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Full text of "The National Collection of Coccidae"

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Technical Series, No. 16, Part I. 

U. S. DEPARTMENT OF AGRICULTURE, 

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



PAPERS ON COCCIDiE OR SCALE INSECTS. 



THE NATIONAL 
COLLECTION OF COCCIDJE, 

/6 -/y - 



By C. L. MARLATT, M. S., 

Entoynologist and Assistant Chief of JBtireau. 



Issued April 23, 1908. 




"WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1908. 



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CONTENTS. 



Historical summary 1 

Kecent work of the Bureau 4 

Preservation of material 5 

The slide collection 5 

Recent additions and present status 6 

Methods of study 7 

The work of the future 9 

A plea for the deposit of types 10 

III 



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U. S. D. A., B. E. Tech. Ser. 16, Pt. I. Issued April 23, 1908. 

PAPERS ON COCCIDJl OR SCALE INSECTS. 



THE NATIONAL COLLECTION OF COCCID^. 

By C. L, Marlatt, M. S., 
Entomologist and Assistant Chief of Bureau. 

The collection of scale insects or Coccidse, accumulated chiefly 
through the agency of the Bureau of Entomology of the U. S. De- 
partment of Agriculture and forming a part of the national collec- 
tion of insects, is undoubtedly the most valuable and complete in 
existence. This has come from the early interest of the Bureau in 
the subject through the work of Prof. J. H. Comstock, and as a result 
of the generosity of many of the describers of new species of later 
years in depositing types or cotypes in the National Collection. It 
may not be out of place, therefore, to give a brief historical summary 
of the National Collection of Coccidse, together with a description 
of its present condition, an account of the methods of installation 
and preservation of material, and a numerical statement of species 
and types represented. 

HISTORICAL SUMMARY. 

The knowledge in America of this important group of insects was 
very scant prior to 1880. During that year Prof. J. Henry Comstock, 
then Entomologist of the Department of Agriculture, in the course of 
an investigation of the insects injurious to orange trees, begun the 
previous year, noted that the greater part of the insect injury to citrus 
fruits was due to scale insects. This led him to make, during 1880, a 
particular study of the family Coccidse, broadening the inquiry to 
include the study of all scale insects affecting cultivated plants in the 
United States. Professor Comstock'personally collected many south- 
ern species in a trip through Florida in January and February, 1880, 
and during the following summer spent three months in the fruit- 
growing sections of California and Utah investigating the subject and 



2 PAPERS ON COCGIDiE OR SCALE INSECTS. 

collecting scale insects. He was assisted also by several of his col- 
leagues in the office, notably Dr. L. O. Howard and Mr. Theo. Per- 
gande. The results of this work appeared as Part II of his Annual 
Report for 1880, in a paper on scale insects, which included descrij)- 
tions of most of the Coccidse then in the collection of the U. S. Depart- 
ment of Agriculture, lack of space preventing the description of all 
the species which had been collected up to that time. Some 45 species 
are listed and described in this report, more than half of which were 
new to science. Previous to this only about '30 species had been de- 
scribed by American writers, the majority of which are to be credited 
to Dr. Asa Fitch. Most of the species referred to in Comstock's 
report were figured, and these figures mark the beginning of really 
careful scientific drawings of Coccidse, and have since been standard 
illustrations of the species concerned. 

Doctor How^ard, while assisting Professor Comstock in the study 
of Coccidse, took up particularly the subject of the chalcidid para- 
sites of these insects, and presented as Part III of Professor Com- 
stock's report a paper entitled " Report on the Parasites of the 
Coccidse in the Collection of this Department." This was the begin- 
ning of Doctor Howard's work on chalcidid parasites and led to his 
becoming the world's authority on the subject. How little was known 
of these parasites previous to the publication of this paper is shown 
by the fact that, with four or five exceptions, the species described 
and figured were new to science. 

Professor Comstock retired from the position of Entomologist in 
1881, but published in Professor Riley's Report for that year a short 
illustrated article on Lac Insects, describing three new species. He 
also gave a brief article on methods of controlling scale insects. A 
much more important article on remedies for scale insects of the 
orange, by Mr. H. G. Hubbard, is published in the same report. In 
this pajDCr some very good work is recorded and some of the more 
important modern methods of control are given first exploitation. 

Professor Comstock's report on Coccidse in the Department Report 
of 1880, as already indicated, was incomplete and did not include all 
of the species collected. His final paper occupied the principal por- 
tion of the Second Report of the Department of Entomology of the 
Cornell University Experiment Station, published in 1883. This 
]3aper is a brief monograph of the subfamily Diaspinse as repre- 
sented in the fauna of North America, together with notes on species 
not at that time observed in the United States. Several new species 
are described in this rej)ort, and the latter concludes with a list of 
the American nondiaspine scale insects. 

This practically concluded Professor Comstock's work on Coc- 
cidse, and for a long period thereafter the systematic study of this 
important group of insects practicall}^ ceased in this country. The 



THE NATIONAL COLLECTION OF COCCID^. 3 

Department Collection contained tyj^es or cotypes representing prac- 
tically all of Comstock's work, and examples of the known species 
collected by him, and Mr. Theo. Pergande undertook, during the 
years following 1880, to identify the material which was sent to the 
Department by comparison with named material in the collection. 
Mr. Pergande's work during this period was of great value to the 
Bureau and to the entomologists in this country, but necessarily 
much that came in could not be identified, and there was a large 
accumulation of unnamed material representing in part old species 
and in part species new to science. 

Mr. H. G. Hubbard, who has already been referred to as having 
published an important paper on remedies for Coccida? in the Annual 
Report of the Department for 1881, continued his investigation of 
orange insects and the means of controlling them, at Crescent City, 
Fla., as an agent of the Department of Agriculture, and published in 
1885 his excellent report entitled " Insects Affecting the Orange." 
This represented nearly four years' study and experiment, and dealt 
very largely with the common scale enemies of the orange. Mr. Hub- 
bard's interest in scale insects, aroused in this study, continued, 
although he published nothing noteworthy afterwards on the subject. 
It was, however, his intention to bring out a new and thoroughly 
revised edition of his Orange Insects, and to take up in connection 
with it a study of the classification of Coccidse; but his untimely 
death prevented the accomplishment of this task. 

About 1891 Prof. T. D. A. Cockerell began to collect and study 
the Coccidi^e of Jamaica, where he was then stationed. For some 
years much of his material came to this office for determination, but 
in a short time he became familiar with the subject, and ultimately 
the world's best authority in the group, and for a period of several 
years much of the new material coming to the Department was sent 
to him for determination and description. Professor Cockerell had 
in the meantime accepted a position in the New Mexico Agricultural 
College, and his collections and descriptions of Coccidse were enriched 
by the inclusion of many new southwestern species as well as by 
material which came to him from all quarters of the world. Very 
fortunately his close cooperation and sympathy with the Bureau of 
Entomology and interest in the National Collection led him to deposit 
his types in Washington, thus verj^ greatly enriching the local collec- 
tion. Professor Cockerell, while diminishing very much his activity 
in this group in later years, has retained his interest. During his 
most active decade his publications on Coccidge were voluminous, 
much exceeding the work of any other investigators in this field with 
the possible exception of Signoret and Maskell. In connection with 
this Bureau he published several papers, notably his bulletin on the 



4 PAPEES ON COCCIDiE OK SCALE INSECTS. 

San Jose scale and its nearest allies,'^ and, through the Smithsonian 
Institution, papers on the food plants of scale insects,^ directions for 
collecting and preserving scale insects,'' and geographical distribution 
of scale insects."' 

RECENT WORK OF THE BUREAIT. 

The resumption of active study of Coccidse in connection with the 
Department of Agriculture began in the early nineties, partly as a 
result of the interest aroused by the San Jose scale and the necessity 
of being able to determine at once and authoritatively all material 
which was coming in supposedlj^ representing this new and danger- 
ous species. It proved, in other words, impracticable to be dependent 
for such determinations on specialists at a distance. Mr. Pergande» 
who had had charge of the coccid collections, had his time so fully 
taken up with other lines of investigation, notably the subjects of 
Aphididse and Formicidse, that he could not keep up with the rapidly 
increasing collection and literature in Coccidae. To meet the need 
and following the incentive of an old interest the writer therefore 
took as his special field in systematic entomology the subject of 
Coccidse. A season spent in studying orange insects in southern Cali- 
fornia accentuated his interest in the subject, and general charge 
of the Department collections in this family was assumed by him in 
1900, and a thorough reorganization of the collection and notes was 
then begun. Before this work had progressed very far it was inter- 
rupted for more than a year by a trip to the Orient, in which, how- 
ever, the scale insects were the special object of study and collection. 

During the twenty years which had elapsed since the work of Corn- 
stock, a great mass of material had accumulated, most of it preserved 
under very bad conditions, and more than half of it unidentified. It 
was no small undertaking to bring all this mass into order, and in 
this work the writer was assisted during a number of years by Messrs. 
J. Kotinsky and E. R. Sasscer. 

The enormous increase in the number of scale insects, and the 
amount of technical work necessary to determine the validity of spe- 
cies and to identify new material, made it early evident that it would 
be impossible for the writer to keep the whole subject as his own- 
field, and in January, 1905, Mr. J. G. Sanders was appointed an 
assistant in the Bureau and became the writer's colleague in the study 
of Coccidse, the important field of the nondiaspine scale insects, to- 
gether with the collections and notes, being turned over to him un- 
reservedly. Mr. Sanders is also charged with the very important 

« Tech. Ser. 6, Div. Ent., U. S. Dept. Agric, 1897. 
&Proc. U. S. Nat. Mus., XIX, No 1122, pp. 725-785, 1897. 
c Bui. 39, Part L, U. S. Nat. Mus., 1897. 
^ Proc. U. S. Nat. Mus., XYII, No. 1026, pp. 615-625, 1895. 



THE NATIONAL COLLECTION OF C0CCIDJ5. 5 

inspection and quarantine work of the new importations made by 
the Department, and in this field has detected and prevented the 
introduction of a number of new scale insects which probably would 
have made very destructive records if established in this country. 

PRESERVATION OF MATERIAL. 

The first effort was to arrange for the proper preservation of the 
type and other material, and a modification of the system followed 
by Green was adopted — namely, the inclosing of material in strongly 
made pasteboard boxes, following a definite unit of length and 
breadth, 3 by 2 inches, so that they can be arranged in five columns 
in the standard insect drawer which has been in use in the U. S. 
National Museum ; '^ these boxes vary in depth, however, to accomo- 
date different quantities of material. All type and valuable material, 
in fact all material not in large quantity, is further protected by being 
first folded in squares of lens paper to retain any loose scales or insects. 
By this system the bulk of the collection was enormously reduced 
and loss of valuable specimens obviated. This system presents also 
the great convenience that additions may be made with the same facil- 
ity as in a card catalogue. 

Special card labels suitable to the nature of the collection were 
secured from the Library Bureau. These are of different colors to 
indicate respectively the genus, the valid species and varieties, and 
the typical material of the species reduced to synonymy. 

THE SLIDE COLLECTION. 

The slide preparations of the different species had become very 
extensive, and now comprise fully 10,000 slides, relating to more 
than a thousand species. A new system of storage for slide material 
was devised, following the plan of the ordinary card catalogue cabi- 
net.^ The older systems of slide storage had proved to be very un- 
satisfactory for so large a collection representing so many different 
species. None of the older systems was sufficiently elastic or allowed 
easy systematic classification, but required card indexes and the con- 
sultation of many different slide boxes to get together material repre- 
senting a single subject. This led the writer to devise a cabinet in 
which the slides could be filed on the card index plan. To enable 
a slide to be thus stored a special envelope or jacket is provided for 
each slide. This jacket is made of strong manila paper and but 
little larger than the standard slide size (slide 1 by 3 inches, jacket 
1^' by 3^ inches), affords an opportunity for sufficient labeling, and 

<* The insect drawers since tlien adopted by tlie National Museum, and in- 
tended for pinned insects, are somewliat sliallower ttian ttiose first used. 
» See Science, Dec. 30, 1904, p. 925. 



6 PAPEES ON C0CCIDJ3 OR SCALE INSECTS. 

can be filed exactly as an index card. The cabinet follows the general 
plan of a library card index, is of five drawers of three partitions 
each, and holds upward of 2,000 slides. Guide cards were devised to 
correspond with the slide envelope, and have on their lower edge an 
extended lip perforated for the countersunk retaining rod which 
runs just below the level of the slides. This rod holds the guide cards 
and gives additional security to the slides, which remain steadily in 
place by their own weight. Each cabinet is arranged with a lock 
which fastens all five drawers. The latter have the retaining device 
so that there is no danger of pulling them entirely out and spilling 
their contents. The envelopes, however, give such protection to the 
slides that the latter will not break even from falling some distance, 
and, furthermore, they protect the slides entirely from dust. The 
slide cabinets, envelopes, and guide cards are all made by the Library 
Bureau at very reasonable cost. The slide jackets cost about $2 a 
thousand, and the cabinets are but little more expensive than the ordi- 
nary index case. 

This system of storing slides allows for additions indefinitely in 
exact classification, with as minute and accurate subdivisions as may 
be necessary. It has now been in use some five years and has given 
complete satisfaction. The work of consulting the slides and han- 
dling the material is reduced at least two-thirds, and consultation is 
so easy that it increases the usefulness of the slide collection many 
times. 

The cabinet described here in some detail is suitable for dry balsam 
mounts, and in the preparation of slides of Coccidse the mount, as 
soon as prepared, is dried over an alcohol flame for a few seconds, 
when it can be put at once into the jacket. In any case where the 
slide can not be thus dried it must be laid in a horizontal position 
until the balsam has become thoroughly hardened. Once thoroughly 
dried, the intense heat of Washington summers does not result in any 
softening or attachment of the slide to the inclosing jacket. 

A card catalogue has been made of all the material which has come 
to the Department, and the accumulated notes have been systemat- 
ically arranged so that they are available for instant reference. This 
work has been done whenever time was available, and has often been 
interrupted for weeks or months together, but has now resulted in 
the collection, notes, and records being in exceptional order and in 
shape for study by the specialist with the greatest economy of time. 

RECENT ADDITIONS AND PRESENT STATUS. 

The deposit of the types of Professor Cockerell's new species repre- 
sent the most important additions since Comstock's time. Following 
this commendable example, most of the other workers in this group 



THE NATIONAL COLLECTION OF COCCIDiE. 7 

in this country have given their types, or at least cotypes, to the Na- 
tional Collection. Among this material are the species described by 
Newell, King, Sanders, Tinsley, Johnson, Townsencl, Cooley, Cole- 
man, and Woglum. Cotypes of the greater number of Green's ori- 
ental species have also been most kindly presented by him to the Na- 
tional Collection, making a large and very valuable addition. Co- 
typical or authentic material representing many of the new species 
of other foreign workers in the family has been received in exchange 
or by gift, including a considerable percentage of Maskell's species 
and those of later students, notably Hempel. 

The collection now includes, besides much unidentified material, 
representatives of 1,072 species of Cocciclse, of which 816 are types or 
cotypes. Of these 366 belong to the subfamily Diaspinse, with 293 
types; and 706 to the non-diaspine scales, with 523 types. The de- 
scribed Coccidse of the world now reach a total of nearly 1,600, but 
of these, from the critical studies which have been in progress now 
for several years, it seems likely that at least one-third will fall as 
synonyms, leaving perhaps a thousand valid species. A correspond- 
ing projDortion of the type and other material in the National Collec- 
tion of Coccidse will fall as synonyms. Nevertheless, the figures given 
indicate a collection of extraordinary richness and value in its accumu- 
lation of species and types. In working out synonymy the type 
specimens of the synonyms, together with the material referred to 
these synonyms by the describers or later workers of authority, are 
kept together, so that they can be easily referred to should any ques- 
tion arise as to their status. 

METHODS OF STUDY. 

A ver}^ large amount of laborious preliminary work has been clone 
which makes no showing, but in the course of this work those who are 
actively conducting it have gained such a familiarit}^ with the subject 
and so broad an acquaintance with the ditferent genera that it should 
now be possible for them to monograph the material far more sat- 
isfactorily and accurately than ever before. The opportunity now 
had for comparison of types leads necessarily to the discovery of a 
large amount of synon3^my, and the task of weighing and definitely 
ascertaining the status of older work is now in progress and repre- 
sents the most difficult part of the undertaking, although a necessary 
preliminary to the description of new species and the monographing 
of genera. 

In attempting to make determinations from published descriptions 
the inadequacy of the characterizations was at once felt, through no 
fault, however, of the early describers, who followed the customary 
methods of the time, just as the descriptions of this day may similarly 



8 PAPERS ON COCCIDiE OR SCALE INSECTS. 

prove to be inadequate fifty years from now. Tlie principal diffi- 
culty, however, was that many structural features of the insect were 
not mentioned in the descriptions, and hence a systematic arrange- 
ment from descriptions Avas impossible, or, if attempted, very faulty. 
To give a standard for comparisons of descriptions, the writer has 
drawn up a scheme to be followed which has proved to be a great 
convenience. It at least insures that no important character is 
omitted, and the discussion of the different parts in regular order 
allows an easy comparison of the same characters for different species. 
Card blanks have been prepared, and following the printed indi- 
cations the descriptions can be made very readily in abbreviated 
form and elaborated as fully as necessary for publication. These 
cards, being of the same size (5 by 8 inches) as the standard note 
card adopted by the Bureau, are filed as a permanent record. The 
card illustrated below is designed especially for the diaspine scale 
insects. Some modification of this card is necessary for the non- 
diaspine species, and Mr. Sanders has worked out such a modifica- 
tion, also illustrated. It is hoped that the describers of new species 
of scale insects will follow the order indicated on these cards, so that 
all future descriptions may be readily compared. 

Order followed in descriMng Diaspince. 
Name : 

Female scale: (Size, shape, density, convexity, color, pellicles, venti'al scale, 

situation, etc.). 
Male scale: (Do.) Male insect : (Unimportant). 

Adult female: (Shape, size, color, chitinization, anterior and marginal char- 
acters, stigmatic pores — if present, relationship). 
Anal plate : ( Shape, size, color, chitinization, etc. ) . 

Lobe: (No., size, shape, chitinization, position, etc.). 
Lateral teeth : (Anterior to lobes and plates). 
Incisions: (i. e., notches between lobes). 

Paraphyses : (Chitinous thickenings at base of lobes and sometimes mar- 
ginal beyond lobes). 
Plates: (No., character, situation). 

Spines: (Position same for all species, as a rule unimportant). 
Anal opening: (Size, shape, position). Genital opening: (Unim- 

portant). 
Paragenitals : (No., massed or scattering, etc.). 
Dorsal pores: (No., size, distribution, etc.). 
Basal thickenings: (Transverse, broken, rod-like elevations near base of 

plate — dorsal). 
Ventral thickenings: (Broad areas of slight chitinization extending from 
tip and longitudinal ridges laterad of vaginal opening). 
Type: (Dept. No., U. S. Nat. Mus. No., host, locality, coll., date). 
Note: (Prominent distinguishing characters, relationships, etc.). 



THE NATIONAIj COLLEGTIOl"^ OF COCCID.E. 
Order foUotced in -descriMng non-diaspine Coccidce. 



Name: 



Male: (Scale or puparium; adult, emergence). 

Female: (Size, shape, color, markings, secretion, maturity, reproduction). 

Derm (cleared) : (Cliaracter, size, outline, color, cliitinization, pilosity, 

color in KOH). 
Antenn.^ : (Location, segments, measurements in ,u, formula, hairs). 
Legs: (Location, development, pilosity, coxa, trochanter, femur, tibia, tar- 
sus, claw, digitules). 
Spiracles: (Location, character, pores). 

Pores: (Character, simple or compound; arrangement, dorsal, ventral). 
Spines — marginal: (Character, length, arrangement). Spiracular : (Char- 
acter, length, arrangement). 
HYPOPYGiAL : (Character, number, arrangement). Surface: (Char- 
acter, . length, arrangement ) . 
Anal plates: (Shape, size, character, location). 

Set^e — fringe: (Number, location, length). Subapical : (Number, 
location, length). 
apical: (Number, location, length). 
Thickenings: (Character, location). 
Anal ring: (Shape, character, number of hairs). 
Type: (Host, locality, date, collector). 
Note: (Prominent distinguishing characters, peculiarities, relationships, etc.). 

THE WORK OF THE FUTUBE. 

It is the intention of the writer and of his colleagues in the study 
of this family to work out the synonymy of described material which 
has accumulated, to prepare synoptic tables of the different genera, 
and describe the species wdiich seem to be new to science. These 
papers will be issued from time to time as parts of the present bulle- 
tin and of later bulletins of the Technical Series of this Bureau. 

The catalogue and bibliography of Coccidse, started in such a com- 
prehensive and thoroughgoing manner by Mrs. Fernalcl, will be 
continued in biennial supplements by Mr. "J. G. Sanders, one such 
supplement having already been issued.'^ 

One of the great difficulties in the study of Coccidse has been the 
inaccessibility of the types of Maskell's many new species, especially 
as the descriptions and figures were in most cases inadequate. It is 
a pleasure, therefore, to be able to announce that, through the cour- 
tesy of Mr. T. W. Kirk, the Department is to be allowed to make a 
critical examination and study of all of Maskell's types and other 
material, which are now making their long journey from Wellington, 
New Zealand, to Washington for this purpose. 

The field of Coccidse is a very large one, and there is plenty of room 
in it for a considerable number of students, and the National Collec- 

"Tech. Ser. No. 12, Part 1, Bur. Ent., U. S. Dept. Agriculture. 1906. 



10 PAPERS ON COCCID.E OR SCALE HsTSECTS. 

tion is open to anyone who wishes to study or consiih it. A good deal 
of exchange material of value has been accumulated, and exchanges 
for species not represented in the Department Collection will be 
gladly made. 

A PLEA FOR THE DEPOSIT OF TYPES. 

. The Coccida? present special difficulties in the way of determination 
on account of their small size and the necessity of making careful 
microscopic preparations. Furthermore, it is not very practicable 
to determine material from descriptions only, and it is necessary in 
most cases to have type, or accurately identified material, for the pur- 
pose of comparison. With Coccidse, therefore, more than with other 
groups of insects, it seems necessary to have some central depot for 
the deposit of types.' where authoritative identifications may be made. 
It is hoped and urged, therefore, that all American describers of new 
species will adopt the course of placing type or cotype material in 
the Xational Collection, where it will be carefully preserved and be 
available for study at any time, 

O 



^fopeny ol tne uniled Stales Goyefriragjil 

Technical Series, No. 16, Part II. 

U. S. DEPARTMENT OF AGRICULTURE, 

BTJRE^XJ OF lS,NTOm:OT-.OG^^. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



PAPERS ON COCCIDJ; OR SCALE INSECTS. 



NEW SPECIES OF 
DIASPINE SCALE INSECTS. 



ByC. L. MARLATT, M. S., 

Entomologist and Assistant Chief of Bureau. 



Issued August 5, 1908. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1908. 



u<s. su' 



CONTENTS 



ILLUSTRATIONS. 



Introductory 11 

Aspidiotus (s. str. ) comperei : ■ - - . , 

Aspidiotus (s. str.) meyeri ' 

Aspidiotus ( Aonidiella) cocotiphagus ; - 

Aspidiotus (Diaspidiotus) africanus : ^ 

Aspidiotus (Diaspidiotus) coursetise 20 

Aspidiotus ( Diaspidiotus) transcaspiensir 21 

Aspidiotus (Hemiberlesia) epigaete - 21 

Aspidiotus (Hemiberlesia) mitchelli 22 

Aspidiotus (Hemiberlesia) popularum 23 

Aonidia juniperi 24 

Aspidiotus (Targionia) chenojjodii 24 

Cliionaspis micropori 25 

Leucaspis indica 26 

Mytilaspis cliilopsidis 27 

Parlatoria mangiferse 28 

Parlatoria pyri 29 

Parlatoria cliinensis - - - 30 



Plate I. Fig. 1. — Aspidiotus (s. str.) comperei. Fig. 2. — Aspidiotus (s. str.) 

meyeri 12 

II. Fig. 1. — Aspidiotus (Aonidiella) cocotiphagus. Fig. 2. — Aspidiotus 

(Aonidiella) cocotiphagus 14 

III. Fig. 1. — Aspidiotus (Diaspidiotus) africanus. Fig. 2. — Aspidiotus 

(Diaspidiotus) coursetite 14 

IV. Fig. 1. — Aspidiotus (Diaspidiotus) transcaspiensis. Fig. 2. — Aspidi- 

otus (Hemiberlesia) epigseee 22 

V. Fig. 1. — Aspidiotus (Hemiberlesia) mitchelli. Fig. 2. — Aspidiotus 

(Hemiberlesia) popularum 22 

VI. Fig. 1. — Aonidia juniperi. Fig. 2. — Aspidiotus (Targionia) cheno- 

podii 24 

VII. Fig. 1. — Cliionaspis micropori. Fig. 2. — Leucaspis indica 26 

VIII. Fig. 1. — Mytilaspis chilopsidis. Fig. 2. — Parlatoria mangiferse 28 

IX. Fig. 1. — Parlatoria pyri. Fig. 2. — ^^Parlatoria chinensis 30 

II 



U. S. D. A., B. E. Tech. Ser. 16, Pt. II. Issued August 5, 1908. 

PAPERS ON COCCID^ OR SCALE INSECTS. 



NEW SPECIES OF DIASPINE SCALE INSECTS. 

By C. L. Marlatt, M. S., 
Entomologist and Assistant Chief of Bureau. 

INTRODUCTORY. 

The species described below as new to science are mostly insects of 
potential economic importance, as indicated by place of origin and 
host plants. The facts concerning each of these species in these 
particulars are given in the notes accompanying the descriptions. 

Of the seventeen species listed, four are apparently native to this 
continent, and the rest are foreign. Of the latter, five species have 
been found on living plants recently imported from foreign countries. 
Two of these, Leucaspis indica and Parlatoria mangiferse, attack 
mango. The former is established in mango plantings in Florida and 
Porto Rico, and the latter is found more or less generally infesting 
the mango nursery stock in the Department greenhouses. A vigor- 
ous effort is being made to exterminate both of these scale insects. 
Parlatoria fyri infested cuttings of apple and soft pear imported 
from Manchuria, and might easily become a pest second only to the 
San Jose scale in importance. All of the infested cuttings were 
burned, and the entire stock was thoroughly fumigated. Parlatoria 
cTiinensis infested XantJioxylon, Thuja, and Zizyphus sp. imported 
from China, and is closely related to 'pyri. Both of these Parla- 
torias were collected by the writer in China in 1901. Aspidiotus 
meyeri infested an Ahies imported from the region of Peking, 
China. The rest of the foreign material came to the Department 
for determination from various collectors as dry, dead specimens, 
and the danger of the introduction of these species is a future one. 
Of the latter, the ones presenting the greatest danger to this country 
are Aspidiotus cocotiphagus , which in Cuba infests the cocoanut 
and trifoliate orange, and is doubtless capable of developing a wide 
range of food plants, and Aspidiotus africanus, which is making in 
South Africa a reputation very similar to that of the San Jose scale 

in this country. 

11 



12 PAPEKS ON COCCID^ OE SCALE INSECTS. 

Credit is due Mr. J. G. Sanders, of this Bureau, who made the photo- 
micrographs of the anal plates of the species described in this paper. 
While it does not seem possible to get a photograph which will show 
satisfactorily all the structural characters of the anal plate, these 
photographs are still very helpful as supplementing the text, and 
give a general pictorial effect which it is impossible to convey by 
description. 

ASPIDIOTUS (s. str.) COMPEREI n. 

(Plate I, fig. 1.) 

Scale of female: About 1 mm. in diameter, strongly coavex, nearly- 
circular. Exuvige covered, but covering secretion easily rubbed off, 
exposing the lemon-yellow to brown exuviae. Secretion covering 
the larval exuvia sometimes more dense and adherent, thus some- 
times giving the scale an annulated appearance due to the second 
exuvia showing through the section surrounding the larval skin. 
Secretionary supplement normally white, sometimes slightly yel- 
lowed. Ventral scale inconsiderable. 

Scale of male: About one-half the size of the female scale, and of 
the normal oval shape. 

Adult female: In balsam mount, nearly circular and hyaline, with 
a diameter of about 0.8 mm.; anal plate broad, rounded, about 
0.2 mm. long by 0.3 mm. broad, for the most part hyaline, apical 
third very slightly suffused or chitinized; three pairs of very small, 
narrow, subequal lobes; median lobes rectangular and more heavily 
chitinized than laterals, and with chitinization, not sharply limited, 
extending into the p3^gidium something more than a lobe's length; 
lateral lobes with distinct outer shoulder and somewhat constricted 
at base, hyaline; a projection is sometimes seen at some distance 
anterior to the third lobe having the appearance of a rudimentary 
fourth lobe; incisions shallow, given a truncate appearance by the 
occurrence of marginal pores; paraphyses wanting; plates about 
lobe length, inconspicuous, apparently two median, two in first 
incision, and three in second incision, with some additional rudi- 
mentary plates beyond second lateral incision; spines normal; anal 
opening oval, very large, larger than the two median lobes together^ 
remote from tip, near center of the segment; paragenitals repre- 
sented onl}'^ by the anterior lateral groups, which consist of from 1 
to 4 (usually 2) glands; dorsal glands small and rather numerous,, 
limited to the apical third of the margin of the segment; basal thick- 
enings wanting; ventral thickenings not present as a divaricating 
band from the tip, but represented by the uniform chitinous suf- 
fusion of the apical third of the segment. 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agricul-ture. 



Plate I. 




Fig. 1.— Aspidiotus (s. str.) comperei. 




Fig. 2.— Aspidiotus (s. str.) meyeri. 
ANAL PLATES OF NEW SCALE INSECTS. 



NEW SPECIES OF DIASPINE SCALE INSECTS. . 13 

Type. — Bureau of Entomology No. 14129. On Hake sp. and an 
undetermined plant. Raventhorpe, West Australia, collected by 
George Compere (Nos. 968 and 990), received at the Department 
February, 1902. 

Note. — This species in scale characteristics and large anal orifice 
reminds one of Aspidiotus camellise Sign.; in its pygidial characters 
it is more like A. destructor Sign. It differs distinctly, however, 
from both of these species, as indicated in the description. It occurs 
massed rather thickly on the smaller twigs of the two food plants. 
It does not seem to coincide with any of the species described by 
Maskell, although, when Maskell's types are examined, it may fall 
to one of his species. 

ASPIDIOTUS (s. str.) MEYEBI n. sp. 

(Plate I, fig. 2.) 

Scale of female: Elongate oval, 1.5 to 2 mm. (or more) in length; 
very convex; exuvias subcentral; larval secretion whitish except as 
obscured by extraneous matter; main or subsequent secretion dull 
resinous brown; larval secretion and exuvia easily lost, exposing the 
large, bright orange colored second exuvia; scale adhering very 
loosely to the leaf and easily dislodged, leaving the insect exposed; 
inner surface of scale covered with snow-white secretion which 
entirely conceals the exuviae. 

Scale of male: Not certainly identified, but apparently the same 
general type as female, except as to size and number of exuviae. 

Adult female: Long, oval, distinctly robust or convex, 0.7 to 
0.9 mm. broad by 1 to 1.4 mm. long; hyaline in balsam mount; 
anal plate broad, rounded, in general hyaline, 0.32 mm. broad by 
0.20 mm. long; two pairs of simple normal lobes, median largest 
and most chitinized, 0.012 mm. in diameter; both lobes with two 
minute shoulders, often worn, and shoulders obscure; some distance 
beyond the second lobe is a very low, rudimentary, lobular projec- 
tion, with difficulty distinguished, terminating in a nipple or pointed 
tip; lateral teeth wanting; incisions moderate, normal; paraphyses 
practically wanting; plates lower than lobes, broad, many-branched; 
2, 2, 3, and, following the rudimentary lateral lobe, 4 or 5 narrow, 
branched plates; spines very short and inconspicuous; anal opening 
large, circular, 0.2 mm. in diameter, larger than a single median lobe 
and about two lobes distant from base of lobes; paragenitals in five 
groups, 2-4, 3-3, 3-4; dorsal pores large and conspicuous, in rows 
counting from the center as follows: 3-4, 5-7, 6-7, with a few addi- 
tional pores in obscure rows of 2, 3, or 4 pores each; basal thicken- 
ings represented by two narrow median transverse rods, and the 



14 PAPERS ON COCCID^E OE SCALE INSECTS. 

widely separated oblique lateral thickenings; ventral thickenings 
normal, but slightly chitinized. 

Type. — Bureau of Entomolog}^ No. 14137. On Ahies sp., im- 
ported by F. N. Meyer, for the Bureau of Plant Industry, from 
Wu Tai Shan, near Peking, China, received April 21, 1908. 

Note. — This insect occurs quite abundantly on the short leaves 
of the Ahies. It is closety allied to ahietis Schrank, but diverges 
from the latter in some easily recognizable sinK'taral features. The 
margin of the anal segment beyond the second rao ision pr(^?ents but 
one lobular projection instead of two, as is the cjise with abieiis; 
and, furthermore, in ahietis these projections are more p>-,,;l,i/.o,l ,= : ■ 
are finely serrate on the oblique outer margin. 

ASPIDIOTUS (AONIDIELLA) COCOTIPHAGTJS n. sp. 

(Plate II, figs. 1, 2.) 

Scale of female: Subcircular, 1.6 to 2 mm. in diameter; light yellow- 
ish brown, with a very minute central white tubercle; exuviae central, 
larval 0.35 mm. in diameter; second stage, 0.75 mm. in diameter. 

Scale of male: Similar in color to the female, oval, a little over a 
millimeter long by 0.7 mm. wide; both male and female secretions 
moderate, translucent. 

Adult female: Subcircular, 1.2 mm. in diameter; the older or 
spent females with the anal segment usually retracted, giving a kidney 
shape to the insect as in aurantii; the inflated cephalo-thoracic and 
abdominal segments, except the anal plate, are also strongly chiti- 
nized as in aurantii; anal flate ^vith basal lobules, 0.4 mm. in diame- 
ter, but slightly chitinized, rounded; median lobes largest, chitinized, 
brown, together measuring a little less than 0.04 mm.; second and 
third lobes much smaller; median lobes with two shoulders; innei 
shoulder of the second and third lobes minute, sometimes absent in 
the case of the third lobe, outer shoulders distinct; three low, serrated 
areas beyond third incision, not always apparent; the posterior point, 
however, of the first area chitinized, and sometimes assuming the 
form of a rudimentary third lateral lobe; incisions normal; paraphy- 
ses shorter than lobes, narrow, the one extending from the inner base 
of the median lobes most prominent; plates in median and first and 
second lateral incisions respectively 2, 2, 3, in length variable, sometimes 
slightly exceeding lobes, deeply forked at tips; three broad plates in 
the third incision, oblique at apex, each terminating at the posterior 
angle in a long spindle-shaped tail; anal opening about the size of 
the first lateral lobe and about two lobes' length distant from tip; 
paragenitals 0-1, 3-8, 3-5, scattering; dorsal pores in two prominent 
rows, extending respectively from the second and third incisions, of 
6 to 10 pores each; there are also some scattering lateral pores; basal 



Tech. Series 16, Part 11, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate II 




Fig. 1.— Aspidiotus (Aonidiella) cocotiphagus. 




Fig. 2.— Aspidiotus (Aonidiella) cocotiphagus. 
ANAL PLATES OF NEW SCALE INSECTS. 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate III 




Fig. 1.— Aspidiotus (Diaspidiotus) africanus. 




Fig. 2.— Aspidiotus i Diaspidiotus i courseti/e. 
ANAL PLATES OF NEW SCALE INSECTS. 



l^EW SPECIES OF DIASPINE SCALE INSECTS. ] 5 

thickenings slight, two short central rods and a larger lateral rod; 
ventral thickenings inconspicuous, represented chiefly by a chitinized 
line on each side below the first lateral incision. 

Tyfe. — Bureau of Entomology No. 14136. Massed on the upper 
surface of the leaves of cocoanut {Cocos nucifera), Santiago de las 
Vegas, Cuba, collected August, 1904, by Wm. T. Home, chief of the 
department of vegetable pathology of the central agricultural experi- 
ment station of Cuba. Also on Citrus trifoliata, collected by Mr. 
Home, Santiago de las A^egas, Cuba, January 13, 1906. 

Note. — This species seems to be an intermediate one between 
Asjndiofus (AonidieUa) aurantii and Aspidiotus (CJirysom/phalus) 
dictyospermi. It, however, differs distinctly from both of these spe- 
cies; from aurantii by notable differences in the pores, plates, and the 
presence of paragenitals, and from dictyospermi in the shape of the adult 
female and of the anal plate, and in the character and shape of the 
plates and lobes. In aurantii all the plates are distinctly subbranched 
and the three broad plates of the third incision have each two promi- 
nent apical branches which are distinctly subbranched. In dictyo- 
spermi the anal plate is much more produced and with much longer 
paraphyses, and of the three plates of the third incision the two pos- 
terior have long, spindle-shaped branches, the anterior plate two 
short branches. There is a distinct difference also in the shape of 
the lobes. 

The specific variation in point of length of plates relative to lobes, 
and size of the lobes, position of the anal opening, etc., is illustrated in 
the two photogTaphs of the anal segment shown on Plate II. The 
upper figure is from type material on cocoanut and the lower from 
trifoliate orange. 

ASPIDIOTUS (DIASPIDIOTUS) AFRICANUS n. sp. 

(Plate III, fig. 1.) 

Scale of female: Dull opaque brown, 1.5 mm. in diameter; circular, 
moderately convex, with fairly prominent central nipple, and slightly 
annulated as in perniciosus; when rubbed, resinous exuviae appear, 
and when old and massed the ashen appearance of perniciosus is 
presented. 

Scale of male: About 1 nnn. long, of normal type. 

Ad.ult female: 1 mm. long, oval. Anal plate 0.32 mm. broad at 
base by 0.2 mm. long, normal, not produced. Median lobes like 
those of ancylus, slightly converging toward tip; inner notch prac- 
tically wanting; lateral lobes wanting or rudimentary, as in the case 
of ancylus; two distinct lateral incisions; paraphyses of first incision 
heavy, subequal, inner slightly largest; paraphyses of second incision 
much smaller; minute but distinct paraphyses also are seen at the 
inner base of the median lobes; plates minute, inconspicuous; spines 



16 PAPERS ON COCCID^ OR SCALE INSECTS. 

rather strongly developed; anal opening circular, near tip; parageni- 
tals often wanting; if present, represented by from 1 to 3 glands only; 
dorsal pores few, scattering; transverse basal thickenings well devel- 
oped, laterals separated from the central line and robust; apical 
ventral chitinization rather strongly developed, brown; ventral 
longitudinal ridges feebly developed. 

Type. — Bureau of Entomology No. 14121. 

Material studied: Original lot from Bloem . — !::. South Africa, 
December, 1907, from the Hon. W. J. Palmer director of agriculture, 
Orange River Colony. Food plants repres<^i; ted; (fieditsia triacan- 
tlios, Scliinus molle, fig (two samples), almond, and nuincf^. T'.-i- 
material was abundantly parasitized, but none of the piir.isi ; 
obtainable for identification. Specimens on Gleditsia triacanih^ ■■ 
followed in description, except as to note on paragenitals. 

Additional lot from Charles P. Lounsbury, government entomolo- 
gist, Cape Colony, with letter of January 29, 1908, representing mate- 
rial on pear. Hex River, Cape Colony; Kiefl'er pear, Prospect Farm, 
Komgha (east of Cape Colony) ; fig, Paarl, Cape Colony, and fig and 
privet from Bloemfontein. 

The original lot, received through the Hon. W. J. Palmer, was sub- 
mitted under the supposition that the insect might prove to be the 
San Jose scale {Aspidiotus perniciosus Comst.), and with the request 
that an authoritative determination be cabled to Bloemfontein. The 
insect proved to be entirely distinct from perniciosus, and evidently an 
undescribed species native to South Africa. The general appearance 
of the scale and insect is strikingly close to the San Jose scale. Even 
the anal plate bears a close resemblance to the San Jose scale, but 
lacks the second pair of lobes and presents some other minor differ- 
ences. In the specimens examined of the first lot paragenitals also 
seemed to be entirely wanting, although later, in the case of three 
specimens, scattering pores were found, in some instances rudimentary. 

The fact that the insect is distinct from the San Jose scale was 
cabled to Mr. Palmer, and a letter giving details was sent. Before the 
receipt of this cablegram the entomologists of South Africa, viz, IVIr. 
Charles P. Lounsbury, Cape Colony; Claude Fuller, Natal; C. W. 
Howard, Transvaal, and H. Neethling, biologist. Orange River 
Colony, had held a conference at Bloemfontein and arrived at the con- 
clusion that the South African insect was distinct from perniciosus. 

The original information given by Mr. Palmer concerning the food 
habits of this insect is as follows: 

In tlie first place the worst infestations met with thus far occur on the mimosa 
{Acacia horrida), acacia {Robinia pseudacacia) , pepper {Schinus molle), and fig; whereas 
pear, pecan, and apple trees growing in the worst infested spots have either not been 
attacked at all, or only to a very slight extent. The distribution over such a large 
area as it has already spread shows that it must have occurred here for a period of at 



NEW SPECIES OF DIASPINE SCALE INSECTS. 17 

least several years, Iiut, in spite of careful search, no tree has been found so badly 
attacked that its life is threatened: no tree has been found to have actually succumbed 
to its attack. 

The scale is apparently kept in check by an insect enemy, although up to the present 
no parasites have actually been discovered. In other countries where the scale has 
1)ecome established, it can hardly be said to be kept down by its parasites. 

In letter of February 3, 1908, transmitting the report of the confer- 
ence of entomologists at Bloemf ontein, Mr. Neethling states that since 
the discovery of the insect it has become very destructive to some 
cultivated plants, notably the fig and privet. The list of plants which 
'' hflri bet'"ti -^ ■•■r-'i ■ '■ -nfest^given in the report referred to, includes the 
)ilowinf: 

Abundantly infested. 

- 'g Do. 

Honey locust (Gleditsia Iriacanthos ) Do. 

Pepper (Schinus molle : • Do. 

Ajaricot Sparsely infested. 

Quince Do. 

Apple - Very rarely and sparsely infested. 

Acacia horrida.. More or less abundantly infested. 

Robinia pseudacacia Sparsely infested. 

Rhus sp Do. 

Mr. Lounsbury's communication of January 29, with accompany- 
ing specimens, gives further interesting information relative to the 
insect, and considerably extends its range in South Africa, as already 
noted. He adds two new food plants for Bloemfontein, namely, 
willow and poplar. He had also found, in the material which he 
had studied, particularly in that from South Africa and more rarely 
in some of the Bloemfontein samples, scattering paragenitals. Fol- 
lowing his suggestion the writer gave a second and most careful 
scrutiny to all the material, with the results in the matter of para- 
genitals and other features indicated in the concluding notes descrip- 
tive of all the material. Mr. Lounsbury states: 

I have known the occurrence on the pear for some years and suspect that the infes- 
tation went with nursery stock sent out years ago by a certain nurseryman. But we 
have failed to find the scale on his place and have come to think that it might have 
Ijeen imported by him with French stocks before the days when we fumigated such 
and that it has since been eliminated. In our notes we have entered the species as 
"near ancylus" and "near perniciosus .'" but the possibility of it being the one at 
Bloemfontein incorrectly identified as perniciosus did not occur to me until I received 
material of the latter. I have not seen the species away from orchards or habita- 
tions, but some Orange River Colony material was found on Rhus and mimosa (native 
plants) under circumstances suggestive of its being indigenous. The pear occurrence 
at Komgha (east of Cape Colony) was on a few trees in one orchard, and cutting them 
out seems to have proved eradicative. A few branches were heavily incrusted. 
The one at Hex River (only 125 miles from here) is also in one orchard only, so far 
as I know, and here some dozens of trees show the injury by the death of inside wood 
and spotting of fruit. I can not compare the species with perniciosus for perniciousness. 



18 PAPERS ON COCCID^E OE SCALE INSECTS. 

The fact that this insect occurs on native plants, as indicated by 
Mr. Lounsbury, and also that it is rather abundantly parasitized, 
further emphasizes the probability that it is an indigenous species. 
Some descriptive notes of the different lots of material follow : 

Original material received from W. J. Palmer, Bloemfontein (letter 
of November 4, 1907) : 

(1) On Gleditsia triacanthos: Type of description. 

(2) On Sciiinus molle (pepper tree): Typical, except for a slightly 
m.ore prominent development of dorsal pores, and a much lighter 
coloring of the scale secretion. When occurring on this food plant 
the scale is light yellowish brown, almost buff', but otherwise normal. 
Such variation in coloring is sometimes seen in the San Jose scale, 
and may be due purely to the food plant. 

(3) On fig: Typical, save for a single anterior lateral circumgenital 
gland in two females out of eight examined. 

(4) On almond:. Typical, save for two rudimentary glands in one 
specimen. These glands in the specimens on fig and almond were 
not noted in the first examination. 

(5) On quince: Typical. 

Material received from Charles P. Lounsbury (letter of January 29, 
1908 V. 

(1) On pear, Plex River, Cape Colony, November 17, 1907, three 
females: Tjq^ical, except that they show scattering paragenitals, as 
follows : 
























2-1 


0-0 


0-1 


1-2 


3-2 


1-1 


0-0 


1-2 


1-1 


1-0 


1-1 


1-1 


1-1 


0-0 



Scales reddish brown, with nipple and ring as in 'perniciosus; when 
rubbed, exposing exuvia as in perniciosus. 

(2) On Kieffer pear. Prospect Farm, Komgha (east of Cape Col- 
ony), September, 1905: Scale similar to last, but with central nipple 
and ring less prominent; pygidium rather produced or triangular; 
dorsal pores numerous and prominent, arranged in two or more irreg- 
ular rows on each side, the first extending from the second incision. 
Paragenitals : 









1 








2-3 


2-4 


2-2 


2-1 


1-1 


2-2 


2-2 


2-2 


2-2 


1-2 



(3) On fig, Paarl, Cape Colony, November 8, 1907; 11 females 
examined: Typical in character of dorsal pores. Paragenitals absent 
or very few. Color of scales modified by the reddish-brown super- 
ficial layers of the bark which are carried over the scale; evidently 



XEW SPECIES OF DIASPINE SCALE INSECTS. 19 

approximating the color of the scales of the other specimens. Para- 
o-enitals : 
















0-0 (two specimens). 




2-1 


1-0 


0-] (three specimens). 


0-0 




2-1 


0-0 


0-0 















1-1 


1-0 




1-1 (two 


specimens). 


1-0 


1-3 




1-1 





(4) On fig, Bloemfontein, November 29, 1907: Substantialh^ tj])- 
ical, one specimen out of six exhibiting two paragenital glands, 
nameh", one each of posterior laterals; with the other five specimens 
no glands present. Mounted material only. 

(5) On privet, Bloemfontein, October, 1907: Tj^pical, except for 
the presence of scattering paragenitals as follows: 









1-2 


3-3 


1-1 


2-2 



Slide mount onh", showing two females. 

All of this material evidently belongs to the same species. The 
five lots originally studied show practically no paragenital glands, 
also general uniformity in dorsal pores, except in the case of speci- 
mens on pepper tree. 

In all the later material received from Mr. Lounsbury scattering 
paragenital glands are found in most of the individuals examined. 
The specimens on fig from Cape Colony and Bloemfontein show a 
general absence of these glands, or, if present, represented mosth^^ by 
single glands. In the case of the other food plants, both from Cape 
Colony and from Bloemfontein, the paragenitals are very scattering, 
with one or more groups often entirely wanting. Taking the reason- 
able supposition that all this material represents the same species — ■ 
and there seems to be no occasion to doubt this judgment — we have 
evidently a case of a species either in a transition stage, in the matter 
of the presence or absence of paragenitals, or slightly variable in this 
feature. The paragenital glands are developed only in the adult 
stage of the female, and it has long been the belief of the writer that 
they can not be given very much specific importance. They are 
always subject to considerable variation, and future studies may 
prove that, in certain instances where species have been separated 
chiefly on the presence or absence of these glands, the separation is 
not justified, as possibly illustrated in the case of latanise and camellise. 
In the matter of dorsal pores, the specimens on Kieffer pear, Cape 
Colony, show a rather unusual development, and the pygidium in the 
examples studied is rather more produced or triangular; otherwise 
the agreement is close. Some slight variation in the number of dor- 



20 PAPEES ON COOCID^ OR SCALE INSECTS. 

sal pores is seen in other specimens, notably in the case of the pepper 
specimens already referred to. This greater development in the 
number of dorsal pores is not a character of great importance, and 
much variation in this respect is seen in other species. The abund- 
ance of dorsal pores seems to be governed to some extent by climatic 
conditions, a marked increase being often noted in specimens from 
arid districts. 

The discovery of scattering paragenitals brir • this species into 
close relationship with the ancylus native to ^ ' h ^.n^erica, and 
places it in a group of species hitherto especial- charact^^-ristic of the 
New World. It is interesting to note also that a related but a much 
more strongly chitinized species, lacking paragenitals, has been re- 
cently received from Transcaspian Russia. We know too little of the 
scale-insect fauna of Asia and Africa to be able to generalize as to 
what types of species are likely to be met with. 

ASPIPIOTTJS (DIASPIDIOTUS) COTJRSETI^ n. sp. 
(Plate III, fig. 2.) 

Scale of female: 1.5 mm. in diameter, nearly circular, of medium 
density, depressed; color grayish, more or less soiled by adhering 
extraneous matter; exuviae sublateral, covered. 

Scale of m,ale: Similar, of normal shape. 

Adult female: Length less than 1 mm., of normal peg-top shape, not 
chitinized; anal 'plate slightly more chitinized than body except for 
lobes and apical and basal thickenings; median lobes only present; 
laterals obsolete as in ancylus; outer shoulder of median lobes 
prominent, inner shoulder minute, sometimes practically wanting; 
paraphyses of two incisions beyond lobes strongly developed; inner 
paraphyses of first incision terminating in a distinct oval knob lying 
at right angles to the line of the thickening; spines normal, fairly 
long; anal opening very minute and in line with the knobbed termi- 
nals of the inner paraphyses of first incision. Paragenitals : Anterior 
laterals, 1-3; posterior laterals, 0-1; dorsal pores few in number, 
inconspicuous ; transverse basal thickenings of dorsal surface in three 
parts, consisting of a long, straight central line, sometimes broken at 
middle, and two stronger lateral thickenings in the form of a double 
curve. The ventral apical divaricating chitinized bands broad and 
undivided. 

Type. — Bureau of Entomology No. 14002. On Coursetia glandu- 
losa, Hermosillo, Mexico; collected by Albert Koebele April 23 and 
24, 1897 (Koebele No. type material, 1713). 

Note. — This species is closely allied to Aspidiotus suhsimilis Ckll. 
The latter differs in having a more produced pygidium and lobes, the 
latter without inner shoulder, anus midway of first thickening, 
paragenitals absent, and in minor characters. This species is 
scarcely more than a variety of suhsimilis. 



NEW SPECIES OF DIASPINE SCALE INSECTS. 21 

ASPIDIOTUS (DIASPIDIOTTJS) TRANSCASPIENSIS n. sp. 

(Plate IV, fig. 1.) 

Scale of female: 1.5 mm. in diameter, subcircular, of a yellowish 
or buff color. Exuviae orange, exposed when rubbed or old. 

Scale of male: Of normal oval shape, similar in color to the female. 

Adult female: Broad, oval, 1 mm. in length; anal ]jlate broad, 
rounded; median lobes only represented; laterals wanting or rudi- 
mentary as in the case of ancylus; median lobes strongly chitinized, 
with very deep outer notch forming a rather distinct lobule; inner 
notch wanting; the two lateral incisions shallow, broad, and with 
very heavy, robust, bordering paraphyses or thickenings; paraphyses 
of ^irst incision particularly large, especially the inner one; of second 
incision, much smaller; plates inconspicuous; spines not especially 
developed, normal; anal opening near tip, very minute; paragenitals 
wanting; dorsal pores few, inconspicuous, scattering; median line of 
basal thickenings subhyaline, inconspicuous ; laterals narrow, well chit- 
inized; two prominent longitudinal thickenings bordering the center 
of the pygidium ; ventral thickenings normal, well chitinized. 

TyiJe. — Bureau of Entomology No. 8216. On some old dried bark 
of Po]Julus , from Transcaspian Russia, supposed to be from C. Ahnger^ 
of Ashkabad, received November, 1898. 

In its rudimentary second and third lobes this species is related 
to ancylus. The important characters are the strongly chitinized 
median lobes, with prominent external lobule and the prominent 
longitudinal thickenings, and the absence of paragenitals. In the 
latter character it falls with the South African species africanus, which, 
however, it does not otherwise closely resemble. 

ASPIDIOTUS (HEMIBERLESIA) ISTIGMM n. sp. 

(Plate IV, fig. 2.) 

Scale of female: 1.5 mm. in diameter, subcircular, elevated, more 
or less conical; exuvia covered but showing as a yellowish spot; 
larval excretion persistent as a central nipple ; color gray, often v/ith 
purplish tinge, more or less annulated with lighter. 

Scale of male: Normal shape, otherwise resembling the female. 

Adult female: 0.6 to 0.7 mm. wide by 0.9 mm. long; normal peg-top 
shape, hyaline in balsam mount; anal plate 0.30 to 0.34 mm. in 
diameter at base, rounded, nearly hyaline ; two pairs of well-developed 
lobes with sometimes a rudimentary third spear-shaped lobe ; median 
lobes much larger than second pair and strongly chitinized, brown, 
the chitinization ending abruptly at base, not extending into segment 
except slightly at inner side, giving the base a somewhat irregular 
oblique termination; median and second pairs of lobes with two 
subequal small notches; second pair of lobes, and third, if present. 



22 PAPEKS ON COCCID^ OE SCALE -IlSrSECTS. 

practically hyaline; median pair of lobes together 0.036 mm. in 
diameter; incisions normal; paraphyses nearly wanting, represented 
by mere points of chitinization; plates very little exceeding lobes, 
not very strongly developed but of the general Hemiherlesia type, two 
median narrow, two broader first notch, then three broad followed by 
two or three simple plates, all plates save last being fingered at tips; 
spines normal; anal opening large, nearly equaling in size one median 
lobe, one and one-half lobe's length distant from base of median lobes; 
paragenitals scattering, linearly arranged, 0-1, 4-7, 4-6; dorsal pores 
very few, scattering; basal thickenings not prominent, but slightly 
chitinized, divaricated ventral thickenings not extending beyond anal 
opening. 

Type. — Bureau of Entomology No. 14135, on the ground laurel 
or trailing arbutus (Epigdea repens), Chain Bridge, Ya. (near Wash- 
ington, D. C), collected by J. G. Sanders, May 6, 1906. Collected 
also on the same food plant. Sugar Grove, Ohio, by Bertha C. Hite, 
and donated by Mr. J. G. Sanders. 

Note. — This species is evidently related to latanise of Signoret, but 
is a smaller species and differs in the notable and peculiar chitiniza- 
tion of the median lobes. It presents considerable variation, some of 
the smaller or incompletely developed specimens of the last stage of 
the female showing less chitinization and approaching rather closely 
in superficial appearance to townsendi, and it might easily be deter- 
mined as this latter species. Its host-plant and occurrence in wood- 
lands indicate that it is a species native to North America. 

ASPIDIOTUS (HEMIBERXiESIA) MITCHELLI n. sp. 

(Plate V, fig. 1.) 

Scale of ■ female: Length, 1.5 mm.; subcircular to broad oval, 
strongly convex, and of the general camellix type; secretionary mat- 
ter rather dense, color dull yellowish, due chiefly to the extraneous 
matter taken up from the surface; exuviae yellowish brown, near the 
anterior end usually covered. Ventral scale a distinct white floccu- 
lent patch, thinnest at the center. 

Scale of male: Similar in general appearance to that of the female, 
but of the normal elongate shape. 

Adult female: Normal top-shaped, 0.75 mm. in diameter; in bal- 
sam hyaline; anal plate a little more yellowed than body, broad, not 
produced; three pairs of lobes; median lobes truncate, not converg- 
ing, with two lateral shoulders, separated by a lobe's width, relativelj^ 
much smaller than the lobes of cameTlise; second lobe minute, spear- 
shaped, often with outer lateral shoulder; third lobe narrow, spini- 
form; thickenings of first and second incisions present, subequal, and 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate IV. 




Fig. 1.— Aspidiotus (Diaspidiotusj transcaspiensis. 




Fig. 2.— Aspidiotus (Hemiberlesia) epig/e/e. 
ANAL PLATES OF NEW SCALE INSECTS. 



Tech. Series 16, Part II, Bureau of Entomology, U. S Dept. of Agriculture. 



Plate V. 




Fig. 1.— Aspidiotus (Hemiberlesia) mitchelli. 




Fig. 2.— Aspidiotus (Hemiberlesia) popularum. 
ANAL PLATES OF NEW SCALE INSECTS. 



NEW SPECIES OF DIASPINE SCALE INSECTS. 23 

together with lobes a Httle more yellow than the pygidium, but not 
strongly chitinized ; plates numerous, long, filiform, central ones sim- 
-pij branched; spines short, inconspicuous; basal dorsal thickenings 
not strongly developed or chitinized; anal opening much larger than 
median lobes, broad oval, about one lobe's length from tip; paragen- 
ital pores wanting; dorsal pores not numerous or conspicuous, but 
with verj^ long (one-half '■■-'dth of pygidium) internal tubes; ventral 
thickening of inte!r-:^v,' j - ^tending from median lobes practically 
^\ anting. 

Type. — Bui :.' ^mology No. 7695. On thick, narrow, 

saghlly obi ail',' • ■ t- leaves about I5 inches long; collected by Mr. 
Charles P. Lounsbury, at Mitchells Pass, South Africa, January 29, 

Note. — This species, in character of scale and last segment, is dis- 
tinctly a Hemiherlesia. The anal opening is relatively as large as in 
camellise, but the median lobes are much smaller, and the chitinization 
of the last segment is distinctly less than in camellias. The scale 
occurred scatteringly on both surfaces of the leaves. 

ASPIDIOTUS (HEMIBERLESIA) POPULARUM n. sp. 

(Plate V, fig. 2.) 

Scale of female: Length, 2 mm.; broad oval, of the camellise, type; 
strongly convex; exuviae near the anterior end, dark brown, the 
larval exuvia nearly black, normally covered; secretionary matter 
yellowish white, very dense. 

Scale of male: Length, 1 to 1.5 mm.; of the normal oval shape, with 
the larval exuvia showing prominently at one end through scanty 
secretionary covering. 

Adult female: Normal top-shaped, in balsam hyaline; L5 mm. in 
diameter; anal plate broad, rounded, tip slightly more yellowed than 
the body; median lobes large, close together, slightly converging, and 
with two distinct lateral shoulders, often lost in old worn specimens; 
second and third lobes of ancylus type or practically wanting; thick- 
enings of first and second incision prominent, subequal; plates rather 
short and inconspicuous, doubtless partly lost or reduced by age and 
use; spines normal; anus very large, broad oval, distinctly larger as a 
rule than median lobes; paragenitals in four groups, anterior groups 
3-9, posterior groups 3-8, often nearly continuous; dorsal pores very 
numerous, chiefly confined to two irregular double lateral rows extend- 
ing from the second incision, and a third inconspicuous incision; basal 
dorsal thickening linear, not especially prominent. 

Type. — Bureau of Entomology No. 8370. On cottonwood, from 
Deming, N. Mex., through Prof. T. D. A. Cockerell, February 9, 1897. 



24 PAPEES OlST COCCID^ OE SCALE INSECTS. 

Additional specimens; also on cottonwood, received from Professor 
Cockerell from Phoenix, Ariz., November 2, 1899. 

Note. — This species is very closely allied to camellise and latanise. 
In the possession of paragenital pores it comes nearest to the latter 
species. It differs notably in the great development of the dorsal 
pores, which seems to be a common characteristic of scale insects in 
very dry climates, and in the sparsity and shortness of the plates. It 
is, furthermore, a rather larger species than either of the two men- 
tioned. 

AONIDIA JUNIPERI n. sp. 

(Plate VI, fig. 1.) 

Scale of female: Thin, wdiitish, slightly tinged with yellow, 1 mm. 
in diameter, circular in outline, slightly convex. First exuvia very 
light yellow, easily lost, showing the much darker second stage as a 
circular central spot through the opening. Second exuvia very large, 
dark resinous, almost reddish, strongly chitinized and inclosing the 
adult female, which can be easily removed through the much thinner_ 
and delicate ventral skin of this stage. 

Scale of male: Unusually large, oval, as large as the second exuvia 
of the female. Coloration and texture as in female. 

Adult female: A little more than half a millimeter in diameter 
(0.54), of normal peg-top shape, not elongated nor chitinized. Anal 
plate, 0.20 mm. broad by 0.14 mm. long; scarcely chitinized, sub- 
hyaline; two small apical, nearly contiguous lobes, laterals wanting; 
lateral teeth wanting save as produced by the indentations containing 
the large peripheral dorsal pores, which form, especially toward the 
apex, distinct incisions; paraphyses and plates wanting; spines 
minute; anal opening remote from tip; paragenitals wanting; dorsal 
pores represented by a large central pore lying between the base of the 
apical lobes and five large lateral pores along the periphery of either 
side; some few other pores present, but inconspicuous ; ventral thick- 
ening inconspicuous save the uniform slight chitinization of the 
pygidium. 

TyjM. — Bureau of Entomology No. 14123. On the fruit of Juni- 
ferus sp.; from E. G. Titus, Logan, Utah. 

This species belongs to the genus Aonidia, and is ovo-viviparous, 
the bodies of the females examined being filled wnth young ready for 
emergence. 

ASPIDIOTTJS ( TARGIONIA ) CHENOPODII n. sp. 

(Plate VI, fig. 2.) 

Scale of female: 1.5 to 2 mm. in diameter, subcircular, convex; light 
buff in color, the secretion covering the larval exuvia whitish; with 
the loss of the larval exuvia, the light-orange second exuvia appears 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate VI. 




Fig. 1.— Aonidia juniperi. 





Fig. 2.— Aspidiotus iTargionia; chenopodii. 
ANAL PLATES OF NEW SCALE INSECTS. 



NEW SPECIES OF DIASPINE SCALE INSECTS. 25 

as a conspicuous spot ; ventral scale attached to the bark, wliite, 
rather abundant. 

Scale of male: Elongate, sides nearly parallel ; length 1 mm., some- 
times slightly more ; same general characters as the female, except 
that the lower secretion remains attached to the upper, forming a 
definite flatfish sac or cocoon which easily separates from the plant. 

Adult female: Nearly circular, 1.3 mm. wide by 1.5 mm. long in the 
larger specimens; hyaline, except anal plate; parastigmatic glands 
wanting. Anal plate about 0.4 mm. in diameter, normally shaped, 
but slightly chitinized; a single pair of rather contiguous narrow 
median lobes, cliitinized, brown, with a more or less distinct basal 
chitinized prolongation; lobes oblique at tip, with a produced inner 
apical angle, closely reproducing the characteristics of the same lobes 
of dearnessi; second and third lobes represented by mere projections; 
lateral teeth represented by the projections just referred to, which are 
homologous with the second and third lobes, and by a minute general 
serration of the edge of the anal plate; first incision deep and contain- 
ing two short but easily distinguishable plates; second and third 
incisions minute ; paraphyses wanting ; plates, except in first incision, 
as noted, minute or wanting; spines normal; anal opening large, oval, 
one-third from tip; paragenitals wanting; dorsal pores minute and 
inconspicuous ; basal thickenings inconspicuous or wanting ; ventral 
thickenings not showing more than the generally suffused chitiniza- 
tion of the anal plate, the longitudinal thickenings distinctly inclosing 
the oval vaginal area. 

Type. — Bureau of Entomology No. 14143. Coolabah, New South 
Wales, on CJienopodiuTn; collector, J. G. Smith. 

Note. — This species is very close structurally to dearnessi Ckll., but 
is much less chitinized. The prominent median lobes are almost ex- 
actly like those of dearnessi. The latter species^ however, has five 
prominent lateral projections instead of the two inconspicuous ones 
exhibited by cTienopodii. The dorsal pores of chenopodii are much 
less conspicuous than in the case of dearnessi; the first incision is 
deep, the plates in this incision are larger and more distinct, and there 
are other minor differences which, taken together with the wide 
separation geographically and the different food plants, perhaps 
warrant giving a specific name to the New South Wales material. 

CHIONASPIS MICROPORI n. sp. 
(Plate VII, fig. 1.) 

Scale of female: 2.5 mm. long, of the normal shape, expanding pos- 
teriorly, usually more or less curved; secretion white, smooth, and 
dense; with distinct ventral scale. 

Scale of male: About 1 mm. in length, normally shaped, with dis- 
tinct median carina. 
46797—08 2 



26 PAPERS OX COCCID^ OE SCALE INSECTS. 

Adult female (dried specimens): Dark purple, vrith the eggs or 
young also dark purple; length 0.9 to 1 mm.; breadth at mdest 
part 0.4 to 0.5 nun., of normal CJrionaspis type, tapering anteriorly; 
body nearly hyaline; parastigmal glands variable, sometimes as 
man}' as 9. Anal i)late broader than long, 0.26 by 0.38 nnn.; in 
older specimens distinctty and generally chitini?ed and brown, nearly 
hyaline in recently matured females; median lobes touching at base 
and widely divaricating toward tips; latter rounded. Inner lobe of 
first lateral pair one-half width of median; outer lobes, same pair, 
very short and much smaller; second lateral pair represented by the 
inner lobe only, which is similar to the same lobe of the first lateral 
pair ; outer lobe apparently practically wanting ; lateral lobes directed 
distinctly toward apex of segment; lateral teeth represented only 
by the projections laterad of the large oblique marginal glands; 
incisions inconspicuous; paraphyses wanting; plate3, including the 
median, 2, 2, 2, 2, 4-5; spines normal; anal opening one-third or less 
from base, circular, about the diameter of the first lateral lobe; para- 
genitals 12-13, 20-29, 18-22 ; oblique marginal pores 1, 2, 2, 2; dorsal 
pores small, nearly circular; inner row or group next to j)aragenitals 
7-9 ; irregular row extending from third incision, more or less in two 
groups, of some 15 pores altogether; followed by an exterior row of 
some 12 pores, number of pores, however, variable; basal thicken- 
ings inconspicuous or wanting; ventral thickenings the suifused 
chitinization of the anal plate already noted, slightly heavier below 
median lobes. 

Type. — Bureau of Entomology No. 14144. Collected on poplar, 
probabl}^ Populus tremula, at Wu Tai Shan, Shansi, China, by F. X. 
J\Ieyer. Received February 26, 1908. 

LETJCASPIS INDICA n. sp. 
(Plate VII, fig. 2. ) 

Scale of female: White, elongate, narrow, convex, flattened at tip, 
easily falling from the insect beneath. 

Scale of male: Not noted. 

Adult female: Entirely inclosed within the swollen, strongly chiti- 
nized second stage, latter 0.5 to 0.6 mm. long, elongate oval, dark 
reddish brown. Adult female hya^line, elongate, sides subparallel. 
Anal plate yqvj minute, about 0.1 mm. in diameter, rounded, longi- 
tudinally striate. Margin with a series of elongate plate-like proc- 
esses; lobes, incisions, and paraphyses wanting. The processes 
referred to are truncate or slightly spatulate at tip, and are of two 
lengths; the long ones are the second, fifth, and ninth from the cen- 
ter and are homologous to the lobes of other species; between the 
median long pair is a central short pair, then beyond the first long 
plate two other short ones, and beyond the second long plate three 



Tech. Series 16, Part II, Bureau of Entomology, U. S, Dept. of Agriculture. 



Plate VII. 




Fig. 1.— Chionaspis micropori. 








% 



-«r -' 






Fig. 2.— Leucaspis indica. 
ANAL PLATES OF NEW SCALE INSECTS. 



XEW SPECIES OF DIASPINE SCALE INSECTS. 27 

short ones, and bej^ond the third three short ones. The marginal 
spines are normal, and within these is a submarginal row of short 
spines; anal opening large, nearer base than tip of segment; parageni- 
tals, dorsal pores, and basal and ventral thickenings wanting. 

The 5 or 6 abdominal segments preceding the anal plate are very 
narrow, rather sharply defined, and covered with cross or longitudi- 
nal strise. 

Tiji^e. — Bureau of Entomology No. 14125. On mangoes imported 
from India, at ]\liami, Fla., collected by Mr. P. J. Wester, of the Sub- 
tropical Laboratory and Garden of the Department of Agriculture, 
and from Mayaguez, P. P., through D. W. May. 

Note. — This mango scale is a very interesting species, apparently 
undescribed, and evidently imported from India with the original 
sending of the variety of mango on which it occurs, namely, Seed 
and Plant Introduction and Distribution No. 7108, the Sundershah. 
It seems to belong to the genus Leucaspis, but the failure to find the 
male scale leaves this reference in some little doubt. It is certainly 
unlike any described species of this genus. It occurs thickly massed 
in the cracks of the bark, and the reddish inflated skins are the sec- 
ond stage, which include the adult insect in a sort of sac, and look like 
minute eggs or seeds, the whitish waxy covering adhering very 
loosely and in many cases having been lost off. It has every appear- 
ance of being a serious pest, from the dense infestation exhibited. 

MYTILASPIS CHILOPSIDIS n. sp. 

(Plate VIII, fig. 1.) 

Scale of female: More than a milhmeter in length, narrow, light 
purplish in color. 

Scale of male: Similar, about one-half the size of the female^ 
Adult female: A little more than a millimeter long, of normal 
Chionaspis shape, expanding notably posteriorly; anterior and 
lateral margins smooth, normal. Anal plate much broader than long^ 
and but slightly chitinized. Lobes hmited to one prominent median 
pair. Laterals obsolete or represented by shght tooth-like projec- 
tions. Margin of pygidium somewhat toothed, caused by the pro- 
jections over each of the large marginal oval pores. Incisions 
inconspicuous; j^araphyses wanting. Plates: 2 median followed on 
either side by a group of 4 or 5, then 2 or .3; plates somewhat longer 
than lobes. Spines normal; anal opening very near base of segment. 
Paragenitals : Anterior 5; anterior laterals 7 to 10; posterior laterals 
7 to 9 (from examination of two females). Large marginal pores 
occurring in three pairs, two each, with a single pair near the base 
of the segment. Large scattering circular or slightly oval pores over 
the general area of the pygidium. Basal tliickenings wanting; ventral 



28 PAPERS ON COCCID.E OE SCALE IK^SECTS. 

thickenings limited to a slighth" cliitinized band about t%\dce the width 
of the median lobes, extending slightly beyond the anal opening. 

Tyi)e. — Bureau of Entomology No. 7218. Scatteringly infesting a 
small section of a plant doubtfulh^ identified as CJiilopsis linearis; 
collected by C. H. T. TowTisend in Tehuantepec City, Mexico, May 
26, 1896. 

Note. — The females of tliis insect were gravid with 3'oung at time 
of collection. The distinguishing feature is the absence of the lateral 
lobes in connection ^^^th the rather large prominent median pair. 

PARLATORIA MANGIFER.ffi n. sp. 

(Plate YIII, fig. 2.) 

Scale of female: Moderateh' dense, dark brown, lighter at the edges, 
oval in outline, usually with slight apical extension; larval exuvia 
dull greenish. Length 1.5 mm.; breadth 1 to 1.25 mm. 

Scale of male: Oval, slightly lighter colored, 1 to 1.3 mm. long by 
.about 0.8 mm. broad. Exuvia more distinctly and purely green than 
in the case of the female, forming quite a striking contrast ^\T.th the 
dull brown of the secretion. 

Adult female: 0.65 mm. by 0.75 mm.; of normal rounded, peg-top 
shape; hj^aline, save the anal plate. A7ial plate 0.45 mm. broad at 
base by 0.25 mm. long; rounded, rather strongly chitinized through- 
out median area and marginally; general structure very similar to per- 
gandei. Lobes of the pergandei type, rather more chitinized. Three 
pairs of lobes with strong outer shoulder, median lobes ^\dth minute 
inner shoulder also; fourth and fifth lobes triangular, terminating in a 
point as in pergandei; lateral teeth represented b}^ irregular serrations 
beyond lobes ; incisions normal ; paraphyses represented by the semi- 
lunar chitinization at the base of the lobes, unusually heavy; plates 
resembling those oi*pergandei in general, i. e., two narrow central 
plates, and two narrow plates in the first and three in the second 
incision, posterior one minute; the third incision bears three broad, 
triangular plates, serrated on the exterior margin; follo\\'ing the 
fourth lobe are four or five similar exteriorl}" serrated plates; spines 
normal; anal opening narrow, halfway between tip of segment and 
vaginal orifice; paragenitals wanting; parastigmals represented by a 
group of 8 to 10 pores on the inner side of each stigma; dorsal pores 
large and numerous, represented by 5 more or less regular rows of 
pores on either side of the center; the central row 3 to 4, the second row 
5 to 6, third 8, fourth and fifth scattering. Marginal oval pores de- 
creasing in size from tij^ to base of segment; basal thickemngs practi- 
cally wanting; ventral tliickenings represented by a broad suffused 
chitinization extending from the tip to the base of segment, and more 
or less alono- the marsin. 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate VIII. 




Fig. 1 .— Mytilaspis chilopsidis. 



,(llj 


1^ 




W^^ 


k. 


1 



Fig. 2.— Parlatoria mangifer/e. 
ANAL PLATES OF NEW SCALE INSECTS. 



NEW SPECIES OF DIASPINE SCALE INSECTS. 29 

Type. — Bureau of Entomology No. 14128. On mango; collected 
in the Department greenhouses, Washington, D. C, Januar}^ 28, 1908, 
by J. G. Sanders, in course of quarantine inspections. Dr. L. Reh, 
of Hamburg, sent this species for determination June 27, 1904, as 
from Singapore. 

Note, — This species is a strongly chitinized pergandei lacking para- 
genital pores and exliibiting a much greater development of dorsal 
pores. In this last feature it resembles some of the varieties of per- 
gandei. It is further distinguished by the more produced, tapering, 
exteriorly serrated lateral plates occurring in the incision posterior to 
the fourth lobe, and also on the margin of the segment anterior to this 
lobe. The large number of small parastigmatic pores is also a marked 
character. 

It occurs massed as thickly as may be on the young mango twigs, 
and evinces the prolificness characteristic of the genus. It is un- 
doubtedly an offshoot of pergandei, and may not deserve more than 
varietal status. 

PARLATORIA PYRI n. sp. 

(Plate IX, fig. 1.) 

Scale of female: 1 mm. to 1.25 mm. in length, oval; exuviae of larval 
and second stage and supplementary secretion normal to the genus, 
the secretion, however, usually terminal, but its position varies some- 
what as affected by irregularities of the bark, sometimes sublateral 
and sometimes more or less inclosing the posterior half at least of the 
second exuvia. Larval exuvia purplish-green; second exuvia dark 
olive, almost black, more or less overlaid with a grayish secretion; 
supplement yellowish. 

Scale of male: Larval exuvia as in female; supplemental area 
broad, circular. 

Adult female: Of the normal peg-top shape, 0.8 mm. long by 0.6 
mm. broad, hyaline. Anal plate distinctly triangular, very slightly 
chitinized; two pairs of lobes, compressed or close together, the 
median more than twice the size of the laterals, and both pairs with a 
deep exterior shoulder; second pair of lobes much lower than median 
pair; edge of pygidium beyond plates, irregularly obtusely serrated; 
incisions shallow and inconspicuous ; paraphyses wanting, or indicated 
by a small point of chitinization at inner end of first and second lateral 
marginal pores; plates short and inconspicuous, not exceeding lobes, 
and extending a short distance laterally beyond lobes; spines normal; 
anal opening fairly large, circular, subcentral or nearer base than tip 
of pygidium. Paragenitals: Anterior group 1 to 4, anterior laterals 
8 to 10, posterior laterals 8 to 12. The large oval marginal pores 
characteristic of the genus are present, 5 to 6 on either margin of the 
anal plate, with one central one between the median lobes, continued 



30 PAPEES ON COCCID.E OE SCALE INSECTS. 

also, somewhat smaller, on all the abdominal segments, with, on these 
segments and also to a less extent on the pygidium, numerous smaller 
oval submarginal pores. Basal and ventral thickenings practically 
wanting. 

Type. — Bureau of Entomology No. 14115. On pear, Chefoo, China, 
collected October 4, 1901, by the writer. Also infesting cuttings of 
apple and soft pear, received from the Province of Liaou Yang, Man- 
churia, through F. N. Meyer (S. P. I. & D. Nos. apple, 20276, 20280; 
soft pear, 20244 and 20280). Cohected by Mr. J. G. Sanders m the 
course of inspection of Department importations, March 21, 1907. 

Note. — In the case of the Meyer importations this insect was 
found very scatteringly present on the cuttings of pear and apple 
referred to, principally the former, and was associated with the San 
Jose scale, which was even more sparsely represented. Curiously 
enough, also, this new ParlaAoria bears a close superficial resemblance 
in the balsam mounted female to the San Jose scale, and partic- 
ularly in the shape of the anal segment and of the produced tip, 
with . two pairs of compressed or closely placed lobes, which are 
almost of the identical size and shape of jjerniciosus. The plates 
also are just about the same size and extent as in perniciosus. While 
distinctly a Parlatoria, this species lacks the abundant and striking- 
development of the plates wdiich is usually characteristic of the 
genus. Its occurrence w^ith the San Jose scale and in somewhat 
greater numbers even than the latter species, and particularly its 
habitat in the North Temperate region, would indicate, in connection 
with its host relations, that it has very great possibilities for evil 
should it gain foothold on this continent. The genus Parlatoria is 
noted for its enormous powers of multiplication, and the possibilities 
of damage as seen in the case of the chaff scale {Parlatoria pergandei 
Comst.) and the date-palm scale {Parlatoria hlancliardi Targ.) indi- 
cate that this Manchurian Parlatoria might develop into a much more 
dangerous insect even than the San Jose scale. Wliile no parasitism 
was noted in the case of the Parlatoria, some of the San Jose scales 
associated with it had been parasitized, the parasite, however, not 
being identifiable from the single fragment found, which Avas, how- 
ever, referred by Doctor HoAvard to the genus Aplielinus. 

PARLATORIA CHINENSIS, n. sp. 

(Plate IX, fig. 2.) 

Scale of female: 1 mm. in diameter or less, subcircular, exuvise 
eccentric, dark metallic olive, similar in general characters to P. 
pyri. 

Scale of male: Not seen. 

Adult female: About 0.6 mm. broad by 0.7 mm. long (probably old 
gravid female considerably larger) ; normally shaped. Anal plate 



Tech. Series 16, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate IX. 




Fig. 1.— Parlatoria pyri. 




Fig. 2.— Parlatoria chinensis. 
ANAL PLATES OF NEW SCALE INSECTS. 



I 



XEW SPECIES CF DIASPIXE SCALE I]S:SECTS. 31 

broad, triangular; widtli 0.3 mm., Isiigtli 0.24 mm.; more chitinized 
than i^yri: lobes in two pairs; median large, each 0.15 mm. broad, 
obtusely pointed; inner margin with one notch, outer margin oblique, 
with four or more fine teeth; second lobe much smaller and lower than 
median, close to the latter, outer margin oblique, with 2 to 4 minute 
teeth; incisions normal; paraphj^ses represented b}' two rounded, 
loiobbed chitinous processes attached to the inner end of each of the 
first and second lateral marginal lunar pores, bringing them at or near 
the outer bases respectively of the median and lateral lobes; plates 
inconspicuous; between the median lobes and in the' first lateral 
incision two very short, scarcely discernible simple plates; three 
slightly longer simple plates occur in the second incision; be3^ond the 
second incision no plates easih^ discernible; spines normal; anal 
opening subcentral, situated just in front of vaginal opening, equaling 
in diameter width of median lobe; paragenitals in four groups of 
about 6 pores each, the anterior group apparentl}^ iiot represented in 
the specimens examined; basal thickenings a distinct narrow line, 
oblique lateral portion of same character as central \nd scarcely sepa- 
rated from latter; ventral thickenings diffused, normal. 

Type. — ^Bureau of Entomology No. 14138. On crab-apple, Tien- 
tsin, China, October 11, 1901; collector, C. L. Marlatt. 

Note. — This anomalous Parlatoria was first collected by the writer 
on a small tree, possibly a crab-apple, growing near the railway station 
at Taira, Japan, August 30, 1901. This tree was thickly infested, 
and this was the only example of this insect found in Japan. In 
China the writer collected it at Chefoo on Hihiscus (October 4, 1901), 
and later, October 11, on crab-apple at Tientsin. The latter lot has 
been indicated as the type material. What is evidently this insect 
was also collected by the writer (March 12, 1902), in Cairo, Egypt, 
on an unknown plant, which it infested in conjunction with Parlatoria 
affinis Newst. The Egyptian variety differs slightly in the exterior 
serration of the median lobes, these serrations or notches being re- 
duced to two or three instead of four or more, and the central lobular 
projection is somewhat larger. The paragenitals also are less abund- 
ant, ranging as follows: 0, 0-3, and 1-2. 

The same insect was found on- plants imported for the Depart- 
ment of Agriculture by F. N. Meyer, from Peking and northern 
China, namely, on Xanthoxylon, received May 2, 1907, on Thuja 
orientalis, received March, 1908, and on jujube, received April 21, 
1908. From these records it is evident that it is a common and 
widespread species in northern China, and probably will be found to 
occur throughout eastern Asia. The Egyptian variety may have 
been a recent importation, or the range of the species may be much 
wider in the Old World than that indicated. 

This insect is remarkably like Parlatoria pyri in general appear- 
ance and characters, and is very likely closelj" related to the latter 



32 PAPERS Ojs^ coccid.^ oe scale ixsects. 

species, perhaps presenting merely a variety. It differs, however, 
very notably in the character of the median lobes, which in pyri 
are of the normal Parlatoria type, with a deep prominent outer 
shoulder on each of the lobes. It also differs from JW^' i^ ^^^^ pres- 
ence of the chitinized processes or paraphyses, which in pyri are 
usually wanting or only faintly indicated, and in the practical absence 
of the smaller pores, which in pyri are found rather abundantly 
near the margin of the anal and other segments. The anal plate is a 
little more heavily chitinized, thus bringing out more prominently 
the basal and ventral thickenings. It occurred very scatteringly, 
altogether but four females being secured. Like pyri, the tip, at 
a superficial glance, reminds one strongly of perniciosus, the effect 
being produced by the very much smaller, lower, and closely placed 
second lobes. This species diverges still farther than pyri from 
the normal Parlatoria characters. The sparsity of plates and the 
practical absence of the branched plates along the margin of the anal 
and other segments, and the sparsity of submarginal and marginal 
pores, are all features which differentiate it from the normal type of 
Parlatoria. The broad lunar marginal pores of the anal segment, 
however, are distinctly of the Parlatoria character, and the secretions 
and exuvias are distinctly those of Parlatoria. 



o 



3 



T) 



Technical Series, No. 16, Part III. 

U. S. DEPARTMENT OF AGRICULTURE, 

BXJRE^TJ OW E:iSrTOM:OL,OGS-Y. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



PAPERS ON COCCIDjE OR SCALE INSECTS. 



CATALOGUE 



RECENTLY DESCRIBED COCCID^-II. 



By J. G. SANDERS, M. A., 

Assista7it. 



Issued December 22, 1909. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE, 

1909. 
C/ 



U..S.SUPT. OFDOf 



CONTENTS. 



Subfamily Monophlebinae.. 

Subfamily Margarodinse 

Subfamily Ortheziinae.-. . . . 
Subfamily Conchaspinse . . 

Subfamily Dactylopinae 

Subfamily Tachardiinae 

Subfamily Coccinae 

Subfamily Diaspinae 

Abbreviations of literature. 



Page. 
33 
34 
35 
35 
35 
43 
43 
48 
59 



U. S. D. A., B. E. Tech. Ser. 16, Pt. III. Issued December 22, 1909. 

PAPERS ON COCCIDii OR SCALE INSECTS. 



CATALOGUE OF RECENTLY DESCRIBED COCCIDiE — II. 

By J. G. Sanders, M. A., Assistant. 

The preparation of a catalogue of this character requires great 
vigilance and constant attention to the voluminous literature of 
entomology, in order that every reference to Coccidse be noted. 
Descriptions of new genera and species appear in all conceivable 
publications in many languages, but the publication of the results 
of technical entomological research in botanical journals or other 
periodicals which do not usually treat of entomological subjects 
can not be too strongly condemned. For decades the publication of 
names for new species without description, nomina nud^., has been 
decried. Still there are entomologists who indulge in this unfortu- 
nate practice, thereby leading to confusion and error. 

In this catalogue, the second of the series, "■ are included references 
to 24 new genera, 195 new species, and 14 new varieties. The writer 
believes the list quite complete to March, 1909, and asks the assist- 
ance of coworkers in adding references which may have been over- 
looked. 

Subfamily MONOPHLEBINiE. 

Monophlebus stebbingi octocaudata Green. 

Monophlebus stebbingi var. octocaudata Green, Mem. Dep. Ag. India, ii, 2, p. 16 (1908). 

Maxwell-Lefroy, Mem. Dep. Ag. India, ii, 7, 
p. Ill (1908). Fig. 
Habitat — Lahore, India. 

On mango; Ficus bengalensis; F. infectoria; F. religiosa; F. glomerata; F. carica; 
Dalbergia sissu; Artocarpus integrifolia. 

Monophlebus tamarindus Green. 

Monophlebus tamarindus Green, Mem. Dep. Ag. India, ii, 2, p. 17 (1908). 
Habitat — India . 
On branches of tamarind. 

«The first catalogue of the series was published as Technical Series 12, Pt. I, of 
this Bureau, and was issued June 5, 1906. 

33 



34 PAPEKS ON COCCID^/OR SCALE INSECTS. 

PalsBococcus pulcher Leonard!. 

Palseococcus pulcher Leon., Annali di Agr., vii, p. 1 (1907). Fig. 
Habitat — ^Java. 
On leaves of Ilex sp. 

Palaeococcus rosae australis (Maskell). 

Icerya rosse var. australis Mask., N. Z. Trans., p. 101 (1893). Fig. 
Habitat — Australia. 
On Hakea gibhosa. 

Palseococcus theobromse Newstead. 

Palseococcus theobromse Newst., Jn. Econ. Biol., ii, 4, p. 154 (1908). 
Habitat — Calabar, West Africa. 
On leaves of cultivated cacao. 

Walkeriana cinerea Green. 

Walkeriana cinerea Green (sine descr.), Mem. Dep. Ag. India, ii, 2, p. 18 (1908). 

Maxwell-Lefroy (sine descr.), Mefii. Dep. Ag. India, ii, 7, p. 121 
(1908). 
Habitat — India. 
On Acacia arabica; Laivsonia alba. 

Icerya minor Green. 

Icerya minor Green, Mem. Dep. Ag. India, ii, 2, p. 17 (1908). Fig. 

Maxwell-Lefroy, Mem. Dep. Ag. India, ii, 7, p. 118 (1908). Fig. 
Lec.chia /estiva Kieffer, Marcellia, vii, p. 163 (1909). Fig. 
Habitat — Bengal. 

On mango. 

a Icerya okadae Kuwana. 

