coloradensis (cont.) 


Fic. 1. Gnathochilarium. X 39.5. 

Fic. 2. Antenna. X 87. 

Fic, 3. Seventh article of antenna, more highly magnified. 
Tingupa ulahensis gen. et sp. nov. 

Fic. 4. Antenna. X 70. 

Fic. 5. Eye. X 87. 

Fic. 6. Eye of an immature specimen. X 87. 

Fic. 7. Female appendages of the second segment. 

Fic. 8. First legs of female. X 70. 


PLATE XXXIV. 
Tingupa utahensis gen. et sp. nov. (cont.) 


Fic. 1. Gnathochilarium. X 87. 
Fic. 2. Caudal end of body, dorsal view. X 47. 


1910] 


FIG. 
FIG. 
FIG. 


FIG. 
FIG. 
Fic. 
FIG. 


oe oe 


© 00ND 


Diplopoda from the Western States 


Tingupa monterea gen. et Sp. nov. 


Eye. X 87. 
Antenna. X 87. 
Gnathochilarium. X 70. 


Caseya irritans sp, NOV. 
Eye. X 8/7. 
Head and first six segments, lateral aspect. 
Caudal end of body, dorsal view. X 18.5. 
Antenna. X 33. 


PLATE XXXV. 


FIG. 


FIG. 
FIG. 
Fic. 
Fiac. 
FIG. 


FIG. 
FIG. 


Ae 


So eG ca bo 


Caseya irritans sp. nov. (cont.) 
Gnathochilarium. X 70. 


Striaria nasinta sp. nov. 
Antenna. X 70. 
First legs. X 47. 
Gnathochilarium, basal sclerites not shown. X 87. 
Labrum. X 70. 
First segment, caudal aspect. X 39.5. 


Fontaria tuobita sp. nov. 


Gonopods, cephalic aspect. X 47. 
Gonopods, caudo-ventral aspect. X 47. 


PLATE XXXVI. 


FIG. 
Fic. 
FIG. 
FiGc. 
Fic. 


FiG. 
Fic. 


Ore Cobo 


Scytonotus piger sp. nov. 
First segment in outline from dorsal aspect. X 33. 
Carina of fourth segment from the ultimate. X 70. 
Carina of fifth segment from the ultimate. X 70. 


Gnathochilarium (proximal portion not shown). X 47. 


Antenna (proximal article not represented). X 39.5. 


Polydesmus (Kepolydesmus subgen. nov.) anderisus Sp. NOv. 
Fic. 6. Antenna. 


ile 
8. 


First leg of male. X 22. 
Second legs of male. X 22. 


Fic. 9. Head and first segment in outline, dorsal aspect. X 


XVII. 
Polydesmus (Kepolydesmus subgen. nov.) anderisus Sp. Nov. 


PLATE XX 


FIG. 
Fic. 
Fic. 
FIG. 


Fic. 
FIG. 
FIG. 


Po 


“ID On 


Third legs of male, caudal aspect. X 22. 
Gonopods of male, caudal aspect. X 33. 
Gonopods of male, cephalic aspect. X 33. 
Gonopods of male, lateral aspect. X 33. 


Polydesmus (Kepolydesmus) sontus sp. nov. 


Antenna. X 33. 
First segment in outline, dorsal aspect. X DPe. 
First legs, caudal aspect. X 33. 


PLATE XXXVIII. 


FIG. 
FIG. 
FiG. 


Fic. 
FIG. 
FIG. 


ts 
2. 
3. 


Polydesmus (Kepolydesmus) sontus Sp. Nov. (cont.) 
Fifth legs, caudal aspect. X 33. 

Gonopods of male, caudo-ventral aspect. X 47. 
Gonopods of male, lateral aspect. X 47. 


Polydesmus amandus sp. Nov. 


Antenna. X 33. 
Head and first two segments in outline, dorsal aspect. 
Gonopods of male, lateral aspect. XxX 70. 


9») Pw) 


aie 


261 


X 33- 


262 Annals Entomological Society of America [Vol. III, 


PLATE XXXIX. 
Polydesmus amandus sp. nov. (cont.) 


Fic. 1. Gonopods of male, cephalo-ventral aspect. X 47. 


Polydesmus sastianus sp. nov. 

Fic. 2. Head and first two segments in outline, dorsal aspect. X 33. 
Fic. 3. Antenna. X 44, 

Polydesmus socarnius sp. nov. . 
Head and first two segments in outline, dorsal aspect. X 39.5. 
Antenna, X 33. 

Paraiulus zakiwanus sp. nov. 
Fic. 6. Posterior gonopods of male, lateral aspect. X 33. 
Fic. 7. Posterior gonopods of male, cephalo-dorsal aspect. X 33. 


PLATE XL. 


Fic. 
FIG. 


ore 


Paraiulus zakiwanus sp. nov. (cont.) 


Fic. 1. Anterior gonopods of male, cephalic aspect. X 33. 
Fic. 2. Anterior gonopods of male, caudal aspect. X 33. 
Fic. 3. Anterior gonopods of male, lateral aspect. X 33. 
Fic. 4. First legs of male, cephalic aspect. X 18.5. 

Fic. 5. Second legs of male, caudal aspect. X 33. 


Paraiulus tiganus sp. nov. 
Fic. 6. First legs of male, cephalic aspect. X 18.5. 
Fic. 7. Second legs of male, caudal aspect. X 33. 
Fic. 8. Posterior gonopods of male. X 22. 
PLaTE XLI. 
Paraiulus tiganus sp. nov. (cont.) 


Fic. 1. Anterior gonopods of male, cephalic aspect. X 18.5. 
Fic. 2. Anterior gonopods of male, caudal aspect. X 18.5. 
Fic. 3. Anterier gonopods of male, lateral aspect. X 18.5 
Fic. 4. Gnathochilarium of male. X 22. 
Paraiulus furcifer sinampus var. nov. 
Fic. 5. Distal portion of sheath of gonopods in immature male, cephalic 
aspect. X 33. 
Fic. 6. The same, lateral aspect. X 33. 
Fic. 7. Gnathochilarium of same specimen. 
Fic. 8. First and second legs of same specimen, lateral aspect. X 33. 


PLATE XLII. 
Paraiulus furcifer sinampus vat. nov. (cont.) 


Fic. 1. Labrum. X 70. 
Fic. 2. First legs of same specimen, caudo-ventral aspect. X 33. 
Paeromopus lysiopetalinus Karsch. 


Fic. 3. First legs of male, caudal aspect. X 18.5. 
Fic. 4. Posterior gonopods of male, cephalic aspect. X 18.6. 
Fic. 5. Anterior gonopods of male, cephalic aspect. X 18.5. 


PLATE XLIII. 
Paeromopus lysiopetalinus Karsch (cont.) 


Fic. 1. Gnathochilarium. X 18.5. 

Fic. 2. Antenna. X 18.5. 

Fic. 3. Second legs of male, showing penial process. X 18.5. 
Paiteya errans gen. et sp. nov. 

Fic. 4. Gnathochilarium. X 70. 

Fic. 5. Eye. X 87. 

Fic. 6. First legs of male, cephalic aspect. X 70. 

Fic. 7. Antenna. X 47. 


ANNALS ES. A. Vou. III,- PLATE: XXX, 


Ralph V. Chamberlin. 


ANNAIS E. S. A 


Ralph V. Chamberlin. 


Vou. III, Phare XXXI. 


+ 4 


Vou. Tl Pr ATE koxOKIT, 


ANNALS E. S. A. 


Ralph V. Chamberlin. 


ANNALS E. 8. A. VoL. II, PLaTe XXXII. 


Ralph V. Chamberlin. 


ANNALS E. S. A. VOL. TID, PLATE XXXIV. 


Ralph V. Chamberlin. 


ANNAIS E. 8S. A, VoL. III, PLATE XXXV, 


Ralph V. Chamberlin. 


ANNALS E. S. A. 


VoL. III, Pratt XXXVI. 


Ralph V. Chamberlin. 


ANNALS E, S. A. VoL. III, PLrare XXXVII. 


Ralph V. Chamberlin. 


Vou. III, PLatre XXXVIII. 


ANNALS E. S. A. 


Ralph V. Chamberlin. 


Von. ID, PLATS. SSI. 


ANNALS E, 8. A. 


Ralph V. Chamberlin. 


VoL. III, PLATE XL. 


. A. 


ANNALS FE. 


Ralph V. Chamberlin. 


ANNALS E. S, A. 


/ 


| 
| 

| 

| 


A | | 


W) 


1 


Ralph V. Chamberlin. 


VoL. III, PLate XLI, 


ANNALS E. S. A. Vou. III, PLATE XLII. 


Ralph V. Chamberlin. 


ANNALS E. S. A. 


Vou. II, PLate XLII. 


i 


Ralph V. Chamberlin. 


THE STRUCTURE AND METAMORPHOSIS OF THE 
ALIMENTARY CANAL OF THE LARVA OF 
PSYCHODA ALTERNATA SAY.* 


By Lreonarp HaAsEMAN. 


The changes which take place in the digestive epithelium 
of insects after feeding, and in connection with the molting 
periods, have been but little studied, though in the cases 
known the phenomena are of great interest. In some insects 
partial, in others, total regeneration of the mid-intestinal epi- 
thelium has been found to occur after each taking of food. 
In other instances it is the changes associated with molting 
which have attracted attention. It was with a view of study- 
ing the morphological changes in the epithelium at the two 
periods and of correlating the results with those of previous 
workers that this study of the larva of Psychoda alternata 
was undertaken. 

A review of the literature shows that but little has been done 
. along these lines with larval forms. Aside from the work of 
Van Gehuchten (’90 and ’93) on Ptychoptera contaminata, 
Mobutsz (97) on Anthrenus and Folsom and Wells (’06) on 
Collembola, we find only scattered references here and there 
in more general papers on the digestive system. Psychoda 
alternata has been selected for the present work because of the 
ease with which it can be secured and bred throughout the year, 
its wide distribution, its convenient size for work and the fact 
that it is one of the most generalized Diptera. 

This study was carried out in the Entomological Labora- 
tories of Cornell University, and to Professor J. H. Comstock, 
I am greatly indebted for valuable suggestions and criticisms. 
The subject was suggested and the work done under the imme- 
diate supervision of Dr. W. A. Riley, to whom I am especially 
obligated for constant advice and help. 


THE STRUCTURE OF THE ALIMENTARY CANAL. 


The alimentary canal of the larva of Psychoda alternata 
is a straight tube extending from the mouth to the anus, with- 
out folds, except during bodily contraction when a slight one 
is formed in the region of the small intestine. Aside from the 
attachment of strong pharyngeal muscles and the Malpighian 


«Contribution from the Entomological Laboratories of Cornell University. 


277 


278 Annals Entomological Society of America  [Vol. III, 


vessels, the surface of the canal for its whole length is unbroken 
(Fig. 1). From the exterior the division into fore-, mid- and 
hind-intestine is distinctly marked. The short, much con- 
stricted fore-intestine extends back to the third thoracic gan- 
glion. Here, by means of a strong esophageal valve, it connects 
with the very large mid-intestine, which in turn extends back to 
the sixth abdominal segment, where the somewhat constricted 
hind-intestine begins. 

The opening of the mouth is on the ventral surface of the 
anterior part of the head. The clypeus and recurved labrum 
form a rostrum over and in front of it. The labrum bears, on 
either side of its anterior recurved margin, a fan of bristling 
hairs (Fig.3 Ir.) The area between these fans is armed with 
numerous posteriorly directed, strong setae and minute serrate 
chitinous plates. Posterior to this area is a strong V-shaped 
plate and laterad of it, along the margin, a pair of mesally 
directed triangular plates. Behind the V-shaped plate comes 
a group of small chitinous plates, arranged in a circle, and a 
series of minute setae. <A pair of small plates, bearing numer- 
ous minute teeth on their mesal margins, lies laterad of this 
chitinous ring. Farther back is a pair of lateral tufts of setae 
and a second pair of oval plates. 

The mandibles are strongly chitinized with numerous 
lateral setae, a single strong spine, with numerous minute teeth 
on its inner margin, and two long plumes (Fig. 3, md.) On the 
inner margin are two series of strong teeth, directed at an angle 
to each other, and a single posterior thumb-like tooth directed 
cephalad. The hind margin of the mandible is deeply concave 
to receive the anterior part of the maxilla. 

The maxillae are but slightly chitinized, with four short 
stout teeth on their inner margin, a small median plume and a 
fan of long setae extending the full length of the lateral margin. 
The palpi are short and thick with six small papillae and num- 
erous short setae. They are circular in outline and fit into the 
depressions in the hind margins of the mandibles (Fig. 3, mx.) 

The labium is a fleshy lobe bearing numerous fine setae and 
papillae. On the anterior margin are two small tufts of short 
setae and behind these a crescent-shaped chitinous plate (Fig. 3, 
la). The sense papillae found upon the labrum, maxillae, palpi 
and labium are so placed as to come in contact with the food 
as it enters the mouth and therefore are very probably organs 


1910] Alimentary Canal of Psychoda Alternata 279 


of taste. The setae and plumes surrounding the mouth are 
curved, near their tips, and so arranged as to sweep food into 
it when the mouth parts are in motion. 

The mouth opens almost immediately into the pharynx, 

which is somewhat expanded dorsally and supported on its 
~ cephalo-lateral margins by curving chitinous bars (Fig. 4, ph.) 
Strong muscles extend from its cephalic, lateral and dorsal sur- 
faces to their points of attachment on the chitinous surface of 
the head. Posteriorly the pharynx tapers rapidly into the nar- 
row esophagus so that their point of union is hardly discern- 
able. The esophagus is slender, uniform in diameter and ex- 
tends from the subesophageal ganglion to a point opposite the 
third thoracic ganglion. The surface of the esophagus is free 
from any visible suspensory attachments and, to the unaided 
eye, shows rather indistinctly the arrangement of the circular 
muscles. 

Lying alongside the esophagus, above the first thoracic 
ganglion, are the salivary glands. They are quite prominent, 
each about the size of one lobe of the brain, and strongly 
reniform. From the ventral end of each gland, a minute duct 
extends downward and forward to a point below the subeso- 
phageal ganglion, where the two from opposite sides unite to 
form the main salivary duct. This duct runs forward below the 
alimentary canal to the mouth, where, as we shall see later, 
it opens between the hypopharynx and labium (Fig. 4, d.) 

Closely enclosing the pharynx are the commissures connect- 
ing the brain with the subesophageal ganglion (Fig. 4, com.) 
These commissures extend forward and slightly downward on 
either side until the level of the pharynx is reached when each 
turns rather abruptly backward and downward to the sub- 
espohageal ganglion. 

Above the second and third thoracic gangha, the narrow 
esophagus is telescoped into the mid-intestine forming the 
espohageal valve. The diameter of the tube in the region of 
the valve is much greater than that of the esophagus but less 
than that of the mid-intestine farther back (Fig. 1, es. v.) The 
mid-intestine normally is a smooth tube tapering slightly 
anteriorly from its middle and posteriorly from a point near its 
union with the hind-intestine. When it is gorged with food, 
however, its diameter may be more uniform throughout. In 
surface view the cellular structure of the mid-intestine stands. 
out very distinctly even with low power magnification. 