Icerya ohadx Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 178 (li907). Fig. 
Ckll., Can. Ent., xli, 2, p. 55 (1909). 
Habitat — Japan. 
On orange tree (Citrus). 

Icerya seychellarum cristata Newstead. 

Icerya seychellarum var. cristata Newst., Liverpool Univ. Quart. Jn., iii, 6, p. 11 (1908). 
Habitat — Madagascar. 

Subfamily MARGARODIN^a:. 

Margarodes raediterraneus Silvestri. 

Margarodes mediterraneus Silv., Bui. Soc. Ent. Ital., xxxviii, p. 140 (1906). Fig. 

Leon., Cherm. Ital., Ease, iv. No. 77 (1908). 
Habitat— Italy. 
On roots of Cynodon sp. 

Genus MATSTJCOCCTJS Cockerell. Type, matsumurx. 

Matsucoccus Ckll., Can. Ent., xli, 2, p. 56 (1909). 

To include Xylococcus matsumurse Kuw., Insect World, ix, 3, March (1905). 
Fig. ; Sanders, Catalogue Coccidae (1906); Kuwana, Bui. Ag. Exp. Sta. Japan, 
I, 2, p. 209 (1907). Fig. 

oThis species is a synonym of Icerya seychellarum, (Westwood), which had been 
previously discovered in China, Formosa, etc. 



CATALOGUE OF RECENTLY DESCRIBED COCCID^ II. 35 

Genus STEINGELIA Nassonow. Type, gorodetskia. 

Steingelia Nassonow, Ann. Mus. Zool. Imp. Acad. Sc. St. Petersbourg, xiii, p. 345 
(1908). 
Ckll., Can. Ent., xli, 2, p. 56 (1909). 

Steingelia gorodetskia Nassonow. 

Steingelia gorodetskia Nassonow, Ann. Mus. Zool. Imp. Acad. Sc. St. Petersbourg, 
XIII, p. 345 (1908). 
Ckll., Can. Ent., xli, 2, p. 56 (1909). Fig. 
Habitat — Russia. 

Xylococcus macrocarpae Coleman. 

Xylococcus macrocarpfe Coleman, Jn. N. Y. Ent. Soc, xvi, p. 198 (1908). Fig. 
Habitat — California . 
On Cupressus macrocarpa. 

Subfamily ORTHEZIINiE. 

Orthezia calif omica Ehrhorn. 

Orthezia californica Ehrh., Can. Ent., xxxviii, p. 329 (1906). 
Habitat — California. 
On Bahia sp. 

Orthezia martelli Leonardi. 

Orthezia martelli Leon., Bol. Zool. Sc. Sup. Portici, in, p. 150 (1908). Fig. 
Habitat — Italy. 
On grasses (Graminacesp). 

Subfamily CONCHASPINiE. 

Genus FAGISUGA Lindinger. Type, triloba. 

Fagisuga Lindgr., Zeitschr. f. wiss. InsektenbioL, v, 4, p. 107 (1909). 

Fagisuga triloba Lindinger. 

Fagisuga triloba Lindgr., Berl. Ent. Zeitschr., iv, 5, p. 107 (1909). Fig. 
Habitat^Chile. 
On Nothofagus dombeyi. 

Subfamily DACTYLOPIINiE. 

Asterolecanium lineare Lindinger. 

Asterolecanium lineare Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. — (1908). 
Habitat — Brazil . 
On cocoanut. 

Asterolecanium. railiaris robusta Green. 

Asterolecanium miliaris var. robusta Green, Mem. Dep. Ag. India, n, 2, p. 19 (1908). 

Maxwell-Lefroy, Mem. Dep. Ag. India, ii, 7, p. 
121 (1908). 
Habitat — B engal . 
On stems of bamboo {Dendrocalamus sp.). 



36 PAPEKS ON COCCID^ OR SCALE INSECTS. 

Lecaniodiaspis baculifera Leonardi. 

Lecaniodiaspis baculifera Leon., Annali di Agr., vii, p. 11 (1907). Fig. 
Habitat — Java. 
On an undetermined plant. 

Lecaniodiaspis frenchii Green. 

Lecaniodiaspis frenchii "Green" Frencli (sine descr.), Victorian Nat., xxni, pp. 184, 
240 (1907). 
Habitat — Victoria, Australia. 

« Genus CERCOCOCCUS Scott. Tjj>e eremobius. 

Cercococcus Scott, Tr. Linn. Soc. Lond., 2d ser., ix, 12, p. 455 (1907). Fig. 
Cerococcus Green, Ent. Mo. Mag., xix, p. 42 (1908). 

Cerococcus eremobius (Scott). 

Cercococcus eremobius Scott, Trans. Linn. Soc. Lond., 2d ser., ix, 12, p. 455 (1907). 

Fig. 
Cerococcus eremobius Green, Ent. Mo. Mag., xix, p. 42 (1908). 
Habitat — Algeria. 
On Helianthemum hahiricum,. 

Cerococcus hibisci Green. 

Cerococcus hibisci Green, Mem. Dep. Ag. India, ii, 2, p. 19 (1908). Fig. 

Maxwell-Lefroy, Mem. Dep. Ag. India, ii, 7, pp. 122, 135 (1908). 
Habitat — Bombay, India. 
On branches of Hibiscus liliflorus and Gossypium, sp. 

Solenococcus muratae (Kuwana). 

Cerococcus muratse Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 180 (1907). Fig. 
iSolenophora muratae GklL, Can. Ent., xu, 2, p. 55 (1909). 

Habitat — Japan. 

On grape; Viburnum odoratissimum. 

Kermes bacciformis Leonardi. 

Kermes bacciformis Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 156 (1908). Fig. 
Habitat — Italy. 
On Quercus suber. 

Kermes himalayensis Green. 

Kermes himalayensis Green, Ent. Mo. Mag., xx, p. 1 (1909). Fig. 
Habitat — Northern India. 
On Quercus incana. 

Kermes miyasakii Kuwana. 

Kermes miyasalii Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 181 (1907). Fig. 
Habitat — Japan . 
On Quercus serrata. 

Kermes rattani Ehrhorn. 

Kermes rattani Ehrh., Can. Ent., xxxvui, p. 329 (1906). 
Habitat — California. 
On Quercus chrysolepis. 

a A synonym of Cerococcus, having been described from immature specimens. 



CATALOGUE OF RECENTLY DESCRIBED COCCID^ II. 37 

Kermes variegatus corticalis Nassonow. 

Kermes variegatus corticalis Nass., Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg, xiii, 
p. 490 (1909). Fig. 
Habitat — Russia . 
On bark of Quercus sp. 

Kermes vastus Kuwana. 

Kermes vastus Kuwana, Bui. Agr. Exp. Sta. Japan, i, 2, p. 181 (1907). Fig. 
Habitat — Japan. 
On Quercus glandulifera. 

Genus ATRIPLICIA Cockerell. Type gallicola. 

Atriplida Ckll., Pr. Ent. Soc. Wash., x, p. 169 (1909). 

Atriplicia gallicola Cockerell & Rohwer. 

Atriplida gallicola Ckll. & Rohw., Pr. Ent. Soc. Wash., x, p. 169 (1909). 
Habitat — Colorado; New Mexico. 
On leaves of Atriplex canescens. 

Eriococcus bahise Ehrhorn. 

Eriococcus bahise Ehrh., Can. Ent., xxxviii, p. 330 (1906). 
Habitat — California. 
On roots of Bahia sp . 

Eriococcus bezzii Leonardi. 

Eriococcus bezzii Leon., Bol. Zool. Sc. Sup. Portici, i, p. 148 (1907). Fig. 
Cherm. Ital., Fasc. iv, No. 83 (1908). 
Habitat — Italy. 
On branches and petioles of Rhododendron ferrugineum. 

Eriococcus catalinee Ehrhorn. 

Eriococcus catalinse Ehrh., Can. Ent., xxxviii, p. 332 (1906). 
Habitat — Cali f ornia . 
On Artemisia sp. 

Eriococcus howardi Ehrhorn. 

Eriococcus howardi Ehrh., Can. Ent., xxxviii, p. 831 (1906). 
Habitat — California. 
On Quercus sp. 

Eriococcus lagerstrcemiae Kuwana. 

Eriococcus lagerstrcemise Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 182 (1907). Fig. 
Habitat — Japan . 
On Ficus carica; Lagerstroemia indica. 

Eriococcus latialis Leonardi. 

Eriococcus latialis Leon., Bol. Zool. Sc. Sup. Portici, i, p. 144 (1907). 
Habitat — Italy. 
On an undetermined plant. 

Eriococcus quercus gUensis Cockerell. 

Eriococcus quercus gilensis Ckll., Pr. Ent. Soc. W^ash., x, p. 167 (1909). 
Habitat — Arizona. 
On Quercus toumeyi. 



38 PAPEES ON COCCID^ OR SCALE INSECTS. 

Genus LEFROYIA Green. Type castanex. 

Lefroyia Green, Mem. Dep. Ag. India, n, 2, p. 21 (1908). 

Lefroyia castaneae Green. 

Lefroyia castanex Green, Mem. Dep. Ag. India, ii, 2, p. 21 (1908). Fig. 
Habitat — Assam. 
On smaller branches of Castanea sp. 

Stictococcus mviltispinosus Newstead. 

Stictococcus multispinosus Newst., Jn. Econ. Biol., ii, 4, p. 150 (1908). 
Habitat — East Africa. 

Dactylopius argentinus Dominguez. 

Dactylopius argentinus Dominguez, Trab. Mus. Farm. Fac. Cienc. Med., no. 17 (1907). 
Dominguez, Bol. Min. Ag., vn, 3, pp. 148, 150 (1907). 
Habitat — Argentina. 
On Cactacese (Opuntia Jicus-indica and 0. aurantiaca). 

Dactylopius ceylonicus (Green). 

Coccus cacti var. ceylonicus Green, Ind. Mus. Notes, iv, 1, p. 7 (1896). 

Mask., N. Z. Trans., xxix, p. 319 (1897). 
Coccus indicus Green, Mem. Dep. Ag. India, ii, 2, p. 28 (1908). Fig. 

Habitat — Ceylon; India. 

On Opuntia dillenii. 

a Spliserococcus draperi Newstead. 

Sphxrococcus draperi Newst., Liverpool Univ. Quart. Jn., i, 2, p. 70 (1906); Agr. News 
West Ind., v, 3, p. — (1906), 
Draper, Scale Insects of Egypt, p. 12 (1907). 
Habitat — Egypt . 
On date palm. 

Genus MACROCEROCOCCUS Leonardi. Type, superhus. 

Macrocerococcus Leon., Bol. Zool. Sc. Sup. Portici, i, p. 151 (1907). 

Macrocerococcus superbus Leonardi. 

Macrocerococcus superhus Leon., Bol. Zool. Sc. Sup. Portici, i, p. 152 (1907). Fig. 

Marchal, Compt. Rend. Ac. Sci., Paris, cxliii, 13, p. 872 
(1909). 
Habitat — Sardinia; Corsica; France. 

On stems of an undetermined grass, and on Lepidium draba; Alyssum mediter- 
raneum; Lotus; Fumaria major; Cetarach officinalis; Antirrhinum latifolium. 

Phenacoccus cholodkovskyi Marchal. 

Phenacoccus cholodkovskyi Marchal, Ann. Soc. Ent. France, lxxvii, p. 245 (1908). Fig. 
Habitat — Southern Russia. 
On Triticum vulgar e. 

a This species must be very close to Phoenicococcus marlatti Ckll., a very common 
pest of the date palm in Algeria, if it is not indeed identical. The description answers 
very well for the latter species. 



CATALOGUE OF KECENTLY DESCEIBED COCCID^ II. 39 

Phenacoccus colemani Ehrhorn. 

Phenacoccus colemani Ehrh., Can. Ent., xxxviii, p. 332 (1906). 
Habitat — California. 
On Bubus sp. 

Phenacoccus formicarum Leonardi. 

Phenacoccus formicarum Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 162 (1908). Fig. 
Habitat — Italy. 
In nest of ants (Pheidole pallidula). 

Phenacoccus graminicola Leonardi. 

Phenacoccus graminicola Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 160 (1908). Fig. 
Habitat — Italy . 
On grasses (Graminacege). 

Phenacoccus hirsutus Green. 

Phenacoccus hirsutus Green, Mem. Dep. Ag. India, ii, 2, p. 25 (1908). Fig. 
Habitat — India ; Tasmania . 

"On terminal shoots of an undetermined shrub, attended by ants (Creviastogaster 
rogenhoferi) , which had constructed shelters around the colonies of insects." 

Phenacoccus iceryoides Green. 

Phenacoccus iceryoides Green, Mem. Dep. Ag. India, ii, 2, p. 26 (1908). 

Maxwell-Lefroy, Mem. Dep. Ag. India, n, 7, p. 129 (1908). 
Habitat — Calcutta, Tanjore, Surat, India. 
On mango; Boswellia; Capparis horrida. 

Phenacoccus insolitus Green. 

Phenacoccus insolitus Green, Mem. Dep. Ag. India, ii, 2, p. 26 (1908). Fig. 
Habitat — Bengal. 
On Sida cordifolia. 

a Phenacoccus ramonae Essig. 

Phenacoccus ramonse Essig, Pomona Jn. Ent., i, 2, p. 44 (1909). Fig. 
Habitat — California . 
On roots of black sage (Ramona stachyoides) . 

Genus COCCTJRA Sulc. Type, comari Kunow. 

Coccura Sulc, Nachrichtsbl. Naturfrsch.-Klubs, Prossnitz, x, 7 pp. (1907); Lindgr., 
Zeitschr. f. wiss. InsektenbioL, iv, 12, p. 475 (1908). 
A new genus for the reception of Coccus {Phenacoccus) comari Kunow, Ent. Nach- 
richten, vi, p. 46 (1880). 

Ceroputo voljmicus Nassonow. 

Ceroputo volynicus Nass., Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg, xiii, p. 471 
(1909). Fig. 

Habitat — Russia . 

On leaves and stems of Dactylis glomerata. 



a This ia undoubtedly an immature stage of Ceroputo yuccsp (Coquillett). The 
lateral patches of spines on each segment, and the incompletely developed 7-iointed 
antennae indicate strongly its connection with the latter species, 
12600— No. 16, pt 3— C9 2 



40 PAPERS ON COCCID^ OR SCALE INSECTS. 

Genus LACHNODIELLA von Ihering. Type, cecropise. 

Lachnodiella von Iher. (sine descr.), Bot. Jahrb., xxxix, 5, p. 680 (1907). 

Lachnodiella cecropise von Ihering. 

Lachnodiella cecropise von Iher. (sine descr.), Bot. Jahrb., xxxix, 5, p. 680 (1907). 
Habitat — Brazil . 
On Cecropia adenopus. 

Pseudococcus agrifolise Essig. 

Pseudococcus agrifolise Essig, Pomona Jn. Ent., i, 2, p. 42 (1909). Fig. 
Habitat — California . 
Beneath bark and in cracks of bark of Quercus agrifolia. 

Pseudococcus artemisiae Essig. 

Pseudococcus artemisise Essig, Pomona Jn. Ent., i, 2, p. 38 (1909). Fig. 
Habitat — California. 
"Under bark of Artemisia calif ornica." 

Pseudococcus citri coleorum Marchal. 

Pseudococcus citri var. coleorum Marchal, Ann. Soc. Ent. France, lxxvii, p. 236 (1908). 

Habitat — France . 

On Coleus. 

Pseudococcus coffeae (Newstead). 

Dactylopius n. sp. Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, p. 74 (1906). 
coffese Newst., Jn. Econ. Biol., m, 2, p. 37 (1908). Fig. 
Habitat — Java. 
On Liberian coffee. 

Pseudococcus cupressi Coleman. 

Pseudococcus cupressi Coleman, Jn. N. Y. Ent. Soc, xvi, p. 197 (1908). Fig. 
Habitat — California. 
On Cupressus macrocarpa. 

Pseudococcus (?) cycliger Leonard!. 

Pseudococcus cycliger Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 168 (1908). Fig. 
Habitat — Italy. 
In nest of ants {Aphenogaster testaceo-pilosus) . 

Pseudococcus elongatus Reuter. 

(t Pseudococcus graminis Reuter, Medd. Soc. Faun. Fenn., p. 66 (1904). 

elongatus Reuter, Medd. Soc. Faun. Fenn., p. 251 (1904). 

elongatus Sanders, Catalogue Coccidse, p. 5 (1906). 
Habitat — Finland . 
On Phleum pratense; Poa pratensis. 

Pseudococcus juniperi Ehrhorn. 

Pseudococcus juniperi Ehrh., Can. Ent., xxxviii, p. 333 (1906). 
Habitat — Arizona. 
On Juniperus virginiana. 

o Specific name preoccupied. 



CATALOGUE OF EECENTLY DESCKIBED COCCID^ II. 41 

Pseudococcus longipes Leonard!. 

Pseudococcus longipes Leon., Bol. Zool. Sc. Sup. Portici, in, p. 166 (1908). Fig. 
Habitat — Italy. 
On Alocasia macrorica. 

Pseudococcus myrmecarius Leonard!. 

Pseudococcus myrmecarius Leon., Bol. Zool. Sc. Sup. Portici, in, p. 164 (1908). Fig. 
Habitat — Sardinia . 
In nest of ants {Camponotus sp.). 

Pseudococcus obscurus Essig. 

Pseudococcus obscurus Essig, Pomona Jn. Ent., i, 2, p. 43 (1909). Fig. 
Habitat — California. 
"On Opuntia and under the bark of Sambucus glauca." 

Pseudococcus saccharifolii (Green). 

Dactylopius saccharifolii Green, Mem. Dep. Ag. India, ii, 2, p. 23 (1908). 

Maxwell-Lefroy, Mem. Dep. Ag. India, ii, 7, p. 124 (1908). 
Fig. 
Habitat — Bengal. 
On leaves of sugar cane. 

Pseudococcus (?) takae (Kuwana). 

Dactylopius takse Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 184 (1907). Fig. 
Pseudococcus takae Ckll., Can. Ent., xli, 2, p. 55 (1909). 

Habitat— Japan . 

On bamboo. 

Pseudococcus theeecola (Green). 

Dactylopius thexcola Green, Mem. Dep. Ag. Ind., i, 5, p. 347 (1907). Fig. 
Green, Mem. Dep. Ag. Ind., ii, 2, p. 24 (1908). 
Habitat — India. 
"On roots of tea plants." 

Pseudococcus virgatus madagascariensis (Newstead). 

Dactylopius virgatus var. madagascariensis Newst., Liverpool Univ. Quart. Jn., in, 
6, p. 9 (1908). Fig. 
Habitat — Madagascar. 
"On an unknown plant." 

Pseudococcus vovae Nassonow. 

Pseudococcus {Dactylopius) vovae Nass., Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg, 
xiii, p. 484 (1909). Fig. 
Habitat — Russia . 
On Juniperus communis. 

Ripersiella kelloggi Ehrhorn. 

Ripersiella kelloggi Ckll. (sine descr.), Pr. Biol. Soc. Wash., xiv, p. 166 (1901). 
Ehrh., Can. Ent., xxxviii, p. 334 (1906). 
Habitat — California. 
On bunch-grass. 



42 PAPERS ON COCCID^ OR SCALE INSECTS. 

Ripersia candidata Cockerell. 

Ripersia candidata CklL, Ann. Mag. N. H. (7), vm, p. 53 (1901). 
"=6-jtd. R. lasii from Massachusetts." — (Ckll. in litt.) 

Ripersia donisthorpei Newstead. 

Ripersia donisthorpei Newst., Ent. Rec. Jn. Variation, xix, p. 5 (1907). Fig. 
Habitat — England. 
In nests of P oner a contracta. 

Ripersia formicarii Newstead. 

Ripersia formicarii Newst., Ent. Rec. Jn. Variation, xix, p. 5 (1907). Fig. 
Habitat — England. 
Associated with Lasiusfavus. 

Ripersia hypogea Leonardi. 

Ripersia hypogea Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 176 (1908). Fig. 

Habitat — Italy. 

Underground. 

Ripersia inquilina Leonardi. 

Ripersia inquilina Leon., Bol. Zool. Sc. Sup. Portici, in, p. 174 (1908). Fig. 
Habitat — Sardinia. 
In nest of undetermined ants. 

Ripersia japonica Kuwana. 

Ripersia japonica Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 186 (1907). Fig. 
Habitat — Japan . 
Under sheathing, base of leaf of Miscanthus sp. 

Ripersia libera Leonardi. 

Ripersia libera Leon., Bol. Zool. Sc. Sup. Portici, in, p. 170 (1908). Fig. 
Habitat — Italy. 
On an undetermined grass. 

Ripersia (?) oryzse Kuwana. 

Ripersia oryzse Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 186 (1907). Fig. 
CklL, Can. Ent., xli, 2, p. 55 (1909). 
Habitat — Japan . 
On roots of rice {Oryza) and other plants. 

Ripersia sardiniae Leonardi. 

Ripersia sardinise Leon., Bol. Zool. Sc. Sup. Portici, ni, p. 173 (1908). Fig. 
Habitat — Sardinia. 
In nest of ants {Solenopsis sp.). 

Genus MICROCOCCUS Leonardi. Type, silvestrii. 
Micrococcus Leon., Bol. Zool. Sc. Sup. Portici, i, p. 135 (1907). 
Micrococcus silvestrii Leonardi. 

Micrococcus silvestrii Leon., Bol. Zool. Sc. Sup. Portici, i, p. 136 (1907). Fig. 
Habitat — Sardinia. 
In nests of Tapinoma erraticum. 



CATALOGUE OP RECENTLY DESCEIBED COCCID^ II. 43 

Micrococcus similis Leonardi. 

Micrococcus similis Leon., BoL ZooL Sc. Sup. Portici, i, p. 143 (1907). Fig. 
Habitat — Sardinia . 
On roots of a cereaL 

Antonina (?) africana Newstead. 

Antonina (?) africana Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, pp. 69, 
72 (1906). 
Draper (sine descr.), Scale Ins. of Egypt, p. 11 (1907). 
Habitat — Egypt. 
On "sunt" tree {Acacia arabica var. nilotica). 

Antonina indica Green. 

Antonina indica Green, Mem. Dep. Ag. India, ii, 2, p. 27 (1908). Fig. 
Habitat — Bengal. 
On bases of leaves of "bariali" grass. 

Antonina phxagmitis Marchal. 

Antonina phragmitis Marchal, Compt. Rend. Ac. Sci., Paris, cxliii, 13, p. 872 (1909). 
Habitat — France. 
On Phragmites gigantea. 

Subfamily TACHARDIIN.^. 

Tachardia argentina Dominguez. 

Tachardia argentina Dominguez (sine descr.). An. Soc. Cient. Argentina, p. 219 (1906). 
Fig. 
Dominguez (sine descr.), Bol. Min. Ag., vii, 3, pp. 148, 150 (1907). 
Habitat — Argentina . 
On Acacia cavenia. 

Tachardia decorella these Green & Mann. 

Tachardia decorella var. theas Green & Mann, Mem. Dep. Ag. India, i, 5, p. 348 (1907). 
Green, Mem. Dep. Ag. India, ii, 2, p. 28 (1908). 
Habitat — India . 
On tea. 

Subfamily COCCIN^. 

Ceronema caudata Froggatt. 

Ceronema caudata Froggatt, n. sp. (?). 

Froggatt, Australian Insects, p. 376 (1907). 
Habitat — New South Wales, Australia. 
On "gum-trees." 

Pulvinaria antigoni Green. 

Pulvinaria antigoni Green, Tr. Linn. Soc. Lond., xii, 2, pp. 199, 204 (1907). Fig. 
Habitat — Seychelles. 
On Antigonon. 

Pulvinaria burkilli Green. 

Pulvinaria burkilli Green, Mem. Dep. Ag. India, ii, 2, p. 31 (1908). Fig. 
Habitat — India. 
On leaves of Croton tiglium. 



44 PAPERS ON COCCIDiE OR SCALE INSECTS. 

Pulvinaria jacksoni Newstead. 

Pulvinaria jachsoni Newst., Jn. Econ. Biol., ii, 4, p. 155 (1908). Fig. 
Habitat — West Africa. 
On cacao and Ficus sp. 

Pulvinaria kuTvacola Kuwana. 

Pulvinaria huvmcola Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 188 (1907). Fig. 
Ckll., Can. Ent., xli, 2, p. 55 (1909). 
Habitat^Japan . 
On mulberry. 

Pulvinaria orientalis Nassonow. 

Pulvinaria orientalis Nass., Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg, xiii, p. 493 
(1909). Fig. 
Habitat — Russia. 
On branches of Haloxylon ammodendron. 

Pulvinaria plucheas Ehrbom. 

Pulvinaria plucheds Ebrh., Can. Ent., xxxviii, p. 334 (1906). 
Habitat — California . 
On Pluchea sericea. 

Genus STOTZIA Marchal. Type, striata. 

Stotzia Marchal, Bui. See. Ent. Fr., p. 143 (1906). 

Stotzia striata Marchal. 

Stotzia striata Marchal, Bui. Soc. Ent. Fr., p. 144 (1906). 
Habitat — Algeria . 
On branches of Ephedra altissima. 

Genus EITPHILIPPIA Berlese & Silvestri. Type, olivina. 

Euphilippia Berl. & Silv., Redia, iii, fasc. 2, p. 396 (1905). 

Euphilippia olivina Berlese & Silvestri. 

Euphilippia olivina Berl. & Silv., Redia, in, fasc. 2, p. 398 (1905). Fig. 
Habitat — Italy. 
On olive. 

Ceroplastes africanus senegalensis Marchal. 

Ceroplastes africanus var. senegalensis Marchal, Bui. Soc. Zool. France, xxxiv, p. 68 

(1909). 
Marchal, Mem. Soc. Zool. France, xxii, p. 168 
(1909). 
Habitat — Senegal. 
On Acacia arabica; A. tortilis. 

Ceroplastes bussei Newstead. 

Ceroplastes bussei Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, pp. 71, 74 (1906). 

Habitat — Cameroon, West Africa. 

On cacao. 

Ceroplastes tenuitectus Green. 

Ceroplastes tenuitectus Green, Tr. Linn. Soc. Lond., xii, 2, p. 204 (1907). Fig. 
Habitat — Aldabra Isl. 
On "bois la fumee." 



CATALOGUE OF EECENTLY DESCRIBED COCCID^ — II. 45 

Ceroplastes theobromae Newstead. 

Ceroplastes theobromas Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, pp. 71, 
74 (1906). 
Newst. (sine descr.), Scale Ins. of Egypt, p. 16 (1907). 
Newst., Jn. Econ. Biol., in, 2, p. 38 (1908). 
Habitat— Cameroon, West Africa. 
On cacao {Theobroma cacao). 

Ceroplastes uvarise Marchal. 

Ceroplastes uvarise Marchal, Bui. Soc. Zool. France, xxxiv, p. 68 (1909). 

Marchal, Mem. Soc. Zool. France, xxii, p. 170 (1909). Fig. 
Habitat — French Guinea. 
On Uvaria sp. (Anonacese). 

Ceroplastes vuilleti Marchal. 

Ceroplastes vuilleti Marchal, Bui. Soc. Zool. France, xxxiv, p. 68 (1909). 

Marchal, Mem. Soc. Zool. France, xxii, p. 165 (1909). Fig. 
Habitat — Senegal . 
On Ormosia laxiflora (Leguminosae). 

Ceroplastodes chiton Green. 

Ceroplastodes chiton Green (sine descr.), Mem. Dep. Ag. India, ii, 2, p. 32 (1908). 
Habitat — India. 
On Cajanus indicu^. 

Coccus gymnospori (Green). 

Lecanium gymnospori Green, Mem. Dep. Ag. India, ii, 2, p. 29 (1908). Fig. 
Habitat — India. 
On Gymnosporia montana. 

Coccus hesperidum javanensis (Newstead). 

Lecanium (Trechocorys) hesperidum var. africanum Newst. (sine descr.), Liverpool Univ. 

Quart. Jn., i, 2, p. 74(1906). 
Lecanium hesperidum var. javanensis Newst., Jn. Econ. Biol., iii, 2, p. 38 (1908). Fig, 
Habitat — Java. 
On Liberian coffee. 

Coccus nicotianae (Newstead). 

Lecanium nicotianse Newst., Liverpool Univ. Quart. Jn., in, 6, p. 8 (1908). Fig. 
Habitat — Madagascar . 
On tobacco. 

Coccus tuberculatus Kotinsky. 

Coccus tuberculatus Kotinsky, Pr. Haw. Ent. Soc, i, pt. 5, p. 168 (1908). Fig. 
Hab itat — Singapore . 
On leaves of unknown tree. 

Mesolecanium perditulum Cockerell & Robbins. 

Mesolecanium perditulum Ckll. & Robb., Can. Ent., xli, p. 150 (1909). 
Habitat — Nicaragua. 
"On bark of small branches of tree No. 2122 "— C. F. Baker coll. 



46 PAPERS ON COCCIDiE OR SCALE INSECTS, 

Akermes montanus (Green). 

Lecanium montanum Green, Mem. Dep. Ag. India, ii, 2, p. 30 (1908). Fig. 
Habitat — Himalaya Region, India. 
On twigs of undetermined shrub. 

Genus PAL.ffiOLECANIUM Sulc. Type, bituberculatum. 
Palseolecanium Sulc, Ent. Mo. Mag., xix, p. 36 (1908). 

Genus PARTHE NO LECANIUM Sulc. Type, coryli. 
Parthenolecanium Sulc, Ent. Mo. Mag., xix, p. 36 (1908). 

Genus SPHLSIROLECANIXJM Sulc. Type, prunastri. 
Sphserolecanium Sulc, Ent. Mo. Mag., xix, p. 36 (1908). 

Globulicoccus n. subgen. of Lecanium auctt. Type, fuscum. 

Lindgr., Ent. Blatter, Schwabacb, m, 8, p. 113; 9, p. 136 (1907). 
Lindgr., Bar. Stat. f. Pflanzenscb. Hamb., ix, p. 7 (1907). 

Lecanium cecconi Leonardi. 

Eulecanium cecconi Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 178 (1908). Fig. 
Habitat — Italy . 
On Menispermum canadense. 

Lecanium glandi (Kuwana). 

Lecanium glandi Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 191 (1907). Fig. 
Eulecanium glandi Ckll., Can. Ent., xli, 2, p. 56 (1909). 

Habitat — Japan. 

On apple, pear, and other trees. 

Lecanium kunoensis (Kuwana). 

Lecanium kunoensis Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 131 (1907). Fig. 
Ckll., Can. Ent., xli, 2, p. 56 (1909). 
Habitat — Japan . 
On Rhar.inus japonica; Prunus mume; Pyrus sinensis. 

Lecanium piilchrum Marchal. 

Lecanium pulchrum King (sine descr.), Allg. Zeits. f. Ent., viii, p. 410 (1903). 
Eulecanium pulchrum Sanders, Catalogue Coccidae, p. 9 (1906). 
Lecanium rubellum Lindgr., Ent. Blatter, Schwabach, iii, 9, p. — (1907). 

Lindgr., Ber. Stat. f. Pflanzenscb. Hamb., ix, p. 7 (1907). 
pulchrum Marchal, Ann. Soc. Ent. France, lxxvii, p. 304 (1908). Fig. 
Habitat — Germany . 
On Calluna vulgaris; Ulex europseus; Rosa arvensis; Rubus sp.; Sarothamnus 

scoparius. 

Lecanium sericeum Lindinger. 

Lecanium sericeum Lindgr., Insekten Borse, xxiii, p. 147, Sept. (1906). Fig. 
Lecanium {Globulicoccus) sericeum Lindgr., Ent. Blatter, Schwabach, iii, 8 and 9 (1907). 

Lindgr., Ber. Stat. f. Pflanzenscb. Hamb., ix, p. 7 
(1907). 
Habitat — Bavaria . 
On Abies pectinata. 



CATALOGUE OF RECENTLY DESCRIBED COCCIDiE II. 47 

Paralecanium cocophyllae Banks. 

Paralecanium cocophyllae Banks, Philippine Jn. Sci., i, 3, pp. 223, 235 (1906). Fig. 
Habitat — Philippines. 
On cocoanut palm. 

"Lecanium" montonoae Newstead. 

Lecanium montonose Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, p. 69 
(1906). 
montonoai Lindgr. (sine descr.), Ent. Wochenblatt, xxiv, 5, p. 19 (1907). 
Habitat — Egypt . 

"Lecanium" nishigaharse Kuwana. 

Lecanium nishigaharx Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 192 (1907). Fig. 
Habitat — Japan . 
On mulberry (Morus sp.). 

Genus HEMILECANIUM Newstead. Type, theobromx. 

Hemilecanium Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, pp. 71, 74 (1906). 
Newst., Jn. Econ. Biol., in, 2, p. 39 (1908). 

Hemilecanium theobromae Newstead. 

Hemilecanium theobromse Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, pp. 
71, 74 (1906). 
Newst. (sine descr.). Scale Ins. of Egypt, p. 16 (1907). 
Newst., Jn. Econ. Biol., iii, 2, p. 39 (1908). Fig. 
Habitat — Cameroon, West Africa. 
On Theobroma cacao. 

« Genus HOUARDIA Marchal. Type, troglodytes. 

Houardia Marchal, Compt. Rend. Soc. Biol., Paris, lxvi, p. 586 (1909). 

Houardia troglodytes Marchal. 

Houardia troglodytes Marchal, Compt. Rend. Soc. Biol., Paris, lxvi, p. 586 (1909). 
Marchal, Bull. Soc. Zool. France, xxxiv, p. 73 (1909). 
Marchal, Mem. Soc. Zool. France, xxii, p. 171 (1909). Fig. 
Habitat — Senegal . 

In tunnels in branches of Balanites (Agialida) segyptiaca inhabited by ants of the 
genus Cremastogaster near kneri Mayr. 

Lecanopsis mirmecophila Leonardi. 

Lecanopsis mirmecophila Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 181 (1908). Fig. 
Habitat — Sardinia. 
In nest of ants ( Tetramorium csespitum) . 

Aclerda biwakoensis Kuwana. 

Aclerda (?) biwakoensis Kuwana, Bui. Ag. Exp. Sta. Japan, i, 2, p. 187 (1907). Fig. 
Ckll., Can. Ent., xli, 2, p. 55 (1909). 
Habitat — Japan . 
On Phragmites communis. 

o An unfortunate name, because of its similarity to Howardia Berlese & Leonardi, a 
genus of Diaspine Coccidae. 



48 PAPEKS ON COCCID^ OR SCALE INSECTS. 

Subfamily DIASPINiE. 

Chionaspis etrusca Leonard!. 

Chionaspis etrusca Leon., Annali di Agr., viii, p. 44 (1908). 

Leon., BoL Zool. Sc. Sup. Portici, m, p. 184 (1908). Fig. 
Leon., Cherm. ItaL, iv. No 88 (1908). 
Habitat — Italy. 
On Tamarix sp. 

Chionaspis gryphsefonnis Newstead. 

Chionaspis gryphseformis Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, p. 71 
(1906). 
crypheaformis Draper (sine descr.), Scale Ins. of Egypt, p. 15 (1907). Cairo. 
Habitat — Egypt . 
" On four unnamed plants; one a conifer." 

Chionaspis inday Banks. 

ct Chionaspis Candida Banks, Philippine Jn. Sci., i, 3, pp. 222, 232 (1906). Fig. 
Chionaspis inday Banks, Philippine Jn. Sci., i, 7, p. 787 (1906). 
, Habitat — Philippines. 
On cocoanut palm. 

Chionaspis manni Green. 

Chionaspis manni "Watt & Mann (sine descr.), Pests and Blights of Tea Plant, 2nd ed., 
p. 309 (1903). 
Green, Mem. Dep. Ag. India, i, 5, pp. 342, 344 (1907). Fig. 
Mann, Mem. Dep. Ag. India, i, 5, p. 351 (1907). 
Green, Mem. Dep. Ag. India, ii, 2, p. 37 (1908), 
Habitat — India. 
On tea; i^icus sp.; Solanum melongena. 

Chionaspis micropori Marlatt. 

Chionaspis micropori Marlatt, Bui. U. S. Bur. Ent., t. s. 16, pt. 2, p. 25 (1908). Fig. 
Habitat — Shansi Province, China 
On Populus tremula. 

Chionaspis vuilleti Marchal. 

Chionaspis vuilleti Marchal (sine descr.), Compt. Rend. Soc. Biol., lxvi, p. 587 (1909). 

Marchal, Bui. Soc. Zool. France, xxxiv, p. 69 (1909). 

Marchal, Mem. Soc. Zool. France, xxii, p. 175 (1909). Fig. 
Habitat — Guinea. 
On gum copal {Copaifera guibourtiana'^) 

Diaspis barbeii Green. 

Diaspis barberi Green, Mem. Dep. Ag. India, ii, 2, p. 35 (1908). Fig. 
Habitat — Tanjore, India. 
On branches of Loranthus sp. 

Diaspis (?) ericicola Marchal. 

Diaspis {Adiscodiaspis) ericicola Marchal, Compt. Rend. Ac. Sci., Paris, cxlviii, 13, 
p. 871 (1909). 
Habitat — France. 
On Erica arbor ea. 

"Specific name preoccupied. 



CATALOGUE OF RECENTLY DESCEIBED COCCID^ — II. 49 

Aulacaspis javanensis Newstead. 

Aulacaspis (Diaspis) javanensis Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 

2, p. 74 (1906). 
Aulacaspis javanensis Newst., Jn. Econ. BioL, iir, 2, p. 35 (1908). Fig. 

Habitat — Java. 

" On various plants." 

Aulacaspis penzigi Leonardi. 

Aulacaspis penzigi Leon., Annali di Agr., vii, p. 14 (1907). Fig. 
Habitat — Java. 
On leaves of Ilex sp. 

Aulacaspis tubercularis Newstead. 

Aulacaspis (Diaspis) tubercularis Newst., Liverpool Univ. Quart. Jn., i, 2, p. 73 (1906). 
Aulacaspis cinnamomi Newst., Jn. Econ. Biol., iii, 2, p. 34 (1908). Fig. 

Habitat — Java. 

On Cinnamomum zeylanicum. , 

Hemichionaspis aspidistrae gossypii Newstead. 

Chionaspis (Hemichionaspis) aspidistrx var. gossypii Newst. (sine descr.), Liverpool 

Univ. Quart. Jn., i, 2, p. 74 (1906). 
Newst. , Jn. Econ. Biol. , m, 2, p. 37 
(1908). 

Habitat — Java. 

On Gossypium hirsutum. 

Hexnichionaspis fici Green. 

Hemichionaspis fid Green, Mem. Dep. Ag. India, ii, 2, p. 37 (1908). Fig. 
Habitat — Bengal . 
On stems of Ficus glomerata. 

Hemichionaspis minima Green. 

Hemichionaspis minima Green, Mem. Dep. Ag. India, ii, 2, p. 38 (1908). Fig. 
Habitat — Bengal. 
On leaves of " Banian" tree (Ficus sp.). 

Hemichionaspis orlandi Leonardi. 

Hemichionaspis orlandi Leonardi, Redia, iii, p. 5 (1906). Fig. 

orlandii Lindgr., Ent. Wochenblatt., xxiv, 5, p. 19 (1907). 
Habitat — Brazil. 
"Underside of leaves of an undetermined plant." 

Pinnaspis javanica Leonardi. 

Pinnaspis javanica Leon., Annali di Agr., vii, p. 17 (1907). Fig. 
Habitat — Java. 
On leaves of Ilex sp. 

Pinnaspis rombica Leonardi. 

Pinnaspis rombica Leon., Annali di Agr., vii, p. 16 (1907). Fig. 
Habitat — Java. 
On leaves of Per sea sp. 



50 PAPERS ON COCCID^ OR SCALE INSECTS. 

Genus ACTENASPIS Leonardi. Type, pusilla. 

Actenaspis Leon., Annali di Agr., vi, p. 25 (1906). Fig. 
A new genus from Leucaspis auctt. 

Genus ANAMASPIS Leonardi. Type, loewi. 

Anamaspis Leon., Annali di Agr., vi, p. 22 (1906). Fig. 
A new genus from Leucaspis auctt. 

Xieucaspis affinis Leonardi. 

Leucaspis affinis Leon., Annali di Agr., vi, p. 4 (1906). Fig. 
Habitat — France. 
On Pinus sylvestris. 

Leucaspis ephedras Marchal. 

Leucaspis ephedrse Marchal, Bui. See. Zool. France, xxxiv, p. 59 (1909). 
Habitat — Algeria. 
On Ephedra corsoniana. 

Leucaspis indica Marlatt. 

Leucaspis indica Marlatt, BuL.U. S. Bur. Ent., t. s. 16, Pt. II, p. 26 (1908). Fig. 
Habitat — Miami, Fla., and Mayaguez, P. R.; imported from ftidia. 
On mango. 

Leucaspis pistacise Lindinger. 

Leucaspis pistacise Lindgr., Jahrb. Hamb. wiss. Anst., xxiii, p. 403 (1906). 

Hab itat — Cyprus . 

On leaves of Pistacia lentiscus. 
o Leucodiaspis Sign.<Lindinger, Jahrb. Hamb. wiss. Anst., xxv (1908). Not valid. 
Adiscofiorinia (n. g. of Fiorinia), Leonardi, Redia, iii, pp. 17, 52 (1906). 
Anamefiorinia (n. g. of Fiorinia), Leonardi, Redia, iii, pp. 17, 48 (1906). 
Trullifiorinia (n. g. of Fiorinia), Leonardi, Redia, iii, pp. 17, 41 (1906). 

Fiorinia atalantiae Leonardi. 

Fiorinia (Adiscofiorinia) atalantise Leon., Redia, iii, p. 54 (1906). Fig. 
Habitat — Ceylon. 
On Atalantia zeylanica. 

« Lindinger resurrects Leucodiaspis Sign, for Leucaspis, and assigns to it all the 
species formerly known under the latter name, and refers to Leucospis Fab., Syst. 
Entom., p. 361, Nr. 114 (1775) (Hymenoptera). The latter name has no reference or 
connection with Leucaspis Targ.-Tozz., a genus of Coccidae erected in 1869. Leuco- 
diaspis credited by Signoret [Ann. Soc. Ent. France, ix, p. 99 (1869)] to Targioni- 
Tozzetti is an error in spelling and of no value. 

The following reference to Leucaspis appears in Atti Soc. Ital. di Sci. Nat., xi, p. 
734 (1869), which establishes its validity: 

"CocciDARUM Catalogus Targ.-Tozz. 

"Gen. 22. Leucaspis noh. Gen. n. 

"Sp. 1. Leucaspis Candida nob. Diaspis Candida nob. 1867. 

"Sp.2. Leucaspis Signoreti nob. n.sp.l868(aD. Signoretirecepta, Pini sp. incola)." 



CATALOGUE OF EECENTLY DESCRIBED COCCID^— II. 51 

Fiorinia diaspiformis Newstead. 

Fiorinia diaspiformis Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 2, p. 74 (1906). 
diaspitiformis Lindgr. (sine descr.), Ent. Wochenblatt., xxiv, 5, p. 20 (1907). 
diaspiformis Newst., Jn. Econ. Biol., iii, 2, p. 35 (1908). Fig. 
Habitat — Java. 
On Piper sp. 

Fiorinia hirsuta Marchal. 

Fiorinia hirsuta Marchal, Bui. See. Ent. Fr., p. 145 (1906). 
Habitat — Algeria. 
On Nephelium longana and other Sapindacese. 

Fiorinia juniperi Leonardi. 

Fiorinia juniperi Leon., Redia, in, p. 39 (1906). Fig. 
Habitat — Ceylon . 
On Juniperus bermudiana. 

Fiorinia rtiacroprocta (Leonardi). 

Trullifiorinia macroprocta Leon., Annali di Agr., vii, p. 19 (1907). Fig. 
Habitat — Java. 
On leaf of Raphis flabelliformis . 

Fiorinia odinse Leonardi. 

Fiorinia odinx Leon., Redia, iii, p. 24 (1906). Fig. 
Habitat — Ceylon. 
On Odina woodice. 

Fiorinia rubrolineata Leonardi. 

Fiorinia (Trullifiorinia) rubrolineata Leon., Redia, in, p. 44 (1906). Fig. 
Habitat — Ceylon. 
On Murraya exotica. 

Fiorinia tumida Leonardi. 

Fiorinia tumida Leon., Redia, iii, p. 38 (1906). Fig. 
Habitat — Ceylon. 
On Grewia sp. 

Aspidiotus africanus Marlatt. 

Aspidiotus (Diaspidiotus) africanus Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 15 
(1908). Fig. 
Habitat — South Africa. 

On fig; apricot; almond; apple; quince; pear; privet; honey-locust (Gleditsia 
triacanthos) ; Schinus molle; Acacia horrida; Robinia pseudacacia; Rhuss]). 

Aspidiotus cecconi (Leonardi). 

Hemiberlesia cecconi Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 188 (1908). Fig. 
Habitat — Sardinia. 
On Osirys alba. 

Aspidiotus cocotiphagus Marlatt. 

Aspidiotus (Aonidiella) cocotiphagus Marlatt, Bui. U. S. Bur. Ent., t. l. 16, Pt. II, p. 14 
(1908). Fig. 
Habitat — Santiago de las Vegas, Cuba. 
On Cocos nucifera; Citrus trifoliata. 



52 . PAPEKS ON COCCIDiE OB SCALE INSECTS. 

Aspidiotus comperei Marlatt. 

Aspidiotus (s. str.) comperei Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 12 (1908). 
Fig. 
Habitat — W. Australia. 
On Hahe sp. and an undetermined plant. 

Aspidiotus corticis-pini Lindinger. 

Aspidiotus corticis-pini Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. — (1908). 
Habitat — Japan . 
On Pinus densiflora. 

Aspidiotus coursetise Marlatt. 

Aspidiotus (Diaspidiotus) coursetise Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 20 
(1908). Fig. 
Habitat — Mexico. 
On Coursetia glandulosa. 

Aspidiotus densiflora Bremner. 

J.spi(?iofMS rfensi^ora Bremner, Can. Ent., XXXIX, p. 366 (1907). Fig. 
Habitat — California . 
On leaves of Quercus densiflora. 

Aspidiotus eglandulosus Lindinger. 

Aspidiotus eglandulosus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. — (1908). 

Lindgr., Ber. Stat. f. Pflanzenscb. Hamb., x, p. 10 (1908). 
Habitat — Guatemala; Panama. 
On Cereus; "auf Kakteen." 

Aspidiotus elaeidis Marchal. 

Aspidiotus elaeidis Marchal (sine descr.) Compt. Rend. Soc. Biol., lxvi, p. 587 (1909). 

Marchal, Bui. Soc. Zool. France, xxxiv, p. 69 (1909). 

Marchal, Mem. Soc. Zool. France, xxn, p. 179 (1909). Fig. 
Habitat — Dahomey. 
On oil palm {Elaeis guineensis) . 

Aspidiotus epigaeae Marlatt. 

Aspidiotus (Hemiberlesia) epigaese Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 21 
(1908). Fig. 
Habitat — Virginia; Ohio. 
On trailing arbutus {Epigsea repens) . 

Aspidiotus herculeanus Doane and Hadden . 

Aspidiotus herculeanus Doane and Hadden, Can. Ent., xli, p. 298 (1909). Fig. 
Habitat — Society Islands. 
On bark of unknown plant. 

Aspidiotus labiatarum Marchal. 

Aspidiotus (Evaspidiotus) labiatarum Marchal, Compt. Rend. Ac. Sci., Paris, cxLvni, 
13, p. 872 (1909). 
Habitat — Corsica. 
On Stachys glutinosa and Teucrium capitatum. 



CATALOGUE OF RECENTLY DESCRIBED COCCIDiE II. 53 

Aspidiotus lectularius Green. 

Aspidiotuslectularius" Green " French (sine descr.), Victorian Nat., xxiii, p. 184 (1907). 
Habitat — Victoria, Australia. 

Aspidiotus meyeri Marlatt. 

Aspidiotus (s. str.) me?/m Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 13 (1908). Fig. 
Habitat — Peking, China. 
On Abies sp. 

Aspidiotus mitchelli Marlatt. 

Aspidiotus (Hemiberlesia) mitchelli Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 22 
(1908). Fig. 
Habitat — Mitchell's Pass, South Africa. 
"On thick, narrow, slightly oblanceolate leaves about 1| in. long." 

Aspidiotus popularum. Marlatt. 

Aspidiotus (Hemiberlesia) popularum Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, 
p. 23 (1908). Fig. 
Habitat — Arizona; New Mexico. 
On Cottonwood (Populus sp.). 

Aspidiotus privignus Lindinger. 

Aspidiotus privignus Lindgr., Zeitschr. f. wiss. Insektenbiol., v, 5, 7 & 8, p. 151 (1909). 
Fig. in No. 5. 
Habitat — Italy; Greece. 
On Hypericum coris; Thymelsea tarton (?). 

Aspidiotus trabuti Marchal. 

Aspidiotus (Hemiberlesia) trabuti Marchal, Bui. Soc. Zool. France, xxxiv, p. 59 (1909). 
Habitat — Algeria. 
On Ephedra altissima. 

Aspidiotus transcaspiensis Marlatt. 

Aspidiotus (Diaspidiotus) transcaspiensis Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, 
p. 21 (1908). Fig. 
Habitat — Transcaspian Russia. 
On Populus sp. 

Aspidiotus vuilleti Marchal. 

Aspidiotus (Hemiberlesia) vuilleti Marchal, Compt. Rend. Soc. Biol., lxvi, p. 587 

(1909). 
Marchal, Bui. Soc. Zool. France, xxxiv, p. 73 

(1909). 
Marchal, Mem. Soc. Zool. France, xxii, p. 178 
(1909). Fig. 
Habitat — Senegal. 
On branches of Balanites sp. 

Aspidiotus yulupaB Bremner. 

Aspidiotus yulupae Bremner, Can. Ent., xxxix, p. 367 (1907). Fig. 
Habitat — California . 
On Quercus lobata. 



54 PAPERS ON COCCID.^ OR SCALE INSECTS. 

Pseudaonidia ferox (Lindinger). 

Selenaspidusferox Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, pp. 4, 7 (1909). Fig. 
Habitat — Gold Coast, West Africa. 
On Euphorbiacese. 

Pseudaonidia gracilis (Lindinger). 

Selenaspidus gracilis Lindgr., Jabrb. Hamb. wiss. Anst., xxvi, pp. 4, 10 (1909). Fig., 
Habitat — Kamerun, West Africa. 
On leaves of Agelaia fragrans and Tricalysia sp. 

Pseudaonidia greeni Marlatt. 

Pseudaonidia greeni Marlatt, Pr. Ent. Soc. Wash., ix, pp. 135, 138, 14(? (1908). 
Habitat — Java. 
On mangosteen and mango. 

Pseudaonidia junctiloba Marlatt. 

Pseudaonidia junctiloba Marlatt, Pr. Ent. Soc. Wash., ix, pp. 135, 138 (1908). 
Habitat — Victoria, Australia. 
On Acacia sp. 

Pseudaonidia kamerunica (Lindinger). 

Selenaspidus Icamerunicus Lindgr., Jahrb. Hamb. wiss. Anst., xxvr, pp. 4, 7 (1909). 
Fig. 
Habitat — Kamerun, West Africa. 
On palms. 

Pseudaonidia lounsburyi Marlatt. 

Pseudaonidia (Selenaspidus) lounsburyi Marlatt, Pr. Ent. Soc. Wash., ix, pp. 134, 136, 
139 (1908). 
Habitat — South Africa. 
On Mesembryanthemuvi edule. 

Pseudaonidia magna (Lindinger). 

Selenaspidus magnus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, pp. 4, 9 (1909). Fig. 
Habitat — Abyssinia. 
On Euphorbia sp. 

Pseudaonidia silvatica (Lindinger). 

Selenaspidus silvaticus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, pp. 4, 10 (1909). Fig. 
Habitat — Kamerun, West Africa; Amani, East Africa. 
On Anacardiaceee ; Rinorea exappendiculata; Ficus indica; Bandeirsea speciosa. 

Genus FURCASPIS Lindinger. Type, biformis. 

Furcaspis Lindgr., Berl. Ent. Zeitschr., lii, pp. 98, 99 (1908). 

Includes Furcaspis {Aspidiotus) biformis (Ckll.) and F. (Aspidiotus) capensis 
(Walk.). 

Furcaspis oceanica Lindinger. 

Aspidiotus biformis Lindgr., Ber. Stat. f. Pflanzensch., vi, p. 44 (1904). In part. 
Furcaspis oceanica Lindgr., Zeitschr. f. wiss. Insektenbiol., v. 5, p. 149 (1909). Fig. 
in Nos. 7 and 8. 

Habitat — Jaluit Islands in Marshall group. 

On fruit hulls of cocoanut. 



CATALOGUE OF EECENTLY DESCKIBED COCCID^ II. 55 

Chrysomphalus barbusano Lindinger. 

Chrysomphalus harhusano Lindgr., Berl. Ent. Zeitschr., lii, p. 101 (1908). 

Lindgr., Ber. Stat. f. Pflanzensch. Hamb., p. 8 (1908). 
Lindgr., Zeitschr. f. wiss. Insektenbiol., v, 4, p. 105 (1909). 

Fig- 
Habitat — Teneriffe, Canary Islands. 
On Phoebe barbusano. 

Chrysomphalus minutus Kotinsky. 

Chrysomphalus minutus Kotinsky, Pr. Haw. Ent. Soc, i, pt. 5, pp. 170-171 (1908). Fig. 
Habitat — Singapore. 
On leaf of undetermined plant. 

Cbrysomphalus propsimus Banks. 

Chrysomphalus propsimus Banks, Philippine Jn. Sci., i, 3, pp. 221, 230 (1906). Fig. 
Habitat — Philippines. 
On leaves of Cocos nucifera. 

Chrysomphalus taxus (Leonardi). 

Aonidiella aurantii (Mask.) Berl. & Leon., Cherm. Ital., Fasc. i, no. 23 (1895). 
taxus Leon., Redia, iir, p. 1 (1906). Fig. 

Leon., Bol. Zool. Sc. Sup. Portici, p. 131 (1907). Fig. 
Habitat — Italy. 
On Taxus baccata. 

Chrysom.ph.alus triglandulosus Green. 

Aspidiotus (Chrysomphalus) triglandulosus Green, Mem. Dep. Ag. India, ii, 2, p. 33 

(1908). Fig. 
Mem. Dep. Ag. India, n, 7, p. 134 
(1908). 
Habitat — Bombay. 
On leaves of undetermined tree. 

Targionia alni Marchal. 

Targionia alni Marchal, Compt. Rend. Ac. Sci., Paris, cxliii, 13, p. 872 (1909). 
Hab itat — France . 
On Alnus glutinosa. 

Targionia chenopodii Marlatt. 

Aspidiotus (Targionia) chenopodii Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 24 
(1908). Fig. 

Habitat — Coolabah, New South Wales. 
On Chenopodium. 

Targionia vitis suberi Leonardi. 

Targionia vitis var. suberi Leon., Bol. Lab. Zool. Sc. Sup. Portici, p. 166 (1907). Fig. 
Habitat — Sardinia. 
On Quercus suber. 

Odonaspis graminis Bremner. 

Odonaspis graminis Bremner, Can. Ent., xxxix, p. 368 (1907). Fig. 
Habitat — California. 
On roots of grass. 



56 PAPERS ON COCCIDiE OR SCALE INSECTS. 

Odonaspis ruthae Ehrhorn. 

Odonaspis ruthse Ehrh. (sine descr.), 2d Bien. Rep. Com. Hort. Cal., p. 28 (1907) 
Habitat — Hawaii; California (at quarantine). 
On grass. 

Pseudoparlatoria chilina Lindinger. 

Pseudoparlatoria chilina Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. — (1908). 
Habitat— Chile. 
On Saxegothasa conspicua. 

Aonidia glandulosa Newstead. 

Aonidia glandulosa Newst., Draper (sine descr.), Scale Ins. of Egypt, p. 11 (1907). 
Habitat — Egypt . 
On "sunt" tree {Acada arabica var. nilotica). 

Aonidia juniperi Marlatt. 

Aonidia juniperi Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 24 (1908). Fig. 
Habitat — Utah . 
On Juniperus sp. 

Aonidia perplexa Green. 

Aonidia perplexa Green, Jn. Bomb. N. H. Soc, xiii, p. 252 (1902). Fig. 

Lindgr., Zeitschr. f. wiss. Insektenbiol., v, 4, p. 109 (1909). . 
Habitat — Ceylon . 
On leaves of Mesua ferrea. 

Aonidia picea Leonardi. 

Aonidia picea Leon., Redia, in, p. 6 (1906). Fig. 
Habitat — Spain . 
On Billartia officinalis. 

Aonidia pinicola Leonardi. 

Aonidia pinicola Leon., Annali di Agr., vi, p. 3 (1906). Fig. 
Habitat — Spain . 
On Pinus sylvestris. 

Gyninaspis clusiae Lindinger. 

Gymnaspis clusise Lindgr., Deutsche Ent. Zeitschr., l, p. 153 (1909). Fig 
Habitat — Jamaica. 
On leaves of Clusia sp. 

Lepidosaphes auriculata (Green). 

Mytilaspis auriculata Green, Tr. Linn. Soc. Lond., xii, pt. 2, p. 205 (1907). Fig. 
Habitat — Seychelles . 
On "croton." 

Lepidosaphes beckii oleae Leonardi. 

Lepidosaphes beckii var. olex Leon., Bol. Zool. Sc. Sup. Portici, iii, p. 193 (1908). Fig. 
Habitat — Sardinia. 
On roots of olive. 

Lepidosaphes beyerise (Green). 

Mytilaspis beyerise "Green" French (sine descr.), Victorian Nat., xxiii, p. 184 (1907). 
Habitat — Victoria, Australia. 



CATALOGUE OF RECENTLY DESCEIBED COCCID^ II. 57 

Lepidosaphes bicolor Newstead. 

Mytilaspis (Lepidosaphes) bicolor Newst. (sine descr.), Liverpool Univ. Quart. Jn., i, 

2, pp. 71, 72 (1906). 
Newst. (sine descr.), Scale Insects of Egypt, p. 15 
(1907). 
Habitat — Egypt . 
"On an unknown plant." 

Liepidosaphes chilopsidis (Marlatt). 

Mytilaspis chilopsidis Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 27 (1908). Fig. 
Habitat — Mexico. 
On Chilopsis linearis. 

Lepidosaphes destefanii Leonardi. 

Lepidosaphes destefanii Leon., Bol. Zool. Sc. Sup. Portici, i, p. 167 (1907). Fig. 
Habitat — Italy. 
On Phillyrea media. 

Lepidosaphes ficifoHae ulmicola Leonardi. 

Lepidosaphes Jicifolise var. ulmicola Leon., Bol. Zool. Sc. Sup. Portici, i, p. 168(1907). 
Fig. 

Habitat — Italy. 
On leaves of Ulmus. 

Lepidosaphes longula Leonardi. 

Lepidosaphes longula Leon., Annali di Agr., vii, p. 20 (1907). Fig. 
Habitat — Java. 
On leaves of Per sea sp. 

Lepidosaphes mcgregori Banks. 

Lepidosaphes mcgregori Banks, Philippine Jn. Sci., i, 3, pp. 222, 233 (1906). Fig. 
Habitat — Philippines. 
On leaves of Cocos nudfera. 

Lepidosaphes ocellata (Green). 

Mytilaspis ocellata Green, Tr. Linn. Soc. Lond., xii, pt. 2, p. 206 (1907). Fig. 
Habitat — Seychelles. 
On fronds of Davallia sp. 

Lepidosaphes piperis (Green). 

Mytilaspis piperis Green, Mem. Dep. Ag. India, ii, 2, p. 34 (1908). Fig. 
Habitat — Madras, India. 
On young stems of Piper nigrum,. 

« Lepidosaphes serrifrons (Leonardi). 

Mytilaspis serrifrons Leon., Riv. Pat. Veg., vi, pp. 118 (276), 121 (279) (1897). Fig. 

Ckll., Bui. 111. St. Lab. N. H., v. Art. xii, p. 397 (1899). 

Leon., Annali di Agr., v, pp. 45, 48 (1903). Fig. 
Habitat — I taly . 
On Croton undulatum and C. majesticum. 

a This species apparently has been omitted in the Fernald catalogue and the writer's 
first supplementary catalogue (1906). 



58 PAPEKS ON COCCID^ OR SCALE UsTSECTS. 

Lepidosaphes unicolor Banks. 

Lepidosaphes unicolor Banks, Philippine Jn. Sci., i, 3, pp. 222, 234 (1906). Fig. 
Habitat — Philippines. 
On leaves of Cocos nudfera. 

Parlatoria chinensis Marlatt. 

Parlatoria chinensis Marlatt, Bui. U. S. Bin-. Ent., t. s. 16, Pt. II, p. 30 (1908). Fig. 
Habitat — China; Japan; Egypt. 
On crab-apple; Hibiscus; Xanthoxylon; Thuja orientalis. 

Parlatoria cinerea Doane & Hadden. 

Parlatoria cinerea Doane & Hadden, Can. Ent., xli, p. 299 (1909). Fig. 
Habitat — Society Islands. 
On bark of orange tree and on cultivated vine. 

Parlatoria greeni Banks. 

Parlatoria greeni Banks, Philippine Jn. Sci., i, 3, pp. 222, 231 (1906). Fig. 
Habitat — Philippines . 
On leaves of Cocos nudfera. 

Parlatoria raangiferae Marlatt. 

Parlatoria mangiferse Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 28 (1908). Fig. 
Is a synonym of P. pseudaspidiotus Lindgr., Insekten Borse, xxii, 33, p. 131 (1905). 

Parlatoria pyri Marlatt. 

Parlatoria pyri Marlatt, Bui. U. S. Bur. Ent., t. s. 16, Pt. II, p. 29 (1908). Fig. 
Habitat — China; Manchuria. 
On apple; pear. 

CORRECTIONS TO CATALOGUE I (1906). 

Page 2. Icerya colimensis CklL, Mem. Soc. Cient. Ant. Alz., xviii, p. 81 (1902). 

Icerya tonilensis CklL, Mem. Soc. Cient. Ant. Alz., xviii, p. 80 (1902). 

Icerya rileyi larrese CklL, Mem. Soc. Cient. Ant. Alz., xvin, p. 82 (1902). 
Page 4. Trionyraus hordei (Lindeman). 

Page 5. Pseudococcus elongatus Reuter, corrected in Catalogue II, p. 40. 
Page 10. Chionaspis angusta Green for Ch. angustata. 
Page 14. Aspidiotus riverx CklL, Ent. News, xvi, p. 161 (1905). 
Page 15. Chrysomphalus malleolus GreeB.= Pseudaonidia malleolus (Green). 
Page 16. Fernaldella for Fernaldiella. 

Attention should be called to the fact that Fernaldella Leonardi is preoccupied in 
Lepidoptera. 



ABBREYIATIONS OF LITERATURE. 

Allg. Zeits. f . Ent. — Allgemeine Zeitschrift ftir Entomologie. 

Agr. News West. Ind. — Agricultural News. West Indies. 

An. Soc. Cient. Argentina. — Anales de la Sociedad Cientifica Argentina. 

Ann. Soc. Ent. France. — Annales de la Societe Entomologique de France. 

Annali di Agr. — Annali di Agricoltura. Portici. 

Ann. Mag. N. H. — Annals and Magazine of Natural History. 

Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg. — Annuaire Musee Zoologique Academic 

Imperiale des Sciences de St. Petersbourg. 
Berl. Ent. Zeitsqhr. — Berliner Entomologiscbe Zeitschrift. 

Ber. Stat. f. Pflanzensch. Hamb. — Bericht der Station fiir Pflanzenschutz, Hamburg. 
Bol. Min. Ag. — Boletm del Ministerio de Agricultura. Argentina. 
Bol. Zool. Sc. Sup. Portici. — Bollettino del Laboratorio di Zoologia generale e agraria 

della R. Scuola Superiore d' Agricoltura di Portici. 
Bot. Jahrb. — Botaniscbe Jahrbiicher. 
Bui. Ag. Exp. Sta. Japan. — Bulletin of the Imperial Central Agricultural Experiment 

Station. Japan. 
Bui. 111. St. Lab. N. H. — Bulletin of the Illinois State Laboratory of Natural History. 
Bui. Soc. Ent. Fr. — Bulletin de la Societe Entomologique de France. 
Bui. Soc. Ent. Ital. — BuUetino della Societa Entomologica Italiana. 
Bui. Soc. Zool. France. — Bulletin de la Societe Zoologique de France. 
Bul.U. S. Bur. Ent. t.s. 16, Pt.II. — Bulletin, Technical Series, Bureau of Entomology, 

U. S. Department of Agriculture. 
Can. Ent. — Canadian Entomologist. 

Catalogue Coccidse.— U. S. Bur. Ent. technical series, Bui. 12, Pt. I (1906). 
Cherm. Ital. — Chermotheca Italica. 
Compt. Rend. Ac. Sci., Paris. — Comptes rendus hebdomadaires des seances de I'Aca- 

demie des sciences. Paris. 
Compt. Rend. Soc. Biol. — Comptes rendus des seances de la Societe de Biologic. Paris. 
Deutsche Ent. Zeitschr. — Deutsche Entomologische Zeitschrift. 
Ent. Mo. Mag. — The Entomologist's Monthly Magazine. 
Ent. Blatter. — Entomologische Blatter, Internationale Monatsschrift fiir die Biologic 

der Kafer Europas. Schwabach. 
Ent. Rec. Jn. Variation. — The Entomologist's Record and Jovu-nal of Variation. 
Ent. Wochenblatt. — Entomologisches Wochenblatt (antea Insekten-Borse). 
Ind. Mus. Notes. — Indian Museum Notes. 
Insekten Borse. — Entomologische Wochenblatt. 

Jahrb. Hamb. wiss. Anst. — ^Jahrbuchder Hamburgischenwissenschaftlichen Anstalte'n. 
Jn. Bomb. N. H. Soc. — ^Jom-nal of the Bombay Natural History Society. Calcutta. 
Jn. Econ. Biol. — The Journal of Economic Biology. 
Jn. N. Y. Ent. Soc. — Joiunal of the New York Entomological Society. 
Liverpool Univ. Quart. Jn. — Institute of Commercial Research in the Tropics, Liver- 
pool University, Quarterly Journal. 
Medd. Soc. Faun. Fenn. — Meddelanden af Societas pro Fauna et Flora Fennica. 
Mem. Dep. Ag. India. — Memoirs of the Department of Agriculture in India. 
Mem. Soc. Cient. Ant. Alz. — Memorias y Revista de la Sociedad Cientifica "Antonio 

Abzate." 
Mem. Soc. Zool. France. — Memoires de la Societe Zoologique de France. 

59 



60 PAPEKS ON COCCIDiE OR SCALE INSECTS. 

NachricMsbl. Naturfrsch.-Klubs, Prossnitz. — Nachrichtsblatt Naturforscher-Klubs, ^ 

Prossnitz (Mahren). 
N. Z. Trans. — Transactions of the New Zealand Institute. 
Pests and Blights of Tea Plant. — The Pests and Blights of the Tea Plant, 2d ed. "Watt 

& Mann, Calcutta, 1903. 
Philippine Jn. Sci. — The Philippine Jom-nal of Science. 
Pomona Jn. Ent. — Pomona Journal of Entomology. Claremont, California. 
Pr. Biol. Soc. Wash. — Proceedings of the Biological Society of Washington. 
Pr. Ent. Soc. Wash. — Proceedings of the Entomological Society of Washington. 
Pr. Haw. Ent. Soc. — Proceedings of the Hawaiian Entomological Society. 
Redia. — Redia. 

Riv. Pat. Veg. — Ri vista di Patologia Vegetale. 
Scale Ins. of Egypt. — Notes on the Injurious Scale Insects and Mealy Bugs of Egypt. 

Cairo, 1907. 
Trab. Mus. Farm. Fac. Cienc. Med. — Trabajos del Museo de farmacologia de la Facul- 

tad de Ciencias Medicas. Buenos Aires, Argentina. 
Tr. Linn. Soc. Lond. — Transactions of the Linnsean Society of London. 
Victorian Nat. — The Victorian Natm-alist. 
Zeitschr. f. wiss. Insektenbiol. — Zeitschrift f iir wissenchaftliche Insektenbiologie. 

o 



v-'Y ^' 



\^ 



Technical Series, No. 16> Part IV. 

U. S. DEPARTMENT OF AGRICULTURE. 

BXJREi^TJ OW EIsTTOMIOLOGS-^". 

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



PAPERS ON COCCIDJ: OR SCALE INSECTS. 



CATALOGUE 



RECENTLY DESCRIBED COCCID^-IIL 



By E. R. SASSCER, 

Scientific Assistant. 



Issued June 8, 1911. 




WASHINGTON : 

GOVERNMENT PRINTING OFFICE. 

1911. 

& 



CONTENTS. 

Page. 

Subfamily Monophlebinse 61 

Subfamily Dactylopiinse 62 

Subfamily Coccinse 66 

Subfamily Diaspinse 68 

II 



U. S. D. A., B. E. Tech. Ser. 16, in. IV. Issued June 8, 1911. 

PAPERS ON G0GGIJ)M OR SCALE INSECTS. 



CATALOGUE OF EECENTIY DESCRIBED COCCID^— III. 

By E. R. Sasscer, Scientific Assistant. 

Since the publication, December 22, 1909, of Tech, Ser. 16, Part III, 
' 'Catalogue of Recently Described Coccidse — II, " by Mr. J. G. Sanders, 
3 new subgenera, 98 new species, and 5 new varieties have been 
recorded by workers on scale insects. In all, some 23 new genera, 9 
new subgenera, 430 new species, and 41 new varieties have been 
recorded from various quarters of the globe since the publication of 
Mrs. Fernald's valuable contribution in 1903. 

Owing to the large amount of literature relating to entomology 
which is published daily and to the fact that some entomologists 
describe species in periodicals which do not relate to insect life, it 
is the earnest request of the writer that all workers on Coccidse send to 
the Bureau of Entomology separates of their papers and as far as pos- 
sible any available material. This list is believed to be fairly com- 
plete up to March, 1911, and any corrections or omissions are solicited 
from coworkers. 

In examining the voluminous literature the writer was greatly 
assisted by Miss Mabel Colcord, librarian of the bureau, and he is 
equally indebted to Prof. T. D. A. Cockerell for his many valuable 
suggestions. 

All species marked with an asterisk are represented in the National 
Collection of Coccidse. 

Subfamily MONOPHLEBINiEo 

Monophlebus sjostedti Newstead. 

Monophlebus sjostedtiNewst., Kilimandjaro-Meru Exp., 12(1 Coccidae), p. 2 (1908). 
Fig. 
Habitat — Kilimandjaro. 

Monophlebus pallidus Newstead. 

Monophlebus pallidus Nevrst., Kilimandjaro-Meru Exp., 12 (1 Coccidse), p. 3 (190S). 
Fig. 
Habitat — Kilimandjaro. 

Walkeriana africana Newstead. 

Walkeriana africana Newst., Kilimandjaro-Meru Exp., 12 (1 Coccidas), p. 4 (1908). Fig. 
Habitat — Kilimandjaro. 
On grass. 
90515°— Nd. 16—11 61 



62 PAPERS ON" COCCIDiE OR SCALE INSECTS. 

Subfamily DACTYLOPIIN^. 

Asterolecanium ceriferunx Green. 

Asterolecanium ceriferum Green, Cocc. Ceylon, iv, p. 324 (1909). Fig. 
Habitat — Ceylon. 
On leaves of palm. 

Asterolecaniiiin ceriferum prominens Green. 

Asterolecanium ceriferum prominens Green, Cocc. Ceylon, iv, p. 326 (1909). Fig. 
Habitat — Ceylon. 
On Oxytenanthera thwaitesii. 

Asterolecanium coronatum Green. 

* Asterolecanium coronatum Green, Cocc. Ceylon, iv, p. 327 (1909). Fig. 
Habitat — Ceylon. 
On Dendrocalamus giganteus and Gigantochloa aspera. 

Asterolecanium exiguum Green. 

Asterolecanium exiguum Green, Cocc. Ceylon, iv, p. 315 (1909). Fig. 
Habitat — Ceylon. 
On under surface of leaves of bamboo. 

Asterolecanium flavociliatum Green. 

Asterolecanium fiavociliatum Green, Cocc. Ceylon, iv, p. 322 (1909). Fig. 
Habitat — Ceylon. 
On Arundinaria sp. 

Asterolecanium lanceolatum Green. 

Planchonia miliaris longa Green (part), Ind. Mus. Notes, iv, p. 5 (1896). 
^Asterolecanium lineare Green, Cocc. Ceylon, iv, p. 336 (1909). Fig. 
lanceolatum Green, Cocc. Ceylon, iv, preface (1909). 

Habitat — Ceylon. 

On leaves of Arundinaria sp. 

Asterolecanium pasaniae Kuwana and Cockerell. 
Asterolecanium pasanise Kuwana and Ckll., Jn. N. Y. Ent. Soc, xvn, 4, p. 152 (1909). 

Fior 

J- J.J,. 

Habitat — ^Japan. 

On Pasania cuspidata. 

Asterolecanium pudibimdum. Green. 

AsterolecaniuTTi pudibundum Green, Cocc. Ceylon, iv, p. 323 (1909). Fig. 
Habitat — Ceylon. 
On Arundinaria sp. 

Asterolecanium pustulans seycheUartim Green. 

Asterolecanium pustulans seychellarum Green, Jn. Econ. Biol., v, 1, p. 3 (1910). Fig. 
Habitat — Seychelles Islands. 
On Hevea brasiliensis. 

^A. lineare was first proposed for this species, but was subsequently changed bj' the author to A. lanceo- 
latum, since the former was preoccupied by a species described by Lindinger on coconut palms from Brazil. 



CATALOGUE OF RECENTLY DESCRIBED COCCID^ III. 63 

Asterolecanium ruDrocorriatiim Green. 

* Aster olecanium ruhrocomatum Green, Cocc. Ceylon, iv, p. 316 (1909). Fig. 

Habitat— Ceylon . 

On upper surface of leaves of Bambusa sp . 

Asterolecanium tenuissimum. Green. 

Asterolecanium tenuissimum Green, Cocc. Ceylon, iv, p. 318 (1909). Fig. 
Habitat — Ceylon. 
On bamboo, both smfaces of leaves. 

Asterolecanium thespesiae Green. 

* Asterolecanium thespesix Green, Cocc. Ceylon, iv, p. 331 (1909). Fig. 

Habitat — Ceylon. 

On TJiespesia populnea. 

Asterolecanium tumidum Green. 

Asterolecanium tumidum Green, Cocc. Ceylon, iv, p. 330 (1909). Fig. 
Habitat — Ceylon. 
On leaves of bamboo. 

Asterolecanium udagamae Green. 

Asterolecanium udagamse Green, Cocc. Ceylon, iv, p. 319 (1909). Fig. 
Habitat— Ceylon . 
On leaf of bamboo. 

Asterolecanium variolosum minor Leonardi. 

Asterolecanium variolosum minor Leonardi (sine descr.), Cherm. Ital., v, No. 107(1909). 
Habitat — Italy. 
On Quercus sp. 

Lecaniodiaspis azadii'aclitee Green. 

Lecaniodiaspis azadirachtx Green, Cocc. Ceylon, iv, p. 298 (1909). Fig. 
Habitat — Ceylon. 
On "margosa" {Azadirachta indica). 

Lecaniodiaspis malaboda Green. 

* Lecaniodiaspis malaboda Green, Cocc. Ceylon, iv, p. 300 (1909). Fig. 

Habitat — Ceylon . 

On "malaboda" {Myristica laurifolia) and Bocagea obliqua. 

Cerococcus albospicatus Green. 

*Cerococcus albospicatus Green, Cocc. Ceylon, iv, p. 308 (1909). Fig. 
Habitat — Ceylon . 
On bark of Symplocos obtusa. 

Cerococcus indicus Green. 

Cerococcus indicus Green, Jn. Econ. Biol., v, 1, p. 5 (1910). Fig. 
Habitat — India. 
On Helicteres isora. 

Cerococcus omatus Green. 

Cerococcus omatus Green, Cocc. Ceylon, iv, p. 306 (1909). Fig. 
Habitat — Ceylon. 
On Coffea arabica and Carissa sp. 



64 PAPERS ON" COCCID^ OR SCALE INSECTS. 

Cerococcus roseus Green. 

*Cerococcus roseus Green, Cocc. Ceylon, iv, p. 310 (1909). Fig. 
Habitat — Ceylon . 
On undetermined tree. 

Stictococcus dimorphus Newstead. 

Stictococcus dimorphus Newst., Bui. Ent. E-es., i, i, p. 63 (1910). Fig. 
Habitat — German East Africa. 
On cacao pods. 

Stictococcus formicarius Newstead. 

Stictococcus formicarius Newst., Jn. Econ. Biol., v, 1, p. 19 (1910). Fig. 
Habitat — Upper Congo, Africa. 

In hollow stems of Barteria fistulosa and Cuviera angoleiisis attended by Sima 
spininoda Andre. 

Phenacoccus oleae Marchal. 

Phenacoccus oleas Marchal, Bui. Soc. Ent. Fr., No. 13, p. 244 (1910). 
Habitat — Tunisie. 
On olive attended by a Cremastogaster. 

Ceroputo ambigua Fullaway. 

Ceroputo ambigua Fullaway, Pr. Haw. Ent. Soc, ii, 2, p. 73 (1909). 

Fullaway, Pr. Dav. Acad. Sci., xii, p. 223 (1910). Fig. 
Habitat — California. 
On pickle weed {Salicornia ambigua). 

Pseudococcus ananassse Kuwana. 

Dactylopius {Pseudococcus) ananassse Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 162 
(1909). Fig. 
Habitat — Japan. 
On pineapple. 

Pseudococcus bakeri Essig. 

Pseudococcus bakeri Essig, Pom. College Jn. Ent., ii, 4, p. 334 (1909). Fig. 
Habitat — California. 

On elder (Sambucus glauca); visdnut {Juglans regia); apple {Pyrus malus); pear 
{Pyrus communis). 

Pseudococcus boiiinsis Kuwana. 

Dactylopius {Pseudococcus) boninsis Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 161 
(1909). Fig. 
Habitat — Japan . 
On sugar cane. 

Pseudococcus coccineus Newstead. 

Dactylopius coccineus Newst., Kilimandjarc-Meru Exp., 12 (1 Coccidse), p. 8 (1908). 
Fig. 

Habitat — Kilimandjaro. 

On Acacia. 

Pseudococcus missionum Cockerell. 

Pseudococcus missionum Ckll., The Entom., xlii, 563, p. 113 (1910). 
Habitat — Argentina. 
On—? 



CATALOGUE OF KECENTLY DESCRIBED COCCID^ ITI. 65 

Pseudococcvis pemiciosiis Newstead and Willcocks. 

Dactylopius perniciosus Newstead and Willcocks, Bui. Ent. Res., i, 2 (Appendix 1), 
pp. 138-140, July, 1910. Fig. 
Willcocks, Bui. Ent. Res., i, 2, pp. 121-137, July, 1910. Fig. 
Hughes, Bui. Ent. Res., i, 2, Appendix II, p. 141, July, 1910. 
Habitat — Egypt. 

Onlebbek {Albizzia lebbek); cotton {Gossypium sp.); Christ's thorn {Zizyphus spina- 
christi); sunt tree {Acacia arabica). 

Ripersia anomala Newstead. 

Ripersia anomala Newst., Kilimandjaro-Meru Exp., 12 (1 Coccidse), p. 9 (1908). 
Fig. 
Habitat — Kilimandjaro. 
Under bark accompanied by Pheidole megacephala. 

Ripersia agasa-warensis Kuwana. 

Ripersia agasawarensis Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 161 (1909). Fig. 
Habitat — Japan . 
On Miscanthus sp. 

Ripersia smithii Essig. 

Ripersia smithii Essig, Pom. College Jn. Ent., ii, 2, p. 218 (1910). Fig. 
Habitat — California. 
" On wild rye {Elymus condensatus) . 

Ripersiella rhizophiUa Fullaway and Kotinsky. 

^ Ripersiella rhizophiUa Fullaway and Kotinsky (sine descr.), Ent. News, xxi, 1, p. 
49 (1910). 

Pollinia ceylonica Green. 

Pollinia ceylonica Green, Cocc. Ceylon, iv, p. 341 (1909). Fig. 
Habitat — Ceylon . 
On under surface of undetermined trees in dense jungle. 

Ceroplastidia Cockerell, n. subg. of Ceroplastes. Type, bruneri Ckll. 

Cockerell, Can. Ent., xlii, 3, p. 76 (1910). 
Ceroplastina Cockerell, n. subg. of Ceroplastes. Type, lahillei Ckll. 

Cockerell, Can. Ent., xlii, 3, p. 76 (1910). 

Ceroplastes ficus Newstead. 

Ceroplastes ficus Newst., Bui. Ent. Res., i, 3, p. 190 (1910). Fig. 
Habitat — German East Africa. 
On bark-cloth (Ficus sp.). 

Ceroplastes lahillei Cockerell. 

Ceroplastes (Ceroplastina) lahillei Ckll., Can. Ent., xlii, 3, p. 74 (1910). Fig. 
Habitat— Argentina . 

Ceroplastes quadrilineatus Newstead. 

Ceroplastes quadrilineatus Newstead, Bui. Ent. Res., i, 3, p. 193 (1910). Fig. 
Habitat — British East Africa. 
On Anona muricata; bark-cloth (Ficus sp.). 

1 Reduced to synonjTn of Geococcus radicum Green by David T. Fullaway, Pr. Haw. Ent. Soc., n, 3, 
p. 108 (1910). 



66 PAPEES ON COCCID^ OR SCALE INSECTS. 

Ceroplastes singiilaris Newstead. 

Ceroplastes singularis Newstead, Bui. Ent. Res., i, 3, p. 188 (1910). Fig. 
Habitat — British East Africa. 
On guava {Psidium guava). 

Ceroplastodes chiton Green. 

Ceroplastodes chiton Green (sine descr.), Mem. Dep. Ag. India, ii, 2, p. 32 (1908). 
Sanders, Catalogue Coccidai — II, p. 45 (1909). 
Green, Cocc. Ceylon, iv, p. 287 (1909). Fig. 
Habitat — India and Ceylon. 
On Cajanus indicus and Cassia sp. 

Ceroplastodes virescens Green. 

Ceroplastodes virescens Green, Cocc. Ceylon, iv, p. 288 (1909). Fig. 
Habitat — Ceylon. 
On terminal shoots of Theobroma cacao. 

Subfamily COGGING. 

Ceroneraa koebeli Green. 

*Ceronema koebeli Green, Cocc. Ceylon, iv, p. 256 (1909). Fig. 
Habitat — Ceylon. 
On Sapium sebiferuvi. 

Pulvinaria cellulosa Green. 

*Pulvinaria cellulosa Green, Cocc. Ceylon, iv, p. 262 (1909). Fig. 
Habitat — Ceylon. 
On Citrus sp. 

Pulvinaria ixorse Green. 

Pulvinaria ixorse Green, Cocc. Ceylon, iv, p. 266 (1909). Fig. 
Habitat — Ceylon. 
On Ixora coccinea. 

Pulvinaria occidentalis subalpina Cockerell. 

^Pulvinaria occidentalis subalpina Ckll., Jn. Econ. Ent., in, 5, p. 428 (1910). 
Habitat — Colorado. 
On Betula glandulosa. 

Pulvinaria thespesiae Green. 

*Pulvinaria thespesise Green, Cocc. Ceylon, iv, p. 259 (1909). Fig. 
Habitat — Ceylon. 
On ' ' suriya " ( Thespesia populnea) . 

Protopiilvinaria longivalvata Green. 

Protopulvinaria longivalvata Green, Cocc. Ceylon, iv, p. 254 (1909). Fk 
Habitat — Ceylon. 
On leaves of Piper nigrum. 

Coccus celtium Kuwana. 

Lecanium (Coccus) celtium Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 162 (1909). Fig. 
Habitat — Japan . 
On Celtis sinensis. 



CATALOGUE OF EECENTLY DESCRIBED COCCID^ III. 67 

Coccus fukayai Kuwana. 

Lecanium (Coccus) fukayai Kuwana, Jn. N. Y. Ent. Soc, xvir, 4, p. 154 (1909). Fig. 

Hab i tat — Japan . 

On vine. 

Coccus muiri Kotinsky. 

^ Coccus tuherculatus Kotinsky, Pr. Haw. Ent. Soc, i, 5, p. 168 (1908). Fig. 

Sanders, Catalogue Coccidae — II, p. 45 (1909). 
Coccus muiri Kotinsky, Pr. Haw. Ent. Soc, ii, 2, p. 37 (1909). 

Habitat — Singapore. 

On leaves of unknown tree. 

Coccus oclinaceae Kuwana. 

Lecanium {Coccus) ochnacex Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 154 (1909). Fig. 
Hab i tat — Japan . 
On Eurya ochnacea. 

Hemilecanium recurvatum Newstead. 

Hemilecanium recurvatum Newst., Jn. Econ. Biol., v, 1, p. 18 (1910). Fig. 
Habitat — Upper Congo, Africa. 
In hollow stems of Plectronia laurentii DeWild attended by Cremastogaster. 

Neolecanium crustuliforme Green. 

Neolecanium crustuliforme Green, Cocc. Ceylon, iv, p. 252 (1909). Fig. 
Habitat — Ceylon. 
On small branches of undetermined tree. 

Lecanium mori var. somereni Newstead. 

Lecanium mori var. somereni Newst., Bui. Ent. Res., i, 3, p. 187 (1910). 
Habitat — British East Africa. 
On Morus sp. 

Lecanitun resinatum Kieffer and Herbst. 

Lecanium resinatum Kieffer and Herbst, Centr. f. Bakt. Par. u. Infekt., zweite 
Abt., XXIII, 1-5, p. 122 (1909). 
Habitat— Chile. 
On Baccharis rosm,arinifolia. 

Saissetia pseudonigrum Kuwana. 

Lecanium (Saissetia) pseudonigrum Kuwana, Jn. N. Y. Ent. Soc, xvn, 4, p. 162 
(1909). Fig. 
Habitat — Japan. 
On "shirotsugi." 

Saissetia sideroxylium Kuwana. 

Lecanium (Saissetia) sideroxylium Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 162 (1909). 
Fig. 
Habitat — ^Japan . 
On Sideroxylonferrugineum. 

Aclerda distorta Green. 

^Aclerda distorta Green, Cocc. Ceylon, iv, p. 290 (1909). Fig. 
Habitat — Ceylon. 
Beneath the enshea thing bases of leaves of Arundinaria sp. 

1 Specific name preoccupied. 



68 PAPERS ON COCCID^ OR SCALE INSECTS. 

Takahashia citricola Kuwana. 

Takahashia citricola Kuwana, Jn. N. Y. Ent. Soc, xvn, 4, p. 153 (1909). Fig. 

Habitat — Japan. 

On Citrus sp. 

Lichtensia japonica Kuwana. 

Lichtensia japonica Kuwana, Jn. N. Y. Ent. Soc, xvn, 4, p. 152 (1909). Fig. 
Habitat — Japan . 
On Thea japonica. 

Inglisia chelonioides Green. 

Inglisia chelonioides Green, Cocc. Ceylon, iv, p. 283 (1909). Fig. 
Habitat — C eylon . 
On Gelonium lanceolatum. 

Inglisia conchiformis Newstead. 

Inglisia conchiformis Newst., Bui. Ent. Res., i, 3, p. 185 (1910). Fig. 
Habitat — West Africa and British East Africa. 
On Averrhoa carambola and Harogana madagascariensis. 

Subfamily DIASPIN^. 

Chionaspis dentilobis Newstead. 

Chionaspis dentilobis Newst., Bui. Ent. Res., i, 3, p. 195 (1910). Fig. 
Habitat — British East Africa. 
"On—?" 

Chionaspis kinshinensis Kuwana. 

Chionaspis kinshinensis Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 155 (1909). Fig. 
Habitat — ^Japan . 
On Quercus sp. 

Chionaspis sassceri Cockerell and Robbins. 

* Chionaspis sassceri Ckll. and Robb., Jn. N. Y. Ent. Soc, xvn, 3, p. 105 (1909). Fig. 

Habitat — California. 

. On orange. 

. Chionaspis substriata Newstead. 

Chionaspis substriata Newst., Bui. Ent. Res., i, 3, p. 197 (1910). Fig. 

Habitat — British East Africa. 

On palms. 

Diaspis africana Lindinger. 

Diaspis africana Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 22 (1909). Fig. 
Habitat — South Africa. 
On leaves of Connarum smeathmanni and Syzygium guineense. 

Diaspis (?) chionaspiformis Newstead. 

Diaspis (f) chionaspiformis Newst., Bui. Ent. Res., i, 3, p. 198 (1910). Fig. 
Habitat — British East Africa. 
On an unknown plant. v 

Diaspis pugionifera Lindinger. 

Diaspis pugionifera Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 24 (1909). Fig. 
Habitat — South Africa. 
On leaves of Mitragyne maerophylla. 



CATALOGUE OF EECENTLY DESCRIBED COCCID^ — III. 69 

Diaspis stilosa Lindinger. 

Diaspis stilosa Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 25 (1909). Fig. 
Habitat — South Africa. 
On leaves of Strychnos cinnabarina . 

Leucaspis indias-orientalis Lindinger. 

^ Leucodiaspis indise-orientalis Lindgr. (sine descr.), Zeitschr. f. wiss. InsektenbioL, 
VI, 11, p. 373 (1910). 
Habitat — India . 
On Pinus sp. 
Cryptodiaspis Lindinger, n. subg. of Diaspis. Type, conservans. 

Lindinger, Jahrb. Hamb. wiss. Anst., xxvi, p. 26 (1909). 

Diaspis conservans Lindinger. 

Diaspis (Cryptodiaspis) conservans Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 26 
(1909). Fig. 
Habitat — South America. 
On leaves of Euphorbiacese. 

Diaspis hamata Lindinger. 

Diaspis (Cryptodiaspis) hamata Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 28 (1909). 
Fig. 
Habitat — South Africa. 
On leaves of Macrolohium zenkeri. 

Diaspis limuloides Lindinger. 

Diaspis (Cryptodiaspis) limuloides Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 30 
(1909). Fig. 
Habitat — South Africa. 
On leaves of Cynometra sp. with Aonidia biafrse Lindgr. 

Aspidiotus arctostaphyli Cockerell and Robbins. 

Aspidiotus arctostaphyli Ckll. and Robb., Jn. N. Y. Ent. Soc, xvii, 3, p. 104 (1909). 
Fig. 
Habitat — California . 
On leaves of Arctostaphylos viscida. 

Aspidiotus fissidens Lindinger. 

Aspidiotus fissidens Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 14 (1909). Fig. 
Habitat — South Africa. 
On leaves of Parinarium gabunense and Strychnos cinnabarina. 

Aspidiotus mseandrius Lindinger. 

Aspidiotus mseandrius Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 15 (1909). Fig. 
Habitat — South Africa. 
On leaf of Dichapetalum sp. 

Aspidiotus mori Herrick. 

* Aspidiotus (Diaspidiotus) mori Herrick, Ent. News, xxi, 1, p. 22 (1910). Fig. 
Habitat — Texas . 
On Morus rubra. 

1 Leucodiaspis is not valid. See Bui. U. S. Btir. Ent., Tech. Ser. 16, Pt. n, p. 50 (1909). 



70 PAPESS OlSr COCCID^ OR SCALE INSECTS. 

Aspidiotus pectinatus Lindinger. 

Aspidiotus pectinatus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 43 (1909). Fig. 
Habitat — South Africa. 
On pear. 

Aspidiotus replicatus Lindinger. 

Aspidiotus replicatus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 17 (1909). Fig. 
Habitat — South Africa. 

On leaves of Ehretia cymosa, Illigera pentophylla, Mitragyne viacrophylla, and 
Anacardiaceae. 

Aspidiotus seurati Marchal. 

Aspidiotus (Hemiberlesia) seurati Marchal, Bui. Soc. Ent. Fr., p. 71 (1911). 
Habitat — South Algeria. 
On Zilla macroptera. 

Aspidiotus spiniger Lindinger. 

Aspidiotus spiniger Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 19 (1909). Fig. 
Habitat — South Africa. 
On leaf of Strombosiopsis tetranda. 

Aspidiotus undulatus Lindinger. 

Aspidiotus undulatus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 20 (1909). Fig. 
Habitat — South Africa. 
On leaves of Acioa pallescens and Strychnos cinnabarina. 

Aonidia biafrse Lindinger. 

AoniSia biafrx Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 40 (1909). Fig. 
Habitat — South Africa. 
On leaves of Schotia humboldtioides, Crudia zenleri, and Cynometra sp. 

Aonidia dentata Lindinger. 

Aonidia dentata Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 1, p. 12 (1911). Fig. 
Habitat — India. 
On leaf of Walsura piscidis. 

Aonidia spinosissima Lindinger. 

Aonidia spinosissima Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, p. 12 (1911). Fig. 
Habitat — Central India. 
On Mimusops hexandra. 

Gymnaspis aberemose Lindinger. 

Gymnaspis aberemox Lindgr., Deutsche Ent. Zeitschr., iv, p. 437 (1910). Fig. 
Habitat — Brazil. 
On Abererrwa rhizantha. 

dirysomphalus portoricensis Lindinger. 

Chrysomphalus (Melanaspis) portoricensis Lindgr., Zeitschr. f. wiss. Insektenbiol., 
VI, 12, p. 441 (1910). Fig. 
Habitat — Venezuela. 
On leaf of Bletia sp. 



CATAIiOGUE OF RECENTLY DESCRIBED COCCID^ III. 71 

Targionia celtis Herrick. 

1 Targionia celtis Herrick, Can. Ent., xliii, 11, p. 373 (1910). Fig. 
Habitat — Texas. 
On Celtis occidentalis. 

Targionia vitis arbutus Leonardi. 

Targionia vitis arbutus Leon, (sine descr.), Cherm. Ital., v, p. 123 (1909). 
Habitat — Italy. 
On Arbutus unedo. 

Pseudoparlatoria cristata Lindinger. 

Pseudoparlatoria cristata Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, p. 10 (1911). 
Fig. 
Habitat — Brazil . 
On leaf of Gnetum leyboldi. 

Cryptaspidiotus mediterraneus Lindinger. 

Cryptaspidiotus mediterraneus Lindgr. (sine descr.), Zeitschr. f. wiss. Insektenbiol., 

Yi, 11, p. 373 (1910). 
Lindgr., Zeitschr. f. wiss. Insektenbiol., vi, 12, p. 437 
(1910). Fig. 
Habitat — Algeria . 
On leaves of Juniperus phcenicea and Callitris quadrivalvis. 

Ischnaspis bipindensis Lindinger. 

Ischnaspis bipindensis Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 32 (1909). Fig. 
Habitat — South Africa 
On leaves of Cyclostemon bipindensis and Strychnos cinnabarina. 

Lepidosapbes aberrans Lindinger. 

Lepidosaphes aberrans Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 33 (1909). Fig. 
Habitat — South Africa. 
On leaf of Cynometra sp. 

Lepidosaphes arii Kuwana. 

Mytilaspis {Lepidosaphes) arii Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 163 (1909). 
Fig. 
Habitat — Japan. 
On Miscanthus sp. 

Lepidosaphes buzenensis Kuwana. 

Mytilaspis (Lepidosaphes) buzenensia Kuwana, Jn. N. Y. Ent. Soc, xvii, 4, p. 155 
(1909). Fig. 
Habitat — Japan. 
On vine. 

Lepidosaphes chitinosus Lindinger. 

Lepidosaphes (Coccomytilus) chitinosus Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, 
p. 34 (1909). Fig. 
Habitat — South Africa. 
On Berlinia. 

1 Not a valid species; synonym of Targionia cueroensis (Ckll.). 



72 PAPBES ON COCCIDiE OE SCALE INSECTS. 

Lepidosaphes crudise Lindinger. 

Lepidosaphes crudise Lindgr., Jahrb. Hamb. wi?s. Anst., xxvi, p. 35 (1909). Fig. 
Habitat — South Africa. 
On leaf of Crudia zenkeri. 

Liepidosaph.es kamertuiensis Lindinger. 

Lepidosaphes kamerunensis Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 37 (1909). 
Fig. 
Habitat — South Africa. 
On leaves of Loranthus sp. 

Lepidosaplies meridionalis Lindinger. 

Lepidosaphes meridionalis Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 38 (1909). 

Fig- 
Habitat — South Africa. 
On Macrolohium palisoti. 

Lepidosaphes tenuior Lindinger. 

Lepidosaphes tenuior Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, p. 40 (1909). Fig. 
Habitat — South Africa. 
On leaf of Cynometra sp. 

Lepidosaphes tuiiloba Kuwana. 

Mytilaspis {Lepidosaphes) uniloba Kuwana, Jn. N. Y. Ent. See, xvri, 4, p. 156 (1909). 
Fig. 
Habitat — ^Japan. 
On Osmanthus. 

Parlatoria hastata Lindinger. 

Parlatoria hastata Lindgr. (sine descr.), Zeitschr. f. wiss. Insektenbiol., vi, 11, p. 375 
(1910). 
Habitat — Not given. 



CORRECTIONS TO CATALOGUE 11—1909. 

Page 38, instead of lines 9 and 10 under "Stictococcus multispinosus Newstead," 
read: 

Stictococcus multispinosus Newst., Kilimandjaro-Meru Exp., 12 (1 Coccidse), p. 6 (1908). Fig. 
Newst., Jn. Econ. Biol., n, 4, p. 150, footnote (1908). 
Habitat — Kilimandjaro, East Africa. 

Page 38, line 12 from bottom, for cxliii read cxlviii. 

Page 46, line 15, for Lecanium cecconi Leonardi read Lecanium cecconi (Leonardi). 
Page 46, line 19, for Lecanium glandi (Kuwana) read Lecanium. glandi Kuwana. 
Page 46, line 24, for Lecanium kunoensis (Kuwana) read Lecanium kunoensis 

Kuwana. 
Page 46, line 13 from bottom. Lecanium ruhellum Lindinger on Calluna vulgaris, 

given as a synonym of Lecanium pulchrum Marchal, has been resurrected as a valid 

species by Mr. J. G. Sanders. 
Page 47 , line 18 . Hemilecanium, theohromse Newstead = Hemilecanium imbricans Green, 

on the authority of Green. 
Page 55, line 16, for cxliii read cxlviii. 



ABBREYIATIONS OF LITERATURE. 

AUg. Zeits. f. Ent. — AUgemeine Zeitschrift fiir Entomologie. 

Agr. News West Ind. — Agricultural News. West Indies. 

An. Soc. Cient. Argentina. — Anales de la Sociedad Cientifica Argentina. 

Ann. Soc. Ent. France. — Annales de la Societe Entomologique de France. 

Annali di Agr. — Annali di Agricoltura. Portici. 

Ann. Mag. N. H. — Annals and Magazine of Natural History. 

Ann. Mus. Zool. Ac. Imp. Sc. St. Petersboiu-g. — Annuaire Musee Zoologique Academie 

Imperiale des Sciences de St. Petersbourg. 
Berl. Ent. Zeitschr. — Berliner Entomologische Zeitschrift. 

Ber. Stat. f. Pflanzensch. Hamb. — Bericht der Station fiir Pflanzenschutz, Hamburg. 
Bol. Min. Ag. — Boletin del Ministerio de Agricultura. Argentina. 
Bol. Zool. Sc. Sup. Portici. — Bollettino del Laboratorio di Zoologia generale e agraria 

della R. Scuola Superiore d 'Agricoltura di Portici. 
Bot. Jahrb. — Botanische Jahrbiicher. 
Bui. Ag. Exp. Sta. Japan. — Bulletin of the Imperial Central Agricultural Experiment 

Station. Japan. 
Bui. Ent. Res. — Bulletin of Entomological Research. 

Bui. 111. St. Lab. N. H. — Bulletin of the Illinois State Laboratory of Natural History. 
Bui. Soc. Ent. Fr. — Bulletin de la Societe Entomologique de France. 
Bui. Soc. Ent. Ital.— Bulletino della Societa Entomologica Italiana. 
Bui. Soc. Zoo. France. — Bulletin de la Societe Zoologique de France. 
Bui. TJ. S. Bur. Ent.,t. s.l6, Pt. II.— Bulletin, Technical Series, Bureau of Entomology, 

U. S. Department of Agriculture. 
Can. Ent.-^Canadian Entomologist. 

Catalogue Coccidse— I.— U. S. Bur. Ent., technical series, Bui. 12, Pt. I (1906). 
Catalogue Coccidge— II.— U. S. Bur. Ent., technical series, Bui. 16, Pt. Ill (1909). 
Centr. f. Bakt. Par. u. Infekt. — Centralblatt fiir Bakteriologie und Parasitenkunde 

und Infektionskrankheiten. 
Cherm. Ital. — Chermotheca Italica. 
Cocc. Ceylon. — Coccidae of Ceylon. 

Compt. Rend. Ac. Sci., Paris. — Comptes rendus hebdomadaires des seances de 1' Aca- 
demie des sciences. Paris. 
Compt. Rend. Soc. Biol. — Comptes rendus des seances de la Societe de Biologic. Paris. 
Deutsche Ent. Zeitschr. — Deutsche Entomologische Zeitschrift. 
Ent. Mo. Mag. — The Entomologist's Monthly Magazine. 
Ent. Blatter. — Entomologische Blatter, Internationale Monatsschrift fiir die Biologie 

der Kafer Europas. Schwabach. 
Ent. News. — Entomological News. 

Ent. Rec. Jn. Variation. — The Entomologist's Record and Journal of Variation. 
Ent. Wochenblatt. — Entomologisches Wochenblatt (antea Insekten-Borse). 
Ind. Mus. Notes. — Indian Museum Notes. 
Insekten Borse. — Entomologische Wochenblatt. 

Jahrb. Hamb. wiss. Anst. — Jahrbuch der Hamb urgischenwissenschaftlichenAnstalten. 
Jn. Bomb. N. H. Soc. — Jom-nal of the Bombay Natural History Society. Calcutta. 
Jn. Econ. Biol. — The Journal of Economic Biology. 
Jn. N. Y. Ent. Soc. — Journal of the New York Entomological Society. 

73 



74 PAPERS ON COCCID^ OB SCALE IlirSECTS. 

Kilimandjaro-Meru Exp. — Schwedischen Zoologischen Expedition nach Dem Kili- 
mandjaro, DemMeru und DenUmgebendenMassaisteppenDeutsch-Ogtafrikas. 

Liverpool Univ. Quart. Jn. — Institute of Commercial Research in the Tropics, Liver- 
pool University, Quarterly Journal. 

Medd. Soc. Faun. Fenn. — Meddelanden af Societas pro Fauna et Flora Fennica. 

Mem. Dep. Ag. India. — Memoirs of the Department of Agricultm-e in India. 

Mem. Soc. Cient. Ant. Alz. — Memorias y Revista de la Sociedad Cientifica "Antonio 
Alzate." 

Mem. Soc. Zool. France. — Memoires de la Societe Zoologique de France. 

Nachrichtsbl. Naturfrsch.-Klubs, Prossnitz. — Nachrichtsblatt Naturforscher-Klubs, 
Prossnitz (Mahren). 

N. Z. Trans. — Transactions of the New Zealand Institute. 

Pests and Blights of Tea Plant.— The Pests and Blights of the Tea Plant, 2d ed. Watt 
& Mann, Calcutta, 1903. 

Philippine Jn. Sci. — The Philippine Jom-nal of Science. 

Pomona College Jn. Ent. — Pomona College Journal of Entomology. Claremont, 
California. 

Pr. Biol. Soc. Wash. — Proceedings of the Biological Society of Washington. 

Pr . Dav. Acad. Sci. — Proceedings Davenport Academy of Sciences. Davenport, Iowa. 

Pr. Ent. Soc. Wash. — Proceedings of the Entomological Society of Washington. 

Pr. Haw. Ent. Soc. — Proceedings of the Hawaiian Entomological Society. 

Redia. — Redia. 

Riv. Pat. Veg. — Rivista di Patologia Vegetale. 

Scale Ins. of Egypt. — Notes on the Injmious Scale Insects and Mealy Bugs of Egypt. 
Cairo, 1907. 

The Entom. — The Entomologist. 

Trab. Mus. Farm. Fac. Cienc. Med. — Trabajos del Museo de Farmacologia de la Facul- 
tad de Ciencias Medicas. Buenos Aires, Argentina. 

Tr. Linn. Soc. Lond. — Transactions of the ?jinngean Society of London. 

Victorian Nat. — The Victorian Natxiralist. 

Zeitschr. f. wiss. Insektenbiol. — Zeitschrift fiir wissenchaftliche Insektenbiologie. 

o 



Technical Series, No. 16, Part V. * 

U. S. DEPARTMENT OF AGRICULTURE. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



PAPERS ON COCCIDJ; OR SCALE INSECTS. 



THE GENUS FIORINIA IN THE 



By E. R. SASSCER, 

Scientific Assistant. 



Issued Decembee 6, 1912, 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1912. 



S. SUPT. C 



CONTENTS. 



Introduction ' 75 

Description of the Genus Fiorinia 75 

Fiorinia ihex Green 76 

Fiorinia fiorinix Targioni Tozetti 79 

Fiorinia fiorinise minor Maskell 79 

Fiorinia Jionnix japonica Kuwana 81 



ILLUSTRATIONS. 



PLATES. Pago. 

Plate X. Anal plate of Fiorinia thex 7d 

XI. Fig. 1. — i^t'onma i/ieas on leaf of Camellia. Fig. 2. — F'iorinia thex on 

leaf of Camellia 76 

XII. Anal plate of Fiorinia fiorinix 80 

XIII. Anal plate of Fiorinia fiorinix minor 80 

u 



ADDITIONAL COPIES of this publication 
xi- may be procured from tlie Supekintend- 
ENT OF Documents, Government Printing 
Ofiice, Washmglou, i). C. at 10 cents per copy 



U. S. D. A., B. E. Tech. Ser. 16, Part V. Issued December 6, 191 

PAPERS ON COCCID^ OR SCALE INSECTS, 



THE GENUS FIOEmiA IN THE UNITED STATES, 

By E. R. Sasscer, Scientific Assistant. 
INTRODUCTION. 

The genus Fiorinia at the present time consists of some 30 described 
species and 4 varieties, and of this number there are only 2 species 
and 1 variety estabhshed in this country, viz, Fiorinia jiorinix 
Targioni Tozetti, F. these Green, and F. jiorinix japonica Kuwana. 
Judging from the available records at hand, it is quite evident thut 
the first two coccids mentioned were imported on ornamental plants, 
and their status in some locations is such as to warrant the use of 
remedial measures. The last-named variety oi jiorinix is a Japanese 
importation, and while it appears to be established in only one State, 
it is not infrequently collected at quarantine on imported evergreens. 

Fiorinia Takgioni. 

This genus is characterized by the adult female being smaller, 
decreasing in size after the second molt, and entirely inclosed in the 
second larval exuvia; puparium elongate, with first larval exuvia 
extending beyond the margin. Male scales white, elongate, with 
or without carinsB, larval exuvia at anterior extremity, Pygidium 
of adult female {Fiorinia s. str.), possessing well-defined median lobes, 
and some species exhibiting a small pair on either side; paragenitals 
normally in five groups, although in some species the median and 
anterior laterals coalesce, forming an arch. 

Leonard! in his paper,^ ''Saggio de Sistematica delle Fiorionise" 
divides Fiorinia sens. lat. into three subgenera, viz, TruUijiorinia 
(paragenitals and lobes present), Anamejiorinia (paragenitals present, 
lobes absent), and Adiscqfiorinia (paragenitals absent), Fiorinia s. 
str. is characterized by possessing paragenitals and plates, and the 
species discussed in this paper come distinctly within this division. 

iRedia, vol. 3, p. IS, 1905. 
67»72»— 12 75 



76 PAPEES ON COCCID^ OR SCALE INSECTS. 

The following table, based on characters exhibited by the adult 
female, is offered as a means for separation of the species occurring 
in the United States. 

I. Spatulate protuberance between antennse present, first pair of lateral lobes rep- 

• resented by serrate thickenings F. these Green. 

II. Spatulate protuberance between antennse absent, first pair of lateral lobes 
prominent. 

(a) Median and anterior lateral paragenitals usually contiguous, 21-23; 

posterior laterals 10-16 F. fioriniae Targ. 

(6) Paragenitals massed and numerous; median 3-11, anterior laterals 16-24, 
posterior laterals 19-31 F. fiorinim var. japonica Kuw. 

THE TEA SCALE. 

Fiorinia these Green. 
Plates X and XI. 

Fiorinia fiorinix var. camelliae RoUe and Quaintance, Coccidse Americange. Decades 
i-iii, No. 6 (1898). 

Fiorinia these Green, Indian Museum Notes, V, No. 1, p. 3 (1900). Fig. 

Fiorinia these Stebbing, Insects that affect forestry. No. 1, p. 133 (1902). Fig. 

Fiorinia these Green, Indian Museum Notes, V, No. 3, p. 102 (1903). Record only. 

Fiorinia these Fernald, Catalogue of the Coccidse of the world, p. 250 (1903). 

Fiorinia these Watt and Mann, The pests and blights of the tea plant, p. 306, Calcutta, 
2d edition (1903). Fig. 

Fiorinia these Leonardi, Redia, III, p. 28 (1905). Fig. "Saggio di Sistematica Delle 
Fiorinise." 

Fiorinia these Green and Mann, Memoirs of the Department of Agriculture in India, 
Entomological Series, I, No. 5, pp. 343, 350 (1907). Bibliography and record only. 

Fiorinia these Green, Memoirs of the Department of Agriculture in India, Entomo- 
logical Series, II, No. 2, p. 39 (1908). Record only. 

Fiorinia thex Stebbing, Manual of forest zoology for India, p. 165 (1908). Fig. 

Fiorinia thex Lindinger, Zeitschr. f. wiss. Insektenbiol., VII, 4, p. 127 (1911). 

Notwithstanding the fact that the tea scale, Fiorinia these, Green, 
has been in the National Collection of Coccidse since 1887 identified 
as F.fiorinisR Targ., it was not described until 1900 by Green on tea 
from Assam and Kangra, India. A thorough study of the material 
in this collection revealed the fact that all the specimens on Camellia, 
with the exception of Comstock's type of F. camellise, which is now 
recognized as a synonym of F. fiorinix Targ., are the tea scale of 
Green, 

In 1897 Cockerell ^ recorded what he considered a distinct variety 
of F. fiorinise Targ. on the underside of a leaf of Camellia japonica 
from Baton Rouge, La., and, judging from his partial description he 
surely had reference to F. these Green. 

A microscopic study of the material in "Coccidse Americanse" 
(Decades i-iii. No. 6 [18981), distributed by Rolfs and Quaintance 

I Bui. Bot. Dept. Jamaica, n. s. 4, p. 149 (1897). 



Tech. Series 16, Part V, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE X. 



Tech. Series 16, Part V, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate XI. 




FlQ. 1.— FlORINIA THE/C ON LEAF OF CAMELLIA. TWICE ENLARGED. (ORIGINAL.) 




Fig. 2.— Fiorinia the/e on Camellia Leaf. Enlarged Five Times. (Original.) 



GEFUS FIORINIA IN THE UNITED STATES. 77 

and labeled F.Jiorinise var. camellise, proves it to be F. these, Green and 
not a variety. 

In March, 1908, j\'Ir. George F. Mitchell, of the Experimental Tea 
Farm at Summerville, S. C, submitted leaves of tea thickly infested 
with Fiorinia these Green and stated that it was becoming a serious 
enemy of this plant. At the request of Dr. R. H. True, Mr. J. G. 
Sanders, at that time an agent of the Bureau of Entomology, visited 
the tea farm and found this scale in abundance on Assam hybrid, 
Darjeeling, and China teas and camellias grown for ornamental pur- 
poses. Mr. Sanders observed that the Darjeeling tea which was 
grown in moist lowland was frequently covered with a brown fungus, 
which was apparently parasitic on the scale. 

It is the behef of Dr. Charles U. Shepard, who is in charge of the 
experimental farm, that the tea affords better food than Camellia 
japonica, and he states that it is seldom if ever that the latter plant 
is killed by attack of this insect alone. 

Since no plants have been introduced from Asiatic regions, all 
being grown from seed, it is extremely probable that its introduction 
was through the agency of the cameHias, which have been for a 
number of years greatly in demand as ornamental plants in this 
country. 

According to Watt and Mann ^ this scale is of common occurrence 
in both Assam and Kangra, and is the most prevalent of all coccids 
in the former Province. Mr. Stebbing ^ records it on the leaves of 
the olive, Olea glandulifera, in the northwestern Himalayas, fre- 
quently causing the leaves to turn yellow and drop off. 

Although this insect occurs on both sides of the leaves of tea and 
Camellia japonica, it is more commonly found on the underside, and 
if present in sufficient numbers it can be easily detected by yellow 
markings plainly to be seen on the upper surface. 

Fiorinia these Green can be readily separated from F.Jiorinise Targ. 
by the larger and darker scale, the presence of a proboscis-like pro- 
jection between the antennse which have no stout spines, and in the 
absence of lateral lobes on the pygidium. 

Scale of female. — Elongate, narrow, dark brown in color, usually 
darker than F. fiorinix, with a distinct dark median longitudinal 
carina; adult insect entirely inclosed in the hardened exuvia of the 
second stage, wliich varies from 1 to 1.24 mm. in length, and does not 
bear a secretionary margin. Length, 1 to 1.40 mm. Breadth, 0.40 
to 0.60 mm. 

Scale of male. — Snow-white, sides nearly parallel, usually indis- 
tinctly tricarinate, pelUcle pale yellow approaching lemon-yellow at 
extreme posterior tip. When present in large numbers the puparia 

1 The Pests and Blights of the Tea Plant (second edition), p. 306 (1903). 

2 Manual of Forest Zoology of India, p. 166 (1908). 



78 PAPERS ON COCCIDiE OR SCALE INSECTS. 

are frequently covered with a flocculent secretion given off by the 
larvae. Length, 0.80 to 1 mm.; breadth, about 0.40 mm. ^ 

Eggs: Arranged in double rows inclosed within the second larval 
exuvia. 

Female} — "Adult female pale yellow, of normal form. Antennae 
close together, on anterior margin; each antenna consisting of an 
irregular tubercle with a single curved bristle on one side. From 
between the antennae springs a stout spatulate process which is not 
chitinous but of the same consistency as the surrounding parts of 
the body. Margin of thorax and abdomen with a series of minute 
spinneret ducts opening on to small conical tubercles. Pygidium 
(PI. I) with a conspicuous median cleft, on the margin of which are 
situated the moderately large serrate median lobes. First pair of 
lateral lobes represented only by small serrate thickenings of the 
margin; second lateral lobes obsolete." [Although the lateral lobes 
are wanting in the adult, they are quite prominent on the second 
larval exuvia and are of the normal F.jiorinia& type.l "Spines nor- 
mal, the dorsal series rather long; one pair springing from within 
the median cleft. Circumgenital glands in five groups; the median 
and upper lateral groups together forming an almost continuous 
arch. Median group with four or five orifices; upper laterals, 10 to 
13; lower laterals, 15 to 18. A very few circular pores with accom- 
panying ducts, on dorsal surface, near the margin. Length, 0.50 
to 0.75 mm." 

DISTRIBUTION. 

The tea scale is recorded on Camellias in Alabama, District of 
Columbia, Florida, Georgia, Louisiana, North Carolina, and South 
Carolina. It has been collected in Ceylon, Philippine Islands, and 
India, occurring in the following localities in the latter country: 
Kangra VaUey, Assam, Bashahr State close to Kilta in the Sutlej 
Valley, Calcutta, and the northwestern Himalayas. 

FOOD PLANTS. 

In the United States this scale has only been collected on camellias 
and tea, but in India it has been recorded on tea, olive {Olea glandu- 
lijera), and citrus. In the Philippine Islands it occurs on a species 
of Caryola, and Mr. R. S. Woglum has collected it on Ostodes in the 
Royal Botanical Gardens of Peradenyia, Ceylon. Although F. these 
seems to show a preference to the tea at Summerville, S. C, it appears 
to be quite a serious pest on the camellias in several of the more 
Southern States, and is not infrequently found associated with Le'pi- 
dosaphes lasianthi Green. 

1 Owing to the accuracy of Mr. E. E. Green's description of this species it has heen deemed wise to give 
it verbatim. (Indian Museum Notes, 5, No. 1, p. 3 (1900).) 



GENUS FIOEINIA IN THE UNITED STATES. 79 

ENEMIES. 

The following predaceous beetles which have been determined by 
Mr. E. A. Schwarz, of the Bureau of Entomology, were found by Mr. 
Sanders preying on the tea scale: CMlocorus livulnerus Muls., Micro- 
weisea misella Lee, and Cyhocephalus nigritulus Lee. As stated 
elsewhere, when on the Darjeeling tea these coccids are frequently 
covered by a parasitic fungus, which doubtless in a measure is effec- 
tive in holding this pest in check. 

Fiorinia fiorinise Tarq. 

Plates XII and XIII. 

Diaspis fiorinisB Targioni, Studii sul. Cocciniglie, p. 14 (1867). 

Chermes arecx Boisduval, Insectologie Agricole, p. 262 (1868). 

Fiorinia pelludda Targioni, Catalogue, p. 42 (1868). 

Fiorinia camellix Comstock, Rep. U. S. Dept. Agr. 1880, p. 329 (1881). 

TJiileria fiorinise. Comstock, 2nd Rep. Dept. Ent. Cornell Univ., p. Ill (1883). 

Fiorinia palmae Green, Ind. Mu3. Notes, IV, p. 5 (1896). 

This coccid is a very cosmopolitan species and is of common occur- 
rence on kentias in greenhouses. A complete bibliography of this 
species is not given, since it has been the subject of numerous papers, 
many of which, however, are of little value either from a scientific 
or economic standpoint. 

DISCUSSION OF SYNONYMY. 

A thorough study of the material on Kentia helmoreana, Cycas revo- 
luta, and Camellia which was described by Comstock ^ as Fiorinia 
camellise proves this species to be none other than Fiorinia Jiorinise 
Targ. In 1896 Green ^ described a form known as Fiorinia falmse., 
but he later reduced it to synonymy. Leonardi,^ in his paper enti- 
tled ''Saggio di Systematica Delle Fiorinise," reduces Fiorinia fiori- 
nise, var. minor Maskell to synonymy, since Maskell in his meager 
description merely states that it was given varietal rank, owing to 
its small size. Assuming that the only character of separation was 
the size of the scale, Leonardi concluded that the difference was too 
inadequate to justify even varietal rank. 

A study of the Maskell material revealed a number of characters 
which are surely of sufficient importance to allow it to remain as a 
good variety if not raised to specific rank. As will be seen by exam- 
ining Plate IV, the median lobes in minor are larger and more chitin- 
ized, the marginal pores more numerous, varying from six to seven, 
and the general shape of the pygidium is different. The material of 
this variety is very scanty and is on the imderside of a leaf which is 

> Rep. U. S. Dept. Agr., 1880, p. 329 (1831). « Redia, IH, p. 16 (1905). 

'Indian Mus. Notes, IV, p. 6 (1S96). 



80 PAPERS ON COCCli)^ OE SCALE INSECTS. 

quite hairy, causing the scales to assume various shapes and possibly- 
preventing them from reaching their normal dimensions. In size 
the pygidium of Fiorinia jiorinise. Targ. and Fiorinia Jiorinise var. 
minor Mask, compare very favorably. 

DISTRIBUTION. 

Fiorinia Jiorinise Targ. has a wide range of distribution and is 
known to occur in the following countries: Algiers, Australia, Bar- 
bados, Brazil, Ceylon, China, Egypt, Europe (Belgium and Spain), 
Jamaica, Japan, Madeira(?), Mauritius, Mexico, New South Wales, 
Peru, United States (Alabama, California, Colorado, District of Colum- 
bia, Hawaiian Islands, Louisiana, Maryland, and Massachusetts), 
West Africa, and Zanzibar. 

FOOD PLANTS. 

The food plants of this insect are numerous, some of the most 
important of which are the following: 

Anthurium acaule, Appolonias canuriensis, Aralia, Areca aurea, 
bamboo, bay, Camellia, Celtis mauritiana, Chamserops liumuli, coco- 
nut palm, Cupressus, Cycas circinalis, C. revoluta, Dracsena indivisa, 
ferns, Ficus elastica, Ficus sp., Garcinia sp., Hedera Tielix, Japanese 
quince, Kentia helmoreana, K. fosteriana, Larix sp., Leptospermum, 
Licula, Livistona, Per sea gratissima, PTioenix canariensis, Phcenix sp., 
Phormium tenax, Phytelephas macrocarpa, Podocarpus, Strelitzia regi- 
nix, tea {TJiea japonica). 

ENEMIES. 

In studying the slides of the National Collection of Coccidse one 
partially mature specimen of the parasite AspidiotipJiagus citrinus 
Craw was discovered inclosed m the body of an adult insect collected 
in Washington, D. C, on CTisemerops Jiumuli and the same chalcid 
has also been reared from material on Persea gratissima from Hono- 
lulu, Hawaii. In addition to the above a species of AspidiotipJiagus 
was reared from this scale on an unknown plant collected at Hong- 
kong by Mr. Koebele, and he also collected Prospaltella aurantii 
How. from this coccid on Ficus sp. at Swatow, Canton, China. 

Among the fungous enemies of this coccid the redheaded fungus 
(SpJiEerostilhe coccophila) has been recorded ' by Mi-. John Parkin 
from Mauritius on infested Camellia. 

DESCRIPTION. 

Scale of female. — Elongate, narrow, revealing a distinct median 
longitudinal ridge; first larval exuvia yellow, small, and extending 
beyond the margin; second larval exuvia ranging from 1.20 to 1.46 

' Ann. Roy. Bot. Gardens Peradeniya, III, p. 67 (1906). 



Tech. Series 16, Part V, Bureau of Entomology, U. S. Dept. of Agriculture. PLATE XII. 




Tech. Series 16, Part V, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate XIII. 




GENUS FIOEIlSriA IN THE UNITED STATES. 81 

mm. in length and varying from brown to dark brown with frequently 
a white secretionary margin, and inclosing the adult insect, which is 
situated in the anterior portion of the scale. Length 1 to 1.25 mm. 
Breadth about 0.50 mm. 

Eggs. — Yellow and arranged in a double row. 

Scale of male. — White, tricarinate, larval exuvia pale yellow, 
Length about 1 mm. Breadth about 0.40 mm. 

Female. — Thin and much contracted after oviposition, elongate, 
possessing rudimentary antennae, which are apparently two-jointed, 
the first consisting of a fleshy tubercle, which bears the second joint 
and a bristle, the second joint occasionally possessing a short lateral 
branch near the tip; anal plate triangular, 0.187 to 0.204 mm. in 
width, slightly truncate at tip, median notch distinct and formed by 
the median lobes, which are oblique and serrated along their entire 
free edge; second lobes prominent both in second larval exuvia and 
adult, incised, about tmce as long as broad and followed by several 
indentations ; margin of the pygidium exhibiting four rather con- 
spicuous tubular pores; between the median lobes are two short 
spines; located on the dorsal surface situated along the lateral 
margin of the median lobe there are two spines, the posterior being 
the larger, one on the outer lobule of the second lobe and two situ- 
ated between the second lobe and the penultimate segment; on 
ventral surface there is a spine corresponding with each dorsal spine, 
except on the first lobe; laterad of each lobe there is an elongate pore 
and two between the second lobule and the penultimate segment; 
anal opening twice the width of one median lobe and far removed 
from the tip; median and anterior lateral paragenitals contiguous, 
forming an arch^ 21 to 23, posterior laterals 10 to 16. 

Fiorinia fiorinise, japonica Kuw. 

Fiorinia fiorinise japonica Kuwana, Proc. Cal. Acad. Sci. (3), III, p. 79 (1902). 

Fiorinia Jiorimse j a ponica Coleman, Journ. N. Y. Ent. Soc, XI, p. 84 (1903). Rec- 
ord only. 

Fiorinia fiorinix japonica Leonard!, Redia, III, Fasc. 1, p. 36 (1905). Bibliography 
and description. 

Fiorinia fiorinix japonica Kuwana, Bui. Imp. Cent. Agr. Exp. Sta., Japan, p. 200 
(1907). Bibliogi-aphy and record only. 

Fiorinia fiorinise japonica Brick, Sta. f. Pflanzenschutz z. Hamburg, XI, p. 6 (1909). 
Record only. 

Fiorinia fioriniae japonica Brick, Jahrb. Hamb. wiss. Anst., XXVII, p. 505 (1909). 
Record only. 

Fiorinia fiorinise japonica Essig, Pom. Journ. Ent., II, 2, p. 207 (1910). Descrip- 
tion and figure. 

Fiorinia fiorinise japonica Lindinger, Zeitschr. f. wiss. Insektenbiol., VII, 4, p. 126 
(1911). Record only. 

Although this variety oi fiorinise, is a native of Japan, it has been 
introduced into this country on numerous occasions on infested ever- 



82 PAPERS ON COCCID^ OR SCALE INSECTS. 

greens. In September of 1908 it was collected on Tsuga sp., at I 
Queens, Long Island, by Mr. F. N. Meyer, where it had apparently 
become established, and subsequently it was collected in New York 
by Mr. E. P. Felt on Japanese hemlock. In June of 1909 it was col- 
lected on the fruit and leaves of Podocar'pus elata received by the 
Bureau of Plant Industry from the Botanical Gardens of New South 
Wales. Mr. Edward M. Ehrhorn, while horticultural commissioner 
of California, collected it at quarantine on Podocar'pus chinensis and 
Pinus sp. Mr. Lindinger in a recent paper ^ records it from, the Philip- 
pine Islands on Podocar'pus nageia and Ahies veitcM and Mr. Brick 
reports it from Japan on Pinus pentaphijlla, P. thunbergi, and Tsuga 
sieboldi. 

DESCRIPTION. 

Scale of female. — Fresh female scales frequently covered with a 
white powdery substance, median carina indistinct and frequently 
not visible, brown to dark brown, reaching a chestnut brown at the 
anterior end of the scale; first larval exuvia yellow, about three-fourths 
of it extending beyond the margin; second larval exuvia 1.80 to 2 
mm. in length and from 0.60 to 0.80 mm. in breadth. When collected 
onTsuga sp., it usually occurs on the underside of the leaves, although 
this does not hold true in the case of the specimens on Podocarpus. 

Scale of 'male. — White, apparently uncarinated, larval exuvia yel- 
low. Length about 1 mm. Breadth about 0.40 mm. 

Female. — Thin, pygidium slightly brownish, antennse rudimentary, 
apparently two-jointed, the second joint bearing a spine; width of 
pygidium 0.187 to 0.255 mm., slightly truncate at tip, median lobes 
serrated along free edge, oblique, forming a distinct notch, second 
lobes conspicuous in both second larval exuvia and adult, incised, 
approximately twice as long as broad, six prominent tubular pores 
on the margin of the pygidium with two very short spines between 
the median lobes; gland spines simple, long, one laterad of each lobe 
and one on the margin near the penultimate segment; on the dorsal 
surface along the lateral margin of the median lobes there are two 
spines, the posterior being the larger, one on the outer lobule of the 
second lobe, and two between second lobe and penultimate segment. 
Anal opening far removed from the tip and twice the width of a 
median lobe. Paragenitals in five groups, arranged as follows: 
Median group 3 to 11, anterior laterals 16 to 24, posterior laterals 
19 to 31. 

» Loc. cit. 

o 



Technical Series, No. 1 6, Part VI. 

U. S. DEPARTMENT OF AGRICULTURE. 

BtlRE^^XJ OW ElSTTOTdlOLOGrY. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



PAPERS ON COCCID^ OR SCALE INSECTS. 



CATALOGUE 



RECENTLY DESCRIBED COCCID^-IV. 



By E. R. SASSCER, 

Scientific Assistant. 



Issued October 19, 1912. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1912. 

V 



CONTENTS. 



Page. 

Subfamily Monophlebinae -. 83 

Subfamily Ortheziinae 84 

Subfamily Dactylopiinae 84 

Subfamily Tachardiinse 87 

Subfamily Coccinse 87 

Subfamily Diaspinge 89 



U. S. D. A., B. E. Tech. Ser. 16, Pt. VT. Issued October 19, 1912. 

PAPERS ON COCCID^ OR SCALE INSECTS. 



CATALOGUE OF RECENTLY DESCEIBED COCCID-ffil— IV. 

By E. R. Sasscer, Scientific Assistant. 

Since the publication of the "Catalogue of Recently Described 
Coccidse — III,"^ 6 new genera, 110 new species, and 4 new varieties 
have been described in various publications by workers on Coccidae. 
This makes a total of 29 new genera, 9 new subgenera, 540 new 
species, and 45 new varieties recorded since the appearance of Mrs. 
Femald's Catalogue in 1903. The writer believes this list to be fairly 
complete to March, 1912, and earnestly requests the assistance of 
coworkers in adding any references that may have escaped his notice. 

All species marked with an asterisk are represented in the National 
Collection of Coccidse. 

Subfamily MONOPHLEBIN.^. 

Monophlebus africanus Newstead. 

Monophlebus africanus Newst., Zool. Anthr. Ergeb. westl. zentr. Siidafr., v, 1, p. 15 
(1912). Fig. 
Habitat — Dutch Southwest Africa. 
On roots of I "kxuibes" — plant. 

Stigmacoccus zimmermanni Newstead. 

Perissopneumon zimmermanni Newst., Mitt. zool. Mus. Berlin, v, 2, p. 157 (1911). 
Habitat — German East Africa. 
On Manihot glazionii. 

Aspidioproctus arxaatus Newstead. 

Aspidioproctus armatus Newst., Mitt. zool. Mus. Berlin, v, 2, p. 160 (1911). Fig. 
Habitat — German East Africa. 
On Acacia. 

Aspidioproctus maximus Newstead. 

* Aspidioproctus mxiximus Newst., Mitt. zool. Mus. Berlin, v, 2, p. 158 (1911). Fig. 
Habitat — German East Africa. 
On M'sasa tree (Brachystegia randii). 

iSasscer, E. R., 'Catalogue of Recently D&scribed Coccidae— lU," Tech. Ser. 16, Part IV, Bur. Ent., 
U. S. Dept. Agr., 1911. 

.83 



84 PAPEKS ON COCCID^ OR SCALE INSECTS. 

Icerya longisetosa Newstead. 

Icerya longisetosa Newst., Mitt. zool. Mus. Berlin, v, 2, p. 155 (1911). Fig. 
Habitat — German East Africa. 
On Acacia. 

Icerya subandina Leonardi. 

Icerya subandina Leon., Bol. Zool. Sc. Sup. Portici, v, p. 238 (1911). Fig. 
Habitat — Argentina. 
On Bulvesia retana. 

Lophococcus mirabUis tricornis Newstead. 

Lophococcus mirabilis tricornis Newst., Zool. anthr. Ergeb. westl. zentr. Siidafr., 
V, 1, p. 17 (1912). Fig. 
Habitat — Dutch Southwest Africa. 
On? 

SubfamUy ORTHEZHN^. 

Orthezia varipes Leonardi. 

Orthezia varipes Leon., Bol. Zool. Sc. Sup. Portici, v, p. 240 (1911). Fig. 
Hab itat — Argentina . 
On Atriplex lampa. 

Subfamily DACTYLOPIIN^. 

Opis those eHs (?) prosopidis Kieffer and Jorgensen. 

Op-isthoscelis prosopidis Kieffer and Jorgensen, Centr. f. Bakt. Par. u. Infekt., xxvii, 
2, p. 417 (1910). Fig. 
Habitat — Argentina. 
In galls of Prosopis adesmioides. 

Asterolecaniiun coffese Newstead. 

Asterolecaniwn coffese Newst., Mitt. zool. Mus. Berlin, v, 2, p. 161 (1911). Fig. 
Habitat — German East Africa. 
On Coffea arabica. 

Lecaniodiaspis africana Newstead. 

Antonina (?) africana Newst. (siue descr.), Liverpool Univ., Quart. Jn., i, 2, pp. 69, 
72 (1906). 
Draper, Scale Insects of Egypt, p. 11 (1907). 
Lecaniodiaspis africana Newst., Bui. Ent. Res., ii, 2, p. 100 (1911). Fig. 
Habitat — Egypt. 
On Acacia arabica and Ficus sp. 

Cerococcus andinus Leonardi. 

Cerococcus andinus Leon., Bol. Zool. Sc. Sup. Portici, v, p. 245 (1911). Fig. 
Habitat — Argentina. 
On Tricycla patagonica, T. cacheuta. 

Cerococcus badius Leonardi. 

Cerococcus badius Leon., Bol. Zool. Sc. Sup. Portici, v, p. 243 (1911). Fig. 
Habitat — Argentiaa. 
On Vendtia calycina, Baccharis rosmarinifolia. 



CATALOGUE OF KECENTLY DESCRIBED COCCID^ IV. 85 

Birchippia americana Leonard!. 

Birchippia americana Leon., Bol. ZooL Sc. Sup. Portici, v, p. 246 (1911). Fig. 
Habitat — Argentina. 
On Larrea divaricata, L. cuneata. 

Kermes shastensis Ehrhom. 

Kermes shastensis Ehrh., Can. Ent., xliii, 8, p. 275 (1911). 
Habitat — California. 
On Quercus chrysolepsis. 

Eriococcus diversispinus Leonard!. 

Eriococcus diversispinus Leon., Bol. Zool. Sc. Sup. Portici, v, p. 249 (1911). Fig. 
Habitat — Argentina. 
On Zuccagnia punctata. 

Eriococcus eriogoni Ehrhorn. 

Eriococcus eriogoni Ehrh., Can. Ent., xliii, 8, p. 276 (1911). 
H abitat — Arizona . 
On Eriogonum stellatum. 

Eriococcus parcispinosus Leonard! . 

Eriococcus parcispinosus Leon., Bol. Zool. Sc. Sup. Portici, v, p. 248 (1911). Fig. 
Habitat — Argentina. 
On Atriplex lampa. 

Eriococcus saUnus Ehrhorn. 

Eriococcus salinus Ehrh., Can. Ent., xliii, 8, p. 276 (1911). 
Habitat — California. 
On roots of grass {Distichlis sp.P). 

Gynmococcus lahillei Leonard!. 

Gymnococcus lahillei Leon., Bol. Zool. Sc. Sup. Portici, v, p. 251 (1911). Fig. 
Habitat — Argentina. 
On Larrea divaricata, L. cuneata. 

Cissococcus ? oahuensis Ehrhorn. 

Cissococcus ? oahuensis Ehrh., Proc. Haw. Ent. Soc, ii, 4, pp. 149, 179 (1912). Fig. 
Habitat — Hawaiian Islands. 
On Opuhe ( Urera sandwichensis) . 

Sphsarococcus cupressi Ehrhom. 

Sphaerococcus cupressi Ehrh., Can. Ent., xliii, 8, p. 277 (1911). Fig. 
Habitat — California. 
"Under dry bark and in crevices on trunk of Cupressus macrocarpa." 

Ceroputo koebelei Ehrhorn. 

Ceroputo koebelei :^rh., Can. Ent., xliii, 8, p. 278 (1911). Fig. 
Habitat — Arizona. 
On Quercus engelmanni. 



86 PAPERS ON COCCID^ OK SCALE INSECTS. 

Genus LACHNODIELLA von Ihering. Type, cecropiss. 

Lachnodiella von Iher. (sine descr.), Bot. Jahrb., xxxix, 5, p. 680 (1907). 
Sanders, Catalogue Coccidje — II, p. 40 (1909). 
Hempel, Rev. Mus. Paul., viii, p. 52 (1911). Descr. 

Lachnodiella cecropise von Ihering. 

Lachnodiella cecropias von Iher. (sine descr.), Bot. Jahrb., xxxix, 5, p. 680 (1907). 

Sanders, Catalogue Coccidse — II, p. 40 (1909). 

Hempel, Rev. Mus. Paul., viii, p. 52 (1911). Descr. 
Habitat — Brazil. 
On Cecropia adenopus. 

Trionymus califomicus Ehrhorn. 

Trionymus califomicus Ehrh., Can. Ent., xliii, 8, p. 279 (1911). Fig. 
Habitat — California. 
On Festuca sp. 

Pseudococcus aridoruia Lindinger. 

Pseudococais aridorum Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 7 (1911). Fig. 
Habitat — Canary Islands. 

On Argyranthevium frutescens, Cytisus prolifer var. palmensis, grass, Tri/olium 
panorinitanum. 

Pseudococcus crotonis Green. 

Dactylopius crotonis Green (sine descr.), Trop. Agr., xxiv, p. 44 (1905). 
Pseudococcus crotonis Sanders, Catalogue Coccidse — I, p. 5 (1908). 
Dactylopius crotonis Green, Jn. Econ. Biol., vi, 2, p. 35 (1911). Fig. 

Habitat — Ceylon. 

On Castilloa elastica, variegated croton (Codissum variegatum), Temtinalia catappa, 
Erythrina lithosperma. 

Pseudococcus ledi Cockerell. 

^Pseudococcus ledi Ckll., Ent. News, xxii, 5, p. 217 (1911). 
Habitat— New York. 
On Ledum groenlandicum. 

Pseudococcus mendozinus Leonardi. 

Pseudococcus mendozinus Leon., Bol. Zool. Sc. Sup. Portici, v, p. 255 (1911). Fig. 
Habitat — Argentina. 
On Hyalis argentea. 

Pseudococcus obtusus Newstead. 

Dactylopius (Pseudococcus) ohtusv^^ Newat., Mitt. zool. Mua. Berlin, v, 2, p. 164 (1911). 
Fig. 
Habitat — German East Africa. 
On Baohdbrinde. 

Pseudococcus percerosus Leonardi. 

Pseudococcus percerosus Leon., Bol. Zool. Sc. Sup. Portici, v, p. 254 (1911). Fig. 
Habitat — Argentina. 
On Gourliea decorticans. 

Genus HELIOCOCCUS Sulc. Tjrpe, bohemicua. 
Heliococcus Sulc, Cas. Spolec. Ent., ix, 1, p. 39 (1912). 



CATALOGUE OF KECBNTLY DESCRIBED COCCID^ IV. 87 

HeKococcus bohemicus Sulc. 

Heliococcus bohemicus Sulc, Cas. Spolec. Ent., ix, 1, p. 39 (1912). Fig. 
Habitat — Bohemia and Moravia. 
On Rohinia pseudacacia. 

Ripersia grandulifera Newstead. 

Ri-persia grandulifera Newst., Zool. anthr. Ergeb. westl. zentr. Siidafr., v, 1, p. 17 
(1912). Fig. 
Habitat — Southwest Africa. 
On Adiantum sp. 

Subfamily TAOHARDnH^. 

Tachardia albizziee Green. 

Tachardia alhizzise Green (sine descr.), Ind. Mus. Notes, v, p. 98 (1903). 

Tachardia albizzise Sanders, Catalogue Coccidae — 1, p. 6 (1906). 

Tachardia alhizzix Green, Jn. Econ. Biol., vi, 2, p. 32 (1911). Fig. Description. 

Habitat — Ceylon. 

On Landolphia sp., Albizzia stipulata, Filicium decipiens, Harpullia cupanioides, 
Croton lacciferum, Theobroma cacao, Sleichera trijuga. 

Tachardia cordaliee Leonardi. 

Tachardia cordalix Leon., Bol. Zool. Sc. Sup. Portici, v, p. 258 (1911). Fig. 
Habitat — ^Argentina . 
On Cordalia lineata. 

Tachardia longisetosa Newstead. 

Tachardia longisetosa Newst., Bui. Ent. Res., ii, 2, p. 102 (1911). ^ 

Habitat — Uganda. 
On bark-cloth (Fiscus sp.). 

Tachardia lycii Leonardi. 

Tachardia lycii Leon., Bol. Zool. Sc. Sup. Portici, v, p. 256 (1911). Fig. 
Habitat — Argentina. 
On Lycium chilense. 

Subfamily COCCINiE. 

Pulvinaria argentina Leonardi. 

Pulvinaria argentina Leon., Bol. Zool. Sc. Sup. Portici, v, p. 260 (1911). Fig. 
Habitat — Argentina. 
On Lycium chilense, Fabiana denudata. 

Pulvinaria plana Lindinger. 

Pulvinaria plana Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 34 (1911). Fig. 
Habitat — Canary Islands. 
On underside of leaves of Laurus canariensis. 

Luzulaspis spinulosa Leonardi. 

Luzulaspis spinulosa Leon., Bol. Zool. Sc. Sup. Portici, v, p. 262 (1911). Fig. 
Habitat — Argentina. 
On Atriplex lampa. 
58242°— 12 2 



88 PAPEEs ojst coccid^ oe scale insects. 

Ceroplastes breviseta Leonard!. 

Ceroplastes breviseta Leon., BoL ZooL Sc. Sup. Portici, v, p. 264 (1911). Fig. 
Hab itat — Argentina . 
On Atriplex lampa. 

Ceroplastes galeatus Newstead. 

Ceroplastes galeatus Newst., BuL Ent. Res., ii, 2, p. 95 (1911). Fig. 
Habitat — Uganda. 
On? 

Ceroplastes irregularis Leonardi. 

Ceroplastes irregularis lieon., Bol. Zool. Sc. Sup. Portici, v, p. 271 (1911). Fig. 
Hab itat — Argentina . 
On Larrea cuneata. 

Ceroplastes longiseta Leonardi. 

Ceroplastes longiseta Leon., Bol. Zool. Sc. Sup. Portici, v, p. 268 (1911). Fig. 
Habitat — Argentina. 
On Fabiana denudata. 

Ceroplastes subrotundus Leonardi. 

Ceroplastes subrotundiis Leon., Bol. Zool. Sc. Sup. Portici, v, p. 266 (1911). Fig. 
Habitat — Argentina. 
On Cercidium andicolum. 

Ceroplastes subsphaericus Newstead. 

Ceroplastes subsphsericus Newst., Mitt. zool. Mus. Berlin, v, 2, p. 166 (1911). 
Habitat — German East Africa. 
On Albizzia lebbek. 

Ceroplastes ugandse Newstead. . 

Ceroplastes ugandse Newst., Bui. Ent. Res., ii, 2, p. 94 (1911). Fig. 
Habitat — Uganda . 
On "Amakebe." 

Ceroplastes vinsonioides Newstead. 

Ceropla.stes vinsonioides Newst., Bui. Ent. Res., ii, 2, p. 96 (1911). Fig. 

Habitat — Uganda . 

On Coffea. 

Ceroplastes go-wdeyi Newstead. 

Ceroplastes gowdeyi Newst., Bui. Ent. Res., ii, 2, p. 98 (1911). Fig. 
Habitat — Uganda. 
On African bark-tree {Ficus sp.). 

Inglisia Castillo ee Green. 

Inglisia castillox Green, Jn. Econ. Biol., vi, 2, p. 29 (1911). Fig. 
Habitat — Ceylon. 

On Castilloa elastica, Grewia microcos, Adenochlsena zeylanica, Solanum sp., Vernonia 
sp., and tea. 

Coccus nyasss Newstead . 

Lecanium nyasx Newst., Mitt. zool. Mus. Berlin, v, 2, p. 162 (1911). Fig. 
Habitat — German East Africa. 
On? 



CATALOGUE OP RECENTLY DESCEIBED COCCID^ IV. 89 

Lecanivuu elegans Leonard!. 

Eulecanium elegans Leon., BoL zooL Sc. Sup. Portici, v, p. 273 (1911). Fig. 
Habitat — Argentina . 
On Larrea cuneata, L. divaricata. 

Lecanium tennivalvatura Newstead. 

Lecanium tenuivalvatum Newst., BuL Ent. Res., ii, 2, p. 92 (1911). Fig. 
Habitat — Uganda. 
On citronella grass. 

Lecaniuxa tremae Newstead. 

Lecanium {Eulecanium) tremx Newst., Mitt. zool. Mus. Berlin, v, 2, p. 162 (1911). 

Fig. 
Lecanium (Eulecanium) tremae Newst., BuL Ent. Res., ii, 2, p. 93 (1911). 

Habitat — German East Africa. 

On Trema guineensis. 

Saissetia silvestrii Leonardi. 

Saissetia silvestrii Leon., BoL ZooL Sc. Sup. Portici, v, p. 275 (1911). Fig. 
Habitat — Argentina . 
Oa Zuccagnia punctata. 

Subfamily DIASPINiE. 

Chionaspis africana Newstead. 

Chionaspis africana Newst., ZooL anthr. Ergeb. westL zentr. Sudafr., v, 1, p. 19 
(1912). Fig. 
Habitat — Southwest Africa. 
On? 

Chionaspis amaniensis Lindinger. 

Chionaspis amaniensis Lindgr., Jakrb. Hamb. wiss. Anst., xxvii, 3, p. 42 (1910). Fig. 
Habitat — German East Africa. 
On imderside of an undertermined dicotyledonous leaf. 

9 

Chionaspis arthrocnemi Lindinger. 

Chionaspis arthrocnemi Lindgr., Zeitschr. f. wiss. InsektenbioL, vii, 11, p. 354 (1911). 
Fig. 
Habitat — Turkey . 
On Arthrocnemum macrostachyum. 

Chionaspis canariensis Lindinger. 

Chionaspis canariensis Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 26 (1911). Fig. 
Habitat — Canary Islands. 

On Plocama pendula, Micromeria ericifolia, M. linJci, M. teneriffx, M. terehinthacea, 
Micromeria sp., Ruta oreojasme, Salsola longifolia, Cneorum pulverulentum, 
Cytisus filiper . 

Chionaspis cassise Newstead. 

Chionaspis cassix Newst., Bui. Ent. Res., ii, 2, p. 89 (1911). Fig, 
Habitat — Uganda . 
On Cassia florihunda. 

Chionaspis longispina Newstead. 

Chionaspis longispina Newst., Bui. Ent. Res., ii, 2, p. 88 (1911), Fig, 
Habitat — Egypt. 
On Justida alba. 



90 ' PAPERS ON COCCID^ OE SCALE INSECTS. 

Chionaspis mytilaspiformis Newstead. 

Chionaspis mytilaspiformis Newst., Zool. anthr. Ergeb. west!, zentr. Siidafr., v, 1, p. 19 I 
(1912). Fig. 
Habitat — Southwest Africa. 
On Rhus lancea. 

Chionaspis nudata Newstead. 

Chionaspis nudata Newst., Mitt. zool. Mus. Berlin, v, 2, p. 170 (1911). Fig. 
Habitat— German East Africa. 
On(?) 

Chionaspis subnudata Newstead. 

Chionaspis subnudata Newst., Zool. anthr. Ergeb. westl. zentr. Siidafr., v, 1, p. 19 
(1912). Fig. 
Habitat — Southwest Africa. 
On "! obarus." 

Diaspis atlantica Lindinger. 

Diaspis atlantica Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 28 (1911). Fig. 
Habitat — Canary Islands. 
On leaf of Juniperus phoenicea. 

Diaspis 'barrancorixni Lindinger. 

Diaspis barrancorum Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, p. 29 (1911). Fig. 
Habitat — Canary Islands. 
On Euphorbia regis-jubse. 

Diaspis parva Lindinger. 

Diaspis parva Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 44 (1910). Fig. 
Habitat — ^German East Africa. 
On leaf and twig of Loranthus undulatus var. sagittif alius. 

Diaspis regularis Newstead. 

Diaspis regularis Newst., Bui. Ent. Res., li, 2, p. 86 (1911). Fig. 
Habitat — Uganda. 
On (?) 

Aulacaspis cinnamomi mangiferae Newstead. 

Diaspis (Aulacaspis) cinnamomi v. mangiferx Newst., Bui. Ent. Res., ii, 2, p. 86 
(1911). Fig. 
Habitat — Egypt (imported from Ceylon). 
On mango. 

Phenacaspis bussii Newstead. 

Chionaspis bussii Newst., Mitt. zool. Mus. Berlin, v, 2, p. 171 (1911). 
Habitat— German East Africa. 
On Macrolobium sp. 

Phenacaspis lutea Newstead. 

Chionaspis lutea Newst., Mitt. zool. Mus. Berlin, v, 2, p. 169 (1911). Fig. 
Habitat — German East Africa. 
On forest tree, under fungus. 

Phenacaspis tangana Lindinger. 

Phenaaispis tangana Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 8, p. 45 (1910). Fig. 
Habitat — German East Africa. 
On leaf of Dracxna sp. 



CATALOGUE OF EECENTLY DESCRIBED COCCID^ IV. 91 

Hemichionaspis chionaspiformis Newstead. 

Diaspis (?) chionaspiformis Newst., Bui. Ent. Res., i, 3, p. 198 (1910). Fig. 
Chionaspis unita Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 43 (1910). Fig. 
Hemichionaspis cMonaspitiformis Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, Bericfeti- 

gung (1910). 
Diaspis (?) chionaspiformis Sasscer, Catalogue Coccidse — III, p. 68 (1911). 

Habitat — British and German East Africa. 

On an unknown plant and Turrasa sp. 

Crypthemichionaspis Lindinger. Type, nigra. 

Crypthemichionaspis Lindgr.,' Zeitschr. f. wiss. Insektenbiol., vii, 5, 6, p. 175 (1911). 

Crypthemichionaspis nigra Lindinger. 

Crypthemichionaspis nigra Lindgr., Zeitschr. f. wiss. Insektenbiol., ^di, 5, 6, p. 175 
(1911). Fig. 
Habitat — South Australia. 
On Acacia salicina; Phyllodien. • 

Leucaspis iadias-orientalis Lindinger . 

Leucaspis indix-orientalis Lindgr., Zeitschr. f . wiss. Insektenbiol., vii, 4, p. 127 (1911). 
Fig. 
Habitat — India. 
On Pinus kasya. 

Fiorinia africana Newstead. 

Fiorinia africana Newst., Bui. Ent. Res., ii, 2, p. 90 (1911). Fig. 
Habitat — Egypt. 
On poplar. 

Fiorinia neo-caledonica Lindinger. 

Fiorinia neo-caledonica Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 5-6, p. 176 (1911). 
Fig. 
Habitat — New Caledonia. 
On Bxckia pinifolia. 

Fiorinia odinae ratiltipora Lindinger. 

Fiorinia odinx muUipora Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 126 (1911). 
Habitat — India. 
On Taxus wallichiana. 

Genus CRYPTASPIDUS Lindinger. Type, nucum. 

(Mfptaspidus Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 43 (1910). 

Cryptaspidus nucum Lindinger. 

Cryptaspidus nucum Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 43 (1910). Fig. 
Hab itat — Madagascar . 
On scales of fruit of Cocos nucifera. 

Aorddia glandulosa Newstead. 

Aonidia glandulosa Newst., Draper (sine descr.). Scale Ins. of Egypt, p. 11 (1907). 

Sanders, Catalogue Coccidse — II, p. 56 (1911). 

Newst., Bui. Ent. Res., ii, 2, p. 103 (1911). Fig. 
Hab itat — Egypt . 
On "sunt" tree {Acacia arabica). 

> Includes Fiorinia acacix Mask., F. lidgetti Green, and Crypthemichionaspis nigra u. sp. 



92 PAPEES ON COCCID^ OE SCALE INSECTS. 

Aonidia longa Lindinger. 

Aonidia longa Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 5, 6, p. 172 (1911). Fig. 
Habitat — New Caledonia. 
s. On Podoearpus grddioides. 

Aonidia (?) paradoza Lindinger. 

Aonidia (?) paradoxa Lindgr., Zeitschr. f. wisa. Insektenbiol., vii, 5, 6, p. 173 (1911). 

Fig. 
Habitat — South Australia. 
On Casuarina glauca. 

Aonidia (?) targioniopsis Lindinger. 

Aonidia (?) targioniopsis Lindgr., Zeitschr. f. wiss. Insektenbiol,, vii, 3, p. 86 (1911). 
Fig. 
Habitat — Burma. 
On Miliusa velutina. 

Aonidia viridis Lindinger. 

Aonidia viridis Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 3, p. 86 (1911). Fig. 
Habitat — India. 
On upper side of leaf of Aglaia minutiflora. 

Cryptaspidiotus aonidioides Lindinger. 

CryptaspidioVus aonidioides Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 21 (1911)» 
Fig. 
Habitat — Canary Islands. 
On leaves of Laurus canariensis, Apollonias canariensis. 

Cryptaspidiotus austro-africanus Lindinger. 

Cryptaspidiotus austro-africanus Lindgr. (Brick), Jahrb. Hamb. wiss. Anst., xxvii, 

p. 505 (1910). 
Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 41 
(1911). Fig. and description. 
Habitat — Natal. 
On Euphorbia sp. 

Aspidiotus argentina Leonard!. 

Aspidiotiis (Eemiberlesia) argentina Leon., Bol. Zool. Sc. Sup. Portici, v, p. 277 (1911). 

Fig- 
Habitat — Argentina. 
On Ophryoporus andinv^. 

Aspidiotus bavaricus Lindinger. 

Aspidiotus (Diaspidiotus) bavaricus Lindgr., Zeitschr. f. wiss. Insektenbiol., viii, 1, 
p. 31 (1912). 
Habitat — Bavaria, Prussia, Austria, Norway, England (?), and Portugal (?). 
On Calluna vulgaris, Erica tetralix. 

Aspidiotus canariensis Lindinger. 

Aspidiotus canariensis Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 12 (191]). Fig. 
Habitat — Canary Islands. 
On Btem and branch of Argyranthemum frutescens. 



CATAJLOGUE OF EECENTLY DESCRIBED COCCID^ IV. 93 

Aspidiotus euphorbise Newstead. 

Aspidiotus (Selenaspidus) euphorbise Newst., Zool. anthr. Ergeb. westl. zentr. Siidafr., 
V, 1, p. 18 (1912). Fig. 
Habitat — South Africa. 
On Euphorbia virosa. 

Aspidiotus fissidens pluridentatus Lindinger. 

Aspidiotus fissidens pluridentatus Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 35 
(1910). 
Habitat — German East Africa, Mozambique. 
On leaves of Bosquiea cerasiflora, Sideroxylon inerme, Chrysophyllum stuhlmanii. 

Aspidiotus flssus Lindinger. 

Aspidiotus fissus Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 35 (1910). Fig. 
Habitat — Abyssinia. 
On Euphorbia sp. 

Aspidiotus furcrseicola Lindinger. 

Aspidiotus furcrseicola Lindgr., Jabrb. Hamb. wiss. Anst., xxvii, 3, p. 36 (1910). Fig. 
Habitat — German East Africa. 
On underside of leaf of Furcrxa gigantia. 

Aspidiotus gymnosporise Lindinger. 

Aspidiotus gymnosporix Lindgr., Jabrb. Hamb. wiss. Anst., xxviii, 3, p. 13 (1911) 
Fig. 
Habitat — Canary Islands. 
On leaf of Gymnosporia cassinoides. 

Aspidiotus lauretorura Lindinger. 

Aspidiotus lauretorum Lindgr., Jabrb. Hamb. wiss. Anst., xxviii, 3, p. 15 (1911). 

Fig- 
Habitat — Canary Islands. 

On Dracsena draco, Gymnosporia cassinoides, Ilex canariensis, I. plafyphylla, Oreo- 
daphne fcet'ens, Picconia excelsa, Smilax canariensis, Hedera helix canariensis, 
Apollonias canariensis, Heberdenia excelsa, Laurus canariensis, and Lauracese. 

Aspidiotus maramillaris Lindinger. 

Aspidiotus mammillaris Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 37 (1910). 
Fig. 
Habitat — Abyssinia. 
On surface of leaves of Aloe eru. 

Aspidiotus reticulatus Newstead, 

Aspidiotus reticulatus Newst., Zool. anthr. Ergeb. westl. zentr, Siidafr., v, 1, p. 17 
(1912). Fig. 
Habitat — Southwest Africa. 
On monocotyledon (?) 

Chrysomphalus samoana Lindinger, 

Chrysomphalus (Melanaspis) samoana Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 5, 6, 
p. 177 (1911). Fig. 
Habitat — Island of Samoa. 
On Myristica hypargyrxa. 



94 PAPERS ON COCCID^ OR SCALE INSECTS. 

Aspidiotus schultzei Newstead. 

Aspidiotus (Selenaspidus) schuUzei Newst., Zool. anthr. Ergeb. westl. zentr. Sudafr., 
V, 1, p. 18 (1912). Fig. 
Habitat — Southwest Africa. 
On succulent plant. 

Aspidiotus taorensis Lindinger. 

Aspidiotus taorensis Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 17 (1911). Fig. 
Habitat — Canary Islands. 
On Euphorbia aphylla, E. regis-jubx. 

Aspidiotus tectarius Lindinger. 

Aspidiotus tectarius Lindgr., Jahrb. Hamb. wiss. Anst., xxvi, 3, p. 20 (1909). Fig. 
Habitat — South America. 
On leaves of Euphorbia sp. 

Aspidiotus tinerfensis Lindinger. 

Aspidiotus tinerfensis Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 18 (1911). Fig. 
Habitat — Canary Islands. 
On Dracxna draco. 

Aspidiotus tsugae Marlatt. 

Aspidiotus (Diaspidiotus) tsugx Marlatt, Ent. News, xxii, 9, p. 385 (1911). Fig. 
Habitat — Japan . 
On Japanese hemlock (Tsuga sp.). 

Aspidiotus varians Lindinger. 

Aspidiotus varians Lindgr., Jahrb. Hamb. wiss. Anst., xxvii, 3, p. 39 (1911). Fig. 
Habitat — German East Africa, Madagascar. 
On leaf and scales of fruit of Cocos nuci/era. 

Targionia (?) carapylanthi Lindinger. 

Targionia (?) campylanthi Lindgr., Jahrb. Hamb. wiss. Anst., xxviii, 3, p. 25 (1911). 
Fig. 

Habitat — Canary Islands. 
On Campylanthus solsoloides. 

Targionia fabianse Leonardi. 

Targionia fabianx Leon., Bol. Zool. Sc. Sup. Portici, v, p. 278 (1911). Fig. 
Hab itat — Argentina . 
On Fabiana denudata. 

Genus PHOTARGIONIA Leonardi. Type, larrex, 

Protargionia Leon., Bol. Zool. Sc. Sup. Portici, v, p. 280 (1911). 

Protargionia larrese Leonardi. 

Protargionia larrex Leon., Bol. Zool. Sc. Sup. Portici, v, p. 280 (1911). Fig. 
Habitat — Argentina. 
On Larrea divaricata, L. cuneata. 

Lepidosaph.es fasciata Green. 

Lepidosaphes { My tilaspis) fasciata Green, Jn. Econ. Biol., vi, 2, p. 31 (1911). Fig. 
Habitat — Ceylon. 
On leaves of Hevea brasiliensis. 



CATALOGUE OP RECENTLY DESCEIBED COCCID^ IV. 95 

Lepidosaphes travancorensia Lindinger. 

Lepidosaphes travancorensia Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 127 
(1911). _ Fig. 
Habitat — India. 
On underside of leaf of Aglaia minutiflora. 

Genus DINASPIS Leonardi. Type, ichesiii 

Dinaspis Leon., Bol. Zool. Sc. Sup. Portici, v. p. 282 (1911). 

Dinaspis ichesii Leonardi. 

Dinaspis ichesii Leon., Bol. Zool. Sc. Sup. Portici, v, p. 282 (1911) * Fig. 
Habitat — Argentina. 
On Ephedra andina. 

Dinaspis lahillei Leonardi. 

Dinaspis lahillei Leon., Bol. Zool. Sc. Sup. Portici, v, p. 283 (1911). Fig. 
Habitat — Argentina. 
On Bulvesia retana. 

Ischnaspis spathulata Lindinger. 

Ischnaspis spathulata Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 127 (1911). 

Fig. 
Habitat — India. 
On underside of leaf of Atica obscura. 

Parlatoria ephedras Lindinger. 

Parlatoria ephedrse Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 129. Fig. 
Hab itat — Persia . 
On Ephedra intermedia, E. nehiodensis, E. nebiodensis var. procera, 

Parlatoria hastata Lindinger. 

Parlatoria hastata Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 129 (1911). Fig. 
Habitat — Borneo. 
On leaf of Gnetum scandens. 

Cryptoparlatorea uberifera Lindinger. 

Cryptoparlatorea uberifera Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 4, p. 126 (1911). 
Fig. 
Habitat — Philippine Islands. 
On leaves of Artocarpus sp. and Mallotus philippinensis . 

Cryptoparlatorea parlatoreiodes Lindinger, 

Cryptoparlatorea parlatoreiodes Lindgr., Zeitschr. f. wiss. Insektenbiol., vii, 3, p. 89 
(1911). Fig. 
Habitat — India. 
On surface of leaf of Xanthophyllum fiavescens. 



CORRECTION TO CATALOGUE HI— 1911, 

On page 66 the heading "Subfamily Coccinse" should be transferred to page 65 to 
precede Ceroplastidia, line 17 from bottom. 



96 PAPERS ON COCCID^ OE SCALE INSECTS 

ABBREVIATIONS OF LITERATURE.* 

Allg. Zeits. f. Ent. — Allgemeine Zeitschrift fiir Entomologie. 

Agr. News West Ind. — Agricultural News. West Indies-. 

An. Soc. Cient. Argentina. — Anales de la Sociedad Cientifica Argentina. 

Ann. Soc. Ent. France. — Annales de la Societe Entomologique de France. 

Annali di Agr. — Annali di Agricoltura. Portici. 

Ann. Mag. N. H.— Annals and Magazine of Natural History. 

Ann. Mus. Zool. Ac. Imp. Sc. St. Petersbourg. — Annuaire Musee Zoologique Acad^mie 

Imperiale des Sciences de St. Petersbourg. 
Berl. ent. Zeitschr. — Berliner entomologische Zeitschrift. 

Ber. Stat. f. Pflanzenscb. Hamb. — Bericht der Station fiir Pflanzenschutz, Hamburg. 
Boi. Min. Ag. — Boletin del Ministerio de Agricultura. Argentina. 
Bol. Zool. Sc. Sup. Portici. — Bollettino del Laboratorio di Zoologia generale e agraria 

della R. Scuola Superiore d'Agricoltm-a di Portici. 
Bot. Jahrb. — Botanische Jahrbiicher. 
Bui. Ag. Exp. Sta. Japan. — Bulletin of the Imperial Central Agricultural Experiment 

Station. Japan. 
Bui. Ent. Res. — Bulletin of Entomological Research. 

Bui. 111. St. Lab. N. H. — Bulletin of the Illinois State Laboratory of Natural History. 
Bui. Soc. Ent. Fr. — Bulletin de la Societe Entomologique de France. 
Bui. Soc. Ent. Ital. — Bulletino della Societa Entomologica Italiana. 
Bui. Soc. Zoo. France. — Bulletin de la Societe Zoologique de France. 
Bui. U. S. Bur. Ent., t. s. 16, Pt. II.— Bulletin, Technical Series 16, Part II, Bureau 

of Entomology, U. S. Department of Agriculture. 
Can. Ent. — Canadian Entomologist. 
Cas. Spolec. Entom. — Casopis Ceske Spolecnosti Entomologicke. Acta Societatia 

Entomologicse Bohemiae. 
Catalogue Coccidae — I. — U. S. Dept. Agr., Bur. Ent., technical series, Bui. 12, Pt. I 

(1906). 
Catalogue Coccidse — II. — U. S. Dept. Agr., Bur. Ent., technical series, Bui. 16, Pt. Ill 

(1909). 
Catalogue Coccidse — III. — U. S. Dept. Agr., Bur. Ent., technical series, Bui. 16, 

Pt. IV (1911). 
Centr. f. Bakt. Par. u. Infekt. — Centralblatt fiir Bakteriologie imd Paraaitenkunde 

und Infektionskrankheiten. • • 
Cherm. Ital. — Chermotheca Italica. 
Cocc. Ceylon. — Coccidse of Ceylon. 
Compt. Rend. Ac. Sci., Paris. — Comptes rendus liebdomadaires des stances de I'Aca- 

demie des sciences'. Paris. 
Compt. Rend. Soc. Biol. — Comptes rendus des seances de la Societe deBiologie. Paris. 
Deutsche ent. Zeitschr. — Deutsche entomologische Zeitschrift. 
Ent. Mo. Mag. — The Entomologist's Monthly Magazine. 
Ent. Blatter. — Entomologische Blatter, Internationale Monatsschrift fiir die Biologie 

der Kafer Europas. Schwabach. 
Ent. News. — Entomological News. 

Ent. Rec. Jn. Variation. — The Entomologist's Record and Journal of Variation. 
Ent. Wochenblatt. — Entomologisches Wochenblatt {antea Insekten-Borse). 
Ind. Mus. Notes. — Indian Museum Notes. 
Insekten Borse. — Entomologisches Wochenblatt. 

Jahrb . Hamb . wiss . Anst . — Jahrb uch der Hamburgischen wissenschaftlichen Anstalten. 
Jn. Bomb. N. H. Soc. — Journal of the Bombay Natural History Society. Calcutta. 

1 This list includes abbreviations of literature used for coecld descriptions referred to in Nos. I, II, III. 
and IV. 



CATALOGUE OF EECENTLY DESCRIBED COCCID^ IV. 97 

Jn. Econ. Biol. — Tlie Journal of Economic Biology. • 

Jn. N. Y. Ent. Soc. — Journal of the New York Entomological Society. 
Kilimandjaro-Meru Exp. — Schwedischen zoologisclien Expedition nach dem Kili- 

mandjaro, dem Meru und den umgebenden Massaisteppen Deutach-Ostafrikaa. 
Liverpool Univ. Quart. Jn. — Institute of Commercial Research in the Tropics, Liver- 
pool University, Quarterly Journal. 
Medd. Soc. Faun. Fenn. — Meddelanden af Societas pro Fauna et Flora Fennica. 
Mem. Dep. Ag. India. — Memoirs of the Department of Agriculture in India. 
Mem. Soc. Cient. Ant. Alz. — Memorias y Re vista de la Sociedad Cientifica "Antonio 

Alzate." 
M^m. Soc. Zool. France. — Memoires de la Society Zoologique de France. 
Mitt. zool. Mus. Berlin. — Mitteilungen aus den zoologischen Museum in Berlin. 
Nachrichtsbl. Naturfrsch.-Klubs, Prossnitz. — Nachrichtsblatt Naturforscher-Klubs, 

Prossnitz (Mahren). 
N. Z. Trans.— Transactions of the New Zealand Institute. 
Pests and Blights of Tea Plants.— The Pests and Blights of the Tea Plant, 2d ed . Watt 

& Mann, Calcutta, 1903. 
Philippine Jn. Sci. — The Philippine Journal of Science. 
Pomona College Jn. Ent. — Pomona College Journal of Entomology. Claremont, 

California. 
Pr. Biol. Soc. Wash. — Proceedings of the Biological Society of Washington. 
Pr. Dav. Acad. Sci. — Proceedings Davenport Academy of Sciences. Davenport, 

Iowa. 
Pr. Ent. Soc. Wash. — Proceedings of the Entomological Society of Washington. 
Pr. Haw. Ent. Soc. — Proceedings of the Hawaiian Entomological Society. 
Redia. — Redia. 

Rev. Mus. Paul. — Revista Museu Paulista. 
Riv. Pat. Veg. — Ri vista di Patologia Vegetale. 
Scale Ins. of Egypt. — Notes on the Injurious Scale Insects and Mealy Bugs of Egypt. 

Cairo, 1907. 
The Entom. — The Entomologist." 
Trab. Mus. Farm. Fac. Cienc. Med. — Trabajos del Museo de Farmacologia de la Facul- 

tad de Ciencias Medicas. Buenos Aires, Argentina. 
Tr. Linn. Soc. Lond. — Transactions of the Linnaean Society of London, 
Trop. Agr. — Tropical Agriculturist. 
Victorian Nat. — The Victorian Naturalist. 

Zeitschr. f. wiss. Insektenbiol. — Zeitschrift fiir wissenchaftliche Insektenbiologie. 
Zool. anthr. Ergeb. westl. zentr. Siidafr. — Zoologische und anthropologiache Ergeb- 

nieee einer Forschungsreise im westlichen und zentralen Siidafrika. 



ADDITIONAL COPIES of this publication 
Xi- may be procured from the Supeeintend- 
ENT OF Documents, Government Printing 
Office, Washington, D. C, at 5 cents per copy 



\ 



Technical Series, No. 16, Part VII. 

U. S. DEPARTMENT OF AGRICULTURE, 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



PAPERS ON COCCIDJ: OR SCALE INSECTS. 



AN INDEX TO CATALOGUES OF RECENTLY 
DESCRIBED COCCID^ 

INCLUDED IN TECHNICAL SERIES Nos. 12 AND 16. 



E. R. SASSCER, 

Scientific Assistant. 



Issued January 23, 1913. 




WASHINGTOK: 

GOVERNMENT PRINTING OFFICE. 

1&13. 



1 ' ^V \^W\i 



BUREAU OF ENTOMOLOGY. 

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

C. L. Marl ATT, Entomologist and Acting Chief in Absence of Chief. 

R. S. Clifton, Executive Assistant. 

W. F. Tastet, Chief Clerk. 

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

A. D. Hopkins, in charge afforest insect investigations. 

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

F. M. Webster, in charge of cereal and forage insect investigations. 

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

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

D. M. Rogers, in charge of preventing spread of moths, field work. 

RoLLA P. Currie, in charge of editorial work. 

Mabel Colcord, in charge of library. 



U. S. D. A., B. E. Tech. Ser. 16, Pt. VII. Issued January 23, 1913. 

PAPERS ON COCCIDJ; OR SCALE INSECTS. 



AN INDEX TO CATALOGUES OF RECENTLY DESCRIBED 
COCCID^ INCLUDED IN TECHNICAL SERIES NOS. 12 
AND 16. 

By E. R. Sasscer, 
Scientific Assistant. 

This index of Coccidse includes all those forms contained in the 
following papers: Technical Series No. 12, Part I, ''Catalogue of 
Recently Described Goccidse" (1906) and Technical Series No. 16, Part 
III, ''Catalogue of Recently Described Coccida?— II" (1909), by 
James G. Sanders; Technical Series No. 16, Part IV, "Catalogue of 
Recently Described Coccidse — III " (1911) and Technical Series No. 16, 
Part VI, "Catalogue of Recently Described Coccidse— IV" (1912), by 
E. R. Sasscer. It is thus mtended that it shall be an index to all 
the forms described and published since the issuance of Mrs. Fernald's 
■'Catalogue of Coccidae" m 1903, makmg it unnecessary to search 
through each of the four separate papers to find the desired citation. 

INDEX. Technical 

Series No. Page. 

aberemose ' 16 70 

aberrans 16 71 

acacise 16 91 

Aclerda 16 47, 67 

Actenaspis .....' 16 50 

Adiscofiorinia 16 50 

africana, Chionaspis 16 89 

africana, Diaspis 16 68 

africana, Fiorinia 16 91 

africana, Antonina (?) 16 43 

africana, Lecaniodiaspis 16 84 

africana, Walkeriana 16 61 

africanus, Aspidiotus 16 51 

africanus, Monophlebus 16 83 

africanus senegalensis 16 44 

a-^jUsawarensis 16 65 

a~rifoli« 16 40 

Akermes 16 46 

alba 12 16 

albizzige 12 6 

16 87 

albospicatus 16 63 

alluaudi 12 4 

99 



100 



PAPEES OlSr COCCIDiE OK SCALE INSECTS. 



alni 

anianiensis 

ambigua 

Amelococcus 

americana 

Anamaspis 

Anamefiorinia 

ananassse 

andersoni 

andinus 

angusta 

angustata 

anomala 

antigoni 

Antonina 

Aonidia 

Aonidiella 

aonidioidcs 

Aonidomytilus 

arbutus s. sp 

Arctorthezia 

arctostaphyli 

argentina, Tachardia . 
argentina, Pulvinaria. 
argentina, Aspidiotus . 

argentinus 

aridorum 

arii 

armatus 

arthrocnemi 

arundinaricie ..... 

Aspidiotus 

aspidistrse gossypii 

Aspidioproctus 

assimiiis 

Asterolecanium 

atalantise, Parlatoria. 
atalantiae, Fiorinia... 

atlantica 

Atriplicia 

Aulacaspis 

aurantiaca 

aurantii 

auriculata 

australis 

australis s. sp 



Technical 
Series No. 

16 


Page. 
55 


16 


89 


16 


64 


12 


4 


16 


85 


16 


50 


16 


50 


16 


64 


12 


5 


16 


84 


16 


58 


12 


10 


16 


58 


16 


65 


16 


43 


12 


5 


16 

12 


43,84 
16 


16 


56 


12 


70,91 
15 


16 


51, 55 


16 


92 


12 


16 


16 


71 


12 


2 


16 


69 


16 


43 


16 


87 


16 


92 


16 


88 


16 


86 


16 


71 


16 


83 


16 


89 


12 


8 


12 


13 


16 


51, 58, 69, 92 


16 


49 


16 


83 


12 


2 


12 


3 


16 
12 


35, 62, 84 
18 


16 


50 


16 


90 


16 


37 


16 
12 


49,90 
6 


16 


55 


16 


56 


12 


5 


16 


34 



IISTDEX TO KECENTLY DESCEIBED COCCID^. 



101 



Technical 

Series No. Page. 

austro-africaiius 16 92 

azadiraclitse ] G 63 

bacciformis 16 36 

baculifera 16 36 

badius 16 84 

bahise 16 37 

bakeri 16 64 

Bambusaspis 12 3 

banksise 12 18 

barberi 16 48 

bambusano 16 55 

barrancorum 16 90 

bavaricus 16 92 

beckii ole^ 16 56 

betheli 12 6 

beyerise 16 56 

bezzii 16 37 

biafrse 16 70 

bicolor 16 57 

bicornis 12 17 

bicruciatus 12 8 

bidens 12 13 

biformis 16 54 

bipindensis 16 71 

Birchippia 16 85 

bituberculatum 16 46 

biwakoensis 16 47 

bohemicus 16 86, 87 

boninsis 16 64 

bornmulleii 12 15 

breviseta 16 88 

bruneri s. sp 12 11 

bupleuri 12 12 

burkilli 16 43 

bussei, Ceroplastes 16 44 

bussii, Phenacaspis 16 90 

buzeneiisis 16 71 

cacti ceylonicus " 16 38 

cserulea 12 6 

calophylli 12 9 

calianthina 12 18 

californica 16 35 

calif omicus, Trionymus 16 86 

califomicus, Aspidiotus 12 13 

campylanthi 16 94 

canaiiensis, Chionaspis 16 89 

canariensis, Aspidiotus 16 92 

Candida, Icerya 12 2 

Candida, Chionaspis 12 10 

Candida, Chionaspis 16 48 

candidata 16 42 

capensis 12 13 

16 54 



102 PAPEES O'N COCCID^ OE SCALE INSECTS. 



capimridis 

capsulatus 

cariuata 

cassise 

cassinise 

castanese 

castillose 

catalinaj 

caudata 

cecconi, Lecaniuni 

cecconi, Aspidiotus 

cecropise 

cellulosa 

celtis 

celtium 

ceratonige 

Cercococcus 

cerifenim, Asterolecanium 

ceriferum prominens, Asterolecanium. 