280 Annals Entomological Society of America  [Vol. III, 


The alimentary canal and especially the mid-intestine is 
closely and firmly enclosed in columns of adipose tissue. Four 
longitudinal lateral strands extend the full length of the larva 
with shorter strands here and there. The adipose tissue and 
Malpighian vessels form a perfect cushion about the canal and 
in this way support it in the central part of the body cavity. 
When the alimentary canal is dissected out the adipose tissue 
adheres to it and severs its connection with the body wall, 
and the respiratory and nervous systems. In living larvae 
numerous fine tracheal branches, from the pair of dorsal longi- 
tudinal tracheae, are seen to ramify the fat bodies and send 
finer branches to the alimentary canal. The nervous connec- 
tions with the adipose tissue and alimentary canal are much 
finer and more difficult to discern than are the tracheal connec- 
tions. Delicate cephalad directed fibers from the large dorsal 
nerve, which extends caudad from the eighth abdominal gan- 
glion, are seen to attach to the ileum just behind the Malpighian 
vessels corresponding to suspensory nerves described by Com- 
stock and Kellogg, ’95, for Corydalis. These fine nervous 
connections from below with the tracheal connections from 
above assist in supporting the alimentary canal. 

From surface view the hind intestine is distinctly divided 
into ileum and colon, while the rectum is less easily made out 
(Fig. 1). The ileum extends from the point of attachmentgof 
the Malpighian vessels, opposite the anterior end of the sixth 
abdominal segment, to the middle of the seventh segment where 
it joins the much expanded colon. It is so much narrower than 
the mid-intestine on the one hand and the colon on the other 
that, in surface view, it stands out in sharp contrast with them. ~ 
The circular muscles of the ileum are strongly developed and 
appear as a distinct transverse surface banding. The walls 
are very thin and elastic. 

The diameter of the anterior end of the colon is twice that 
of the ileum, while posteriorly it tapers to the rectum where it 
is about equal that of the ileum. The walls of the colon are 
very thick and but slightly elastic. The circular muscles are 
strongly developed and are imbedded in pits in the epithelium. 

The rectum is quite short, extending from a point opposite 
the anterior end of the eighth abdominal segment, where the 
canal bends ventrad, to the anus. It is narrower than the colon 
and the circular muscles are not imbedded in pits. Exteriorly 


1910} Alimentary Canal of Psychoda Alternata 281 


it is developed into four distinct papillae which surround the 
ematise. (ig? cls ac sp.) 

The Malpighian vessels join the alimentary canal at the 
union of the mid-intestine and ileum. They are five in number, 
two on the latero-ventral side, two on the dorso-lateral side, 
and the fifth on the dorsal surface of the canal (Fig. 1, m. t.) 
The vessels all have an expanded reservoir-like basal portion, 
which in living larvae is snow-white, but 1n alcoholic specimens 
fades out (Fig. 27, c.) The two latero-ventral vessels extend 
far forward to a point opposite the middle of the mid-intestine 
where they make an abrupt turn backward for a short distance. 
The two dorso-lateral ones start forward but in the middle of 
their expanded basal reservoirs they turn abruptly upon them- 
selves and their distal ends become closely associated with the 
reproductive organs opposite the middle of the colon. The 
dorsal tubule follows the same course, its tip lying free above 
the middle of the colon. Beyond the expanded basal reservoir, 
the vessels are much constricted and heavily charged with a 
reddish-brown pigment in life. The vessels are slightly motile. 

Wheeler (93) claims that the primitive number of Mal- 
pighian vessels in insects is six. Where there are less than six, 
he claims that the reduction is due to a loss or fusion of certain 
ones, while where there are more than six, he explains it as being 
due to a division of some of them. However this may be, it is 
well known that the vessels are usually paired, that is, there is 
an even number of them. It is rare to find an odd number of 
Malpighian vessels in insects. Wheeler reports seven for a 
Neuropterous larva, probably Chauliodes, and the same number 
is found in the larva of Corydalis cornuta. Five Malpighian 
vessels have been found in Culex and Psychoda. I find this 
same number in Pericoma. In Psychoda alternata there seems 
to be no difference in form or structure between the single dor- 
sal vessel and the other four paired vessels. They all join the 
alimentary canal singly and are about equi-distant. The same 
number and arrangement are found in the pupal and adult 
stages. 

Surrounding the anal opening are four prominent fleshy - 
lobes. These are direct prolongations from the rectum and have 
a respiratory function. Numerous fine tracheae extend down- 
ward and forward to these, from a constriction in the large 
longitudinal tracheae just caudad of the vent. These lobes are 


282 Annals Entomological Society of America  [Vol. III, 


slightly protractil, becoming especially prominent in larvae 
that have been submerged for a considerable length of time. 
They are homologous with the anal tracheal gills of other 
insects. Vaney ('02) gives a figure showing the connection 
of the tracheae with these lobes in a Psychoda larva where the 
tracheae are far more prominent than in Psychoda alternata. 

The chitinous intima of the mouth cavity is quite thick and 
continuous with that of the mouth parts and surface chitin 
of the head (Fig. 5,in.) The epithelium is thin except beneath 
the minute serrate plates on the anterior surface of the labrum 
where the cells are very long and slender. In longitudinal sec- 
tions the hypopharynx and epipharynx are very prominent. 
The epipharynx is closely associated with the back part of the 
labrum and bears setae and taste papillae (Fig. 5, eph.) The 
hypopharynx lies immediately aboye the labium. Its intima is 
very thick and dark and near its base are strong setae protrud- 
ing into the mouth of the pharynx (Fig. 5, hy.) 

From the back part of the mouth the food passes directly 
into a large chamber, the anterior part of the pharynx (Fig. 
2, ph.) The dorso-cephalic wall of this chamber is very thin 
and flexible to accommodate varying quantities of food. On 
either side of this chamber near its anterior end are two strong 
dorsally directed curving chitinous bars. These bars from 
opposite sides almost meet above and below and serve as a sup- 
port for the flexible walls and at the same time bear a series of 
long yellow ventro-mesally directed chitinous setae which give 
them a comb-like appearance (Fig. 6, ph. b.) These comb-like 
structures across the entrance to the alimentary canal function 
as strainers in keeping out large foreign bodies. The pharyngeal 
chamber is always more or less filled with food and probably 
functions as a crop in which the food is mixed with the salivary 
secretions, by the action of the circular and pharyngeal muscles 
and these comb-like chitinous structures. The epithelium in 
this part of the pharynx is quite thin, while the chitinous intima 
is thick. The circular muscles in this region are by no means so 
strongly developed as in the caudal part of the pharynx. 

The posterior portion of the pharynx is constricted so that 
its diameter is about equal that of the esophagus. In transverse 
section it appears pear-shaped, the broader portion being ven- 
trad (Fig. 9.) It is provided with very powerful circular muscles 
which lie so closely together as to form almost a continuous 


1910} Alimentary Canal of Psychoda Alternata 283 


layer. The epithelium consists of very large cells, with large 
oval nuclei, whose nucleoli and chromatin material stain ~ 
intensely with haematoxylin. The cavity of the pharynx in this 
region has the shape of a Roman I, a narrow dorso-ventral slit 
and transverse dorsal and ventral arms (Fig. 9, 1.) The intima 
of the dorso-ventral slit is very thick and the underlying epi- 
thelial cells are very large, cubical and with distinct cell walls. 
The intima lining the dorsal and ventral arms of the cavity is 
much thinner and the underlying epithelium is equally reduced 
in thickness. 

The powerful circular muscles serve as constrictors for clos- 
ing the lumen of the pharynx, while four sets of strong phar- 
yngeal muscles serve as relaxers. On either side of the median 
portion of the pharynx are two horizontal rows of strong muscles, 
each row consisting of four separate muscles and a single very 
large anterior one. .These muscles extend laterad and attach 
at the sides of the head. Their inner ends pass between the 
circular muscles and are attached to the large cubical epithelial 
cells by means of numerous radiating, zig-zag strands (Fig. 
8, ph. m.) From the dorsal surface of the pharynx at the tips 
of the comb-like bars is a series of four strong muscles which 
extend dorsad to the surface of the head, joining it at its caudal 
margin. Low down on the dorso-cephalic margin of the 
expanded pouch-like portion of the pharynx are a number of 
strong muscles which extend dorso-cephalad to the surface of 
the head. The caudal portion of the pharynx with its heavy 
chitinous intima and strong constrictor and retractor muscles, 
must serve as a mill for crushing the food as it is passed on to 
the esophagus. 

The pharynx ends and the esophagus begins beneath the 
caudal end of the brain (Fig. 4, es.) The circular muscles of 
the anterior portion of the esophagus completely clothe it, but 
the individual muscles are less strongly developed than those 
of the pharynx or those of the posterior part of the esophagus. 
Longitudinal muscles have not been found in any of the prepara- 
tions. The epithelium is very thin. The cells forming it are 
without distinct walls and their small nuclei are arranged in 
definite rows along the dorsal, lateral and ventral surfaces of 
the esophagus. The large lateral cells of the pharynx give way 
in the esophagus to small cells which protrude into the lumen to 
form a longitudinal ridge on either side (Fig. 10, ep.) The 


284 Annals Entomological Society of America [Vol. III, 


slight mid-dorsal and ventral protrusions. of the epithelium in 
the pharynx, persist in the anterior part of the esophagus, 
forming a slight dorsal and a ventral longitudinal ridge. In the 
median portion of the esophagus these ridges almost disappear 
while in the caudal region there are two dorsal and two ventral 
ridges (Fig. 11, ep.) These dorsal and ventral ridges are simply 
thickenings of the cells opposite the nuclei while the lateral ones 
are formed by an elevation of the cells from the muscular coat 
and a protrusion of them half way in to the center of the lumen. 
The chitinous intima of the esophagus is well developed al- 
though much less strongly than in the pharynx. It becomes 
much folded and is usually severed from the epithelium by fix- 
ation. On the inner surface of the infolded intima of the esoph- 
ageal valve, there are to be seen, with high magnification, a 
number of strong spine-like setae, but these are absent in the 
anterior portion of the esophagus (Fig. 17). They are so di- 
rected as to resist the passage of food from the mid-intestine 
back into the esophagus. 

The salivary glands are quite prominent, each about the 
size of one of the lobes of the brain (Fig. 12). They are strongly 
reniform so that their free ends lie almost directly above their 
proximal ends. When the glands are not secreting their lumina 
may be very large, but during active secretion they are nearly 
obliterated. The cells are very large and protrude into the 
lumen so that in sections there are deep grooves between them. 
They have a coarse granular and vacuolate appearance, and are 
usually set off by distinct walls, except during active secretion 
when these may become obliterated. The nucleus is very large 
and centrally located, with deeply staining chromatin material 
and a very large nucleolus. The chitinous intima is quite 
thick but obscured by the granular nature of the underlying 
cells and the secretions in the lumen. The limiting basement 
membrane is very delicate, but stands out distinctly. 

From the ventral anterior end of the glands, ducts pass for- 
ward and downward to a point beneath the sub-esophageal 
ganglia where the two join to form the main salivary duct. 
This main duct passes forward to the mouth where it opens on 
the surface of the labium between it and the hypopharynx. 
The intima of this duct is quite delicate posteriorly while the 
epithelium is more prominent, but near its anterior end both the 
intima and epithelium are prominent. Just before the duct 


1910} Alimentary Canal of Psychoda Alternata 285 


reaches the labium it is provided with a distinct press. The 
lower surface of the duct is attached to a strong chitinous process 
from the intima of the labium while a pair of very strong 
muscles is attached to the dorsal surface (Fig. 6, pr.) These two 
muscles extend laterad and slightly dorsad to their point of 
attachment to a prong of the tentorium. By their contrac- 
tion the dorsal wall of the duct is drawn away from the ventral 
and the salivary secretions are permitted to escape into the 
mouth. 

The circular muscles of the esophagus become greatly 
enlarged for a short distance in front of the esophageal valve 
and for some distance back into it (Figs. 14, 15 and 16, sp.) 
They thus function as a sphincter. The valve is strongly 
developed, the length of the inflexed portion of the esophagus 
being about equal to the diameter of the valve. The epi- 
thelium and intima of the inflexed portion are unmodified while 
those of the reflexed portion are much thinner and in longitudi- 
nal section appear as two heavy lines with here and there a 
nucleus in the epithelium (Figs. 14 and 15.) The reflexed por- 
tion extends a short distance cephalad to the anterior end of 
the mid-intestine and so forms a second, a caudally directed, 
loop. Circular muscles are absent from the caudal portion 
of the inflexed esophagus and from all of the reflexed portion. 
Between the inflexed and reflexed portions there is a large blood 
space, filled with the granular blood plasma and scattered blood 
corpuscles and traversed by numerous delicate fiber-like out- 
growths of the reflexed epithelial cells (Fig. 16, s.) A number 
of these fibers and anteriorly protruding longitudinal muscles 
from the mid-intestine, meet the esophagus opposite the an- 
terior end of the sphincter. 

The point of union of the fore- and mid- intestine is plainly 
set off by the striking contrast between the epithelium of the 
two regions. The thin epithelium, of the reflexed part of the 
esophagus, gives way to a girdle of large deeply-staining cells, 
which extend about half the length of the esophageal valve. 
In longitudinal section from six to twelve large cells are seen to 
comprise this group on either side (Fig. 15, gl.) The anterior 
ones are short, polygonal and lie transversely, but farther back 
they become strongly club-shaped and are directed meso- 
caudad. The outer ends of the cells are closely appressed while 
their inner ends are expanded greatly and form a smooth secre- 


286 Annals Entomological Society of America [Vol. III, 


tory surface. The nuclei are large, with large nucleoli and 
coarse, deeply-staining chromatin granules, and lie in the 
expanded inner portion of the cells. These cells are glandular 
in nature. They stain far more intensely with haematoxylin 
than any other cells of the alimentary canal. They are granular 
and finely vacuolate toward their inner margins, where a thin 
layer of granular substance is to be seen (Fig. 18, p.m.) The 
inner margin of this granular layer hardens to form a delicate 
elastic peritrophic membrane which is continually fed back into 
the mid- and hind-intestine to envelope the food material. 
This membrane may be seen as a delicate envelope surrounding 
the excrement as it is thrown off. In some preparations the 
formation of the peritrophic membrane from the granular 
secretion can be seen very distinctly with the 1-12 immersion 
objective. 

There are several views as to the origin of ite peritrophic 
membrane in insects. By Pagenstecher (’64), it was thought 
to be a product of the salivary glands. Frenzel (’85) thought 
it was a product of the mid-intestinal epithelium. Simpson 
(00) in an unpublished paper described it as a product of the 
. mucus cells in Clisiocampa. Schneider (’87) considered it a 
chitinous prolongation from the esophagus. Van Gehuchten 
(90) and Cuenot (’95) claimed that a ring of cells just cephalad 
of the mid-intestine produces it. In the larva of Psychoda alter- 
nata there can be no doubt as to the origin of this membrane. 
The girdle of cells at the anterior margin of the mid-intestine 
secretes it. 