Cerococcus 

Ceronema 

Ceroplastes 

Ceroplastidia s. g 

Ceroplastiua s. g 

Ceroplastodes 

Ceroputo 

ceylonica, Pollinia 

ceylonica s. sp., Hemichionaspis 

ceylonicus 

chelonioides 

chenopodii 

chilina 

chilopsidis 

chinensis 

Chionaspis , 

chionaspiformis 

cMonaspitiformis , 

clusiae 

chitinosus , 

chiton 

cholodkovskyi 

Chrysomplialus 

cinerea 

cingala 

cinnamomi, Chionaspis 

cinnamomi mangiferge, Aulacaspis 

Cissococcus (?) 

cisiidoides 

citri coleorum 

citricola 



Technical 
Series No. 

12 




Page. 
8 


12 




13 


12 




16 


16 




89 


12 




17 


16 




38 


16 




88 


16 




37 


16 




43 


16 




46 


16 




51,72 


16 




40,86 


16 




66 


16 




71 


16 




66 


12 




12 


16 




36 


16 




62 


16 




62 


16 




63,84 


16 




43,66 


12 




7 


16 


44, 65, 88 


16 




65 


16 




65 


16 




45,66 


16 
16 


39, 64, 85 
65 


12 




12 


16 




38 


16 




68 


16 




55 


16 




5b 


16 




57 


16 




58 


12 




10 


16 48 


58 


68,89 


16 




68.91 


16 




91 


16 




56 


16 




71 


16 




45,66 


16 




38 


12 




15 


16 
16 


58 


70,93 

34,58 


12 




18 


12 




10 


16 




90 


16 




85 


12 




15 


16 




40 


16 




68 



INDEX TO EECEJSTTLY DESCKIBED COCCIDtE. 103 



CoCCINiE 

coccineus 

coccophylas 

coccura 

Coccus 

cockerelli 

cockerelliana 

cocotiphagus 

coffese 

colemani 

coleorum 

colimensis 

coloradensis 

comari 

comperei 

C0NCHASFIN.E 

Conchaspis 

conchiformis 

concolor, Physokermes 

concolor, Aonidomytilus.. 

coniferarum shastae 

conservans 

cordaliae 

cordise 

coronatum 

coronifera 

corrugata 

corsa 

corticalis s. sp 

corticis-pini 

corticoides s. sp 

coryli 

coulteri 

coursetiae 

crenulata 

cristata s. sp., Icerya 

cristata, Pseudoparlatoria. 
crotonis 

crudise 

crustuliforme 

cryphseformis 

Cryptaspidiotus 

Cryptaspidus 

Crypthemichionaspis 

Cryptodiaspis s. sg 

Cryptoparlatorea 

Cryptophyllaspis 



Technical 
Series No. 


Page. 


12 


6 


16 


43, 66, 87, 95 


16 


64 


16 


47 


16 


39 


12 


8 


16 


45, 67, 88 


12 


4 


12 


16 


16 


51 


16 


40,84 


16 


39 


16 


40 


12 


2 


16 


58 


12 


7 


16 


39 


16 


52 


12 


3 


16 


35 


12 


3 


16 


68 


12 


10 


12 


16 


12 


13 


16 


69 


16 


87 


12 


11 


16 


62 


12 


11 


12 


17 


12 


12 


16 


37 


16 


52 


12 


14 


16 


46 


12 


6 


16 


52 


12 


16 


16 


34 


16 


71 


12 


5 


16 


86 


16 


72 


16 


67 


16 


48 


16 


71,92 


16 


91 


16 


91 


16 


69 


12 


18 


16 


95 


12 


15 



104 



PAPEES ON^ COCCID^ OE SCALE INSECTS. 



Technical 

Series No. Page. 

CtenocMton 12 7 

cupressi, Leucaspis 12 12 

cupressi, Pseudococcus 16 40 

cupressi, Sphgerococcus 16 85 

cuculis 12 13 

curculiginis 12 15 

curtisi 12 8 

cycliger 16 40 

Dactylopiin^ 12 3 

16 35, 62, 84 

Dactylopius 16 38 

Dactylopius 16 64, 86 

dealbata s. sp 12 4 

decorella these 16 43 

decurvata 12 11 

densiflora 16 52 

dentata 16 70 

dentilobis 16 68 

destefanni 16 57 

Diaspidiotus s. g 12 13 

16 51, 92, 94 

diaspiformis 16 51 

DlASPIN^ 12 10 

16 48, 68, 89 

Diaspis » 12 .11 

16 48, 68, 90 

dimorplius 16 64 

Dinaspis 16 95 

discrepans 12 9 

distorta 16 67 

diversjpes = 12 8 

diversispinus 16 85 

dordsthorpei 16 42 

draper! 16 38 

dudleyi 12 5 

ebeni 12 16 

echinata 12 16 

eglandulosus 16 52 

ehrhorrd 12 13 

elseidis 16 52 

elastica 12 4 

elegans 16 89 

elongatus, Pseudococcus 12 ' 5 

16 40, 58 

elongatus s. sp. , Cryptophyllaspis 12 15 

ephedrae var. , Pseudococcus 12 5 

epliedrse, Leucaspis 16 50 

ephedrge, Parlatoria 16 95 

epigsese 16 52 

eremobius 16 36 

ericicola 16 48 

Eriococcus 12 3, 4 

16 37; 85 



INDEX TO EECENTLY DESCRIBED COCCID^. 



105 



Technical 

Series No. Page. 

erigoni 16 85 

Eriopeltis , 12 7 

etrusca 16 48 

Eucalymnatus 12 7 

Eulecanium , . . . . 12 8 

16 89 

Euparlatoria s. g * . . . 12 18 

Euphilippia 16 44 

euphorbise 16 93 

Evaspidiotus 12 14 

16 52 

exiguum 16 62 

expansum javanicum 12 9 

expansum metallicum 12 9 

expansum quadratum 12 9 

expansum rotundum 12 9 

fabianse 16 94 

Fagisuga 16 35 

fasciata 16 94 

Fernaldella 16 58 

12 16 

Fernaldiella -. 16 58 

ferox 16 54 

festiva 16 34 

fici, Hemichionaspis 16 4^ 

fid, Tachardia. 12 6 

ficifolise ulmicola 16 57 

ficifolii 12 17 

ficus 16 65 

Fiorinia ■ 12 13 

16 50, 91 

fissidens, Aspidiotus 16 69 

fissidens pliu-identatus, Aspidiotus 16 93 

fissus 16 93 

flavociliatum 16 62 

florencise 12 13 

fluminensis 12 3 

folsomi 12 9 

formicarii 16 42 

formicarius 16 64 

formicarum 16 39 

formosa 12 11 

frenchii 16 36 

frontalis 12 8 

f ukayai 16 67 

Furcaspis 16 54 

furcreeicola 16 93 

fuscum 16 46 

galapagoensis 12 3 

galeatus 16 88 

gallicola 16 37 

gilensis s. sp. . 16 37 

68144°— 13 2 



106 



PAPEES ON COCCID^ OK SCALE INSECTS. 



IJ 

glandulosa 

gleditsiae 

Globulicoccus s. g 

glomerella 

goethei 

gorodetskia 

gossypii s. sp 

gowdeyi 

grabhami 

gracilis 

graminicola . - - 

graminis 

graminis, Odonaspis 

grandulifera 

greeni, Asterolecanium . 
greeni, Pseudaonidia. . . 

greeni, Parlatoria 

gryphseformis 

Gymnaspis 

Gymnococcus . . . , 

gymnospori 

gymnosporiae. 

hamata - 

hastata 

Heliococcus 

Hemiberlesia s. g 

Hemichionaspis 

Hemilecanium 

bempeli 

iierculeanus 

hesperidum javanensis. 

hibisci 

himalayensis 

hirsuta 

hirsutus 

hordei 

liowardi 

Houardia 

Howardia 

hymenantherse. 

hypogea 

Icerya 

iceryoides 

ichesii 

imbricans 

immaculatus 

incisus 



Technical 
Series No. 


Page. 


16 


46; 72 


16 


56,91 


12 


11 


16 


46 


12 


6 


12 


6 


16 


35 


16 


49 


16 


88 


12 


6 


16 


54 


16 


39 


16 


40 


16 


55 


16 


87 


12 


3 


16 


54 


16 


58 


16 


48 


12 


16 


16 


56,70 


16 


85 


16 


45 


16 


93 


16 


69 


16 


72,95 


16 


86 


12 


13 


16 


52,92 


12 


12 


16 


49,91 


16 


47, 67, 72 


12 


2 


16 


52 


16 


45 


16 


36 


16 


36 


16 


51 


16 


39 


12 


4 


16 


37 


16 


47 


12 


11 


16 


47 


12 


17 


16 


42 


12 


2 


16 


34, 58, 84 


16 


39 


16 


95 


16 


72 


12 


13 


12 


8 



Technical 

Series No. 


Page.' 


16 


48 


12 


16 


16 


69,91 


16 


43 


16 


50 


16 


63 


12 


8 


16 


68,88 


12 


6 


16 


42 


12 


10 


16 


39 


12 


17 



INDEX TO RECENTLY DESCRIBED COCCIDiE. 107 



mday 

indentata 

indise-orientalis 

indica, Antonina 

indica, Leucaspis 

indicus 

inflatum 

Inglisia '. 

innumerabilis betheli 

inquilina 

insolens 

insolitus 

intermedia victorise. 

irregularis 16 88 

Ischnaspis 16 71, 95 

ixorse 16 66 

jacksoni 16 44 

japonica, Ripersia 16 42 

japonica, Lichtensia 16 68 

javanensis, Aonidia 12 16 

javanensis, Opuntiaspis 12 18 

javanensis s. sp., Coccus 16 45 

javanensis, Aulacaspis 16 49 

javanica 16 49 

javanicum s. sp 12 9 

junctiloba 16 54 

juniperi, Pseudococcus 16 40 

juniperi, Fiorinia 16 51 

kamerunensis 16 72 

kamerunica, Pseudaonidia '. 16 54 

kamerunicus, Pseudaonidia 16 54 

kelloggi, Leucaspis 12 12 

kelloggi, Ripersiella 16 41 

kermanensis ,. 12 12 

Kermes, = 16 ' 36, 85 

kinshinensis 16 68 

koebelei 16 85 

koebeli 16 66 

kunoensis 16 46, 72 

kuwacola 16 44 

kuwanae 12 4 

labiatariun 16 52 

Lachnodiella 16 40, 86 

lagerstroemige 16 37 

lahillei, Ceroplastes 16 65 

lahillei, Gymnococcus 16 85 

lahillei, Dinaspis , 16 95 

lanceolatum 16 62 

larrese s. sp., Icerya 12 2 

16 58 

larrese, Protargionia 16 94 

latialis 16 37 

lauretorum 16 39 



108 



PAPEES ON COCCID^ OE SCALE INSECTS. 



Lecanium 

Lecaniodiaspis 

Lecanopsis 

lectularius 

ledi 

Lefroyia 

leonardi 

Lepidosaphes 

Leucaspis 

leucaspis 

Leucodiaspis 

libera 

Lichtensia 

lidgetti 

lilacinus 

limnanthemi 

limuloides 

lineare 

littoralis tonilensie 

liistneri 

loewi 

lobulata 

longa s. sp 

longa, Aonidia 

longipes 

longiseta 

longisetosa, Tachardia 

longisetosa, Icerya 

longispina 

longivalvata 

longula 

Lophococcus 

lounsburyi 

lutea 

Luzulaspis 

lycii 

macrocarpse 

Macrocerococcus 

macroprocta 

madagascariensis s. sp 

mseandrius 

magna 

malaboda 

malleolus 

mammillaris 

mangiferse, Parlatoria 

mangiferee s. sp., Monophlebiis. 



Technical 
Series No. 

12 


Page. 
10 


16 


46, 67, 


16 
16 


72, 88, 89 
36,84 

47 


16 


53 


16 


86 


16 


38 


12 


12 


12 


16 


16 
12 


66, 71, 94 

12 


16 
12 


50, 69, 91 

18 


16 


50,69 


16 


42 


16 


68 


16 


91 


12 


5 


12 


10 


16 


69 


16 
12 


35,62 

2 


12 


9 


16 


50 


12 


11 


16 


62 


16 


92 


16 


41 


16 


88 


16 


87 


16 


84 


16 


89 


16 


66 


16 


57 


16 


84 


16 


54 


16 


90 


16 


87 


16 


87 


16 


35 


16 


38 


16 


51 


16 


41 


16 


69 


16 


54 


16 


63 


12 


15 


16 


58 


16 


93 


16 


58 


12 


2 



INDEX TO EECENTLY DESCKIBED COCCID^. 



109 



mangiferse s. sp. , Aulacaspis 

manni 

Margarodes 

Margarodin.e 

mai-supialis 

martelli 

Matsucoccus 

matsumurse 

matumuree ". 

maxima 

maxixaus 

mcgregori 

mediterraneus, Margarodes 

mediterraneus, Cryptaspidiotus 

Melanaspis s. g , 

mendozinus 16i 

meridionalis 16 

Mesolecanium 12 

le 

metallicum s. sp 12 

ineyeri 16 

micropori 16 

Micrococcus — . 16 

miliaris longa 16 

miliaris robusta 16 

Mimosicerya, n. sec. Icerya 12 

minima 16 

minor, Icerya 16 

minor s. sp., Asterolecaniiim. . 16 

minutus 16 

mirabiUs tricornis 16 

mirmecophila 16 

missionum ' 16 

mitcbelli 16 

miyasakii 16 

MONOPHLEBIN^ 12 

16 

Monophlebus 12 

16 

montanus 16 

montonose 16 

moreirai 12 

mori, Aspidiotus 16 

mori somereni, Lecanium 16 

muiri 16 

multipora, Lepidosaphes 12 

multipora s. sp. , Fiorinia 16 

multispinosus 16 

muratee 16 

myrmecarius 16 

myrtus 12 



Technical 
Series No. 


Page. 


16 


90 


16 


48 


16 


34 


12 


2 


16 


34 


12 


8 


16 


35 


16 


34 


16 


34 


12 


2 


12 


6 


16 


83 


16 


57 


16 


34 


16 


71 


16 


70,93 



72 

8 
45 

9 
53 
48 
42 
62 
35 

2 
49 
34 
63 
55 
84 
47 
64 
53 
36 

2 
33, 61, 83 

2 
33, 61, 83 
46 
47 
14 
69 
67 
67 
17 
91 
38 
36 
41 
IS 



110 PAPEES 0]Sr COCCIDiE OK SCALE INSECTS. 



mytilaspiformis 

Mytilaspis 

Mytilella 

nanus 

neo-caledonica 

Neolecanium 

nicotianse 

nigra 

nishigaharse 

nucum 

nudata 

nyasse 

oaliuensis 

obtusus 

obscurus 

occidentalis subalpina 

occultus elongatus 

oceanica 

ocellata 

ochnacese 

octocaudata s. sp 

odinss, Fiorinia 

odinse multipora, Fiorinia. 
Odonaspis 

ohioensis 

okadae 

olese, Phenacoccus 

olese, Lepidosaphes 

olivacea 

olivina 

opacus s. sp 

Opisthoscelis (?) 

Opuntiaspis 

orientalis 

orlandi 

ornatus 

Ortbezia 

Ortheziin^ 

ortholobis bruneri 

oryzse 

oxycoccus 

Palseococcus 

Palgeolecanium 

pallidas 

paradoxa 

Paraiecanium 

parcispinosus 

parlatorioides 



Technical 

Series No. 


Page. 


16 


90 


12 


17 


16 


56,94 


12 


16 


12 


5 


16 


91 


16 


67 


16 


45 


16 


91 


16 


47 


16 


91 


16 


90 


16 


88 


16 


85 


16 


86 


16 


41 


16 


66 


12 


15 


16 


54 


16 


57 


16 


67 


16 


33 


16 


51 


16 


91 


12 


16 


16 


55 


12 


14 


16 


34 


16 


64 


16 


56 


12 


3 


16 


44 


12 


7 


16 


84 


12 


18 


16 


44 


16 


49 


16 


63 


12 


3 


16 


35,84 


12 


2 


16 


35,84 


12 


11 


16 


42 


12 


14 


16 


34 


16 


46 


16 


61 


16 


92 


12 


9 


16 


47 


16 


85 


16 


95 



INDEX TO RECENTLY DESCRIBED COCCID^. 



Ill 



Parlatoria 

parlatorise 

Parthenolecanium. . . 

parva 

pasanise 

pectinatus 

penzigi 

perditulum 

pedronis 

penicillata 

peradeniyense 

percerosus . 

pergandii phyllanthi.. 

perniciosus 

perplexa 

Phenacaspis 

Phenacobryum 

Phenacoccus 

philippina s. sp 

phragmitis 

phyllanthi, Targionia. 

phyllanthi s. sp 

Physokermes 

picea 

piceus 

pinicola 

Pinnaspis 

piperis 

pisai 

pistacige 

plana 

pluchese 

plurid^ntatus s. sp 

Pollinia 

polygonata 

Polyocellaria 

popularum 

portoricensis 

privignus 

prominens s. sp 

propsimus 

prosopidis 

Pro targionia 

proteus 

prunastri 

Pseudaonidia 

pseudaspidiotus 

Pseudococcus 



Teclmieal 
Series No. 


Page. 


12 


18 


16 


58, 72, 95 


12 


18 


16 


46 


16 


90 


16 


62 


16 


70 


16 


49 


16 


45 


12 


15 


12 


16 


12 


9 


16 


86 


12 


18 


16 


65 


16 


56 


12 


12 


16 


90 


12 


3 


12 


4 


16 


38,64 


12 


7 


16 


43 


12 


16 


12 


18 


12 


10 


16 


56 


12 


14 


16 


56 


16 


49 


16 


57 


12 


14 


16 


50 


16 


87 


16 


44 


16 


93 


16 


65 


12 


6 


12 


2 


16 


53 


16 


70 


16 


53 


16 


62 


16 


55 


16 


84 


16 


94 


12 


18 


16 


. 46 


12 


15 


16 


54 


12 


18 


12 


5 


16 


40, 58, 86 



112 PAPEES OlS" COCCID^ OE SCALE IIs^SECTS. 



pseud onigrum 

Pseudoparlatoria 

pseudospinosus 

psidii, Saissetia 

psidii philippina, PuUdnaria 

pudibundum 

pugionifera 

pulcher 

pulchra 

pulchrum 

Pulvinaria 

punctulifera 

pusilla, Aonidia 

pusilla, Actenaspis 

pustulans, Sphserococcus 

pustulans, Aspidiotus 

pustulans sambuci, Asterolecanium 

pustulans seychellarum, Asterolecanium. 

pyri ■ 

quadratum s. sp 

quadriclavatus 

quadrilineatus 

quercus gilensis 

ramonse 

rattani 

recurvatum 

regularis 

tehi, Asterolecanium 

tehi, Pulvinaria 

teplicatus 

resinatum 

reticulatus 

rhizopMlla 

Rhopaloaspis 

liccee 

fiieyi larreae 

Ripersia 

Ripersiella 

ripersioides 

riverse 

robusta s. sp 

rombica 

rosse australis 

roseus 

rotundiun s. sp 

rubellum 

rubrocomatum 



Technical 
Series No. 

16 


Page. 
67 


16 


56,71 


12 


14 


12 


10 


12 


7 


16 


62 


16 


68 


16 


34 


12 


16 


12 


9 


16 


46,72 


12 


6 


16 

12 


43, 66, 87 
10 


12 


16 


16 


50 


12 


4 


12 


14 


12 


3 


16 


62 


16 


58 


12 


9 


. 12 


15 


16 


65 


16 


66 


16 


37 


16 


39 


16 


36 


16 


67 


16 


90 


12 


3 


12 


7 


16 


70 


16 


67 


16 


93 


16 


• 65 


12 


11,13 


12 


11, 13 


12 


2 


16 


58 


16 
16 


42, 65, 87 
41.65 


12 


4 


12 


14 


16 


58 


16 


35 


16 
16 


49 
34 


16 


64 


12 


9 


16 
16 


46,72 
63 



INDEX TO EECENTLY DESCEIBED COCCIDiE. 113 



nibrolineata 

rubrovittatiis 

ruthte 

saccharif olii 

Saissetia 

salinus 

sambuci s. sp 

samoana 

sanguineus 

sardiniae 

sassceri 

schrottky i 

schultzei 

Selenaspidus s. sp 

sericeum 

sen-atus 

senifrons 

seurati 

seychellarum s. sp., Asterolecanium. 

seychellarum cristata, Icerya 

shastse 

shastensis 

sideroxylium 

signiferus 

silvatica 

silvestrii, Micrococcus 

silvestri, Saissetia 

sitnilis 

simplex dealbata 

singularis 

sjostedti, Stictococcus 

sjostedti, Monophlebus 

smithii 

Solenococcus 

Solenophora 

solidaginis 

somereni s. sp 

sorb usee s. sp 

sordidus •. . 

spathulata 

Sphaerococcus 

Sphaerolecanium . , 

spiniger 

spinomarginata 

spinosissima 

spinulosa 

squamosus 

stebbingii mangiferse 

stebbingi octocaudata 

Steingelia 



Technical 
Series No. 

16 


Page. 
51 


12 


17 


16 


56 


16 


41 


12 


9 


.16 


67,89 


16 


85 


12 


3 


16 


93 


12 


7 


16 


42 


16 


68 


12 


7 


16 


94 


16 
16 


54,93 
46 


12 


7 


16 


57 


16 


70 


16 


62 


16 


34 


12 


13 


16 


85 


16 


67 


12 


8 


16 


54 


16 


42 


16 


89 


16 


43 


12 


4 


16 


66 


12 


8 


16 


61 


16 


65 


16 


36 


16 
12 


36 
3 


16 


67 


12 


7 


12 


3 


16 


95 


12 


4 


16 


38,85 


16 
16 


46 
70 


12 


16 


16 


70 


16 


87 


12 


11 


12 


2 


16 


33 


16 


35 



114 



PAPEES ON COCCID.^ OR SCALE INSECTS. 



Stictococcus 

Stigmacoccus 

stilosa 

Stotzia 

striata 

Btrobilanthi 

Bubalpina s. sp 

subandina 

subcorticalis 

euberi s. sp 

subfervens 

Bubnudata 

subrotundus 

subtessellatus 

subrubescens corticoides 

subsphgericus 

substriata 

superbus 

sylvatica 

Tachardia 

TACHARDIINiE 

takse 

TakahasMa 

tamarindus 

tangana 

taorensis 

taprobanus 

Targionia s. g 

targioniopsis 

tayabanus, Aspidiotus 

tayabanus, Pseudococcus 

taxifoliae 

taxTis 

tectarius 

tsnebricopbilum 

tenuior 

tenuissimum 

tenuitectus 

tenuivalvatum 

theee, Euparlatoria 

theee s. sp., Tachardia 

theae ceylonica, Hemichionaspis. 

thetecola 

theobromae, Paleeococcus 

theobromee, Ceroplastes 

theobromse, Hemilecanium 

tbesi^esiee, Asterolecanium 

thespesise, Pulvinaria 

tinerf ensis 



Technical 
Series No. 

12 


Page. 

7 


16 


38,64 


16 


83 


16 


69 


16 


44 


16 


44 


12 


" 12 


16 


66 


16 


84 


12 


11 


16 


55 


12 


14 


16 


90 


16 


88 


12 


7 


12 


14 


16 


88 


16 


68 


16 


38 


12 


11 


12 


6 


16 


43,87 


12 


6 


16 


43,87 


16 


41 


16 


68 


16 


33 


16 


90 


16 


94 


12 


15 


12 


14,16 


le 


55, 71, 94 


16 


92 


12 


15 


12 


5 


12 


10 


16 


55 


16 


94 


12 


10 


16 


72 


16 


63 


16 


44 


16 


89 


12 


18 


16 


43 


12 


12 


16 


41 


16 


34 


16 


45 


16 
16 


47,72 
63 


16 


66 


16 


94 



IISTDEX TO EECEISTTLY DESCEIBED COCCIDiE. 115 



tonilensis s. sp., Icerya 

townsendi, Monophlebulus 

townsendi, Hemichionaspis 

trabuti 

Ti-abutina 

transcaspiensis 

travancorensis 

tremge 

tricarinatus 

triglandulcsus 

triloba 

Trionymus '. 

troglodytes 

TruUifiorinia 

tsugae 

tubercularis 

tuherculatus 

tumida 

tumidum 

tyleri 

uberifera 

Tidagama? 

Tigandse 

Tilmicola s. sp 

Ultracoelostoma s. g 

undulatus 

ungulata 

unicolor 

uniloba 

unita 

uvarise 

vagabundus 

varians 

variegatus corticalis 

variolosum minor 

varipes 

vastus 

verrucosee 

virescens 

virgatus var., Pseudococcus 

vii'gatus madagascariensis, Pseudococcus. 

victorise 

vinsonioides 

viridis 

vitis arbutus, Targionia 

vitis opacus, Pulvinaria 

vitis sorbusae, Pulvinaria 

vitis suberi, Targionia 

vitis verrucosae, Pulvinaria 

volynicus 

vovse 



Technical 

Series No. 

12 


Page. 
2 


16 


58 


12 


2 


12 


V 12 


16 
12 


53 

4 


16 


53 


16 


95 


16 


89 


12 


4 


16 


55 


16 


35 


12 


4 


16 


58,86 


16 


47 


16 


50 


16 


94 


16 


49 


16 


45,67 


16 


51 


16 


63 


12 


7 


16 


95 


16 


63 


16 


88 


16 


57 


12 


2 


16 


70 


12 


17 


16 


58 


16 


72 


16 


91 


16 


45 


12 


5 


16 


94 


16 


37 


16 


63 


16 


84 


16 


37 


12 


7 


16 


66 


12 


5 


16 


41 


12 


17 


16 


88 


16 


92 


16 


71 


12 


7 


12 


7 


16 


55 


12 


7 


16 


39 


16 


41 



116 



PAPERS 0]Sr COCCID^ OR SCALE INSECTS. 



Technical 

Series No. Page. 

vuiJleti, Aspidiotus 16 53 

vnilleti, Ceroplastes 16 45 

vuilleti, Chionaspis 16 48 

Walkeriana 16 34, 61 

Websteriella s. g 12 18 

wilga 12 17 

Xylococcus 12 2 

16 35 

yulupse 16 53 

zimmermanrii 16 83 

zonatum 12 9 



ADDITIONAL COPIES o f this publication 
-^ may be procured from the Superintend- 
ent OF Documents, Government Printing 
Office, Washington, D. C. , at 5 cents per copy 




Technical Series, No. 17, Part I. 

U. S. DEPARTMENT OF AGRICULTURE, 

BTJRE^XJ OF E]SrTO]M:OL,OG-^^. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



CONTRIBUTIOliS TOWAED A MONOGRAPH OF 
THE SCOLYTID BEETLES. 



I. THE GENUS DENDROCTONUS. 



A. D. HOPKINS, Ph. D., 

In Charge of Forest Insect Investigations. 



Issued June 30, 1909. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 

1909. 

0/ 



LETTER OF TRANSMITTAL. 



U. S. Department of Agriculture, 

Bureau of Entomology, 

Washington, D. C, March 16, 1909. 

Sir: I have the honor to transmit herewith manuscript of the first 
part of a bulletin of the technical series to be entitled ''Contributions 
toward a Monograph of the Scolytid Beetles." This family of beetles 
includes some of the most important enemies of North American 
forests, as well as of crude forest products, and therefore must demand 
special attention in future systematic and economic investigations 
by this Bureau and by forest entomologists connected with other 
pubhc institutions and private enterprises. It is necessary, as a foun- 
dation for such work, that the heretofore-described species should be 
accurately identified, that those new to science should be described, 
and that tliis information, together with other systematic data 
based on original research by this Bureau, should be made available 
in the form of contributions to a monograph. This work has been 
undertaken by Doctor Hopkins, of tliis Bureau, and the greater 
part of the collecting and working up of the material has been com- 
pleted. Delay in pubHcation will be avoided, and it is beUeved 
that the pubhshed results will be more useful in future systematic 
and economic investigation if the results relating to groups of species 
which have similar characters and characteristics and similar rela- 
tions to given economic problems are pubhshed as parts of a bulletin 
rather than in one undivided pubhcation. These technical parts 
are to be supplemented in a hke manner by parts of a bulletin of 
the regular series, giving information of immediate practical impor- 
tance to the forester and owners and managers of private forests. 

The first part of this bulletin is entitled "The Genus Dendroc- 
tonus." It embodies the results of extensive systematic investiga- 
tions of the genus, carried on by Doctor Hopkins during the past 
seventeen years, and is of especial interest and importance from the 
fact that it deals with a small group of beetles which are the most, 
destructive enemies of the principal coniferous forest trees of North 
America. 

The discussions and illustrations relating to anatomical and tech- 
nical details are necessary as a basis for the correct description. 



IV LETTER OF TRANSMITTAL. 

interpretation, and recognition of generic and specific characters, on 
which depends the future success of economic work on tlie scolytid 
beetles, and as aids in the preparation of technical and economic 
contributions to the monograph. 

I recommend the publication of this paper as Part I of Technical 
Series, No. 17, of the Bureau of Entomology. 

Respectfully, L. O. Howard, 

(Jhief of Bureau. 
Hon. James Wilson, 

Secretary of Agriculture. 



< 



( 



PREFACE TO BULLETIN. 



During the writer's investigations of extensive insect depredations 
in the forests of West Virginia, from 1890 to 1902, he was forcibly 
impressed with the importance of the forest-insect problem in any 
future efforts toward the successful management of the forests of 
this country, and was thus led to give special attention to the sub- 
ject. It was soon realized that among the principal groups of insect 
enemies of forest trees the scolytid bark and wood boring beetles 
must occupy first rank, both in economic importance and in system- 
atic interest. Subsequent investigations in West Virginia, in con- 
nection with the work of the West Virginia Agricultural Experiment 
Station, and in all of the principal forest regions of the country, in 
connection with the work of the Bureau of Entomology, have served 
to confirm these first impressions. 

In these investigations special efforts have been made to acquire 
information on habits and seasonal history of the various species, 
and other facts relating to them, and to collect an abundance of mate- 
rial for systematic study — all to form a basis for conclusions in regard 
to the principal enemies of American forests and practical methods 
for their control. 

The large amount of material has been pretty thoroughly worked 
over and identified, and synoptic tables and descriptions for the 
greater part have been completed for some time. Delay in publish- 
ing the results of the systematic part of the investigations has seemed 
necessary, in order that sufficient time might be given for the deter- 
mination of taxonomic details as a basis for reliable conclusions in a 
comprehensive treatment of the group as a whole, but with increas- 
ing duties in the general work on forest insect investigations, and the 
more and more limited time available for systematic work, it is realized 
that these taxonomic studies can not be completed for many years. 

To avoid further delay in the publication of data of immediate 
interest and importance, the writer has decided to postpone the dis- 
cussion of the taxonomic and other subjects of a philosophical nature, 
required for a completed monograph, and for the present to issue 
separate contributions, each part to be restricted to one genus, or, at 



VI THE SCOLYTID BEETLES. 

most,- to a few closely allied genera. These parts will include synop- 
tic tables, the necessary revisions of old descriptions of species and 
genera, and descriptions of genera and species which appear to be 
new to science. 

It is proposed to follow these technical contributions with parts of 
a bulletin in the regular series, to include the determined bionomic 
and economic facts. 

A somewhat comprehensive treatment of the anatomical details, 
fully illustrated, is given in Part I of this technical bulletin in order 
that it may serve as a basis for reference and comparison in the sub- 
sequent treatment of the other genera and groups of the family. 

LABELS AND RECORDS OF TYPE AND OTHER MATERIAL. 

A single specimen (a female, if possible) is designated as the type 

of a described species by a printed red label ("Type No. , 

U.S.N.M.")j with the type catalogue number of the U. S. National 
Museum written in the blank. When additional specimens are avail- 
able, the type, with one other specimen representing the opposite sex, 
labeled " i type" (or "9 type") on red label, without type number, 
together with revision types °' and other specimens showing range in 
variation, constitute the type series which is deposited in the type 
collection of the U. S. National Museum. Other paratypes and typ- 
ical examples of revised descriptions, comprising one or more speci- 
mens of each species. described, are marked with small red labels, and 
together with the duplicate collection of pinned, alcoholic, and bio- 
logic material, are kept in the reference collection of the Branch of 
Forest Insect Investigations, Bureau of Entomology, U. S. Depart- 
ment of Agriculture. 

All pinned, alcoholic, and biologic material collected or received 
from correspondents are referred to in the field or laboratory records 
and bears number labels, each number referring to a consecutively 
numbered record of the observations made at the time the specimens 
were collected or received. 

Material collected by the writer during his connection with the 
West Virginia Agricultural Experiment Station between 1890 and 
1902 is designated by ''Hopk. W. Va.," number labels. Material 
collected by the writer during liis temporary employment on special 
explorations and trips of investigation for the U. S. Department of 
Agriculture between 1899 and July, 1902, as well as that collected 
during the investigations of forest insects subsequent to April, 1902, 
or received from correspondents, is distinguished by a ''Hopk. U. S." 
number label. In addition to the note number label each completely 

a The term "revision type" is used to designate the specimens, male and female, 
on which a revised description is based. 



PREFACE. VII 

labeled mounted specimen bears labels which supply the following 
data: Collector or correspondent, locality, collecting or rearing date, 
host, and sex. The numbered notes are permanently bound in vol- 
umes of 1,000 numbers, each note relating to one or more species and 
to one or more specimens. The "Hopk. W. Va." numbers begin with 
1 and were limited in June, 1902, to 7,791, and in January, 1907, to 
7,793. The ^'Hopk. U. S." numbers began with 1 in April, 1899, and 
will be Hmited to the period during which the writer is in charge of 
the Branch of Forest Insect Investigations in the Bureau of Ento- 
mology. 

MATERIAL STUDIED. 

Unless otherwise mentioned, the material which forms the basis of 
information and study, so far as it relates to matter in this bulletin,, 
is that bearing the ''Hopk. U. S." or "Hopk. W. Va." note numbers. 
The former is in the forest insect collection of the Bureau of Ento- 
mology, U. S. Department of Agriculture; the latter belongs to the 
collection of the West Virginia Agricultural Experiment Station, but 
at present is in charge of the author, and forms a distinct part of the 
forest insect collection of the Bureau of Entomology. 

ABBREVIATIONS. 

The abbreviations adopted in this publication in referring to mate- 
rial in the different collections examined and that identified by the 
writer are as follows: 

D. A. — Division and Bureau of Entomology, U. S. Department of Agriculture, 
Washington, D. C, other than Hoph. U. S. 

Hopk. U. S. — Branch of Forest Insect Investigations, Bureau of Entomology, U. S. 
Department of Agriculture, Washington, D. C. 

Hopk. W. Va. — W. Va. Agricultural Experiment Station, Morgantown, W. Va. 

U. S. N. M. — U. S. National Museum, Washington, D. C. 

H. & S. — H. G. Hubbard and E. A. Schwarz collection in the U.S. National Museum. 

B. & S.— Collected by H. S. Barber and E. A. Schwarz for the U. S. National 
Museum. 

Soltau. — H. Soltau collection in the U. S. National Museum. 

Lee. — Le Conte collection, Museum of Comparative Zoology, Cambridge, Mass. 

Horn. — Horn collection, American Entomological Society,. Philadelphia, Pa. 

A. E. S. — American Entomological Society, Philadelphia, Pa. 

A. M. N. H. — American Museum of Natural History, New York, N. Y. 

N. Y. S. M.— New York State Museum, Albany, N. Y. 

Harris. — Harris collection, Boston Society of Natural History, Boston, Mass. 

Fitch. — Asa Fitch collection, as represented in the U. S. National Museum. 

Wickham. — Wickham collection of Scolytidte in the U. S. National Museum. 

ACKNOWLEDGMENTS. 

The writer desires to acknowledge the indispensable assistance and 
encouragement, during his earlier studies of the scolytid beetles 



VIII THE SCOLYTID BEETLES. 

(1890-94), rendered by Mr.'E. A. Schwarz, of the U. S. National 
Museum, Oberf orster W. Eichhoff , of Strasburg, Germany, Mr. W. H. 
F. Blandford, of London, England, and M. L. Villard, of Lyon, 
France. Messrs. Eichhoff and Schwarz especially were most kind 
and generous in furnishing identifications of species and in liberally 
loaning and donating specimens. Finally, the writer wishes to ac- 
knowledge the help of his associates and assistants during the prose- 
cution of the work, and especially Mr. W. E. Rumsey, of the West 
Virginia Experiment Station, and the office and field force of the 
Branch of Forest Insect Investigations, in the Bureau of Entomology. 

A. D. H. 



CONTENTS. 



Page. 

Introductory : ' 1 

Historical 3 

Original description of genus 4 

Translation • 4 

Synonymy 5 

Revisional notes : 5 

Revised description of genus 5 

Anatomical 5 

Nomenclature 7 

Illustrations 9 

External characters of the imago 9 

Xhe head 11 

The thorax 22 

Divisions of the thoracic segment 23 

Elements of the adult thorax 26 

The prothorax .' 26 

The mesothorax 28 

Mesotergum 28 

Mesoplura 29 

Mesosterna 30 

The metathorax. 30 

Metatergum 30 

Metapleura 34 

Metasterna 35 

The abdomen. .*. 35 

Abdominal tergites 36 

Abdominal pleurites 38 

Abdominal sternites 38 

Spiracles 39 

The legs 39 

The wings 40 

Mesothoracic and metathoracic wings 42 

Metathoracic or hind wings 43 

Mesothoracic wings or elytra 49 

Internal anatomy 51 

Digestive system 52 

Secondary sexual characters 52 

Pupa 53 

Larva 57 

External characters 57 

Digestive system 64 

Eggs 64 

Physiological characteristics 64 

IX 



X THE SCOLYTID BEETLES. 

ragp. 

Specific distinctions -66 

Range or limits of specific variation 66 

Progressive modifications 67 

Distinction of major and minor divisions of the genus 68 

Plan of synoptic treatment , * 69 

Synopses oi morphological and physiological character.? 69 

Synopsis of adult characters 69 

Synopsis of secondary sexual characters 73 

Synopsis of pupal characters 73 

Synopsis of larval characters 75 

Synopsis of gallery characters 76 

Table of distribution 77 

Table showing relation of species to host trees 78 

Table of host trees 79 

Revision and systematic notes, with descriptions of new species 79 

Distinctive generic characters 79 

Bibliography and synonymy of genus 80 

Division 1 81 

Subdivision A 81 

1. Dendroclonus brevicomis Le Conte 81 

2. Dendroclonus barberi n. sp 85 

3. Dendrodonus convexifrons n. sp 87 

4. Dendrodonus frontalis Zimmerman 90 

5. Dendrodonus arizonicus n. sp 95 

6. Dendrodonus mexicanus Hopkins 97 

7. Dendrodonus imrallelocollis Chapuis 99 

8. Dendrodonus approximatus Dietz 101 

Subdivision B 105 

9. Dendrodonus monticolx Hopkins 105 

10. Dendrodonus ponderosse Hopkins 109 

11. Dendrodonus jeffreyi n. sp 114 

DiAdsion II 116 

Subdivision C 117 

12. Dendrodonus simplex Le Conte 117 

13. Dendrodonus pseudotsugx Hopkins 121 

14. Dendrodonus piceaperda Hopkins 126 

15. Dendrodonus engelmann in. sp 130 

16. Dendrodonus borealis n. sp 133 

17. Dendrodonus obesus (Mannerheim) 135 

18. Dendrodonus rufipennis (Kirby) 138 

19. Dendrodonus murrayanse n. sp 140 

20. Dendrodonus punctatus Le Conte 142 

21. Dendrodonus micans (Kugelann) 143 

Subdivision D ] 46 

22. Dendrodonus terebrans (Olivier) 147 

23. Dendrodonus ralens Le Conte 151 

24. Dendrodonus adjunctus Blandford 157 

Bibliography - 159 



ILLUSTRATIONS 



PLATES. • 

Page. 
Plate I. Classification of the genus Dendroctonus, showing technical and com- 
mon names and species numbers 1 

II. Map of world, showing geographical distribution of the genus Den- 
droctonus 80 

III. Dendroctonus adults. Fig. 1. — D. brevicomis. Fig. 2. — D. harheri. 

Fig. 3. — D. convexifrons. Fig. 4. — D. frontalis. Fig. 5. — D. ari- 

zonicus. Fig. 6. — D. mezicanus. Fig. 7. — D. parallelocollis 164 

IV. Dendroctonus adults. Fig. 8. — D. approximatus . Fig. 9. — D. monti- 

colse. Fig. 10. — D. ponder OS iv. Fig. 11. — D. jeffreyi 164 

V. Dendroctonus adults. Fig. 12. — D. simplex. Fig. 13. — D. pseudo- 

tsugx. Fig. 14. — D. piceaperda. Fig. 16. — D. borealis 164 

VI. Dendroctonus adults. Fig. 17. — D. obesus. Fig. 18. — D. rufipennis. 

Fig. 20. — D. punctatus. Fig. 21. — D. micans 164 

YII. Dendroctonus Skdults. Fig. 22. — D. terebrans. Fig. 23. — D.valens.... 164 
VIII, Denrfroctonws larvae. Fig. 1. — D. brevicomis. Fig. 19. — D.murrayanx. 

Fig. 23.— Z>. valens 164 

TEXT FIGURES. 

Fig. 1. Dendroctonus valens: Adult, dorsal aspect 6 

2. Dendroctonus valens: Adult, ventral aspect 8 

3. Dendroctonus valens: Adult, lateral aspect 9 

4. Dendroctonus valens: Head, dorsal and lateral aspects 12 

5. Dendroctonus valens: Head, ventral aspect, and mouthparts 13 

6. Dendroctonus valens: Head, oral aspect, epistoma, etc 14 

7. Pterostichus californicus: Head, dorsal and ventral aspects 15 

8. Pissodes strobi: Head, ventral aspect, and mouthparts 16 

9. Pissodes strobi: Head, dorsal aspect, and mandibles 17 

10. Dendroctonus: Epistomata .- 19 

11. Dendroctonus: Antennae 20 

12. Dendroctonus: Antennae 20 

13. Dendroctonus: Antennae 21 

14. Dendroctonus valens: Mandible 22 

15. Dendroctonus: Eyes 23 

16. Dendroctonus valens: Areas of pronotum 23 

17. Dendroctonus valens: Areas of prothorax, ventral aspect 24 

18. Dendroctonus valens: Mesothorax, ventral aspect 24 

19. Dendroctonus valens: Mesotergum and mesopleurum 28 

20. Dendroctonus valens: Metatergum and metapleurum 31 

21. Dendroctonus valens: Metatergum, inner aspect 33 

22. Dendi octonus valens: Abdominal tergites 35 

23. Dendroctonus valens: Male, abdominal tergites 7 and 8 36 

24. Dendroctonus valens: Female, abdominal tergites 7 and 8 37 

XI 



XII THE SCOLYTID BEETLES. 

Page. 
Fig. 25. Dendroctonus valens: Abdominal stemites, ventral and lateral as- 
pects 38 

26. Dendroctonus valens: Tibia, tarsus, articulation, etc 40 

27. dendroctonus: Left tibiae, dorsal and ventral aspects 41 

28. Dendroctonus: Left tibiae, dorsal and ventral aspects 42 

29. Dendroctonus: Left tibiae, dorsal and ventral aspects 43 

30. Dendroctonus valens: Diagram of basal area of hind wing 44 

31. Dendroctonus valens: Right elytron, ventral aspect 45 

32. Dendroctonus valens: Basal process of right elytron 46 

33. Dendroctonus valens: Declivital section of elytra 46 

34. Dendroctonus valens: Diagram of elytron, showing striae, interspaces, 

and tracheae 47 

35. Dendroctonus valens: Digestive organs of adult 62 

36. Dendroctonus valens: Fore intestine, showing details 53 

37. Dendroctonus valens: Pupa, dorsal, lateral, and ventral aspects. . . 54 

38. Dendroctonus valens: Pupa, lateral and anal aspects 55 

39. Dendroctonus valens: Larva, dorsal, lateral, and ventral aspects 58 

40. Dendroctonus -valens: Head of larva. 59 

41. Dendroctonus valens: Mouthparts of larva 60 

42. Dendroctonus valens: Mandibles of larva 63 

43. Dendroctonus valens: Digestive organs of larva 65 

44. Dendroctonus brevicomis: Egg galleries 82 

45. Dendroctonus brevicomis: Bark showing pupal cells, exit burrows, and 

pitch tubes 83 

46. Dendroctonus brevicomis: Distribution map 84 

47. Dendroctonus barberi: Egg galleries 86 

48. Dendroctonus barberi: Distribution map 87 

49. Dendroctonus convexifrons: Egg galleries and lar^■al mines 89 

50. Dendroctonus convexifrons: Distribution map 90 

51. Dendroctonus frontalis: Egg galleries and larval mines 91 

52. Dendroctonus frontalis: Termination of egg galleries 92 

. 53. Dendroctonus frontalis: Beginning of egg galleries 92 

54. Dendroctonus frontalis: Bark showing pitch tubes, etc 92 

55. Dendroctonus frontalis: Old egg galleries in living tree 93 

56. Dendroctonus frontalis: Egg gallery in living tree, the resulting wound 

in process of healing 93 

57. Dendroctonus frontalis: Distribution map 94 

58. Dendroctonus arizonicus: Distribution map 97 

59. Dendroctonus mexicanus: Section of egg galleries 98 

60. Dendroctonus mexicanus: Distribution map 99 

61. Dendroctonus parallelocollis: Section of egg gallery 100 

62. Dendroctonus parallelocollis Distribution map 101 

• 63. Dendroctonus approximatus: Single egg gallery 102 

64. Dendroctonus approximatus: Egg galleries 103 

65. Dendroctonus approximatus: Distribution map 104 

66. Dendroctonus monticolae: Egg galleries and larval mines in bark 107 

67. Dendroctonus monticolae: Egg galleries and larval mines grooved in sur- 

face of wood 108 

68. Dendroctonus monticolae: Distribution map 109 

69. Dendroctonus ponderosse: Egg galleries and larval mines 112 

70. Dendroctonus ponderosse: Tree with bark removed, showing egg gal- 

leries grooved and marked on surface of wood 113 

71. Dendroctonus ponderosse : Distribution map 114 



ILLUSTRATIONS. XIII 

Page. 

Fig. 72. Dendroctonus jeffreyi: Distribution map \ 116 

73. Dendroctonus simplex: Egg galleries and larval mines 119 

74. Dendroctonus simplex: Distribution map 120 

75. Dendroctonus pseudotsugx: Egg galleries and larval mines 122 

76. Dendroctonus pseudotsugse: Egg gallery and larval mines 123 

77. Dendroctonus pseudotsugse: Section of log with bark removed, showing 

brood galleries marked and grooved on surface of wood 124 

78. Dendroctonus pseudotsugse: Distribution map 125 

79. Dendroctonus piceaperda: Egg gallery and larval mines 128 

80. Dendroctonus piceaperda: Distribution map 129 

81. Dendroctonus engelmanni: Egg gallery and eggs in living bark 132 

82. Dendroctonus engelmanni: Distribution map 133 

83. Dendroctonus borealis: Distribution map 134 

84. Dendroctonus obesus: Distribution map 137 

85. Dendroctonus rufipennis: Distribution map 140 

86. Dendroctonus murrayanse: Distribution map 141 

87. Dendroctonus punctatus: Distribution map 143 

88. Dendroctonus micans: Egg galleries and larval chamber 145 

89. Dendroctonus micans: Distribution map 146 

90. Dendroctonus terebrans: Distribution map 150 

91. Dendroctonus valens: Egg galleries and larval chamber 1-52 

92. Dendroctonus valens: Work in bark at base of stump 153 

93. Dendroctonus valens: Basal wound in living tree resulting from primary 

injury by this species 154 

94. Dendroctonus valens: Distribution map 155 

95. Dendroctonus adjunctus: Distribution map 158 



Tech. Series 17, Part I, Bureau of Entomology, U, S. Dept. of Agriculture. 



Pl-xte I. 



> 

s 



CO 

02 02 






a^ 



|j4 









i! 

o C 
O 



i 

Ph 




1. hrevicomis Lee. 

2. barberi Hopk. 



1. Western Pine 70,81 

Beetle. 

2. Southwestern Pine 70, 85 

Beetle. 



3. convexifrons Hopk. 3. Roundheaded Pine 70, 87 
Beetle. 



4. frontalis Zimm. 

5. arizonicus Hopk. 

6. mexicanus Hopk. 



4. Southern Pine 70,90 

Beetle. 

5. Arizona Pine 70,95 

Beetle. 



6. Smaller Mexican 70, 97 
Pine Beetle. 

7. par allelocolUs ChsiTp. 7. Larger Mexican 70,99 

Pine Beetle. 

8. approximatus Dietz. 8. Colorado Pine 70,101 

Beetle. 



9. monticolse Hopk. 
LO. ponderosiE Ho; 
11. jeffreyi Hopk. 



9. Mountain Pine 71, 105 
Beetle. 

10. ponderosie Hopk. 10. Black Hills Beetle 71, 109 

11. Jeffrey Pine Beetle 71,114 



12. simplex Tuec. 12. Eastern Larch 71,117 

Beetle. 

13. pseudotsugse Hopk. 13. Douglas Fir Beetle 71, 121 

14. piceaperda Hopk. 14. Eastern Spruce 71,126 

Beetle. 

15. engelmanni Hopk. 15. Engelmann Spruce 71, 130 

Beetle. 

16. borealis Hopk. 16. Alaska Spruce 72, 133 

Beetle. 

17. o&esMS Mann. 17. Sitka Spruce Beetle 72,135 



18. rufipennis Kirhy. 18. Eed winged Pine 72,138 

Beetle. . 

19. murrayanse Hopk. 19. Lodgepole Pine 72, 140 

Beetle. 

20. punctatus Lee. 20. Allegheny Spruce 72, 142 

Beetle, 

21. micans Kug. 21. European Spruce 72, 143 

Beetle. 



22. terebrans Oliv. 

23. valens Lee. 



Position doubtful- 



22. Black Turpentine 72, 147 

Beetle. 

23. Red Turpentine 72, 151 

Beetle. 

157 



24. adjunctus Blandf. 24. Guatemala Beetle 

Classiflcatiou of the Genus Dendroctonus, Showing* Technical and Common 
Names and Species Numbers. 

This diagram will enable the reader to refer at once to the technical and common names 
of any species number mentioned in the text, and will show at a glance the position 
and relations of thp divisions, subdivisions, sections, subsections, series, and species 
into which the genus is divided. 



U. S. D. A., B. E. Tech. Ser. 17, Pt. I. F. 1. 1., June 30, 1909. 

CONTRIBUTIONS TOWARD A MONOGRAPH OF THE 
SCOLYTID BEETLES. 



I. THE GENUS DENDROCTONUS. 

By A. D. Hopkins, 
In Charge of Forest Insect Investigations. 

INTRODUCTORY. 

The active work on forest insects conducted by the West Virginia 
Agricultural Experiment Station in 1890-91, and by the Division and 
Bureau of Entomology of the U. S. Department of Agriculture since 
1899, has resulted in the accumulation of a mass of systematic and 
biological data on the principal described and undescribed insect ene- 
mies of forest trees and forest products of the United States. When- 
ever an attempt has been made, however, to work up the material 
relating to a given species, -or group of species, it has been apparent 
that the publication of anything without first describing the new 
species and revising the data in both the s^^stematic and economic 
literature would contribute to confusion rather than to advancement. 
Indeed, it becomes more and more evident that in order to give reliable 
information on applied entomology we must have at our command tJie 
Tiuowledge gained hy careful teclinical, or systematic, studies of the insects 
with which we have to deal. Therefore, when we find, as we do in many 
cases, that the published results of systematic work on a given genus 
or species are meager or otherwise unsatisfactory, it becomes necessary 
to revise and verify the descriptions and biological records, and to 
adjust the classification to meet the requirements of the newly dis- 
covered facts relating to the described and undescribed species. 

The genus Dendroctonus presents a striking example of the need of 
systematic study as a basis for economic investigation. It is both 
the most important group of insect enemies of the coniferous forest 
trees of North America and one of the most difficult for systematic 
study. Le Conte (1876) expressed the difficulty met with in a study 
of the species when he said in his later revision : 

If I have failed to indicate more strongly the differences between these species, it 
is because they are not distinguishable by any prominent or definite characters; and 
the student who may have difficulty in identifying the species as here defined would 
have almost equal difficulty if the specimens in my collection were before him. 

1 



2 THE SCOLYTID BEETLES. 

Until within recent years little progress had been made toward the 
discovery and clear definition of the specific and sexual characters. 
In consequence the identification of the species was both difficult and 
uncertain and has led to much confusion in both systematic and eco- 
nomic literature. With our prasent knowledge of the genus, based 
on an exhaustive study of the systematic and biologic details, most, 
if not all, of the difficulties have been removed, so that the identifi- 
cation of the species is comparatively easy. 

It is the purpose of this paper to revise and bring up to date the 
available information on the described species, to describe those that 
appear to be new to science, and to record the results of original inves- 
tigations relating to the more technical details that can not well be 
included in the paper w^hich is to follow as a part of a bulletin in the 
regular series and which will give full information on the bionomic 
features. 

The material which has served as a basis for the study of this genus 
consists mainly of the notes and specimens taken by the writer in the 
field during his connection with the West Virginia Agricultural 
Experiment Station, between 1890 and July, 1902, includmg special 
investigations for the Division of Entomology, U. S. Department of 
Agriculture, in 1899, 1900, and 1901, and those taken during the 
investigations by this Bureau between July, 1902, and July, 1908. 
In addition to the large amount of material thus accumulated the 
writer has studied the tjrpe and other specimens in the larger collec- 
tions of this country. 

The writer desires to acknowledge, in this connection, the assist- 
ance rendered by the following gentlemen m providing facilities for 
the study of specimens in the collections of which they have charge : 
Mr. Samuel Henshaw, in charge of the Le Conte collection in the 
Museum of Comparative Zoology, Cambridge, Mass., and of the Harris 
collection, Boston Society of Natural History; Dr. Henry Skinner, 
in charge of the Horn collection of the American Entomological 
Society and the general collection of the American Entomological 
Society, Philadelphia, Pa.; Mr. E. A. Schwarz, honorary custodian of 
Coleoptera in the Division of Insects, U. S. National Museum; Dr. 
W. G. Dietz, who loaned type and other specimens from his collec- 
tion, and Mr. C. O. Waterhouse, of the British Museum, who com- 
pared specimens with the type of Dendroctonus rufipennis Kirby. 

It also gives the writer pleasure to acknowledge the efiicient assist- 
ance of Messrs. J. L. Webb, H. E. Burke, and W. F. Fiske in the field 
and office w^ork, of Mr. E. J. Kraus in the more recent office work, and 
of Messrs. J. F. Strauss and R. E. Snodgrass in the preparation of the 
illustrations for this part of the bulletin. 



THE GENUS DENDEOGTONUS. . 3 

HISTORICAL. 

The genus Dendrodonus was described by Dr. W. F. Erichson (18-36) 
to include (BostricJius) micans Kug., (Scolytus) terebrans Oliv., 
(Dermestes) piniperda L., (Hylesinus) minor Hartig, and (Hylesinus) 
minimus Fab. 

Eichhoff (1864) revised the genus and referred D. piniperda (L.) and 
D. minor (Hartig) to BlastophagusJLichh. and later (1879) to Myeloplii- 
lus Eichh., and I), minimus (Fab.) to CarpJiohorus Eichh., leaving D. 
micans (Kug.) as the type. 

Lacordaire (1868) referred to the synonymy and revised the descrip- 
tion, including Dendrodonus junipiri Doeb. { = Phlceosinus junipiri 
(Doeb.)j, B. valens Lee, and Z). similis Lee. 

Zimmerman (1868) divided the genus into three groups, placing 
D. hifurcus (= CarpJiohorus hifurcus Eichh.) in the first, none in the 
second, and D. terebrans (Oliv.) and D. frontalis Zimm. in the third 
group. 

Le Conte (1868) revised the classification for the North American 
species to include D. terebrans (Oliv .), Hylurgus obesus Mann., Hylurgus 
rujipennis Kirby, D. frontalis Zimm., and two new species, D. pundatus 
Lee, and D. simplex Lee. He here referred D. valens Lee. to D. tere- 
brans (Oliv.), and D. similis Lee. to D. obesus (Mann.) . He recognized 
two divisions, Division B represented by D. frontalis, and Division A 
by the other five species. 

Chapuis (1869) included D. micans (Kug.), D. valens Lee, D. tere- 
brans (Oliv.), D. obesus (Mann.), and added one new species, D. paral- 
lelocollis Chap., but did not recognize D. frontalis Zimm. 

Le Conte (1876) included D. terebrans Lac. ( = 01iv.), D. similis 
Lee, D. rufipennis (Kirby), B. pundatus Lee, D. simplex Lee, B. fron- 
talis Zimm., and one new species, B. brevicomis Lee He here 
restored B. similis and omitted D. obesus. 

Dietz (1890) in his ''Notes on the Species of Dendroctonus of 
Boreal America," revised the classification, principally on the char- 
acter of the epistoma, which he considered of primary importance 
in separating the species. He included B. terebrans, with varieties 
a, b, c, d, B. rufipennis, D. similis Lee, B. simplex Lee, B. frontalis, 
added one new species, Z>. approximatus Dietz, and referred D. pundi- 
collis Lee to D. rufipennis (Kirby) and B. brevicomis Lee to B. 
frontalis Zimm. 

Blandford (1897) mentioned D. terebrans (Oliv.), D. parallelocoUis 
Chap., and an undescribed species from Texas — probably D. terebrans 
(Oliv.) — and added one new species, B. adjunctus Blandf. 

The writer (Hopkins, 1899a) referred to B. terebrans, D. rufipennis 
(Kirby), B. simplex Lee, and B. frontalis Zimm., with descriptions of 
79980—09 2 



4 THE SCOLYTID BEETLES. 

different stages, habits, etc., of B. frontalis, and larvae and habits of 
D. terebrans (=B. valens). In 1901 he described D. piceaperda in. 
all stages in connection with an account of habits, seasonal histoiy, 
etc., and referred to the type of D. rufipennis (Kirby). In 1902 he 
described D. ponderosse, in all stages, in connection with an account of 
habits, seasonal history, etc. In 1902, under " Some Notes on the Genus 
Dendroctonus," he referred to a statistical method of determining 
natural positions of the species, and gave a list of described species 
and manuscript names of undescribed species, as follows: D. pinicida 
MSS., D. arizonicus MSS., D. monticola MSS., D. ponderosse, Hopk. 
MSS., I). Iceeni MSS., D.jietcheri MSS., D. piceaperda Hopk., D. dietzi 
MSS., D. californicus MSS., D. sliosTione MSS., D. wickJiami MSS., 
and D. horealis MSS. He restored D. hrevicomis Lee. and D. punctatus 
Lee. from Dietz's synonymy, and recognized D. ohesus (Mann). In 
1905 he described D. pseudotsugse, and D. monticola in connection 
with accounts of habits, seasonal history, etc. 

ORIGINAL DESCRIPTION OF GENUS. 

Dr. W. F. Erichson (1836) described the genus Dendroctonus as 

follows : 

Dendroctonus. 

[p. 52] Antennae funiculo 5-articulato, capitulo 4-annulato, suborbiculari, com- 
presso. Tibise extus denticulatse. 

Palpi maxillares articulo primo brevissimo, secundo maximo, sequentibus duobus 
sensim minoribus. Labium fortiter compressum. Palpi labiales articulo primo 
longiore, subclavato, secundo tenuiore, cylindrico, minuto, tertio obtuse subulato. 
Antennae breves, scapo clavato, funiculi articulo primo breviter clavato, secundo 
obconico, reliquis brevibus transversis; capituli segmentum ijrimuni reliquis con- 
junctis Bequale, politum. [p. 53] Corpus oblongum, cylindricum. Rostrum brevis- 
simum. Prosternum antice obsolete impressum. Coxae anticse approximatae. Tibiae 
compressae, extus denticulatae. Tarsi articulo tertio dilato, bilobo. Elytra margine 
antico elevate. 

[Translation.] 

Antennae with 5-jointed funicle; the club suborbiculate, com- 
pressed, with four segments (annulse). Tibiae externally denticulate. 

Maxillary palpi with the first joint very short, the second the 
longest, the two following gradually smaller. Labium strongly com- 
pressed. The labial palpi with the first joint rather long, subclavate, 
the second joint more slender, cylindrical, small, the third obtusely 
subulate. Antennae short, scape clavate, first joint of funicle shortly 
clavate, second joint obconical, the remaining joints short, trans- 
verse; first segment of club equal to the others conjointly, polished. 

Body oblong, cylindrical. Beak very short. Prosternum ante- 
riorly obsoletely impressed. Anterior coxae approximate. Tibiae 
compressed, externally denticulate. Tarsi with the third joint 
dilated and bilobed. Elytra with the anterior margin elevated. 



THE GENUS DENDROCTOISrUS. 5 

SYNONYMY. 

The following species were included, all but two of which were 
subsequently referred by Eichhoff (1864) to other genera: 

Bostrichus micans Kugelann = Dendroctonus micans (Kugelann) . 

(Type of genus.) 
Scolytus terebrans Olivier = Dendroctonus terebrans (Olivier). 
Dermestes 'pini'perda Linnaeus = MyelopTiilus piniperda (Lin- 
naeus). 
Hylurgus minor Hartig= MyelopMlus minor (Hartig). 
Hylesinus minimus Fabricius = Carphohorus minimus (Fabri- 
cius.) 

REVISIONAIi NOTES. 

The generic characters mentioned by Erichson in the original 
description are recognized in the type and other species except that 
the maxillary palpi are not 4-jointed. The first or basal joint has a 
basal ring or outward curved basal margin for the attachment of the 
membrane connecting the joint with the palpiger. This might have 
been mistaken for the "very short first joint" referred to, but it is 
evident that this or any other structure does not represent such a basal 
joint. In the type species the first joint of the club is equal to the 
others, but ranges from shorter to longer in the other species. 

Le Conte's added characters in his revision of 1868 and 1876 are 
generally correct, except that the antennal club is not always concave 
on one (external) side or anterior face, the sutures are more often 
curved than straight, and in some species only two sutures are 
visible on one side of the club. The prosternum is sometimes flat, the 
fifth joint of the tarsus is never longer than the others united, and the 
ventral segments are only approximately equal in length, the last 
one being usually as long as the two preceding combined. 

Dietz (1890) called attention to the unreliability of the sutures and 
joints of the antennal club in dried specimens, and laid special stress 
on the value of the epistoma in distinguishing the species. It appears, 
however, that while the form of the epistoma is a good generic and 
subdivisional character, it is of little or no value in distinguishing the 
species. 

The additional generic characters recognized by the writer will be 
found described under external and internal anatomy, and the char- 
acters distinguishing the major and minor divisions will be found in the 
synoptic tables. 

REVISED DESCRIPTION OF GENUS. 

ANATOMICAL. 

The following discussion of anatomical details includes the imago, 
larva, and pupa, and is based primarily on the results of original 
dissections and anatomical investigations by the author during the 



6 



THE SCOLYTID BEETLES. 



past eighteen years, and of those conducted by assistants under his 
immediate supervision during the past three years. 




2 'i '. 










'O c3 












S "O 






•" m 






a a 






03 ~ 








■S Vi 




5^ 


•5,^ {S. 




^■.~ 




00 










% 


^ a 



3 3 
<1 



Dendroctonus valens Lee. has served as the principal subject for 
dissection, comparison, and illustration, both on account of the 



THE GENUS DENDEOCTONUS. 7 

abundance of material at hand and because of the comparatively 
large size of the individuals of this species. Sufficient comparative 
studies have been made, however, of the other species of the genus 
and of representatives of other genera of the family and suborders to 
form a reliable basis for the interpretations and conclusions relating 
to the more important taxonomic characters and the significance of 
their modifications in the distinction of species, genera, etc. 

Through the assistance of Mr. R. E. Snodgrass an extensive investi- 
gation has also been made of the thoracic segments of representa- 
tives of all of the principal orders of insects. The results have 
served as additional data and evidence on which to base conclusions 
in this paper, and will be utilized by Mr. Snodgrass as a basis for 
a more detailed discussion in a paper entitled "The Thorax of Insects 
and the Articulations of the Wings," to be published later. This 
will include a quite complete bibliography and references to the 
principal systems of nomenclature proposed or adopted by the lead- 
ing authors, thus rendering it unnecessary to include extensive 
bibliographic references in the present paper. 

In all of this anatomical work the object of the author has been to 
acquire direct information on the facts as they exist in the subjects ex- 
amined; such information to furnish a basis for the determination, 
naming, description, and illustration of the anatomical elements as 
represented in the scolj^tid beetles, and at the same time to serve 
as a guide to the determination of further facts relating to insect 
anatomy in general. 

The literature on insect anatomy has been utilized as a guide in 
securing additional information on the facts and principles involved, 
and with the idea of adopting such interpretations and nomen- 
clature as appeared to conform more nearly to the facts and con- 
tribute to uniformity. No attempt is made to discuss the -merits 
of opposing opinions or theories, or to prove or disprove them. 

In this presentation of the results of independent investigation 
and discussion of the facts as interpreted by the author, it is hoped 
that something has been accomplished toward the advancement of 
information on the general subject of insect anatomy, and that its 
special reference to the anatomy of the scolytid beetles will make 
the future systematic study of this troublesome group less difficult 
and more accurate, and thus lead to the determination of bionomic 
and economic data of immediate practical importance. 

NOMENCLATURE. 

There is yet much confusion in the literature and considerable 
difference of opinion among the best authorities in regard to ana- 
tomical nomenclature as applied to the structure of insects in gen- 
eral and especially to representatives of different orders. There is 



THE SCOLYTID BEETLES. 






111 






,Coxc 
Coxa 







.re 



Hum 



' Exocoxal piece 

Epimervm 

Stcniellar area 

Coxa 

Sternum 

Episternum 

Median line 

Sternellar piece 

^_^^--Coxa 
Epimenim 

^ntercoxal process 
'~ '■ 2d suture 

'~'"Sternite3 



"^^^V. 



Fig, 2.—Dendroctonus valens: Adult, ventral aspect, c, Sternellar area. (Original.) 



THE GENUS DENDEOCTONUS. 



9 



evidently much room for improvement in the line of uniformity in 
names and interpretations. In the present paper the writer has 
endeavored to adhere to the more generally accepted names proposed 



-HEAD 




3 




J 

A 




^■# ■ 


5 CD 



Fig. 'i.—Dendroctonus valens: Adult, lateral aspect, a, Pleural elavicula; 6, pregena. (Original.) 

by Audouin and other writers for the principal parts, and to suggest 
only such revisions and new names as the immediate requirements of 
clear definition in comparative anatomy and taxonomy appear to 
demand. 



10 THE SCOI.YTID BEETLES. 

ILLUSTRATIONS. 

The figures are intended to be sufficiently complete to leave little 
to be added in the way of description, except to emphasize and 
elucidate some of the more important features, or to call attention 
to the variation within the genus or species. 

EXTERNAL CHARACTERS OF THE IMAGO. 

The structural details and general external anatomy and sculpture 
are shown in figures 1, 2, and 3. The principal characters peculiar 
to the genus are found in the large, prominent head, the epistomal 
process (figs. 2, 3, 4, 6, 10) (referred to by Dietz as the median seg- 
ment of the epistoma), the form of the antenna (figs. 11, 12, 13), 
the approximate or subcontiguous anterior coxse (fig. 2), and the 
strongly recurved hypopleural sutures 4, 5, and 6 of the abdominal 
sternites (fig. 25). 

Length and relative proportions. — The length of the imago ranges 
from 2.5 mm. in D. frontalis to 9 mm. in D. valens. There is con- 
siderable range in length within the limits of some of the species, 
while in others the length is more constant. The relative propor- 
tions of the width of the head, width and length of the prothorax, 
width and length of the elytra, or a composite of the ratios, serve 
as a taxonomic index for the classification of the species, and, together 
with other characters, serve to distinguish the major and minor 
divisions and, to a certain extent, the species. The progressive 
modification appears to be from a head nearly as broad as the pro- 
notum and the latter as broad as the elytra, with the sides nearly 
parallel, to a head much narrower than the pronotum, the latter 
slightly narrower than the elytra, with the sides narrowed and con- 
stricted toward the head; also, from a slender, subelongate, to a 
stout body. 

Color. — The color ranges from pale yellowish-red to brown and 
deep black, but is fairly constant in the matured individuals of a 
species. The immature individuals are always lighter, and some of 
those of the black species are reddish. In some species the head, 
prothorax, and ventral surface of the body are darker than the 
elytra, while in others little or no difference is noticeable. 

Vestiture. — The body is more or less clothed with short to long 
hairs, the presence or absence of which on different areas is of far 
more taxonomic significance than was at first recognized. Except 
in old rubbed specimens, the vestiture serves as one of the important 
characters distmgTiishing the major, as well as some of the minor, divis- 
ions. See synopsis. Divisions I and II, sections al and a2 (PI. I) . 

Sculpture.— Within the genus and also within each species there 
is considerable variation in the sculpture of the front, pronotum, 
and elytra. Nevertheless, such characters as the presence or absence 



THE GENUS DENDEOCTONUS. 11 

of frontal grooves and tubercles serve to distinguish . some of the 
minor divisions of the genusj while the presence or absence of a 
posterior median impression in those species without a frontal groove 
is of considerable importance in distinguishing some of the minor 
divisions. The relative size, density, and arrangement of the punc- 
tures of the pronotum, while variable within the species, is of con- 
siderable taxonomic value. The character of the rugosities of the 
interspaces and the punctures of the striae are also variable within 
the species and are of secondary value in distinguishing minor divi- 
sions. The sculpture of the elytral declivity is of special specific 
and sexual importance, and in some cases the characters are of value 
in distinguishing minor divisions. 

THE HEAD. 

The general characters and details of the external skeleton and 
appendages of the head are shown in figures 4, 5, and 6. It will be 
noted that the elements which in some other Coleoptera and other 
insects are more or less clearly defined are quite completely fused and 
obscured in this genus as in other rhynchophorous beetles. The 
labrum and clypeus are obsolete. The epistoma, or "post-clypeus," 
or ''pre-front," as recognized by different authors, is not separated 
from the front by a line or suture, but is quite clearly defined, and the 
epistomal process is far more prominent than in other allied genera. 
The front is completely fused with the epicranium, which in turn is 
fused with the genge, the latter joined beneath with a single gular 
suture. Anterior to the gular suture there are three quite clearly 
defuied sclerites, which may be designated as pregula, pregena, and 
hypostoma (fig. 5, E). By comparing the head of Dendroctonus 
with that of a carabid beetle, PterosticJius (fig. 7), and a typical curcu- 
lionid beetle, Pissodes strohi (figs. 8, 9), the striking diflterence in struc- 
ture and relative position of the corresponding elements and their 
extreme modification are at once apparent. 

Labrum. — The labrum is not present as a distinct element, but may 
be represented by a part of the anterior margin of the epipharynx 
beneath the anterior median section of the epistoma (fig. 6, A). 

Clypeus. — The clypeus is not represented unless it is by the pro- 
duced anterior margin of the epistoma, and by the long epistomal 
bristles. 

Epistoma (figs. 2, 3, 4, 5, 6, 7, 10, 40, B, D, £".)— The epistoma is 
apparently represented in both the larvae and adults of all true man- 
dibulate insects, but is more distmctly defined in some than m others.- 
In some insects it is separated from the clypeus by a suture, Ime, or 
articulating membrane, while in others there is no evidence of sepa- 
ration or the clypeus is not represented. Its separation from the 
front is often defined by a line, impression, elevation, or otherwise, 
although sometimes it is so completely fused that the line of junction 



12 



THE SCOLYTID BEETLES. 



is entirely obscured, as in Pterostichus. It serves the important 
function of a rigid bridge over the oral foramen and support for the 
clypeus, labrum, and epipharynx, and at its lateral angles provides 
the necessary rigid support for the dorsal articulation of the man- 



Mandible 




Maxiltanj palpus^^^\ ^-' 
.Labial palpuA 



Fig. 4. — Dendroctonus valens: Head, dorsal and lateral aspects. A, Dorsal aspect of head; 5, lateral aspect 
of head; C, dorsal aspect of right mandible; D, ventral aspect of right mandible; a, dorsal area; 6, dorsal 
impression; c, anterior condyle; d, median fossa; e, median condyle;/, posterior fossa; g, basal ridge; h, 
apical tooth; i, acute margin; j, subapical tooth; k, median tooth; I, molar; m, anterior condyle; n, me- 
dian fossa; 0, posterior condyle; p, lateral area; r, dorsal bristles of mandible; s, lateral bristles of man- 
dible; t, epistomal bristles; u, lateral angle of epistoma. (Original.) 

dibles. In fact the latter function serves to distinguish it from other 
parts. The median area is variously and sometimes greatly modified 
in insects of the same order or family, and it appears that m Coleop- 
tera generally this element is of much greater taxonomic value than 
has been usually recognized heretofore. 



THE GENUS DENDEOCTONUS. 



13 



Epistomal process. — In Dendroctonus the epistomal process serves 
to clearly distinguish the genus from other genera of the suborder to 
which it belongs. This process is usually composed of a median and 




suture .;»S . .^ ^^ 





Fig. 5. — Dendroctonus valens: Head, ventral aspect, and mouthparts. A, Labium; B, maxilla, intemo- 
lateral aspect; C, same, externo-lateral aspect; D, hypostomal region, dorsal aspect; E, head, ventral 
aspect; a, basal fossa of mentum; 6, joints; c, basal membrane; d, palpiferal area; e, stipal area; /, sub- 
galeal area; g, fossa; ft, muscle processes; k, median condyle; I, lateral fossa; m, anterior condyle; n, 
median fossa; a, posterior condyle; p, hypopharyngeal bracon; q, submental process; r, maxillary con- 
dyle; s, gular apodeme; m, oral foramen; v, occipital apodeme; ?«, postgular piece. (Original.) 



two lateral sections and is fringed anteriorly with thickly set, long 
bristles which completely cover the anterior median epistomal area. 
Hypostoma (fig. 5). — This, as here interpreted, is a ventral piece 
or area which corresponds in general function to the epistoma in 
forming a rigid ventral rim of the oral foramen for the support of the 



14 



THE SCOLYTID BEETLES. 



articulatory accessories of the labium and maxillae, and at the lateral 
angles supports the ventral articulations for the mandibles. It 
seems to the writer that this part or area, whenever sufFicientlj^ dis- 
tinct to be recognized, should be designated as the hypostoma, not 



, Epistoma 



ticidoiion 




~ " a" " 



Epislomal process ,-' 



.-Lateral section 




Gular suture' n 

Fig. 6. — Dendroctonus valens: Head, oral aspect, epistoma, etc. A, Ventral aspect of epistomal region; 
B, dorsal aspect of epistomal region; C, oral aspect of head; a, median impression and longitudinal ele- 
vations; b, median condyle; c, lateral fossa; d, posterior fossa; dd, projection over median condyle; e, 
base of epistomal bristles; /, median section of epistomal process; ff, lateral angles of epistoma; g, ante- 
rior fossa; h, anterior condyle of ventral articulation ; i , posterior condyle of ventral articulation ; j , median 
fossa of ventral articulation; k, bj'popharyngeal bracon; I, maxillary condyle; m, submental processes; 
n, ventral view of hypopharynx; o, oral foramen; p, dorsal articulation of mandible; q, ventral articu- 
lation of mandible; r, apex of hypopharynx. (Original.) 

on account of any theory of origin from a primitive segment, but 

because its location and function are similar to those of the epistoma. 

Front (figs. 3, 4). — The front is not defined by sutures or lines, but 

is fused anteriorly with the epistoma and laterally and posteriorly with 



THE GENUS DENDROCTONUS. 



15 



the epicraniuni. It is represented by a frontal area, however, which 
not only in this genus but in other scolytids presents characters of 
special value in distinguishing major and minor divisions, species, 
sexes, etc. The significance of frontal characters in this genus is 
defined in the synopses of adult and secondary sexual characters 
and shown in the figures. 

Antennae (figs. 1-6, 11-13). — The characters of the antennae are 
clearly shown in the figures. The scape, funiculus, and club are 




Fig. 7 .—Pterostichus californicus: Head, dorsal and ventral aspects, and maxillae. A, Ventral aspect; B, 
dorsal aspect; C, dorsal aspect of left maxilla; D, lateral aspect of left maxilla; E, ventral aspect of left 
maxilla; a, stipal foramen; 6, muscle processes; c, lacinial digitus; d, cardo fossa; e, basal membrane;- 
/, palpiferal bristle; g, stipal bristle; ft, median fossa of ventral articulation of mandible; i, labral bristles; 
7, clypeal bristles. (Original.) 

nearly equal in length. The scape toward the apex is clavate — cy- 
lindrical to angular. The funiculus is 5-jointed and always slightly 
longer than the club. The first joint (or pedicel of some authors) is 
of the usual form and as long or longer, rarely shorter, than the 
second. The second joint is as long as the third, fourth, and fifth 
together, or slightly shorter in some species, and the second to fifth 
increase in width toward the club, which is broad, thickened toward 
the base and compressed toward the apex, and has three or four 
distinct segments, with two or three slightly to strongly curved 



16 



THE SCOLYTID BEETLES, 



sutures. The relative concavity or convexity of the anterior face, 
as well as the relative lengths of the segments on the opposite faces,' 



Palpi 



Palpift 

Mentum- 



Submentum-<\z ~ 




~ >Maxillce 








— "'l- --2-3- -4> 



nul^S 



Gular suture 



Occipital area ^' 



Fig. 8.-Pissodesstrobi: Head, ventral aspect, and mouthparts. A , Ventral aspect of apical region of beak- 
B, ventral aspect of head; C, interno-lateral aspect of maxilla; D, externo-lateral aspect of maxilla" a 
apical tooth; 6, subapical tooth; c, lateral arm of hypostoma; d, pleurostoma; e, mandibular scrobe'/ 
hypostomal area; <7,lacinial bristles; h, antennal groove; i, joints; j, cardo fossa; k, hypostomal punctui-e 
(Origmal.) ^ 

contour of the suture, etc., are shown in figures 11 to 13, but often 
appear different in dried specimens. The articulatory attachment 



THE GENUS DENDEOCTONUS. 



17 



of the scape is in a rather deep scrobe (figs. 4, 5), situated in front of 
the eye near the base of the mandible and lateral angle of the epistoma. 




c 

Fig. 9. — Pissodes strobi: Head, dorsal aspect, and mandibles. A , Dorsal aspect of left mandible; B, ventral 
aspect of left mandible; C, dorsal aspect of head; a, apical tooth; 6, subapical tooth; c, median tooth; 
d, molar; e, median condyle;/, lateral muscle process; gf, lateral condyle; A, lateral fossa; i, extensor tendon; 
j, pharyngeal bracon; k, retractor tendon; I, ventral area; m, dorsal area; n, median condyle; o, anterior 
fossa; p, anterior section of beak; q, posterior section of beak; s, dorsal area; t, anterior condyle; u, lateral 
fossa. (Original.) 

Epicranium,. — The epicranium is not defined from the front or gena 
by sutures or lines, but the area is quite clearly indicated by the 



18 THE SCOLYTID BEETLES. 

smoother surface and by the presence of the compound eye, which is 
situated on the side of the head near the base of the antennae. The 
anterior end of the epicranial suture defines the anterior dorsal limit 
of the epicranium, designated as the vertex, while the gena is repre- 
sented by the ventral area between the eyes and the gular suture. 
The epicranial suture is more distinct in some species than m others. 

Eyes. — The eyes vary from slightly oblong oval to oblong ovate 
and are obliquely placed in the anterior angle of the epicranial area, 
just posterior to the base of the antennae. The variation in form 
within the genus and within the same species is shown in figures 1 to 
6 and 15. There are about four hundred facets, which are small and 
densely placed. 

Occiput (figs. 4, 5). — The occiput is not clearly defined, as it is in 
Pterostichus , but the posterior area of the cranium to the occipital 
foramen may be designated as the occipital region or area. 

Occipital foramen (fig. 5). — The posterior opening in the head, or 
occipital foramen, is small as compared with the oral foramen. The 
invaginated wall forms a part and posterior support to the tentorium, 
and the dorsal apodeme is continuous with the epicranial suture. 

Gula. — The gula is not represented by a space defuied by two 
longitudinal sutures, as in most Coleoptera other than the Rhyn- 
chophora. The gular apodemes are present (fig. 5, D), but the gular 
space is invaginated, so that there is but a single gular suture. 

Pregula.— In Dendroctonus there is a small sclerite immediately 
anterior to the gular suture (figs. 5, 6) which is distinctly separated 
from the gula and gense by an invaginated apodeme, laterally from 
the pregena by an evident external line, and anteriorly from the 
hypostoma by a ridge wliich defines the anterior margin. In the 
rostrate beetles this is extended with the pregena and forms a more 
or less distinct gular space of the rostrum to a similar anterior space 
which supports the so-called gular peduncle, or submentum. There- 
fore it appears that the term pregula should serve to distinguish this 
important element, which is also more or less distinctly represented 
in Coleoptera other than the Rhynchophora. (Compare figs. 6, 7, 8.) 

Gena. — The gena is not defined by lines, but it is represented by 
the ventral area between the gular suture and the epicranial area, 
as described under epicranium and gular suture. 

Pregena (figs. 5, 6). — The pregena is a distinct pleural area situ- 
ated between the base of the antennae and the pregula, bounded 
posteriorly by the genal area and anterior angle of the epicranium, 
and anteriorly by the hypostoma. 

Submentum (figs. 5, 6). — The submentum is represented by a 
bifid process or median extension of the h3^postoma, and is sup- 
ported by two stout braces rising from the large transverse rostral 
apodeme beneath the posterior angle of the pregula. 



THE GENUS DENDROCTONUS. 



19 



Lahium proper (figs. 2, 5, A). — In Dendrodonus and other rhyn- 
chophorous beetles, the mentum, palpifer, glossse, and paraglossa, 
while more or less clearly indicated, are not represented as separate 
elements of the labium. Mentum: The mentum articulates with 
the bifid submentum and completely surrounds the basal portion of 
the labrum, being subcylindrical, with the anterior ventral area 
strongly retuse. Palpifer: The palpifer is represented by the area 
between the ventral impression of the mentum and the row of 
palpiferal bristles which define the anterior limit of the mentum. 
Palpi: The labial palpi are distinct, 3-jointed, and as long as the 
mentum, or longer, with the first joint longer than the other two, or 
rarely equal. Ligula: The ligula 
is situated between the palpi, is 
thickly set with long lacinial teeth, 
and occupies the greater part of 
the dorsal area. It is evident 
that tliis hgular area represents 
the glossse and paraglossa of other 
insects, and that it is homolo- 
gous with the galea and lacinia 
of the first maxilla. 

MaxilU (figs. 2, 5, B, 0).— The 
maxillse (fig. 5, B) have the char- 
acteristic form of those of all other 
rhynchophorous beetles and are 
strikingly different from those of 
other Coleoptera. The form and 
relative proportions are shown in 
the figures. Cardo: The cardo is 
the stout basal section which 
articulates with a condyle on 
the maxillary process of the hy- 
postomal apodeme. Stipes: The 
stipes articulates mth the cardo and, while it does not appear as a 
separate piece, it is represented by the posterior ventro-lateral and 
externo-lateral part of the median section of the maxilla. Palpifer, 
galea, subgalea, and lacinia: The palpifer is fused with the stipes 
and is represented by the anterior part of the median section (fig. 5). 
The palpifer and stipes are also fused with the subgalea on the 
exto-lateral area, but on the interno-lateral area the fine separating 
the palpifer from the subgalea is distinct, as is also the suture be- 
tween the latter and the lacinia and galea, which are fused, the latter 
being represented by a narrow chitinous margin next to the palpus 
and palpifer. The lacinia is armed on the inner edge with stout 
lacinial teeth. The length of the base of the subgalea from the apex 

79980—09 3 




Fig. 10.— Dendroctonws.- Epistomata. a to I, D. val- 
ens; m to o, D. simplex; p to s, D. pseudotsugse. 
(Original.) 



20 



THE SCOLYTID BEETLES. 



to the posterior angle is usually greater than tl>e length of the palpifer 
and stipes, but is sometimes equal and rarely shorter. The ventral 




Fig. 11. — Bendroctonus: Antennae. 1, hrevicomis; 3, convexifrons $,• 3a, convexifrons cf ; Jf, frontalis <f; 5, 

arizonicus c? ; 5a, arizonicus ? ; 6, mexicanus; 7, parallelocoUis; 8, approximatus 5 . (Original.) 

1 — a, posteriorface of club when antenna is extended at right angles to head, joints 1, 2, 3; 6, anterior face, 

joints 1, 2, 3, 4; d. funiculus, joints 1, 2, 3, 4, 5; c, scape. 

chitinous area of the palpifer and stipes together is always a third 
or more longer than the combined chitinous parts of the joints of 




Fig. 12.— Dendroctonus: Antennte. 9, monticolx; 10, ponderosx: 12, simplex; IS, pseudotsugx ? ; U,picea- 
perda 5; 15, engclmanni: 17, obesus if"; 17a, ohesus $. (Original.) 

the palpi. Palpi: The palpi are 3-jointed, the joints connected j 
with each other and with the palpifer b}^ flexible membrane which ' 
allows for a certain amount of telescoping, but not adapted for free 



THE GENUS DENDROCTONUS. 



21 



lateral movements as in Pterostichus (fig. 7). The relative lengths 
of the joints vary considerably in the species of the genus, and more 
or less in the individuals of a species. The first joint is usually 
longer than the other two together, but is sometimes equal or shorter; 
the second joint is usually longer than the third, but is sometimes 
shorter. 

Mandibles (figs. 4, 14). — The mandibles are prominent, stout, 
triangular, and especially adapted for burrowing in the bark. The 
inner edges are acute, with a subapical and a median tooth toward 
the middle and a molar on the basal angle. The lateral area toward 
the base has a large impression and there is usually a less evident 
one on the dorsal area, each bearing one or two bristles. The dorsal 




Pig. 13. — Dendroctonus: Antennse. 18, rufipennis; 20, puncf.afus; SI, micans; 22, terebrans; 23, valens <?; 
23a, valens 9; S3b, valens <?; 23c, valens 9- (Original.) 

articulation with the epistoma is especially adapted to meet its 
several requirements. The peculiar trochlear mechanism of the 
articulating condyles and fossa are illustrated in figure 14; that of 
the dorsal condyle appears to be common to other rhynchophorous 
beetles, but apparently not represented in other Coleoptera, including 
those with similar bark and wood boring habits. The ventral 
articulation also appears to be different from that in other Coleop- 
tera, but to a less degree. A detailed comparative study of the 
mandibles may reveal specific characters, but as a rule such charac- 
ters are unsatisfactory from the fact that in comparisons the man- 
dible must be viewed from exactly the same position to avoid error 
in conclusions. 