Behind and laterad to this girdle of cells comes a second 
girdle of glandular cells. In longitudinal sections seldom more 
than two cells are to be seen (Figs. 13, 15 and 46, c. gl.) They 
are very large, among the largest found in the alimentary canal, 
coarsely vacuolate, and have a straw-yellow color when stained 
with iron-haematoxylin. The nuclei are large, with rather small 
nucleoli and fine deeply staining chromatin granules. The cells 
pour their straw yellow secretion, which has a uniform finely 
granular consistency, into a pouch which opens widely into the 
lumen and extends latero-cephalad between the two rings of 
cells as seen in longitudinal sections. This pouch-like structure 
is simply a deep invagination on the inner surface of the gland. 

Caecae are usually present in the larvae of Diptera, espec- 
ially in the more specialized forms. They may be long, as in 


1910] Alimentary Canal of Psychoda Alternata 287 


the larvae of the blow-fly (Calliphora), but are more often short 
and inconspicuous. The number of caecae may be two (Cteno- 
phora), four (Tipula, Simulium, etc.) or many as Miall and 
Hammond (’00) figure for the larva of the Harlequin fly. Vaney 
(02) says that -in Anthomyia there are four caecae, in Chir- 
onomus many, while in Stratiomyia and Tanypus they may be 
absent. In Tanypus he finds and figures a group of very large 
vacuolate cells with ectal diverticula which he claims function 
as caecae. This group of cells is similarly located, and the cells 
themselves, except for the ectal diverticula, closely agree with 
those described above for Psychoda. In Psychoda I am con- 
vinced that this group of cells is the rudimentary caecal glands. 
I find it also present in Pericoma. Caecae are said to be entirely 
absent in the larvae of Lepidoptera, which in this respect would 
seem to show a relationship between the Psychodidae and the 
Lepidoptera. There are not wanting a number of other char- 
acters which indicate such an affinity. 

Behind the caecal gland comes a group of cells very similar 
to those which secrete the peritrophic membrane (Figs. 14 and 
15). The cells stain quite deeply, are elongate and extend 
meso-caudad. They have numerous vacuoles which usually 
contain large spherical concretions (Fig. 19, con.) The nuclei 
are large with distinct nucleoli and intensely staining chromatin 
granules. The posterior extent of this group is marked by a 
few greatly elongated cells which project into the lumen (Fig. 
14, ep.) It might be supposed that this group of cells assisted the 
anterior one in the production of the peritrophic membrane 
but from repeated examination of my specimens no signs of 
this have been found. 

Caudad of this region comes a long stretch of fairly uniform 
cells. They,are very large and cubical or polygonal (Fig. 2, ep.) 
The ones in front and behind are cubical, while those in the 
median region are longer than deep. Scattered among the large 
cells are to be found small slender ones (Fig. 21, ep.) The 
cells of this region are stained but slightly with 1ron-haema- 
toxylin, are finely vacuolate, and possess a narrow but distinct 
striate border. The nucleus is large and the nucleolus and 
chromatin are sharply differentiated. These cells are secretory 
but do not take on the greatly elongated condition found in 
those farther back during most active discharge of secretion. 


288 Annals Entomological Society of America  [Vol. III, 


These large cells grade off posteriorly into a region of tall 
slender secretory cells (Fig. 2, ep.), comprising the posterior 
third of the mid-intestinal epithelium. These slender cells are 
finely vacuolate and have a very strongly developed striate 
border (Fig. 24, st. b.) The nuclei are small, usually centrally 
located, with deeply staining nucleoli and chromatin material. 
During active secretion, groups of these cells increase greatly in 
heighth and protrude into the lumen, giving to the inner margin 
of the epithelium a decidedly uneven surface (Fig. 23, ep.) 
In this respect, this region stands out in sharp contrast with the 
anterior region of large cells, where their inner margins form 
almost a straight line. Here and there among these slender 
cells as well as the larger anterior ones are to be found huge, 
coarsely vacuolate cells which protrude into the lumen. These 
have been described as mucus cells (Fig. 46, m. c.) 

The striate border is more or less developed throughout the 
whole length of the secretory part of the mid-intestine, but its 
greatest development is to be found in the posterior region 
during active secretion and just before molting. Some of the 
earlier workers considered the striate border as a chitinous inti- 
ma perforated by many fine canals. Later work goes to show, 
however, that it is simply a ciliate border, consisting of non- 
vibratile cilia, except perhaps in case of Chironomus larvae 
(Vignon '99). In Psychoda alternata it grades gradually from 
the vaculate tips of the cells into a clear homogenous border 
formed by closely appressed fine cilia. So gradual is the grada- 
tion and so intimate is the relation between the cell body and 
the border that I am inclined to agree with Van Gehuchten 
(90) that the cilia are continuations of the cells rather than 
secretions deposited by them upon their inner margin. 

The basement membrane is quite distinct throughout the 
whole length of the mid-intestine (Figs. 22 and 24, b. m.) 
In places it is considerably thickened. With perfect fixation 
the basement membrane remains closely connected with the 
epithelium. It bears on its outer surface, or more often in 
slight depressions of it, the numerous strands of circular muscles. 

In the region of the esophageal valve, and for a short dis- 
tance caudad, the circular muscles are weak and the strands far 
apart (Fig. 20, c.m.) But beginning with the region of very 
large epithelial cells and extending back along it a short distance 
the bands of circular muscles become much broader and more 


1910] Alimentary Canal of Psychoda Alternata 289 


regularly placed, with intervening spaces equal to the breadth 
of the bands. These broad bands give way to narrower and 
more widely separated ones, which continue back to the region 
of tall slender epithelial cells. Sections of these strands show 
their breadth to scarcely exceed the diameter of the nuclei of 
the underlying epithelial cells. At the point where the large 
epithelial cells give way to the tall slender ones, the strands 
of circular muscles are greatly crowded together, having the , 
appearance of a slight sphincter (Fig. 22,c.m.) This crowding 
of the circular muscles covers but a short area and posteriorly 
gives way to a greater spacing of them, which extends back to 
the base of the Malpighian vessels. 

The longitudinal muscles are not strongly developed in the 
mid-intestine. They consist of from ten to twelve narrow, flat 
strands, which fit closely to the surface of the circular muscles 
@hic 2 and: 23, 1. mt)» Opposite the point, where the circular 
muscles are crowded together, these longitudinal strands are 
greatly thickened, almost circular in cross section and stand out 
from the circular bands more distinctly. 

Just at the union of the mid- and hind- intestine are the 
five Malpighian vessels. In cellular structure they much more 
closely resemble the epithelium of the mid-intestine than that 
of the hind-intestine, but their glandular function will account 
for this, however. From embryological studies it has been 
shown that these vessels are outgrowths from the hind-intestine. 
At the point of attachment of these vessels with the alimentary 
canal they have a short, compact and slightly constricted 
region (Fig. 26, m. t.) Here the cells are large and alternately 
from opposite sides extend into the lumen of the vessels giving 
it a zig-zag course. The nuclei of these cells are oval with small 
nucleoli and fine chromatin granules. This constricted portion 
gives way to the much expanded, thin-walled reservoir. The 
epithelium of this region is very thin, with its inner surface del- 
icately fibrillate and with valve-like projections here and there 
extending into the lumen (Fig. 27, c.) Beyond the reservoir 
comes the true glandular portion of the vessel which is much 
constricted and, in life, 1s charged with a reddish brown pig- 
ment (Fig. 27, a and b). Here the lumen is greatly reduced, 
being almost closed by the finely granular and vacuolate cells. 
These cells are very long, as can be seen from longitudinal sec- 
tions, and as shown by some transverse sections, each cell 


290 Annals Entomological Society of America  [Vol. III, 


almost completely girdles the tubule, and forces the lumen 
toward one side, but more often the lumen is centrally located, 
the cell completely encircling the tubule. The basement mem- 
brane is strongly developed throughout the whole length of the 
tubule. The chitinous intima is slightly developed in the distal 
glandular portion, but seems to be absent in the basal portion. 

The change from the mid-intestine to the ileum of the hind- 
intestine 1s a most striking one. The large secretory cells of 
the mid-intestine give way to the very small cells of the ileum 
(Fig. 26 and 28, ep.) These first cells are so very small that 
with the 1-12 immersion they appear as mere thickenings of 
the epithelium. The nuclei are very small, deep-staining, and 
without prominent nucleoli. A distinct, thin chitinous intima 
is present. This thin epithelium extends caudad to a point 
slightly before the middle of the ileum, where it develops a dis- 
tinct fold, which extends into the lumen to form an ileal valve 
(Fig. 2, 29 and 31, il. v.) From this point to the union of ileum 
and colon, the epithelial cells are much larger, with enormous 
nuclei, in which the small nucleoli and coarse chromatin gran- 
ules stain deeply. The ileum is well supplied with muscles. 
The delicate circular muscles of the mid-intestine are replaced 
in the ileum by very strong bands which completely clothe it. 
For a short distance, immediately behind the Malpighian ves- 
sels, the circular muscles are absent (Figs. 28-29.) From the 
point where the circular muscles begin, back to the ileal valve, 
there are two layers of them. The inner layer of delicate bands 
serves as a sphincter for the ileal valve, while the outer layer 
is continuous throughout the hind-intestine (Figs. 29 and 30). 
From the ileal valve back to the colon the individual bands of 
the circular muscles are much stronger and more oval than those 
of the anterior part of the ileum. The longitudinal muscles are 
only slightly developed in the hind-intestine. From longitu- 
dinal sections it would appear that, at the point where the first 
circular muscles appear, the longitudinal muscles divide to 
form an inner and an outer layer, but from cross sections only 
outer longitudinal muscles appear in front of the ileal valve. 
Three strong, and two or three smaller strands appear clustered 
together along either side of the canal. From the ileal valve 
to the colon these same lateral clusters of outer longitudinal 
muscles appear as well as scattered inner strands between the 
circular muscles and epithelium (Fig. 32, 1. m.) 


1910] Alimentary Canal of Psychoda Alternata 291 - 


Posteriorly the constricted ileum projects into the expanded 
colon so as to form a slight valve (Figs. 2 and 33, v.) Just 
in front of this valve is a strong sphincter composed of five or 
six strong strands of circular muscles. The colon is slightly 
shorter than the ileum but its diameter is twice as great. Its 
epithelium is very thick, consisting of large cubical cells, of unt- 
form finely granular nature, with distinctly striate inner curv- 
ing margins which project into the lumen (Figs. 35 and 36, ep.) 
The cell walls are seldom distinct to the base.. The nuclei are 
very large, and oval, with large nucleoli and fine chromatin 
granules. The basement membrane is very delicate and diffi- 
cult of detection in most preparations. The circular muscles 
are strongly developed and are so fitted into pockets in the epi- 
thelium that their outer surface is just even with that of the 
epithelium (Fig. 36, c. m.) Longitudinal muscles have not 
been observed either in longitudinal or cross sections of the 
colon as well as the rectum. Sections of close-fitting nerve 
fibers were at first mistaken for longitudinal muscles. The 
chitinous intima is but slightly developed and closely adheres 
to the epithelium. Along the surface of the intima and espec- 
ually down in the grooves between the cells is a layer of a finely 
granular deposit which stains blue with haematoxylin. This 
deposit is present in all of my preparations, which together with 
the enormous development of the epithelium, would suggest 
some association with absorption and I am of the opinion that 
in this larva much of the absorption of digested food takes 
place here. . 

The circular muscles of the antefior end of the rectum are 
much more strongly developed than are those of the colon 
(Fig. 37, c. m.) They lie on the surface of the epithelium and 
form a continuous layer. Farther caudad the individual bands 
decrease in strength rapidly. These circular muscles form a 
sort of sphincter about the rectum. The nuclei of the circular 
muscles are very large and are surrounded by undifferentiated 
protoplasm. The basement membrane is more strongly devel- 
oped than in the colon. The epithelium is greatly reduced 
especially in the caudal portion where the cells are long and thin 
except opposite the nuclei, where they are thickened so as to 
protrude slightly into the narrow lumen. In the anterior region 
of the rectum the epithelium is thrown into six longitudinal 
folds (Fig. 37, ep.) The epithelium of the rectum is continuous 


292 Annals Entomological Society of America  [Vol. III, 


with that of the four slightly eversible anal papillae, where it 
becomes much thicker and the cells more vacuolate. The 
chitinous intima of the rectum is much heavier than that of the 
colon and its thickness increases toward the anus where it is 
continuous with the chitin of the anal papillae (Fig. 2, a. p.) 


MORPHOLOGICAL CHANGES IN THE EPITHELIUM OF THE 
MID-INTESTINE. 

In insects the changes, which the epithelium of the mid- 
intestine undergoes, are associated either with the pouring out of 
digestive secretions at the time of feeding or with the larval 
molting periods. These changes are marked by total or partial 
epithelial degeneration and regeneration. In the present work, 
special efforts have been made to get as complete series of 
larvae as possible in order to show the changes accompanying 
feeding and molting. By means of carefully timed starvation 
and feeding experiments, all desired stages associated with the 
feeding period were easily obtained. But it was much more 
difficult to get all the stages at the molting period, since the 
morphological changes in the mid-intestine are largely completed 
before the skin is cast and there is practically no forewarning 
as to just when the transformation begins. Only by means of 
a large series of specimens taken during the inactive period 
preceding and following the casting of the skin was it possible 
to get.all the desired stages of transformation. 


Changes Accompanying Feeding. 

The extent of degeneration and regeneration of the epithe- 
lium of the mid-intestine, at the time of feeding depends upon 
the type of secretion found in the epithelial cells. - Where the 
holocrine type of secretion prevails, partial or even total degen- 
eration and renewal of the epithelium will follow each full 
feeding, while with the merocrine type only slight or no renewal 
will be found since the epithelial cells are not destroyed by the 
discharge of secretions. 

The nymphs of dragonflies and adult water beetles, in which 
may be found the holocrine type of secretion, are excellent 
examples for the study of epithelial regeneration following ~ 
feeding. Needham ('97) found that, in case of dragonfly 
nymphs, after prolonged fasts of two months, the cells became 
so gorged with digestive secretions that the epithelium became 
twice the height of the normal and when food was taken in 


1910] Alimentary Canal of Psychoda Alternata. 293 


practically all of the cells were destroyed by the process of 
excretion and a complete new epithelium formed from scattered 
nidi or regenerating centers. Frenzel (85), Vangel (’86), 
Bizzozzero (’93) and Rengel (’97) described even more striking 
cases of epithelial degeneration and regeneration in adult water 
beetles of the genera Hydrophilus, Hydrobius, Hydrous and 
Cybister. These beetles like the nymphs of dragonflies, feed 
intermittently, being able to fast for periods when food is not 
at hand and then when food is abundant, devouring great 
quantities of it. In these bettles the entire epithelium with its 
basal chitinous membrane 1s described as being cast off after 
each full feeding. The new epithelium is quickly developed 
from highly specialized regenerating centers or nidi, which pro- 
trude through the muscular coat of the canal, to form distinct 
diverticula. 

The merocrine type of secretion is found in the mid-intestinal 
epithelium of those insects which feed continuously and which 
require a continuous supply of digestive fluids. In these forms 
the cells are always active, the extent of the secreting activity 
depending upon the extent of feeding. But since the cells are 
not destroyed by secretion and excretion there is no occasion 
for any regeneration in connection with the intake of food and 
digestion. This type of epithelial secretion was described by 
Van Gehuchten (90 and ’93) for Ptychoptera contaminata, 
there being no epithelial regeneration associated with digestion. 
Folsom and Wells (’06) describe this as the prevailing type of 
secretion in the mid-intestine of Collembola, there being no 
regeneration at the time of feeding. 