22 



THE SCOLYTID BEETLES. 
THE THORAX. 



The thorax, as usual, consists of three distinct segments. The 
pro thorax freely articulates with the mesothorax, but the pleurites 
and sternites of the mesothorax and metathorax are rigidly con- 
nected. The combined length of the ventral areas of the three 
thoracic segments is shghtly greater than that of the ventral area of 
the abdominal segments, while the combined length of the dorsal 




Fig. li.—Dendroctonus valens: Mandible, a, Apical tooth; 6, subapical tooth; c, median tooth; d, 
molar tooth; e, dorsal area; /, dorsal bristles or setas; g, dorsal impression; h, transverse ridge; i, pha- 
ryngeal process; j, retractor tendon; k, lateral area; 1, lateral bristle; m, lateral impression; n, anterior 
condyle; o, basal ridge; p, median fossa; q, median condyle; r, posterior fossa; s, condyle of ventral 
articulation; i, basal foramen; «, retractor disk; !>, extensor disk; w, extensor tendon. (Original.) 

areas of the thoracic segments is about equal to that of the area of 
the abdominal segments, or slightly longer. The pronotum is as 
long as both the mesotergum and the metatergum together. The 
prosterna and mesosterna are about equal in length, and both 
together about equal to the metasterna, while the combined length 
of the thoracic pleura is shghtly greater than that of the abdominal 
pleura. The anterior dorsal margin of the pronotum and the pos- 
terior margin of the metatergum are greatly extended anteriorly 



THE GENUS DENDEOCTONUS. 



23 



beyond the ventral margin of the same segments, while the posterior 
dorsal margin of the pronotum and the anterior dorsal margin of the 
mesonotum are not produced beyond 
the corresponding sternal margin. 



Divisions op the Thoracic Segment. 

The divisions and other characters 
peculiar to the thoracic segments of a 
scolytid beetle are shown in figures 16, 
17, 18, 19, and 20. 

It will be noted that while the usual 
divisions or sclerites are quite clearly de- 
fined in the metathorax, corresponding 
divisions are less distinct in the mesotho- 
rax, and are obsolete or completely fused 
in the prothorax. The taxonomic signifi- 
cance of this wide range in the modifica- 
tion of similar parts or areas in the three 
thoracic segments of the same insect is 




Fig. 15.— Dendroctonus: Eyes. 1, brevi- 
comis; 2, barberi; 5, arizonicus; 8, ap- 
proximatus; 10, !0a, pondcrosse; 12, 
siTtiplex; 13,pseudotsugse; 14, piceaperda; 
15, 15a, engelmanni; 20, punctatus; 22, 
22a, b, c, terebrans; 23, 23a, b, c, d, e,f, 
g, h, i, j, k, I, valens. (Original.) 



combined 
any two 



realized when we compare these parts 
with corresponding segments in repre- 
sentative species of other families, sub- 
orders, and orders of insects. It will be seen that each segment has 
characters peculiar to the order or minor group to which the species 

belongs, and that in like 
^ manner the 

characters of 
or all three in the same 
insect present many fea- 
tures peculiar to the groups, 
thesuborder, family, genus, 
or species represented. 

It is also significant of 
the influence of a domi- 
nant principle or plan of 
structure and order of mod- 
ification that one or more 
thoracic segments of prac- 
tically any insect examined 
will show certain divisions 
more or less clearly defined, 
which are common to all 
other insects, and that 
when we compare the segments of different stages of insects of all 
orders, we find that a composite segment would represent a system 
of four longitudinal and four transverse divisions. The longitudinal 




B 



C 



Fig. 16.— Dendroctonus valens: Areas of pronotum. A, an- 
terior area; B, median area; ,C, posterior area; D, lateral 
area; E, dorsal area; a, anterior angle; aa, anterior margin; 
6, posterior angle; d, basal margin; e, posterior declivity; 
/, anterior section of lateral area; g, anterior section of dorsal 
area; h, median section of lateral area; i, median section of 
dorsal area; j, posterior section of lateral area; k, posterior 
section of dorsal area; I, posterior margin or vertex. 
(Original.) 



24 



THE SCOLYTID BEETLES. 




/Pre'episternal area 
■Prestcrnal area 

-Slenial area 
Epislernal area 
, Intercoxal or 
^ster-nellar area 
"Exocoxal area 

Epimeral area 



^Poslslernellar area 
Fig. 17. — Dendroctonus valens: Areas of prothorax, ventral 
aspect, a, Anterior margin ; 6, posterior margin; c, an- 
terior angle; d , posterior angle; e, anterior en to thoracic 
fold for attachment of intersegmental membrane; /, an- 
terior foramen; g, posterior foramen; h, coxal cavity. 
(Original.) 



divisions are one dorsal, two lateral, and one ventral; the transverse 
divisions are one anterior, two median, and one posterior. 

Audouin (1824) recognized the four longitudinal divisions and 
named them sternum, pleurae, and tergum. He also recognized two 
divisions of the pleura and named them episternum and epimerum, 
and four transverse divisions of the tergum, which he named pres- 

cutum, scutum, scutellum, 
and postscutellum. These 
names have been adopted 
with but slight modifica- 
tion by most of the leading 
writers on insect anatomy, 
including McLeay, 1830; 
Newport, 1839; Kolbe, 
1889; Amans, 1885; Corn- 
stock, 1902; and Voss, 
1905. The same divisions 
have been recognized by 
many other authors, who 
have designated them by 
different names. 

The first, second, third, and fourth transverse divisions of the 
ventral or sternal area were recognized by McLeay in 1830, and 
named, in order, presternum, sternum, sternellum, and poststernellum. 
All of the divisions and subdivisions mentioned as having been 
recognized by Audouin and McLeay are here recognized by the writer, 
and, in addition, the first and fourth transverse divisions of the pleura; 
thus four longitudinal and four 
transverse divisions of each 
segment in the adult insect 
have been recognized. The 
writer has also recognized the 
same or a similar system of 
division in the thoracic and 
abdominal segments of larvae 
and pupae. Wliile the taxo- 
nomic significance of the char- 
acter and modification of these 
primary and secondary divisions as represented in one or more thoracic 
segments of the same insect, or in one or more segments in insects of 
different orders and minor groups, has been recognized, there has been 
wide difference of opinion as to the origin or homology of these divisions 
and in their interpretation or definition. This has naturally resulted 
in much confusion in the adoption and application of the nomencla- 
ture proposed by different authors, and, more than anything else 



^\ Presternum 


ut"" 


yA^ /''"'^^. 


J/-\\-Pre'epistemum 


^ — ^y. -^ 


j y Sicrmim 


N. '^/Episternum 


flO^?-?^ 


X^^/^-^Epimerum 


Stemellar area'' / 


-^ 


Poststemellar piece^ 





Fig. 18. — Dendroctonus valens: Mesothorax, ventral 
aspect, a, Preepisternal process; b, transverse Lm- 
pression; c, coxal cavity. (Original.) 



THE GENUS DENDEOCTONUS. 25 

connected with the anatomical problem, it was this state of confusion 
which led the writer to make a study of the subject in order to deter- 
mine the facts and principles involved and to establish a basis for 
his future systematic and economic work on the scolytid and other 
beetles. 

There appear to be two opposing ideas regarding the origm and 
evolution of the primary and secondary elements of the insect seg- 
ment. One involves the principle of reduction of several primitive 
segments into one, on the theory that the transverse divisions rep- 
resent modifications of several primitive segments. The other involves 
the principle of complex modification from a simple undivided primi- 
tive segment into many primary and secondary divisions, on the 
theory that this has been brought about more or less independently 
through the influence of the requirements of function to meet the 
demands of peculiar life activity in different forms or species, and 
that this plan of modification has been controlled and limited by 
the fundamental plan of structure in the hexapodal type of organism, 
and by the principle of relative proportions and correlation of parts, 
so as to conform to the general modification of the entire body in 
the evolution of the species. 

The writer does not deem it advisable, in this connection, to dis- 
cuss the relative merits of these theories or any of the other theories 
advanced by different authors. He is inclined to believe that while 
it is important to utilize any good evidence relating to the probable 
origin and homology of parts, it is more important for present needs 
to deal with the facts as they are found in existing forms and stages 
and to so name and define the major and minor divisions or elements 
of the segment that they may be readily recognized and utilized in 
any comparative study of their modification and in the description 
and identification of species, genera, and larger groups. Therefore 
the writer's interpretation of the recognizable elements in the thoracic 
segments of Dendroctonus does not involve any theory of origin or 
evolution, but is based on the recognition of a dominant tendency 
in the insect segment to represent a system of four longitudinal and 
four transverse divisions, any one or all of which may or may not be 
clearly represented in one or more segments of the same insect. 

With tliis conception of a prevailing principle as a guide to the 
location of the possible primary and secondary divisions of the ana- 
tomical elements as they are indicated in any given segment, and to 
the recognition of the possible range of modification and distinction 
as manifested in the different segments of the same insect or in the 
corresponding segment of different insects, many of the difficulties 
and confusing factors relating to the proper definition of parts and 
application of names are eliminated. 



26 THE SCOLYTID BEETLES. 

In the following discussion of the thoracic segments reference is 
made to the named parts as represented or not represented, as the case 
may be, rather than to say that they are present or not present, 
because in some cases where they are not defined on the external 
surface they may be indicated by apodemes or lines on the inner 
surface of the body wall, while in other cases their position or rela- 
tive areas are indicated only by some character of surface sculpture 
or vestiture. 

In the adult Dendroctonus there is a wide range of difference in the 
representation of parts in the prothorax, mesothorax, and meta- 
thorax. In the pupa there is a similar but not so marked difference 
between the three thoracic segments, the divisions being less evident 
in the mesothorax and metathorax than in the adult, but in the 
abdominal tergites the divisions are quite plainly indicated. In the 
larvae there is not only less difference in the three thoracic segments, 
but these are only slightly different from the first to seventh abdomi- 
nal segments. In the thoracic segments the prescutal and scutel- 
lar divisions are clearly represented, with evidence of the scutal divi- 
sion on the sides. The sternal and sternellar divisions are also 
clearly represented, with evidences of the presternal and poststernellar 
divisions in the prosternum, and the latter clearly defined in the 
mesosternum and metasternum. The pleurites are also represented 
by pleural lobes. In the abdominal tergites 1 to 6 the prescutal, 
scutal, and scutellar divisions are clearly represented and the sternal, 
sternellar, and poststernellar divisions are in like manner repre- 
sented in abdominal sternites 1 to 8, inclusive. Whether or not these 
divisions or lobes are homologous with divisions occupying rela- 
tively the same positions in the pupa and adult may be a subject for 
difference of opinion, but the names here applied to what appear to 
be corresponding parts should serve as a reliable guide to their recog- 
nition and accurate definition and description in comparative studies 
and identification of species. 

Elements op the Adult Thorax. 

The primary and secondary elements as represented in the thoracic 
segments of an adult Dendroctonus beetle are sho^^'^l in the figures 
and are interpreted, named, and described as follows:'* 

THE PROTHORAX. 

In this genus, as in rhynchophorous beetles generall}^, the tergal, 
pleural, and sternal areas are fused into a continuous band. The 

(I Notum and tergum. — While the names notum and tergum are synonymous, the 
former has been applied more specifically to the dorsal division of the prothorax, 
especially in beetles, and is here utilized in that sense. The name tergum is here 
used to designate the dorsal areas of the mesothorax, metathorax, and abdomen, on 
accoimt of the use of the term tergite to designate a subdivision. 



THE GENUS DENDROCTONUS. 27 

primary and secondary divisions are not indicated by lines or sutures, 
but the corresponding areas are suggested by peculiar characters of 
sculpture and vestiture, which are of more or less taxonomic impor- 
tance, and thus may be arbitrarily indicated, as in figures 16 and 17, 
to serve as guides to the location of characters in comparative study 
and description. 

Pronotum.—T]ie pronotum is the dorsal or tergal area of the pro- 
thorax, as defined by the anterior, posterior, and lateral margins. 
There is considerable specific variation in its structure, sculpture, and 
relative proportions. It ranges from about one-fourth to about one- 
third broader than long, with about the same range of difi^erence in 
the width of the posterior and anterior areas. In some species the 
lateral margins are nearly parallel, while in others they are distinctly 
convergent and constricted anteriorly. The anterior margin is 
broadly sinuate, while the vertex or dorsal margin of the posterior 
declivity is bisinuate. The anterior area is broadly transversely 
impressed, except in the females of some species, where the median 
section of the area is transversely elevated. The posterior declivity, 
which perhaps represents the postscutellum, is more distinctly 
exposed and defined in this genus than it is in allied genera and is 
therefore an important character of generic distinction. The pleural 
and sternal areas are indicated in figure 17. 

Episternal area. — The episternal area is limited dorsally by the 
lateral margins of the notum, ventrally by the smooth exocoxal 
area, posteriorly by the epimeral area, and anteriorly by a preepis- 
ternal impression or in some species by a ridge. The sculpture of 
this area is quite variable and in some species furnishes characters of 
considerable value. 

Epimeral area. — The epimeral area is represented by a flattened, 
smooth space situated between the roughened episternal area and the 
posterior margin of the prothorax and between the coxae and the 
basal angle of the notal area. 

Sternal area. — The entire sternal area between the anterior and 
basal margins is largely occupied by the coxal cavities, which are 
separated by the very narrow intercoxal or sternellar piece. The 
elevated anterior margin evidently represents the presternum, while 
the sternum is quite clearly defined by a nearly vertical flat to con- 
cave space between the presternum and the coxae, the lateral limit 
being indicated by the smooth, shiny exocoxal area between the coxge 
and the episternal area. The sternum proper is quite variable, rang- 
ing from concave, smooth, and shiny, without trace of a median lon- 
gitudinal line to nearly flat, roughened, or with a median subcari- 
nate line; but apparently none of these minor characters is suffi- 
ciently constant, even within the same species, to be of much taxo- 
nomic value. 



28 



THE SCOLYTID BEETLES. 



Poststernellar area — The poststernellar area is well defined and 
serves to completely inclose the coxal cavity. It has been referred 
to as the epimeron, but since the epimeral is so clearly defined as a 
lateral area, it appears to more correctly represent the poststernellum, 
which in the mesosternum and metasternum is not evident, or is 
modified to accommodate the large coxal cavities. 



THE MESOTHORAX. 



The mesothorax (figs. 18, 19) is short and partiall}'' hidden 
from view by the prothorax, which covers the anterior third of the 
sterna, pleurites, and tergites, while the base of the elytra covers 




Tergites 



Fig. 19. — Dendroctonus valens: Mesotergum and mesopleurum. a, Lateral arm of prephragma and 
preseutum; b, wing root or connecting membrane; c, basal margin of elytra; d, radial plate; e, flexor 
plate; /, median plate; g, scapular plate; h, subscapular plate; i, sutural or anal margin; j, preepistemal 
process; k, clavicle condyle; I, coracoidal condyle; m, scutellum; n, lateral margin of elytra; o, arm of 
preepistemal process ; strife 2-10; interspaces 2-11; p, lateral arm of postphragma; g, pleural clavlculus. 
(Original.) 

the posterior third and dorsal area of the tergum, leaving but a 
small triangular area exposed between the thorax and inner angles 
of the elytra. Upon removing the prothorax and elytra this segment 
is found to represent most of the primary and secondary divisions 
of the normal segment. 

MESOTERGUM. 

The mesotergum (fig, 19) is rectangular in form, with the pre- 
seutum occupying two-thirds of the area, while the scutum, scutellum, 
and postscutellum are less clearly defined or rudimentary. 

Preseutum. — It is evident that the large subtriangular dorsal sec- 
tion represents the preseutum, as indicated by the evident prescutal 
lobe and prescutal process, attachment of wing accessories, etc. 



THE GENUS DENDEOCTONUS. 29 

PrepTiragma.— The prephragma is strongly flexed beneath the 
median area, but the anterior arms, in conjunction with the anterior 
angles of the prescutum, are prominent and strongly produced. 

Scutum. — The oblique, impressed, triangular section situated 
beneath the posterior margin of the prescutum evidently represents 
the scutum. ^ 

Scutellum. — The hornlike process situated at the apical angle of 
the prescutum apparently belongs to the scutellum and corresponds, 
perhaps, to the structure which forms the scutellar groove of the 
metathorax. This, with the posterior lateral section, represents the 
median and lateral sections of the scutellum. 

Posts cutellum. — The postscutellum apparently is not represented 
by an external piece, but by an evaginated fold beneath the scutellum 
and by the lateral arms of the postphragma (fig. 19). 

Postphragma. — The postphragma is represented by the posterior 
invagination or fold beneath the scutellum and by the lateral arms, 
as evidenced by the attachment of the scutal muscle to the arm and 
the connection of the arm with the invaginated phragma. 

MESOPLEURA. 

The episternum, preepisternum, epimerum, and postepimerum are 
all represented and together occupy an area greater than that of the 
sternal and slightly greater than that of the tergal. 

Preepisternum. — The preepisternum occupies the area in front of 
a transverse impression and is quite prominent. The anterior mar- 
gin bears the preepisternal process (fig. 19, j), which is common to 
most of the Rhynchophora, and is more or less represented in certain 
other Coleoptera, such as Carabidse, Cicindelidae, Scarabasidse, etc., 
but it is surprising to find that it is not represented in Ips ( Tomicus) 
and allied genera of the Scolytidse. It is peculiar to the mesothorax 
and is progressively modified in character and function through 
various groups of Coleoptera from an obscure accessory of the clavi- 
cle disk to a prominent process. In Dendroctonus it is fused with 
the anterior margin of the preepisternum and has an arm extending 
to the dorsal angle to form an accessory to the wing process. It 
Ijears a set of powerful muscles, which are attached to the inner wall 
of the postscutellar and scutellar areas of the pronotum, thus form- 
ing a powerful muscle connection between the two segments. The 
mesothoracic spiracle is situated in the angle between this process 
and the sternal area and is covered by the epicranial area of the 
prothorax. 

Clavicle disJc. — The slender plate situated in front of the arm of 
the preepisternal process appears to correspond with the clavicle 
disk of the metapleurum. It is connected by a ligament to the head 
of the elytra, and its muscle is attached to the rudimentary ante- 
coxal piece. Both the preepisternal process and the clavicle plate, 



30 THE SCOLYTID BEETLES. 

as here defined, are probably modifications of the episternal paraptera 
of Audouin. 

Episternum. — ^The episternum is the clearly defined, large, exposed 
triangular pleurite situated between the posterior margin of the 
preepisternum and the epimerum, with the dorsal angle dilated, pro- 
duced, and flexed ventrally at the apex and with the apex of the 
epimerum and preepisternal process forming the pleural claviculus 
with its clavicle and coracoidal condyle for the articulation of the 
elytra. The posterior angle of the episternum is acute, and the 
suture between it and the sternum is obliquel}^ sinuate. The epi- 
sternal impression is clearly defined by the elevated posterior margin 
of the preepisternum, and is usually covered by the posterior margin 
of the prothorax. 

Epimerum. — The epimerum is exposed at its posterior ventral 
half and has its p^Dsterior margin fused with the metasternum and 
metepisternum and the produced anterior dorsal angle with its 
coracoidal condyle is covered by the episternum. 

Postepimerum. — The postepimerum is represented by a small de- 
clivous area beneath the posterior dorsal angle, where it covers the 
metathoracic spiracle. 

MESOSTERNA. 

Presternum (fig. 18). — The presternum is quite clearly represented 
by the narrow, slightly elevated, anterior margin joined directly with 
the anterior ventral angle of the preepisternum. 

Sternum.. — The sternum is short, flat, and subdeclivous, with the 
posterior angle (exocoxal piece) extending around the coxal cavity to 
its junction with the anterior angle of the metasternum. 

Sternellar area. — The sternellar area appears to be represented by 
the elevated and rather broad intercoxal piece, whfle the poststernel- 
lum is apparently represented by a poststernellar piece. 

THE METATHORAX. 
METATERGTJM. 

From a systematic and taxonomic point of view, the metatergum 
is by far the most important and interesting part of the thorax of 
beetles. We find in it not only evidence of the four transverse 
divisions, but examples of the possible extremes in modification to 
meet the requirements of wing articulation and wing motion. 

Transverse sutures. — By a comparison of the metatergum of repre- 
sentatives of different orders of insects and of the larvse, pupse, and 
adults of some insects, as in Dendroctonus, we find that the prevailmg 
principle of division involves three transverse external lines, sutures, 
or impressions, and three corresponding entothoracic ridges, apo- 
demes, or invaginated ectoderm, which define more or less clearly 
the four divisions, viz, prescutum, scutum, scutellum, and postscu- 



THE GENUS DENDEOCTONUS. 



31 



tellum. We also find that these transverse sutures are subject to 
o-reat variation in position, contour, character of surface, manifesta- 




FiG. 20.— Dendroctonus valens: Metatergum and metapleumm. B, Anterior aspect of prephragma and 
prescutum; D, metatergum and metapleurum; E, posterior aspect of postptiragma and postscutel- 
lum; a, lateral arm of prephragma; &, dorsal band; c, scapular plate; d, subscapular plate; e, flexor 
plate; /, radial plate; g, medial plate; h, lateral arm of postphragma; i-i, metatergal costse; j, anterior 
disk; fe, scutellar groove; I, posterior ridge; o, exposed triangular plate of postepimeron; p, pleural 
hinge; pc, pleural clavicula; q, posterior ventral angle of epistemum; r, anterior ventral angle of 
epistemum; s, pleural suture; t, postscutellar process; u, clavicle condyle; v, coracoidal condyle; 
w, attachment and articulation of scapular plate; x, dorsal area of postscu tellum; y, pleural disk; 
2, elevated acute margin; zl, membranous area; z5, villotis area. AVing veins: C, costa; Sc,sub- 
costa; R, radius; M, media; Cu 1, cubitus 1; Cu 2, cubitus 2; A, anai. (Original.) 

tions, etc., to correspond with the enormous range of modification to 
which one or all four of the transverse divisions are subject. These 
external evidences of separation of parts are here referred to as sutures, 



32 THE SCOLYTID BEETLES. 

although in some cases they may be but faintly or obscurely indicated. 
The lateral and median sections of the anterior suture separatmg the 
prescutum from the scutum have a tendency to curve posteriorly, 
and especially the median section, which has a very strong tendency 
in this direction, and is often manifested to such an extent as to 
separate the scutum into two lateral sections, as shown in figure 20. 
The median suture has a reverse tendency, the lateral sections 
extending anteriorly, the submedian section posteriorly, and the 
median strongly anteriorly. Thus we often find, as shown in figures 
20 and 21, that the two sutures overlap and form external longitu- 
dinal ridges and internal oblique apodemes, with an external median 
longitudinal impressed area. It appears that the dorsal groove may 
belong to either the scutellum or prescutum or represent a combina- 
tion of the two, but for the sake of uniformity in its definition the 
name scutellar groove is here adopted. The posterior suture is 
usually distinct in the metatergum, especiall}^ in that of Coleoptera, 
and is much less subject to striking modifications in contour, etc., 
than the two preceding. Thus, it clearly defines the postscutellum, 
as in figure 20. 

Transverse divisions. — The writer's interpretations of the modifi- 
cations and position of the four transverse divisions as represented 
in Dendroctonus are demonstrated in figure 20. 

Prescutum (fig. 20). — The area designated as the prescutum is that 
involving the attachments of the principal sternotergal muscles and 
the anterior lateral process for the attachment and articulation of 
the scapular plate. The anterior limit is defined by the prephragma, 
its posterior limits by the anterior suture and apodeme and the 
posterior limit of the prescutal lobes, and laterally by the anterior 
angle or limit of the lateral emargination. The most important 
features are the prominent prescutal lobes and anterior lobes for the 
attachment of the depressor muscles, the prescutal disk for the small 
muscle connecting it with the pleural clavicula, and the triangular 
prescutal process for the attachment and articulation of the scapular 
plate. (See, also, figure 21 for the entotergal characters and anterior 
apodeme.) 

Prephragma (fig. 20, B). — The prephragma is the median section of 
the anterior vertical area of the prescutum. Its dorsal and lateral 
limits are defined by the line of attachment of the intersegmental 
membrane. The lateral arms in conjunction with the anterior process 
of the anterior lobe of the presternum are greatly extended ventrall}^ 

Scutum. — The scutum is represented by the large scutal lobes situ- 
ated each side of the scutellar groove. These lobes are for the ante- 
rior attachment of the large scutal muscles with the posterior attach- 
ment to the lateral arms of the postphragma. The lateral margm of 
the scutum is defined by the lateral emargination and elevated 
scutellar ridge which terminates in the scutellar process, and poste- 



THE GENUS DENDEOCTONUS. 



33 



riorly by the oblique sinuated line of the median suture and the 
lateral section of the posterior ridge. 

Scutellum. — The scutellum is represented externally by the area 
posterior to the oblique line of the median suture and by the sublat- 
eral and lateral ridge which terminates in the scutellar process, as is 
indicated by the character of the entotergum and by comparison 
with the less modified scutellar division in other insects. The 
median longitudinal groove appears to represent the median produced 
section of the scutellum rather than a part of the scutum or prescu- 
tum, as indicated by the character of the entotergum and the widely- 
separated apodemes of the median suture which extend to and join 



Anterior apodeme 
/ \ J PrcphrarjmCC 



Membranous area 
? ^y^ ^Anterior lohej 

icidal disc 

■cscutal lobe- 



Prosciital pj'ocfW 



Pi^csculnm 



Scutumi-^ 




alcral emarfjination 
7':>cu,ldla)- prnccus 

Sciilal lobe 
^Median apodeme, 

Poslsriilellar process 



'^'-Mciaplcural hook 



Scutellum 



Poslplirmjina 



Fig. 21. — Dendroctonus valens: Metatergum, inner aspect, a, Lateral arm of prephragma; 6, dorsal 
band; c, scapular articulation; d, posterior angle of prescutum; e, posterior arm of postphragma; /, pos- 
terior apodeme; g, anterior disk; h, posterior margin of scutellum; x, ventral wall of postscutellum. 
(Original.) 

with the anterior apodeme (fig. 21), thus defining a large median 
triangular area which is evidently scutellar. 

Postscutellum. — The postscutellum is the exposed dorsal and lateral 
area between the clearly defined posterior suture and the line of 
attachment of the first abdominal tergite. It is firmly connected 
with the scutellum toward each side at a point near the base of the 
oblique apodeme of the middle suture, otherwise the connection is 
membranous. The anterior angles support the metapleural hooks 
(fig. 20, 2?, It), which fit into a fold in the dorsal margin of the postepi- 
merum (fig. 20, p). 

Postphragma. — The postphragma is an invagination of the pos- 
terior section of the postscutellum and, with the produced posterior 
disks and arms, serves as important posterior attachments for the 
longitudinal, tergal, and oblique scutal muscles. 



34 THE SCOLYTID BEETLES. 



METAPLEURA. 



The metapleiirum is well developed and distinctly represented by 
the two longitudinal sclerites, episternum and epimerum (figs. 3, 
20), mth their anterior dorsal angles greatly produced to form the 
pleural clavicula with its clavicle and coracoidal processes. 

Pleural suture and apodeme. — The pleural suture marks the line of 
division between the episternum and epimerum, and extends from the 
dorsal angle of the coxse to the apex of the pleural clavicula and 
between the clavicle and coracoidal process. That tliis is the true 
pleural suture is indicated by the corresponding prominent pleural 
apodeme. It is also quite evident that the episternum corresponds 
to the hypopleurites and the epimerum to the epipleurites of the 
abdominal segments (figs. 3, 22). 

Episternum. — The episternum is exposed when the elytra are 
closed (fig. 2). The suture between it and the sternum is distinct 
and nearly straight, with the anterior end curved toward the coxa. 
The posterior ventral angle is oblique and joins the posterior dorsal 
angle of the sternum; from here the posterior margin is oblique to 
its acute junction with the epimerum and the dorsal angle of the 
coxal cavity. From here the dorsal margin is acutely elevated to 
fit into the anterior lateral groove of the elytron, and is nearly parallel 
with the ventral margin to the preepisternum. 

Preepisternum. — The preepisternum appears to be represented by 
the narrow declivous anterior section of the episternum connected 
with the anterior basal area of the pleural clavicula and is apparently 
involved in the formation of the clavicle process. The clavicle disk 
evidently represents one or both of the paraptera of certain other 
insects and belongs to the prepleura. It is situated immediately 
anterior to the preepisternum. It is large, prominent, and partially 
exposed, and is connected by a chitinous tendon to the side of the 
clavicle process. This disk supports the set of large clavicular or 
sterno-pleural muscles, the opposite ends of which are attached to the 
sternum and sternellum. 

Epimerum. — The epimerum is situated between the pleural suture 
and the tergum. With the exception of the extreme posterior ven- 
tral angle of the postepimerum it is covered by the elytra. The 
anterior dorsal angle is strongly produced to form the coracoid pro- 
cess. The ventral area is chitinous and is joined to the episternum 
by the pleural suture, while the dorsal area is submembranous tc 
membranous to its junction with the base of the wing membrane. 

Postepimerum. — The posterior ventral angle and posterior lateral 
section represent the postepimerum, as is indicated by its articula- 
tory junction with the poststerneUum (fig. 20, p). The posterior 



THE GENUS DENDEOCTONUS. 



35 



ventral angle or ventral section of the postepimerum, which might 
be mistaken for a postepisternum, is indicated by the pleural apodeme 
and pleural suture which here join the dorsal angle of the coxa. It 
is not impossible, however, that this plate may represent a combined 
postepisternum and postepimerum. 



METASTERNA. 



The metastema (figs. 2, 3) form a broad rectangular plate separated 
into two lateral sections by a median longitudinal line. The pre- 
sternum and poststernellum are not represented by external parts. 




\^zs^\ .Spiracle I 
IV-i -\-Epipleurite 1 

( if5>--1" Spiracla2 
_^\ "J Epipleurite 2 

V ^ ~Z.T~ Epipleurite 3 

// /=i - - A- Spirach 4 

-Epipleurite 4 

- » Spiracle 5 

^Epipleurite 5 

■ Spiracle 6 
'Epipleurite 6 

"Spiracle 7 
Epipleurite 7 



SlriJulating scraper 



Fig. 22. — Dendroctonus valens: Abdominal tergites. a, Anterior arm of epipleurite 7; b, posterior arm of 
epipleurite 7; c, membranous lobes. (Original.) 

_ Sternum. — The sternum is evidently represented by the large con- 
tinuous area between the mesocoxse and the small median plate and 
the shghtly acclivous area anterior to the metacoxa. 

Sternellum. — The sternellum is evidently represented by the pos- 
terior median plate and the posterior acclivous areas (fig. 2, a). The 
relation of the latter to the sternellar area is indicated by the attach- 
ment of the posterior pair of clavicular muscles. 



THE ABDOMEN. 

The abdominal terga, pleura, and sterna, and their relative pro- 
portions, are shown in figures 1-3 and 22-25. 

79980—09 4 



36 



THE SCOLYTID BEETLES. 



Abdominal Tergites. 

The eight abdominal tergites are normally covered by the elytra. 
The apparent difference in the relative proportions, as indicated by 
figures 3 and 22, is due to the flexible intersegmental membrane and 
the fact that figure 20 is from a balsam mount. The integument 
of 1 to 6, inclusive, is more or less membranous, while that of 7 and 8, 
with the exception of the finely sulcate membranous lobes of 7, is 
chitinous. In the female, 8 is covered by 7, and forms the so-called 




Fig. 23.— Dendroct.onus valens: Male, abdominal tergites 7 and 8. ^,Tergite8(pygidium), ventral; 5, same, 
dorsal; C, same, ventral, showing position of stemite 8; D, tergite 7 (propygidium), dorsal; E, same, 
ventral; a, anterior arm of epipleurite 7; b, posterior arm of epipleurite 7; c, membranous lobe; d, 
epipleural process; e, hypopleural arm of sterniteS; /, pleural opening; g, anal space; h, ventral fold; i, 
rudimentary spiracle 8; j, epipleural disk; fc, apical spine; I, stridulating process; in-, sensory bristles; 
n, enlarged section of lateral area; o, enlarged section of membranous lobe; p, stridulating scraper. 
(Original.) 

"pygidium," while in the male 8 is distinct and together with 7 forms 
the so-called "divided pygidium." 

Pygal tergites of the male. — The pygal tergites of the male are shown 
in figure 23, A, B, C, D, E. Tergite 7 is the propygidium and as a 
bearer of generic and sexual characters is the most important of the 
entire series. In the male the posterior margins between the epi- 
pleurites converge toward the apex, which is produced into a bifid 
process and supports the stridulating scrapers. The posterior area 
of the tergite is thickened and strengthened to meet the requirements 
of stridulation. There is a broad ventral fold (fig. 23 E, 7i) of the 
integument which may serve a sinular function to that of a sounding 
board. 



THE GENUS DENDEOCTONUS. 



37 



The median area is triangular in form and covered with bristles 
and hairs rising from variously formed bases. On its face and some- 
times on the posterior area there are a few irregularly arranged 
truncate tubercles (J), m), each bearing a short, stiff bristle. These 
may possibly function as sense organs. 

The membranous lobes are subovate, finely sulcate, and thickly 
clothed with reclining microscopic spines (o). The exact function 
of these lobes is not known to the writer. Tergites 4, 5, and 6 have 
similar lobes. The other dorsal and ventral characters are made 
sufficiently clear in D and E. 















Fig. 24. — Dendroctonus valens: Female, abdominal tergites 7 and 8. a, Anterior arm of epipleurite 7; 
6, membranous area surrounding spiracle; c, membranous lobe; d, epipleural process; e, hypopleural 
processor stemiteS;/, pleural opening; g, anal space; h, ventral fold; i, rudimentary spiracle; j, epipleural 
disk; k, median membranous connection of the lateral sections of stemite 8. (Original.) 

Tergite 8 (A, B) is the pygidium. This, in the male, is always 
larger and more exposed beyond the margin of tergite 7 than in the 
female. The relative proportions, as compared with 7, and the dorsal 
and ventral characters are clearly shown in A and B. The lateral 
arms serve as attachments for pleural muscles and articulating mem- 
brane and ligaments. In C the abdominal sternite is added to show 
its relative position and proportions. 

Pygal tergites of tJie female. — The pygal tergites of the female are 
shown in figure 24, A, B, C, D, E, F. 

Tergite 7 (propygidium) is much more simple in structural details 
in the female than in the male, and tergite 8 (pygidium) is also more 
simple and shorter, being almost or entirely covered by 7 when in 
normal position. 

The characters of sternite 8 are shown in D, the most important 
of which is the median membranous area. 



38 



THE SCOLYTID BEETLES. 



Abdominal Pleurites. 

At the lateral ends of the abdominal tergites and sternites there 
are well-defined areas (figs. 3, 22, 25), which may be designated as 
pleurites. Those situated immediately above the pleural suture and 
bearing the spiracles may be referred to as e-pipleurites , while those 
of the sternites which are immediately below the -pleural suture may 
be designated as liypopleurites; both series are well defined in 
Dendrodonus. 

In a lateral view (fig. 3) seven epipleurites and five liypopleurites 
are clearly defined, with the eighth epipleurite and the second hypo- 
pleurite indicated, and when the abdomen is removed both the first 
and second of the latter series are quite distinct. 



UyfO'phv.Tites 



Co.ral cavittf 



Intercoxal 




Fig. 25. — Dendrodonus valens: Abdominal sternites, ventral and lateral aspects. A, Lateral; S, ventral; 
a, stemite ^, faintly indicated; b, sternite ^, faintly indicated. (Original.) 

Abdominal Sternites. 

The characters of the abdominal sternites are shown in figures 2, 
3, and 25. There are eight, corresponding to the eight tergites, but 
only five are exposed, viz, 3 to 7, which are densely chitinized and 
clearly defined by four sutures. 

Sternites 1 and 2 and the anterior portion of 3 are covered and 
obscured by the large metacoxa and form the posterior wall of the 
coxal cavity. They are fused, but the sutures are indicated by faint 
lines. Sternite 3 (first ventral segment of some writers) has the 
median area produced anteriorly, and with faintly indicated median 
portions of sternites 2 and 1 it forms the intercoxal process, the apex 
of which forms a junction with the metasternellar piece. The 
anterior exposed margin forms the posterior margin of the coxal 
cavity, but the junction with the preceding segment is but faintly 



THE GENUS DENDEOCTONUS. 39 

indicated in the wall of the coxal cavitj. Sternites 4, 5, and 6 are 
nearly of equal length, while sternite 7 is nearly as long as 5 and 6 
together, with the posterior margin broadly curved and forming 
the apex of the exposed series. Sternite 8 (figs. 23, C, and 24, D) is 
entirely covered by 7, and is represented in the male by a narrow 
cliitinous rim below the anal opening, while in the female the median 
section of this sternite is membranous. 

Suture 3, between sternites 3 and 4, is the first visible suture, and 
is rigid and straight throughout, while sutures 4, 5, and 6 are slightly 
flexible and are strongly recurved toward and between the hypo- 
pleurites, thus presenting an important generic character. 

Spiracles. 

There are 9 well-developed spiracles, 2 thoracic and 7 abdominal, 
with the rudiments of a tenth. The large mesothoracic spiracle is 
located in the intersegmental membrane between the prothorax and 
mesothorax, and lies between the preepisternal process and the 
anterior ventral angle of the preepisternum. It overlaps the anterior 
margin of the latter for half its length, but is completely covered and 
obscured by the epimeral area of the prothorax. The metathoracic 
spiracle is situated in the intersegmental membrane between the 
metathorax and mesothorax, and concealed beneath the dorsal 
margin of the mesepimerum. The abdominal spiracles 1-7 are con- 
spicuous; 1 is very large and situated in the epipleurite just posterior 
to the pleural hook of the metapostscutellum; 2-7 are situated in 
their respective epipleurites, as shown in figures 3, 22, 23, and 24, while 
8 is evident, but; rudimentary. 

THE LEGS. 

The structures and characters of the parts of the legs are so well 
illustrated in the figures (figs. 3, 26-29) that they do not require 
detailed description. The procoxse and mesocoxse are large, globose, 
and prominent, the former subcontiguous and the latter widely 
separated by the elevated intercoxal or sternellar piece, while the 
metacoxae are oblong, oval, and separated by the process of the 
third abdominal sternite. There is no striking difference in the 
anterior, middle, and posterior trochanters, femora, tibiae, and 
tarsi. The trochanters are small; the femora are moderately stout, 
and each is as long as its tibia, which is dilated toward the apex and 
armed on its outer lateral margin with stout teeth. The anterior 
dorsal area has a distinct tarsal groove for the retractile tarsus, as 
shown in figures 26 to 29. The tarsi are each more than half as long 
as their tibiae, and have five joints; joint 1 is always longer than 2, 
but never as long as 2 and 3 together; 3 is distinctly bilobed, the 



40 



THE SCOLYTID BEETLES. 



lobes slightly longer than joint 4; joint 5 from tip of lobes of 3 is 
never as long as the others (1 to 3) together,, but sometimes shorter 
than 1 and rarely equal to 2 and 3. In the males this joint is often 
longer than in the female. 

The trochlear articulation of the tibia with the tarsus is shown in 

figure 26, in which the 
other more important char- 
acters are shown and 
named. 

THE WINGS. 

Notwithstanding the vast 
amount of pubhshed data 
on the wings of insects,there 
is yet much difference of 
opinion among the leading 
authors in regard to some 
of the details, and much 
confusion exists, due to 
different interpretations of 
the homologies of the ele- 
ments of the wing and its 
articulatory accessories. A 
detailed investigation has 
been made of the basal 
areas of the wings of rep- 
resentatives of different 
orders of insects, to deter- 
mine facts relating to the 
fundamental plan of devel- 
opment and modification, 
and the system of organi- 
zation of the elements as 
represented in Dendroc- 
tonus. 

Nomenclature. — While 
the more generally accepted nomenclature has been adopted, it has 
seemed necessary to revise and more defhiitely define the appHcation 
of some of the old names and to introduce some new ones to desig- 
nate the elements heretofore obscurely defined. 

Attachments and articulations.— There are certain elements in the 
structure, mechanism, attachments, and articulation common to 
the wings of all insects, but within defined Umits and according to a 
definite'' system modifications, additions, and reductions occur. 
Therefore the presence or absence of a given element should be 
detected in any form of wing. 




Fig. 2G.—Dendroctonus valens: Tibia and tarsus, articulation, 
etc. A, Left tibia, ventral view; B, left tibia, dorsal view; 
C, left femora, ental view; D, left tibia, dextral view of base; 
E, left tibia, sinistral view of base; F, tarsus; a, apical tooth; 
6, subapical tooth; c, tarsal groove; d, subapieal ridge; e, 
marginal teeth; /, sinistral margin; g, dextra! margin; h, 
median fossa; z, lateral condyle of tibia; J, anterior fossa; fc, 
lateral condyle of femur; I, lateral fossa of femur; ?n, median 
condyle of femur; m, attachment of extensor muscle; o, basal 
foramen; p, attachment of flexor muscle; q, basal groove; r, 
tibial groove. (Original.) 



THE GENUS DENDROCTONUS. 



41 



Primary elements of the Dendroctonus wing. — The primary wing 
elements and their relations to each other as represented in Den- 
droctonus may be summarized as follows: TJie structure consists of a 
dorsal and ventral membrane or chitinous integument. The primary 
trachex are costal, subcostal, radial, cubital, and anal. The primary 
veins are costa, subcosta, radius, media, cubitus, and anal. The 
wing plates are scapular, subscapular, flexor, subflexor, radial, and 
medial. The wings are attached by membrane to the tergum and 




VENTRAL 
Fig. 21 .—Dendroctonus: Left tibise, dorsal and ventral aspects. 1, brevicomis; 2, barbcri; 3, convcxifrons, 
4, frontalis; 5, arizonicus; 6, mexicanus; 7, parallelocolUs; 8, approximatus; 9, monticolx; 10, ponderosse. 
(Original.) 

pleurum, and by ligaments and tendons to tergal and pleural proc- 
esses and muscle disks. The pleural processes are the clavicle and 
coracoid processes, which together form the pleural clavicula. The 
tergal processes are the prescutal, scutellar, and postscutellar. 

Elements of wing motion. — The elements of wing motion are the 
clavicular disk and clavicular muscle, pleural disk and pleural muscle, 
flexor and flexor muscles, prescutal disk and muscles, anterior pres- 
cutal lobe and anterior sternotergal muscles, posterior prescutal lobe 
and posterior sternotergal muscles, scutal lobe and scutal muscles, 



42 



THE SCOLYTID BEETLES. 



scutellar lobe, postscutellar processes, prepliragma, and postphragma; 
also the pleural clavicula, clavicle, coracoidal, tergal, prescutal, scu- 
tellar, and postscutellar processes, and connecting ligaments. 

Mesothoracic and Metathoracic Wings. 

While there is a wide difference in the appearance and structural 
details of the elytra and the hind wings of beetles, they are evidently 
homologous and differ only in their modification in structure and 
function. 




VENTRAL 
Fig. 28.~Dendroctonus: Left tibiae, dorsal and ventral aspects. 12, simplex; IS, pseudotsugas; I4, piceaperda; 
15, engelmanni; 16, borealis; 17, obesus; 18, rufipennis. (Original.) 

Structure. — The wing consists of two layers of integument, the dor- 
sal evidently rising from the tergites and the ventral from the epi- 
pleurum (epimerum) . Between these layers there is a system of tra- 
cheation and circulation. The integument of the mesothoracic wing 
or elytron is chitinous throughout, while that of the metathoracic 
or hind wing, with the exception of the veins and basal pieces, is 
membranous. 

TracTieation. — The same system of primary tracheae prevails in 
both the elytra and the hind wings. In the former it corresponds in 



THE GENUS DENDEOCTONUS. 



43 



general position to that of the primary veins in the latter, thus con- 
forming to the prevailing system in fully developed wings of all 
insects. The primary tracheae are costal, subcostal, radial, medial, 
cubital, and anal. In the elytron these occupy the marginal and 
alternating interspaces between the longitudinal striae or rows of 




DORSAL 




VENTRAL 

Fig. 29. — Dendroctonus: Left tibise, dorsal and ventral aspects. ^0, punitatus; 21, micans; 22a, 22b, 

terebrans; 23, valens. (Original.) 

punctures, while in the hind wing they follow approximately the 
primary veins. 

Metathoracic or Hind Wings. 

The hind or metathoracic wings (figs. 1, 20, and 30) are a third 
longer than the elytra or mesothoracic wings, under which they are 
folded when at rest. In consequence the veins toward the middle 
of the wings are flexible and adapted to the requirements of folding 
and unfolding. 



44 



THE SCOLYTID BEETLES. 



Basal area (fig. 30). — The basal area is that in which the basal 
plates and head, of the veins occur. In this area there are four 
axillary plates, which are more or less common to insect wings in 
general. These appear to belong to the wing rather than to the 
body structures, and are here designated as scapular, subscapular, 
flexor, radial, and medial plates. They are discussed in greater 
detail under wing articulation. 

Veins. — The six primary veins represented in the hind wing are 
costa, subcosta, radius 1 and 2, media 1 and 2, cubitus 1 and 2, and 
anal, which last is rudimentary. 




Fig. 30. — Dendroctonus valens: Diagram of basal area of hind wing, a, Scapular plate; 6, subscapular 
plate; c, flexor plate; d, radial plate; e, medial plate; /, tendon attachment; ;/, articulation; 7i, subcostal 
head; i, costal head; J, costal tongue; k, radial head: 1, medial head; m, cubital head; n, anal head; 
0, scapular condyle; p, scapular arm; q, scapular base; r, articulatory margin; s, connection of costa 
with subcosta; J, flexor arm; m, connection ofmedial plate with flexor; C, costa; Sc, subcosta; -K, radius; 
JV/,media; Cm, .Z, 5, cubitus 1 and 2; JL, anal. (Original.) 

Costa. — In Dendroctonus and most beetles the vein which corre- 
sponds to the costal trachea is confined to the basal area, and forms 
the anterior basal angle. The head is produced beyond the head of 
the subcosta, with which it is fused to form the articulating fossa. 
The produced head of this vein appears to function as an important 
accessory of the clavicle muscles in extending and depressing the 
wing, since it is connected with the clavicle condyle and clavicle disk 
by tendons. The vein proper extends outward but a short distance 
to its submembranous connection with the subcosta, and from that 
point the costal margin is occupied by it and the subcosta to its 



THE GENUS DENDEOCTONTJS. 



45 



junction with the radius, which appears to form the broad chitinous 
costal area to near the apex, with branch 2 as a distinct rein. 

Suhcosta. — The subcosta is the principal vein of the wing. It gives 
rigidity to the base, and with the head of the costa forms the articu- 




lating head. It is broadest where it is joined by the costa, and is 
strongly narrowed to its junction with the radius. 

Radius. — The radius is an important vein in givmg additional rigid- 
ity to the median, distal, and costal areas and in forming the folding 



46 



THE SCOLYTID BEETLES. 



hinge. It arises through the radial plate from the subscapula, and 
joins and fuses with the head and posterior edge of the subcosta to 




Fig. 32. — Dendroctonus valens: Basal process of right elytron. A, Ventral; B, latero-sinistral; C, dorsal; 
D, latero-dextral; a, foramen of articulatory head; 6, subcostal head; c, costal head; d, costal fold; e, 
costal margin; /, callus; g, basal foramen; h, anal fold; i, muscle disk; j, radial plate; k, posterior ven- 
tral area; I, anal margin; m, posterior dorsal area; Sc, subcostal area. (Original.) 




Fig. 33.— Dendroctonus valens: Declivital section of elytra, a, Ventral shoulder; 6, dorsal shoulder; c, 
sutural tongue; d, ventral lip; e, dorsal lip;/, sutural groove; g, rugose surface of apical wing-lock; ?i, 
stridulating rasp; i, apical wing-lock; j, suture; A, anal canal; Cu. cubital canal; M, medial canal; E 
radial canal; Sc, subcost&l canal; C, costal canal; D, transverse section; E, ventral aspect; F, trans- 
verse section through stridulating rasp; G, right elytron; H, left elytron. (Original.) 

the point where it becomes broadened and obliquely rugose. Here 
it separates from the subcosta and joins the media by a short cross- 
vein, and thence proceeds, as shown in the figure. 



THE GENUS DENDEOCTONUS. 



47 




} Strife ._ 

1 Interspaces 

— Tracked 



Media. — The media is distinctly connected with a basal, irregular, 
flexible medial plate, which is jomed to the flexor, radial, and sub- 
scapular plates by membrane and flexible chitin. Near the distal 
limit of the basal area there is an evident fold or cross vein connecting 
the base of the media with the base of the cubitus, from which it 
proceeds outward to the hinge, and 
from this pomt two branches ex- 
tend to the anal margin. 

Cubitus. — The cubitus rises from 
the outer border of the basal area 
and apex of the flexor and has two 
branches. Branch 1 extends to the 
margm. Branch 2 is short and 
more or less rudimentary. 

Anal. — The anal vein is evidently 
represented by the broad, short spur 
arising from the base of the cubitus, 
and does not extend to the margin. 

Wing attachment. — The wing is 
attached to the body by chitinous 
dorsal and ventral integument, the 
latter arising from the dorsal mar- 
gin of the epimerum, and the for- 
mer from the lateral margins of the 
prescutum, scutum, and scutellum, 
as indicated in the pupa. The heads 
or roots of the veins are attached 
by a system of connecting chitinous 
tendons and ligaments to the pleural 
and tergal processes and disks. 

Wing articulation. — The principal 
articulation of the wing is between 
the wing head formed by the costa 
and subcosta and the condyles of 
the clavicle and coracoid processes 
together with the scapular plate, 
nected with the prescutal process by articulating membrane and 
ligaments. 

Pleural clavicula (^'clavicula thoracique," Chabrier, 1820). — The 
position and function of the articular processes of the episternum 
and epimerum (fig. 20, pc), as represented more or less distinctly in all 
insects, are in Dendroctonus s-o strikingly analogous with that of the 
clavicle and coracoid in winged vertebrates as to suggest to the writer 
the same names. The giving of these names conforms with the prac- 
tice of adopting for insect anatomy such of the nomenclature of ver- 



Fig. 34. — Dendroctonus valens: Diagram of ely- 
tron, showing striae, interspaces, and tracheae. 
Striae 1-10; interspaces 1-11. Tracheaj: C, cos- 
tal; Sc, subcostal; it!, radial; 31, medial; Cu, 
cubital; A, anal. (Original.) 

The scapular plate is also con- 



48 THE SCOLYTID BEETLES. 

tebrate anatomy as is applicable to parts in insects having the same 
or similar functions. Subsequently the writer's attention was called 
to the fact that the same idea was suggested to Chabrier. 

In Dendroctonus the clavicle and coracoid processes are prominent 
and clearly defined, the condyle of the former being definitely con- 
nected with that of the head of the subcostal vein, which articulates 
between the condyles of the clavicle process and scapular plate. 
Chabrier's name clavicula has been adopted for the combined clavicle 
and coracoid processes. 

Tergal processes (fig. 20). — The processes of the tergal area, which 
have more or less important functions as articulatory accessories, 
may be designated as prescutal, scutellar, and postscTitellar. 

Prescutal process. — The prescutal process is represented by a tri- 
angular extension of the posterior angle of the prescutum, and is of. 
primary importance as an accessory to the scapular plate. 

Scutellar process. — The scutellar process is represented by the thick- 
ened lateral margin of the lateral impression and by the produced, 
acute, anterior angle. It is attached along its lateral margin to a 
pleural tendon connecting the pleural disk to the flexor and sub- 
scapula. This process is also accessory to the flexor. 

Postscutellar process. — The postscutellar process is an extension or 
arm of the anterior angle of the postscutellum, and has its apex 
attached to the pleural disk and to the pleural hook. 

Lateral emargination. — What is termed the lateral emargination 
is the emargination in the side of the scutum between the posterior 
angle of the prescutum and the scutellar process. It is present in 
most of the insects and appears to facilitate the functions of the flexor 
muscle. 

Lateral impression. — The lateral impression is an impression to 
accommodate the flexor plate when the wing is at rest. 

Basal elements. — The basal elements of the wing which fTinction as 
articulatory accessories are here referred to as head of costal vein, 
scapular, subscapular, radial, medial, and flexor plates (fig. 30). 

Head of costal vein (i) . — The head of the costal vein is produced 
beyond its fused connection with the head of the subcosta. It is 
connected to the head of the clavicle by a ligament, and evidently 
functions in extending the wing forward as well as in contributing to 
other motions. 

Scapular plate (a) . — In form and function the fundamental basal 
plate, which we here call scapular plate, is very suggestive of the 
scapula of vertebrates, but its peculiar functions require quite differ- 
ent tergal connections. It is joined by ligaments to the prescutal 
process and lateral margin of the prescutal lobe in such a manner as 
to facilitate part of its functions — that of unfolding, elevating, and 
depressing the wing. Its cond^de articulates directly with the 



THE GENUS DEISTDEOCTONUS. 49 

dorsal or inner edge of the head of the subcosta and with the clav- 
icle condyle. 

Subscapular plate (b) . — The subscapular plate is more complicated in 
its structure than the scapular plate, to which it is a direct accessory, 
the two being closely joined by articulating ligament. It functions 
as an intermediate patella-like connection of the system of tendons 
which connect the pleural disk at the head of the pleural muscle with 
the flexor, head of scapular plate, head of coracoidal process, head of 
subcosta, etc. Therefore it must be of fundamental importance in 
wing motion. 

Radial plate (d) . — The radial plate is represented hj a thin chitinous 
piece connecting the radius with the subscapula. 

Medial plate (e) . — The medial plate is of flexible chitin connecting 
the media with the flexor and subscapula. 

Both the radial plate and flexor plate evidently function as articu- 
lating accessories. 

Flexor plate (c) . — The flexor plate comes next to the scapular plate 
in its fimdamental importance in wing connection and articulation, and 
is especiall}^ fitted in structure and muscular connection for its pri- 
mary function of flexing and longitudinally folding the wing, as well 
as in the reverse action of contributing to its outward extension and 
rigidity during flight. 

Mesothoracic Wings or Elytra. 

The form and general structure of the elytra are shown in figures 1 
and 31. They are oblong, rigid shields, with a subacute apex and a 
truncate declivous base and a produced articulating head. The 
structure, like that of the metathoracic wing, consists of two layers 
of integument inclosing the tracheal and circulatory system, but 
instead of the dorsal and ventral layers being partially composed of 
flexible membrane, they are chitinous throughout. The ventral 
layer is thin and smooth, while the dorsal one is thick and deeply 
sculptured. 

Tracheation. — The six primary trachese (figs. 31, 34) occupy the 
marginal and the alternating longitudinal spaces between the rows 
of punctures. Each has numerous fine lateral branches passing be- 
tween the punctures into the intervening interspaces, producing a 
network of fine tracheae, with the punctures representing the mesh. 
Thus we have a probable explanation of the primary cause of the 
system of punctures in the elytra and the longitudinal and tranverse 
thickened spaces between them. The thickened and elevated areas 
are due to a concentration of chitin over the tracheal and circulatory 
canals, while the punctures and grooves are the points of adhesion or 
junction of the two layers to form the walls between the canals. 



50 THE SCOLYTID BEETLES. 

Sculpture. — The dorsal chitinous layer presents many and varied 
characters of sculpture, the principal elements of which are the 
striae, including the longitudinal impression and rows of punctures. 
The interspaces are longitudinal spaces between the striae. The 
rugosities of the interspaces and strige and the elevated rugose basal 
margin are all characteristic elements of sculpture. There are ten 
striae and eleven interspaces. For convenience in referring to the 
variable characters, these are numbered, beginning with those next to 
the dorsal suture, when the elytra are closed, or with the posterior or 
anal margin when the elytra are open. Thus we have interspaces 1 
to 11, and striae 1 to 10 (figs. 31, 33). 

Interspaces. — In an ideal system (fig. 34) interspaces 1 to 5 are 
continuous toward the apex with 11 to 7, leaving 6 independent 
between 5 and 7. The primary tracheae occupy interspaces 1, 3, 5, 7, 
9, and 11, There is, however, more or less variation and modifica- 
tion in the elytra of beetles from this ideal arrangement and espe- 
cially upon the distal ends and their junctions with each other on 
the declivital area. In Dendroctonus interspace 1 is usually more 
elevated and continuous to apex, where it joins the very narrow 
marginal 11; 2 is less elevated to flat, narrowed toward apex, and 
joins the very narrow and obscure submarginal 10, which becomes 
broader and distinct toward the base; 3 joins the distinct 9; 4 joins 
6 around the apex of 5, and also joins 8 around the apex of 7. 

StrisB. — In the ideal arrangement (figs. 1, 31, 34), striae 1 to 5 are 
continuous with striae 10 to 6, but the usual arrangement on the 
declivity in this genus is 1 to 3 continuous with 10 to S, while 4 is 
continuous with 5, and 6 with 7. The strial punctures range from 
small to coarse and from very distinct to obscure, and are some- 
times variable in size and appearance in the same species. The pre- 
vailing condition, however, of relative obscurity or distinctness in 
different species is of considerable specific importance. The strial im- 
pressions also vary within the genus from scarcely to distinctly or 
deeply impressed, and the prevailing condition within the species is 
of considerable value. The elytral declivity, as is usual in the scoly- 
tid beetles, bears some of the more important specific and second- 
ary sexual characters. 

The other character-bearing areas of the elytra are the lateral, 
median, and the dorsal toward the vertex and base. 

Vestiture. — The elytra are more or less distinctly clothed with 
short or long hairs. The length, size, arrangement, and areas occu- 
pied furnish important taxonomic characters in distinguishing the 
major and some of the minor divisions, as shown in the synoptic 
table. A progressive modification in. vestiture is from ver}" short 
hairs over the entire surface to longer hairs and sparsely arranged 



THE GENUS DENDROCTONUS. 51 

bristles toward the vertex of the declivity and on the declivity 
itself, or to fine and coarse long hairs over the entire surface. 

Lateral fold or costal groove (fig. 31, n, o). — In the costal edge of 
the elytron, from near the base to the median section, there is a 
lateral or costal groove for the reception, when the elytra is closed, 
of the corresponding produced and acute dorsal edge of the epi- 
sternum. There is also a deeper and broader groove in the median 
section of the costal area, for the reception of the produced dorsal 
edges of hypopleurites 3 and 4. According to LeConte and other 
writers, this lateral groove is an important subordinal character. 

Sutural tongue and groove (fig. 33, a-f). — In the sutural edge of the 
left elytron there is a deep lateral groove and produced ventral edge 
for the reception of the corresponding produced lateral edge or 
tongue of the right elytron, thus forming a tongue and groove suture. 
Toward the apex both the ventral edge of the left and the tongue of 
the right are dilated to facilitate the locking of the elytra when they 
are closed. 

Stridulating accessories. — In the male there is a transversely and 
microscopically sulcated area on the ventral surface toward the 
suture and apex of each elytron (fig. 33). When the elytra are closed 
this forms a continuous filelike surface situated directly above the 
stridulating scraper of the seventh abdominal tergite or propygid- 
ium. A peculiar, independent, upward and backward motion of 
the propygidium brings the scraper in contact with the file, and 
thus produces a peculiar chirping sound which is quite audible to the 
human ear. 

The exact location of the organs of hearing in these beetles has 
not been determined. 

Basal and pleural elements. — The basal process, or articulatory arm 
(fig. 32) of the elytron appears to represent the fused heads of the 
costa, subcosta, and radial veins. The usual scapular, subscapular, 
fiexor, and medial plates are quite definitely represented, and occupy 
the same relative positions as in the metathoracic wing. The pleural 
clavicula are represented in the mesothorax by the clavicle and cora- 
coidal processes, which are fused beneath the anterior dorsal angle 
of the episternum to form the condyles (fig. 19). The clavicle disk 
is not represented, unless it is by a narrow free piece attached to the 
costal angle of the elytral process, and represents the parapterum or 
extensor plate, to which the extensor muscle is attached. 

INTERNAL ANATOMY. 

While some study has been made of the internal anatomy of these 
beetles it has not been sufficient to warrant a detailed discussion in 
this connection. 

79980—09 5 



52 



THE SCOLYTID BEETLES. 



DIGESTIVE SYSTEM. 

The general character of the digestive system is shown in figure 
35, and no furtiier explanation is necessary in this connection than 
that given in the legend under figures 35 and 36. In figure 36 some 
of the details of the internal anatomy of the fore intestine are shown, 

and especially the structures and 
elements of the proventriculus and 
'^<Esovhagus hypopharyux. 

SECONDARY SEXUAL CHARAC- 
TERS. 



~ Prnrcntriculus 



Ve^itriculus 




While there are certain clearly 
defined secondary sexual characters 
in these beetles, they have not been 
recognized by other writers, and 
they were not found by the present 
writer until after much detailed 
study of the genus. When they 
were determined it was surprising 
how such prominent characters 
could have been overlooked. Thus 
we have another example of how 
thoroughly familiar one must be 
with a given group ia order to rec- 
ognize and properly interpret the 
significance of characters in struc- 
ture, sculpture, vestiture, etc. In 
the first subdivision of the genus 
the females are distinguished by a 
transverse ridge across the anterior 
area of the pronotum, while the 
males are distinguished by the ab- 
sence of this ridge and by more 
prominent frontal tubercles which 
are separated by a deeper frontal 
groove. 

In the second subdivision the 
females are distinguished by the 
smoother and more shining elytral 
declivity. In the third subdivision 
the females have the elytral declivity distinctly more rugose, while 
that of the males is smooth and shining. Thus we have a reversal 
of the secondary sexual characters within the same genus, which is 
an unusual occurrence. 



, -'Colon 



Fig. 35. — Dendroctonus valens: Digestive organs 
of adult. A, Fore intestine; B, mid intestine; 
C, hind intestine; a, anterior section of mid in- 
testine; 6, median section of mid intestine; c, 
posterior section of mid intestine; d, coecal 
glands; e, ileum or small intestine; /, base of 4 
malpighian tubes; g, base of2malpigliian tubes. 
(Original.) 



I 



THE GENUS DENDEOCTONUS. 



53 



In the fourth subdivision the sexes are more difficult to recognize, 
but the males are distinguished by stouter, more opaque mandibles, 
broader front, and by a narrower and more elongate antennal club. 
Whenever there is doubt as to the sex of an individual it can be settled 
by examining the pygal segments for the characters shown in figures 
23 and 24. 



Ht/popharypx ff\. 




Pig. 3&.~Dendroctonus valens: Fore intestine, showing details. A, Fore intestine; B, transverse section 
ofproventriculus; C, longitudinal section of same; D, single plate; £, teeth; i?', plates; (r, hypopharynx; 
a, muscles; 6, proventricular teeth; c, posterior condyle of mandibular articulation; d, median fossa 
of same; e, anterior condyle of same; /, hypopharyngealbracon; .9, anterior margin; ft, hypopharyngeal 
papilli. (Original.) 

PUPA. 

The general structure, proportions, and anatomical details of the 
Dendroctonus pupa are shown in figures 37 and 38. 

Among the distinctive generic characters are the large prominent 
head and broad pronotum, while among the divisional and specific 
characters are the sculpture, armatures, etc., of the head, pronotum, 
elytra, and abdominal segments, as shown in the figures and defined 
in the synoptic tables and descriptions. 



54 



THE SCOLYTID BEETLES. 



Head. — The elements of the adult head recognizable in the young pupa 
are the antennae, mandibles, maxillae, labium, and what appears to 
be a well-developed labrum, which extends to the middle of the man- 
dibles. Evidently, however, this does not represent the labrum or 
even the clypeus, but is a pad to accommodate the development of 






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the long epistomal bristles. The frontal spines in examples repre- 
senting different divisions and species are variable in size and position 
and are of considerable taxonomic importance. The antennae do not 
extend to the base of the pronotum or scarcely beyond the mouth 
parts, and the club does not extend beyond the lateral margins of the 
pronotum. 



THE GENUS DENDEOCTONUS. 



55 



Proihorax — The form of the pronotuni corresponds to that of the 
adult and its relative proportions are of some value in distinguisliing 
the species. The number and position of the frontal spines are 
fairly constant in a species, although they vary in prominence with 



rcfijite t 



Stern, 
Slernitc 

S/crnileS— 



'ffi-qile 2 



; Tirr/i/c 3 



i-ijife 4 




Epiplcnrile 6"" 
Hypoplivrile G^ 
Epiplcnrile 7^ 
Hypopkimle 



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Slc.rnite 7' 
Slernite 8 
Sternitc 9 



\Tergite l(f 
\Pleurik'Epipleurite 10 



\StcmitelO 
9 



Fig. 38. — Dendroctonus valens: Abdomen of pupa, lateral and anal aspects, a, Prescutal lobe; 6, scutal 
lobe; c, scutellar lobe; d, postscutellar area; e, supraanal lobe; /, paranal lobe; g, Infraanal lobe; q, pleural 
spines; r, caudal spine or epipleural spine of the 9tli segment; s, lateral spines; t, dorsal spines; u, anal 
opening; 10, tenth segment. (Original.) 

the age of the individual, as do other spines of the body, being much 
less prominent in the older or preimaginal stage. 

Mesothorax. — The mesotergum is subrectangular and without 
special characters, except, perhaps, in the number and arrangement 



56 THE SCOLYTID BEETLES. 

of the spines. The median process of the scutellum is prominent and 
the posterior or scutellar ridge is distinct. The base of the elytra is 
oblique and elevated, and its integument continuous with that of the 
tergum and scutellar ridge. The sternum is situated between the 
mesocoxee and the trochantins of the prothoracic leg. 

Metathorax. — The metatergum is prominent and has the usual dor- 
sal or scutellar groove. The transverse posterior or scutellar ridge is 
distinct and joined at its ends with the basal angle of the wing pads. 
Each of the scutal lobes bears a pair of spines. It differs from the 
metatergum of the adult mainly in the absence of the prescutum and 
postscutellum, as defined by external elements. 

Abdominal tergites (fig. 38). — There are nine tergites visible dor- 
sally and a very small tenth visible ventrally. Tergites 3 to 6 are 
armed more or less distinctly with dorsal, lateral, and pleural spines. 
The dorsal spines are located each side of a narrow dorsal groove; 
the pleural spines on the epipleura posterior to the spiracles, and the 
lateral spines are situated between the dorsal and pleural. The size 
of the pleural spines and the size and number of the dorsal and lateral 
ones are quite variable and of considerable importance in defining 
the minor divisions. Tergites 7 and 8 are usually unarmed, but, as 
in the adults, show sexual differences in their relative prominence; 
9 has the median lobe short, but the pleurites are greatly enlarged 
and each is armed with a prominent caudal spine. 

The four transverse divisions of the segments are quite clearly indi- 
cated in tergites 1 to 6. Tergites 7 and 8 show two divisions, the first 
representing prescutal and the second the scutal and scutellar com- 
bined, while tergites 9 and 10 are undivided. It is interesting to 
note that the dorsal and pleural armatures are borne by what is evi- 
dently the scutellar division, and that the spiracles are in the prescutal 
division, thus indicating that the prescutal represents the first pri- 
mary division and the combined scutal, scutellar, and postscutellar 
represent the second primary division. 

Abdominal sternites. — There are eight exposed abdominal sternites. 
These are sternites 3 to 10, 1 and 2 being concealed beneath the 
metacoxse, as shown in figure 38. 

Abdominal pleurites. — Epipleurites and hypopleurites 1 to 8 are 
clearly defined in the removed abdomen (fig. 38), but in 9 and 10 only 
the epipleurites are represented, as indicated by the pleural suture. 
The pleural suture is distinct to the ninth segment, where it joins the 
lines marking the dorsal and ventral limits of the pleural division. 

Spiracles. — There are nine spiracles in each side of the body, one 
large mesothoracic spiracle situated between the posterior lateral 
margin of the prothorax and the anterior ventral angle of the elytral 
pad, and eight abdominal ones, each in its respective epipleurite. 



THE GENUS DENDROCTONUS. 57 

The metathoracic spiracle is not represented. Spiracles 3 to 8 are 
exposed when the elytral pad is in normal position, but spiracles 7 
and 8 are very small and obscure. Thus the pupa has the same 
number of spiracles as the larva, while in the adult there is an addi- 
tional one, although that of the eighth abdominal tergite is apparently 
rudimentary. The larva has one thoracic spiracle, apparently in the 
prothoracic segment. The pupa has one in the mesothoracic, and the 
adult has one in the mesothorax and one in the metathorax. 

Legs. — The front and middle legs are exposed, while the hind legs 
are partially concealed beneath the elytra and wing pads. The front 
coxge are large and contiguous, the anterior fourth covered by the 
maxillse and labium, and the posterior margin extends over the 
anterior margin of the mesosternum. The middle coxae are partially 
hidden by the apex of the front tibia and its tarsus. The hind 
cox^ are for the most part exposed, and distinctly separated by an 
intercoxal area. The positions of the different parts of the legs in 
their relation to the exposed structures are shown in figure 37, and 
are of considerable taxonomic importance. The apical and subapical 
spines of the femora are also of considerable importance as distinc- 
tive characters. 

LARVA. 

The structure and general characters of the larva are shown in 
figure 39. It is of the subcylindrical, wrinkled, legless type common 
to all of the true Rhynchophora, and also has the form of mouth parts 
characteristic of the larvae of this suborder. There are three thoracic 
and ten abdominal segments, the tenth being represented by the 
anal lobes. The four longitudinal divisions, viz, one sternal, two 
pleural, and one tergal, are clearly represented in all of the segments. 
The tergal division occupies nearly one-half of the circumference, 
the two pleural divisions together about one-fourth, and the sternal 
division slightly more than one-fourth. The head is much narrower 
than the first thoracic segment and but slightly longer. The three 
thoracic segments together, or the thorax, is about one-third as long 
as the abdomen. With the exceptions of the scattering hairs on the 
head and on the scutellar lobes of the thoracic and abdominal seg- 
ments, the body is without distinguishing vestiture. 

EXTERNAL CHARACTERS. 

Head (figs. 40, 41). — The head is by far the most important part of 
the body as a bearer of taxonomic characters in the larva. The gen- 
eral structure is shown in figure 40, and the anatomical details in 
figures 41 and 42. All of the primary elements of the adult head are 
represented, but they are much more simple in their structural details. 
The more strikino; differences in the larval head are found in the 



58 



THE SCOLYTID BEETLES. 



presence of clearly defined front, clypeus, and labrum, in the articu- 
lation of the niandibles, and in their rudinientary hypostoma. 

Labrum (figs. 40, 41). — The labruni is prominent, the dorsal area 
twice as broad as long, about one-third narrower than the clypeus, 
but nearly as long, with the apical margin broadly rounded, truncate 



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\ / 

/ 




or faintly emarginate, and with several apical papillae. The median 
dorsal area bears several long hairs and two slightly elevated dark spots 
where the epipharyngeal bracons are attached. The latter somewhat 
resemble the mandibular hooks of dipterous larvae, and may or may 
not represent paired elements of the head of a primitive arthropod. 



THE GENUS DENDEOCTONUS. 



59 



Whatever their origm may have been, the present function is to sup- 
port the epipharynx and also serve as chitinous attachments for the 
depressor muscles of the labrum. They are covered by the epipharynx 
and extend down and back to the oesophagus and to a point beneath 
the base of the clypeal area. 



■££i<rm);mm) 




'OccifUat foramen 



- '~* -Occiput' 



Fig. iO.—Dendroctonusvalens: Head of larva. A, Transverse section; B, oral aspect; C, ventral aspect; D, 
longitudinal section; E, dorsal aspect; a, muscles; 6, submental lobe; c, maxilla; d, condyle of dorsal articu- 
lation of mandible; e, pleurostoma;/, hypopharyngeal bracon; (/.fossaof ventral articulation of mandible; 
h, maxillary condyle; i, gularplate; j, maxillary foramen; jj, oral foramen; fc, attachment of epipharyngeal 
bracons (labral hooks); Z,pregenalarea; m, gular area; 7ni,gularapodeme; n, attachment of labial muscle; 
0, frontal apodeme; p, integumental attachment; q, occipital apodeme; r, frontal apex; s, frontal suture; 
t, prescutal lobe of mesothoracic segment; u, scutellar lobe of prothoracic segment: v, prescutal lobe; w, 
hjrpopharyngeal bracon; x, sternal section of prothoracic segment; y, stemellar section of mesothoracic 
segment, both distorted; z, apical papilli. (Original.) 

Clypeus. — The so-called clypeus evidently does not represent an 
entirely distinct element, but a produced dilated preepistomal area 
or extension of the epistoma proper. In Dendroctonus larv^ it is 
twice as broad as long and narrowed toward the apical margin, 
which is usually slightly emarginate. The basal connection with the 
epistoma is continuous and rigid, and bears two widely separated 
bristles near the base. The sides are rounded to a rather acute im- 
pressed basal angle at the mandibular condyle. 



60 



THE SCOLYTID BEETLES. 



Epistoma. — The epistojna is quite clearly defined as a thickened 
transverse area between the clypeus and the frontal area. It does 
not extend laterally to the frontal sutures, but the ends, where they 
join the pleural ridge or pleurostoma, bear the condyles for the dorsal 
articulation of the niandibles. As in the adults, this area is quite vari- 

a5^ qT *-t oT ^ o 



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o- S 



able within the genus. It is elevated to fiat, with the anterior margin 
ranging from curved to nearly straight and the lateral angles elevated 
or slightly produced so as to form the rigid support for the dorsal 
condyles. 

Hypostoma. — The hypostoma is not represented by an exposed 
piece, but by the apodeme which forms the thickened lateral and sub- 



THE GENUS DENDROCTONUS. 61 . 

lateral margins of the jiiaxillary foramen. The anterior end supports 
the fossa {g) of the ventral articulation for the mandible, and the 
ventral end supports the condyle for the articulation of the maxillary 
cardo. It is connected across the gular space by the entogular plate. 

Pleurostoma (fig. 40, e). — The pleurostoma is represented by the 
thickened lateral margin of the oral foramen. The dorsal end con- 
tributes to the rigid support of the dorsal articulation for the mandible 
and the ventral end to that of the ventral articulation. 

Front. — The front is situated posterior to the epistoma and between 
two oblique sutures which converge from the anterior angles to the 
epicranial suture. The median area is quite variable within the genus. 
It may be flat and smooth to elevated. In the latter case it may be 
small, smooth, and convex, or prominent, transverse, and rugose. 

Epicranium. — The epicranium is represented by the dorsal areas 
of the two large lobes each side of the distinct epicranial suture and 
frontal area. These lobes are continuous throughout the occipital 
and genal areas and accommodate the very large retractor muscles 
of the mandible. The genal areas are connected by the broad ento- 
gular plate. 

Occipital foramen. — The occipital foramen is situated in the pos- 
terior ventral section of the head and occupies about one-half of the 
ventral area. It is bounded posteriorly and laterally by a broad 
entoccipital rim and anteriorly by a subchitinous rim. The occipital 
apodeme arises from the posterior margin, and extends anteriorly 
inimediately beneath the epicranial suture. 

Entogular plate (fig. 40, i). — The entogular plate is the subchitinous 
plate which forms the entocranial connection between the genal areas 
and anteriorly between the lateral sections of the hypostoma. It is 
covered by the submental lobe, part of the muscles of which are at- 
tached to the posterior angles and posterior margin. 

Maxillx (fig. 41). — The maxillae are quite simple in structural 
details. The cardo is present and distinct. Its basal articulation 
and attachment are by ligaments and a fossa to a condyle supported 
by the hypostomal apodeme. Its anterior attachment to the stipes 
is by articulating membrane. The median section is not divided into 
stipes, subgalea, and palpifer, but is one continuous piece mth the 
anterior inner angle produced into a lacinial lobe which is armed with 
a number of papillae situated on a membranous integument. The 
palpus is 2-jointed and telescopic as usual. The relative proportions, 
sculpture, and vestiture are shown in the figure. 

Lahium. — The labium of the larva is very different in structure 
from that of the adult. The submentum is represented by a lobe 
which is very broad and differs but slightly from the sternal lobe of 
the prothoracic segment with which its posterior integument is 
directly connected. The lateral integument is continuous with that 



.62 THE SCOLYTID BEETLES. 

of the maxilla and the anterior angles are extended forward to the 
base of the palpi. The nientum is represented by the median tri- 
angular chitinous plate, the posterior section of which is produced 
and narrowed, and the anterior median section is produced ante- 
riorly between the palpi and supports the ligula. The short, conical, 
2-jointed palpi are situated on the anterior angles of the mentum 
and are scarcely longer than the simple lobelike ligula which bears a 
few simple papillae. 

Hypopharynx and epipTiarynx. — The position and character of these 
important elements of the oral opening are shown in figures 40, D, 
and 41, B. 

Mandibles (fig. 42). — The mandibles are stout, with the laterodorsal 
surface rugose, except toward the apex, which is produced into an 
apical tooth; the inner edge toward the apex is provided with a sub- 
apical tooth and two small irregular medial teeth. The condyle of 
the ventral angle is globular and fits into the concave fossa (fig. 40, g) 
of the hypostoma, while in the dorsal articulation the fossa is borne 
by the mandible and the condyle by the epistoma (fig. 40, d). The 
small extensor muscle is attached to the outer basal margin midway 
between the condyles, while the large and powerful retractor muscle 
is attached to the margin of the more produced inner angle, thus 
giving a direct lateral motion to the mandibles. 

Tergites (fig. 39). — The dorsal area of the tergum of the prothoracic 
segment is undivided, but evidently represents the scutum (e) and 
scutellum (/) . The dorsal area of the mesothoracic and metathoracic 
segments has two divisions. The anterior division evidently repre- 
sents the prescutal lobe (g). There is evidence of a scutal lobe (e) on 
the lateral area of both segments, as indicated in the abdominal ter- 
gites, where the scutal lobe appears between the anterior and the 
posterior lobe. Thus the latter evidently represents the scutellar 
division, or scutellar lobe (f). 

Sternites. — The sternum of each of the segments has three sections, 
anterior, median, and posterior, or sternal (h), sternellar (i), and post- 
sternellar (j). In the thoracic segments the sternal is the larger and 
projects posteriorly over the middle of the sternellar lobe, which is 
represented by a coxal lobe each side of the sternal section. In 
some of the species these lobes have a median chitinous spot or foot 
callus at the point where a foot occurs in the legged larvae of other 
Coleoptera. The abdominal sternites have the same number of sec- 
tions, but the sternellar section i's not covered by the sternal. 

Pleurites. — -The pleurum of each segment is divided longitudinally 
by an irregular pleural groove or suture (w) . The lobe immediately 
below the groove at the end of the sternites may be referred to as the 
hypopleural (k) and that immediately above it as the epipleural (I). 
The hypopleura of the thoracic segments represent the episternum, 



THE GENUS DENDEOCTONUS. 



63 



and the epimerum is obscurely represented by the epipleura, both 
of which are but httle, if at all, different from those of the abdomen. 
The epipleural lobe of the prothoracic segment has a spiracle, while 
those of the mesothoracic and metathoracic segments are without a 
spiracle, but has lateral lobes or areas for the embryonic wing. 

Spiracles. — It will be noted in figure 39 and Plate VIII that the 
prothoracic segment has a spiracle situated on the epipleurite near 




Fig. 42. — Dendroctonus valens: Mandibles of larva. A, Latero-dorsal aspect; B, dorsal aspect; C, lateral 
aspect; D, apical aspect; a, apical tooth; 6, subapical tooth; c, median tooth; d, molar tooth; e, extensor 
tendon; /, retractor muscle disk; g, retractor muscle; h, extensor muscle; i, dorsal fossa; j, dorsal condyle; 
k, ventral condyle; I, basal ridge. (Original.) 

the epipleurite of the mesothoracic segment. The writer is not cer- 
tain as to whether or not this really belongs to the prothoracic seg- 
ment or, as in the abdomen, to the anterior or prescutal division of 
the mesothoracic segment. The metathoracic segment is plainl}^ 
without spiracles, but the abdominal segments 1 to 8, inclusive, have 
spiracles which are more or less distinct, being rather obscure in the 
first division and in section a3 (see PI. I) and without lateral tuber- 
cles, while in section a4 and subdivision D both the spiracles and 



64 THE SCOLYTID BEETLES, 

spiracular tubercles (Plate VIII) are distinct. The ninth segment is 
without spiracles. 

Chitinous plates. — In some species (division I) there are no distinct 
chitinous plates or tubercles, while in others, section a4 and subdi- 
vision D, they are present and, excepting Dendroctonus micans, 
become more distinct toward and including subdivision D, in which 
the dorsal plates of the eighth and ninth abdominal segments are 
distinctly armed. 

DIGESTIVE SYSTEM. 

The peculiar characters of the digestive system of the larva are 
illustrated in figure 43, showing, at right, a median longitudinal sec- 
tion through the body from the oral to the anal opening. In every 
respect the anatomical details of the digestive system are much more 
simple in the larva than in the adult. The same primary divisions 
of fore, middle, and hind intestine are represented and there is the 
same number of malpighian tubes, but the fore intestine is very simple 
as compared with that of the adult, the crop and proventriculus being 
scarcely different in general details from the oesophagus. 

EGGS. 

The eggs of Deudroctonus have not been studied in detail, but they 
are short, oval to oblong-oval, pearly white and shining, and appar- 
ently without distinctive generic or specific characters. 

PHYSIOLOGICAL CHARACTEEISTICS. 

In addition to the morphological characters which serve to distin- 
guish the genus, there are certain physiological characteristics pecul- 
iar to the species of the genus which serve as additional evidence of 
distinction. Indeed, it becomes more and more evident that a cor- 
rect interpretation of natural groups of individuals, termed species, 
and natural groups of species, termed genera, must be based not alone 
on a common plan of structure or similarity in one or more anatomical 
elements, but that, in order to come nearer the truth, the morpho- 
logic evidence of specific distinction must be supplemented by physio- 
logic and bionomic evidence. Some of the physiological features 
common to the species of this genus, and more or less peculiar to them, 
are found in the character of their brood galleries, in their habit of 
attacking living trees, in their concentration of effort to overcome the 
resistance exerted by the tree attacked, and especially by their ability 
to manipulate and to dispose of the quantities of resin which flow 
into their burrows in the living bast and cambium; lastly, in their in- 
timate bionomic relations to definite genera and species of conifers.*^ 

a See, also, physiological characteristics of the species, as given in the forthcoming 
Bulletin No. 83, Part I, which deals with the bionomic and economic features, and 
other characteristics peculiar to the major and minor divisions as defined in the syn- 
optic tables of galleries, host trees, and distribution in the present paper (pp. 76-79).- 



THE GENUS DENDKOCTONUS. 



65 




EpiMoma 

Clypeus 

Lahrum-r- 
a 



sophagus 



Crop-""'' 

i-^ .j'^S'-Prnuentriculus"^' 



/ \ 

/ \ 
/ \ 

f b ^ '"Ventncul'usi /, 








Fig. AZ.—Dendroctonus valens: Digestive organs of larva. 
A , Fore intestine; B, mid intestine; C, hind intestine; 
a, mouth; 6, anterior section of mid intestine; c, median 
section; d, posterior section; e, small intestine;/, coecal iQlh' 
glands; g, anus; Oth, ninth segment; 10th, tenth segment; \ '*; 

ft, base of the 2 malpighian tubules; i, base of the 4 I ^ 

malpighian tubules. (Original.) 







66 THE SCOLYTID BEETLES. 

SPECIFIC DISTINCTIONS. 

In the literature on Scolytidse, and, for that matter, on almost any 
group of insects of special systematic and economic importance, there 
is much confusion, due to different interpretations of specific distinc- 
tion. Some authors have combined many described species into one, 
while others have recognized many distinct forms among those here- 
tofore included in one species, and have proposed as many different 
names for them. It is evident that whenever "lumping" or '^ split- 
ting" is necessary for the clear definition and recognition of a species 
it should be done, but it is equally evident that neither should be 
attempted without an adequate knowledge of at least the genus rep- 
resented, in order that the true characters of specific distinction may 
be recognized from those which serve to distinguish the genus or the 
major and minor divisions of higher rank than the species. 

RANGE OR LIMITS OF SPECIFIC VARIATION. 

The determination of the range or limits of variation in characters 
utihzed for the distinction of a species is one of the most troublesome 
questions with which the systematist has to deal. With one or a few 
specimens the line separating one recognized species from another may 
be distinct and definite, but as the number of specimens from different 
localities increases' the line of distinction from allied forms often 
becomes less and less distinct until it is almost or quite obscure. 
Here is where expert judgment, based on experience and a technical 
knowledge of the special group involved, is required in order to decide 
whether or not two heretofore recognized and closely allied species 
should be kept separate or be combined. The recognition of pre- 
vailing variants or constants, or of forms having abnormal or normal 
morphologic and physiologic characters, is of special importance in 
this connection, as is also the recognition of the disturbing factor of 
parallel modification in characters and habits among species of the 
same genus, as well as among those of different genera. 

If the variants connecting two allied groups comprise only a small 
percentage of the individuals, they may be considered as departures 
from the constants of the species more nearly represented, and thus 
the groups so slightly connected will serve the purposes and require- 
ments of species and neither of them should, in the writer's opinion, 
be designated as a named subspecies, race, or variety; but if, on the 
other hand, the connecting variants comprise a large percentage of 
individuals, and no other characters sufficiently distinct and constant 
can be found by which individuals may be readily referred to one or 
the other of the heretofore recognized species, it would indicate that 
the two are not specifically distinct, 



THE GENUS DENDEOCTONUS. 67 

PROGRESSIVE MODIFICATIONS. 

The writer has been forcibly impressed with the prevaiHng princi- 
ple of progressive modification in relative proportions in form and 
structural details in scolytid and other beetles. Whenever these 
modifications in relative proportions are available for the statistical 
method of analysis it is often possible to express in numbers the dif- 
ference between species and to indicate clearly the lines of modification 
and rates of departure among the species of a genus or larger group. 

There are some good examples of this principle of progressive modi- 
fication in the genus Dendroctonus, which is manifested not alone in 
the adults, but in the pupae, larvae, and character of work, and it is 
most interesting and significant to note that the modifications are in 
the same general direction in all cases. When the species are ar- 
ranged in the order indicated by these modifications and other char- 
acters, the species of the first division to the last of those of the 
second division are found to be modified from small to larger size, 
the extremes being represented by D. frontalis, with the minimum 
length of 2.5 mm., to D. valens, with the maximum length of 9 
mm. Naturally we find the same rate of difference in size of the 
immature stages and galleries. This same tendency toward increased 
size is manifested within each subdivision, section, or minor group of 
alUed forms and appears to be a prevailing principle throughout the 
Scolytidae, and thus serves, in connection with other lines of modifica- 
tion, as one of the first guides to a natural arrangement or classifica- 
tion of the species. In Dendroctonus the progressive modification of 
characters other than size is sho^vn or indicated as follows: 

Progressive Modification op Characters in the Genus Dendroctonus. 



Primary characters. 
Body slender to stouter. 
Head large to smaller. 
Prothorax long to shorter. 

Pronotum with sides nearly parallel to distinctly narrowed or constricted anteriorly. 
Pronotum as broad as elytra to narrower. 

(A mean composite ratio of the above gives a number which expresses the relative 
proportions and serves as a species index.) 

Front grooved and tuberculate to convex and smooth. 

Elytra without long hairs to long hairs over entire surface. 

Tibia from slender to broader with a tendency to dilate toward the apex. 

Funiculus of antenna with second joint long to shorter. 

Secondary sexual characters. 

Front of head with sexual differences to similar or alike in both sexes. 
Pronotum with sexual differences to alike in both sexes. 
Elytra! declivity without sexual differences to distinct differences. 
Declivity rugosities small to coarse, smooth in female to coarse in male or reversed. 
Mandibles alike or similar in both sexes, to much stouter in the male, 
79980—09 6 



68 THE SCOLYTID BEETLES. 



Front of head grooved to convex. 
Body spines small to coarse. 

LARV^. 

Body simple, without chitinous plates or hairs, to distinct chitinous plates and more 
prominent hairs. 

Eighth and ninth abdominal segments without chitinous plates to with plates, these 
last unarmed to armed. 

Spiracles simple to complex, smooth to tuberculate. 

Epiplem-ites without tubercle, to prominent tubercle. 

GALLERIES. 

Long and winding to short and straight. 

Eggs isolated to grouped and massed. 

Larval mines hidden to exposed and short to long.' 



From one genus to many genera, and from one species to many species. 
D. brevicomis is found in pine only, while Z>. raZe?is infests Pinus, Picea, and Larix; 
D. simplex infests Larix only, and D. pseudotsugx infests Pseudotsuga and Larix. 

DISTINCTION OF MAJOR AND MINOR DIVISIONS OF THE GENUS. 

In a comparative study of the species of the genus to determine 
their relative positions, as indicated by degrees of resemblance or dif- 
ference, they are found to fall according to progressive modification of 
characters into major and minor divisions, which may be designated as 
divisions, subdivisions, sections, subsections, series, and subseries, to 
the smallest practicable minor division of the genus, viz, the species. 

In this classification of the genus the rank of a primary division may 
be that of the subgenus of some authors and the lower series of closely 
allied species may be recognized by some systematists as occupying 
the rank of subspecies, races, or varieties; but the writer has been 
guided by the belief that the principle of a less restricted range of 
generic and more restricted range of specific distinction will contribute 
toward a more correct knowledge of the forms of life than if the 
reverse principle is followed. 

The classification of the species of a genus into major and minor 
divisions is necessarily arbitrary, and is subject to changes as may 
be suggested by increased knowledge and the addition of species. 
To a more limited extent, the designation of a species is arbitrary 
and with additional material and information is subject to revision; 
but since the species, next to the individual, is the constant or unit 
of classification and investigation, it should represent tlie lowest prac- 
tical division of a genus that is recognizable from a description of a 
typical form or by comparison with the type on which the description 
was based. 



THE GENUS DENDEOCTONUS. 69 

PLAN OF SYNOPTIC TREATMENT. 

The plan here appHed for the classification and synoptic treatment 
of the species of the genus is one which appears to be most available 
and practicable for the clear definition of the progressive modification 
of taxonomic characters and for indicating the relative systematic 
positions and limits of the major and minor divisions and the species. 
It is not radically different from some of the more generally adopted 
dichotomous systems, and it conforms to the primary objects of a 
synopsis' in that it provides (a) for a direct comparison of opposing 
characters, (b) for a direct line of references leading down to the 
specific characters, or vice versa. 

With this method of indicating the supposed natural relation of 
the species, the described characters of the major and minor divisions 
and sections, together with those of specific distinction, serve as a 
description of the species. Thus, division I, subdivision A, section a2, 
subsection b2, series c2, defines the characters common to species 
6, 7, and 8, which are separated by their respective specific char- 
acters. Some additional advantages of this method are the con- 
secutive arrangement of letters and figures which throughout a given 
table are not duplicated. The Roman numerals indicate at once the 
primary divisions, the capital letters the subdivisions, and the com- 
bined small letter and Arabic numeral the sections, subsections, 
series, etc., to any desired limit. The reference from right margin 
to center, instead of to left margin, is also an advantage in defining 
the linfits of a major and minor division. It also provides for full 
paragraphs, thus economizing space and cost of printing. 

SYNOPSES OF MORPHOLOGICAL AND PHYSIOLOGICAL 
CHARACTERS. 

SYNOPSIS OF ADULT CHARACTERS. 

Pronotum somewhat elongate and as broad as elytra; not distinctly narrowed ante- 
riorly except in subdivision B; anterior dorsal half of elytra without long hairs. 

Division I, pages 69, 81. 

Pronotum stout; usually narrower than elytra, and distinctly narrowed and constricted 
anteriorly; anterior dorsal half of elytra normally with long hairs, except in tere- 
brans Division II, pages 71, 116. 

DIVISION I. 

Body somewhat slender, pronotum but slightly narrowed anteriorly; elytral declivity 
with second stria straight, second interspace not distinctly broader or narrowed 
toward apex; head with frontal groove and tubercles except in convexifrons. 

Subdivision A, pages 69, 81. 
Body stout; pronotum distinctly narrowed and constricted anteriorly; elytral decliv- 
ity with second stria curved, second interspace broad and distinctly narrowed 
toward apex; head without frontal tubercles or groove. 

Subdivision B, pages 71, 105. 
Subdivision A. 

Elytral declivity without long hairs Section al, page 70. 

Elytral declivity vdth long hairs Section a2, page 70. 



70 THE SCOLYTID BEETLES. 

Section al. 

Length 3 to 5 mm., browoish to nearly black; elytral rugosities fine, densely placed, 
the striae obscure and but slightly or not at all impressed. California, Oregon, 

Washington, and Idaho, in Pinus 1. brevicomis Lee, page 81. 

Length 2.5 to 4.7 mm., brownish to black; elytral striae moderately to distinctly 
impressed ; interspacial rugosities moderately coarse and obtuse to coarse and acute. 
Arizona, New Mexico, southern Colorado, southern Utah, and southei'n California. 

2. barberi n. sp., page 85. 
Section a2. 

Head loithout frontal tubercles or groove, but with posterior impression. 

Subsection bl, page 70. 

Head with more or less prominent frontal tubercles each side of a distinct median 

groove Subsection b2, page 70. 

Subsection bl. 

Length 4 to 6 mm.; reddish-brown to black, shining; body slender; elytral rugosities 
moderately coarse but not densely i^laced, except toward base, the striae faintly 
impressed, with rather coarse indistinct punctures; pronotum with long, erect 
hairs on the entire lateral area; declivity shining. Arizona, New Mexico, south- 
ern Colorado, and southern Utah, in Phius 3. convexifrons n. sp., page 87. 

Subsection b2. 

Elytral striae distinctly punctured Series cl, page 70. 

Elytral striae not distinctly punctured Series c2, page 70. 

Series cl. 

Length 2.5 to 4 mm.; brownish to black; elytral rugosities obtuse, moderately coarse, 
not very densely placed, and but moderately coarser toward the base and vertex 
than elsewhere; pronotum usually with a few long hairs on the anterior section 
of the lateral area. Pennsylvania to Florida, westward to Ohio and Texas, in 
Pinus and Picea 4. frontalis Zimm., page 90. 

Length 4 to 5 mm.; brownish to black; elytral rugosities subacute, moderately coarse 
and distinctly coarser toward the base and vertex; pronotum with long erect hairs 
on the anterior half of the lateral areas; elytra with long hairs confined to declivity 
and posterior areas. Central Arizona, in Pinu^ 5. arizonicus n. sp., page 95. 

Series c2. 
Striae distinctly impressed. 

Length 3 to 4.5 mm.; brownish to black; elytral rugosities distinctly coarser 
toward the base and vertex, and with an evident row of acute rugosities on the 
first interspace; pronotum clothed with stout reclining hairs and the entire 
lateral area with long, erect hairs; elytral strias moderately distinct. Southern 

Mexico, in Pinus 6. mexicanus Hopk., page 97. 

Striae not distinctly impressed. 

Length 5 to 6 mm.; black; elytral rugosities obtuse, rather densely placed, mod- 
erately coarse toward dorsal suture, fine and less evident toward side; striae 
obscure, especially on the side; pronotum distinctly pubescent, with very long 
erect hairs on the entire lateral area; punctures moderately coarse. Southern 

Mexico, in Pinus 7. parallelocollis Chap., page 99. 

Length 4 to 7.4 mm.; black; elytral rugosities rather coarse and sparse; the striae 
toward the side rather distinct; pronotum moderately pubescent, with long 
hairs on the anterior two-thirds of the lateral area; punctures usually fine. 
Arizona, New Mexico, southern Colorado, and Utah, in Pinus. 

8. apjproximatus Dietz, page 101. 



THE GENUS DENDKOCTONUS. 71 

Subdivision B. 
Pronotum with deep punctures. 

Length 3.7 to 6.4 mm.; brownish to black; elytra with striae moderately impressed, 
punctures usually small or moderately coarse and distinct. North and west of 
northwestern Colorado, southward to Yosemite National Park, Caliioinia, in 

Pinus 9. moniicolse Hopk., page 105. 

Length 4.5 to 7 mm.; black; elytral stride distinctly impressed, punctures dis- 
tinct and coarse. Black Hills, South Dakota, southward through Colorado 
and southern LTtah, into New Mexico and Arizona, in Pinus and Picea. 

10. ponder osse Hopk., page 109. 
Pronotum with small shalloiv punctures. 

Length 6 to 8 mm.; black; punctures of elytral striae distinct throughout; pronotum 
distinctly shining. Yosemite National Park, California, in Pinus jeffreyi and 
P. ponderosa 11. jeffreyi n. sp., page 114. 

DIVISION II. 

Front usually ivith posterior impression; pronotum with large and small punctures 

intermixed Subdivision C, pages 71, 117. 

Front without posterior impression; pronotum with regular punctures. 

Subdivision D, pages 72, 146. 
Subdivision C. 

Elytral declivity with striae deeply impressed; epistomal process narrow, flat, with 
lateral sections nearly parallel; pronotum with punctures moderately regular, 

and with long hairs on dorsal and lateral areas Section a3, page 71. 

Elytral declivity with striae not deeply impressed; epistomal process broad, concave, 
and the lateral sections oblique; punctures of pronotum distinctly irregular. 

Section a4, page 71. 
Section aS. 

Pronotum with coarse punctures. 

Length 3.5 to 5 mm.; reddish to brown; pronotum with short dorsal and lateral 
hairs; apex of epistomal process not extending beyond the anterior frontal 
margin. New Brunswick to Michigan and West Virginia, in Larix. 

12. simplex Lee, page 117. 
Pronotum with fine, shallow punctures. 

Length 4 to 7 mm.; reddish to dark brown; pronotum shining, with long dorsal 
and lateral hairs; apex of epistomal process usually extending beyond the 
anterior margin of epistoma. British Columbia southward to Texas, westward 
to California, in Pseudotsuga and Larix 13. pseudotsugse Hopk., page 121. 

Section a4. 

Posterior half of proepisternal area not distinctly punctured. Subsection b3, page 71. 
Posterior half of proepisternal area distinctly punctured Subsection b4, page 72. 

Subsection hS. 

Length 4.7 to 6 mm.; body stout; reddish to black; elytral striae quite distinctly im- 
pressed toward sides, with punctures coarse and distinct; interspaces convex; 
rugosities acute, rather closely placed, irregular. New Brunswick, through 
Canada, New England, and Michigan, in Picea. . . 14. piceaperda Hopk., page 126. 

Length 5 to 7 mm.; reddish to black, shining; punctures of prothorax and elytra 
coarser; striae of lateral area not distinctly impressed, the interspaces scarcely 
convex or rugose. Idaho and Black Hills, South Dakota, to New Mexico and 
westward to California, in Picea 15. engelmanni n. sp., page 130. 



72 THE SCOLYTID BEETLES. 

Length 6 mm.; elytra reddish; pronotum darker; elytral striae moderately im- 
pressed, punctures of strise of dorsal area small, obscure, and those of the lateral 
area moderately coarse; interspaces flat, with rugosities moderately coarse, obtuse, 
and rather densely placed; pronotum with punctures more regular in size than 
usual. Alaska 16. borealis n. sp., page 133. 

Length 6 to 7 mm. ; black; elytral strise but faintly impressed; punctures rather coarse; 
interspaces flat; rugosities moderately coarse, not dense, acute toward vertex; 
pronotum with fine punctures, moderately dense, coarser toward sides. Pacific 
coast, Oregon to Sitka, in Picea sitchensis 17. obesus (Mann.), page 135. 

Subsection b4- 

Strise of declivity with fine punctures Series c3, page 72. 

Strise of declivity with coarse punctures Series c4, page 72. 

Series c3. 

Length 5 to 7.3 mm.; elytra red; pronotum brown to black; body rather stout; elytral 
strise scarcely impressed, except on dorsal area; strial punctures moderately 
coarse; rugosities of interspaces moderately coarse, acute, not dense; pronotal 
punctures coarse, deep, moderately dense; elytral declivity with interspaces 
moderately punctured. Lake Superior, in Pinus strobus. 

18. rufipennis (Kirhy), page 138. 

Length 5.4 to 6.5 mm.; elytra dark reddish; pronotum dark; body somewhat slender, 
as compared with species 18; elytral strise moderately impressed; punctures 
coarse, shallow; pronotal punctures coarse, not distinctly irregular and mod- 
erately dense; elytral declivity with interspaces finely densely produced. 
Yellowstone National Park and Keystone, Wyoming; Alberta, and British Colum- 
bia, in Pinus 19. murrayanae n. sp., page 140. 

Series c4. 

Length 6.5 mm.; brownish; elytral striae impressed; punctures coarse, interspaces 
narrow and moderately coarse, acute, rugosities coarser toward the suture. 
New York to West Virginia, in Picea 20. punctatas Lee, page 142. 

Length 7 to 8 mm.; brownish; elytral strise not impressed on dorsal or lateral areas, 
punctures moderately coarse, shallow; interspaces broad, flat, shining; rugosities 
small, obtuse, sparse. Europe, etc., in Picea, Larix,? and Abies? 

21. micans (Kug.), page 143. 

Subdivision D. 

Pronotum somewhat elongate, slightly narrower than elytra, moderately constricted 
toward head, median and posterior dorsal areas without long hairs, but hairs pres- 
ent on anterior and lateral areas; head broad, epistomal process usually broad, 
concave, with sides strongly oblique; elytral rugosities variable, usually coarse. 
Body black. . 

Length 5 to 8 mm.; black; epistomal process moderately broad, with apical 
angles usually tub ere ulate; pronotal punctures very coarse, regular, moderately 
dense, scarcely decreasing in size toward base; elytra without long hairs toward 
base. Long Island, New York, south to Florida, west to West Virginia and 

Texas, in Pmi6S 22. terebrans {Oliw), page 147. 

Body reddish. 

Length 5.7 to 9 mm.; reddish, never black; epistomal process broad, with apical 
angles obtuse, never tuberculate; pronotum with punctures smaller and denser 
toward base; elytra with long hairs toward base. Eastern United States and 
Canada, north of mountains of North Carolina, westward to Pacific coast, south 
from British Columbia into Mexico, in Pinus and Picea. 

23. valens Lee, page 151. 



THE GENUS DENDEOCTONUS. 73 

SYNOPSIS OF SECONDARY SEXUAL CHARACTERS. 
DIVISION I. 

Females: Anterior pronotal area with transverse ridge Subdivision A. 

Females: Anterior pronotal area without transverse ridge Subdivision B. 

Subdivision A. 

Females: With transverse ridge across the anterior area; elytral declivity slightly 

smoother, more shining, and less rugose Species 1 to 8, inclusive. 

Males: Without transverse ridge across the anterior area, but with broad impression. 

Species 1 to 8, inclusive. 
Subdivision B. 

Femnles: Elytral declivity with interspaces more shining, rugosities less prominent. 

Species 9 to 11, inclusive. 
Males: Elytral declivity with interspaces more opaque, rugosities more prominent. 

Species 9 to 11, inclusive. 
DIVISION II. 

Elytral declivity with distinct sexual characters in both sexes Subdivision C. 

Elytral declivity without distinct sexual characters in either sex Subdivision D. 

Subdivision C. 

FeTnales: Interspaces of elytral declivity rugose. Males: Interspaces of elytral 

declivity smooth. 
Striae of elytral declivity impressed; interspaces convex in both sexes. 

Section a3, Species 12 and 13. 
Striae of elytral declivity not impressed; interspaces flat Section a4. 

Section a4. 

Females: Striae of elytral declivity slightly impressed; interspaces subconvex. 

Species 14 to 21, inclusive. 
Males: Striae of elytral declivity not impressed; interspaces flat. 

Species 14 to 17, inclusive. 

Males: Striae of elytral declivity slightly impressed; interspaces subconvex, but 

smoother than in female Species 18 to 21, inclusive. 

Subdivision D. 

Females: Head with front moderately broad; mandibles shining, moderately stout; 

antennal club broader Species 22 and 23. 

Males: Head with front broad; mandibles opaque, stout; antennal club narrow, more 

elongate and less compressed Species 22 and 23. 

SYNOPSIS OF PUPAL CHARACTERS. 

Vertex of head distinctly to faintly grooved, and with two small or prominent frontal 
spines on or toward the vertex each side of groove Division I. 

Vertex of hed^d faintly impressed, flat or convex, and with two small, widely separated 
frontal granules toward vertex Division II. 

DIVISION I. 

Frontal spines small; elytral pads smooth; abdominal tergites 2 to 6 with or without 
small pleural spines Subdivision A. 

Frontal spines large, prominent; elytral pads rugose; abdominal tergites 2 to 6 with 
prominent pleural spines Subdivision B, 



74: THE SCOLYTID BEETLES. 

Subdivision A. 

Anterior and middle femora smooth; abdominal tergites 3 to 6 with small pleural spines; 

1 and 2 without distinct dorsal and lateral spines. Section al, Species 1, 2, and 3. 

Anterior and middle femora ivith small apical spines Section a2. 

Section a2. 

Abdominal tergites 2 to 6 without distinct pleural spines; 7 and 8 with small gran- 
ules Species 4. 

Abdominal tergites 2 to 6 with small pleural spines of equal size; 7 and 8 with small 

granules Species 5. 

Abdominal tergites 1 to 6 with small pleural spines, increasing in size; 7 and 8 smooth. 

Species 8. 
Subdivision B, 

Apex of anterior and middle femora with two spines Species 9. 

Apex of anterior and middle femora with one spine; abdominal spines and elytral 

rugosities coarser than in species 9 Species 10. 

Apex of anterior and middle femora with two spines; abdominal spines apparently less 

prominent than in species 9 and 10 Species 11. 

DIVISION II. 

Vertex of head fattened ov faintly impressed; apex of front and middle femora smooth 
or with minute granule; abdominal tergites with pleural and dorsal spines moder- 
ately prominent Subdivision C. 

Vertex of head convex; front and middle femora each with a minute pilated, subapical 
granule; abdominal tergites with less distinct pleural and dorsal spines. 

Subdivision D. 
Subdivision C. 

Tergal spines of abdomen with pale or white tips; segment 7 smooth Section a3. 

Tergal spines of abdomen with pale, dark, or black tips; segment 7 with a few setigerous 
granules Section a4. 

Section aS. 

Anterior and middle femora with minute apical spines; abdominal tergites 2 to 6 with 
stout, prominent pleural spines; 3 to 6 with prominent dorsal spines. Species 12. 

Anterior and middle femora without apical spines; abdominal tergites 2 to 6, with mod- 
erately stout pleural spines, and 3 to 6 with small dorsal ones Species 13. 

Section a4. 

Vertex of head moderately impressed; anterior and middle femora without apical spines; 

abdominal tergites 2 to 6 with very small pleural spines; 4 to 6 with small dorsal 

spines, all with pale tips Species 14. 

Vertex of head distinctly impressed; abdominal tergites 2 to 6 with distinct pleural 

spines, and 3 to 6 with distinct dorsal ones Species 15. 

Abdominal tergites 2 to 6 with very small pleural, lateral, and dorsal spines, all with 

dark tips Species 17. 

Subdivision D. 

Abdominal tergites 1 to 6 with moderately small pleural spines, 2 to 6 with small dorsal 

and lateral ones, all with pale tips Species 22. 

Same as preceding, except spines have darker to black tips Species 23. 



THE GENUS DENDKOCTONUS. 75 

SYNOPSIS OF LARVAL CHARACTERS. 

Abdominal tergites 8 and 9 without dorsal plates Division I. 

Abdominal tergites 8 and 9 loith dorsal plates, except in section a3 Division II. 

DIVISION I. 

Front with or without median convexity Subdivision A. 

Front with transverse elevation slightly more prominent toward the sutures. 

Subdivision B. 
Subdivision A. 

Front without median convexity Section al. 

Front ivith median convexity Section a2. 

Section al. ^ 

Clypeus with apex subacutely emarginate Species 1. 

Clypeus with apex broadly emarginate Species 2. 

Section a2. 

Prothoracic lobes without foot calli Subsection bl. 

Prothoracic lobes with distinct foot calli Subsection b2. 

Subsection bl. 

Front smooth, with shining convexity; clypeus with apex broadly emarginate. 

Species 3. 

Subsection b2. 

Front smooth, with median smooth convexity; clypeus with apex deeply emargi- 
nate Species 4. 

Front with anterior third transversely rugose and with transverse median convexity, 
produced toward apex; clypeus with apex broadly emarginate Species 5. 

Front with broad convexity; clypeus with apex truncated Species 8. 

Subdivision B. 

Front of head with posterior apex subacute; frontal elevation moderately stout in the 
middle; clypeus with faint median tubercle toward the base Species 9. 

Front of head with posterior apex subobtuse; frontal elevation stout slightly posterior 
to the middle; clypeus with a faint median groove and elevation toward base. 

Species 10. 

Front of head with posterior apex subacute; frontal elevation narrow, situated in the 
middle, not more distinctly elevated toward suture; clypeus with faint median 
groove, without elevation Species 11 . 

DIVISION II. 

Abdominal tergites 8 and 9 without dorsal plate, or with unarmed plates. 

Subdivision C. 
Abdominal tergites 8 and 9 with armed plates Subdivision D. 

Subdivision C. 

Abdominal tergites 8 and 9 loithout dorsal plates Section aS. 

Abdominal tergites 8 and 9 -with dorsal plates, excepting micans, in which 8 is un- 
armed Section a4. 



76 THE SCOLYTID BEETLES. 

Section aS. 

Front with indistinct transverse elevation; clypeiis with distinct median impression. 

Species 12. 
Front with distinct transverse elevation; clypeus with median impressed line. 

Species 13. 

Section a4. 

Abdominal tergites 8 and 9 with rugose but unarmed plates; front with faint trans- 
verse elevation anterior to middle; clypeus with median groove Species 14. 

Frontal elevation near the middle Species 15. 

Frontal elevation distinctly anterior to tniddle Species 16. 

Front with lateral impressions toward the anterior angles, and fi'ontal elevation ante- 
rior to middle Species 19. 

Abdominal tergite 9 with dorsal plate; frontal elevation anterior to middle; clypeus 
with median groove Species 21. 

Subdivision D. 

Abdominal tergites 8 and 9 each with dorsal plate and armed with three permanent 
teeth; front of head without elevation; clypeus with faint median groove. 

Species 22 and 23. 

SYNOPSIS OF GALLERY CHARACTERS. 

Egg galleries winding to straight; eggs isolated or in approximate groups, but never in 
masses; larval mines exposed or concealed in inner bark Division I. 

'EjgggdMexies longitudinal, straight to slightly winding; eggs in groups or masses; larval 
mines and pupal cells exposed in inner bark Division II. 

DIVISION I. 

Pupal cells in outer hark; eggs isolated, never in groups; egg galleries winding; larval 
mines short, narrow to broad, exposed or concealed Subdivision A. 

Pupal cells in inner bark; eggs in approximate groups; egg galleries slightly winding 
to straight; larval mines short, broad, always exposed Subdivision B. 

Subdivision A. 

Larval mines concealed in inner bark Section al. 

Larval mines exposed or concealed Section a2. 

Section al . 

Egg galleries subtransverse, winding Species 1. 

Egg galleries distinctly transverse Species 2. 

Section a2. 

Larval mines exposed. 

Egg galleries longitudinal, winding Species 3. 

Egg galleries sublongitudinal, winding Species 4. 

Larval mines concealed. 

Egg galleries evidently transverse, or subtransverse, winding, with concealed 
larval mines Species 5. 

Egg galleries subtransverse, winding Species 6. 

Egg galleries longitudinal, winding, with transverse branches. . Species 7 and 8. 

Subdivision B. 

Egg galleries winding to straight Species 9. 

Egg galleries usually straight .• . . Species 10 and 11. 



THE GENUS DENDROCTONUS. 77 

DIVISION II. 

Larval mines separate, especially beyond the middle Subdivision C. 

Larval mines contiguous, forming broad larval chamber Subdivision D. 

Subdivision C. 

Egg galleries slightly winding to straight; eggs in groups, but larval mines separate 
from the beginning Section a3. 

Egg galleries broad, nearly straight; eggs in small to large groups, the larval mines 
usually contiguous toward the egg gallery ": Section a4. 

Section a.3. 

Larval mines normally short and broad Species 12. 

Larval mines normally long Species 13. 

Section a4. 

Subsection b3. 

Larval mines separated beyond the middle Species 14, 15, and 16. 

Subsection b4. 

Larval mines usually not separated beyond the middle, but forming a common chamber. 

Species 19 to 21. 
Subdivision D. 

Egg galleries broad to very broad, short to very long, straight to winding; larval mines 
forming a large common chamber Species 22 and 23. 

TABLE OF DISTRIBTJTION.a 

America, north of Guatemala, and in northern Europe The genus. 

DIVISION I. 

Subdivision A. 

North American continent, in South Atlantic and Gulf States and Southwestern 
States, southward to Guatemala and northward in Sierra Nevada and Cascade 
Mountains to British Columbia Sections al and a2. 

Section al. 

West of western Montana and southwestern Idaho, and southward to Santa Barbara 
County, California Species 1, page 81. 

South of southern Colorado and Utah, into Texas and Mexico, and westward into 
southern California, and possibly in the Coast Range and northward along the 
forested foothills into northern California Species 2, page 85. 

Section a2. 

Southern Colorado and Utah, southward into Mexico Species 3, page 87. 

Atlantic and Gulf States, south westward into Texas Species 4, page 90. 

Central to southern Arizona Species 5, page 95. 

Southern Mexico Species 6 and 7, pages 97, 99, 

Central Colorado and southern Utah, southward to southern Arizona, and New Mexico. 

Species 8, page 101. 

a For exact and probable distribution see maps under description of each species. 



78 THE SCOLYTID BEETLES. 

Subdivision B. 

"West of western Montana and southwestern Idaho, south through the Sierra Nevada 
Mountains of California Species 9, page 105. 

Western South Dakota and southern Wyoming, southward through Utah, Colorado, 
to southern Arizona and New Mexico Species 10, page 109. 

Northern California, southward in Sierra Nevada, into San Bernardino County, Cali- 
fornia Species 11, page 114. 

DIVISION II. 

Subdivision C. 

Maine to western Michigan, southward into northwestern West Virginia. 

Species 12, page 117. 
Northern Idaho and Montana, south to southern Arizona and New Mexico, and north- 
ern Washington, south into Santa Barbara County, California. Species 13, page 121. 

Section a4. 

Maine to northeastern Minnesota, southward to central Pennsylvania. 

Species 14, page 126. 
Northern Idaho, east to western South Dakota, southward to southern Arizona and 

New Mexico Species 15, page 130. 

Alaska.. Species 16, page 133. 

Alaska?, along the coast to northwestern California Species 17, page 135. 

Lake Superior region • Species 18, page 138. 

Western Montana southeast to central Colorado Species 19, page 140. 

Highermountainsof New York, Pennsylvania, and West Virginia. Species 20, page 142. 
Central Europe to Denmark, Russia, and eastward into Siberia. Species 21, page 143. 

Subdivision D. 

Atlantic States south of Massachusetts, to Tampa, Florida, westward to western West 
Virginia and Texas Species 22, page 147. 

Mountains and foothills of North Carolina, northward into Maine and northwestern 
Washington, southward into Guatemala Species 23, page 151. 

Guatemala Species 24, page 157. 

TABLE SHOWING RELATION OF SPECIES TO HOST TREES. 

DIVISION I. 

Section al. 

Species 1. Pinus lambertiana, ponderosa. 

Species 2. Pinus ponderosa var. scopulorum, edulis; Pseudotsuga taxifolia. 

Section a2. 

Species 3. Pinus scopulorum. 

Species 4. Pinus strobus, tseda, rigida, virginiana, pungens, echinata, glabra, palustris; 

Picea rubens and excelsa. 

Species 5. Pinus scopulorum. 

Species 6. Pinus teocotl?, lejophilla, and ayacahuite'^ 

Species 7. Same as 6. 

Species 8. Pinus arizonica, scopulorum, and cidhuahuana. 

Subdivision B. 

Species 9. Pinus moniicola, lambertiana, ponderosa, murrayuna; Picea engelmanni. 
Species 10. Pinv^ fiexilis and strobiformis. 
Species 11. Pinus lambertiana and ponderosa. 



THE GENUS DENDEOCTOlSruS. 



79 



DIVISION II. 

Section aS. 

Species 12. Larix laricina. 

Species 13. Larix occidentalis; Pseudotsuga taxifolia and macrocarpa. 

Species 14. Picea mariana, rubens, canadensis. 

Species 15. Picea canadensis and engelmanni. 

Species 16. Picea canadensis. 

Species 17. Picea sitchensis. 

Subsection b4. 

Species 18. Pinus strobus. 

Species 19. Pinus murrayana; Picea engelmanni. 

Species 20. Picea rubens. 

Species 21. Pinus, Picea, Abiesf, Larix. 

Subdivision D. 

Species 22. Pinus strobus, taeda, rigida, serolina, echinata, palustris. 

Species 23. Pinus strobus, monticola, lambertiana, strobiformis, ponderosa, scopulorum, 

jefreyijChihuahuana, murrayana, radiata, rigida, virginiana; Larix laricina; 

Picea rubens, canadensis, excelsa. 

TABLE OF HOST TREES. 



Dendroctonus spe- 
cies number. 

Pinus strobus 4, 18, 22, 23. 

monticola 9, 23. 

lambertiana 1, 9, 11, 23. 

Jiexilis 10. 

strobiformis 10, 23. 

edulis 2,23. 

arizonica 8, 23. 

ponderosa 1,9,11,23. 

scopulorum 2, 3, 5, 8, 10, 23. 

jeffreyi 11, 23. 

chihuahuana 3, 8, 23. 

murrayana 9, 10, 19, 23. 

radiata 23. 

tseda 4, 22. 

rigida. 4,22,23. 

serotina 22. 

virginiana 4, 23. 

pungens 4. 



Dendroctonus spe- 
cies number. 

Pinus echinata 4, 22. 

glabra 4. 

palustris 4, 22. 

sylvestris 23. 

teocotl 6,7. 

lejophilla 6,7. 

ayacahuite 6,7. 

Larix laricina 12, 23. 

occidentalis 13. 

Picea mariana 14. 

rubens 4, 14, 20, 22, 23. 

canadensis 14, 15, 16, 23. 

engelmanni 9,10,15,19. 

sitchensis 17. 

excelsa 4,23. 

Pseudotsuga taxifolia 2, 13. 

macrocarpa. . 13. 



REVISION AND SYSTEMATIC NOTES, WITH DESCRIPTIONS OF 

NEW SPECIES. 

DISTINCTIVE GENERIC CHARACTERS. 

Adult (figs. 1, 2, 3). — Antennal funiculus 5-jointed; club broad, 
thickened at base, compressed toward apex, and usually with. 4 dis- 
tinct segments, the sutures curved or nearly straight; tarsi with joint 
3 bilobed; tibia with inner angle produced and armed with a single 
tooth; outer angle oblique and armed with 3 or more stout teeth; 
distinct dorsal impression toward apex for the retractile tarsus. 



80 THE SCOLYTID BEETLES. 

Anterior coxae approximate or subcontiguous. Abdominal sternites 
with ends of sutures 4, 5, and 6 strongly recurved. Body cylindrical, 
subelongate to stout, ranging in color from reddish and brown to 
deep black. Head prominent and large, with distinct epistomal proc- 
ess at anterior margin of front. Eyes transversely placed and 
oblong-oval to oblong-ovate. Antennal insertion in front of ventral 
end of eye. Pronotum with sides nearly parallel to narrowed and 
constricted toward head, one-fourth to one-third broader than long. 
Elytra with base elevated and rugose, remaining surface rugose, with 
punctured striae and the declivity convex to subconvex. 

Pupa (figs. 37, 38) . — The pupa is of the general size of the adult, and 
is distinguished by its broad prominent head, and the form of the pro- 
thorax. The sculpture and armature vary with the age of the specimen. 
In the preimaginal stage the granules and spines become more obscure. 

Larva (text fig. 39; PI. VIII) .^The body of a matured larva 
of a given species is somewhat longer than the adult or pupa, and 
is cylindrical, deeply wrinkled, legless, and with a few long hairs on 
each segment, becoming longer on the posterior ones. The head 
is moderately large, shining, yellowish, and with a few hairs on the 
scutellar lobes. Front distinct; antennae present, but obscure; eye 
spots not present. The thoracic segments are larger and more prom- 
inent in some species than in others. Abdominal segments 1 to 9 are 
of about equal width and length; 10 is represented by the anal lobe. 

Egg. — Short, oval to oblong-oval, pearly white and shining, and 
apparently without sculpture and specific characters, except in rela- 
tive size, corresponding with the size of adult representatives of the 
species. 

Galleries. — The primary or egg galleries are excavated in the inner 
bark and sometimes mark or groove the wood and vary in their course 
in the bark of the tree from transverse and winding to longitudinal 
and straight, and normally are of the single unbranched type. 

Distribution (PI. II). — Eastern continent: Central and northern 
Europe, from Denmark into Siberia. Western continent : Guatemala, 
northward through the United States into Alaska and Labrador. 

Host trees. — Pinus, Picea, Pseudotsuga, Larix, and Ahies, the latter 
rarely, if at all. 

BIBLIOGRAPHY AND SYNONYMY OF GENUS. 

Erichson, 1836, pp. 52-53, original description, to include i). micans (Kug.) (type), D. 
terebrans (Oliv.) (cotype), Myelophilus pinij^erda (L.), Myelophilus minor (Hartig), and 
Carphoborus minimus (Fab.). Eichhoff, 1864, pp. 26-27, PL I, figs. 5, 6, 7, tibia, maxilla, 
labium, revised description to include the single Em'opean species, D. micans (Kug.). 
Lacordaire, 1866, pp. 360-361, revision to include D. m,icans (Kug.) and D. terebrans 
(Oliv.). Zimmerman, 1868, pp. 148-149, revision to include C'arp/«o6orMs6t/'u?'eitsEich., 
D. terebrans (Oliv.), and D. frontalis Zimm. Le Conte, 1868, pp. 172-173, revision to 
include i). terebrans Lac, D. obesus (Mann.), D. rufipennis (Kirby), D. punctatus Lee, 



Tech. Series 17, Part I, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate II 



O 




THE GENUS DENDROCTONUS. 81 

D. simplex Lee, D. frontalis Zimm., eliminating Carphoborus bifurcus Eichh. Eich- 
hoff, 1881, pp. 125, brief revision, including D. micans (Kug.). Le Conte and Horn, 
1883, p. 523, in synoptic table. Dietz, 1890, pp. 27-28, revision and synopsis to 
include D. terebrans (Oliv.), D. rufipennis (Kirby), D. similis Lee, D. simplex Lee, 
D. approximatus n. sp., and D. frontalis; revised description of species, with syn- 
onymy, figs. 1-6, epistomata and antennae. Hopkins, 1894b, p. 280, author's extra, p. 7, 
sexual characters of D. terebrans (Oliv.), D. frontalis Zimm. Blandford, 1897, p. 143, 
synoptic table. Lovendal, 1898, pp. 86-87, PI. II, fig. 7, antenna, fig. 8, tarsus. 
Hopkins, 1902a, pp. 3-4, secondary sexual characters, statistics, etc. Hopkins, 1906b, 
pp. 143-147, larval mouthparts. 

DIVISION I. 

The distinctive characters common to the species of the first division 
are as follows : 

Adult. — Prothorax somewhat elongate and as broad as elytra; 
anterior dorsal half of elytra without long erect hairs. 

Pupa. — ^Vertex of head distinctly to faintly grooved, and with two 
prominent to small frontal spines or granules on or toward the vertex 
each side of groove. 

Larva. — Abdominal tergites 8 and 9 without dorsal plates. 

Galleries. — Egg galleries winding to straight; the eggs isolated or 
in approximate groups, but never in masses; larval mines exposed 
or concealed in inner bark. 

Subdivision A. 
(Species numbers 1 to 8, inclusive.) 

The distinctive characters common to the species of the first sub- 
division are as follows: 

Adult. — Body somewhat slender; prothorax with sides but shghtly 
narrowed and not constricted toward the head; elytra with second 
stria of declivity straight; second interspace not distinctly broader 
in middle: 

Sexes. — Female: Pronotum untJi transverse ridge across anterior 
area. Male: Pronotum without transverse ridge, but usually with 
anterior area broadly impressed. 

Pupa. — Elytra smooth; vertex of head faintly grooved and with 
two small, widely separated frontal tubercles. 

Larva. — Front with or without median convexity. 

Galleries. — Egg galleries winding; larval mines exposed or con- 
cealed. Pupal cells in outer bark; eggs isolated, never in groups. 

1. Dendroctonus brevicomis Le Conte. 
(PI. ni, fig. 1.) 

Adult. — Typical female: Length, 4.6 mm.; light brown. Elytral 
dechvity without long hairs. Head with front convex, slight eleva- 
tions each side of a faint median groove. Elytral rugosities fine, 
densely placed; striae obscure and but faintly or not at all impressed. 
79980—09 7 



82 



THE SCOLYTID BEETLES. 



Pronotum with a few reclining long hairs on the anterior half, remain- 
ing areas with very few and stout pubescence. Secondary sexual 
characters: Pronotum with transverse ridge extending across the 
anterior area to near the sternum. Elytral decHvity moderately 
smooth and sliining; interspaces finely and densely punctured; striae 
fine, with punctures scarcely visible. 

Typical female labeled 'Hype of revision," name label, "Hopk. 
4/18/02," second name label, "Hopk. Jan 16/08, Pinus ponderosa, 
Hopkins, collector, 3/24/99, Grant's Pass, Or., ? , Hopk. U. S. 34." 




Fig. 44. — Dendroctonusirevicomis: Egg galleries. (Original.) 

Type male in Le Conte collection, labeled "Cal.," examined by 
writer. 

Typical male: Length 4.6 mm. Agrees with female, except in the 
more prominent frontal tubercles each side of a distinct groove and 
a transverse impression instead of an elevation across the anterior 
area of the pronotum, and the elytral declivity slightly less shining. 

Typical male labeled 'Hype of drawing," name label, "Hopk. 
1/16/08, Pinus ponderosa, Hopkins, collector, 3/24/99, Grant's Pass, 
Or., $ , Hopk. U. S. 34." 



THE GENUS DENDEOCTONUS. 



83 



Variations. — Length, 3 to 5 mm.; color light brownish to nearly 
black; punctures of pronotum ranging from moderately coarse to 
very fine, elytral striae not at all impressed in some examples, in 
others the impression and punctures more distinct, but never as 
distinct as in the majority of D. harheri. The elytral rugosities" 
also vary from very fine to moderately coarse. The front varies 
greatly, from convex without tubercles to deeply grooved and with 
prominent tubercles. The epistomal process varies from the nor- 
mal concave form with angles elevated and lateral margins strongly 
oblique to flat with rounded apex and lateral margin suboblique. 

Distinctive characters. — 

/irnerBarrA ^ 
OurerBa. 



The adults of this species 
are at once distinguished 
from its nearest ally, D. 
harheri, by the finer rugo- 
sities of the elytral inter- 
spaces and the much less 
distinctly impressed strise. 

Revisional notes. — The 
labeled type in the Le 
Conte collection agrees 
with the description, ex- 
cept that the prothorax is 
not nearly twice as broad 
as long. It is certainly 
distmct from D. frontalis. 
The specimens in the Horn 
collection under D. fron- 
talis that evidently repre- 
sent part of the material 
on which Doctor Dietz 
based his revision, include 
one specimen of D. frontalis labeled with red disk, one speci- 
men of D. hrevicomis labeled "Cal.," one specimen of D. harheri 
labeled "Williams, Ariz. 7. 28," one specimen of D. arizonicusC^) 
"Williams, Ariz. 7. 28," and one specimen without locality label. 
One specimen was also received from Dietz, under D. frontalis, 
labeled "Arizona," which proved to be D. harheri. In 1898 D. hrevi- 
comis Leo. was not represented in the U. S. National Museum. There- 
fore it appears that up to 1899 there were only two specimens of the 
species in the large collections of the country. 

Pupa. — In addition to the divisional and subdivisional characters, 
the pupae range in length about the same as adults; the apices of the 
front and middle femora are smooth; abdominal tergites 3, 4, 5, and 




Fig. 45.— -Dendroclonus brevicomis: Bark showing, a, pupal 
cells; 6, exit burrows; c, pitch tubes. (From Webb.) 



84 



THE SCOLYTID BEETLES. 



6 with small pleural spines, 1 and 2 without distinct dorsal and lateral 
spines. Pupal type labeled "Hopk. U. S. 34." 

Larva (PL VIII, fig. 1). — In addition to "the divisional and subdivi- 
sional characters the larvas range slightly longer than the pupae. 
Epistonia distinctly elevated. Front without median convexity; 
clypeus with apex subacutely emarginate; thoracic segments with 
prominent sternal lobes; the sternellar lobes with distinct foot calli; 
apex of abdomen subtruncate. Larval type labeled '' Ilopk. U. S. 25a." 

Galleries (figs. 44, 45). — Egg galleries subtransversely mnding; 
eggs isolated; larvae concealed in inner bark; pupal cells iu outer bark. 

Distribution (fig. 46). — (Hopk. U. S.) California: McCloud, Badger, 
Chester, Wawona, Sterling, Summerdale, Yosemite, Ballard. Montana: 



N?t|:|f|-.j.,_ 


-T-=--T*^ 


V ^'^//> 


/.■■'■■■ '■':! .^>,.;-.>.- •. 




^^^^^^rT"'^ ^^ v^ ! \/ 


1 ■ ■'•■ ' •>' ■ ' ' — 


7 4AJ 

-^-j )x--i 


w 


^^ ''' ' 




\ '""X """'"' "^\ 


\ i'--- 1 


\\\ 


--\ y--^ 


'r''~^^^ \ 


vW 


\ f 


V \ 


\ I \ 




xj 



Fig. 40. — Dendroctonus brevicomis: Distribution map. (Original.) 

Mssoula. Oregon: Grants Pass, Joseph. Washington: Buckeye (near 
Spokane), Chelan. Idaho: Moscow, Smiths Ferry, Centerville, Stites, 
Kooskia (Harris Ridge), Pioneer (Grimes Pass), Garden Valley, 
Placerville, Cedar Mountains, Troy. 

Host trees. — Pinus ponderosa and P. lambertiana. 

Identified specimens. — Le Conte, 1; Horn, 1; Hopk, U. S., several 
hundred, including all stages and work. 

Bibliography and Synonymy. 

'Dendroctonus brevicomis Le Conte, 1876, p. 386, original description, synopsis, 
localities. Packard, 1887, p. 177, Le Conte quoted. Packard, 1890, p. 722, 
Le Conte quoted. Hopkins, 1899a, p. 395, reference. Hopkins, 1899b, pp. 13, 
20, 26, first records of habits, hosts, etc. Hopkins, 1901b, p. 66, habits, galleries, 
etc. Hopkins, 1902a, p. 3, name restored. Hopkins, 1902c, p. 21, note. Hop- 



THE GENUS DENDEOCTONUS. 85 

kins, 1903b, p. 281, mentioned. Hopkins, 1904, p. 18, habits, distribution, etc. 
Webb, 1906, bulletin, Pis. II, III, figs. 7-12, stages and work illustrated, full 
account of habits, life history, etc. Hopkins, 1908, pp. 162-163, depredations. 

Dendroctonus frontalis (not of Zimm.) Dietz, 1890, p. 32 (in part), California. 

Bendroctonus brevicornis Dietz, 1890, p. 32 (in part), California. 

2. Dendroctonus barberi n. sp. 

(PI. Ill, flg. 2.) 

Adult. — Type of species, female: Length, 4.5 mm.; very dark 
brown. Elytra and elytral declivity without long hairs. Head with 
front convex, with slight elevation each side of a faint median groove. 
Elytral rugosities moderately coarse and dense; strise distinctly 
impressed. Pronotum with a few long hairs on the anterior half of 
the lateral area, the remaining area with very fine and short pubes- 
cence. The secondary sexual characters are the same as in the 
preceding species. 

Type labeled ''Type No. 7444, U.S.N.M.," name label, "Hopk. 
1/15/08, ? , individual /, Barber & Schwarz, Collectors], Williams, Ar., 
7.6" ( = June7). i 

Male type: Length, 4 mm. Front with prominent tubercles each 
side of a distinct groove. Pronotum without transverse ridge across 
the anterior area, but with a broad transverse impression. The 
elytral declivity shining, with strial punctures distinct and interspaces 
slightly more rugose than in the female. 

Male type labeled " $ type," otherwise same as female, except type 
number. 

Variations. — Length 2.5 to 4.7 mm., color from brownish to black. 
The frontal and prothoracic variations are similar to those of D. 
Irevicomis. The elytra vary from rugosities moderately coarse and 
obtuse to distinctly coarse and acute, and striae from moderately to 
very distinctly impressed, and the punctures from obscure to distinct. 

Distinctive characters. — The adults of D. harleri are at once distin- 
guished from the next species by the absence of long hairs on the ely- 
tral declivity, and from D. hrevicomis, to which it is closely allied, by 
the prevailing coarser rugosities of the elytral interspaces and the 
distinctly impressed striae. Some systematists might concede these 
characters of no more than subspecific or varietal value, but it must 
be remembered that in this genus there is a close general resemblance 
of allied species arid that the characters which in other genera would 
be of special value in specific distinction are so variable and incon- 
stant in this as to be of no value. Therefore any constant or prevail- 
ing character of distinction, even if it does seem insignificant, is of 
vastly higher value than would otherwise be allowable, especially 
when it is supported by differences in physiological characteristics. 



86 



THE SCOLYTID BEETLES. 



Revisional note. — This species is not represented in the Le Conte 
collection. One specimen labeled "Williams, Ariz., 7.28, 151," and 
one without label found in the Horn collection, both under D. frontalis, 
and one specimen labeled "Ariz.," received from Doctor Dietz under 
D. frontalis, all belong to D. larheri. These were evidently before 
Doctor Dietz when he prepared his revision of D. frontalis. Hid. 

Pwpa. — In addition to the generic, divisional, and subdivisional 
characters, the apex of the front and middle femora is smooth; abdom- 
inal tergites 3 to 6 with very small dorsal, lateral, and pleural spines; 
1 and 2 without dorsal or lateral spines ; 3 to 6 with minute dorsal and 
lateral spines ; 7 and 8 smooth, and 9 with small pleural spines. Pupal 
type, labeled "Hopk. U. S. 5030" (m alcohol), differs from pupa of 
D. Irevicomis in the absence of lateral spines on abdominal tergites 
3 and 4; but these, with other pupal characters, are so variable that 




Fig. 47.— Dendroctonus barberi: Egg galleries. (Original.) 

not much reliance can be placed on any of them to separate closely 
allied species. 

Larva. — In addition to the generic, divisional, and subdivisional 
characters, the clypeus has the apex broadly emarginate instead of 
subacutely emarginate, as in D. hrevicomis. It also differs in the 
more rounded apex of the labrum and in the more distmctly rugose 
mandibles. Larval type labeled "Hopk. U. S. No. 5129" (mounted 
and alcoholic). 

Galleries (fig. 47). — In addition to the divisional and subdivisional 
characters, the egg galleries are usually distinctly transversely wind- 
ing, thus differing from D. hrevicomis; otherwise there is little 
difference. 

Distribution (fig. 48). — (Hopk. U. S.) Arizona: Williams, Flagstaff, 
San Francisco Mountams, Grand Canyon, Walnut Canyon, Dead Man's 



THE GENUS IDENDROCTOiSTtlS. 



Si 



Flat, Show Low, Santa Catalina Mountains. New Mexico: Vermejo, 
Santa Fe, Meeks, Capitan (Mountains), Cloudcroft. Texas: Davis 
Mountains. Colorado: Fort Garland and Monte Vista. Utah: Esca- 
lante and Panguitch. Additional localities from other collections. 
(H. & S.) Chiricahua Mountains, Ariz. 

Host trees. — Pinus ponderosa var. scopulorum (very common), P. 
edulis (rare), and Pseudotsuga taxifolia (rare, probably abnormal). 

Identified specimens. — Horn collection, 2; Dietz, 1; U. S. N. M., 
H. & S., 2, B. & S., 62; Hopk. U. S., more than 400, including all 
stages and work. 




Yia.^S.—Dendroctonusharhcri: Distribution map. (Original.) 



BlBLIOGRAPPIY AND SyNONYMY. 

Dendroctonus frontalis (not of Zimm.) Dietz, 1890, p. 32 (in part), Arizona. 
Dendroctonus arizonicus Hopkins, 1902a, p. 3 (in part), manuscript name only. 
Dendroctonus n. sp. Hopkins, 1904, pp. 42, 44, habits, host, distribution, etc. 
Dendroctonus brevicomis var. barheri Hopkins, 1906b, p. 147, PI. IV, fig. 9, anatomy 
of larval head, manuscript name. 

3. Dendroctonus convexifrons n. sp. 

(PI. Ill, flg. 3.) 

Adult. — Type of species, female: Length, 6 mm.; reddish-brown, 
shining. Ely tral declivity with long hairs. Head with front convex; 
without median frontal groove or tubercles, but with posterior impres- 
sion. -Elytral rugosities moderately coarse, but not densely placed, 
except toward the base ; striae faintly impressed and with rather coarse, 
indistinct punctures. Pronotum with long, erect hairs on the entire 



88 THE SCOLYTID BEETLES. 

lateral area, the remaining area with short, erect, and sparse pubes- 
cence. Secondary sexual characters: Pronotum with transverse 
ridge across the anterior area, extending to the sternum. Elytral 
declivity shining; striae very slightly impressed; punctures obscure; 
interspacial granules sparse and small, but distinct. 

Type labeled ''Type No. 7445 U.S.N.M.," name label, ''Hopk. 
1/16/08, Pinus fonderosa, Hopkins, Coir., 9/8/02, WiUiams, Ariz., 9 , 
Hopk. U. S. 1109." 

Male type: Length, 5.6 mm. Front convex and with faint anterior 
and posterior impressions, but without frontal groove or tubercles. 
Pronotum without transverse elevation. Elytral declivity less shin- 
ing; striae more impressed, with more distinct punctures and the inter- 
spacial granules distinctly coarser than in the female. 

Male type labeled " $ type;" otherwise same as female, except 
type number. 

Variations. — The length varies from 4 to 6 mm., with the average 
at about 5.5 mm. ; the color ranges from reddish to black, with red- 
dish-brown prevailing; front ranges from completely convex without 
median impression to more or less distinctly impressed, but never 
with frontal tubercles. The punctures of the pronotum vary as 
usual, and the dorsal area ranges from the absence of a longitudinal 
median space or line to a distinct elevated line. 

Distinctive characters. — The adults of I), convexifrons are at once 
distinguished from D. approximatus by the more slender form and 
shining appearance, and the prevailing convex front in both sexes, 
which latter character also distinguishes small examples of the species 
from large examples of D. arizonicus. The long hairs on the elytral 
declivity render it absolutely distinct from D. larheri, to which it 
appears to be more closely allied on account of pupal characters. 

Revisional notes. — This species is not represented in the Le Conte 
collection, but among the three specimens in the Horn collection, 
under D. ajjproximatus Dietz, there was one labeled "N. M.," which 
certainly must be referred to it. 

Pupa. — In addition to the divisional and subdivisional characters, 
the front and middle femora are smooth or with minute apical 
granule; abdominal tergites 1 and 2 without dorsal spines but with 
one or two lateral granules ; 2 to 6 with very small lateral spines, and 
3 to 6 with very small dorsal and lateral spines, becoming more 
prominent toward the sixth; 7 and 8 smooth and 9 with widely 
separated caudal spines. Pupal type labeled "Hopk.- U. S. No. 
5090." 

Larva. — In addition to the divisional characters, the front has a 
median smooth, shining convexity; mandibles distinctly rugose on 
lateral area toward base; labrum with apex broadly rounded and 



THE GENUS DENDEOCTONUS. 



89 



clypeus with apex broadly emarginate. Sternellar lobes of thoracic 
segment without foot calli. Larval type labeled ''Hopk. U. S. No. 
5078." 

Galleries (fig. 49). — ^Egg galleries longitudinally winding; eggs 
isolated; larval mines exposed in inner bark; pupal cells in outer bark. 

The galleries of this species are at once distinguished from those 
of I), harieri and D. approximatus by the exposed larval mines. 




Fig. 49. — Bendroctonus convcxifrons: Egg galleries and larval mines. (Original.) 

These three species usually infest the same tree, and often all of 
their galleries are represented in a piece of bark. 

Distribution (fig. 50). — (Hopk. IT. S.) Arizona: WilHams, Flagstaff, 
Show Low, Paradise, Santa Catalina Mountains, and Chiricahua 
Mountains. Colorado: Las Animas County, La Veta, Fort Garland, 
and Monte Vista. New Mexico: Cloudcroft, Meeks, Capitan, Fort 



90 



THE SCOLYTID BEETLES. 



Wingate Military Reservation, Vermejo, Lincoln National Forest, 
and Sierra Blanca Mountains. Utah: Escalante, La Salle National 
Forest, and Panguitch Lake. Additional locality from other collec- 
tions: (B. & S.) Las Vegas, N. Mex. 

Host trees. — Pinus ponderosa var. scojjulorum (very common) 
and P. chihuahuana (rare). 

Identified specimens. — Hopk. U. S., more than 100 specimens, 
including all stages and work; Horn, 1 specimen under D. approxi- 
matus, labeled "N. M."; U.S.N.M., B. & S., 7 specimens. Las Vegas, 
N. Mex., 17-8, No. 164, and Williams, Ariz., 5-6, 5-7, and 5-10. 

Bibliography. 

Dendroctonus approximatus Dietz, 1890, p. 31 (in part), New Mexico. Schwarz, 
1902, p. 32 (in part). 




Fig. 50. — Dendroctonus convexifrons: Distribution map. (Original.) 



4. Dendroctonus frontalis Zimmerman. 

(PL in, flg. 4.) 

Adult. — Typical female: Length, 3.6 mm.; reddish-brown. Ely- 
tral dechvity with long hairs. Front with a moderately prominent 
tubercle each side of a distinct median groove. Elytral strise dis- 
tinctly punctured; interspacial rugosities moderately coarse, obtuse, 
not very densely placed, and not very distinctly coarser toward the 
base and vertex. Pronotum with a few long hairs on the anterior 
section of the lateral area. Secondary sexual characters: Pronotum 
with transverse ridge across the anterior area, the elytral dechvity 
shining and with finely granulate interspaces, the strise distinctly 
impressed, but the punctures obscure. 



THE GENUS DENDEOCTONUS. 



91 



Typical female labeled 'Hype of revision, type of drawing," name 
label, ''Hopk. 4-18-02, 9 , H. S. 58, S. C." 

Type in Le Conte collection labeled "Type 1, B. frontalis (Fab.) 
Zimm.," red disk (= CaroUnas). 

Male type: Length, 3.5 mm. Front with prominent frontal 
tubercle each side of a broad, deep frontal groove. Pronotum 
without transverse elevation or impression across the anterior area. 
Elytral decHvity with striae more impressed and the interspacial 
granules coarser and more sparse than in the female. 




Fig. 51. — Bcndroctonus frontalis: Egg galleries and larval mines, a, Entrance; 6, entrance burrow; c, egg 
gallery; d, normal larval mine; e, abnormal larval mine;/, terminal; g, ventilating burrows. (Original.) 

Typical male labeled with name, "Hopk. 1-16-08, ^ , U.S.N.M. 
37, sp., N. C." 

Variations. — The length varies from 2.5 to 4 mm., with the average 
at about 3.2 nim. The color ranges from light brown to nearly black; 
the epistoma, front, prothoracic punctures, and elytral rugosities vary 
as usual. The anterior area of the pronotum, which is glabrous in 
the typical females and males, usually has a few long hairs. 

Distinctive characters. — This species is distinguished from D. ari- 
zonicus, its nearest ally, by its smaller size, broader pronotum, with 
finer punctures, and finer and less acute rugosities of the elytra. 

Revisional Twtes. — The type series in the Le Conte collection is repre- 
sented by three specimens — one labeled "Type 1, D. frontalis (Fab.) 



92 



THE SCOLYTID BEETLES. 



Zimm./' red disk ( = Carolinas) , which should stand as the type, and 
the other two labeled ''Type 2" and "Type 3," same locality. 
There are also two specimens, one labeled "Specimen 4, Lake Supe- 
rior," and the other labeled "Specimen 5, Va." Both of these agree 
with the West Virginia form. In addition, there are three specimens 
labeled with an orange disk ( = Georgia) , one with light green disk 
( = Middle States) , all of which were doubtless before Le Conte when 
he drew up his revised description. 

The series in the Horn collection, which was evidently before Doc- 
tor Dietz when he drew up his revised description, includes but one 



||iililllillliifii!iiriiniii|, 




c 



N 



>v 



Fig. 52.—Dendrocto- 
nus frontalis: Ter- 
mination of egg gal- 
leries. (Original.) 




Fig. 53.~Dendroctonus froii- 
talis: Beginning of egg galler- 
ies; a, in liviag bark; 6, in 
dying bark; c, marked on 
surface of wood (white area 
represents normal appear- 
ance of wood preserved by 
resin). (Original.) 




Fig. 54. — Dendroctonus frontalis: 
Bark showing, a, pitch tubes; 6, 
entrance biurow; c, egg gallery; 
d, ventilating burrow; e, pupal 
cells; /, exit burrows; g, inner 
liark; A, outer bark. (Original.) 



specimen labeled with red disk ( = Carolinas) , The specimens men- 
tioned, together with two or three specmiens in the United States 
National Museum, were probably all that were in the larger collec- 
tions of the country up to 1890. 

The form which in 1891 and 1892 extended northward from the 
normal range of the species into Virginia and West Virginia is repre- 
sented in the revision series by specimens labeled "Hopk. W. Va. 9 
individual 1" and "Hopk. W. Va. S individual 1." The West Vir- 
ginia female differs from the typical South Carolma female in a more 



THE GENUS DENDEOCTONUS. 



93 



rugose front, prothorax slightly narrow in front, with a few short and 
long hairs on sides and the punctures of anterior dorsal surface 
coarser toward base and fine toward anterior margin; elytral rugosi- 
ties more acute; strial punctures coarser and more distinct; elytral 
declivity with longer and more numerous hairs. The West Virginia 
male differs from the typical North Carolina male type in the more 
shining front, with punctures more distinct, while the pronotum and 
elytra show the corresponding differences mentioned under the 
female individual 1. At one time it seemed to the writer that this 




Fig. 55. — Dendroctonus frontalis: Old egg galleries 
in living tree, with surrounding callus of new 
wood. (Original.) 




Fig. 56.~Dendroctonus frontalis: Egg gal- 
lerjr in living tree marked on surface of 
wood six j^ears before block was cut 
from tree, a, Mark of gallery on origi- 
nal surface; 6, resinous wood; c, surface 
scar six years later; d, original surface 
or 7-year-old annual laj^er of wood; e, 
six subsequent annual layers of wood 
formed over original wound. (Origi- 
nal.) 



northern form was worthy of specific 

distinction, and the manuscript name 

of D. pinicida was proposed for it, but 

it was later found that some southern 

examples showed the same, and even 

greater, variations from the type. Therefore, since the species had 

disappeared from its northern range, it was decided that it would not 

be advisable to recognize it as specifically distinct. 

Pupa. — In addition to the divisional and subdivisional characters 
the apices of the front and middle femora are armed with small 
apical spines or granules. Abdominal tergites 1 to 6 without pleural 
spines; 1, 2, and 3 without distinct dorsal and lateral spines; 4 to 6 
with a pair of dorsal spines and one or two lateral ones; 7 and 8 with 



94 



THE SCOLYTID BEETLES. 



a few dorsal granules, and 9 with medium-sized caudal spines. Pupal 
type labeled ''Hopk. U. S. No. 2968." 

Larva, — In addition to the divisional characters, the front has a 
distinct, median, smooth, shining convexity produced toward vertex ; 
clypeus short, broad, with apex deeply emarginate; prothoracic seg- 
ments very large and sternellar lobes with distinct foot calli. Apex 
of abdomen truncate. Larval type labeled " Hopk. W. Va. No. 5976." 

The frontal convexity is more rugose in some specimens than in 
others. 

Galleries (figs. 51-56). — Egg galleries sublongitudinal, winding; the 
eggs isolated; larval mines exposed; pupal cells in outer bark. 

Distribution (fig. 57). — (Hopk, W. Va.) West Virginia: Hamp- 
shire, Monongalia, Hardy, Pendleton, Randolph, Pocahontas, Tucker, 
Kanawha, Raleigh, Greenbrier, and Wood counties. Virginia: Port 




Tig. 57. — Dendroctonus frontalis: Distribution map. (Original.) 

Republic. Work observed m southern Pennsylvania, Maryland, and 
Virginia (author). (Hopk. U. S.) Alabama: Calhoun, Montgomery. 
ArJcansas: Hampton. Georgia: Clyo, Thomasville, Demorest. Louisi- 
ana: Singer, Wilson. Maryland: Near Cumberland. North Carolina: 
Tryon, Pisgah Ridge, Fletchers, Boardman, Pink Beds, Biltmore. 
South Carolina: Chicora (Pregnalls). Tennessee: Ducktowh. Texas: 
Call, Deweyville, Earbyville, Beaumont. Virginia: Green Bay, Glen 
Allen, Auburn Mills, Virginia Beach. District of Columbia: Washington. 
Additional localities rejyresented in other collections: (Le Conte) Lake 
Superior, Michigan. (There may have been some mistake in regard to 
the labeling of this specimen, since it is not likely the species will ever 
be found that far north.) (H. & S.) Haw Creek, Florida; Cobbs Island, 
Virginia; (A. M. N. H.) Black Mountains and Mount Graybeard, 
North Carolina; (Barber) "western Indian territory" [Oklahoma]. 



THE GENUS DENDKOCTOlSrUS. 95 

Host trees. — Pinus strohus, P. txda, P. rigida, P. virginiana, P. pun- 
gens, P. ecJiinata, P. glabra, P. palustris, Picea rubens, and P. excelsa. 

Identified specimens. — Le Conte collection, 9; Horn, 1; U.S.N.M., 
H. & S., 5; D. A., 7; Barber, 2; Hopk. W. Va., 68; Hopk. U. S., more 
than 150, including all stages and work. 

Bibliography and Synonymy. 

Dendroctonus frontalis Zimmerman, 1868, p. 149, original description, type, locality, 
Carolina. Le Conte, 1868, p. 173, synopsis and reference to p. 149. Le Conte, 
1876, p. 386, revision, synopsis, synonymy excluded, bibliography, localities. 
Packard, 1887, p. 177, Le Conte quoted. Packard, 1890, p. 722, Le Conte quoted. 
Dietz, 1890, p. 32, in part, fig. 6, antennaand epistoma. Hopkins, 1892a, pp. 64-65, 
depredationsinW. Va. Hopkins, 1892b, p. 353, importation of enemy. Schaufuss, 
1892, p. 316, introduction of enemy. Hopkins, 1893a, pp. 187-189, habits, etc. 
Hopkins, 1893b, p. 143, No. 77 and index, habits, host, distribution, enemies. 
Hopkins, 1893c, p. 213, No. 301, same as 1893b. Hopkins, 1893d, pp. 123-129, 
habits and enemy. Lintner, 1894, p. 292, reference to Hopkins. Hopkins, 
1894a, pp. 71-76, same as Hopkins 1893d. Hopkins, 1894c, p. 348, insects, birds, 
and forests. Lintner, 1895, same as 1894. Hopkins, 1896, pp. 246-250, disap- 
pearance in W. Va., occurrence in N. C. Hopkins, 1897a, pp. 29-41, importation 
and distribution of enemy. Hopkins, 1897b, pp. 35-36, enemy, etc. Hopkins, 
1897c, pp. 79, 94, 95, PL I, dead trees, fig. IV, Clerus formicarius L., discussion of 
habits, etc., pp. 147, 151, reprint from 1896. Chittenden, 1897, pp. 67, 75, fig. 
43, adult, destructive habits. Schwarz, 1898, p. 81, habits and disappearance. 
Hopkins, 1898b, pp. 104-105, habits, etc. Hopkins, 1899a, pp. 394-414, etc. (see 
index), full account, all stages and work illustrated and described, natural ene- 
mies, hosts, distribution, bibliography, etc. Hopkins, 1899b, pp. 11, 13, 14, 
reference to habits. Hopkins, 1899c, p. 343, disappearance in W. Va. due to freez- 
ing. Chittenden, 1899, pp. 55, 56, fig. 5, habits, etc. Ulke, 1902, pp. 36, 56, host, 
habits, etc., in D. C. Hopkins, 1903a, p. 59, occurrence and habits in southern 
States. Hopkins, 1903b, pp. 270, 275, figs. 26, 27, 28-32, stages and work, with 
account of habits, life history, etc. Hopkins, 1904, pp. 41, 42, 44, PL I, fig. 2, 
PI. VII, figs, a, b, stages and work, account of distribution, habits, etc. Felt, 

1905, p. 6, reference. Hopkins, 1906c, p. 81, mentioned in comparison. Webb, 

1906, pp. 20, 22, mentioned. Hopkins, 1908, p. 163, depredations. 
Bostrichus frontalis (not of Fab.) Zimmerman, 1868, p. 149, synonymical reference. 

Le Conte, 1868, p. 173. 
Dendroctonus pinicida Hopkins, 1902a, p. 3, manuscript name only. 
Dendroctonus frontalis Zimm. var. destructor Hopkins, 1902b, p. 21, note. Hopkins, 

1902c, p. 20, habits, etc. 
Dendroctonus brevicornis Dietz, 1890, p. 32 (in part). 

5. Dendroctonus arizonicus n. sp. 

(PL III, fig. 5.) 

Adult. — Type of species, female: Length, 3,7 mm., dark b^o^^^l. 
Elytral decUvity with long hairs confined to declivity and posterior 
lateral areas. Head with front convex, shining, and with distinct 
frontal tubercle each side of a broad median groove. Elytral striie 
distinctly punctured; the interspaces with subacute, moderately 
coarse rugosities, distinctly coarser toward the base and vertex. 
Pronotum with long erect hairs on the anterior half of the lateral 
area. Secondary sexual characters same as in D. frontalis. 



96 THE SCOLYTID BEETLES. 

Type labeled "Type No. 7446, U.S.N.M./' name label, "Hopk. 
1/15/05, 9, Barber & Schwarz Colllectors], Williams, Ar., 7.6" 
( = June 7). 

Male type: Length, 3.3 mm. Front with prominent tubercles each 
side of a deep median groove. Pronotum with transverse impression 
and faint transverse elevation across anterior area. Elytral declivity 
less sliining; striae more distinctly impressed and punctured and the 
interspaces more convex and distinctly rugose than in the female. 

Male type labeled, name label, "Hopk. Jan. 15/08, Pinus ponderosa, 
Webb, Collr., 8-22-04, Flagstaff, Ariz., S , Hopk.U. S. No. 5118." 

Variations. — Length 3.5 to 3.9 mm., average about 3.6 mm.; color 
from light brown to black. The short hairs of the lateral area 
of the elytra range from obscure to distinct, and the long hairs of 
the elytral declivity range from short and sparse to very long and 
numerous. The variation in the size of the punctures of the pronotum 
is less marked than in other species. 

Distinctive characters. — The coarse punctures of the pronotum, and 
especially the very coarse shallow rugose ones, together with the 
coarser rugosities of the elytra, serve to distinguish specimens of 
tliis species from D. harieri, and the absence of short reclining hairs 
on the pronotum with the more distinctly impressed elytral striae 
and less evident short and long hairs on the elytra serve to distinguish 
it from D. mexicanus, wliich is its nearest ally. It is also distinguished 
from D. frontalis by its larger size, coarser punctures of the pronotum, 
and coarser rugosities of the elytra, as it is from small examples of D. 
convexifrons by the grooved front, and from small examples of D. 
approximatus by the impressed elytral striae and the shorter, more 
yellow, and less numerous hairs of the declivity. 

Revisional notes. — This species is not represented in the Le Conte 
collection and not in the Horn collection unless it is one specimen 
found under D. frontalis labeled "Wilhams, Ariz., 7.28, 152," which 
the writer has not seen since D. arizonicus has been recognized as a 
distinct species. If this specimen is D. arizonicus, it was evidently 
before Doctor Dietz when he prepared his revised description of D. 
frontalis. 

Pupa. — In addition to the generic and divisional characters, the 
apices of the front and middle femora have each a small spine. Ab- 
dominal tergites 2 to 6 with small pleural spines and 4 to 6 with small 
dorsal, lateral, and pleural spines, increasing in size toward 6; 7 and 
8 with a pair of dorsal granules, coarser on 7. Pupal type labeled 
"Hopk. U. S. No. 3129." 

Larva. — In addition to the divisional characters, the front has a 
median transverse and rugose convexity produced toward apex; cly- 
peus with apex broadly emarginate; labrum broad, mth apex broadly 
rounded. Thoracic segments with foot caUi on sternellar lobes. 
Larval type labeled "Hopk. U. S. No. 5156." 



THE GENUS DE]SrDEOCTO]SrUS. 



97 



Oallerles. — While the galleries of this species have not been defi- 
nitely recognized from those of D. harheri, with which they are nearly 
always present, it is evident that they are quite similar in general 
character to those of D. harheri, especially in the concealed larval 
mines. 

Distrihution (fig. 58). — (Hopk. U. S.) Arizo7ia: Williams, Flag- 
staff, Santa Catalina Mountains (rare). 

Host tree. — Pinus ponderosa var. scopulorum. 

Identified specimens. — Horn collection, 1 (?), B. & S., 2; Hopk. 
U. S., more than 50 specimens, including adults, larvae, and pupse. 

Bibliography. 
Dendroctonus arizonicus Hopkins, 1902a, p. 3 (in part), manuscript name. 



7T--.i. 





vT- ••■""•••■"' ■ • • ~-r 1 L—r 

.;Y. • :2}^:-l ■ : ■; :■ r — i V ^ 






Fig. 58. — Dendroctonus arizonicus: Distribution map. (Original.) 

6. Dendroctonus mexicanus Hopkins. 
(PI. Ill, fig. 6.) 

Adult. — Type of species, female: Length, 4 mm.; dark brown. 
Elytra with long hairs on posterior half and declivity. Head with 
front convex, shining, with small frontal tubercles each side of the 
short, shallow groove; elytral striae distinctly impressed, but not dis- 
tinctly punctured; elytral rugosities distinctly coarser toward the 
base and vertex, and with an evident row of acute rugosities on the 
first interspace. Pronotum clothed with short reclining hairs on 
entire lateral area. Secondary sexual characters: Pronotum with 
transverse ridge across the anterior area; elytral declivity shining, 
striae distinct, obscurely punctured; interspaces roughened with 
sparsely exposed granules, becoming coarser toward the vertex in 
lateral area. 

^ 79980— 09 8 



98 



THE SCOLYTID BEETLES. 



Type labeled ''Type No. 7518, U.S.N.M., individual l/'name label, 
''n. sp., Hopk., 4/18/02,'' name label, "1/15/08, A. L. Herrera, col- 
lector, 9 , Sacramento, Amecameca, Mexico." 

Male type: Length, 4 mm.; front with moderately prominent fron- 
tal tubercles each side of the distinct groove. Pronotum with rather 
distinct transverse elevation across the entire area similar to that of 
the female; elytral declivity more uniformly convex; striae more 
impressed and deeply punctured and interspaces more convex and 
distinctly rugose, 

Male type labeled ''Type No. 7518, U.S.N.M., individual 5," remain- 
ing labels same as on female except sex label. 

Variations. — Length from 3 to 4.5 mm.; aver- 
age about 3.8 mm.; color from brown to black. 
The prescutal ridge is more prominent in some 
females than in others and is present in some 
males and absent in others. 

Distinctive characters. — This species is more 
closely allied to D. arizonicus than to any of the 
other species of the division to which it belongs 
and is distinguished from it by the presence of 
short, reclining hairs on the pronotum, less dis- 
tinctly impressed elytral strise, and more evident 
short pubescence of the elytra. 

Revisional notes. — It is evident that this spe- 
cies was not represented in the material studied 
by Blandford. The size comes near that of his 
D. adjunctus, but the characters as given by 
him to distinguish this species from D. paraUel- 
ocollis at once distinguish it from D. mexicanus. 
The pupa and larva have not been seen. 
Gallery (fig. 59). — A small section of the gal- 
lery, evidently of this species, was received with the specimens of 
adult from Prof. A. L. Herrera. This indicates a winding egg gallery, 
with the eggs isolated, the larval mines concealed in the inner bark, 
and the pupal cells in the outer bark. 

Distribution (fig. 60) and host trees. — The specimens received at dif- 
ferent times from Prof. A. L. Herrera and Dr. S. J. Bonansea were 
evidently from Pinus teocotl, P. lejo])hiTla, and P. ayacahuite in Ame- 
cameca, Michoacan, and Tacubaya, Mexico. 

Identified specimens. — Thirty-six specimens of adults and one speci- 
men of work were identified for Professor Herrera and Doctor Bonan- 
sea, a set of which were returned to them and the remainder retained 
for the forest insect collection of the Bureau of Entomology. 




Fig. 59. — Dendroctonus mex- 
icanus: Section of egg gal- 
leries. (Original.) 



THE GENUS DENDROCTONUS. 
Bibliography. 



99 



Dendroctonus viexicanus Hopkins, 1906c, p. 80, original description, both sexes, 
host, etc. 




Fig. go. — Dendroctonus mexicanus: Distribution map. (Original.) 

7. Dendroctonus parallelocollis Chapuis. 
(PI. Ill, flg. 7.) 

Adult. — Tyi^ical female: Length, 6.1 mm., deep black; elytral decliv- 
ity with long hairs; head with front convex, shining, without dis- 
tinct frontal tubercles each side of the shallow median groove; elytral 
strise not impressed or distinctly punctured except at vertex; elytral 
rugosities obtuse, rather densely placed, moderately coarse toward 
suture, fine and less evident toward and on the lateral area; striae 
obscure, especially on the lateral area. Pronotum with numerous 
short', reclining hairs and with very long, erect hairs on the entire 
lateral area, punctures distinct. Secondary sexual characters: Pro- 
notum with transverse ridge across the anterior area ; elytral declivity 
subconvex, shining; strise distinct, slightly impressed, punctures 
rather distinct; interspaces with a few granules and sparse punctures, 
the lateral areas more rugose and coarsely punctured. 

Typical female labeled, name label, "Hopk., January 16/08, 
Michoacan, Mexico, A. L. Herrera, collector, 9 , Hopk. U. S. 2896b." 

Typical male: Length, 6 mm. Head with front convex and with 
prominent frontal tubercles each side of a deep groove. Pronotum 
with transverse impression across anterior area; elytral declivity 



100 



THE SCOLYTID BEETLES. 



subconvex; striae distinct, slightly impressed, and the interspaces 
more distinctly rugose than in female. 

Typical male labeled, name label, ''Hopk. 1/16/08, type of draw- 
ing, Pinus, Michoacan, Mexico, A. L. Herrera, collector, $ , Hopk. 
U. S. 2896a." 

Variations. — Length from 5 to 6.1 mm., average about 5.5 mm.; 
color, dark brown to black, with the usual variation in epistoma, pro- 
notal punctures, pubescence, and median line. 

Distinctive characters. — This species is distinguished from D. 
approximatus by the noticeably less shining and more pubescent pro- 

notum, the deeper punctures, the 
noticeably and constantly less im- 
pressed elytral* striae, and the less 
distinct punctures. 

Revisional notes.- — Agrees with 
original description, except that 
the specimens before the writer are 
smaller, the type being given as 6§ 
mm. The median line is flat in all 
but one specimen, in which it is im- 
pressed toward the anterior and 
posterior margin. In Blandford's 
revision the length of the tyi^e is 
given as 6.3 mm. Blandford states 
that it differs from D. approxima- 
tus Dietz by its smaller size, more 
elongate shape, more shining and 
strongly punctured prothorax, and 
indistinct elytral striae. The last dif- 
ference holds, but the others do not. 
In the writer's reference to this 
species (1905a, p. 81) it was con- 
sidered to be the same as D. approximatus Dietz, but I am now con- 
vinced that the two are specifically distinct. 
The pupae and larvae have not been observed. 

Galleries (fig. 61). — A short section of the egg gallery, from which 
an adult was taken, indicates that it is quite similar, in its winding 
character and the absence of exposed larval mines, to that of D. 
approximatus. 

Distribution (fig. 62). — Recorded from Mexico by Chapuis (1869) 
and Blandford (1897). Specimens were received with D. mexicanus 
from Prof. A. L. Herrera, labeled Mexico and Michoacan, Mexico, 
with his statement that it destroys the forests of lonacatapec, Morelos, 
and many regions of Mexico. 




Fig. 61. — Dendroctonus parallelocollis: Section of 
egg gallery. (Original.) 



THE GENUS DENDKOCTONUS. 



101 



Host trees. — According to evidence from correspondence with 
Professor Herrera, it would appear that this species, like D. mexi- 
canus, attacks Pinus teocotl (?), P. lejopMlla, and P. ayacahuite, 
although there is no definite statement to that effect. It is quite 
evident, however, that this species is associated with D. mexicanus 
in the same manner as D. approximatus in its association with D. 
arizonicus and D. harheri. 

Identified specimens. — Ten specimens of adults and one specimen of 
work identified for Prof. A. L. Herrera. 




Fig. C2. — Dendroctonus parallclocollis: Distribution map. (Original.) 

Bibliography. 

Dendroctonus parallelocollis Chapuis, 1869, p. 36, original description. Chapuis, 
1873, p. 244, reprint. Blandford, 1897, p. 147, synopsis, revision, distribution, 
bibliography. Hopkins, 1906c, p. 81, systematic notes. 

8. Dendroctonus approximatus Dietz. 

(PI. IV, fig. 8.) 

Adult. — Typical female: Length, 6.5 mm., reddish brown; elytral 
declivity with long hairs; head with front convex, shining, and with 
obscure frontal tubercles each side of short, inedian groove; elytral 
striae not impressed or distinctly punctured; elytral rugosities rather 
coarse, sparse; striae on lateral area rather distinct; pronotum with 
long hairs on anterior two-thirds of lateral area, but not on posterior 
section. Secondary sexual characters: Pronotum with transverse 
ridge across the anterior area; elytral declivity subconvex, shining; 



102 



THE SCOLYTID BEETLES. 



striae very distinct, impressed, and distinctl}' piinctiiredj interspaces 
sparsely tuberculate, and with a few fine punctures; lateral area 
shining, with a few subacute tubercles, and rather coarse, distinct 
punctures. 

Typical female labeled, name label, "Hopk. 1/16/08, 9 , individual 
c, Barber & Schwarz, Coll[ectors], Flagstaff, Ar. 2.7" ( = July 2). 

Type of species, 9 , in Horn collection, A.E.S., Philadelphia, 
labeled "Type D. approximatus n. sp., Colorado]." 

Male type: Length, 5.7 mm. Head 
with front convex, shining, a prominent 
frontal tubercle each side of a deep 
groove. Pronotum with broad impres- 
sion across the anterior area; elytra! de- 
clivity same as female, except that the 
interspaces are more densely rugose. 

Male type labeled, name label, "Hopk. 
1/16/08, ^ type of revision, type of 
drawing, Williams, Ar., 7.6" ( = June 7) 
" $ , Barber & Schwarz, Coll[ectors]." 

Variations. — The length varies from 4 
to 7.4 mm., the average about 6 mm. 
The color ranges from reddish-brown to 
black. The epistomal, frontal, pronotal, 
and elytral sculpture and vestiture vary 
as usual. The greatest variation is in 
the strial and interspacial punctures of 
the declivity. 

Distinctive characters. — The characters 
which distinguish this species from D. 
parallelocollis , its nearest ally, are the 
noticeably more shining and less pubes- 
cent pronotum, with the punctures finer 
and more shallow, the elytral striae con- 
stantly more impressed, and the punc- 
tures distinct. 

Revisional notes. — The original description (Dietz, 1890, p. 31), was 
based on "four specimens, two males and two females, from New 
Mexico and Colorado in Doctor Horn's collection." The specimen 
labeled type in the Horn collection is a female from Colorado, and 
agrees with the description, but one specimen labeled "N. M." is a 
female of D. convexifrons Hopk., and one other specimen with the 
type, but without locality label, is quite a different thing from either 
of the other two. The specimen has not been examined since the 
more distinctive characters of D. monticolx Hopk. and D. ponderosss 
Hopk. have been recognized, but it evidently belongs to one of these 
species. The fourth specimen mentioned by Dietz was not in the 




i 



Fig. 63. — Dendroctonus approximatus: 
Single egg gallery. (Original.) 



THE GENUS DENDEOCTONUS. 



103 



collection when examined by the writer on January 12, 1900. The 
reference in the original description to the front of the head "coarsely 
granulated, channeled" applies to the type, and at once distinguishes 
it from the other two specimens which are without a median frontal 
channel or groove. The "strong transverse impression about one- 
fourth from the an- 
terior margin" of 
the prothorax is as- 
sociated with the 
more distinct trans- 
verse elevation 
which the author 
failed to mention. 
In the writer's re- 
marks (Hopkins, 
1905, p. 81) under 
D. paraUelocoIlis it 
is stated that D. 
paraUelocoIlis Chap . 
and D. approxima- 
tus Dietz are evi- 
dently the same, 
but further com- 
parative studies 
have convinced 
him that they are 
specifically distinct. 

Pupa. — In addi- 
tion to the generic, 
divisional, and sub- 
divisional charac- 
ters, the apex of the 
front and middle 
femora have each a 
minute subapical 
s p in e ; abdominal 
tergites 1 to 6 with 
small pleural spines, 
1 without distinct 
dorsal and lateral spines; 2 to 6 with distinct dorsal and lateral 
spmes, increasing in length and prominence to and including tergite 
6; 7 and 8 smooth; 9 with long pleural spine. Pupal type labeled 
"Hopk. U. S. No. 5777." 

Larva, — In addition to generic and divisional characters, the front 
has a broad, but not prominent, median elevation; clypeus short, 
broad, with apex truncate; prothoracic segments with prominent 




Fig. 64. — Dendroctonus approximatus: Egg galleries. (Original.) 



104 



THE SCOLYTID BEETLES. 



sternellar lobes, each with a distinct foot callus. Larval type labeled 
"Hopk. U. S. No. 5024." 

Galleries (figs. 63, 64). — Egg galleries sublongitudinal, branched, 
slightly winding; eggs isolated; larval mines concealed; pupal cells in 
outer bark. 

Distribution (fig. 65). — (Hopk. U. S.) Colorado: Brookville, Glen 
Haven, Las Animas County, La Veta, Monte Vista, and Palmer Lake. 
VtaJi: Escalante, Kamas, Panguitch. New Mexico: Capitan Moun- 
tains, Cloudcroft, Lincoln National Forest, Santa Fe. Arizona: 
Black Mesa Forest Reserve, Chiricahua Mountain^ Flagstaff, Para- 
dise, Rincon Mountains, Santa Catalina Mountains, Show Low, 
Tucson. Additional locality from correspondence: Glenhaven, Colo. 




Fig. 05. — Dendroctonus approximatus: Distribution map. (Original.) 

Host trees. — Pinus ponderosa scopulorum, (common), P. arizonica 
(rare), and P. cTiihuahuana (rare). 

Identified specimens. — Horn, A. E. S., 1 specimen, the type; 
U.S.N.M., 31; E[opk. U. S., more than 200 specimens, including all 
stages and work. 

Bibliography and Synonymy. 

Dendroctonus approximattis Dietz, 1890, p. 31, fig. 5, antenna and epistoma; original 
description (applies to type only). Blandford, 1897, p. 147, reference to charac- 
ters. Hopkins, 1899a, p. 392, fig. LVIII, reference. Schwarz, 1902, p. 32 (in 
part), destructive to pine at Flagstaff, Ariz, (see also convexifrons) . Wickham, 
1902, p. 310 (in part), list, Dietz quoted. Hopkins, 1903a, p. 60, reference to 
habits, etc. Hopkins, 1903b, p. 281, mention. Hopkins, 1904, p. 44, habits, dis- 
tribution, hosts, etc. Hopkins, 1905, p. 11, some distinctive characters. Hopkins, 
1906c, p. 81, referred to D. parallelocollis. 

Dendroctonus (parallelocollis) var. approximatus (Dietz) Fall, 1907, p. 218, list, 
localities. 



THE GENUS DENDROCTONUS. 105 

Subdivision B. 
(Species numbered 9 to 11, inclusive.) 

The distinctive characters common to the species of the second sub- 
division are as follows : 

Adult. — Body somewhat stout, prothorax with sides distinctly nar- 
rowed and constricted toward the head; elytral declivity with second 
stria distinctly curved, second interspace broader toward middle, nar- 
rowed toward apex. 

Sexes. — Female: Elytral declivity with interspaces smoother and 
more shining, mandibles less stout. Male: The reverse. 

Pupa. — Elytral pads roughened with granules, vertex of head 
deeply grooved and with two prominent forward curved frontal spines 
toward the vertex; abdominal tergites 2 to 6 with long and prominent 
pleural spines. 

Larva. — -Front with prominent transverse and rugose elevation, 
more prominent toward the sutures. 

Galleries. — Pupal cells in inner bark, eggs in approximate groups, 
egg galleries slightly winding to straight, larval mines always exposed 
in inner bark. 

9. Dendroctonus monticolae Hopkins. 

(PI. IV, fig. 9.) 

Adult. — Type of species, female: Length 5.6 mm., light brown; 
elytral declivity with a few long hairs; head with front convex, with- 
out median elevation or groove, but with faint posterior impression; 
elytral rugosities coarse, rather dense, coarser on dorsal area; striae 
not distinctly impressed, except toward suture, punctures small. 
Pronotum with the anterior area transversely impressed, but with 
moderately long hairs on the lateral area, slightly longer on the ante- 
rior section. Secondary sexual characters : Elytral declivity convex, 
subopaque; striae distinct, and impressed, with fine obscure punc- 
tures; interspaces with rather coarse granules, in approximate rows; 
lateral area with obscure punctures and subacute rugosities. 

Type labeled "No 7447 U.S.N.M.," name label, "Hopk. 1/16/08, 
Pinus monticola, Hopkins, collector, 5/27/99, Kootenai, Idaho, 9 , 
Hopk. U. S. 205." 

Male type: Length 4.9 mm. Head with front less distinctly punc- 
tured and more rugose than female; elytral declivity with striae more 
distinctly punctured, interspaces more opaque, and rugosities slightly 
coarser, the mandibles stouter than in female. 

Male type labeled same as female except sex label. 