Psychoda alternata in its larval state is not an intermittent 
feeder but is most at home at the mouth of open sewers, in 
cultures of decaying horse manure or other vegetable ferments, 
where it finds abundance of food always present. An occasional 
plankton form may be found mingled with the food but the 
bulk of the intestinal contents consists of bits of decayed 
vegetation. The larva is a voracious feeder and extremely 
active, scarcely refraining from feeding long enough to molt. 
The canal always has a great quantity of food material in it, 
even at the molting periods. From careful study of the epithe- 
lium of the mid-intestine of this form, I find the merocrine type 
of secretion prevailing and an absence of epithelial regeneration 
associated with feeding. 


294 Annals Entomological Society of America  [Vol. III, 


In hopes of being able to study to better advantage the type 
of secretion and of better understanding the relation of its 
discharge to the intake of food, I undertook a series of feeding 
experiments. Larvae were placed in small glass vessels with 
clean tap-water and kept there without food for periods 
varying from one to twelve days. The larger larvae invariably 
pupated before the experiments could be completed, so it was 
necessary to use larvae which were less than half grown. Larvae 
that had fasted for varying periods up to twelve days were 
killed and sectioned for determining what changes, if any, the 
mid-intestinal epithelium underwent during the fasts. Like- 
wise larvae starved for two, three, four, seven and twelve days 
were removed to vessels containing abundance of food, and then 
after feeding for periods varying from half an hour up to seven 
hours, they were killed and sectioned for studying the epithelial 
changes accompanying feeding. 

From a study of the preparations made from these larvae 
it was found that where the fast was prolonged more than four 
days, the mid-intestinal epithelium became so reduced and 
weakened that it failed to recover even though the larva sur- 
vived the fast and fed freely for a time (Fig. 40). Simpson (’01, 
in an unpublished paper) found a similar degeneration of the 
epithelial cells of the mid-intestine of Clisiocampa larvae that 
had fasted for five days. When the fast is prolonged four days 
the cells take on a distinct vacuolate appearance associated 
with a heavy discharge of secretion (Fig. 39). After six or 
seven days this vacuolate condition has disappeared and the 
cells become greatly reduced in height. They almost cease to 
excrete digestive fluids and the striate border is then more 
conspicuous. Aside from a continued reduction in the height of 
the epithelium, and a greater shriveling of the cells, no further 
marked changes are brought about, the cells finally dying. At 
no time, even after a twelve day fast, is the mid-intestine 
completely free from food material. As a consequence there 
is a constant stimulus to the excretion of the digestive secretion 
and as the fast is prolonged the finely granular secretion is 
poured out and soon comes to fill most of the space between the 
reduced epithelium and the food content of the canal (Fig. 39, 
d. sec.) In place of the secretion being stored up in the cells 
as in the nymphs of dragonflies, it is continually poured into the 
lumen. This process soon drains the cells of their vitality, 


1910] Alimentary Canal of Psychoda Alternata 295 


since but little real nourishment can be absorbed from the 
already thoroughly digested content of the canal. 

With larvae that had fasted less than four days no marked 
changes were found in the mid-intestinal epithelium. The food 
content of the canal is sufficient, apparently, to sustain the 
larva and to stimulate the cells of the epithelium to normal 
activities for this period so that when a fresh supply of food is 
taken in normal digestion is continued (Fig. 41). From the 
results of these expreiments, which comprised upwards of fifty 
separate lots of larvae, it was found necessary to turn to the 
preparations of the normal feeding larvae, for the determination 
of the type of secretion and the relation of its discharge to the 
intake of food. 

For some time, the changes in the mid-intestinal epithelium 
just preceding the casting of the larval skin were mistaken for 
signs of an excessive production and storing of secretions, but 
when series of larvae preparing to molt were studied the mistake 
was detected. In this larva there are four sources from which 
the digestive secretions of the mid-intestine are derived. 

As previously noted, the gastric caecae, usually present in 
the higher Diptera, are replaced in Psychoda alternata by a 
greatly reduced caecal gland which girdles the mid-intestine 
in the region of the esophageal valve. The large cells of this 
gland produce a yellowish digestive secretion similar to that of 
the gastric caecae of other insects. During most active secre- 
tion of these caecal glands, the cells become swollen and finely 
vacuolate (Figs. 18 and 47, c. gl.) The pouch, into which the 
secretion is poured, becomes so completely filled that its neck 
appears to flare, leaving a wide exit of the secretion into the 
lumen. During less active secretion and excretion the cells 
become somewhat reduced, the vacuolation less distinct and the 
pouch less distended with secretion or even almost empty. This 
is one source of digestive secretion. 

A second source is the long stretch of very large cells caudad 
of the esophageal valve (Fig. 2, ep.) These cells during active 
secretion and excretion give off a finely granular secretion. 
This secretion is not given off as globules or droplets but simply 
oozes through the thin striate border of the large finely vacuolate 
cells and comes to fill all the space between the epithelium and 
the peritrophic membrane (Fig. 42). This is perhaps the most 
abundant supply of digestive secretion found in the mid-intes- 
tine of this form. 


296 Annals Entomological Society of America  [Vol. III, 


Just caudad of the point where the region of large cells gives 
way to that of the tall slender ones, comes a group of cells which 
during active secretion becomes greatly elongate and protrude 
into the lumen. These cells are active in throwing off small 
spherical bodies of their protoplasm by the pinching off of their 
tips, which float out into the lumen to dissolve and form a 
third supply of digestive secretion (Figs. 43 and 44, sec. p.) 

The fourth source of digestive secretion is the less elongate 
epithelial cells farther caudad. These cells are more granular 
and vacuolate next to the striate border and throw off distinct 
globules which collect along the margin of the striate border 
(Fig. 45, sec. g.) These globules seem to burst and give out a 
secretion similar to the finely granular material of the anterior 
part of the mid-intestine. The striate border becomes less 
distinct where the cells are actively secreting and excreting but 
is never completely obliterated. 

The mucous cells of Leidig (’57), Frenzel (’82 and ’85), 
Korotneff (’85), Beauregard (’86), Fausseck (’87), Mingazzini 
(’89), Balbiani (90), and Simpson (’00 in unpublished paper) 
are present in this larva. They are scattered among the large 
secretory cells of the anterior region as well as among the tall 
slender cells of the posterior region. As compared with the 
surrounding cells they are truly monsters (Fig. 46, m.c.) They 
are connected with the basement membrane by a constricted 
pedicle while the distal end is enormously expanded and pro- 
trudes into the lumen. They are coarsely vacuolate throughout 
but especially so toward the inner end. The nucleus is scarcely 
larger than those of adjoining cells, and does not differ from 
them in the arrangement of nucleolus and chromatin material. 
Some writers have attributed to these cells the function of 
secreting the peritrophic membrane, others claim that their 
secretion serves as a lubricant. In the larva of Psychoda alter- 
nata these cells certainly have nothing to do with the secretion 
of the peritrophic membrane. The fact that the secretion of 
these cells resembles that from surrounding cells microscopically, 
that their greatest activity corresponds with that of the sur- 
rounding cells and the fact that the food, enveloped in the peri- 
trophic membrane, is continually floated, so to speak, in diges- 
tive fluids, cause me to hesitate to accept the view that the secre- 
tion of these cells is merely a lubricant. I am inclined to be- 
lieve that here we have simply specialized digestive cells, 


1910] Alimentary Canal of Psychoda Alternata 297 


The slender cells, of the region immediately caudad of the 
caecal glands, have a coarse granular and finely vacuolate 
structure. In some preparations a slight discharge of granular 
secretions is to be seen although this is more frequently absent. 
The striate border is only slightly developed. Between the 
middle and distal margin of these cells are to be found vacuoles 
of various sizes which inclose dark brown concretions (Figs. 
19 and 47, con.) These concretions are spherical, almost com- 
pletely filling the vacuoles and very dense. I have not found 
them in any of the other epithelial cells of the mid-intestine 
of this larva. They closely: resemble those found by Folsom 
and Wells in the mid-intestinal epithelium of Collembola, and are 
probably waste products which are secreted and held by these 
cells until the molting period when they are discharged along 
with the cells. If my interpretation of these concretions is 
correct these cells have a definite excretory function. 

Passing forward to the anterior region of the mid-intestine 
we find the cells which produce the peritrophic membrane. 
These cells are especially active during feeding. They appear 
long, club-shaped and become finely vacuolate on their inner 
margin. From them is discharged a finely granular secretion, 
which forms a narrow inner border to the cells (Figs. 18 and 47, 
p. m.) It is difficult to tell just where the cell-body ends and 
the layer of secretion begins. On the inner surface of this 
border a portion of it hardens to form the thin uniform peri- 
trophic membrane which closely resembles the chitinous intima 
of the esophagus. From the reaction of these cells to stain it 
is evident that they are secreting continuously; but the under- 
lying border of secretion is especially prominent at certain 
times. The course of most active secretion seems to pass along 
the canal like a wave, following the entrance of a new supply 
of food somewhat as found in dragonfly nymphs. By the time 
the discharge of secretions in the posterior part of the mid- 
intestine has reached a maximum it has begun to decrease in 
the anterior portion. This is the case with the cells producing 
the peritrophic membrane as well as those of the caecal glands. 


Changes at the Molting Periods. 

It is well known that, with each larval molt, the entire outer 
cuticula, as well as the intima of the fore- and hind-intestine and 
the. tracheal system, are cast. Sharp (’95) maintains that the 
facilitation of growth by the casting of the chitinous coat is 


298 Annals Entomological Society of America [Vol. III, 


only of secondary importance. The discharge of the chitinous 
coat is merely an outward sign that internal changes have been 
going on. Eisig (’87) suggests that since chitin is a nitrogenous 
substance, perhaps the casting of the chitinous coat is a means 
of disposing of nitrogenous wastes, to which Sharp adds that 
perhaps the same is true of carbonaceous wastes. If the casting 
of the exuviae has an excretory function associated with it and 
if the epithelium of the mid-intestine has an ectodermal origin, 
as believed by some writers, then it will be easier to explain 
certain morphological changes which occur in the mid-intestinal 
epithelium at the time of molting. The striking morphological 
changes undergone by insects at the time of pupation and emerg- 
ing of the adult, are well known and have been given much 
attention by workers, but there are similar although perhaps 
less intensive changes at each larval molt, which have been 
almost entirely overlooked. We shall now consider the changés 
in the mid-intestinal epithelium at the time of molting. 

Folsom and Wells’ (06) work on epithelial degeneration, 
regeneration and secretion in the mid-intestine of Collembola 
deals primarily with the changes in the digestive epithelium at 
the molting periods. The Collembola are among our most 
primitive forms of insects, lacking Malpighian vessels and devel- 
oping without obvious metamorphosis. With this type of 
development we should expect to find equivalent morphological 
changes at the different molting periods. Conditions found in 
the study of a form of this generalized type will not necessarily 
be like those in higher forms but are suggestive for one making a 
study of any more specialized form. 

The normal epithelium of the mid-intestine of Collembola 
was found to be composed of very large granular and finely 
vacuolate cells. At the approach of an ecdysis the cells begin 
elongating and the inner half becomes more vacuolate. As 
this vacuolation continues, any concretions and gregarine 
parasites present in the cells and about half of the nuclei migrate 
from the more dense portions of the cells into the inner, more 
vacuolate region. Then comes a cutting off of the inner half 
of the epithelium. During these changes cell walls are obliter- 
ated. A new striate border forms along the line of constriction 
and the inner portion of the epithelium is cast into the lumen 
where it is digested. To replace the nuclei lost by the sloughing 
off, those remaining undergo mitotic division, cell boundaries 


1910] Alimentary Canal of Psychoda Alternata 299 


reappear and the normal epithelium is again restored. Since 
the forms studied lacked Malpighian vessels, Folsom & Wells 
concluded that this periodical discharge of half of the epithelium 
with concretions, inclusions and parasites had an excretory 
function similar to that of the Malpighian vessels in other 
insects. The short inactive period preceding each molt made 
it possible to get any desired stage of transformation. Here 
we find only partial epithelial regeneration at each molting 
period. 

We shall now consider a more specialized type of insect, 
one with complete metamorphosis, and see how the changes in 
the mid-intestinal epithelium at the time of the larval molts 
compare with those in Collembola at the molting periods. 
Mobitisz (’97) studied the changes in the mid-intestinal epithe- 
lium of Anthrenus during the larval molts and found that the 
same changes appeared then as at pupation, only less intensive. 
In the study of his preparations, which included larvae in var- 
ious stages of transformation, he found a large mass of material 
in the lumen of the mid-intesine which he first mistook for 
parasites. Further studies revealed to him that this inclusion 
was simply the remains of the sloughed off epithelium. He was 
able to find various stages in the throwing off of the old cells 
from that in which, by muscular contraction and development 
of the new epithelium, the old with its basement membrane was 
separated from the muscular coat, to that in which it was cast 
into the lumen and digested leaving its residue enveloped in the 
resistant basement membrane. Mobtsz takes exception to 
Rengel’s work on Tenebrio molitor (’96) where he describes 
the nidi as being cast off with the old epithelium and basement 
membrane and later making their way through the basement 
membrane and regaining an attachment to the muscular coat. 
Rengel must certainly have had faulty preparations. Mobisz 
found the nidi retaining their attachment with the muscular 
coat and concluded that either, the cells of the nidi being 
embryological, no basement membrane was produced beneath 
them, or if produced it was dissolved so that it would slip off 
over them when it was cast off with the old epithelium. By 
multiplication the cells of the nidi quickly produce the layer of 
new epithelium. In fact these first changes are produced with 
such rapidity that Mobtsz failed to catch them in any of his 
specimens. The new epithelium soon developes its basement 


300 Annals Entomological Society of America [Vol. III, 


membrane and striate border and in time the nidi take on their 
normal condition and the new epithelium is mature. Here we 
find the epithelium cast off bodily and not merely in part. In 
conclusion Mobtsz maintains that: . 

(1) At each larval molt there is a total regeneration of the 
epithelium of the mid-intestine. 

(2) The transformation of the mature larva to the pupa, 
in the holometabola, is simply an intensified larval molt and we 
may expect the same morphological changes, although to vary- 
ing degrees, perhaps, at these periods. 

(3) He believes that there is not only total regeneration 
of the mid-intestinal epithelium but also that there are greater 
or less changes in the epithelium of the fore- and hind-intestine 
and in the muscular and adipose tissues of the larva. 

The larva of Psychoda alternata on preparing to molt, 
ceases feeding for a short time and to a marked degree dis- 
charges the food content from the alimentary canal. Then 
comes a short period of inactivity followed by pronounced 
peristalic contractions. In.the thoracic region a strong ven- 
tral flexure occurs and then peristaltic waves of contraction pass 
forward from the posterior end of the body. This may be 
repeated a number of times before the chitin bursts on the dor- 
sum of the first two thoracic segments and the larva crawls out. 
As soon as the larva is free it becomes quite active and begins 
to feed. These outward signs of transformation are simply the 
culmination of the metamorphosis. The internal changes are 
practically completed before the larval skin is cast. 