Variations. — The length varies from 3.7 to 6.4 mm,, averaging about 
5.5 mm. The color ranges from light brown to black; the sculpture 
and vestiture of the epistoma, front, pronotum, and elytra vary as 
usual, but the character of the strial punctures is fairly constant. 



106 THE SCOLYTID BEETLES. 

The greatest variation is in size and in the presence or absence of 
dorsal Hne of the pronotum. 

Distinctive characters. — The characters which distinguish this spe- 
cies from the next following, to which it is more closely allied, are 
the average smaller size and prevailing moderately impressed elytral 
striae with distinct but moderately coarse punctures. While the strial 
punctures vary in size in different individuals, they are never so coarse 
as in the average D. ponderosse. It is also distinguished from 
D. jeffreyi by the much smaller average length of body and the pre- 
vailingly coarser punctures of the pronotum. 

Revisional notes. — A brief description (Hopkins, 1905, p. 11) was 
published under the name D. monticola, but the name should read 
D. monticolse, which, as the manuscript name indicates, was origi- 
nally intended. The species is represented by three specimens from 
California in the Le Conte collection, with the type series, under 
D. similis, bearing specimen numbers 4, 12, and 13, and therefore 
may have been included in the revision by Le Conte (1876, p. 385), 
although the locality (California) is not given in that correction. 
There is also one specimen in the Horn collection, labeled "Cal.," 
under D. similis. It is probable that the one under similis was before 
Dietz when he prepared his revision (1890, pp. 30-31) under the name 
similis. These specimens are evidently the only ones which may have 
been involved in Le Conte's or Dietz's revisions on the literature pre- 
vious to Hopkins, 1899b, pp. 14 and 26. 

Pupa, — In addition to the generic, divisional, and subdivisional 
characters, the apex of the front and middle femora is armed with 
two small apical spines; abdominal tergites 2 to 6 with long and prom- 
inent pleural spines, 1 is without distinct dorsal and lateral spines, but 
2 to 6 have distinct dorsal and lateral ones, 2, 3, and 6 with a pair, 
and 4 and 5 with three lateral spines each side of the dorsal ones; 
7 has two dorsal spines, while 8 is smooth and the pleural spines of 9 
are long and prominent. Pupal type labeled "Hopk. U. S. No. 196." 

There is the usual variation in the arrangement and number of 
minor spines, and one specimen from Yosemite National Park is quite 
different in the widely separated frontal spines, with the dorsal ones 
of the abdominal tergites coarser and tergite 7 with a pleural spine. 
It is possible that this may be a pupa of an unrecognized species. 
The characters of the pupa of D. monticolx, which serve to distinguish 
it from that of D. ponderosse, are the less coarse spines of the abdom- 
inal segments, the less densely granulated elytral pads, and the pres- 
ence of two apical spines on the front and middle femora. 

Larva. — In addition to the generic and divisional characters, the 
front has the posterior angle subacute and a moderately stout, rugose 
elevation situated in the middle, becoming shghtly thickened and 
elevated toward the suture. Clypeus shining, with faint median 



THE GENUS DENDROCTONUS. 



107 



groove and usual elevation; apex broadly emarginate, labrum with 
apex broadly rounded, and the sternellar lobes of the thoracic seg- 
ments with distinct foot calli. Larval type labeled "Hopk. U. S. 
No. 196." 

There is some variation in the frontal elevation, but generally it is 
situated slightly anterior to the middle or in the middle, while the 
posterior angle of the front is distinctly subacute. These serve as 




Fig. 66. — Bendroctonus monticolx: Egg galleries and larval mines in bark. (Original.) 

the most distinctive characters to separate the larva of tliis species 
from that of I), 'ponderosx. 

Galleries (figs. 66, 67). — The egg galleries are longitudinal, distinctly 
to sHghtly winding or straight, usually grooved on the surface of 
the wood and deeply grooved in the bark, the larval mines and pupal 
cells exposed in the inner bark; the eggs are placed in approximate 
groups, and the larval mines are short and broad. The egg galleries 
differ from those of D. fonderosx in smaller size, more winding form, 
and are often of a much greater length. 



108 THE SCOLYTIP BEETLES. 

Distribution (fig. 68). — (Hopk. U. S.) California: Alder Creek, 
Chester, Fulda, Lake Tenaya, Mariposa Grove, Millwood, Siskiyou 
County, Soda Springs, Summerdale, Tioga Road, Wawona. Idaho: 
Boise County, Cedar Mountain, Centerville, Coeur d'Alene National 




Fig. 67.—Dendroctonus monticolx: Egg galleries and larval mines grooved In surface of wood. (Original.) 

Forest, Collins, Helena, Kootenai, Moscow Mountains, Sand Point, 
Smith's Ferry, Weiser. Montana: Big Fork, Columbia Falls, Iron 
Mountain, Lolo, Lewis and Clark National Forest, Missoula, Saltese. 
Oregon: Ashland, Grants Pass, Joseph, Pokegama, Wallowa. Waslir- 
ington: Longmires Springs, Mount Rainier National Forest, Pialschie, 



THE GENUS DENDROCTONUS. 



109 



Washington National Forest. Wyoming: North Fork Shoshone 
River, Wapiti. Additional localities from other collections: (A. M. 
N. H.) Millwood, CaL; (U.S.N.M.) Columbia Falls, Mont. 

Host trees. — Pinus lamhertiana, P. monticola, P. murrayana and P. 
ponderosa (common); Picea engelmanni (rare). 

Identified S'pecimens. — Le Conte, M. C. Z., 3; Horn, A. E. S., 1; 
A. M. N. H., Webb collection, 1; Henry Edwards collection, 1; 
U.S.N.M., 2; D. A., 11; Webb collection, 14; Hopk. U. S., over 500, 
including different stages and work. 




Fig. 68. — Dcndroctonus monlicolx: DistriljuUon map. (Original.) 
Bibliography and Synonymy. 

Dendroctonus similis (not of Le Conte, 1860) Le Conte, 1876, p. 385 (in part), locality. 
Dendroctonus n. sp. Hopkins, 1899b, pp. 14, 26, first record of habits and hosts. 
Dendroctonus monticola Hopkins, 1901b, p. 67, referred to as new species but not 

described, habits, galleries. Hopkins, 1902c, p. 21, notes. Hopkins, 1905, 

p. 11, first description, distribution, characters, very brief. Webb, 1906, p. 22, 

mentioned. 
Dendroctonus monticolse Hopkins, 1902a, p. 3, manuscript name. 
Dendroctonus n. sp. (mountain pine Dendroctonus). Hopkins, 1904, pp. 19, 42, 45, 

habits, hosts, distribution, etc. 
Mountain pine beetle. Hopkins, 1908, p. 162. 

10. Dendroctonus ponderosse Hopkins. 

(PI. IV, fig. 10.) 

Adult. — Type of species, female: Length, 6 mm., black; elytral 
declivity with a few long hairs. Head with front convex, without 
median elevation or groove, but with faint posterior impression; 
elytral rugosities moderately coarse and moderately dense, becoming 



110 THE-SCOLYTID BEETLES. 

much finer on the lateral area and coarser toward and on the vertex. 
Pronotum with moderately long, erect hairs on the lateral area, con- 
siderably longer toward the anterior section; punctures of elytral 
striae distinct and coarse. Secondary sexual characters same as in 
preceding species. 

Type labeled ''Type No. 7448 U.S.N.M./' name label, "Hopk. 
1/16/08, Pinus imnderosa, Spearfish, S. D., 7/1/00, A. D. Hopkins, 
collector, ? , Hopk. U. S. 434." 

Male type: Length 5.5 mm. Characters same as in female, except 
pronotum with very dense,, subrugose punctures toward the anterior 
margin, the elytral rugosities finer and less dense; elytral dechvity 
with coarser interspacial granules, and the strial punctures shghtly 
more distinct. 

Male type labeled same as female, except sex label. 

Variations. — ^The length varies from 4.5 to 7 mm., with the average 
about 6 mm. The color ranges from brown (in young specimens) to 
black in matured. The sculpture and vestiture of the epistoma, front, 
pronotum, and elytra vary as usual, and there is a quite noticeable 
variation from a somewhat slender form to a shorter and stouter one. 
The greatest variation is in length and in the size and density of the 
punctures of the pronotum and of the strise of the elytra. 

Distinctive characters. — The characters which distinguish tliis species 
from the one following are its average smaller size, less shining pro- 
notum, with coarser and deeper punctures, and from the preceding 
by its average larger size and somewhat stouter form, with the elytral 
striae more distinctly impressed and the punctures distinctly coarser. 
There is a considerable range of variation in these characters, but the 
specimens with less distinctly impressed striae and finer punctiu-es 
which might be mistaken for D. monticola are exceptional, and should 
cause no confusion as long as the range of distribution of the two 
species is so distinct. 

Revisional notes. — In March, 1902, the writer (Hopkins, 1902a, 
p. 3) published the manuscript name, D. i)onderosse, without descrip- 
tion of any kind, and in April of the same year (Hopkins, 1902b, p. 10) 
he gave a brief description under B. ponderosa, but, as indicated by 
the manuscript name, it was intended that the name should relate to 
the host tree, Pinus ponderosa, therefore the name D. ponderosse, under 
which it is here fully described, should stand. The species is repre- 
sented in the Le Conte collection by one immature example, which, 
in 1900, was in the D. rujipennis series, labeled ^'Specimen 8, Col." 
In March, 1907, this specimen was again examined by the -writer, and 
identified as D. pomderosx. It is evident that it was in Le Conte's 
collection when he prepared his latest revision (1876), but there is 



THE GENUS DENDROCTONXJS. Ill 

evidence that the characters and locality were not involved in the 
revised description under D. rujipennis. It is probably represented 
in the Horn collection (A. E. S.) by one specimen, without locality 
label, found by the writer in 1900 as the third specimen in the type 
series, under D. ai)proximatus. Two other specimens, labeled ''Colo.," 
were found in the Horn collection and one specimen in the general 
Academy of Natural Sciences collection under D. rujipennis. The 
two in the Horn collection were evidently before Dietz when he pre- 
pared his revision under D. rujipennis and included in the Colorado 
locality if not in the revised description. The writer (1902, p. 10) 
refers in a footnote to wrong identifications under D. rujipennis and 
D. terebrans. This probably includes all of the published references 
in which this species has been in any manner involved in revisions 
or systematic notes. 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the front and middle femora are armed each with one small 
apical spine. Abdominal tergites 2 to 6, with long and prominent 
plem-al spines; 1 is without distinct dorsal and lateral spines, but 2 to 
6 have distinct dorsal and lateral ones, and all of them have a pair of 
dorsal, and 2, 3, and 6 have a pair, and 4 and 5 have three lateral 
spines each side of the dorsal ones; 7 has two dorsal spines, while 8 
is smooth, and the pleural spines of 9 are long and prominent. Pupal 
type labeled ''Hopk. U. S. No. 623." 

In a number of individuals, the usual variation in the arrangement 
and number of minor spines is found, and between the younger and 
older examples there is a wide range of variation. 

The characters which, seem to distinguish the pupa of this species 
from that of I), monticolx are the coarser spines of the abdominal ter- 
gites, the more densely granulate el3^tral pads, and the presence of but 
one apical spine on the front and middle femora. 

Larva. — In addition to the generic and divisional characters, the 
front has the posterior angle subobtuse, and a stout prominent 
rugose transverse elevation situated slightly behind the middle and 
elevated and broad toward the sutures ; clypeus with the base sliining 
and bearing a faint median elevation, and the apex broadly emargi- 
nate; labrum less than half as long as broad, with the apex truncate; 
sternellar lobes of the thoracic segments with indistinct foot calli. 
Larval type labeled "Hopk. U. S. No. 755." 

There is some variation in the frontal elevation, but generall}^ it 
is situated behind the middle, and the posterior angle of the front is 
more obtuse than in the preceding, which latter serves as the most 
distinctive character separating the larva of this species from that of 
D. monticolx. 



112 



THE SCOLYTID BEETLES. 



Galleries (figs. 69, 70). — Tlie egg galleries are longitudinal, and 
usually nearly straight. The}^ are usually grooved on the surface of 
the wood and deeply grooved in the inner bark, with the larval mines 
and pupal cells exposed. The eggs are placed in approximate groups. 




Fig. 69. — Dcndroctonus pondcrosx: Egg galleries and larval mines. (Original.) 

and the larval mines are short. The egg galleries differ from those 
of D. monticolsd in the larger diameter and straighter, shorter form. 

Distribution (fig. 71). — (Hopk. U. S.) Arizona: Chiricahua Na- 
tional Forest, Flagstaff, Fredonia, San Francisco Mountains. Colo- 
rado: Bailey, Brookvale, Cascade, Cat Mountain, Cochetopa, Colo- 



THE GENUS DENDEOCTONUS. 



113 



rado Springs, Fort Garland, Glenwood Springs, Gunnison National 
Forest, Green Mountain Falls, Halms Peak, Husted, Indian Creek, 
Larlvspur, La Yeta, Manitou, Medicine Bow National Forest, Meeker, 
Monte Vista, Ouray National Forest, Palmer Lake, Pikes Peak, Pine, 
Ponclia Springs, San Isabel National Forest, San Juan National For- 
est, Sequache, Uncompahgre National Forest, White River National 
Forest. New Mexico: 
Gila National Forest, 
Vermejo. South Da- 
Jcota: Custer, Elmore, 
Hill City, Lead, Nemo, 
Piedmont, Sylvan Lake. 
Utah: Escalante, Ka- 
mas, Kanab, La Salle 
National Forest, Pan- 
guitch, Provo. Wyo- 
ming: Downington, En- 
campment, Keystone. 
Additional locality from 
other collectors : (Brown) 
Las Animas County, 
Colo. Localities report- 
ed hy correspondents: 
Eagle, Florissant, Idaho 
Springs, Kennedy Sta- 
tion, La Veta, Montrose, 
Pagosa Springs, Porter, 
San Juan,Ute Pass, and 
West Cliff, Colo. 

Host trees. — Pinus 
ponderosa scopulorum, 
P. jiexilis, P. murray- 
ana, P. strohiformis , and 
Picea engelnfianni. 

Economic relation to 
forests. — This species is 
exceedingly destructive 
to the pine forests of 
the central and southern Rocky Mountain region, having caused a 
loss of forest resources worth many milHons of dollars. The rock 
pine, or bull pine (Pinus ponderosa var. scopulorum), is its favorite 
host tree, but it attacks and kills the other pines and infests the 
spruce, though apparently not destructive to the latter. 

Identified specimens.— Le Conte (M. C. Z.), 1 specimen; Horn (A. E. 
S.), 2 specimens; A. N. S., 2 specimens; U.S.N.M., 1 specimen; 
79980—09 9 




Fig. 70.— Dendroctonus ponderosx: Tree with bark removed, show- 
ing egg galleries grooved and marked on surface of wood. (Orig- 
inal.) 



114 



THE SCOLYTID BEETLES. 



D. A., 14 specimens; Gillette, 2 specimens; Hopk. U. S., more 
than 10,000 specimens, including all stages and work. 

Bibliography and Synonymy. 

Dendroctonus similis (not of Le Conte, 1860) Le Conte, 1876, p. 385 (in part) (in col- 
lection 1890). 

Dendroctonus approximatus Dietz, 1890, p. 31 (in part?) (specimen with type 
series) . 

Dendroctonus ponderosx Hopkins, 1902a, p. 3, manuscript name only. Hopkins, 
1903b, pp. 275, 282, p. xxix, figs. 28, 32, stages and work figured, and full account 
of habits, life history, methods of control, etc. Hopkins, 1904, pp. 41, 43, 44, 
PL I, fig. 1, Pis. Ill, VIII, IX, XII, fig. 2, stages and work illustrated, habits, 
host, distribution, etc. Hopkins, 1905, pp. 1-24, full account of history, hab- 
its, life history, work, methods of control, etc.. Pis. I, II, figs. 1-6, stages and 
work. Hopkins, 1906a, p. 4, old work. Hopkins, 1906b, p. 147, Pis. IV, V, 
figs. 1-5, anatomy of larA'al head. Fall, 1907, p. 218, list, locality. Hopkins, 
1908, p. 162, depredations. 

Dendroctonus ponderosa Hopkins, 1902b, p. 10, brief original description, adult, etc., 
galleries and work illustrated, fig. 1, Pis. I, III, IV, VII, full account of habits, 
methods of control, etc. Hopkins, 1902c, p. 21, habits. Hopkins, 1903a, p. 59, 
habits, etc. 




Fig. 71. — Dendroctonus -ponderosx: Distribution map. (Original.) 

11. Dendroctonus Jeffrey! n. sp. 

(PI. IV, fig. 11.) 

Adult. — Type of species, female: Length 7.5 mm., black; elytral 
declivity with a few long hairs. Head with front convex, with shght 
anterior and posterior impressions, without frontal elevations ; elytral 
rugosities moderately coarse and dense, becoming much finer on lat- 
eral area and coarser toward the vertex. Pronotum shining, sides 



THE GENUS DENDEOCTONUS, 115 

distinctly constricted toward the head, with moderately long, erect 
hairs on the lateral area, longer and denser on the anterior surface; 
punctures of elytral striae distinct and coarse, the striae more dis- 
tinctly impressed on the dorsal area. Secondary sexual characters 
same as in preceding species. 

Type labeled "Type No. 7449 U.S.N.M.," name label, '^Hopk. 
1/22/08, Pinus jeffreyi, Little Yosemite, Cal., H. E. Burke, col- 
lector, 9 , Hopk. U. S. 4394a." 

Male type: Length 7 mm., same characters as female, except ely- 
tral declivity is more opaque and with distinctly coarser interspacial 
rugosities. 

Male type labeled same as female, except sex label. 

Variations. — The length varies from 6 to 8 mm., with the average 
about 7 mm., and the color from brown in young specimens to deep 
black when matured. The sculpture and vestiture of the epistoma, 
front, pronotum, and elytra vary as usual. There is apparently less 
variation in size and in other characters than is found among the 
individuals of the other species. 

Distinctive characters. — This species is at once distinguished from 
either of the two preceding by its average larger size and shining pro- 
notum, with its fine and shallow punctures. 

Note. — This species is not represented in any of the collections 
examined by the writer, and nothing has heretofore been published 
about it. 

Pupa. — In addition to the generic and divisional characters, the 
apices of the front and middle femora are armed with two distinct 
spines; abdominal tergites 2 to 6 with long and prominent pleural 
spines; 1 is apparently without dorsal spines, 2 and 6 are evidently 
with dorsal and lateral spines, but in the single poor specimen the 
relative size and number can not be made out; they appear, how- 
ever, to be less prominent than in either of the preceding species; 7 
has two small dorsal, three distinct lateral spines, and a small pleura] 
one; 8 is smooth, and 9 is with prominent pleural spines, as usual. 
Pupal type labeled '^Hopk. U. S. No. 4412a." 

The above description is based on a single specimen, which is dam- 
aged, therefore is subject to revision when more specimens are exam- 
ined. It seems to be quite different from the unique pupa mentioned 
under D. monticola. 

Larva. — In addition to the generic, divisional, and subdivisional 
characters, the front has the posterior angle subacute, and a narrow 
transverse elevation situated in the middle and not broadened or 
more elevated toward the sutures; clypeus shining, with median 
groove toward the base and with the apex broadly emarginate ; labrum 
with apex broadly truncate ; sternellar lobes of the thoracic segments 



116 



THE SCOLYTID BEETLES. 



with obscure foot scars. Larval type labeled "Hopk. U. S. No. 
6204b." 

There is very Uttle variation in the four specimens before the 
writer, and the narrow smoother frontal elevation, not elevated and 
broadened toward the suture, is the most distinctive character sepa- 
rating the larva of this species from those of the two preceding 
species. 

Galleries. — The egg galleries are longitudinal, nearly straight, 
grooved on surface of the wood, and deeply grooved in the inner 
bark, with the larval mines and pupal cells exposed. The eggs are 
placed singly and in approximate groups. The egg galleries and lar- 
val mines of this species differ from those of the two preceding in 




Fig. 72.— Dendroctonusjeffreyi: Distribution map. (Original.) 

being much coarser; otherwise they appear to be more like those of 
monticolse. 

Distribution (fig. 72). — (Hopk. U. S.) California: Chester, Little 
Yosemite, Nevada City, Pinogrande, Sterling, Tallac, Yosemite 
(Yosemite National Park), and Seven Oaks (San Bernardino National 
Forest). 

Host trees. — Pinus jejfreyi. P. ponderosa, and P. lambertianxi. 

Identified specimens. — Hopk. U. S., over 160 specimens, including 
adults, pupae, and larvse. 

DIVISION II. 

The distinctive characters common to the species of the second 
division are: 

Adults. — Prothorax stout, usually narrower than elytra, distinctly 
narrowed or constricted toward head; elytra with anterior dorsal 



THE GENUS DENDEOCTONUS. 117 

half bearing long hairs, except in B. terebrans and badly rubbed speci- 
mens. 

Pupa. — Vertex of head faintly impressed, fiat or convex, and with 
two small widely separated frontal granules toward vertex. 

Larva. — Abdominal tergites 8 and 9 with dorsal plates except in. 
simplex and pseudotsugx, and 8 without plate in micans. 

Galleries. — Egg galleries longitudinal, straight to slightly winding; 
eggs in groups or masses; larval mines and pupal cells exposed in 
inner bark. 

Subdivision C. 
(Species Nos. 12 to 21, inclusive.) 

The distinctive characters common to the species of this third sub- 
division are: 

Adults. — Front usually with posterior impression, pronotum with 
large and small punctures intermixed. Pronotum with long hairs on 
dorsal and lateral areas. 

Sexes. — Female: Interspaces of elytral declivity more roughened 
and the striae more distinctly impressed. Male: The reverse. 

Pupa. — -Vertex of head flattened or faintly impressed; apices of 
front and middle femora smooth; abdominal tergites with moderately 
prominent pleural and dorsal spines. 

iarva.— Abdominal tergites 8 and 9 without dorsal plate in simplex 
and pseudotsugce and with unarmed dorsal plate in the remaining 
species. 

Galleries. — Egg galleries slightly winding to straight; eggs in 
groups, but larval mines separated from the beginning, or beyond the 
middle, except in D. Tuicans. 

12. Dendroctonus simplex Le Conte. 
(PI. V, fig. 12.) 

Adult. — Typical female: Length, 3.9 mm., dark reddish-brown. 
Head with front distinctly convex, with faint posterior impression. 
Pronotum with distinctly coarse and fine punctures intermixed; ely- 
tral declivity with strias deeply impressed ; epistomal process narrow, 
flat, the sides nearly parallel, apex not extending beyond the anterior 
frontal margin; elytral rugosities moderately coarse, becoming finer 
on the lateral areas, sparse, coarser, and more acute on the dorsal 
area and vertex; striae toward suture impressed, not impressed on 
lateral area; strial punctures coarse and distinct. Pronotum with 
moderately long reclining hairs on lateral area. Secondary sexual 
characters: Elytral declivity with interspaces more rugose and the 
striae more distinctly impressed. 

Typical female labeled, name label, "Hopk. 1/22/08, IT. S. 41, 9 , 
Grand Ledge, Mich., 21.4" ( = April 21). 

Typical male: Length, 8.8 mm.; elj^tra more shining, less rugose; 
declivit}^ shining, interspaces convex and smooth, with fine, distinct 



118 THE SCOLYTID BEETLES. 

punctures and with striae distinctly impressed and finely punctured ; 
otherwise as in female. 

Typical male labeled, name label, "Hopk. 1/22/08, H. S. 40, $ , 
Grand Ledge, Mich., 20.5" ( = May 20). 

Variations. — The length varies from 3.5 to 5 mm., with the average 
about 4.7 mm. The color varies from reddish to reddish-brown, with 
the head and thoracic segments ranging from light to dark brown, 
and nearly black. The epistomal characters are more constant in 
this and the next species, otherwise the usual variation in the sculp- 
ture and vestiture of the head, pronotum, and elytra prevails. The 
greatest variation, other than size, is found in the punctures of the 
pronotum and in the strial punctures and interspacial rugosities of 
the elytra. 

Distinctive characters. — The characters which distinguish this 
species from the following, to which it is more closely allied, are the 
much smaller average size of the individual and the coarser and 
deeper punctures of the pronotum. 

Revisional notes. — The original description was based on two male 
specimens labeled "Canada," which have been examined by the writer 
and found to agree with the common species which lives in the 
eastern larch. The comparison in the description with D. ohesus 
must refer to D. pseudotsugse Hopk., representatives of which were 
then confused with the true D. ohesus (hisum.). The smooth intervals 
on the declivity referred to is a male character. The reference to a 
much deeper sutural stria relates to what is now recognized as 
stria 1. The revision relates to the type specimens. With our 
present knowledge of the specific characters, those given in Le Conte's 
tables are onl}'^ partially applicable. It is evident that no other 
species were confused with this one in Le Conte's description and 
revision, but it was involved in the revision under D. ohesus (1868) 
andD. similis (1873). In 1900 it was in Le Conte's collection under 
D. rujipennis, labeled "Lake Superior" and "Tex.," and under 
D. similis, labeled "Lake Superior" and "Can." In Dietz's revi- 
sion, the specimens from Colorado and California were evidently 
D. pseudotsugx, but did not involve any confusion in the description, 
except in the length, 6.2 mm., which was evidently based on a 
specimen of D. fseudotsugx. 

The species was found to be represented in the Horn collection by 
one specimen, under simplex, labeled "Can.," and two specimens 
under D. similis, labeled "Can." 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the apex of the front and middle femora has a minute 
subapical granule. Abdominal tergite 1 with very small and 2 to. 6 
with stout, prominent pleural spines, 1 without dorsal or lateral, 2 
without dorsal, 3 to 6 with prominent dorsal spines, becoming larger 
toward 6, 2 to 6 with two lateral spines each side, becoming more 



THE GENUS DENDEOCTONUS. 



119 



prominent toward the latter, 7 with two minute dorsal hairs, 8 
smooth, 9 with prominent widely separated spines. Pupal type, 
labeled ''Hopk. U. S. No. 6444b." 

The usual variation in arrangement and number of minor spines 
prevails, but the pupa of this species is easily distinguishable from 
that of the following by its smaller size and the more prominent 
dorsal, lateral, and pleural spines. 




Fig. 73.—Dendroctonus simplex: Egg galleries and larval mines. (Original.) 

Larva. — In addition to the generic and divisional characters, the 
front is convex, opaque, with the posterior angle obtuse, and the 
middle with an indistinct transverse elevation, and transversely 
rugose. The clypeus is prominent, shining, with a distinct median 
impressed line and the apex broadly emarginate; the labium is short, 
with the apex broadly rounded. The sternellar lobes of the thoracic 
segments are prominent and with distinct foot calli. Larval type 
labeled "Hopk. U. S. No. 6444b." 

Galleries (fig. 73). — Egg galleries longitudinal, slightly winding, 
and sometimes branched and grooved on the surface of the wood, as 



120 



THE SCOLYTID BEETLES. 



well as deeply grooved in the inner bark; larval mines and pupal 
cells exposed in the innfer bark. Eggs are placed in groups of three 
to five or more, and the larval mines, which are short, are separated 
from the start. The galleries differ from those of the next species 
by their smaller size, more elongate and winding form of the egg 
galleries, and the much shorter larval galleries arranged in much 
smaller groups. 

Distribution (fig. 74). — (Hopk. U. S.) Maine: Cupsuptic. Michi- 
gan: Grand Island, Munising, Mackinac Island, Seney. (Hopk. W. 
Va.) West Virginia: Cranesville. Additional localities from other col- 
lections: (U.S.N.M.) Ungava Ba}^ Canada; Agricultural College, 
Mich. (H. & S.) Marquette, Grand Ledge, and Port Huron, Mich. 




Fig. 74. — Dendroctonus simplex: Distribution map. (Original.) 

(D. A.) West Stewartstown, N. H. One specimen in the Le Conte 
collection under D.ohesus, labeled "Texas" (must be an erroneous 
locality, resulting from some mistake) . 

Host tree. — Larix laricina. 

Identified s'pedmens. — Le Conte, 10 specimens (2 under D. simplex, 
3 under D. rufipennis, and 5 under D. similis); Horn, 3 specimens 
(2 under simplex, 2 under similis); U.S.N.M., 6; H. & S., 10; D.A., 
5 specimens; Hopk., W. Va., 157, and Hopk. U. S., over 150 
specimens. 

Bibliography and Synonymy. 

Dendroctonus simplex. .Le Conte, 1868, p. 173, original description, synopsis, 
localities. Le Conte, 1876, p. 385, revision, synopsis, bibliography, localities. 
Packard, 1887, p. 177 (Le Conte quoted). Schwarz, 1888, p. 175, synonymy, 
habits in larch. Packard, 1887, p. 177, Le Conte quoted. Packard, 1890, p. 722 



THE GENUS DENDEOCTONUS. 121 

(ibid). Dietz, 1890, p. 31 (in part), Michigan and Lake Superior, fig. 4, antenna 
and epistoma. Harrington, 1891, p. 27, habits and host. Hopkins, 1898a, p. 69, 
distinct from rujipennis (piceaperda) . Hopkins, 1898b, in larch in W. Va. 
Hopkins, 1899a, p. 392, etc., fig. Iviii, adult, revisional notes, etc., host, distribu- 
tion. Hopkins, 1899c, p. 343, good species, habit, host, etc. Felt, 1906, p. 752 
(in part), bibliography. 

Dendroctonus similis (not of Le Conte, 1860) Le Conte, 1876, p. 385 (in part), revi- 
sion, synonymy, bibliography, localities. Dietz, 1890, p. 31, Canada. 

Dendroctonus rufipennis (not of Kirby) Le Conte, 1876, p. 385 (in part) (in collec- 
tion 1900-1907, under rujipennis) . 

Dendroctonus sp. Harrington, 1884, p. 218. Packard, 1890, p. 903. 

13. Dendroctonus pseudotsugae Hopkins. 

(PI. V, flg. 13.) 

Adult. — Type of species: Length 5.75 mm.; reddish brown, with 
the prothorax darker. Head with front convex, with faint median 
and posterior impression; elytral declivity with striae deeply im- 
pressed; epistomal process narrow, slight, with sides nearly parallel, 
the apex scarcely projecting beyond the anterior margin. Pronotum 
with punctures fine, and moderately regular in size; elytral rugosities 
moderately coarse, finer on lateral area, coarse and more acute on 
dorsal area and vertex; strise of dorsal area distinctly impressed, not 
impressed on lateral area; punctures coarse and distinct. Pronotum 
with moderately long hairs on the lateral area. Secondary sexual 
characters: Elytral declivity convex, with interspaces rugose and 
the striae distinctly impressed and punctured. 

Type labeled ''Type No. 7450 U.S.N.M.," name label, ''Hopk. 
1/22/08, Pseudotsuga taxifolia, Hopkins, collector. Grants Pass, Or., 
9 , Hopk. U. S. 39." ' 

Male type: Length 5.75 mm.; elytral declivity with interspaces 
strongly convex and smooth, shining, sparsely punctured; striae 
deeply impressed, punctures obscure, otherwise as in female. 

Male type labeled " S type," name label, ''Hopk. 1/22/08," other- 
wise same as female. 

Variations. — The length varies from 4 to 7 mm., with the average 
about 6 mm. The color ranges from light reddish to nearly black. 
The usual variation in sculpture, vestiture, etc.^- "prevails. The 
greatest variation is in the size and color, and in the size of the 
punctures of the pronotum. The epistomal process varies consid- 
erably, so that in some specimens it extends beyond the epistomal 
margin, while in others it does not. 

Distinctive characters. — This species and the one preceding are 
at once distinguished from all of the other species of the genus by the 
characters of the epistoma and from each other by the sculpture of 
the pronotum and the difl^erence in average size of representative 
individuals. D. pseudotsugse is distinguished especially by the 
shining and finely punctured pronotum. 



122 



THE SCOLYTID BEETLES. 



Revisional notes. — This species has been an element of much con- 
fusion in descriptions, revisions, and identifications, under the 
names D. similis, D. obesus, D. rufipennis, etc. It is represented in 
the type series in Le Conte's collection under D. similis, but the 
specimen which bears the name label, and therefore the type of 
D. similis, is a true D. obesus (Alarm.). Le Conte (1868, p. 173) referred 
D. similis to obesus on account of the intervals of the elytra being 
''rough for their whole extent," which is the case in the type of 
D. similis, and in all females of D. obesus. Later he evidently com- 




FiG. 75. — Dendroctonus pscudotsugx: Egg galleries and larval mines, a, Beginning or basal sections of egg 
galleries in bark; 6, entrance; c, egg gallery; d, ventilating hole; e, egg nest; /, abnormal branch; g, lar- 
val mines; ^, egg gallery packed with borings; i, subsequent passage or inner gallery through borings. 
(Original.) 

pared the type of D. similis with a single male specimen of D. obesus 
in his collection, which, accordmg to Mr. Henshaw, is from the 
Maruierheim collection, and finding that this differed from his D. 
similis in the smooth elytral declivity he restored D. similis (Le Conte, 
1876, p. 385) and called attention to the roughened interspaces of the 
declivity as a distinctive character ; all of which makes it quite clear 
that he considered the specimen bearing the name label as the type 
of his D. similis and that therefore this name must fall as a synonym 
of D. obesus Mann. Thus the other specimens of the type series are 
left to represent a distinct species as here described. In 1900 it was 



THE GENUS DENDEOCTONUS. 



123 



represented in the Le Conte collection by two specimens labeled "Or." 
and three specimens labeled "Van." under D. similis, and one speci- 
men labeled "Garland Pass, Col,," under D. rufipennis. In 1900 
it was represented in the Horn collection by one specimen labeled 
"Col." and one specimen labeled "Cal.," and in the A. E. S. collec- 
tion by three specimens from Oregon under D. similis and two speci- 
mens labeled "Col." under D. rufipennis. Dietz's revision under 
D. similis (1890, pp. 30-31) includes the characters of D. pseudo- 
tsugsd, as represented by the Oregon, Colorado, and California speci- 




FiG. 76. — Dendroctonus pseudotsugie: Egg gallery and larval mines, a, Egg gallery in bark and grooved 
in surface of wood; 6, larval mines in bark; c, larval mines marked and slightly grooved on surface ol 
wood. (Original.) 

mens, while the specimens from Canada represented D. simplex, and 
one from California referred to in the note is D. monticolx. 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the apices of the front and middle femora are smooth; 
abdominal tergite 1 is without a pleural spine, 2 with small, and 3 
to 6 with rather stout ones ; 1 and 2 are without dorsal and lateral 
spines, while 3 to 6 have a pair of dorsal and a pair of lateral ones 
each side ; 7 and 8 are smooth, and 9 has the usual prominent pleural 
spines. Pupal type labeled "Hopk. U. S. No. 2298." 



124 



THE SCOLYTID BEETLES. 



The usual variation in number and arrangement of minor spines 
prevails in other specimens, but they are easily distinguished frojn 
those of D. simplex by their larger size and less prominent dorsal, 
lateral, and pleural spines. 

Larva. — In addition to the generic and divisional characters, the 
front is opaque, except toward the apex, where it is more shining, 
and the apex is subacute; the middle has a transversely rugose 

elevation, slightly more ele- 
vated and broader toward 
the suture. The clypeus is 
prominent, shining, and with 
an impressed line from the 
middle to the anterior mar- 
gin, which is broadly emar- 
ginate; labium short, with 
the apex broadly rounded; 
the sternellar lobes are mod- 
erately prominent, with in- 
distinct foot calli. Larval 
type labeled "Hopk. U. S. 
No. 2289." 

Galleries (figs. 75-77). — 
The egg gallery is longitudi- 
nal, short, but slightly wind- 
ing, sometimes branched, 
slightly grooving the surface 
of the wood, and deeply 
grooved in the inner bark. 
The larval mines and pupal 
cells are exposed in the inner 
bark, and the eggs are rather 
closeh^ placed in groups of 
three to ten or more, but the 
larval mines are separated 
from the start and are usu- 
ally extended for some dis- 
tance from the egg gallery. 
The galleries of this species 
differ from those of the one preceding by their larger size and shorter 
form of the egg gallery and the much larger larval mines, which are 
arranged in larger groups. 

Distribution (fig. 78). — (Hopk. U. S.) Arizona: Chiricahua National 
Forest, Flagstaff, San Francisco Mountains, Santa Catalina National 
Forest. California: Fieldbrook, Guerneyville, McCloud, San Mateo 
County (Big Basin). Colorado: Colorado Springs, Fort Garland, Gun- 
nison National Forest, Indian Creek, Leavenworth Valley, Moffat, 




Fig. 77.— Dendroctonus pseudotsugse: Section of log with 
bark removed, showing brood galleries marked and 
grooved on surface of wood. (Original.) 



THE GENUS DElSTDROCTOlSrUS. 



125 



Ouray, Palmer Lake, San Isabel National Forest, San Juan National 
Forest, Saguache. Idaho: Beaver Canyon, Centerville, Bailey, 
Henrys Lake National Forest, Kooskia, Kootenai, Pioneerville, 
Priest River, Sand Point, Smiths Ferry, Stites. Montana: Belton, 
Bozeman, Middle Creek (Gallatin County), Ovando. New Mexico: 
Capitan, Cloudcroft, Sacramento National Forest, Santa Fe, Ver- 
mejo. Oregon: Corvallis, Detroit, Grants Pass, Newport, Slate 
Creek, St. Helena. Utah: Panguitch. Washington: Ashford, Buck- 
eye, Des Moines, Dole, Gray's Harbor City, Hoquiam, Junction, 
Kent, Keyport, Meredith, New London, North Bend, Orting, 
Pialschie, Port Angeles, Port Williams, Pullman, Puyallup, Rock 




Fig. 78. — Dcndroctonus pseudotsugx: Distribution map. (Original.) 

Creek, Satsop. Additional localities from other collections: (Le Conte) 
Vancouver, B. C. (U.S.N.M.) Easton, Wash. . ( H. & S.) Hood 
River, Oreg; Beaver Canyon, Idaho. (D. A.) Mount Angel, Oreg. 
(Soltau) Seattle, Wash. (Wickham) Leavenworth Valley and 
Kalispell, Mont. 

Host trees. — Pseudotsuga taxifolia, P. Tnacrocarpa, and Larix 
occidentalis. 

Identified specimens. — Le Conte, 6 specimens (1 under rufipennis, 
5 under D. similis); Horn, 2; A. E. S., 5; Dietz, 2; U.S.N.M., 7; 
H. &. S., 2; Soltau, 3 ; Webb, 21; Wickham, 2; Laurent, 1; D. A., 5; 
Hamilton, 1 ; Hopk, U. S., over 700 specimens, including all stages and 
work. 



126 THE SCOLYTID BEETLES. 

Bibliography and Synonymy. 

Dendroctonus similis (not of Le Conte, 1860) Le Conte, 1876, p. 385 (in part), revision, 
synonymy, bibliography, localities. Le Conte, 1878, p. 469, listed, Leavenworth 
Valley, Colo. Packard, 1887, p. 177, Le Conte quoted. Packard, 1890, p. 722, 
Le Conte quoted. Dietz, 1890, pp. 30-31, from Oregon, California, Colorado, fig. 3, 
antenna, epistoma. Hopkins, 1899a, p. 392, fig. Iviii, adult. Hopkins, 1899b, pp. 
10, 11-15, 21, 22, 26, first records, habits, hosts, etc. Wickham, 1902, p. 310, list 
and localities. Hopkins, 1903a, p. 61, synonymy. Fall, 1907, p. 218, in list, 
locality. 

Dendroctonus rufipennis (not of Kirbj') Le Conte, 1876, p. 385 (in part) (in collection 
1900-1907, under rufipennis). Le Conte, 1878, p. 469 (in part). Packard, 1887, 
pp. 177, 243 (in part?). Hopkins, 1899b, p. 15, localities, note. Hopkins, 1904, 
p. 19, reference. 

Dendroctonus simplex (not of Le Conte) Dietz, 1890, p. 31 (in part), Colorado, Cali- 
fornia. Wickham, 1902, p. 310 (on Dietz's authority). 

Dendroctonus pseudotsugx Hopkins, 1901b, p. 67, brief description of adult, galleries, 
habits, distribution, etc. Hopkins, 1903a, p. 60, habits, comparison with D. simi- 
lis, which=Z). obesus. Hopkins, 1905, pp. 10, 11, brief description, habits, etc. 
Hopkins, 1906a, p. 4, old work. 

Dendroctonus n. sp. (Douglas spruce beetle.) Hopkins, 1904, pp. 19, 45. 

14. Dendroctonus piceaperda Hopkins. 

(PI. V, fig. 14.) 

Adult. — Type of species, female: Length, 5.75 mm.; elytra red, 
thorax, head, and abdomen black; head with front convex and with 
faint median and posterior impression and anterior elevated line. 
Elytral declivity with striae not deeply impressed; epistomal process 
broad, concave, with the lateral section oblique; punctures of prono- 
tum distinctly irregular; posterior half of proepisternal area not 
punctured; elytral striae distinctly impressed in dorsal and lateral 
areas, with punctures rather coarse and distinct; interspaces slightly 
convex; rugosities acute, rather closely placed, irregular. Secondary 
sexual characters: Declivity convex; striae very faintly impressed, 
with fine indistinct punctures; interspaces nearly flat, shining, with 
approximate row of fine granules. 

Type labeled ''Type No. 7451 U.S.N.M.," name label, "Hopk. 
4/23/02, compared with Kirby type rujipennis, does not agree, Picea 
canadensis, Hopkins, collector. Camp Caribou, Me., 9 type, Hopk. 
U. S. 326." 

Male t3rpe: Length, 5.6 mm.; elytra dark reddish-brown; thorax 
and head darker; elytral declivity convex; striae not impressed; 
punctures obscure; interspaces flat, shining, and finely punctured, 
with very small granules toward vertex. 

Type labeled, " $ type, type of drawing," name label, "Hopk. 
1/22/08, Picea canadensis, Hopkins, collector. Camp Caribou, Me., 
$ type, Hopk. U. S. 326." 

Variations. — The length varies from 4.7 to 6 mm., with the average 
about 5.5 mm. The color ranges from uniform light red to black, to 
the head, thorax, and abdomilial sternites dark to black, with the ely- 



THE GENUS DENDEOCTONUS. 127 

tra lighter or red. The sculpture and vestiture of the epistoma, front, 
pronotum, and elytra vary as usual. The greatest variation is in size 
and color. 

Distinctive characters. — The characters which distinguish this spe- 
cies from the next are its smaller average size, slightly less elongate 
form, less shining elytra, with the striae more distinctly impressed on 
the sides and the interspaces slightly more convex and more acutely 
rugose. The difference is not so perceptible in comparing single indi- 
viduals as when many individuals of both species are compared. It 
differs from D. horealis by the noticeably more elongate and narrower 
pronotum, and from species 17 to 21 it is distinguished by the dis- 
tinctly impressed lateral striae of the elytra, except D. punctatus Lee, 
which is at once recognized by the coarse punctures of the declivital 
striae. It is at once distinguished from D. rufipennis (Kirby) by its 
smaller size and coarsely punctured and impressed lateral striae. 

Revisional notes. — Probably no species of the genus has been in- 
volved in so much confusion as this. It has been extensively dis- 
cussed under D. rufipennis, and confused in collections with several 
other species under this name. There are three specimens in the 
Le Conte collection labeled "Anticosti," which were evidently the 
ones referred to in his revision (1876, p. 385). There are also two 
specimens without locality labels, which may have been the ones from 
Colorado, while the one from Alaska is here referred to D. horealis. 
The smoother and more shining declivity referred to by Le Conte as a 
distinguishing character relates to the males only. It is represented 
in the Horn collection by two specimens labeled ''Canada," under 
D. rufipennis, which were therefore evidently included in Dietz's 
revision. It is also very probable that the specimens from New 
Brunswick belonged to this species. 

It is very evident that the barkbeetle referred to under D. rufipen- 
nis by Peck, Packard, Hough, and other authors as depredating 
on the spruce of New Brunswick, Canada, New England, New York, 
and Pennsylvania was D. piceaperda. 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the apices of the front and middle femora are smooth; 
abdominal tergites 2 to 6 with very small pleural spines; 1 without 
distinct dorsal or lateral spines; 2 to 3 without dorsal, but with two 
small lateral spines each side; 4 to 6 with a pair of very small dorsal 
and three or four small lateral spines each side; 7 and 8 smooth; 9 with 
usual pleural spines. Pupal type labeled "Hopk. U. S. 377." 

The usual variation in minor details prevails. It is distinguished 
from the pupa of Z>. engelmanni by the less impressed vertex of the 
head and the generally smaller spines and the absence of dorsal 
spines on the third abdominal tergite. 

Larva. — In addition to the generic, divisional, and subdivisional 
characters, the front has a slight transverse, rugose elevation situ- 



128 THE SCOLYTID BEETLES. 

ated slightly in front of the middle, the anterior surface, including the 
elevation, opaque. The area behind the elevation is impressed and 
shining; clypeus short, broad, with median groove and its apex sub- 
acutely emarginate ( in dried specimens) ; labrum prominent, its ante- 
rior margin subtruncate; mandibles opaque, with slight dorsal impres- 
sion or elevation; sternellar lobes of thoracic segments moderately 




ViG. 79.— Dendroctonus piceaperda: Egg gallery and larval mines, a, Egg gallery; 6, boring dust packed 
in gallery; c^ entrance and subsequent or inner gallery; d, larval mines. (Author's illustration.) 

prominent and with distinct foot calli. Larval type labeled "Hopk. 
U. S. No. 318." 

The most distinctive characters are the opaque mandibles with 
moderate impression and elevation, and the distinctly elevated ante- 
rior margin of the epicranium. 

Galleries (fig. 79). — The egg galleries are short, broad, longitudinal, 
grooving the surface of the wood and deeply grooved in the inner 



THE GENUS DENDEOCTONUS. 



129 



bark, the larval mines exposed and the pupal cells partially to entirely 
exposed. The eggs are closely placed in large groups, and the larval 
mines are at first contiguous or nearly so, near the egg gallery, but 
soon become separated and when completed are often as long as the 
egg gallery or longer. The egg galleries differ from those of all of the 
species of subdivisions A and B in being very much broader than the 
diameter of the beetle's body. This broad groove is packed with 
borings, through which a central gallery is excavated by the parent 
beetle after the eggs have been deposited. 

Distribution (fig. 80). — (Hopk. U. S.) Maine: Beaver Pond, Camp 
Caribou, Cupsuptic, Meadows. Michigan: Grand Island, Munising. 
New HaTupsJiire: Waterville. Additional localities from sj^ecimens 




Fig. 80. — Dendroctonus piceaperda: Distribution map. (Original.) 

identified in other collections: (Le Conte) Anticosti, Canada. (Horn) 
Canada. (U.S.N.M., H. & S.) Isle Royale, Mich. (D. A.) Colebrook 
and West Stewartstown, N. H. (Wenzel) Ricketts, Pa. 

It is evident that this species follows the distribution of the spruce 
from the higher mountains of central Pennsylvania northward and 
eastward into New York, New Hampshire, Maine, New Brunswick, 
and Canada, and westward to the Lake Superior region. 

Host trees. — Picea rubens, P. canadensis, and P. mariana. 

Identified specimens. — Le Conte, 3 specimens from Anticosti, 2 with- 
out label, under D. rufipennis, 1 labeled "N. Y." under D. punctatus 
(Mar. 11, '07), 2 specimens without locality labels doubtfully referred 
to this species; Horn, 2 specimens labeled "Can.," under D. rufipen- 
nis; U.S.N.M., H. & S., 1 specimen labeled ''Isle Royale;" Weed and 
79980—09—10 



130 THE SCOLYTID BEETLES. 

Fiske, 12 specimens; Hopk. U. S., over 300 specimens, including all 
stages and work. 

Bibliography and Synonymy. 

Hylurgus rujipennis (not of Kirby) Peck, 1876, pp. 283, 301, destruction of spruce in 
New York (evidently the work of D. piceajyerda Hopk.). Peck, 1879, pp. 32-38, 
ravages in spruce in northern wilderness (same as 1876?). Packard, 1890, pp. 814- 
815 (quotes Peck, ibid.). Hough, 1882, pp. 259-263, insect ravages in spruce 
forests of Maine. 

Dendroctonus rujipennis (not of Kirby) Le Conte, 1876, p. 385, revision, synopsis, 
localities. Lintner, 1885, p. 54, destruction of spruce in New York. Fletcher, 
1887, pp. 39-40, habits. Dietz, 1890, p. 30 (in part), Canada and New Brunswick. 
Packard, 1887, pp. 177-243 (in part). Packard, 1890, p. 722 (in part), quotes 
Le Conte. Harvey, 1898, p. 176, depredations on spruce in Maine; p. 98, host, 
distribution, etc. Hopkins, 1898a, p. 69, distinct from simplex. Weed and Fiske, 
1898, pp. 67-69, report on investigations. Chittenden, 1898, p. 96, doubt as to 
Kirby's species. Smith, 1899, p. 364, Lakewood, N. J. Hopkins, 1899a, pp. 349- 
393 (in part), reference. Hopkins, 1899c, p. 343 (in part?), reference. Chitten- 
den, 1899, p. 56 (in part?), reference. Johnson, 1901, p. 92, habits in Pa. 
Hopkins, 1905, p. 6, reference to wrong determination. Felt, 1906, p. 753 (in 
small part), bibliography. 

Polygraphus rujipennis (not of Kirby) Packard, 1890, p. 721, [fig. 251=Polygraphus 
rujipennis (Kirby)], Le Conte quoted (includes several species). 

Xyloterus bivittatus (not of Kirby) Packard, 1890, p. 823, fig. 276 (in part), adult?; PI. 
XXIV, fig. 1, larva?, 1 a, pupa?, destruction of spruce. 

Barkbeetles. Packard, 1890, pp. 811-824 (inpart), destruction of spruce, New Bruns- 
wick to New York. 

Dendroctonus (Polygraphus) rujipennis (not of Kirby) Gary, 1900, pp. 52-54, depre- 
dations on spruce, methods of control. 

Dendroctonus piceaperda ■ Hoipkins, 1901a, p. 16, PL II, larvae, pupse, adult, etc., 
original description, different stages and galleries, with full account of habits, host, 
natural enemies, methods of control, etc. (see also index and Pis. I-V, XIV, XV). 
Hopkins, 1902b, p. 21, mention. Hopkins, 1902a, p. 3. Hopkins, 1902c, p. 22, 
habits, etc. Hopkins, 1903b, pp. 266, 270, 281, PL XXVII, figs. 23-25, stages 
and work figured, revised account of habits, life history, methods of control, etc. 
Hopkins, 1904, p. 26, PL I, fig. 3, Pis. V, XII, fig. 1, Pis. XIII, XIV, XV, stages 
and work (reprints), habits, hosts, distribution, etc. Hopkins, 1905, pp. 10, 11, 
distinctive characters, brief. Felt, 1905, pp. 6, 7, habits and work. Felt, 1906, 
pp. 379-385, fig. 85 b, history, habits, etc. Hopkins, 1908, pp. 160-161, depreda- 
tions. 

15. Dendroctonus engelmanni n. sp. 

Adult. — Type of species, female: Length 6.2 mm., black. Head 
with front convex, faint median and posterior impression and faint 
anterior line. Elytral declivity with striae not deeply impressed; 
punctures of pronotum distinctly irregular; posterior half of proepi- 
sternal area not punctured ; punctures of prothorax and elytra rather 
coarse; strias moderately impressed; interspaces moderately convex, 
and scarcely rugose, except on dorsal area. Secondary sexual char- 
acters: Declivity convex; striae rather distinctly but not deeply 
impressed; punctures distinct; interspaces with rows of granules. 



THE GENUS DENDEOCTONUS. 131 

Type labeled "Type No. 7452 U.S.N.M.," name label, "Hopk. 
1/22/08, Picea engelmanni, Capitan, N. M., W. F. Fiske, collector, 
9 , Hopk. U. S. 3958." 

Male type: Length 5.5 mm. Front without anterior line. Elytra 
with striae less distinctly impressed and interspaces less convex than 
in female; declivity convex, with strife and strial punctures obscure; 
interspaces flat, shining, finely but distinctly punctured and without 
granules except on vertex. 

Type labeled, " $ type," name label, "Hopk. 1/22/08, Picea engel- 
manni, Capitan, N. M., W. F. Fiske, collector, $ , Hopk. U. S. 3958." 

Variations. — The length varies from 5 to 7 mm., with the average 
at about 6.5 ]5lm. The color ranges from uniform light to dark red 
and black, to black head, thorax, and abdomen, and red elytra. The 
sculpture and vestiture of the epistoma vary as usual, with the 
greatest variation in size, color, and punctures. 

Distinctive characters. — The characters which serve to distinguish 
this species from the one preceding are the larger average size, 
slightly more elongate form, more sliining elytra, with the lateral 
strige somewhat less impressed, the punctures usually coarser, and the 
interspaces less acutely rugose. It is more closely allied to D. obesus, 
from which it is distinguished by the commonly darker prothorax, 
and more distinctly impressed lateral striae of the elytra, with coarser 
punctures. 

Revisional notes. — This species, like the preceding, has been involved 
in the confusion in revisions and collections under D. rufipennis. 
The species under this name was represented in the Le Conte collec- 
tion by two specimens labeled "Alta, Ut." and "Colo.," by two in 
Horn's collection labeled "H. B.," and "Alta, Ut.," and by one 
specimen from Doctor Dietz, labeled "Ut." These were doubtless 
involved in Le Conte 's and Dietz's revisions. 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the apices of the front and middle femora are smooth; ab- 
dominal tergites 2 to 6 with small pleural spines; 1 and 2 without 
dorsal but with lateral spines, and 3 to 6 with dorsal and lateral spines ; 
7 and 8 smooth; 9 with pleural spines, as usual. Pupal type labeled 
"Webb No. 2." 

The usual variation in minor details prevails, but the pupa of this 
species is distinguished from that of the preceding one by the more 
distinctly impressed vertex of the head, the generally coarser spines, 
and the presence of dorsal spines on the third abdominal tergite. 

Larva. — In addition to the generic, divisional, and subdivisional 
characters, the front has a transverse rugose elevation situated near 
the middle; the anterior surface, including the elevation, is opaque; 
the area behind the elevation is impressed and shining, clypeus 



132 



THE SCOLYTID BEETLES. 



broad, with median dorsal groove and the apex subacutely emar- 
ginate; labrum prominent, its anterior margin truncate; mandibles 
opaque toward base, more shining toward apex, with a distinct dorsal 
impression and oblique ridge near the middle; sternellar lobes of the 
thoracic segments moderately prominent and with distinct foot calli. 

Larval type labeled '^Webb 
No. 2." 

The most distinctive charac- 
ters separating the larva of this 
species from that of the preced- 
ing one are the more shining 
mandibles, with much deeper 
dorsal impression and more 
prominent oblique ridge and 
the much less distinctly ele- 
vated anterior margin of the 
epicranium. 

Galleries (fig. 81). — The gal- 
leries of this species are very 
much the same as those of the 
preceding one, except that the 
larval mines are more distinctly 
contiguous for a greater dis- 
tance from the egg galleries. 
Distrihution (fig. 82). — 
(Hopk. U. S.) Arizona: Cliiri- 
cahua Mountains. Colorado: 
Clyde, Boulder, Craig, Fort Col- 
lins, Glenwood Springs, Gunni- 
son, Hahn's Peak, Holy Cross 
National Forest, Meeker, Ou- 
ray National Forest, San Isabel 
National Forest, Steamboat 
Springs, White River National 
Forest. Neiv Mexico: Capi- 
tan Mountains, Sierra Blanca 
Mountains, Sacramento Na- 
tional Forest. South DaJcota: 
Spearfish Canyon, Black Hills. 
Utah: Ephraim. ^yyoming: En- 
campment. Additional localities from specimens in other collec- 
tions : (Horn) "H. B." (Northwest Territory, probabl}^ in Macken- 
zie Eiver region) and Alta, Utah. (Wickham) Argentine, Leadville, 
and Silver Plume, Colo. (Cockerell) Las Vegas, N. Mex. (H. & S.) 
Calgary, Alberta; Glacier, British Columbia. (Webb) Collins, Idaho. 




Fig. 81.— Dendroclonus engelmanni: Egg gallery in liv- 
ing bark. A, Normal; B, boring dust removed; a, 
entrance; 6, basal section; c, boring dust packed in 
gallery; d, subsequent or inner gallery; e, venti- 
lating burrow;/, egg nest, with and without eggs; g, 
freshly hatched larvae; h, pits in roof of gallery. 
(Original.) 



THE GENUS DENDEOCTONUS. 



133 



Host trees. — Picea engelmanni and P. canadensis. 

Identified specimens. — Le Conte collection, 2 specimens ; Horn, 2 ; 
Dietz, 1 (Utah); U.S.N.M. (H. & S.), 4; Wickham, 4; Cockerell, 7; 
Webb, 25; Hopk. U. S., more than 200 specimens, including all 
stages and work. 




Fig. S2.—Dendroctonus engelmanni: Distribution map. (Original.) 



Bibliography and Synonymy. 

Dendroctonus rufi-pennis (not of Kirby) Le Conte, 1876, p. 385, revision (?). Le 

Conte, 1878, p. 469 (in part) (in collection 1900-1907, under D. rufipennis). 

Packard, 1887, pp. 177-243 (in part). Packard, 1890, p. 721 (in part), p. 722 (in 

part), quotes Le Conte. Dietz, 1890, p. 30, Colorado, Utah. Wickham, 1902, p. 

309, list, localities. 
Dendroctonus dietzi Hopkins, 1902a, manuscript name only for variation (Utah). 
Dendroctonus californicus Hopkins, 1902a, p. 3, manuscript name only, locality. 
Dendroctonus wickhami Hopkins, 1902a, p. 3, manuscript name only. 
Dendroctonus piceaperda (not of Hopkins) Wickham, 1902, p. 310, in list, locality, 

host, reference to synonymy. Hopkins, 1906a, pp. 4, 5, old work on Pike's Peak. 
Dendroctonus piceaperda var. engelmanni (Hopk.) Fall, 1907, p. 218, manuscript name, 

list, localities. 
Dendroctonus piceaperda (not of Hopkins) var. Fall, 1907, p. 218, list, Cloudcroft, 

N. Mex. 
The Engelmann spruce beetle. Hopkins, 1908, pp. 161-162, depredations. 

16. Dendroctonus borealis n. sp. 

(PI. V, fig. 16.) 

Adult. — Type of species, female: Length 6 mm., nearly black. 
Head with front convex, with faint anterior and posterior impressions 
and faint anterior line; elytral declivity with striae not deeply im- 



134 



THE SCOLYTID BEETLES. 



pressed; punctures of pronotum distinctly irregular; posterior half 
of proepisternal area not punctured; punctures of pronotum and 
elytra moderately coarse; elytral striae scarcely impressed except in 
dorsal area; interspaces scarcely coarser and but faintl}?^ rugose, 
except toward base and vertex. Secondary sexual characters : Elytral 
declivity convex; strise faintly impressed, with punctures moderately 
distinct; interspaces slightly convex, with distinct row of granules. 

Type labeled ''Type No. 7453 U.S.N.M.," name label, "Hopk. 
1/22/08, U.S.N.M. 22, Alaska, 9 , U.S.N.M. Ace. 25431." 

Male type: Length 6 mm., elytra red; thorax, head, and abdomen 
much darker; other characters the same as in female, except elytral 
declivity, which is shining, the strise and strial punctures obscure; 




' S type," name label, 
Alaska, W. H. Osgood, 



Fig. 83. — Dcndroctonus horealis: Distribution map. (Original.) 

interspaces flat, shining, and 1 and 2 without granules except toward 
vertex. 

Male type labeled "type of drawing," 
"Hopk. 1/22/08, Picea canadensis, Eagle 
collector, $ , Hopk. U. S. 1170a." 

Variations. — There is scarcely any variation in the four specimens 
in the collections, but the color varies from nearly black in the type 
to the head, thorax, and abdomen dark, and the elytra red in the other 
specimens. 

DistinguisTiing characters. — The short, stouter form, short and broad 
pronotum, with the punctures more uniform in size, the punctures of 
the dorsal strise of the elytra finer and less distinct, serves to distin- 
guish this species from all of the allied forms. It appears to be more 
closely allied to D, ohesus, hnt is distinguished from it by its shorter 
pronotum and elytra, and the other characters mentioned. 



THE GENUS DENDROCTONtJS. 135 

Revisional notes. — It is quite evident that the specimens described 
byMannerheim(1853, p. 238) under Z>. rujlpennis' sive D. horealis. The 
single specimen in Le Conte's collection under D. rufipennis, labeled 
" Hylurgus rujipennis Kjrby," and locality Kenai, is evidently from 
Mannerheim's collection, and probably one of the specimens before 
him when he prepared his description under that name. Superfi- 
cially, this specimen resembles D. piceajjerda, which led Le Conte to 
identify his Anticosti and Canada specimens as D. rujipennis, and is 
evidently the one which represented the Alaska locality in his revi- 
sions (1868 and 1876). 

The immature stages and galleries of this species have not been 
observed. 

Host tree. — Picea canadensis. 

Distribution (fig. 83). — AlasJca: Eagle, (?) Kenai Peninsula. 

Identified specimens. — Le Conte, 1 specimen; U.S.N.M., 2; Hopk. 
U. S., 2, collected by W. H. Osgood, of the Biological Survey, U. S. 
Department of Agriculture, at Eagle, Alaska, August, 1903, from 
white spruce. 

Bibliography and Synonymy. 

Hylurgus rujipennis (not of Kirby). Mannerheim, 1853, p. — (in part). 
Dendroctonus rujipennis (not of Kirby). Le Conte, 1868-1876 (in part). 
Dendroctonus borealis Hopkins, 1902a, p. 3, manuscript name only. 

17. Dendroctonus obesus (Mannerheim). 

(PI. Vl.flg. 17.) 

Adult. — Typical female: Length 6.5 mm., nearly black. Head with 
front convex, with faint anterior and posterior impression and mod- 
erately distinct anterior line. Elytral declivity with striae not deeply 
impressed; punctures of pronotum distinctly irregular; posterior half 
of proepisternal area not punctured; punctures of pronotum and 
elytra moderately coarse; elytral strige scarcely impressed; inter- 
spaces flat, finely, sparsely rugose on dorsal area and toward base of 
vertex. Secondary sexual characters: Declivity convex, subopaque; 
striae faintly impressed, with punctures moderately distinct; inter- 
spaces faintly convex, with distinct row of granules. 

Typical female labeled, name label, "Hopk, 1/22/08, Picea sitcli- 
ensis, Queen Charlotte II., Keen [collector], 9 ." 

Typical male: Length 6.7 mm., black. Front convex, with faint 
anterior impression and distinct anterior line. Agrees with female, 
excepting that the punctures of elytral striae and interspacial rugosi- 
ties are coarser; declivity subopaque; strial impressions and punc- 
tures obscure; interspaces flat, faintly punctured, and with a few 
granules toward vertex; pronotum with distinctly elevated line. 

Typical male labeled, name label, " 1/22/08, Picea sitcJiensis, Queen 
Charlotte II., J. H. Keen, Collr., S ." 



136 THE SCOLYTID BEETLES. 

Variations. — The length varies from 6 to 7 mm., with the average 
about 6.5 mm. The color ranges from uniform light red to brown in 
young specimens, to uniform black in matured ones, it being exceed- 
ingly rare to find examples with the pronotum darker than the elytra, 
which is so characteristic in the three preceding species. The sculp- 
ture and vestiture of the epistoma, front, pronotum, and elytra vary 
as usual. The greatest variation is in the punctures of the pronotum 
and in the presence and absence of the dorsal line; the presence or 
absence of a frontal carina is also an important variation, and in some 
examples the body is noticeably more elongate than in others. 

Distinctive characters. — The characters which serve to distinguish 
this species from the three preceding are the uniform black color of 
the matured adults and the prevailingly less impressed elytral striae, 
especially those of the lateral area, and also the prevailing slightly 
more elongate form. Its host tree and distribution also serve as 
distinguishing characters, except, perhaps, in the case of horealis, 
which may be found in the Sitka spruce. 

Bevisional notes. — There can be little or no doubt that the material 
under observation represents Mannerheim's species, whose varieties 
a, i, and c were evidently immature specimens. D. similis Lee. is 
to be referred to this, which fact was recognized by Le Conte in his 
1868 paper, but the beetle was subsequently confused with the 
species discussed in the present paper under D. fseudotsugse. D. ohesus 
is represented in the Le Conte collection by two specimens, one 
specimen from Mannerheim's collection, labelled D. ohesus (''Speci- 
men 5" under D. rujlpennis in 1900), and one specimen, the type 
of D. similis. It is possible that "specimen 2" under D. ruji'pennis 
is also D. ohesus, but was not recognized by the writer when ex- 
amined in 1900. Dietz, 1890, did not recognize or mention D. ohesus, 
and it was not found by the writer in the Horn collection or that of 
the Academy of Natural Sciences. 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the apices of the front and middle tibiae are smooth or 
rarely with a single granule, abdominal tergites 2 to 6 with very 
small pleural spines, 1 without dorsal but with small lateral spines, 
and 2 to 6 with dorsal and lateral ones, 7 and 8 smooth, 9 with 
prominent pleural spine as usual. Pupal type labeled ''Hopk. U. S. 
No. 4049a." 

The usual variation in minor details prevails, but the pupa of 
this species is distinguished from that of the three preceding by the 
more evident lateral spines of the first abdominal tergite and the 
prevailing darker tips to the abdominal spines. 

Larva. — In addition to the generic, divisional, subdivisional, and 
sectional characters, the front has a faint transverse elevation dis- 



THE GENUS DENDEOCTOlSrUS. 



137 



tinctly in front of the middle and the pronotal area is fiat to apex. 
The clypeus is sliort and broad, with an anterior dorsal groove and 
the apex broadly emarginate, labium prominent, slightly longer than 
the clypeus, with the apex broadly rounded. Larval types, labeled 
"Hopk. U. S. Nos. 4081, 4046a, and 4049a." 

The most distinctive characters appear to be the anteriorly placed 
transverse elevation of the front ; the characters of the mandibles and 
anterior margin of the epicranium more nearly approach those found 
in D. engelmanni. 

Galleries. — The galleries of this species are of the same, or of 
similar character to those of D. piceaperda, as given in divisional, 




Fig. 84.— Dendroctonus obesus: Distribution map. (Original.) 

sub divisional, or sectional characters, but differ in the more extended 
common larval chamber which precedes the independent larval 
mines which are usually so confused by crossing each other that they 
are difficult to follow. 

Distribution (fig. 84). — (Hopk. U. S.) Oregon: Newport. Wash- 
ington: Hoquiam, Aberdeen. Additional localities from other col- 
lections: (U.S.N.M.) (H. & S.) Vancouver, British Columbia; (Rev. 
Keen) Queen Charlotte Islands, British Columbia. 

Host tree. — Picea sitcJiensis. 

Identified specimens. — Le Conte collection, 2 specimens : U.S.N.M., 
1 from Doctor Fletcher; H. & S., 3; Hopk. U. S., more than 120 
specimens, including all stages. 



138 THE SCOLYTID BEETLES. 

Bibliography and Synonomy. 

Hylurgus obesus « Mannerheim, 1843, p. 296, original description. Mannerheim, 
1852, p. 356, spec. 474, list, variety b and variety c, brief descriptions. Man- 
nerheim, 1853, p. 238, separate p. 146, list, var. d described. Le Conte, 1868, 
p. 173, mentioned, synonymy. 

Dendroctonus similis Le Conte, 1860, p. 59, description (from one specimen ■which= 
D. obesus). Le Conte, 1868, p. 173, mentioned as synonymous with Z). o&esws Mann. 
Hopkins, 1902a, p. 3, recognized as synonymous with D. obesus (Mann.). 

Dendroctonus obesus (Mann.) Le Conte, 1868, p. 173 (in part). Chapuis, 1869, p. 35; 
1873, p. 243, revised description. Hamilton, 1894, p. 35 (in part). Hopkins, 
1899b, pp. 15, 21, habits, host, etc. Schwarz, 1900a, p. 537, author's reprint p. 
185 (in part), list. Hopkins, 1902a, p. 3, species recognized as distinct from D. 
nifipennis Kirby. Hopkins, 1902c, p. 22, habit and host. Hopkins, 1903a, p. 
60, reference. 

Dendroctonus rufipennis (not of Kirby). Le Conte, 1868, p. 173 (?=Z). obesus). 
Le Conte, 1876, p. 385, revision, synonymy, bibliography, localities (in collection 
1900-1907, under rufipennis). Packard, 1887, pp. 176, 243 (in part?). 

Dendroctonus rufipennis (obesus Mann.) Harrington, 1890, p. 189, author's extra, p. 19. 

Dendroctonus heeni Hopkins, 1902a, p. 3, manuscript name only on variation. 

Dendroctonus fletcheri Hopkins, 1902a, p. 3, manuscript name only on variation. 

18. Dendroctonus rufipennis (Kirby). 

(PI. VI, fig. 18.) 

Adult. — Typical female: Length 6.2 mm. Elytra red; thorax, 
head, and abdomen reddish brown. Head convex, with faint ante- 
rior and posterior impression and short anterior line; elytral declivity 
with striae not deeply impressed; punctures of pronotum distinct, 
irregular; posterior half of proepisternal area punctured; strise of 
elytral declivity with fine punctures; elytral strise scarcely impressed, 
except toward suture; strial punctures moderately coarse; inter- 
spaces with rugosities moderately coarse, sparse, and acute; pronotal 
punctures coarse, deep, moderately dense. Secondary sexual char- 
acters: Elytral declivity convex; striae faintly impressed; punctures 
obscure; interspaces faintly convex, with row of fine granules. 

Typical female labeled "type of drawing," name label, "Hopk., 
4/25/02. Agrees with Kirby's type, compared by C. O. Waterhouse, 
H. S. 28, 9 , White Fish Point, L[ake] S[uperior]." 

Typical male: Length 6.7 mm. Elytra dark red, thorax and head 
reddish brown. Agrees with female, excepting that the elytral 
declivity is more shining, the strial punctures are less distinct, and thej 
interspaces have less distinct rows of granules. 

Typical male labeled, name label, "Hopk, 1/22/08, Pinus strobus, 
Grand Island, Mich., W.F.Fiske, collector, <? , Hopk. U. S. 376L" 

Variations. — The length varies from 5 to 7.3 mm., with the average 
at about 6.5 mm., the head, thorax, and ventral segments froml 

a This was Eschscholtz's manuscript name, published in Dejean Cat., 3me. Edit.,] 
p. 331, but Mannerheim published the first description. 



THE GENUS DENDEOCTONUS. 139 

darker reddish brown to nearly black, while the elytra are light to 
dark red. The greatest variation noted in the few specimens under 
observation is in size, with less variation in sculpture and vestiture 
than in the preceding species, Nos. 14, 15, and 17. 

Distinctive characters. — The characters which serve to at once dis- 
tinguish this species from the preceding allied ones are the coarse 
punctures of the posterior section of the proepisternal area, the more 
distinctly red elytra, the bright-red hairs, and the much less distinctly 
impressed elytral strisB of the lateral area. 

Revisional notes. — There is quite extensive literature under the 
name Hylurgus rujipennis Kirby and Dendroctonus rujipennis Kirby. 
Apparently no part of it except the original description refers to 
Kirby's species, and even the type series in the British Museum repre- 
sents at least one other species. It appears that up to the fall of 1906 
the only representative of the species in the collections of this country 
was a female specimen in the Hubbard & Schwarz collection, U. S. 
National Museum, labeled ' ' White Fish Point, L. S. " This, with other 
specimens of the Dendroctonus ol the U. S. National Museum and Hub- 
bard & Schwarz collections, was submitted to the writer in December, 
1898, for study, and was then labeled "H. S. 28." In 1900 this speci- 
men, together with another labeled ''H. B." (Northwest Territory) 
from the National Museum collection, and some specimens collected 
by the writer from the spruce in Maine, were sent to the British 
Museum for comparison with Kirby's type of D. rujipennis. They 
were compared by Mr. Charles O. Waterhouse who, in a letter dated 
November 1, 1900, wrote as follows: 

I have examined your species, but am only concerned with your two largest speci- 
mens. We have three of the specimens which Kirby had before him, all marked 
exactly alike. The one to which he attached his ticket is a dark-brown variety (unless 
it is stained with grease), but in all other respects agrees with your H. S. 28, with 
fairly equally distributed punctuation on the thorax. Kirby's two other specimens 
have red elytra and agree with your H. B. 7401, 824, and have a closely punctured 
impression or flattening at the base of the thorax. 

The specimen in the type series which bore the name label when the 
comparison was made should be recognized as the type. Thus it is 
quite certain that our H. S. 28 from White Fish Point, Lake Superior, 
is a true representative of the species, while the two other specimens 
with which our H. B. 7401, 824, etc., agree evidently represent D. en- 
gelmanni and are probably the specimens referred to by Kirby as 
coming from Lat. 65°. The writer is informed by Mr. Schwarz that 
our H. B. (No. 7401) specimen came from about the same latitude. 
The other specimens from Maine, which were so different from the 
type as to be at once recognized as distinct, were representatives of 
D. piceaperda Hopk. 

The pupse, larvae, and galleries have not been observed. 



140 



THE SCOLYTID BEETLES. 



Distribution (fig. 85). — Michigan: White Fish Point and Grand 
Island. 

Host tree. — Pinus strohus. 

Identified specimens. — U.S.N.M.,H. & S., 1 specimen; Hopk. U. S., 
14 specimens of adults, collected by W. F. Fiske at Grand Island, Mich. 

Bibliography. 

Hylurgus rufipennis Kirby, 1837, p. 195, No. 261, original description. Packard, 
1887, p. 176, note. Hopkins, 1899c, p. 343 (in part?). Schwarz, 1900a, p. 537, 
author's copy, p. 185 (in part?). Hopkins, 1901a, p. 16, reference to distribution, 
characters, first time recognized since description. 




Fig. 85. — Dcndroctonus rufipennis: Distribution map. (Original.) 
. 19. Dendroctonus murrayanae n. sp. 

Adult. — Type of species, female: Length, 6.9 mm; elytra red; 
thorax, head, and abdomen nearly black. Head convex, with obscure 
impression; elytral declivity with striae not deeply impressed; punc- 
tures of pronotum distinct, coarse, irregular; posterior half of pro- 
episternal area punctured; striae of el3^tral declivity with fine 
punctures; elytral striae scarcely impressed; punctures moderately 
coarse; interspaces with rugosities moderately coarse, acute, and 
rather coarsely placed; pronotal punctures coarse, deep, moderately 
dense. Secondary sexual characters: Eh'tral declivity convex; 
strige distinctly impressed; punctures obscure; interspaces convex, 
with irregular punctures and rows of granules. 

Type labeled "No. 7454 U.S.N.M.," name label, "Hopk. 1/22/08, 
Pinus murrayana, Keystone, Wyo., J. L. Eebmann, collector, 9 , 
Hopk. U. S. 2690." 



THE GENUS DENDEOCTONUS. 



141 



Male type: Length, 6.5 mm. Agrees with female, except that the 
elytral declivity is more shining and the interspaces are more dis- 
tinctly punctured and less rugose. 

Type labeled " $ type," otherwise same as female. 

Variations. — The length varies from 5.4 to 6.5 mm., with the aver- 
age about 6 mm.; the head, thorax, and ventral segments are dark 
reddish brown to black, while the elytra are light to dark red. The 
greatest variation noted in the few specimens under observation is in 
the size, with less variation in sculpture and vestiture than in species 
Nos. 14, 15, and 17. 

Distinctive characters. — The characters which serve to distinguish 
this species from B. rujipennis, to which it is more closely allied, are 
its slightly smaller size and more distinctly impressed elytral striae 




Fig. 86. — Dendroctonus murrayanx: Distribution map. (Original.) 

on the dorsal and lateral areas and the finely, densely punctured 
interspaces of the declivity in the male. 

It appears that this species has not been referred to in literature. 

The .pupa has not been observed. 

Larva (PI. VIII, fig. 19). — In addition to the generic, divisional, 
and subdi visional characters, the front has an impression toward the 
anterior angles and a faint transverse elevation in front of the middle. 
The clypeus is longer than the labrum and marked with a median 
groove, and with the apex broadly emarginate; labrum with apex 
faintly emarginate, and the mandibles with distinct dorsal impres- 
sion. Larval type labeled "Hopk. U. S. No. 2690c." 

Galleries. — Only fragmentary specimens of the galleries have been 
observed, but they appear to come between piceaperda and valens, 
the larvae, for the most part, excavating a common chamber. 



I 

142 THE SCOLYTID BEETLES. 

Distribution (fig. 86). — (Hopk. U. S.) Wijoming: Homestake, Sar- 
atoga, Keystone, Cheyenne National Forest, and Big Horn National 
Forest. Colorado: Jefferson. Additional localities from other col- 
lections: (U.S.N.M.) (H. & S.), National Park, Wyoming. 

Host trees. — Pinus murrayana and Picea engelmanni. 

Identified specimens. — H. & S., 5; Hopk. U. S., over 100 specimens, 
including adults, larvae, and work. 

Bibliography. 
Dendroctonus shoshone Hopkins, 1902a, p. 3, manuscript name only. 

20. Dendroctonus punctatus Le Conte. 

(PI. VI, fig. 20.) 

Adult. — Typical female: Length 6.5 mm., dark reddish brown. 
Head convex, with moderately distinct anterior impression. Elytral 
declivity with striae distinctly impressed. Punctures of pronotum 
distinct, coarse, irregular; posterior half of proepisternal area punc- 
tured; striae of elytral declivity with coarse punctures; elytral striae 
distinctly impressed; punctures coarse and distinct; interspaces 
rather narrow, convex, moderately rugose on dorsal area, but nearly 
smooth on lateral area. Secondary sexual characters: Declivity 
convex, shining; striae distinctly but not deeply impressed; punc- 
tures coarse and distinct; interspaces narrow, convex, with row of 
granules. 

Typical female labeled 'Hype of drawing," name label, ''Hopk. 
4/25/02, Picea rubens, Randolph County, W. Va., A. D. Hopkins, 
collector, 9 , Hopk. W. Va. 6312." 

Variations. — There appears to be very little variation in the few 
specimens observed. 

Distinctive characters. — This species is at once distinguished from 
all of the preceding by the coarse punctures of the el3"tral striae, 
especially on the declivity. Its nearest ally is D. micans, of Europe, 
from which it differs in its smaller size, more elongate form, and' 
more distinctly impressed elytral striae. 

Revisional notes. — While the original description does not include 
the more distinctive characters, it was based on a type which was 
readily recognized as a distinct species. Dietz referred the specimen 
in the Horn collection, labeled D. punctatus Lee, to D. rvfipennis 
(Kirby), but it is certainly distinct from what is now recognized as 
D. rufipennis. 

Host tree. — Picea rubens. 

Pupae, larvae, and galleries of this species have not been observed. 

Distribution (fig. 87). — (Hopk., W. Va.) 1 specimen collected by 
the writer May 21, 1892, in the high mountains of Randolph County, 



THE GENUS DENDKOCTONUS. 



143 



West Virginia, under bark on spruce stump. (Le Conte, M. C. Z.) 
3 specimens labeled ''New York"; (Horn, A. E. S.) 1 specimen 
labeled ''Pa." 

Bibliography and Synonymy. 

Dendroctonus punctatus Le Conte, 1868, p. 173, original description, synonymy, 
locality. Le Conte, 1876, p. 385, revision, synonymy, bibliography, localities. 
Packard, 1887, p. 177, Le Conte quoted. Packard, 1890, p. 722, Le Conte quoted. 
Hopkins, 1899a, p. 447, habit, etc.. West Virginia. Hopkins, 1902a, p. 3, recog- 
nized as a good species and restored. 

Dendroctonus rufipennis (not of Kirby) Dietz, 1890, p. 30, Pennsylvania. 




Fig. 87. — Dendroctonus punctatus: Distribution map. (Original.) 



21. Dendroctonus micans (Kugelann). 

(PI. VI, fig. 21.) 

Adult. — Typical female: Length 7.25 mm., dark reddish brown. 
Head convex, with faint anterior impression, without anterior line; 
elytral declivity with striae not deeply impressed; punctures of 
pronotum distinct, coarse, irregular; posterior half of proepisternal 
area punctured; striae of elytral declivity with coarse punctures; 
elytral striae not impressed, punctures moderately coarse, interspaces 
broad and fiat, sparsely and finely rugose. Secondary sexual charac- 
ters: Elytral declivity convex, subopaque; strige faintly impressed; 
punctures coarse and distinct; interspaces moderately convex, with 
sparse, irregular granules. 

Typical female labeled "type of drawing," name label, "Hopk. 
4/25/02, 9 . Determination No. 20, Eichhoff, Sachsen, Horrung." 



144 THE SCOLYTID BEETLES. 

Typical male: Length 6 mm., black. Agrees with female in every 
respect, except that it may have stouter mandible, the club of antenna 
smaller, more elongate ; the declivity of elytra with strial punctures 
finer, and the interspaces without granules. 

Typical male labeled ^^ $ , from Dr. Severin." 

Variations. — Length 7 to 8 mm., average about 7.5 mm.; uniform 
reddish brown to nearly black, with usual variation in sculpture and 
vestiture. 

Distinctive characters. — This species is more closely allied to D. 
punctatus than to any of the other species of the genus, from which 
it is distinguished by its larger size and stouter form, with the striae 
scarcely at all impressed, and the punctures smaller. 

The male of this species appears to be far more rare than in the 
other species, from the fact that among 83 specimens examined only 
2 males were found. While the declivity is somewhat more shining 
and smoother in the male, this character is by no means as striking 
as in the other species of the section to which it belongs. 

The pupa has not been studied by the writer, but is evidently 
similar in general character to that of D. piceaperda. 

Larva. — ^Abdominal tergite 8 without, 9 with, small dorsal plate, 
which is not rugose. Front with distinct elevation. In addition to 
the generic, divisional, and subdivisional characters, the frontal 
elevation is subopaque, transversely wrinkled, situated in front of 
the middle and joined to the epistoma; lateral angles are curved 
back to their junction with the frontal sutures, which are broadly 
curved toward the apex. The area behind the elevation is broad, 
flat, and more shining. Clypeus broad, with faint median groove 
and the apex broadly emarginate. Labrum small, rather stout, with 
broad dorsal impression, the "apex subtruncate; mandibles shining, 
with a distinct dorsal impression and oblique ridge near the middle; 
sternellar lobes of the thoracic segments moderately prominent and 
with distinct foot calli. 

Type. — One of a large series of larvae received from Dr. G. Severin, 
conservateur, Royal Museum of Natural Histor}^, Belgium. 

The larva of this species is at once distinguished from that of all 
of the other species of the genus, so far as observed, by the faint 
dorsal plate of the ninth abdominal segment, by the absence of a 
plate on the eighth, by the frontal elevation connected with the 
epistoma, and by the greater number and more distinct hairs on the 
scutellar lobes of the thorax and abdomen. 

Galleries (fig. 88). — The galleries are evidently quite similar to 
those of D. terebrans and D. valens, especially in the fact that the 
larvae live together in a common chamber exposed in the inner bark. 
According to Dr. G. Severin, the egg gallery is vertical, frequently 



THE GENUS DENDEOCTONUS. 



145 



curved and somewhat irregular, sometimes doubly inflected and 
from 12 to 20 cm. long. The female here deposits from 20 to 25 
eggs in several places. The larvae eat very close together, growing 
equally in size and age and making a common cavity underneath 
the bark. In order to go through the pupal stage, they return to 
the large space which they left behind them and which is now filled 
with excrement and resin. Departing from their common cavity, 
they eat out isolated galleries, and at the end of these they pupate. 




Fig. 8S.—Dendroctonus micans: Egg galleries and larval chamber. ^, Basal sections of egg galleries; B, 
advanced stage of worlc; a, entrance burrow; b, excavated July 8-16; c, excavated July 8-29; d, eight 
days old; e, three weeks old; /, basal section; g, boring-dust; h, subsequent or inner gallery ("mother 
gallery"); j,egg nest with eggs scattered about in boring-dust; k, social chamber excavated by larvse; 
I, boring-dust and resin; vi, larvae at work. (Adapted from Pauly Forstlich-natur wissenschafthche 
Zeitschrift, I Jahrgang, figs. 3 and 4.) 

Distribution (fig. 89). — According to the literature, this species 
ranges from central to northern Europe and from Denmark and 
Russia eastward into Siberia. 

Host trees. — It is said to infest Pinus, Picea, Abies, and Larix. 

Identified specimens. — The writer has examined 1 specimen received 
from W. Eichhoff, 8 from B. W. Schlick, Denmark; 2 with specimens 
of work from Reitter's collection, collected in Bohemia, and about 70 
specimens of adults, as well as larvae and specimens of work from 
Dr. G. Severin, of the Musee Royale d'Histoire Naturelle, Brussels, 
Belgium. 



79980—09- 



-11 



146 



THE SCOLYTID BEETLES. 



Bibliography and Synonymy. 

IBostrichus ligniperda Herbst, 1793, p. 107 (in part). 

Bostrichus micans Kugelann, 1794, p. 523, original description. 

?Hylesinus ligniperda Gyllenhal, 1813, pp. 335, 336 (in part). 

Dendroctonus micans (Kug.) Erichson, 1836, p. 53, type of genus. Bach, 1849, p. 144. 
Stein, 1854, pp. 277-279, habits, destructiveness. Kollar, 1858, pp. 23-28, habits, 
control. Eichhoff, 1864, p. 27, pi. 1, fig. 5, tarsi, fig. 6, maxilla, fig. 7, labium, in 
revision of genus. Lacordaire, 1866, p. 360, in revision of genus. Chapuis, 1869, 
p. 35; 1873, p. 243, revised description. Lindemann, 1875, pp. 213, 221, pi. 1, figs. 
1-10, male reproductive organs described and illustrated. Eichhoff, 1881, 
pp. 125-128, fig. 23, adult, fig. 24, galleries, revision, bibliography, account 
of habits, distribution, etc. Altum, 1881, pp. 262-266, description, biology. 
Judeich and Nitsche, 1889, pp. 458-462, life history, habits, importance, remedy. 
Pauly, 1892, pp. 315-327, 4 figs, of galleries, habits. Verhoeff, 1896, pp. 124-133, 
anatomy. Menegaux & Cochon, 1897, pt. 2, p. 120, habits, etc. Severin, 1902, 
p. 145, habits in Belgium. Weber, 1902, p. 108, fig. 5, enemy. Brichet et 
Severin, 1903, pp. 244-258, habits, etc. Baudisch, 1903, pp. 151-152, habits, etc. 
Quairiere, 1904, pp. 626-628. Niisslin, 1905, pp. 175-178, habits, description, 
importance. Quievy, 1905, pp. 334, 335. Severin, 1908, pp. 1-20, description, 
habits, depredations, control. 

Hylesinus micans (Kug.) Batzeburg, 1839, p. 217, Taf. VII, fig. 3, adult. 

Hylesinus {Dendroctonus) micans (Kug.) Ratzeburg, 1839, p. 217, Taf. VII, fig. 3, 
adult; Taf. VIII, figs. 1, 2, 3, galleries, young larvse, pupae. 




Fig. %%.— Dendroctonus micans: Distribution map. (Original.) 
Subdivision D. 