Shortly before the time of molting the mid-intestinal epi- 
thelium begins to undergo striking changes. The cells begin 
to elongate and protrude into the lumen. Their distal halves 
become distinctly vacuolate (Fig. 48). There is to be noted 
in the nuclei themselves some change. The chromatin usually 
disappears or comes to closely envelope the nucleolus. At this 
stage the cytoplasm and enclosed secretions of the cells stain 
intensely with iron haematoxylin which can be removed only 
by prolonged differentiation. 

This stage in the metamorphosis was first observed in a 
larva that had fasted for three days and was thought to be due 
to the storing up of digestive secretions, but when other speci- 
mens from the same lot were examined and no signs of such 
development found and when larvae, that were known to be 


1910] Alimentary Canal of Psychoda Alternata S01 


ready to molt, were examined and the same identical condi- 
tions found, the theory of stored up secretions was discarded. 
This condition of degeneration of the mid-intestinal epithelium 
has also been found in some specimens collected at random for 
general structural work, they having been caught in the act 
of transformation. 

A later stage shows these epithelial cells becoming more 
broken up by large vacuoles. The entire distal portion of the 
cells becomes vacuolate with strands of cytoplasm surrounding 
the vacuoles (Fig. 49). The vacuoles at first circular in sec- 
tion become irregular in shape as their size increases, while 
the nuclei are found moving out into the distal more vacuolate 
portion of the epithelium. All signs of cell boundaries are lost 
and the epithelium resembles a mass of ragged cytoplasm with 
nuclei scattered about through it. In the cytoplasm small 
round concretions are usually present in great abundance, es- 
pecially in the more vacuolate portion (Figs. 48 and 49, con.) 
Gregarine parasites are seldom found within the epithelium. 
The nuclei degenerate unevenly. Some near the basement 
membrane may retain a more healthy appearance with chroma- 
tine granules and large nucleolus, while those beginning to mi- 
grate lose their chromatin. At this stage of degeneration, there 
appears in the lumen of the mid-intestine, surrounding the peri- 
trophic membrane and enveloped food contents, a great mass 
of finely granular and deeply staining secretion which is evident- 
ly the product of the degenerating cells of the epithelium. 
This deposit seems to be confined largely to the posterior half 
of the mid-intestine where the epithelium consists of the tall 
slender cells. In this bluish-staining deposit are to be seen 
scattered remains of degenerating nuclei, especially nucleoli, 
which seem to be especially resistant. Scattered through this 
deposit are also to be found deeply staining bodies, probably 
concretions, and a quantity of uniform yellowish secretions, 
evidently a product of the degenerating cells. 

This degeneration of the mid-intestinal epithelium begins in 
the posterior portion where the cells are long and slender and 
as metamorphosis advances, the process of degeneration may 
be traced forward to the esophageal valve (Figs. 50 and 51). 
When the large cells of the anterior half of the mid-intestine 
have become strongly vacuolate, those of the posterior part 
have almost completely broken down and their contents cast 


302 Annals Entomological Society of America  [Vol. III, 


into the lumen. The method, by which the midintestinal 
epithelium, in this form, is cast off, in no wise resembles that 
described for Hydrophilus, after feeding, where the epithelium 
with its basal chitinous layer is cast off bodily; nor is it so com- 
plete and regular as that described by Mobusz for Anthrenus. 
It more nearly resembles the degeneration of the mid-intestinal 
epithelium of dragonfly nymphs after feeding. The individual 
cells themselves break down and are sloughed off rather than 
the epithelium in mass being cast into the lumen. 

The extent to which this degeneration takes place is not 
always easy to determine. It can be followed to where practi- 
cally all of the old nuclei have migrated out into the lumen and 
degenerated (Fig. 51). And where practically all of the cell 
content is dissolved, although for a time, at least, a quantity 
of the cytoplasm remains in contact with the basement mem- 
brane, which is not cast off. I am inclined to believe that this 
is as far as the degeneration of the mid-intestinal epithelium 
proceeds in this form. The old adhering cytoplasm is appar- 
ently absorbed by the newly developing epithelium. 

Between the epithelium of the mid-intestine and its under- 
lying basement membrane are to be found numerous very small 
regenerating cells from which the new epithelium is developed 
(Figs. 48, 49, 50, 51, r. c.) These small cells are extremely 
difficult of detection except at the time of transformation. Cell 
division takes place in these embryonic cells while their diameter 
is yet scarcely one-fifth that of the nucleus of mature epithe- 
lial cells, so that it is extermely difficult to make out the type 
of division, but from a study of my preparations it would seem 
that here we have direct cell division (Fig. 54). They may be 
collected in small groups or in pairs or scattered singly along 
the basement membrane. The first structure that can be defi- 
litely made out is the deeply staining nucleus, but as the degen- 
ration of the old epithelium advances, a definite cell form can be 
distinguished. The cytoplasm at the beginning is a mere border 
about the nucleus, but as development advances it rapidly 
increases in bulk and comes to form a distinct cell body about the 
nucleus. By the time the old epithelium is nearly all sloughed 
off these new cells have come to protrude up into the adhering 
cytoplasm of the old cells, which they seem to absorb in part 
(Ris 761 "re 


1910] Alimentary Canal of Psychoda Alternata 303 


These conditions of degeneration and regeneration of the 
mid-intestinal epithelium are found in larvae previous to the 
casting of the larval skin. Changes at this point take place 
with very great rapidity so that it is easy to see how former 
workers came to overlook, almost completely, this degeneration 
and regeneration of the mid-intestinal epithelium at the larval 
molting periods. 

In larvae, taken immediately after the casting of the skin, 
the new epithelium of the mid-intestine is already developed. 
At first the cells are low and stain deeply with iron haematoxy- 
lin (Fig. 52, ep.). The nucleoli are large and stain deeply, 
while the chromatin material is not distinctly made out. The 
cytoplasm is coarsely granular and has the appearance of being 
Simavued= next to the lumen. ~ The striate borderus developed 
early and appears even more distinct than in later stages when 
the cells have begun to secrete (Figs. 52 and 58, st. b.) 

The contents of the mid-intestine, not cast off before meta- 
morphosis began, is still found to be enclosed in the resistant 
peritrophic membrane and occupies the central portion of the 
lumen. Between this central mass and the new epithelium, 
there is still to be found a great quantity of the finely granular 
yellowish product from the degenerate epithelium. Occasion- 
ally the sloughed off intima of the esophagus is found coiled 
about in the mid-intestine, but the degenerate epithelium 
of the mid-intestine is so completely dissolved that only the 
slightest traces of it are to be found after the skin has been cast. 

In this way the epithelium of the entire mid-intestine, 
including the caecal glands and the group of cells which produce 
‘the peritrophic membrane, is renovated at each larval molt. 
As soon as the metamorphosis is completed and the skin cast 
off the larva resumes activity and begins to feed again. The new 
cells of the mid-intestinal epithelium at once respond to the 
demand made upon them for secretions and the larva enters 
upon the next stage of its development. 


CONCLUSIONS. 


The alimentary canal of the larva of Psychoda alternata is a 
generalized one in conformity with the habits of the larva. 
Both the fore- and hind-intestine are relatively short while 
the mid-intestine makes up at least two-thirds of the entire 
length of the canal. 


304 Annals Entomological Society of America  [Vol. III, 


Longitudinal muscles are absent in the fore-intestine and 
in the colon and rectum. 

Gastric caecae are replaced by a caecal gland, which girdles 
the median portion of the esophageal valve. 

There are five Malpighian vessels of similar form and struc- 
ture, two extending cephalad and three caudad. 

The rectum is prolonged into four slightly eversible papillae, 
which are homologous with the anal tracheal gills of Culex and 
related forms. 

The epithelium of the mid-intestine is divided into five 
distinct regions; an anterior one giving rise to the peritrophic 
membrane, a caecal gland, a small area of long glandular cells, 
a long stretch of large polygonal cells, and the posterior region 
of tall slender cells. The so-called mucus cells are abundant. 
There is no regeneration of the epithelium of the mid-intestine 
at the time of feeding. During a fast, secretions are not stored 
up in the epithelial cells but are continually poured out and if 
the fast be prolonged the cells may completely break down. 
The peritrophic membrane is secreted by a definite group of 
cells at the anterior end of the mid-intestine. 

At each larval molt the epithelium of the mid-intestine 
is completely renewed. The old cells are not cast off as a whole, 
but the degeneration begins next to the lumen and continues 
until the whole cell is broken down. This degeneration begins 
in the posterior region of the mid-intestine and proceeds forward. 

Regeneration is from scattered embryonic cells which lie 
in the angles at the base of the epithelial cells. . The regenerated 
epithelium of the mid-intestine is mature by the time the larval 
skin is cast off. 


1910} Alimentary Canal of Psychoda Alternata 30: 


Ot 


BIBLIOGRAPHY. 


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Cuenot, L., 99. La région absorbante dans l’intestin de la Blatta. Critique d’un 
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Deegener, P., 02. Ammerkung zum Bau der Regenerationscrypten des Mittel- 
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87. Beitrage zur Histologie des Darmkanals der Insecten. Z. wiss. Zool. 
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93. Contribution a l’etude du mecanisme de 1’Excretion Cellulaire. Cellule, 
IX, pp. 95-116, one plate. 
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Leger, L., and Duboscq, O., 02. Sur la régénération épithéliale dans 1l’intestin 
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Miall, L. C., 93. | Dicranota; a carnivorous Tipulid larva, Tr. ent. Soc. Lon- 
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X, pp. 190-208, pl. V. 


LIST OF ABREVIATIONS. 


a. p., anal papillae. 

b. c., blood corpuscles. 

b. m., basement membrane. 

br., brain. 

c., concretion bearing cells. 

c. gl., caecal gland. 

‘¢. m., circular muscles. 

col., colon. 

com., commissure. 

con., concretion. 

d., salivary duct. 

d. c., degenerating cells. 

d. sec., digestive secretion. 

ep., epithelium. 

eph., epipharynx. 

es., esophagus. 

es. v., esophageal valve. 

f., food. 

gl., gland which secretes the peritro- 
phic membrane. 

hy., hypopharynx. 

il., ileum. 

il. v., ileal valve. 

in., intima. 

1., lumen. 

la., labium. 

1. m., longitudinal muscles. 


Ir., labrum. 
m., mouth. 
m. c., mucus cell. 


‘“md., mandible. 


m. int., mid-intestine. 

m. t., Malpighian tubule. 
mx., maxilla. 

mx. p., maxillary palpus. 

n., nucleus. 

nl., nucleolus. 

n. m., nuclear membrane. 
p., caecal gland pouch. 

ph., pharynx. 

ph. b., pharyngeal bars. 

ph. m., pharyngeal muscles. 
pr., press of salivary duct. 
T. c., regenerating cells. 
rec., rectum. 

s., blood space. 

sec. c., secretory cells. 

sec. p., secretory proliferations. 
s. g., salivary gland. 

sp., sphincter. 

st. b., striate border. 

vac., vacuole. 

v., valve between ileum and colon. 


2p , =) 


1910] Alimentary Canal of Psychoda Alternata 307 


EXPLANATION OF PLATES. 


PEATE xell LVe 


Alimentary canal in surface view. 
Longitudinal section of alimentary canal. 
Mouth parts. 
Anterior end of larva with alimentary canal and nervous system in situ. 
Longitudinal section through the anterior portion of the head. 
Transverse section through the anterior end of the pharynx. Plane of 
section sloping slightly dorso-caudad. 
Transverse section through the median portion of the pharynx. Plane 

of section sloping slightly dorso- caudad. 

8. Portion of the transverse section through the pharynx in the region of the 
large lateral pharyngeal muscles. 

9. Transverse section through the caudal region of the pharynx. 

10. Transverse section through the anterior region of the esophagus. 

11. Transverse section through the anterior end of the esophageal valve. 
Plane of section sloping slightly dorso-caudad. 


es Cred) es 


PLATE XLV. 


12. Longitudinal section of salivary gland. 

13.. Transverse section through the middle of the esophageal valve. Plane 
of section sloping slightly cephalad to the right. 

14. Longitudinal section of esophageal valve. 

15. A portion of longitudinal section of esophageal valve. 

16. Longitudinal section of cephalic end of esophageal valve Seon a por- 
tion of the blood space. 

17. Longitudinal section through the tip of the infolded portion of the Sone 
agus, showing chitinous spines. 

18. A portion of a longitudinal section through the region of the caecal gland. 

19. Cells with enclosed concretions, from the region immediately caudad of 
the caecal gland. 

20. Longitudinal section through the region at which the concretion-bearing 
cells give way to the large polygonal digestive cells. 


IPVATE el Vele 


21. Transverse section of mid- intestine through the region of large polygonal 
digestive cells. 

22. Longitudinal section through the region at which the polygonal cells 
give way to the tall slender ones. 

23. Transverse section of mid-intestine through the region of tall slender 
cells. 

24. Longitudinal section through the region of tall cells. 

25. Transverse section through the region at which the Malpighian vessels 
join the alimentary canal. 

26. Longitudinal section through the same region. 

27. Transverse sections of Malpighian vessels. (a and b) Through the pig- 
mented secretory portions, (c) through the basal reservoir. 

28. Transverse section of the ileum immediately caudad of the Malpighian 
vessels, showing absence of circular and longitudinal muscles. 

29. Longitudinal section of the anterior portion of the ileum. 


308 Annals Entomological Society of America  [Vol. III, 


PLATE XLVII. 


30. Transverse section through the ileum just cephalad of the ileal valve. 
31. Longitudinal section through the ileal valve. 

32. Transverse section through ileum, caudad of the ileal valve. 

33. Longitudinal section through the valve between the ileum and colon. 
34. Transverse section through the valve between the ileum and colon. 

35. Transverse section through the colon. 

36. Portion of longitudinal section through the colon. 

37. Transverse section through the rectum. 

38. Longitudinal section through the rectum and anal papillae. 


XLVIII. 


Epithelial secretion and regeneration (with 1-12 immersion.) 


39. Epithelial cells of the mid-intestine of a larva after a four days’ fast. 

40. (a) Epithelial cells of the mid-intestine of a larva after a seven days’ 
fast. (b) Same after feeding half an hour. 

41. Epithelial cells of mid-intestine of a larva that had fasted two days 
then fed for half an hour. 

42. A single large polygonal digestive cell from the anterior portion of the 
mid-intestine. 

43. Tall slender epithelial cells discharging digestive proliferations. 

44. Same, from region slightly cephalad of (438). 

45. Epithelial cells from the posterior region of the mid-intestine discharging 
globules of digestive secretion. 

46. Mucus cells from posterior region of mid-intestines. 

47. Longitudinal section through the caecal gland. 


48. Epithelial cells from the anterior portion of the mid-intestine shortly _ 


before molting, showing the first stage of epithelial degeneration. 

49. Epithelial cells showing a later stage of degeneration. 

50. Epithelial cells from anterior portion of mid-intestine showing the process. 
of degeneration well under way. 

51. Epithelial cells from the same larva, but from the region of tall slender 
cells with degeneration almost complete. 

52. Newly regenerated epithelium of the mid-intestine of a larva that had 
just emerged from the larval skin. 

53. Epithelium of the mid-intestine of a larva that had molted less than a day 
previous to killing. 

54. Embryonic cells in various stages of development. 


ANNALS E. S. A. 


L. Haseman. 


m.int.~ {> 


Vou. III, PLATE XLIV. 


Vou. III, Phare XLV. 


A. 


S. 


ANNALS E, 


Haseman. 


L. 


ANNALS E. S. A. VOL. III, PLATE XLVI. 


L. Haseman. 


ANNALS E. S. A, 


Vou. ITI, Plate XLVII. 


NM 


(oA EE 
i 


cou HEA 9 


‘ 


rec..-] 


38 


J. Haseman. 


ANNALS E. S. A. VoL. III, PLATE XLVIII. 


- p.m. 


L. Haseman. 


A KEY TO THE GENERA OF THE SUBFAMILY 
APHIDINAE AND NOTES ON SYNONOMY. 


By H. F. Witson, 
Oregon Agricultural College. 


The following paper is not as complete as the author would 
like to have it as type species of four recently formed genera are 
not available. Several attempts have been made to secure the 
desired species without success and as there does not seem to 
be much chance of securing them in the near future copies of 
the original keys are included in this paper. All of them were 
made by Del Guercio and three of them are given as a division 
of the so called genus A phis. 

The first three are given in Redia (vol. 4, pp. 190-192, 1907) 
with the following key 

1. Femine partenoge ae attire ed alate sprovviste di codetta 0 con 

questa affatto rudimentdle. 

Gen. ANURAPHIS Del Guercio 
(Sp. tip. Aphis pyri Koch). 
Femmine attere ed alate con codetta distinta................ 2 

2. Codetta costantamente piu corta dei sifont. 

Gen. Apuis Linne 
(Sp. tip. Aphis sambuci L.) 
Codetta constantamente eguale, o piu lunga dei sifont...... 3. 

Codetta piu lunga dei sifont. 


— 


Gen. Urapuis Del Guercio 
(Sp. tip. Aphis genistae Kalt.) 
Codetta della stessa lunghezza dei sifoni, 
Gen. MicrostpHon Del Guercio 
(Sp. tip. Aphis tormentillae Pass.) 


The fourth was published two years later (Rivista di patolo- 
gia X Vegetale anno IV Num. 11. Agosto 1909. pp. 4-5.) This 
species is given as the dividing line between Anuraphis Del 
Guercio and Mysus Passerini. 


A. Codetta verricuforme, rudimentale o nulla. 

I. Femmine attere con tubercoli frontali nulli, in ogni caso evident- 
emente piu corti della meta della fronte; antenne corte col 
primo articolo subeguale al secondo, 

Gen. ANURAPHIS Del Guercio. 

II. Femine attere con tubercoli frontali piu. lunghi della meta del 
margine della fronte fra essi compressa: primo articolo del 
antenne, che sono lunghe, due volte piu lungo del secondo. 

Gen. MaccutaTieLLta Del Guercio. 

AA. Codetta hen distinta 


Gen. Myzus Passerint. 
314 


1910} Key to the Genera of Aphidinae 315 


In working up synonomy from a standpoint of literature 
there seems to be a great difference of opinion. However since 
an International Code has been established there seems to be 
but one correct plan and if followed in all cases the troubles 
of the systematist must soon disappear. One of the rules of 
the code in substance states that if a number of species belonging 
to distinct different genera are arranged under a certain genus 
without a type being set by the maker of the genus then the 
type may be any one of the original species. If all the species 
but one are removed without a type being set then the last 
species is the type. 

In 1758 Linnaeus formed the genus Aphis without setting 
a type. In this genus he included a large number of species 
which have been placed in various different genera, and Aphis 
sambuct 1s now generally taken as the type of the original 
genus. In 1801 Lamarck™ set Aphis ulmi as the type and A phis 
sambuct was not set until a year later by Latreille.t 

Aphis ulmi L. is the species generally known as Tetraneura 
ulmi DeGeer. ‘This species was first described and figured by 
Reaumer previous to the 10th Edition of Linnaeus. Linnaeus 
and DeGeer give the same citation, so the species does not 
belong to DeGeer. Schizoneura ulmi Linn. is the species 
designated by himself as Chermes ulmt. 

The author of this paper will not at this time attempt to 
say that any of the above names should be changed, but it 
seems that a decision of some kind as to the disposal of the 
above should be made. 

In making the key type species of each genera were used 
and more or less detailed descriptions taken of the external 
characters. In several instances wide variations were noticed 
among the species included in certain genera and some were 
separated to form new genera. 

In studying closely related genera the development of exter- 
nal characters may be placed in five divisions.. 1. The antennae 
and spur. 2. The antennal tubercles. 3. The development 
of the nectaries. 4. The development of the cauda. 5..The 
development of the wing venation. In a group of insects as 
pliable as the present one, any one or two of these characters 


* Systeme des animaux sans vertebres, ou Tab. gen. classes, des orders et generes animaux 
A. Paris, an. 1X, p. 300. 1801. 
+ Histore Naturelle Gen. et par. Des Crus. et Ins. Tome III, p. 264. 1802. 


316 Annals Entomological Society of America  [Vol. III, 


may be either under or over developed and it is necessary to 
place the genera according to the greatest development. Of 
all the characters which show this variation the wings show 
what may be true of all the characters. 

Along these lines the subfamily has been divided into three 
tribes as follows: 


A. Nectaries nearly as long as the body and with long hairs...... Trichosiphini. 
Nectaries without hairs and not more than half as long as the body By 

B. Nectaries variable in length but at least one fourth the length of the body. 
Antennae as long as the body or longer and set on distinct antennal tuber- 

cles. The antennae of Phorodon are shorter but the tubercles are greatly 
GEVELOP EG Fs re ie 6 ai§ wing vis aes Weed ripe ee ee AG _. Macrosiphini. 
Nectaries variable in length; antennae usually shorter than the body and not 

set on distinct antennal tubercles. When tubercles are present antennae 
and mectaries, are much reduced: <2... 0. «syne em ne ek ele ae Aphidini. 


Tribe TRICHOSIPHINI 


This tribe is limited to two genera both of which are found 
in the Asiatic Islands. The nectaries are quite remarkable in 
that they are covered with long hairs. In other respects they 
are not so different from the genera of the other two tribes. 


I. Antennae not longer than the body and the cubital vein twice forked. 
Trichosiphum. 
II. Antennae longer than the body and the.cubital vein with but one fork. 
Greenidea. 


Gen. Trichosiphum Pergande 
type anonae Pergande. 


Antennae about as long as the body, six, segmented, and fixed 
on large tubercles; spur of sixth segment shorter than third 
segment; first segment gibbous. Antennal tubercles slightly 
tapering but not gibbous being oblique to the forehead which is 
nearly flat. Body elongate and narrow; stigma of wing long 
and slender; cubitus twice forked. Nectaries as long as the 
body cylindrical, and standing out at right angles to the body. 
Cauda short and triangular; anal plate short, half moon shaped. 
Antennae, legs, and nectaries as well as the body covered with 
long bristle like hairs. 


A pterous female:—Body short, robust, with antennae one 
half the length of the body; six segmented, with small antennal 
tubercles. Nectaries one fourth the length of the body and 
vasiform. Cauda short and broadly rounded; anal plate broad 
and slightly emarginated at the center. Antennae, nectaries, 
legs and body exceedingly hairy. 


1910} Key to the Genera of Aphidinae Sty 


Gen. Greenidea Schouteden. 
type S. artocarpt Westwood. 


Antennae longer than the body and six segmented; spur of 
sixth shorter than the third segment; first segment slightly 
gibbous on the inner side. Antennal tubercles large and taper- 
ing, front of head wide and flat. Body long and slender, wings 
long, venation variable but usually the cubitus is but once 
forked. Nectaries almost as long as the body, slender and 
nearly cylindrical. Cauda short, broader than long and rounded 
at the tip; base slightly constricted; anal plate rounded. Body 
and appendages exceedingly hairy. ; 


Tribe MACROSIPHINI 


The author has divided this group from the following in 
order that the key may not contain too many conflicts. The 
division is made between species having distinct antennal 
tubercles and those having none or at the most indistinct 
tubercles. However should a certain species have distinct 
antennal tubercles with the other characters wanting then it 
would have to go in the next tribe. 


I. Antennal tubercles tapering and very large, not gibbous on the inner side. 
IIe 
— Antennal tubercles gibbous or toothed on the inner side.............. me lull: 
Il. Nectaries nearly half as long as the body, cylindrical and curved.. . Illinoia. 
— Nectaries at least one fourth the length of the body and vasiform. 


Nectarosiphon. 

III. Antennal tubercles large, and as long on the outer side as on the inner; 

upper inner angle more or less gibbous. Nectaries variable.......... IV. 

= Antennal tubercles prominent but not large and the inner side longer than 
ihe outer orelse-outer side isi but, avlines eee ee ee WAL 

IV. Nectaries tapering longer than the cauda which is sickle shaped. Wings 
regular with twice forked cubitussis een eee Macrosiphum. 

== Nectaries and cauda variable, wing venation irregular but very striking. 
Veins either wanting or else combineds.cean taken atk At Mammen tele We 


V. Antennal tubercles with sharp upper inner angle, cauda shorter than the 
nectaries and tapering. Stigmal and cubital vein almost meet in a broad 
dark band, giving the wing the appearance of having a closed triangular 
ell eee ee ea eae nS AM ges A Mr eee ee ey amie oie Idiopterus. 

== Antennal tubercles with small round tubercle at upper inner angle. Necta- 
ries tapering and slightly longer than tapering cauda. Wing venation 
variable but usually with the cubitus once forked, and the hind wing with 

Joie, eisGyeated (EMT) Vie fbn Wer Seo ansbondobundacusnocuauscdouee Microparsus. 

= Antennal tubercle with small rounded tubercle at upper inner angle. Nec- 
taries constricted at the middle and tip. Wing venation variable but 

usually the stigmal and cubital veins are partly joined and form a distinct 


Glosed.cell with founsides i.e ssc cat oc nee ee eee Pentalonia. 

VI. Antennal tubercles short but not wedge shaped. Cauda large and long. 
VII. 

— Antennal tubercles wedge shaped with the inner side formed into a tubercle 
ori tooth.) “Cauda shontzne py speek oe eee ene ee tre non WAUUIES 


VII. Antennal tubercles slightly gibbous. Antennae much longer than the 
body, nectaries strongly clavate. Wings with cubitus twice forked. 
Amphorophora 


318 Annals Entomological Society of America  [Vol. III, 


Antennal tubercles as in the above genus. Nectaries tapering, wings with 

the cubital vein having but a single fork..................... Toxoptera 

VIII. Antennal tubercles with a strong tooth on the inner side. Nectaries 
almost cylindrical but tapering slightly. Cauda short and tapering. 

Phorodon 

— Antennal tubercles with a prominent blunt projection forming the inner 

side. Nectaries slightly clavate, cauda tapering and with a knobbed tip. 

Rhopalosiphum 

Antennal tubercles with a distinct but not prominent blunt projection 

forming the inner side of the tubercle (more prominent in the apterous 

forms). Nectaries cylindrical and slightly curved at tip. Cauda short 

and tapering, being almost triangularinform...................... Myzus 


Gen. IIlinoia new genus 
type M. liriondri Monell 
Antennae longer than the body, six segmented, and situated 
on prominent tubercles; spur of sixth segment as long or longer 
than the third; first segment strongly gibbous on inner side. 
Antennal tubercles and inner side not gibbous but tapering. 
Body elongate, wings long and medium stout; cubitus twice 
forked. Nectaries almost as long as the abdomen, broad at 
the base and tapering, being strongly constricted for a short 
distance back of the tip. Tip curved outwardly. Cauda 
ensiform, about one third the length of the abdomen and 
constricted in the middle. Anal plate broad and bluntly 
angled. 
Gen. Nectarosiphon Schouteden 
type M. rubicola Oestlund. 


Antennae longer than the body, six segmented, and placed 
on fairly prominent tubercles; spur of sixth segment shorter 
than the third segment, first joint slightly gibbous on inner side. 
Front of head flat and narrow, Body elongate, wings large, 
nectaries about one fourth the length of the body with distal 
half dilated. Cauda one half the length of the nectaries and 
tapering. Anal plate triangular. 


Gen. Macrosiphum Passerini 
type A. rosae Linn. 


Antennae six segmented, longer than the body and situated 
on fairly prominent tubercles; spur of sixth segment longer than 
third, first segment about as long as broad. Antennal tubercles 
slightly gibbous on the inner side. Inner surfaces at right 
angles to the head; forehead flat. Wings long, cubitus with two 
forks. Nectaries about one fourth the length of the body and 
tapering. Cauda as long as the nectaries and ensiform. 


1910] Key to the Genera of A phidinae 319 


Gen. Idiopterus Davis 
type nephrelepidis Davis. 

Antennae about one and a half times as long as the body and 
placed on prominent tubercles; segments one and two large, 
the remaining segments long and slender, spur of sixth segment 
longer than the third. Antennal tubercles much larger than 
the second segment and with the upper inner side sharply 
angled. Body elongated, wings with the stigmal vein deeply 
rounded almost touching the discoidal and both lying in a 
broad dusky band. This gives the wing the appearance of 
having a closed triangular cell. Nectaries about one fifth the 
length of the body, straight and cylindrical. Cauda nearly as 
long as the nectaries and tapering. 


Gen. Microparsus Patch 
type variabilis Patch. 

Antennae longer than the body and placed on prominent 
tubercles, spur of sixth segment longer than the third. First 
segment large and gibbous on the inner side. Antennal tuber- 
cles close together, head flat in front and merging into the 
tubercles. Wings long, cross veins heavy and cubitus usually 
with a single fork; stigma long and slender, hind wing with but 
a single cross vein. Nectaries about one fifth the length of the 
body and tapering. Cauda as long as the nectaries and broadly 
tapering. Anal plate slightly broader than the base of the 
cauda and triangular. 


Gen. Pentalonia Coquerel 
type nigronervosa Coq. 


Antennae slender, longer than the body and placed on promi- 
nent tubercles; spur of sixth segment longer than the third, 
first segment strongly gibbous. Antennal tubercles gibbous 
at the upper inner edge. Body short and robust, wings long 
and with the oblique veins nearly at right angles to the costa; 
cubitus twice forked and at or slightly beyond the second fork 
combines with the radial sector, thus forming a closed cell; 
veins dark, hind wing with but a single cross vein. Nectaries 
about one-fifth the length of the body, and constricted in the 
middle and just behind the tip. Cauda but a knob situated on 
a broad base as in Callipterini. Anal plate broad with apex 
parallel to the base. 


320 Annals Entomological Society of America  [Vol. III, 


Gen. Amphorophora Buckton 
type ampullata Buckton. 


Antennae longer than the body and placed on prominent 
tubercles, third segment shorter than the spur of the sixth; first 
segment slightly gibbous on the inner side. Antennal tubercles 
on inner side rounded. Head broad in front and almost flat. 
Body elongate, abdomen stout. Wings large and regular, 
nectaries about one fourth the length of the body and very 
strongly vasiform. Cauda two thirds the length of the nec- 
taries and tapering. Some species with cauda constricted about 
the middle. 


Gen. Toxoptera Koch 
type aurantiae Koch. 

Antennae about as long as the body and placed on small 
but distinct tubercles, third segment shorter than the spur of 
sixth; first segment gibbous on the inner side. Body elongate, 
wings long and slender with the cubital vein having but a single 
fork. Nectaries one sixth the length of the body and cylindrical. 
Cauda nearly as long as the nectaries and ensiform. Anal 
plate broadly rounded. 


Gen. Phorodon Passerini 
type A. humult Schrank 


Antennae nearly as long as the body and placed on prominent 
tubercles, third segment longer than the spur of the sixth; first 
segment strongly gibbous. Antennal tubercles bearing a long 
tooth on the inner side, in the apterous forms this is more 
developed. Forehead narrow with a small tubercle bearing 
a frontal ocellus. Body elongate, wings long and broad with 
cubitus twice forked. Nectaries about one sixth the length of 
the body, cylindrical, and with a slight taper at the base. Cauda 
one half the length of the nectaries, broad at the base and 
tapering to a sharp point. Anal plate broadly rounded. 


Gen. Rhopalosiphum Koch 
type A. persicae Sulzer. 

Antennae slightly longer than the body and placed on 
prominent wedge shaped tubercles, third segment about the 
same length as the spur of the sixth; first segment gibbous on 
the inner side. Antennal tubercles with a blunt tooth on the 


1910] Key to the Genera of Aphidinae a2 | 


inner side, the whole tubercle appearing as a wedge inserted 
between the head and the first antennal segment. Forehead 
narrow, outer edge of tubercles appear constricted. Body 
elongated, wings long. Nectaries about one fourth the length 
of the body and clavate at the outer end, cauda short, anal lobe 
broadly rounded. 


Gen. Myzus Passerini 
type A. cerast Fabricius. 


Antennae about as long as the body and situated on distinct 
tubercles, third segment as long as spur of sixth; first segment 
strongly gibbous. Antennal tubercles strongly gibbous on the 
inner side. Body robust, wings long and broad with cubitus 
twice forked. Nectaries about one fourth the length of the 
body, cylindrical, slightly constricted near the tip and slightly 
curved outward. Cauda less than half the length of the 
nectaries and tapering. Anal plate broadly rounded. 


Tribe APHIDINI 


The characters which separate this tribe from the previous 
are taken as follows. Antennae shorter than the body, or when 
as long as the body nectaries and cauda very short. Antennal 
tubercles, when present, are indistinct or else the nectaries and 
cauda are small. When nectaries are very long or large the 
development is limited and the other characters are used to 
place the genera. -Liosomaphis has large nectaries but the 
antennae and cauda are very short. Mastopoda has the anten- 
nae more developed than other genera but the tubercles are 
wanting. 


lM Nectaries “strongly: “claviatet. (20) «yam ere ee ner ees ie ee IES 
= INectaries taperingian dilon pies .<, Sia oce setae ayes ate ee ren ie Ws 
— Nectanies cylindmicalvand longs (pyre neers ys eer ens een ae ne IW 
-- Nectaries very short or obscure AT CS Aes ee nis ta Rete en Sere We 


II. Antennae shorter than the body, spur of sixth segment not longer than 
third segment. Nectaries long and strongly clavate on one side. 

Liosomaphis 

= Antennae slightly shorter than the body and with the spur of sixth segment 

much longer than third segment. Nectaries slightly swollen throughout 


GMEROULET HM ALE: *) oh. A ye Yee ok coer esha rene eaenete ert cee ee aS Hyadaphis 
III. Antennae about as long as the body, nectaries long and tapering and longer 
Gliasar the) Ganda 50a. va ty steele eee ee Ie eR eae Aphis 


= Antennae shorter than the body, nectaries short and tapering and nearly 
half as wide as long. Cauda short but as long as the nectaries. 

Pergandeida 

IV. Antennae as long as the body, nectaries nearly one fourth the length of the 

body.) Larst ‘andy claws natrophied.-a nt ae Aheee coe Mastopoda 


322 Annals Entomological Society of America  [Vol. III, 


Antennae much shorter than the body, nectaries slightly constricted at 


the tips cic where ih Oe aes es ot ae ee Coloradoa 
Antennae shorter than the body and with but five segmented antennae. 
Nettaries’ shortand cylindrical... hci + secs ere cen ane Cerosipha 

V. Nectaries prominent and readily distinguishable...................... VI. 
— Nectaries very small and‘obsctite. 24.0. sae es ts oe de ae ee VII. 
VI. Antennae shorter than the body, nectaries very stout and tapering. Cauda 
short. ands irck:). <6 et onan «tyne han ere ees _.. Pergandeida 
Antennae shorter than the body, nectaries very short in above but slender 

and swollenin the middle ss©. 21... 5.45 whats. leg ome ee oe Hyalopterus 


VII. Antennae much shorter than the body, nectaries cylindrical and as broad 
as long. Cauda much longer than the nectaries and tapering. . Brachycolus 
Antennae longer than the body, nectaries barely distinguishable, the end 

being trumpet shaped. Cauda not as long as wide and triangular. 


Microsiphum 
Antennae shorter than the body, nectaries little more than raised rings, 
and not flanged like the preceding genus.................... Cryptosiphum 


Gen. Liosomaphis Walker 


type A. berberidis Kaltenbach 


Antennae shorter than the body and with indistinct anten- 
nal tubercles, spur of sixth segment about the same length as 
the segment; first segment gibbous on the inner side. Body 
elongate, abdomen robust, wings long and broad, cubitus twice 
forked. Nectaries about one fourth the length of the body 
and strongly clavate. Cauda less than one half the length of 
the nectaries and sharply tapering. Anal plate broadly rounded. 


Gen. Hyadaphis Kirkaldy 
Syn. Siphocoryne passerini preoccupied. 


Antennae nearly as long as the body, third segment shorter 
than the spur of the sixth. Head with a slight projection at 
the inner base of each antenna. Body elongate, wings long 
and broad, venation regular. Nectaries about one fourth the 
length of the body and clavate in the middle. Cauda triangular 
short and rounded at the tip. Anal plate broad half moon 
shaped. 

Gen. Aphis Linnaeus 
type A. sambuci Linn. ? 

Antennae as long as the body, spur of sixth segment three 
times as long as the segment and as long as the third segment. 
Body elongated, abdomen robust. Wings broad and stout, 
with cubitus twice forked. Nectaries about one fourth the 
length of the body, somewhat slender and tapering. Cauda 
one fourth the length of the nectaries, stout and set on a broad 
base. Tip broadly rounded. 


1910} Key to the Genera of Aphidinae 323: 


Gen. Mastopoda Oestlund 
type pteridis Oestlund. 

Antennae about as long as the body, third segment shorter 
than the spur of the sixth; first segment gibbous on the inner 
side. Forehead nearly flat, body elongate, abdomen robust. 
Wings long and slender, nectaries about one fourth the length 
of the body and cylindrical. Cauda very short and conical, 
anal plate broadly rounded. The atrophied tarsi and claws of 
this genus are the most distinguishing characters. 


Gen. Coloradoa new genus 
type A. rufomaculata Wilson. 

Antennae about two thirds the length of the body, third 
segment longer than the spur of the sixth which is only about 
twice as long as the sixth. Wings moderately long, nectaries 
one sixth as long as the body, cylindrical at the base and slightly 
clavate at the end as in Rhopalosiphum. Cauda two thirds as 
long as the nectaries and tapering; tip blunt. Caudal plate 
broadly rounded. 


Gen. Cerosipha Del Guercio 
type passerinina Del Guerc. 

Antennae shorter than the body and with but five segments 
and spur, third segment longer than the spur of the fifth. 
Forehead nearly flat but slightly raised in the center. Body 
elongate, abdomen the widest part of body. Wings long, 
medium wide, wing venation regular with twice forked cubitus. 
Nectaries about one tenth the length of the body and cylindrical, 
with a very slight taper. Cauda slightly shorter than the 
nectaries and tapering to a round point. 


Gen. Pergandeida Schouteden 
type ononidis Schout. 


Antennae about as long as the body, spur of sixth segment 
as long as the third segment. Forehead with a prominent 
tubercle supporting the frontal ocelli, and slightly raised at the 
base of the antennae. Body elongated, stout; wings low and 
broad with twice forked cubitus. Nectaries short stout and 
tapering. Cauda as long as nectaries and broadly rounded at. 
the tip. Anal plate broadly rounded. 


324 Annals Entomological Society of America  [Vol. III, 


Gen. Hyalopterus Koch 
type aurantiae Koch. 


Antennae shorter than the body, third segment as long as 
the spur of the sixth; first segment strongly gibbous on the 
inner side. Frontal ocelli prominent, forehead slightly raised 
at the base of each antenna. Body long and slender, wings 
also long and slender with cubitous twice forked. Nectaries very 
short and slender, being slightly swollen in the middle. Cauda 
twice as long as the nectaries and tapering to a blunt point, 
slightly constricted near the middle. Anal plate broadly rounded. 


Gen. Brachycolus Buckton 
type A. stellariae Hardy. 


Antennae shorter than the body, third segment as long as 
the spur of the sixth; first antennal joint slightly gibbous. Body 
slender, wings long and slender and with the cubitus twice 
forked. Nectaries very short, being as long as broad and 
cylindrical. Cauda about twice as long as the nectaries and 
slender, tapering to a blunt point. Anal plate rounded and 
not distinct from the body. 


Gen. Microsiphum Cholodkovsky 
type ptarmicae Cholod. 


Antennae about as long as the body and set on distinct 
tubercles, spur of sixth segment six times as long as the segment 
and longer than the third segment, first segment slightly 
gibbous on the inner side. Forehead broad, body large and 
robust. Wings long and broad, cubitus twice forked. Nec- 
taries very short, being slightly shorter than the second antennal 
segment and flanged at the end like that of a horn. Cauda 
wide, exceedingly short and triangular in form. Anal 
plate barely distinguishable as a broad elevation. The only 
American form of this genus is the species which T. A. Wil- 
liams called Cryplositphum canadense, taken on Artemisae 
ludovicianae.* 


*Special Bull. No. 1. Dept. of Entomology, University of Nebraska, July 8, 
1891. 


(ow) 
bo 
Or 


1910] Key to the Genera of A phidinae 


Gen. Cryptosiphum Buckton 
type artemisae Buck. 


Antennae shorter than the body, spur of sixth segment 
shorter than the third segment and about one and one half 
times as long as the sixth. Forehead flat, frontal ocelli not 
-prominent. Body robust, wings long with the cubitus twice 
forked. Nectaries little more than pores with the edges 
slightly raised. Cauda short and tapering, anal plate rounded. 

All of the genera given in this paper may be found with the 
original citation listed in the Entomological News for April 
1910, pp. 147-156. 


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INDEX TO VOLUME III. 


Aecanthaclisis, 40. 
fundatus, 44. 
subtendeus, 44. 

Acronycta hasta, 116. 
hastulifera, 116. 
leporina, 116. 
oblinita, 116. 
occidentalis, 116. 

Adela, 123. 
degeerella, 124. 

Aegeriidae, 100, 122. 

Alsophila pometaria, 113. 

Amendments passed, 8. 

Amphion nessus, 109. 

Amphorophora, 320. 
ampullata, 317, 320. 

Anarsia lineatella, 23. 

Angoumois grain moth, 
host of Pediculoides, 19. 

Anisota, 110. 

Anosia plexippus, 106. 

Anthomyia, 287. 

Anthonomus eugenii, 23. 
grandis, 23. 

Anthophora retusa, 20, 22. 

Anuraphis, 314. 

Apantesis parthenice, 119. 

Apatelodes, 109, 124. 
angelica, 109. 
torrefacta, 109. 

Aphidinae, 314. 

Aphidini, 321. 

Aphis, 314, 315. 
pyri, 314. 
sambuci, 314. 
tormentillae, 314. 
ulmi, 315. 

Aplodes sp., 113. 


Apparatus for the Determination of 


Optimums of Temperature and 
Moisture, 147. 

Aranea, 173, 175. 
emnculatawl76, Lid, 179; 18. 
frondosa, 173, 177, 179, 181. 
gigas, 179. 
ocellata, 177. 

Araneinae, 175. 

Arctiidae, 99. 118. 

Arctiinae, 118. 

Argynnis cybele, 106. 

Argyresthia goedartella, 124. 

Artace pemectistriga, 110. 

Assets, 4. 

Athysanini, 215. 

Atypus bicolor, 168. 


‘ 


Banks, N., article by, 40. 


327 


Basilarchia disippus, 106. 
Basilona, 110. 
Bombycia or, 112. 
Bombycidae, 107, 111, 124. 
Bombyx mori, 111. 
Boston meeting, Minutes, 1. 
Botys polygonalis, 124. 
Brachycolus, 

stellariae, 
Bradley, C. J., title of paper, 9. 
Brephos nothum, 113. 


Cacoecia, 122. 
Cacoecia cerasivorana, 124. 
California Rodent Fleas, 
of, 61. 
Calliphora, 287. 
Callipodidae, 233. 
Callistoleon, 40, 42. 
erythrocephalum, 42. 
Calocampa, curvimacula, 116. 
Cambalidae, 258. 
Carpocapsa pomonella, 124. 
Caseya irritans, 241. 
Cash receipts, 4. 
Caterpillars, structure of, 94. 
Gatocalaicaraululi7e 
Catocalinae, 117. 
Cecidomyia resinicola, 48. 
Cecidomyia resinicoloides, 
Anatomy of larva, 45. 
Ceramica picta, 117. 
Ceratophyllus acutus, 61, 62, 63, 65, 67, 
CO leo: 
canis, 70. 
fasciatus, 63, 70, 76, 81, 82. 
musculi, 70. 
Cerosipha, 322, 323. 
passerinina, 323. 
Ceratomyia amyntor, 108. 
Cercyonis alope, 106. 
@erura, 12: 
Chamberlin, R. V., article by, 233. 
Chauliodes, 281. 
Chermes ulmi, 315. 
Chigger, reference to, 15. 
Chironomus, 287, 288. 
Cilix glaucata, 113. 
Citheronia, 110. 
Chrysoleon, 40, 48. 
punctatum, 43. 
Clisiocampa, 286, 294. 
Cicada septendecem, 23. 
Cnidocampa flavescens, 120. 
Coleophora, 124. 
Collembola, 277, 293, 298, 299. 


Bionomics 


328 Index to Volume pes 


‘oloradoa, 322, 323. 

i rcconedae 1, 323. 
Yomstock, J. H., article by, 161. 
‘onotyla coloradensis, 237. 

deseretae, 235. 

Sorydalis, 280. 

cornuta, 281. 

‘osmopteryx scribaiella, 124. 
Sossidae, 121. 

Sossus cossus, 122 

‘osymbia lumenaria, 113. 
‘rambus falsellus, 124. 
‘raspedpsomidae, 235. 

*rosby, C. R., title of paper, 9. 
‘ryptosiphum, 322, 325. 

artemisae, 325. 

Ctenucha virginica, 119. 
Ctenophora, 287. 
Cucullia, 116. 

Culex, 281, 304. 
Cybister, 293. 


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Darapsa myron, 109. 
Datana integerrima, 112. 
ministra, 112. 

Davis, J. J., title of paper, 9. 

Deilephila gallii, 109. 

Deltocephalinae, 215. 

Deltocephalus. 214, 215. 

Depressaria, 122, 123. 

putridella, 124. 

Demas coryli, 115. 

Daicrisia virginica, 119. 

Diapheromera femorata, structure of 
the egg, and significance of resem- 
blance of eggs to seeds, 83. 

Diplopoda from the Western States, 233 

Distoleon, 40, 42. 

‘ bistrigatus, 43. 

verticalis, 42. 

Ditropinotus aureoviridis, 37. 

Dolomedes fontanus, 181, 182. 

Dorydini, 205. 

Drasteria crassiuscula, 117. 

erechtea, 117. 
Drepanidae, 101, 112. 
Drepana arcuata, 113. 

falcataria, 113. 
Dolba hylaeus, 109. 
Dysdera, 167. 

interrita, 167. 


Zarias chlorana, 116. 

Edwards, W. H., Resolutions on 
Death of, 10. 

Elrod, M. J., title of paper, 9. 

Endromidae, 174. 

Endromis versicolor, 111. 

Endrosis, 122, 123, 

lacteella, 124. 
Estigmene acraea, 119. 


Eubaphe nigricans, 118. 
Euchaetias egle, 118. 
Euclea delphinii, 120. 
Eucleidae, 120. 
Euphoeades troilus, 105. 
Euphydryas phaeton, 106. 
Euploeinae, 106. 

Euproctis chrysorrhea, 115. 
Eutettix, 215. 

Euvanessa antiopa, 106. 
Ewing, H. E., article by, 209. 


Executive Committee Report, 2. 


Expenditures classified, 3. 


Feltia, 117. 
Fidonia truncataria, 113. 
Field, W. L. W., 
Filistata, hibernalis, 162. 
Fleas, California Rodent, 
of, 61. 

trapping of, 66. 

.Fontaria tuobita, 2438. 


Forbes, W. T. M., article by, 94. 


title of paper, 9. 
Formicaleo, 40. 

dispersus, 43. 

marginalis, 44. 

septus, 44. 
Frenatae, 104. 

key to families, 102. 


Galleria mellonella, 124. 
Geometridae, 100, 107, 113. 
Glenurus, 40. 
dissolutus, 40. 
pulchellus, 40. 
stigmatus, 40. 
Gluphisia, 107. 
Gracilaria, 122. 
alchimiella, 124. 
Greenidea, 316, 317. 
artocarpi, 317. 
Gymnobates, 211. 


Habrosyne derassa, 112. 
Hadena turbulenta, 117. 
Halesidota caryae, 118. 
maculata, 118. 
tesselaris, 118. 
Hapalia indicans, 154. 


Hammar, A. G., title of paper, 9. 


Haseman, L., article by, 277. 
title of paper by, 9. 

Headlee, T, H., article by, 147. 
title of paper by, 9. 

Hecatera, 154. 

Hemerocampa leucostigma, 115. 

Hepialus, 96. 

Hesperiinae, 105. 

Heterocampa guttivittata, 112. 

Heterogynidae, 120. 


title of paper, 6. 


Bionomics 


Index to Volume ITI 


Heterogynis paradoxa, 120. 
Heteropus ventricosus, 15. 
Homoeosoma, 122. 
nebulella, 124. 
Hoplopsyllus anomalus, 78. 
Hormaphis hamamelidis, 
life cycle of, 144. 
Hyadaphis, 321, 322. 
Hyalopterus, 322, 324. 
aurantiae, 324. 
Hydria undulata, 115. 


Hydrocampa nymphaeata, 124. 


Hydrophilus, 293. 
Hydrobius, 293. 
Hydrous, 293. 
Hypeninae, 118. 
Hypena humuli, 118. 
Hyperchiria io, 111. 
Hyphantria textor, 119. 
Hypochilus, 169. 
thorelli, 164, 169. 


Idiopterus, 317, 319. 
nephrelepidis, 319. 
Illinoia, 317, 318. 
liriondri, 318. 
Incurvaria, 123, 
koerneriella, 124. 
Ingura, 117. 


Insects and Entomologists, 12. 


Isia isabella, 119. 
Isosoma grande, 28, 29. 
hordei, 23. 
tratiei 0) oo. 


Jasoniades glaucus, 105. 
Jugatae, 104. 


Kepolydesmus, 246, 247. 


Lacosoma, 124. 

chiridota, 114. 
Lacosomidae, 114, 124. 
Laertias philenor, 105. 
Lagoa crispata, 120. 
Lasiocampidae, 107, 110, 124. 
Leptotrachelus dorsalis, 23. 
Lepthyphantes minuta, 183. 
Libythia celtis, 105. 
Libytheidae, 100, 105. 
Libytheinae, 105. 
Linyphyia, 170, 172, 175. 

phrygiana, 170, 173. 
Linyphiidae, 170. 
Liosomaphis, 321, 322. 

berberidis, 322. 
Lithosiinae, 118. 
Lithosia complana, 118. 


Loemopsylla cheopis, 61, 70, 76, 81. 


Lophodonta, 112. 

Loxosceles, 165, 167. 
Loxosceles rufescens, 175. 
Lycaenidae, 101, 105. 

Lycia cognataria, 113. 

Lyman, H. H., title of paper, 5. 
Lymantriidae, 114. 
Lysiopetalum mutans, 233. 


Macchiatiella, 314. 


Macgillivray, A. D., title of paper, 5, 9. 


Macrosiphini, 217. 

Macrosiphum, 317, 318. 
rosae, 318. 

Malacosoma, americana 110. 
disstria, 110. 

Mamestra species of, 154. 
circumcincta, 158. 
illaudabilis, 154, 155, 157. 
laudabilis, 154, 155, 157. 
marinitincta, 157. 
olivacea, 158. 
restora, 156, 157. 
spiculosa, 157. 
StrictawlommMose: 
strigicollis, 157. 
tenisca, 158. 

Mastopoda, 321, 328. 

pteridis, 323. 

Matheson, R., title of paper, 5. 

Megalopygidae, 120. 

Melalopha, 99, 124. 

apicalis, 112. 

Melalophinae, 112. 

Melittia cucurbitae, 122. 

Meromyza americana, 22. 

Mesamia, 214. 

Microlepidoptera, 122. 

Microparsus, 317, 319. 
variabilis, 319. 

Microsiphon, 314. 

Microsiphum, 322, 324. 
ptarmicae, 324. 

Minutes, Boston Meeting, 1. 

Mirabilis jalapa, 89. — 


Mite, Predaceous, noxious to man, 15. 


Mitzmain, M. B., article by, 61. 


Moisture apparatus, 147. 
Mordellistina ustulata, 24. 


Morgan, Miss A. H., title of paper, 5. 


Morsvanwech: Et and Shull: 
article by, 144. 
Myrmeleon, 40. 
eroceicollis, 44. 
pictifrons, 44. 
uniseriatus, 44. 


Jano tes 


Myrmeleonidae from Australia, 40. 


Myzus, 314, 318, 321. 
cerasi, 321. 


330 Index to Volume III 


Nadata, 107. 
Nadata gibbosa, 112. 
Nectarosiphon, 317, 318. 
rubicola, 318. 
Nepticula, 123. 
pomivorella, 124. 
Noctua, 117. 
Noctuidae, 97, 100, 107, 115, 122. 
Noctuinae, 115. 
Nolidae, 100, 120. 
Nola cucullata, 121. 
Notodontidae, 101, 107, 111, 124. 
Notodontinae, 112. 
Nycteolinae, 117. 
Nycteola revayana, 117. 
Nymphalidae, 105. 
Nymphalinae, 106. 


Officers elected for 1910, 6. 
Opheltes glaucopterus, 186, 197. 


Optimums of Temperature and Mois- 


ture, 147. 
Oribatidae, 209. 
Oripoda, 210. 

elongata, 211. 
Orneodes, 1238. 

hexadactyla, 124. 
Oxyptilus hieracii, 124. 


Pachygnatha, 169, 170. 
Paeromopidae, 257. 
Paeromopus lysiopetalinus, 257. 
Paiteya, 258. 
errans, 258. 
Paniscini, North American, 186. 
Paniscus, alaskensis, 197. 
albotarsatus, 198. 
albovarigatus, 198. 
geminatus, 186, 198, 201, 
medius, 198. 
melanostigma, 199. 
nigripectus, 199. 
ocellatus, 199. 
subfuscus, 200. 
texanus, 200. 
tinctipennis, 201. 
Panopoda rufimargo, 117. 
Panorpa, 123. 
Panorpata, 99. 
Pantheinae, 115. 
Paonias myops, 109. 
Papilio polyxenes, 105. 
Papilionidae, 100, 105. 
Papilionina, 104, 105. 
Paraiulidae, 253. 


Paraiulus furcifer, var. sinampus, 256. 


tiganus, 254. 
zakiwanus, 253. 

Pediculoides noxious to man, 15. 
tritici, 15, 16, 17, 19. 
yventricosus, 15, 18, 19, 20, 27. 


Pentalonia, 317, 319. 
nigronervosa, 319. 
Pergandeida, 321, 323. 
ononidis, 323. 
Phalonia, 123. 
alcella, 124. 
Phlegethontius celeus, 109. 
sextus, 109. 


Pholisora catullus, 105. 
Pholus pandorus, 109. 
Phorodon, 318, 320. 
humuli, 320. 
Phyllium crurifolium, 86. 
scythe, 85. 
Physogaster larvarum, 20. 
Pieridae, 105. 
Pieris brassicae, 105. 
rapae, 105. 
Pisauridae, 181. 


_Platymetopius, Am. species of, 214. 


acutus, 217, 219; 221. 
var. dubius, 220. 
angustatus, 217, 223. 
brevis, 218, 230. 
cinereus, 217, 223. 
cuprescens, 217, 221. 
dorsalis, 216, 224. 
elegans, 216, 219. 
frontalis, 217, 225. 
fuscifrons, 217, 230. 
fulvus, 217, 224, 225. 
hyalinus, 216, 218. 
irroratus, 218, 227. 
latus, 217, 221. 
loricatus, 217, 226. 
magadelensis, 215. 
majestus, 216, 219. 
modestus, 230. 
nanus, 217, 226. 
nasutus, 217, 225. 
nigriviridis, 216, 218. 
var. dixianus, 218. 
obscurus, 217, 227, 229. 
oregonensis, 217, 220. 
osborni, 218, 229. 
rostratus, 214. 
scriptus, 218, 228. 
tenuifrons, 216, 231. 
slossoni, 217, 222. 
undatus, 214. 
verecundus, 216, 218, 227. 
vittatus, 214. 
Polydesmidae, 244. 
Polydesmus amandus, 249. 
anderisus, 246. 
sastianus, 251. 
socarnius, 252. 
sontus, 247. ; 
Polygonia interrogationis, 106. 
Porthetria dispar, 114. 


Index to Volume III 


Protoplectron, 40. 
costatus, 41. 
pallidum, 41. 
Pseudosphinx tetrio, 108. 
Psyche zelleri, 121. 
Psychidae, 97, 100, 121. 
Psychinae, 121. 
Psychoda, 281. 
Psychoda alternata, structure and met- 
amorphosis of the alimentary canal 
of larva of, 277. 
Pterophoridae, 123. 
Ptychoptera contaminata, 277, 293. 
Pulex irritans, 61, 63, 64, 65, 69, 70, 81. 
Pygerinae, 112. 
Pyralididae, 100, 122, 123. 
Pyrophila pyramidoides, 117. 
Receipts, cash, 4. 


Report of Auditing Committee, 7. 
Executive Committee, 2. 
Committee on Resolutions, 7. 
Resolutions on death of Wm. H. Ed- 
wards, 10. 
On death of M. V. Slingerland, 11. 
Rhodophora, 117. 
Rhopalosiphum, 318, 320. 
persicae, 320. 
Rodent Fleas, Bionomics of, 61. 


Sarcopsylla, penetrans, 15. 
Sarcoptides psoriques, 15. 
Saturniidae, 107, 111, 124. 
Saturniina, 110, 124. 
Scaphoideus, 215. 
Schizoneura ulmi, 315. 
Schizura badia, 112. 
concinna, 112. 
unicornis, 112. 
Scopelosoma, 116. 
Scytonotus, granulatus, 245. 
piger, 244. 
Sesiidae, 122. 
Severin, H. H. P., article by, 83. 
Shrullaeneweh sand Morgan, i) sb 
article by, 144. 
Simaethis, 122. 
Simaethis oxyacantha, 124. 
Simulium, 287. 
Simyra henrici, 116. 
Siphocoryne, 322. 
Sitotroga cerealella, 19, 29, 30, 33. 124. 
Slingerland, M. V., Resolutions on 
death of, 11. 
‘Smith, J. B., Abstract of Address by, 12. 
Article by, 154. 
Solenobia, 123. 
Solenobia pineti, 121. 
Solenobiinae, 121. 
Sphingidae, 107, 108, 124. 


dol 


Sphinx gordius, 108. 
Spiders, palpi of male, 161. 
Stratiomyia, 287. 

Striaria nazinta, 242. 
Striariidae, 242. 

Suriana maritima, 89. 
Synchlora, aerata, 113. 
Syntomidae, 119. 


Tanypus, 287. 
Telea polyphemus, 111. 
Temperature, apparatus, 147. 
Tenebrio molitor, 299. 
Tetragnatha extensa, 169. 
Tetraneura ulmi, 315. 
Thaumetopoeidae, 114. 
Thaumetopoea, 114. 
Thecla ilicis, 105. 
Thyatiridae, 100, 101, 107, 112. 
Thyridopteryx ephemeraeformis, 121. 
Thyris, 123. 
Thyris vitrina, 124. 
Tineidae, 99. 
Tineina, 100. 
Tineola, 123. 

biselliella, 124. 
Tingupa, 238. 

monterea, 240. 

utahensis, 238. 
Tipula, 287. 
Tortricidae, 99, 122, 123. 
Toxoptera, 318, 320. 

aurantiae, 320. 
Trichoptera, 99. 
Trichosiphini, 316. 
Trichosiphum, 316. 

aonae, 316. 
Triggerson, C. J., title of paper, 5. 
Tropaea luna, 111. 


Van Duzee, E. P., article by, 214. 


Walking Stick, structure of egg, 83. 
Washburn, title of paper, 6. 
Webster, F. M., title of paper, 5. 
article by, 15. 
Wheeler, W. M., title of paper, 5. 
Whitmarsh, R. D., article by, 186. 
Williams; F. X., article by, 45. 
Wilson, H. F., article by, 314. 


Xystodesmidae, 243. 


Yponomeuta, 122, 123. 
cagnagellus, 124. 


Zerene catenaria, 113. 
Zygaenidae, 120. 
Zygaena trifoli, 120. 
Zygaenina, 119. 


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CHAMBERLIN, RALPH V.—Diplopoda from the Western 4 af 
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Hasreman, L.—The Structure and Metamorphosis of he if 
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Attermsta Say SC ky Si Pik an Sota nae 27h ae 
Wi1son, H. F.—A Key to the Genera of the Subfamily , ee 
Aphidinae and Notes on Synonomy..-...:.----- 314 7 
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