The distinguishing characters common to the species of the fom'th 
subdivision are: 

Adult. — Front without median or posterior impression. Prono- 
tum somewhat elongate, shghtly narrower than elytra, moderately 
constricted toward head, with regular punctures or without coarse 
and fine punctures intermixed, long hairs absent on median and 
posterior dorsal areas, present on anterior and lateral areas; head 



THE GENUS DENDROCTONUS. 147 

broad, convex; epistomal process and elytral rugosities variable 
within the same species. 

Sexes. — Females with front of head moderately broad; mandibles 
shining, moderately stout; antennal club broad and stout; elytral 
declivity slightly more rugose; striae impressed with distinct punc- 
tures. 

Males with front of head distinctly broader; mandibles opaque, 
stout; antennal club narrow, more elongate, and the elytral declivity 
slightly less rugose; strise less distinctly impressed, and the punctures 
more obscure. 

Pupa. — Vertex of head convex; front and middle femur each with 
a minute subapical spine; abdominal tergites with moderately dis- 
tinct spines. 

Larva. — Abdominal tergites 8 and 9 with distinct dorsal plates, 
each armed with three prominent teeth. 

Galleries. — Egg galleries slightly winding to nearly straight ; larval 
mines not separated, except very rarely near the outer extremity, 
but forming broad common larval chambers. 

22. Dendroctonus terebrans (Olivier). 

(PI. VII, fig. 22.) 

Adult. — Typical female: Length 5.6 mm., black. Front convex, 
without impressions; epistomal process moderately broad, lateral 
angles tuberculate. Pronotal punctures very coarse, regular, mod- 
erately dense, scarcely decreasing in size toward base; elytra with- 
out long hairs toward base. 

Typical female labeled, name label, "Hopk. 1/22/08, Pinus ecJii- 
nata, Hopkins, collector, Tryon, N.C., 9 , Hopk. U. S. 530aa." 

Typical male: Length 5.6 mm. Differs from female in stouter 
mandibles and slightly coarser rugosities of elytral declivity. 

Typical male labeled same as female. 

Variations. — The length varies from 5 to 8 mm., with the average 
about 7 mm. The color ranges from piceous to deep black, the latter 
prevailing. Immature specimens are reddish, but fully matured 
ones are always darker than the darkest D. valens. The greatest 
variation is in size, and while the usual variation prevails in some of 
the other characters, it is much less so than in D. valens. In New 
Jersey, Pennsylvania, Virginia, and West Virginia, where there is an 
overlapping of the range of D. valens and D. terebrans, specimens are 
sometimes found which appear to be hybrids, but it appears that the 
more dominant characters of D. terebrans prevail in such hybrids, so 
that the darker color and coarse punctures of the pronotum serve to 
distinguish them as being more closely allied to the latter species. 

Distinctive characters. — The characters which serve to distinguish 
this species from D. valens, to which it is more closel}^^ allied, are its 



148 THE SCOLYTID BEETLES. 

prevailing black or dark color, the more uniform and coarser punctures 
of the pronotum, the narrower epistomal process, with the angles more 
tuberculate, and the less evident long hairs on the anterior dorsal 
area of the elytra. 

Revisional notes. — While the type of this species has not been seen 
by the writer, it is clearly evident from Olivier's description and 
figures (Olivier, 1795) that the large black form common to the 
southern United States represents the species described. The only 
distinctive specific character mentioned, however, is the reference to 
the Black Scolytus and to the body being black, brown, or brownish- 
black. The confounding of Dendroctonus valens and D. terebrans under 
the latter name has resulted in much confusion in the literature. With 
our present knowledge, however, it is not difficult to clear up some 
of the confusion and to revise and correct the literature so that it 
may be known in many cases whether or not one or both species was 
included in a given reference. Erichson, 1836, Lacordaire, 1866, and 
Chapuis, 1869, evidently did not compare D. valens and D. terebrans. 
While a specimen of this species has been in the Harris collection since 
1839, Harris apparently made no reference to its characters. Zimmer- 
man, 1868, page 149, did not mention D. valens, but evidently had the 
two species confused in his revised description. Le Conte, 1868, page 
173, referred D. valens to D. terebrans, and in 1876, pages 384-385, 
confuses the characters and distribution of the two species. Dietz, 
1890, page 29, included this species under his variety a, and (p. 30) 
evidently includes two specimens from Florida under his revision of 
D. rujipennis. In subsequent literature up to 1906 there is more or 
less confusion of this species with D. valens. The writer, 1906c, page 
81, restored D. valens Lee. and called attention to the characters dis- 
tinguishing Z>. terebrans (Oliv.) . In 1900 the writer found one specimen 
in the Harris collection, under Hylurgus terebrans, under his No. 99, 
referred to in his note as "Dark specimen abundant under bark of 
pitch pine, October 27, 1839," but it appears that no reference was 
made to this dark specimen in any of his publications. The locality 
is not given, but it is presumably Cambridge. It appears that this 
species was not represented in the Horn collection under I), terebrans 
when Doctor Dietz prepared his revision, and that the only example 
involved in the revision under D. terebrans was the one in the Ulke 
collection from Pennsylvania, designated as "variety a." Two exam- 
ples were found in the Horn collection under D. rujipennis, labeled 
"Fla.," and it was evidently on these that Doctor Dietz based his 
Florida locality in his revision of D. rujipennis. In 1907 this species 
was represented in the Le Conte collection by 9 specimens and 8 addi- 
tional specimens in the general collection of the Museum of Com- 
parative Zoology, 



THE GENUS DElSTDEOCTOlSrUS. 149 

Pupa. — In addition to the generic, divisional, and subdivisional 
characters, the front and middle femora are armed each with a minute 
apical spine; abdominal tergites 1 to 6 have moderately small pleural 
spines, 1 is without dorsal spine, but with distinct la:teral ones; 2 to 6 
have small dorsal and lateral spmes, the former increasing in size to 6, 
All have pale tips; 7 and 8 unarmed; 9 with usual stout pleural spine. 
Pupal type labeled "Hopk. W. Va. 7701." 

The usual variation prevails in the number and arrangement of 
minor spines and between the young and older examples. 

The character which in general serves to distinguish the pupa of 
this species from the preceding is found m the paler tips of the body 
spines. 

Larva. — In addition to the generic, divisional, and subdivisional 
characters, front of head with posterior angle and median area not 
elevated but transversely rugose except near apex, where it is smooth ; 
epistoma flat, opaque, smooth, with straight anterior margin; clypeus 
broad, promment, convex, with faint median longitudinal line, sides 
rounded, apex broadly emarginate; labrum short, with sides nearly 
parallel and apex tuberculate. Prothoracic tergum with two broad, 
shining dorsal plates separated by a rather broad median space, and 
a smaller lateral plate each side ; sternellar lobes each with a faint foot 
callus; mesoterga and metaterga with shining plates on the lateral 
lobes. Abdommal scutellar lobes with a rather prominent tubercle 
on each epipleurum. Larval type labeled "Hopk. U. S. 1201." 

The larva of this species is scarcely to be distinguished from that of 
D. valens. 

Galleries. — The egg galleries are generally longitudinal, more or less 
winding, and vary greatly in length, sometimes being very long. 
They are irregular in width, sometimes with branches, and are slightly 
grooved in the surface of the wood. The eggs are placed in masses at 
intervals along the sides and m the inner bark; the larvse excavate 
broad chambers which vary in size from a square inch to many square 
feet. The galleries of this species do not differ materially from the 
following, and have a wide range of variation in size and general 
character. 

Distribution (fig. 90). — (Hopk. U. S.) Alabama: Calhoun. Dela- 
ware. District , of Columbia: Takoma. Georgia: Cornelia, Thomas- 
ville. New Jersey: Lakewood, New Brunswick. New Yorlc: Islip 
(Long Island). South Carolina: Chicora, Lumber, New Landing, 
Pregnall. Texas: Austin, Call, Deweyville, Ktrbyville, Tarkington. 
Virginia: Glen. West Virginia: Kanawha Station. (Hopk. W. Va.) 
West Virginia: Crow, Marion County, Morgantown, Romney. Addi- 
tional localities from other collections: (Le Conte) Georgia, North 
Carolina, New Hampshire, Pennsylvania. (M. C. Z.) Texas, South 



150 



THE SCOLYTID BEETLES. 



Carolina, Maryland. (A. E. S.) New Jersey, Delaware. (A. N. S.) 
Marion County, Fla. (U.S.N.M.) Lakewood, N. J. (D. A.) Islip, 
Long Island, N. Y. (Laurent) Pennsylvania. 

Host trees. — Pinus palustris, P. rigida, P. txda, P. serotina, P. 
strobus, P. ecTiinata, Picea rubens. 

Identified specimens. — Le Conte, 9 specimens; M. C. Z., 8; Horn, 7; 
U.S.N.M., 2; H. & S.,4; D. A.,7; Hopk. U. S., about 400 specimens, 
including adults, larvas, and work. 





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Fig. 90. — Dendroctonus terebrans: Distribution map. (Original.) 



Bibliography and Synonymy. 



Scolytus terebrans Olivier, 1795, p. 6, PI. I, fig. 6, adult; original description, locality 
southern U. S. (Southern Georgia, Schwarz). 

Dendroctonus terebrans (Oliv.) Erichson, 1836, p. 53 (cotype of genus). Lacordaire, 
1866, p. 360. Zimmerman, 1868, p. 149 (in part). South Atlantic States. Le 
Conte, 1868, p. 173 (in part). Chapuis, 1869, pp. 35-36. Chapuis, 1873, pp. 
243-244, revision, Texas. Schwarz, 1878, p. 469, list, Florida. Packard, 1887, 
p. 177 (in part), Le Conte quoted. Packard, 1890, p. 721 (in part). Dietz, 
1890, p. 29 (in part). Hopkins, 1893b, p. 143, No. 76 (in part), Hampshire and 
Monongalia counties, W. Va. Hopkins, 1893c, p. 213, No. 300 (in part), list, 
host, etc. Smith, 1899, p. 364 (in part), distribution, habits. Chittenden, 
1899, p. 56 (in part). Smith, 1901, p. 92, destructive to pine, Lahaway [Borden- 
town], N. J. Ulke, 1902, pp. 36-56, list, habits, etc. Hopkins, 1902b, p. 10, 
footnote, mention. Hopkins, 1906c, p. 81, distinguishing characters. Felt, 
1906, pp. 342-345, Long Island. 

Dendroctonus terebrans (Lacordaire) Le Conte, 1876, p. 385 (= Oliv. in part), bibli- 
ography, distribution, systematic note, on specimens from Georgia. 

Dendroctonus rufipennis (not of Kirby) Dietz, 1890, p. 30, Florida. 

? Dendroctonus sp. Blandford, 1897, p. 147, reference to black form from Texas. 



THE GENUS DENDEOCTONUS. 151 

23. Dendroctonus valens Le Conte. 
(PL VII, fig. 23.) 

Adult. — Typical female: Length, 8.7 mm. Head with front broad, 
convex, and with broad anterior impression. Epistomal process 
broad, with lateral sections oblique. Pronotum with punctures 
moderately coarse, much smaller and denser toward base. Elytra 
with a few long hairs toward base; declivity convex; striae impressed, 
with small distinct punctures; interspaces moderately convex, dis- 
tinctly rugose. 

Typical specimen labeled, name label, "Hopk. 1/22/08, Pinus 
ponderosa, Hopkins, collector, McCloud, Cal., 9 , Hopk. U. S. 18a." 

Typical male: Length 7.6 mm. Differs from female in stouter 
mandibles, narrower antennal club, more opaque declivity, less dis- 
tinctly impressed striae, and more obscure punctures. 

Typical male labeled same as female. 

Variations. — The length ranges from 5.7 to 9 mm., with the aver- 
age about 8 mm. The color of the elytra, pronotum, and vertex 
of head ranges from light to dark red, but is never black, while the 
ventral part of the body varies from light red to black. The great- 
est variation is in size, but there is a wide and remarkable range 
in the epistoma and front and in the sculpture of the pronotum 
and elytra, as in almost every other character. Thus a large num- 
ber of individuals may be easily selected, each of which might be 
considered as representing a good species, but when a large series 
of specimens is examined from any given faunal region or locality 
no sufficiently distinctive and constant characters have been found 
by which they can be readily recognized as distinct from those of 
any other faunal region, so that those examples from Maine can 
not be distinguished from examples collected in the mountains of 
Mexico. 

Distinctive characters. — The characters which serve to distinguish 
this species from D. terebrans, to which it is closely allied, are its 
prevailing red color, the less uniform and more densely placed 
pronotal punctures, the much broader epistomal process, with the 
lateral sections less angular and more oblique, and the more evident 
and numerous hairs on the anterior dorsal area of the elytra. 

Revisional notes. — See same heading under D. terebrans . 

The typical specimen on which Le Conte' s original description 
(Le Conte, 1860) was based is from California and represents the 
common red form which has heretofore been confused with D. tere- 
brans. The distinguishing character, ''finer and denser punctures of 
the thorax," mentioned by Le Conte is undoubtedly a good specific 
character when taken with color and other characters. The species 
is represented in the Le Conte collection by the type and 9 speci- 



152 



THE SCOLYTID BEETLES. 



mens, and in the collection of the Museum of Comparative Zoology 
by over 50 specimens. 

Pwpa. — In addition to the generic, divisional, and subdivisional 
characters, the front and middle femora are armed each with a 
minute apical spine; abdominal tergites 1 to 6 with moderately 
small pleural spines; 1 is without dorsal spines, but with distinct 
lateral ones; 2 to 6 have small dorsal and lateral spines, the former 
increasing in size to 6, and all have dark tips; 7 and 8 are unarmed; 
9 has the usual stout pleural spine. Pupal type labeled "Hopk. 
U. S. 2824." 




Fig. 91. — Dcndroctonus valcns: Egg galleries and larval chamber. A , Incomplete egg galleries with boring 
dust removed; B, normal gallery; C, advanced stage of work; a, entrance burrow; 6, basal section; 
c, ventilating burrow; d, egg nest with eggs; e, boring dust; /, subsequent or inner galleries; g, larvae 
at work; li, pupal cell in boring dust mixed with resin. (Original.) 

The usual variation prevails in the number and arrangement of 
minor spines and between the young and older examples. 

The character which in general serves to distinguish the pupa of 
this species from the preceding is found in the darker tips to the 
body spines. 

Larva (text fig. 39 and PI. VIII, fig. 23). — In addition to the ge- 
neric, divisional, and subdivisional characters, front of head with pos- 
terior angle, median area not elevated but transversely rugose, except 
near apex, where it is smooth; epistoma flat, opaque, smooth, mth 
straight anterior margin ; clypeus broad, prominent, convex, with faint 
median longitudinal line, sides rounded, apex broadly emarginate; 



THE GENUS DENDEOCTONUS. 



153 



labium short, sides nearly parallel, and apex tuberculate. Protho- 
racic tergum with two broad, shining dorsal plates separated by a 
rather broad median space, and a smaller lateral plate each side; 
sternellar lobes each with a faint foot callus; the mesoterga and 
metaterga with shining plates on the lateral lobes. Abdomen with a 
rather prominent tubercle on each epipleurum. Larval type labeled 
"Hopk. U.S. 2824." 




Fig. 92.—Dendroctonus valens: Work in bark at base of stump, a, Entrance and pitch tube; 6, egg gallery; 
c, boring dust and resin; d, pupal cell; e, pupa; /, larvae at work feeding on inner living bark; g, exit 
burrows; h, resulting old scar or basal wound, often referred to as basal fire wound; i, inner bark with 
outer corky bark removed. ( Original . ) 

The larva of tliis species is scarcely to be distinguished from the 
preceding. 

Galleries (figs. 91-93). — The egg galleries are generally longitudinal, 
more or less winding, and vary greatly in length, sometimes being 
very long; they are irregular in width and sometimes with branches, 
and are shghtly grooved into the surface of the wood. The eggs 
are placed in masses at intervals along the sides in the inner bark, 
and the larvae excavate broad chambers, which vary in size from a 



154 



THE SCOLYTID BEETLES. 



square inch to many square feet. The galleries of this species do 
not differ materially from the preceding, and have a wide range of 
variation in size and general character. 

Distribution (fig. 94). — (Hopk. U. S.) Arizona: Chiricahua Moun- 
tains, Flagstaff, Fredonia, Grand Canyon, Paradise, Ramsey Canyon, 
Rincon Mountains, Santa Catalina National Forest, Williams. Califor- 
nia: Alder Creek, Berkeley, Chester, Del Monte, La Moine, Little Yosem- 
ite, Madera County, Merced, McCloud, Nevada City, Pacific Grove, 
Palo Alto, Pinogrande, San Bernardino, Sterling, Summerdale, 
Wawona, Yosemite. Canada. Colorado: Fort Garland, Manitou 
Park, Palmer Lake, San Isabel National Forest. Guatemala: Cabon. 




Fig. 93. — Dendroctonus valens: Basal wound in living tree resulting from primary injury by this species. 
Often mistaken for fire wound. (Original.) 

Idaho: Cedar Mountain, Centerville, Grimes Pass, Harris Ridge 
(Kooskai), Pioneerville, Priest River, Smiths Ferry. Kansas. 
Maine: Brunswick, Casco Bay, Limerick, Peak Island, Portland. 
Massachusetts: Lynn, Wyoming. Mexico: Chalco, Chihuahua, Mex- 
ico City, Michoacan, Ponada, Satazin. Michigan: Grand Island. 
Montana: Missoula. Nevada. New Hampshire: Durham, Webster. 
New Mexico: Capitan, Capitan Mountains, Cloudcroft, Lincoln 
National Forest, Vermejo, ^Yliite Mountains. New YorJc: Ithaca. 
North Carolina: Biltmore, Pink Beds. Oregon: Albany, Hood River, 
Joseph, Slate Creek. Pennsylvania: Milford, Philadelphia. South 



THE GENUS DENDEOCTONUS. 155 

Dakota: Black Hills, Elmore, Lead, Spearfish. Utali: Escalante, 
Kamas, Kanab, Panguitch Lake. Vermont: Fairlee. Virginia. 
Washington: Buckeye, Easton, Pullman, Skykomish. West Vir- 
ginia: Crow, Hampsliire County, Hardy County, Kanawha Station, 



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Fig. 94. — Dendroctonus valens: Distribution map. (Original.) 

Monongalia County, Morgantown, Pendleton, Raleigh County, 
Romney, Roosevelt, Tucker. Wisconsin: Ashland, Madison. (Hopk. 
W. Va.) West Virginia: Bretz, Cranesville, Deckers Creek, Dellslow, 
Mayfield Hill, Moorefield, Morgantown, Pendleton, Randolph, 



156 THE SCOLYTID BEETLES. 

Romney. Additional localities from other collections: (M. C. Z.) New 
Mexico; California; Pennsylvania; Idaho; Maine; Massachusetts; 
Cambridge, Mass. (Le Conte) Middle States; Connecticut; Garland, 
Colo.; California. (Horn) North Carolina, Oregon, Wisconsin, Maine, 
California, New Mexico, and Pennsylvania. (A. E. S.) Kansas, Idaho, 
Nevada, Colorado, Canada, Virginia, Illinois, New Jersey. (W. & F.) 
Durham, N. H. (U.S.N.M., H. & S.) Marquette, Mich.; Cambridge, 
Mass.; Helena, Mont.; Garland, Colo.; Coeur d'Alene, Idaho; Hood 
River, Oregon; Chiricahua Mountains, Arizona; Sisson,Cal.; Coldridge, 
N. Mex. (U.S.N.M., B. & S.) Las Vegas, N. Mex.; Bright Angel, 
Prescott, Flagstaff, and Williams, Ariz. (Gillette) Colorado; Bailey, 
Colo. (U.S.N.M.) Easton, Wash. ; Placer, Colo. ; Shasta County, Cal. ; 
Powder River, Colorado; Lake Superior; Skokomish River, Wash- 
ington; New Mexico; Siskiyou County, Cal.; Ozumba Mountain, 
Mexico. (Weed & Fiske) Durham, N. H. (Webb) Pullman, Wash. 
(Dietz) Pennsylvania, California, Arizona, Washington [State], 
(D. A.) Dunsmuir, Cal.; Custer County, Cal. (Chittenden) Duluth, 
Minn.; Ithaca, N. Y.; Grangeville, Idaho; Cahfornia. (Baldwin) 
Ventura County, Cal. (Soltau) Colorado. 

Host trees. — Pinus ponderosa, P. strohus, P. radiata, P. rigida, 
P. lambertiana, P. murrayana, P. strohiformis, P. cJiihuahuana, P. 
edulis, P. jeffreyi, P. sylvestris, P. virginiana, P. arizonica, P. sp.; 
Picea canadensis, P. excelsa, P. rubens; Ahies concolor; Larix laricina. 

Identified specimens. — Le Conte, 10 specimens; M. C. Z., 55; Horn, 
14; A. E. S., 8; U.S.N.M., 12; H. & S., 20; Dietz, 5; D. A., 3 speci- 
mens. This species is also represented in the forest insect collection 
of this Bureau by about 5,000 specimens, including all stages and 
work. 

Bibliography and Synonymy. 

Scolytus terebrans (not of Oliv.) Harris, 1826, pp. 169, 170, character and habits. Har- 
ris, 1862, p. 86, footnote, name only. Zimmerman, 1868, p. 149, as synonym for 
D. terebrans (in part). 

Eylurgus terebrans (not of Oliv.) Harris, 1841, p. 72, brief description of larva? and 
habits. Harris, 1842, pp. 72, 73, repeated. Harris, 1852, p. 76, repeated. Fitch, 
1858, pp. 728-729, description of adult and larva, habits? Harris, 1862, p. 86, 
repeated, fig. 42, adult. Harris, 1863, pp. 84-86, fig. 42, adult, account repeated. 
Thomas, 1876, p. 146, brief description of adult and habits of larva. Smith, 1877, 
p. 52, work in pine. 

Dendroctonus valens Le Conte, 1860, p. 59, original description. Le Conte, 1868, p. 
173, mentioned as synonym of D. terebrans Lacordaire. Chapuis, 1869, p. 35; 
Chapuis, 1873, p. 243. Le Conte, 1878, p. 472, list, Atlanta, Idaho. Hopkins, 
1903a, p. 61, reference to habits, etc. Hopkins, 1904, p. 19, PI. VII, figs, a, b, g 
(reprint). Powell, 1904, anatomy. Powell, 1905, ibid. Hopkins, 1905, pp. 6, 
11, 17, distinctive characters. Hopkins, 1906b, p. 147, PI. IV, figs. 6, 8, anatomy 
of larval head. Hopkins, 1906c, p. 81, mentioned as a good species, host, 
localities, etc. Fall, 1907, p. 218, list, localities. 



THE GENUS DENDROCTONUS. 157 

Dendroctonus terebrans (not of Oliv.) Zimmerman, 1868, p. 149 (in part), revision. Le 
Conte, 1868, p. 173 (in part), synopsis, bibliography. Le Baron, 1871 (in part), 
economic reference. Le Conte, 1876, p. 385 (in part), revision, synopsis; bibli- 
ography, locality. Packard, 1887, pp. 175, 243 (in part). Packard, 1890, p. 721, 
fig. 250 (in part), adult, quotes Harris and Le Conte. Dietz, 1890, p. 29 (in 
part), Eastern States, California, Washington, fig. 1, antenna and epistoma; var. 
b, Washington, California, Colorado, Arizona, and Idaho; var. c. New Mexico, 
Arizona, Nevada, California; var. d, Pennsylvania. Hopkins, 1892a, pp. 64-65 
(in part), habits, etc. Hopkins, 1893b, p. 143, No. 76 (in part), and index, 
habits, distribution, host, enemies, etc., in West Virginia, all except part of two 
references from Hampshire and Monongalia counties. Hamilton, 1895, pp. 346, 
378, list, etc. Wickham, 1896a, p. 169, listed. Lake Superior. Wickham, 1906b, 
p. 170, list, Coolidge, N.Mex.; Walnut and Williams, Ariz. Hopkins, 1897a, p. 
41, habits, etc. Blandford, 1897, pp. 146-147, synopsis, redescribed, distribu- 
tion in Mexico, bibliography. Wickham, 1898, p. 312, list, Arizona. Hopkins, 
1899a, pp. 392-393, 415^21, and index (in greater part), different stages, galleries, 
etc., described and illustrated (except egg and pupa), with full account of habits, 
and distribution in West Virginia, etc. Hopkins, 1899b, pp. 14, 15, habits, hosts, 
etc. Hopkins, 1899c, p. 343 (in part), stridulation, etc. Chittenden, 1899, p. 
56 (in part). Wickham, 1902, p. 309, list, locality. Felt, 1903, pp. 480-481, 
figs. 1-3 (in part), adult, larva, pupa. New York State except Long Island (in 
part?), habits, hosts, enemies, etc. Felt, 1906, pp. 342-345 and index, fig. 64, 
adult; 65, pupa; 66, larva (in part), habits, host, etc. 

Dendroctonus terebrans (not of Lacordaire) Le Conte, 1868, p. 173. 

Dendroctonus obesus (not of Mann.) Packard, 1877, p. 803, description and probable 
habits in Colorado, compared with D. terebrans (not of Oliv.). 

f Dendroctonus rufipennis (not of Kirby) Packard, 1887, p. 176, in pitch pine. 

fDendroctonus similis (not of Lec.)Slosson, 1902, p. 319, list, locality. 

Dendroctonus valens Lee. var. occidentalis Hopkins, 1902b, p. 12, manuscript name for 
variety. 

Dendroctonus (terebrans) var. valens (Lee.) Wickham, 1902, p. 309, Garland, Colo. 

24. Dendroctonus adjunctus Blandford. 
ORIGINAL DESCRIPTION. 

''Long. 4.6-5.3 millim. 

"Oblong, less elongate than D. parallelocollis, slightly depressed, 
piceous-black, with apex of elytra lighter. Median segment of 
epistoma shorter than the lateral segments, its sides very oblique and 
elevated, its apical border concave ; front covered with close granules 
and punctures, with an interrupted median impressed line deepest at 
its junction with a slight transverse subocular impression; vertex 
finely punctured; antennae piceous, the scape with rounded club, 
second joint of funiculus" scarcely longer than first, club transverse 
oval, its sutures curved toward apex. Prothorax more transverse 
than in D. parallelocollis, constricted in front, the apical emargination 
and basal bisinuation distinct; surface impressed behind apex and 
somewhat flattened, shining, the punctures sparse and not strong, the 
median line obsolete. Elytra feebly striato-punctate, shining, except 



158 



THE SCOLYTID BEETLES. 



at the closely granulate base, with somewhat scattered granules, 
obsolete on the declivity, interstices with rows of distinct long dark 
hairs from middle to apex, the first with a single series of stronger 
granules. Legs piceous. 




Fig. 95. — Bendroctonus adjunctus: Distribution map. (Original.) 

^^Hah. [See fig. 95.] Guatemala, Totonicapam {Cliampion). 
''A distinct species, resembling a MyelopJiilus rather than a Ben- 
droctonus. But two specimens were taken." 

Bibliography. 
Dendroctonus adjunctus Blandford, 1897, p. 147. 



BIBLIOGRAPHY. 

1793. Herbst, J. F. W. — Natursystem aller bekannten in- und auslandischen Insek- 

ten-Kafer, fiinfter Theil, p. 107, Taf. 48, fig. 5. 

1794. KuGELANN.— Schneid., Mag., 5, p. 523-12. 

1795. Olivier, A. G. — Entomologie, ou histoire naturelle des insectes, Vol. IV, Genus 

78, p. 6, No. 6, PI. I, fig. 6, a, h. ■ 
1813. Gyllenhal, L. — Insecta Suecica, Tom. I, Pars III, pp. 335-336. 
1820. Chabrier, J. — Essai sur le a'oI des insectes. <Mem. du Mus. d'Hist. Nat., 

VI, pp. 410-472, pis. 18-21. 
1824. AuDouiN, Victor. — Recherches anatomiques sur le thorax des animaux 

articules, et celui des insectes hexapodes en particulier.<Ann. Sci. Nat., I, 

pp. 97-135, 416-432. 
1826. Harris, T. W.— Trees. < New England Farmer, V, December 22, pp. 169-171. 
1830. MacLeay, W. S. — Explanation of the comparative anatomy of the thorax of 

winged insects, with a review of the present state of the nomenclature of its 

parts. <Zool. Journal, V, pp. 145-179, Pis. V, VI. 

1836. Erichson, W. F. — Systematische Auseinandersetzung der Familie der Bor- 

kenkafer (Bostrichid8e).<Archiv fiir Naturgeschichte v. A. F. Wiegmann, 
z welter Jahrgang, erster Band, pp. 45-65. 

1837. KiRBY, Wm. — Fauna Boreali Americana; or the zoology of the northern parts 

of British America. Part IV, Insecta, p. 195. 
1839. Ratzeburg, J. T. C— Die Forst-Insekten, Vol. I, p. 217. 
1839. Newport, George. — Insecta. Cyclopaedia of anatomy and physiology (R. 

B. Todd), II, pp. 853-994, figs. 329-439. 

1841. Harris, T. W. — A report on the insects of Massachusetts injurious to vegeta- 

tion, p. 72. 

1842. Harris, T. W. — A treatise on some of the insects of New England which are 

injurious to vegetation, pp. 72-73. 

1843. Mannerheim, G. G. — Beitrag zur Kaefer-Fauna der Aleutischen Ineeln, Insel 

Sitka und Neu-Californiens.<Bull. Soc. Imp. Nat. Mosc, Tom. XVI, II, 

pp. 175-314. Moscou. 
1849. Bach, M.— Kafer,. II, p. 144. 
1852. Stein, Friedrich. — Beitrage zur Insektenkunde.<Tharand Jahrbuch, T. 8, 

p. 235. 
1852. "Mannerheim, G. G. — Zweiter Nachtrag zur Kaefer-Fauna der nordamerika- 

nischen Laender des Russischen Reiches.<Bull. Soc. Imp. Nat. Mosc, Tom. 

XXV, II, pp. 28.3-387. Moscou. 

1852. Harris, T. W. — A treatise on some of the insects of New England which are 

injurious to vegetation, p. 76. 

1853. Mannerheim, G. G. — Dritter Nachtrag zur Kaefer-Fauna der nordamerika- 

nischen Laender des Russischen Reiches.<Bull. Soc. Imp. Nat. Mosc, Tom. 

XXVI, II, pp. 95-273. Moscou. 

1854. Stein, F. — Jahrbuch der Akademie fiir Forst- und Landwirthe zu Tharand, 

III, pp. 277-279. 
1858. KoLLAR, ViNCENZ. — Beitrage zur Naturgeschichte des grossen Fichten Bast- 
kafers, Hylesinus (Dendroctonus) micans Kug., aus den Beobachtungen des 
Herrn Conr. Leinweber, k. k. Hofgartner, zusammengestellt von Vincenz 
KoUar, pp. 23-28. 

159 



160 THE SCOLYTID BEETLES. 

1858. Fitch, Asa. — Fourth report on the noxious, beneficial and other insects of the 
State of New York.<Trans. N. Y. St. Agr. Soc, XVIII, pp. 728-729. 

1860. Le Conte, J. L. — Reports of explorations and surveys * * * from the 
Mississippi River to the Pacific Ocean, p. 59; 

1862. Harris, T. W. — A treatise on some of the insects injurious to vegetation, p. 86. 

1862. DoBNER. — Einige Bemerkungen iiber schadliche Forstinsekten . <Allgemeine 

Forst- und Jagd-Zeitung, neue Folge, 38 Jahrgang, pp. 275-277. 

1863. Harris, T. W. — A treatise on some of the insects injurious to vegetation, 

Flint edition, pp. 84-86. 

1864. EiCHHOFF, W. J. — tjber die Mundtheile und die Fiihlerbildung der euro- 

paischen Xylophagi sens, strict. <Berliner entomologische Zeitschrift 

pp. 17-46. 
1866. Lacordaire, T. — Genera des Coleopteres, VII, p. 360. 
1868. EiCHHOFF, W. J. — Berliner entomologische Zeitschrift, Heft I und II (pp. 

i-viii, 1-176, 313-368. Published May, 1868). Heft III und IV (pp. ix-xiv, 

177-312, 369-432. Published March, 1869). Note especially p. xi. footnote. 

Reference to scolytids, pp. 145-152, 273-282. 
1868. Le Conte, J. L. — Synopsis of the Scolytidse of America north of Mexico. 

Appendix. <Trans. Amer. Ent. Soc, XV, pp. 150-178. 

1868. Zimmerman, C. — Synopsis of the Scolytidse of America north of Mexico. 

<Trans. Amer. Ent. Soc, XV, pp. 141-150, September. 

1869. Chapuis, F. — Synopsis des Scolytides.<Mem. Soc. Sci. Liege, author's extra 

issued 1869, pp. 1-61. (See also Chapuis, 1873.) 
1871. Le Baron, W. — Means against larvf« in timber; answers to correspondents. 

<Prairie Farmer, vol. 42. 
1873. Chapuis, F. — Synopsis des Scolytides.<Mem. Soc. Sci. Liege, III, pp. 

213-269. 

1875. Lindemann, C. — ^Vergleichend-anatomische Untersuchung iiber das mann- 

liche Begattungsglied der Borkenkafer.^Bull. Soc. Imp. Nat. Moscou (Bull. 
Mosc), XLIX, pp. 196-252, Pis. I-V. 

1876. Le Conte, J. L. — The Rhynchophora of America north of Mexico. <Proc 

Amer. Phil. Soc, pp. 384-386, December, 1876. 
1876. Peck, C. H.— The black spruce. <Trans. Albany Inst., VIII, pp. 283-301. 

1876. Thomas, Cyrus. — Sixth report of the State Entomologist on the noxious and 

beneficial insects of the State of Illinois, p. 146. 

1877. Smith, E. A. — Shade trees, indigenous shrubs and vines, by J. T. Stewart, 

M. D., and insects that infest them, by Miss Emma A. Smith. Peoria, 111., 
1877, p. 52. 

1877. Packard, A. S. — Report on the Rocky Mountain locust and other insects now 

injuring or likely to injure field and garden crops in the "Western States and 
Territories. <9th Ann. Rep. U. S. Geol. and Geog. Surv. Terr., pp. 589-810. 

1878. Schwarz, E. A. — The Coleoptera of Florida. <Proc. Amer. Phil. Soc, XVII, 

p. 469. 

1878. Le Conte, J. L. — The Coleoptera of the Alpine Rocky Mountain Regions, 

Part I.<Bul. U. S. Geol. and Geog. Surv., IV, pp. 469, 472, November 30. 

1879. EiCHHOFF, W. — Ratio, descriptio emendatio eorum Tomicinorum. Extrait 

des Memoires de la Societe Royale des Sciences de Liege, 2^ serie, VIII. 
1879. Peck, C. H.— Report of the Botanist. <28th Ann. Rep. N. Y. State Mus. Nat. 

Hist., pp. 32-38. 
1881. Altum, B.— Forstzoologie. III. Insekten, I. Abtheil., pp. 262-266. 

1881. EiCHHOFF, W. J. — Die europaischen Borkenkafer. Berlin, pp. 37, 125-128. 

1882. Hough, F. B. — Report on Forestry, submitted to Congress by the Commissioner 

of Agriculture, Part VIII, Insect Ravages, pp. 259-274. 1. The injmies done 
to spruce and other coniferous timber by insects. 



THE GENUS DENDROCTONUS. 161 

1883. Le Conte and Horn. — Classification of the Coleoptera of North America, p. 523. 

1884. Harrington, W. Hague. — Canadian Entomologist, XIV, p. 218. 

1885. LiNTNER, J. A. — Second report on the injm-ious and other insects of the State 

of New York, p. 54. 
1885. Amans. — Comparaisons des organes du vol dans la serie animale.<Ann. Sci! 

Nat., Ser. 6, Zool., XIX, pp. 9-222, Pis. I-VIII. 
1887. Fletcher, J. — ^The spruce barkbeetle (D. rufipennis). Appendix to the report 

of the Minister of Agriculture. Report of the Entomologist and Botanist, 

pp. 39, 40. 

1887. Packard, A. S. — Insects injurious to forest and shade trees. <Bul. 7, U. S. Ent. 

Comm., pp. 175, 176, 177, 243. 

1888. ScHWARZ, E. A. — Proc. Ent. Soc. Wash., I, p. 175, November 1. 

1889. Judeich, J. T., und Nitsche, H. — Lehrbuch der mitteleuropaischen Forst- 

insektenkunde, pp. 458-462. 

1889. KoLBE, H. J. — Einfiihrung in die Kenntnis der Insekten. 

1890. Packard, Alpheus S. — Insects injurious to forest and shade trees. Revision 

and enlarged edition of Bui. 7.<5th Rep. IT. S. Ent. Comm., U. S. Dept. Agr., 
pp. 721, 722, etc. 
-1890. Harrington, W. Hague. — On the lists of Coleoptera published by the Geolog- 
ical Survey of Canada, 1842-1888. <Can. Ent., XXII, No. 9, p. 189 (author's 
extra, p. 19). 

1890. Dietz, W. G. — Notes on the species of Dendroctonus of Boreal America. <Trans. 

Amer. Ent. Soc, XVII, pp. 27-32. 

1891. Harrington, W. Hague. — General notes. <Can. Ent., XXIII, No. 2, p. 27. 

Ibid., Science, XX, pp. 256-257, November 4, 1892. , 

1892. Pauly, a. — Uber die Brutpflege und jahrliche Geschlechterzahl des Riesen- 

bastkafers, Hylesinus micans Ratz.<^Forstlich-naturwissenschaftliche Zeit- 

schrift, I. Jahrgang, pp. 315-327. 
1892a. Hopkins, A. D. — Notes on a destructive forest tree scolytid.<Science, XX, 

pp. 64-65, July 29. . 
1892b. Hopkins, A. D.— Proc. Ent. Soc. Wash., II, p. 353, October 6. 

1892. ScHAUPUSS, C. F.— Barkbeetle destroyer.<Can. Ent., XXIV, No. 12, p. 316. 

1893. CocKERELL, T. D. A. — The entomology of the Mid-Alpine Zone of Custer 

County, Colorado. <Trans. Amer. Ent. Soc, XX, p. 336. 
1893a. Hopkins, A. D. — Damage to forests by the destructive pine barkbeetle. 

<Insect Life, V, No. 3, pp. 187-189, January. 
1893b. Hopkins, A. D. — Catalogue of West Virginia Scolytidse and their enemies. 

<Bul. 31, W. Va. Agr. Exp. Sta., No. 76 [=p. 143], April. 
1893c. Hopkins, A. D. — Catalogue of the West Virginia forest and shade tree insects. 

<Bul. 32, ^Y. Va. Agr. Exp. Sta., May. 
1893d. Hopkins, A. D. — Destructive scolytids and their imported enemy. <;insect 

Life, VI, No. 2, pp. 123-129, December. 

1894. LiNTNER, J. A. — Gardening, II, p. 292, May 15. 

1894. Hamilton, John. — Catalogue of the Coleoptera of Alaska, with the synonymy 

and distribution. <Trans. Amer. Ent. Soc, XXI, p. 35. 
1894a. Hopkins, A. D. — Destructive scolytids and their imported enemy. <;24th Ann. 

Rep. Ent. Soc. Ontario, pp. 71-76. 
1894b. Hopkins, A. D. — Sexual characters in Scolytidse. <Can. Ent., XXVI, pp. 274- 

280. 
1894c. Hopkins, A. D.— The relations of insects and birds to present forest conditions. 

<;Garden and Forest, VII, p. 348. 

1895. Hamilton, John. — Catalogue of the Coleoptera of southwestern Pennsylvania, 

with notes and descriptions. <Trans. Amer. Ent. Soc, XXII, pp. 346, 378, 
October. 
79980—09 12 



162 THE SCOLYTID BEETLES. 

1896a. WiCKHAM, H. F. — A list of the Coleoptera from the southern shore of Lake 

Superior. <Proc. Davenport Acad. Nat. Sci., VI, p. 169. 
1896b. WiCKHAM, H. F. — A list of some Coleoptera from the northern portion of New 

Mexico and Arizona. <Bul. Lab. Nat. Hist. St?te L'niv. Iowa, Vol. Ill, No. 

4, p. 170. 
1896. Hopkins, A. D. — Some notes on insect enemies of trees. <Can. Ent., 

XXVIII, No. 10, pp. 246, 250, October. 

1896. Verhoeff, Carl. — tjber das Abdomen der Scoiytiden.<Arch. f. Naturg., 

Jahrgang 62, I Bd., pp. 109-144, Pis. VII, VIII." 

1897. Menegaux, A., et Cochon, J. — Sur la biologic de I'Hylesine brillante.<C. R. 

Acad. Sc. Paris, Tome 124, No. 4, pp. 206-209. (Abstr. Journ. R. Micr. 

Soc. London, pt. 2, p. 120.) 
1897a. Hopkins, A. D. — Report of Entomologist. <6th Ann. Rep. W. Va. Agr. Exp. 

Sta. for fiscal year ending June 30, 1893, pp. 29-42. 
1897b. Hopkins, A. D. — Report of Entomologist. <7th Ann. Rep. W. Va. Agr. Exp. 

Sta. for fiscal year ending June 30, 1894. 
1897c. Hopkins, A. D. — Report of the Entomological Department. <9th Ann. Rep. 

' W. Va. Agr. Exp. Sta. for fiscal year ending June 30, 1896, pp. 79, 94-95, 147, 

151, PI. I. 
1897. Blandford, Walter F. H. — Fam. Scolyticl8e.<Biol. Centr.-Amer., Coleopt., 

IV, pt. 6, August, 1897, pp. 146, 147. 

1897. Chittenden, F. H. — Insect injury to chestnut and pine trees in Virginia and 

neighboring States. Some Miscellaneous Results of the Work of the Division 
of Entomology. <Bul. 7, n. s., Div. Ent., U. S. Dept. Agr., pp. 67-75, fig. 43. 

1898. Harvey, F. L. — Notes on insects of the year.<13th Ann. Rep. Maine Agr. 

Exp. Sta., Orono, Maine, 1897. Part II of the Annual Report of the Uni- 
versity of Maine, p. 176. Note -on depredations on spruce in Maine. 

1898. Howard, L. 0.,and Chittenden, F. H. — Notes from correspondence. Some 
Miscellaneous Results of the Work of the Division of Entomology. <Bul. 10, 
n. s., Div. Ent., U. S. Dept. Agr., p. 98. 

1898a. Hopkins, A. D. — Proceedings of the Tenth Annual Meeting of the Association 
of Economic Entomologists. <Bul. 17, n. s., Div. Ent., IT. S. Dept. Agr., p. 69. 

1898b. Hopkins, A. D. — Insects detrimental and destructive to timber and timber 
products. ^Proceedings of the Nineteenth Annual Meeting of the Society for 
the Promotion of Agricultural Science, held at Boston, Mass., 1898, pp. 
104, 105. 

1898. ScHWARZ, E. A.— Proc. Ent. Soc. Wash., IV, No. 2, pp. 81, 82, March 21. 

1898. Weed, Clarence M., and Fiske, W. F. — Notes on spruce bark-beetles. Pro- 
ceedings of the Tenth Annual Meeting of the Association of Economic Ento- 
mologists. <Bul. 17, n. s., Div. Ent., U. S. Dept. Agr., pp. 67-69. 

1898. Chittenden, F. H. — Recent injury by bark-beetles. A correction. Some 
Miscellaneous Results of the Work of the Division of Entomology, III.<Bul. 
18, n. s., Div. Ent., U. S. Dept. Agr., p. 96. 

1898. WiCKHAM, H. F. — The beetles of southern Arizona. <Bul. Lab. Nat. Hist. 
State Univ. Iowa, Vol. IV, No. 3, p. 312. 

1898. Johnson, Chas. W. — Report on insects injurious to spruce and other trees. 
Chap. V.<3d Ann. Rep. Pa. Dept. Agr., Part II, 1897, issued in 1898 after 
September, pp. 69-110. 

1898. LovENDAL, E. A.— De Danske barkbiller (Scolytidse et Platypodidse Danicte), 
pp. 86-91. 

1899a. Hopkins, A. D. — Report on investigations to determine the cause of unhealthy 
conditions of the spruce and pine from 1880-1893. <Bul. 56, W. Va. Agr. Exp. 
Sta., April. 



THE GENUS DENDEOCTONUS. 163 

1899b. Hopkins, A. D. — Preliminary report on the insect enemies' of forests in the 

northwest. <Bul. 21, n. s., Div. Ent., U. S. Dept. Agr., pp. 10, 11, 13, 14, 

15, 20, 21, 22, 26. 
1899c. Hopkins, A. D. — Notes on Dendroctonus .<^Vroc . Ent. Soc. Wash., IV, No. 3, 

p. 343, May 24. 
1899. Smith, J. B. — Insects of New Jersey. <Supplement to the 27th Annual Report 

of the State Board of Agriculture, p. 364. 

1899. Chittenden, F. H. — Insect enemies of the white pine. The white pine {Pinus 

strobus Linnaeus). <Bul. 22, Div. For., U. S. Dept. Agr., Y)p. 55, 56, figs. 5. 
1900a. ScHWARZ, E. A. — Papers from the Harriman Alaska Expedition, XVIII. 

Entomological results (12); Coleoptera.<Proc. Wash. Acad. Sci., II, p. 537, 

December 20. 
1900b. ScHWAEZ, E. A. — Coleoptera of the expedition. <Ibid., Author's extra, p. 185. 

1900. Gary, Austin. — Insect damage to spruce timber in Maine and New Hamp- 

shire. <The Forester, VI, No. 3, pp. 52-54. 

1901. Smith, J. B.— Doings of societies. <Ent. News, XII, No. 3, p. 92, March. 

1901. Johnson. — Doings of societies. <Ent. News, XII, No. 8, p. 92, March. 
1901a. Hopkins, A. D. — Insect enemies of the spruce in the northeast. <Bul. 28, n. s., 

Div. Ent., U. S. Dept. Agr., pp. 1-48, Pis. I-V, XIV, XV (issued October 1, 
1901). 
1901b. Hopkins, A. D. — Some insect enemies of living trees. -< Proceedings of the 
Twenty-Second Annual Meeting of the Society for the Promotion of Agricul- 
tural Science held at Denver, Colo., pp. 66-69. 

1902. Slosson, a. T. — Additional list of insects taken in the Alpine Region of Mount 

Washington. <Ent. News, pp. 319-321, December, 1902. 
1902. Comstock, J. H., and Kellogg, V. L. — Elements of insect anatomy. Fourth 

edition. 
1902. Severin, G. — Le Dendroctonus micans (Kugelann) en Belgique.<Bull. Soc. 

Centr. forestiere Belg., Vol. IX, pp. 72-81. 
1902. Severin, G. — L'invasion de I'Hylesine geante.<Bull. Soc. Centr. forestiere 

Belg., Vol. IX, pp. 145-152. 
1902. Ulke, Henry. — A list of the beetles of the District of Columbia. <Proc. U. S. 

Nat. Mus., XXV, pp. 1-57. 
1902a. Hopkins, A. D. — Some notes on the genus Dendroctonus .-^Froc . Ent. Soc. 

Wash., V, No. 1, pp. 3-4, May 17, 1902 (author's extra published March 28, 

1902). 
1902b. Hopkins, A. D. — Insect enemies of the pine in the Black Hills Forest Reserve. 

<Bul. 32, n. s., Div. Ent., U. S.'Oept. Agr., pp. 1-24, Pis. I-VII, figs. 1-5 

(issued April 29, 1902). 
1902c. Hopkins, A. D. — On the study of forest entomology in America. Proceedings 

of the Fourteenth Annual Meeting of the Association of Economic Entomolo- 

gists.<Bul. 37, n. s., Div. Ent., U. S. Dept. Agr., pp. 5-32. 
1902. ScHWARZ, E. A.— Proc. Ent. Soc. Wash., V, No. 1, p. 32, May 17. 
1902. Weber, L. — Zur Biologie von Rhizophagus grandis Gyllh.<Allg. Zeitsch. f. 

Ent., VII, p. 108. 

1902. WiCKHAM, H. F. — A catalogue of the Coleoptera of Colorado. <Bul. Lab. Nat. 

Hist. State Univ. Iowa, Vol. V, No. 3, pp. 217-310. 
1903a. Hopkins, A. D. — Forest insect explorations in the summer of 1902. <Can. 

Ent., XXXV, No. 3, pp. 59-61, March. 
1903b. Hopkins, A. D. — Some of the principal insect enemies of coniferous forests in 

the United States. < Yearbook U. S. Dept. Agr., for 1902, pp. 265-282. 

1903. Felt, E. P. — Insects affecting forest trees. <7th Rep. Forest, Fish and Game 

Comm. State N. Y., pp. 480-481, 2 plates, figs. 1-3. 



164 THE SCOLYTID BEETLES. 

1903. Brichet, 0., et Severin, G. — Le Dendroctonus micans, degats moyens pre- 
ventifs et destructifs.<Bull. Soc. Centr. Forestiere Belg., X, pp. 244-258. 

1903. Baudisch, Fr. — tjber Dendroctonus micans Kug.<Centralbl. ges. Forstwesen, 

Jahrg. 29, pp. 151-152. 

1904. Hopkins, A. D. — Catalogue of exhibit of insect enemies of forests and forest 

products at the Louisiana Purchase Exposition, St. Louis, Mo., 1904. <BuL 
48, Div. Ent., U. S. Dept. Agr. 
1904-5. QuAiRiERE, C. — Le Dendroctonus' micans. <CBul[. Soc. Centr. Forestiere 
Belg., XI, pp. 626-628;- XII, pp. 183-186. 

1904. Powell, P. B. — The development of wings of certain beetles, and some 

studies of the origin of the wings of insects. <Journ. X. Y. Ent. Soc, XII, 
pp. 237-243, Pis. XI-XVII. 

1905. Ibid, (continued), XIII, pp. 5-22. 

1905. Hopkins, A. D. — The Black Hills beetle, with further notes on its distribu- 
tion, life history, and methods of control. <Bul. 56, Bur. Ent., U. S. Dept. 
Agr., pp. 1-24, Pis. I, II, figs. 1-6. 

1905. QuiEVY, Prosper. — Dendroctonus micans, invasion. < Bui. Soc. Centr. Fores- 
tiere Belg., T. 12, pp. 334-335. 

1905. Felt, E. P. — Insects affecting park and woodland trees. <N. Y. State Mus., 
Mem. 8, Vol. 1, pp. 6-7. 

1905. NussLiN, 0. — Leitfaden der Forstinsektenkunde, pp. 175-178. 

1905. Voss, Friedrich. — Uber den Thorax von Gryllus domesticus, mit besonderer 

Berucksichtigung des Fliigelgelenks und dessen Bewegung.<Zeitschr. f. 

wiss. Zool., LXXVIII, pp. 268-521, 654-759, Pis. XV, XVI, XXIV. 
1906a. Hopkins, A. D. — Barkbeetle depredations of some fifty years ago in the Pikes 

Peak region of Colorado. <Proc. Ent. Soc. Wash., VIII, Nos. 1-2, pp. 4-5, July 

13, 1906. 
1906b. Hopkins, A. D. — Notes on scolytid larvae and their mouthparts.<Proc. Ent. 

Soc. Wash., VII, Nos. 2-3, pp. 143-149, Pis. IV, V, January 10. 
1906c. Hopkins, A. D. — Notes on some Mexican Scolytidse, with descriptions of some 

new species. <Proc. Ent. Soc. Wash., VII, Nos. 2-3, pp. 71-81, January 10. 

1906. Webb, J. L. — Some insects injurious to forests. The western pine-destroying 

barkbeetle. <Bul. 58, Pt. II, Bur. Ent., U. S. Dept. Agr., pp. 17-30, Pis. II, 
III, figs. 7-12, August 18. 

1906. Felt, E. P. — Insects affecting park and woodland trees. <N. Y. State Mus., 

Mem. 8, Vol. 2. 

1907. Fall, H. C, and Cockerell, T. D. A. — The Coleoptera of New Mexico. 

<Trans. Amer. Ent. Soc, XXXIII, pp. 145, 218. 

1908. Severin, G. — Le genre Dendroctonus.<CBu\l. Soc. Centr. Forestiere Belg. 

Author's extra, pp. 1-20. Received May, 1908. 
1908. Hopkins, A. D. — Notable depredations by forest insects. < Yearbook L^. S. 
Dept. Agr., for 1907, pp. 149-164. 

o 



Tech. Series 17, Part I, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate III. 




Dendroctonus Adults. 

Fig.l.—D.brevicomis. Fig.2.—D.barben. Fig.Z.—D. convexifroyis. Fig. 4. — D. frontalis. Fig. 5.- 
D. arizonicus. Fig. 6. — I), mexicamts. Fig. 7. — D. paraUelocollis. (Original.) 



Tech. Series 17, Part I, Bureau of Entomology, U. S. Dept, of Agricultur 



Plate IV, 






Dendroctonus Adults. 

Fig. 8.-D. approximatm. Pig. 9.-Z>. monticolx. Fig. IQ.-D. ponderosx. Fig. 11 -D jeffrevi 

(Original.) "^ ' 



Tech. Series 17, Part I, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate V. 




Dendroctonus Adults. 

Fig. 12.— D. simplex. Fig. 13.— D. pseudotsugx. Fig. 14.— Z>. piceaperda. Fig. 16.— D. borealis. 

(Original. 1 



Tech. Series 17, Part I, Bureau of Entomolofry, U. S. Dept. of Agriculture. 



Plate VI 




Dendroctonus Adults. 

Fig. n.—D. obesus. Fig. 18.— i>. ruflpennis. Fig. 20.— Z>. punctatus. Fig. 21.— i>. micans. 

(Original.) 



Tech. Series 17, Part I, Bureau of Entomology, U. S, Dept. of Agriculture. 



Plate VII. 




Dendroctonus Adults. 

Fig. 22. — D. terebrans. Fig. 23. — D. valens. (Original.) 



Tech. Series 17, Part I, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate VII 




Dendroctonus LARV/E. 

Fig. 1. — D. brevicomix; la, spiracle, enlarged; 16, spiracle, greatly enlarged, showing simple 
bifid processes. Fig. 10. — D. ponderosee, spiracle, greatly enlarged. Fig. 13. — D. psru- 
dotsugse, spiracle, greatly enlarged. Fig. 14. — D. piceape'rda, spiracle, greatly enlarged. 
Fig. 19. — D. murrayame; 19a, dorsal aspect of abdominal segments 8 and 9, showing 
plates; 196, anal aspect; 19c, 19r?, and 19e, different aspects of spiracular tnbercle, much 
enlarged Fig. 23. — D. valens: 'Ma, dorsal aspect of thoracic segments; 236, dorsal aspect 
of abdominal segments; 23c, dorsal aspect of abdominal segments 8 and 9, showing 
armed plates; 2'id, ventral aspect of thoracic segments; 23f, ventral aspect of abdominal 
segment; 23/, anal aspect of abdominal segments 8, 9, and 10: 23(7, ^3/;, 23/, different 
aspects of spiracular tubercles, moderately enlarged; 23J, spiracle and spiracular tubercle, 
greatly enlarged. (Original.) 



I 



Technical Series, No. ^7, Part II. 

U. S. DEPARTMENT OF AGRICULTURE. 

BTJRE^^XJ OW EISTTOIMOLOGJ-Y. 

L. 0. HOWARD, Entomologist and Chief of Bureau. 



CONTRIBUTIONS TOWARD A MONOGRAPH OF 
THE SCOLYTID BEETLES. 



IL PRELIMINARY CLASSIFICATION OF THE 
SUPERFAMILY SCOLYTOIDEA. 



A. D. HOPKINS, Ph. D., 

In Charge of Forest Insect Investigations. 



Issued January 9, 1915. 




WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 

1^5. 



''^■■•"^- : .A 

B UREA U OF ENTOMOLOG Y. 

L. 0. Howard, Entoviologist and Chief of Bureau. 
C. L. Marlatt, Entomologist and Acting Chief in Absence of Chief. 
R. S. Clifton, Chief Clerk and Executive Assistant. 

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

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

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

F. M. Webster, in charge of cereal and forage insect investigations. 

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

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

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

RoLLA P. Currie, in charge of editorial work. 

Mabel Colcord, in charge of library. 

Forest Insect Investigations. 

A. D. Hopkins, in charge. 

H. E. Burke, August Busck, W. D. Edmonston, Josef Brunner, T. E. Snyder> 
F. C. Craighead, J. M. Miller, S. A. Rohwer, A. G. Boving, C. T. Greene, 
W. S. Fisher, Jacob Kotinsky, B. T. Harvey, entomological assistants. 

Carl Heinrich, A. B. Champlain, Wm. Middleton, MaryE. Faunceo, Mary C. 
Johnson, scientific assistants. 

Morris Chrisman, Geo. Hofer, J. D. Riggs, P. D. Sergent, J. J. Sullivan, 
Albert Wagner, W. E. Glendinning, J. C. Evenden, F. B. Herbert, P. F. 
Keen, J. H. Pollock, entomological rangers. 

Esther H. Hart, artist. 

Margaret Fagan, Elizabeth PaTcniE, Maude Taylor, Edna 0. Haslup, Pearl 
L. Boone, preparators. 



a Resigned April 1, 1914. 



CONTENTS, 



Page. 

Introduction 165 

Position of the Scolytoidea 167 

General anatomy 168 

Taxonomy 169 

Morphological characters 169 

Superfamily characters. 169 

Family characters 170 

Subfamily characters 170 

Generic characters 171 

The antennae 171 

The eyes 174 

The mouthparts 174 

Characters in general 175 

The body 175 

Form 175 

Color 176 

Vestiture 176 

Sculpture 176 

Armatures 176 

Head 177 

Prothorax 177 

Meso thorax 177 

Metathorax 178 

Legs 179 

Elytra : . . 179 

Posterior wings 179 

Abdominal tergites ISO 

Abdominal spiracles 181 

Abdominal sternites 182 

Stridulating accessories 182 

Internal anatomy 183 

Digestive system 183 

Proventriculus 183 

Reproductive organs 184 

The reproductive organs of the male 184 

Posterior elements 185 

Anterior and median elements 190 

Variations and combinations of the elements 190 

Terminology of the reproductive organs 192 

Classifications based on the reproductive organs 194 

Secondary sexual characters 195 

The pupse 196 

The larvae 197 

The eggs 197 

The embryo 197 

III 



IV CONTENTS. 

Taxonomy — Continued. Page. 

Physiological characteristics 198 

General habits 198 

Classification according to habits 198 

Food habits of the adults 199 

Food habits of the larvae 199 

Pupal habits 199 

Flight habits 200 

Social habits 200 

Galleries 201 

Terminology of the galleries 202 

Classification of the galleries 204 

Egg galleries 204 

Larval mines 204 

Types and subtypes of egg galleries 205 

Relation of types of galleries to the subfamilies and families. . 206 

Taxonomic relation between the beetles and their host plants 207 

Part of plant selected by the beetles 207 

Condition of the plant 208 

Relation of the species of beetles to the systematic position of 

their host 208 

Range of host plants 208 

Associations of species of beetles and species of plants 209 

Summary of taxonomic evidence furnished by host relations .... 209 

Geographical distribution in its bearing on taxonomy 210 

Review of taxonomic characters and characteristics 211 

Morphological characters 212 

Physiological characteristics 213 

Correlation of morphological and physiological elements 213 

Parallel modifications 214 

Reversals 214 

Progressive modifications 215 

Examples of progressive modification 215 

Statistical taxonomy 216 

The species 216 

Varieties, races, etc 216 

The genus 217 

Nomenclature - 218 

Types of genera 219 

Genus Scolytus Geoffrey : 219 

Genus Ips De Geer 220 

Genus Tomicus Latreille 220 

Genus Hylastes Erichson 221 

Genus Cryphalus Erichson 221 

Genus Hypothenemus Westwood 221 

Genus Polygraphus Erichson 222 

Genus Lepisomus Kirby 222 

Description of a new genus and species 222 

Preliminary classification of the Superfamily Scolytoidea 224 

Key to the families 224 

Division 1 224 

Subdivision A 224 

Family Ipidse 224 



CONTENTS. V 

Preliminary classification of the Superfamily Scolytoidea — Continued. 
Key to the families — Continued. 

Division I — Continued. Page. 

Subdivision B 225 

Family Scolytidse 225 

Family Scolytoplatypodidse 225 

Division II 225 

Subdivision C 225 

Family Platypodidae 225 

Family Ipidag. Key to the subfamilies 225 

Division 1 225 

Subdivision A 225 

Subfamily Cryphalinse 225 

Subfamily Ipinse 225. 

Subfamily Corthylinae 225 

Subfamily Micracinae 225 

Subfamily Webbinse 225 

Subdivision B 225 

Subfamily Xyloctoninae 225 

Division II 225 

Subdivision G 225 

Subfamily Crypturginse 225 

Subfamily Phloeotribinse 225 

Subdivision D 225 

Subfamily Hylesininae , 225 

Subfamily Phloeoborinae 225 

Family Scolytidse. Key to the subfamilies 225 

Division 1 225 

Subfamily Goptonotinae 225 

Division II 225 

Subdivision A-. 225 

Subfamily Hexacolinse 225 

Subfamily Bothrosterninse 225 

Subfamily Hyorrhynchinse 225 

Subdivision B 225 

Subfamily Camptocerinse 225 

Subfamily Scolytinse 225 

Family Scolytoplatypodidse - 225 

Family Platypodidse. Key to the subfamiUes 225 

Division 1 225 

Subfamily Platypodinse 225 

Division II 225 

Subfamily Genyocerinse 225 

Subfamily Ghapuisinae 225 

Position of the principal described genera in the preliminary classification 226 

Genera described in 1911-12, not included in foregoing list, but here pro- 

\isionally assigned to the subfamilies 227 

Genera of doubtful position 228 

Bibliography ,. 229 



ILLUSTRATIONS 



PLATES. 

Page. 
Plate IX. Figs. 1-18. — Typical tarsi and tibiae of the subfamilies of Scoly- 

toidea 170 

X. Figs. l-5a. — Antennae of typical species of the subfamilies of Scoly- 

toidea 170 

XI. Figs. 7-lOa. — Antennee of typical species of the subfamilies of 

Scolytoidea - 170 

XII. Figs. 12-18b. — Antennae of typical species of the subfamilies of 

Scolytoidea 170 

XIII. Figs. l-9b. — Tibiae and tarsi of typical species of the subfamilies of 

Scolytoidea 178 

XIV. Figs. 10-lOb. — Tibiae and tarsi of typical species of the subfamilies 

of Scolytoidea 178 

XV. Figs. 12-15b. — Tibiae and tarsi of typical species of the subfamilies 

of Scolytoidea 178 

XVI. Figs. 16-18b. — Tibiae and tarsi of typical species of the subfamilies 

of Scolytoidea 178 

TEXT FIGURES. 

Fig. 96. The antenna in scolytoid beetles: Diagram and terminology 172 

97. Abnormalities in the antennal funicle of Xyleborus tacky graphus 173 

98. Tps emarginatus: Proventricular plate 183 

99. Scolytus scolytus: Proventricular plate 183 

100. Crossotarsus lecontei: Proventricular plate - 183 

101. Dendroctonus '2;aZens; Membranous and nonchitinous elements of male 

reproductive organs 184 

102. Xyleborus saxeseni: Chitinous elements of male reproductive organs . . 184 

103. Ips emarginatus: Chitinous elements of male reproductive organs 185 

104. Pityophthorus s^.: Chitinous elements of male reproductive organs. . 186 

105. Pityophthorus bellus: End plates and seminal rod of male reproductive 

organs 186 

106. Micracis suturalis: Posterior elements of male reproductive organs. . . 186 

107. Dendroctonus valens: Posterior elements of male reproductive organs. 187 

108. Scolytus rugulosus: Posterior elements of male reproductive organs. . 189 

109. Scolytus quadrispinosus : Posterior elements of male reproductive 

organs 189 

110. Crossotarsus lecontei: Posterior elements of male re}) reductive organs. . 191 

111. Dendroctonus valens: Female reproductive organs 194 

112. Classification of galleries of Scolytoidea 205 

VI 



U. S. D. A., B. E. Tech. Ser. 17, Pt. II. F. I. I., January 9, 1915. 

CONTRIBUTIONS TOWARD A MONOGRAPH OF THE 
SCOLYTID BEETLES. 



II. PRELIMINARY CLASSIFICATION OF THE SUPERFAMILY 

SCOLYTOIDEA.'^ 

By A. D. Hopkins, Ph. D., 
In Charge of Forest Insect Investigations. 

INTRODUCTIOSr. 

The object of this contribution is to discuss the taxonomy and 
present a preliminary classification of the families and subfamilies 
of the scolytoid beetles of the world. The discussion and classifi- 
cation are based on a study of representatives of about 122 described 
and undescribed genera and about 1,000 species of North America 
and other countries, in the collections of the U. S. National Museum 
and certain other museums and institutions of this country. 

The types of North American species described by Zimmermann, 
LeConte, Fitch, Harris, Schwarz, Ulke, and, with a few exceptions, 
those of other North American authors, have been studied by the 
writer. Nearly all of the North American species described by 
Eichhoff are represented in the U. S. National Museum collections 
by specimens sent by him to the writer and to Dr. C. V. Riley. 
Many of these specimens are from the type series not only of North 
American but of many foreign species described by Eichhoff. Cen- 
tral and South America, the West Indies, the Eastern Continent, 
Australia, and Oceania are represented to a greater or less extent 
by described and undescribed species, of which 31 genera and 96 
species are from Europe, principally from the collections of Eichhoff 
of Germany, Blandford of England, and Villard of France; 

The material in the forest insect collections of the Bureau of 
Entomology and the West Virginia Agricultural Experiment Sta- 
tion, collected by the writer and his associates, includes nearly all 
of the described North American species and in addition a large 

°- See Hopkins, A. D. List of generic names and their type-species in the coleop- 
terous superfamily Scolytoidea. Proc. U. S. Nat. Mus., vol. 48, No. 2066, pp. 115-136, 
December 16, 1914. 

165 



166 THE SCOLYTID BEETLES. 

number of undescribed species and a great amount of new biological 
material. With this amount of material at hand and a special study 
of this group of insects extending over a period of twenty years, the 
writer feels justified in proposing a revised classification and in 
defining the characters which it appears to him are of special taxo- 
nomic importance. 

In presenting the preliminary classification of the Scolytoidea the 
author does so with no idea of criticising the systems proposed by 
other authors. The whole presentation is simply to stand as the 
writer's interpretation of the characters and characteristics repre- 
sented by the material before him, and to serve, so far as it may, as 
a step in the evolution of artificial systems of classification toward 
the ideal or natural. Each comprehensive system of classification 
proposed since that of Linnaeus has contributed something toward 
the evolution of better systems. Some of them have been progres- 
sive, others in part retrogressive, and a few have been revolutionary 
in their character. But, as in most conflicts of opinion, general 
progress results. Therefore those investigators of the present and 
the future who, without prejudice as to any opinion or theory, can 
sift out the true from the erroneous in that which has been published, 
and add new truths from their own observations, wiU make the 
most rapid progress toward the attainment of the ideal. 

The anatomical investigations conducted by the writer and out- 
lined in Part I of this bulletin have revealed heretofore unrecorded 
facts relating to structural characters, which, in connection with a 
greatly increased knowledge of the physiological characteristics of 
the stages of development and of the habits, host relations, and dis- 
tribution of described and undescribed forms, seem to warrant a 
somewhat different classification from those proposed by other 
writers. 

It seems that a study of the facts as revealed by modifications in 
morphological characters and physiological characteristics of exist- 
ing forms, without any attempt to explain their origin or phylogeny, 
will lead to a more correct interpretation of natural affinities than 
any amount of speculation on hypothetical ancestral forms from 
which present species may have evolved. Indeed, it would seem 
that we have, in the progressive modification of the more funda- 
mental and dominant characters and characteristics, a better taxo- 
nomic basis on which to correlate the characters and construct a 
so-called natural system than can be found in those characters which 
are subject to special modification through similar use and influ- 
ences of environment.^ 



oin this connection see Part I of this bulletin, p. 25, second paragraph, and pp. 
67-68. 



PKELIMHSTARY CLASSIFICATION OF SCOLYTOIDEA. 167 

POSITION OF THE SCOLYTOIDEA. 

The contrasting characters which indicate the position of the super- 
family Scolytoidea in the order Coleoptera are as follows: 

Order COLEOPTERA. 

Maxilla undivided, the palpi rigid and with not more than three joints; larvae leg- 
less Suborder Rhynchophora. 

Maxilla divided, more or less flexible, and with a flexible palpus, usually 4-jointed; 
larvae rarely without legs All other Coleoptera. 

Suborder RHYNCHOPHORA. 

Head without prominent rostrum or beak and the submentum never produced into 
a gular peduncle Superfamily Scolytoidea. 

Head usually with prominent rostrum or beak and the submentum always more or less 
produced into a gular peduncle All other Rhynchophora. 

It is evident to the writer that, in consideration of the extremes in 
morphological characters to be found in the order Coleoptera, the 
superfamily Scolytoidea occupies a position opposite to that of the 
Carabidss, and that in the Rhynchophora it occupies an opposite 
position to that of the Apionid^e. 

It seems desirable to place the Scolytoidea first in the classifi- 
cation of the Coleoptera, not because of any theory as to their origin 
or phylogeny but because their elements of structure seem to form a 
better and more correct basis from which to proceed in the interpre- 
tation of the progressive modification of the characters which serve 
to distinguish the major and minor divisions and groups. Such a 
method should not conflict with other methods because it should 
make little difference whether we begin with the Scolytoidea or the 
Carabidse. If the interpretation of the progressive modifications 
and natural relations is correct, the relative positions of the various 
groups will be the same, or similar. It is simply a matter of 
choosmg between the two directly opposite methods to attain the 
same result and of adopting the one which seems to be more in accord 
with the natural course of change or evolution from one extreme to 
the opposite. 

If we begin with the scolytoid beetles we find throughout the sub- 
order Rhynchophora two dominant or constant characters, namely, 
the rigid maxillae and the legless larvee. We find also certam change- 
able characters repeatedly paralleled in the various major and minor 
groups, and that the general progressive modification of these charac- 
ters, as, for example, the tendency toward a prolongation of the head, 
represents a separate and greatly diverging line of morphological 
expression or evolution from that prevailing in the other divisions of 
the Coleoptera. 

In the other divisions of the Coleoptera the divided maxilla, with 
flexible palpi, and the legged larvee are the prevailing and more domi- 



168 THE SCOLYTID BEETLES. 

nant characters. Here^ again, there are a number of changeable and 
frequently parallel characters, the progressive modification of which 
shows many diverging lines toward the extreme, as, for example, 
the greatest development of jointed and differentiated appendages "■ 
in the larvse, the large size, the odd shapes, and the extraordinary 
elements of structure and habit. 

It is evident that the scolytoid beetles could not have been derived 
from the highly differentiated carabids or scarabaeids and that these 
groups could not have been derived from any existing group of tlie 
Rhynchophora. It is also evident that the species in the two primary 
divisions of the Coleoptera which may be the nearest representatiA^es 
of their primitive ancestors are themselves so highly differentiated 
from one another and from ancestral forms of Coleoptera that they 
can not be recognized, or, if they could, they would be of little or no 
use as a basis for speculation on the origin and evolution of the order; 
then, too, we have no material in fossils on which to base a reliable 
hypothesis, because only the highly specialized forms have been pre- 
served. Therefore we must rely on facts as they exist and endeavor 
to discover and interpret the elements of distinction and relationship 
which have survived m the course of their evolution, under the influ- 
ence of natural selection, dominant differentiation, and progressive 
and parallel evolution. The Avriter's interpretation of these facts, 
so far as they apply to morphological characters and the distinction 
of groups and species, will be expressed m the tables of families, sub- 
families, genera, and species, in the succeeding contributions toward 
a monograph, as will also the progressive modification of characters 
indicative of the lines of divergence from one extreme toward the 
opposite. 

GEN-ERAL AIJ'ATOMY. 

The discussion, illustrations, and terminology of the anatomical 
elements of Dendroctonus given in Part I of this buUetm \^tl11 serve 
as a basis for comparing the anatomy of representatives of other 
genera. The writer has made a detailed study of the entire anatomy 
of only a few representatives of other genera, but the more important 
elements of the external and internal anatomy of representatives 
of all of the genera in the local collections have been studied, involv- 
ing the preparation of over 1,200 microscope slides. This, together 
with the work of other investigators, notably Lmdemann, Niisslin, 
Fuchs, and Eichhoff, gives us a basis for a somewhat comprehensive 
view of the subject. There remains, however, an immense amount 
of anatomical work to be done before a knowledge of the subject is 
anything hke complete. 

o The writer holds that it is by no means proven that larvse with jointed append- 
ages are more primitive than those without such appendages. 



PEELIMIl^AEY CLASSIFICATION OF SCOLYTOIDEA. 169 

TAXONOMY. 
MORPHOLOGICAL CHARACTERS. 

The principal morphological characters adopted by the writer to 
distinguish the families^ subfamilies, and genera, and the major and 
minor divisions of each are to be found in the external anatomy of 
the imago. However, some consideration has been given to the 
location of correlated characters in the internal anatomy and in the 
physiological characteristics of the different stages of development 
from the egg to the imago. 

It has seemed to the writer that in order to have a classification 
which would indicate natural positions and lines of modification, 
and at the same time be simple and practicable, we should endeavor 
to locate and utilize, as far as possible, external characters which 
are readily available for examination by a hand lens or the micro- 
scope without serious mutilation of the specimens by those who make 
use of the systematic tables and descriptions. The small size of most 
of the scolytoid beetles renders it difficult and tedious to examine 
the minute and obscure elements of the external and internal anat- 
omy, such as the mouthparts and the digestive and sexual organs. 
It is very important, in fact essential, that the taxonomist should 
study in detail, and comprehensively, the various elements of exter- 
nal and internal anatomy in order to have a broad basis for his con- 
clusions, but the general student and investigator should not be 
required to go to such extremes in order properly to mterpret the 
conclusions. Therefore it has been the object of the writer to con- 
form to the simple method of expression rather than to that involving 
a complexity of detail. 

The principal charactei -bearing parts of the external anatomy 
which appear to serve as the best taxonomic guides toward a natural 
system of classification are mentioned in the following pages merely 
as a basis for the prehminary classification, which is subject to 
revision in the more detailed treatment of the several subfamilies. 

SUPERFAMILY CHARACTERS. 

The superfamily is at once separated into two primary divisions 
by the relative length of the first tarsal joint, and the subdivisions, 
sections, and famifies are distinguished by characters of the tibia 
(PL IX). It is interesting to note that while the most dominant 
character is found in the first joint of the tarsus, the modification 
of this joint withm the famifies, subfamilies, and genera is of minor 
importance. The same, to a lesser degree, may be said of the tibia, 
in which the character of the apical angles is constant within each 
subdivision of the superfamily but the modification of these angles 



170 THE SCOLYTID BEETLES. 

throughout the minor groups to the species, except ia a few cases, 
is of httle or no importance. 

The complete opposition in the characters of the apical angles of 
the tibia, as shown in the subdivisions, is of special interest. There 
is no apparent explanation on any theory of use or natural selection, 
but it does appear to signify widely diverging lines of descent in 
which this reversed element in the two subdivisions has remained 
as the dominant and distinctive character. 

FAMILY CHARACTERS. 

In the classification of the famihes Ipidse and Scolytidse into sub- 
famihes the principal characters used to distinguish the divisions 
are found in the head (2)", pronotum (4), and tarsi (2). 

The subdivisional characters are found in the antennae (2), tibia (2), 
and abdominal sternites (2). 

The sectional characters are found in the antennae (2), eyes (2^. 
pronotum (2), tibia (4), and form of body (2). 

The subfamily characters are found in the pronotum (4), antennae 
(2), seventh abdominal sternite (2), elytra (2), and third tarsal 
joint (2). 

It will be seen that in the classification of the two families into 
subfamihes the principal characters are found in the head (8), pro- 
notum (10), and tibia (7). 

SUBFAMILY CHARACTERS. 

In the classification (in manuscript) of the 16 subfamilies of Ipidae 
and Scolytidae into genera the principal character-bearing parts for 
the divisions, subdivisions, and sections are as follows: 

Divisions: Head (1), antennal club (3), funicle (3), eyes (^), epis- 
toma (2), maxiUa (1) — total for head (14); pronotum (5), tarsi (3), 
and elytra (2). 

Subdivisions: Antennal club (5), eyes (4), funicle (2), epistoma (1), 
maxiUa (1) — total lor head (13); third tarsal joint (2), anterior 
coxae (3), seventh abdominal sternite (2), and pygidium (2). 

Sections: Antennal club (4), funicle (2), -eyes (1) — total (7). 

It will be seen that in the classification of the subfamilies the 
characters are found in the head (35), tarsi (7), pronotum (5), 
elytra (4), coxae (3), seventh abdominal sternite (3), pygidium (2), 
and tibia (1). 

Subsections and series to genera: The dichotomous characters of the 
subsections and series and the distmctive characters of the genera 
are as follows: Antennal club (138), funicle (110), scape (7) — total for 

a The number following the name of a structure indicates the relative importance 
or the number of times it figures in the dichotomy. 



Tech. Series 17, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate IX. 




9 10 12 13 15 16 











Typical Tarsi and Tibi/e of the Subfamilies of Scolytoidea. 

Fig. 1.— Cryphalinffi {Cryphalus asperatus). Fi?. 2.— Ipin;B (7p.s tyjxjgraphus). Fig. 3.— Corthylinse 
( Corihylus columbianus). Fig. 4. — Micraeinee {Micracis suturalis). Fig. 5. — Webbinse ( Webbia dip- 
terocarpi). Fig. G.— Xyloctoninse {Xyloctonus scolytoides) (adapted from Eichhofl). Fig. 7.— 
Crypturginse ( Cryplurgus pusillus). Fig. 8.— PhloeotribiiiEe {Phlceotribus olex). Fig. 9.— Hylesininas 
{Hylesmus crenatus). Fig. 10.— Phloeoborinse {Phlceoborus rudis). Fig. 12.— Hexacolinge {Hexacolus 
sp.). Fig. 13.— BothrosterniiiEe (Bothrosternus sculpturatus) . Fig. 1.5.— Camptocerinas ( Camp- 
tocerua leneipennis). Fig. 16.— Scolytinse (Scolytus scolytus). Fig. 17.— Scolytoplatypodinee {^coly- 
toplatypus sp.). Fig. 18. — Piatypodrnge (Platypus cylindrus). (Original.) 



Tech. Series 17, Part II, Bureau of Entormology, U. S. Dept. of Agriculture. 



Plate X. 







Antenn/e of Typical Species of the Subfamilies of Scolytoidea. 

Fig. 1.— Crnphalus asperatus (Cryphalinffi), anterior aspect of left antenna. Fig. la.— Posterior aspect 
of left antenna of same. Fig. 2.—Ips typographus (Ipinfe) , anterior aspect of left antenna. Fig. 2a.— 
Posterior aspect of left antenna of same. Fig. Z.— Corthylus columbianus (Corthylinfe), female, 
anterior aspect of left antenna. Fig. 3a.— Female of same, posterior aspect of left antenna. Fie. 
3b.— Male of same, club. Fig. i.— Micracis suturalis (Micracinee), female, anterior aspect of left 
antenna. Fig. 4a.— Female of same, posterior aspect of left antenna. Fig. 46.— Male of same, scape. 
Fig. 5.— Webbia dipterocarpi (Webbinae), anterior aspect of left antenna. Fig. oa.— Posterior aspect 
of left antenna of same. (Original.) 



Tech. Series 17, Part II, Bureau of Entomology, U. S, Dept. of Agriculture. 



Plate XI. 










Antenn/e of Typical Species of the Subfamilies of Scolytoidea. 

rig. 7. — Crypturgus jjusillus (Crj'ptiirginge), anterior aspect of left antenna. Fig. 7a. — Posterior aspect of 
left antenna of same. Fig. 8. — Phlaotribus oZcas (Phloeotribinse), anterior aspect of left antenna. Fig. Sa. — 
Posterior aspect of left antenna of same. Fig. 9. — Hylesinus crenatus (HylesintniB), anterior aspect of 
left antenna. Fig. 9o. — Posterior aspect of left antenna of same. Fig. 10. — Phlceoborus rudis (Phlceo- 
borinse), anterior aspect of left antenna. Fig. lOo. — Posterior aspect of left antenna of same. (Original.) 



Tech, Series 17 Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate XII. 




Antenn/e of Typical Genera of the Subfamtlies of Scolytoidea. 

Fig. 12. — Hexacolus sp. (HexacolinEc), anterior aspect of left antenna. Fig. 12a.— Po.sterior aspect of left 
antenna of same. Fig. IZ. — Bothrosternus sailpturatus (Bothrosterninse), anterior aspect of left antenna. 
Fig. 13a. — Posterior aspect of left antenna of same. Fig. 15. — Campfocerus seneipcnnis (Camptocerinfe), 
anterior aspect of left antenna of female. Fig. 15o.— Club of female of same. Fig. 15b.— Scape of male of 
same. Fig. lC>.—Scohjtus scohjtus (Scolytina;), anterior aspect of left antenna. Fig. Ifin.- Posterior 
aspect of left antenna of same. Fig. 17. Scolytoplaty pus sp. (ScolvtoplatAi)odin8e), anterior aspect of 
left antenna. Fig. 17a.— Posterior aspect of left antenna of same. Fig. 18.— Pteh/D«.s cylindrus (Platy- 
podinaj), anterior a.=pect of left antenna of female. Fig. ISa.— Posterior aspect of left aiitenna of same. 
Fig. 18&. — Male scape of same. (Original. ) 



PEELIMINAEY CLASSIFICATIOlSr OF SCOLYTOIDEA. 



171 



antennas (145); eyes (143), epistoma (6), front of head (2) — total 
for head including antennae (297); pronotum (29), elytra (12), tarsi 
(8), tibia (6), form of body (5), pygidium (2), abdominal sternites (2), 
and coxse (5). Thus it is shown that the most important generic 
characters are found in the antennae and eyes. 

A summary of the principal character-bearing parts, as recognized 
by the writer in his preliminary classification, may be tabulated as 
follows : 

Table I. — Summary of principal character-bearing parts in the superfamily Scolytoidea. 



Characters. 



In the su- 
perfamily. 



In the 
families. 



In the sub- 
families. 



In the sub- 
sections 

and series 
to the 
genera. 



Tarsi, first joint 

Tibia, apical angles 

Head 

Pronotum 

Elytra 

Third tarsal joint 

Coxag 

Seventh abdominal sternile . 
Pygidium 



Timesused. 
2 
3 



Timesused. 



Timesused. 
7 
1 
35 
5 
4 
7 

3 
2 



Timesused. 



6 

297 
29 
12 
8 
5 
2 



GENERIC CHARACTERS. 

The Antenna. 
.(Fig. 96; Pis. X-XII.) 



ANTENNAE FUNICLE. 

The antennal funicle is perhaps the most important taxonomic 
element of the scolytoid beetles. It is one of the first things to be 
looked for as a guide to the combination of characters which distin- 
guish the genus and, at the same time, indicate its position in the 
classification. While the same number of joints may be paralleled 
many times in the genera of the same subfamHy and in different sub- 
families, the writer holds that, with very few exceptions, there must 
be the same number of joints in the funicle of all of the species of a 
genus. The exceptions are found in Hypotlienemus and StepTiano- 
deres, in which the males are smaller than the females and the an- 
tennal funicle has a less number of joints. The males in Xylehorus 
and allied genera are also smaller than the females, but usually have 
the same number of joints. Occasionally there is a smaller number 
or other abnormal developments. (See fig. 97.) 

The writer's conclusions are based on the study of balsam mounts 
of representatives of all of the genera which have been available to 
him, and of a large percentage of the commoner species. It has 
been found that a normal variation in the number of joints m the 



172 



THE SCOLYTID BEETLES. 



same sex of the species of a well-defined genus is exceedingly rare. 
Therefore, an odd number of joints in examples of the \lominating 
sex either mdicates an abnormal development or a different genus. 
If abnormalities occur, they may be easily detected by the combina- 
tion of other generic characters. The number of joints in the anten- 
nal funicle of the scolytoid beetles is limited to seven, and there is 

probably no normal 
exception to this in 
the superfamily, and 
probably not in the 
entire Rhynchophora, 
or, if so, it will be ex- 
ceedingly rare. 

According to the 
number of j oints m th e 
funicle, the 221 gen- 
era, includmg many 
new ones, are distrib- 
uted as follows: 

Table II. — Funicular 
joints in the superfamily 
ScolytoideaA 

1 joint in 3 genera rep- 
resenting 1 subfamily. 

2 joints in 6 genera rep- 
resenting 2 subfamilies. 

3 joints in 13 genera rep- 
resenting 4 subfamilies. 

4 joints in 44 genera rep- 
resenting 7 subfamilies. 

5 joints in 73 genera rep- 
resenting 9 subfamilies. 

6 joints in 31 genera rep- 
resenting 9 subfamilies. 

7 joints in 51 genera rep- 
resenting 9 subfamilies. 

Total, 221 genera repre- 
sentins: 16 subfamilies. 




Fig. 96. — The antenna in scolytoid beetles: Diagram and terminology. 
Scrobe: a, Fossa. Scape: b, Condyle; c, basal section or neck; d, elbow; 
e, median section;/, apical section; /a, dorsal fringe; g, apical fossa; ga, 
apical margin. Funicle: h, Base or condyle; ?', basal section; j, me- 
dian section; k, apical section; fee, apical angle; kb, dorsal lobe; I, 
ventral angles of joints; in, dorsal angles; ma, fimicular fringe or 
dorsal bristles; n, apical joint. Club: o, Basal annulation; p, apical 
or sutural annulation of joint 1; q, ventral septum or single septum; 
r, dorsal and ventral septa or double septa; s, chitinous suture; t, 
ventral fringe; ta, ventral bristle; tb, procumbent bristle; tc, re- 
cumbent bristle; u, setaceous annulation or sutural annulation; v, 
sensitive pores; w, sensitive and setal granules; wa, sensitive area; 
wb, chitinous area. Club: A, solid; B, amiulated; C, conical; D, 
separated joints or sublamellate; E, compressed; F, thickened at 
base; O, obliquely truncate; H, antenna extending at right angles 
to the head. (Original.) 



Among eight genera 
of doubtful position 
the number of joints 
in the funicle is not given m the description of one genus. Three 
genera have 5, one has 6, and three have 7 joints. 

Within the subfamilies the progressive modification in generic 
characters and characteristics appears to be associated with the 

«This table includes eleven recently described genera which, have been provi- 
sionally referred to subfamilies. 



PRELIMINARY CLASSIFICATIOlSr OF SCOLYTOIDEA. 



173 



increase in the number of joints in the funicle. The exception to 
this rule appears to be shown in the Corthylmse where the one-jointed 
and two-jointed f unicles are characteristic of genera which, in certain 
hnes, have progressed further than genera in other subfamiUes with 
five, six, and seven joints. But it is by no means certain that these 
Corthyhnse are not highly specialized survivors of one of the most 
ancient groups in which a one-jomted funicle became the fixed and 
dominant character. 

It is interesting to note in Table II that the five-jointed funicle 
is common to the greater number of genera. AU but two of the 
classified genera with a five-jointed funicle fall in the subfamilies of 
the Ipidse and the 
large majority of 
them in the first di- 
vision ; while the 
genera with a seven- 
jointed funicle faU 
in the last part of 
Division II of the 
Ipidse and in the 
family Scolytidse. 



ANTENNAL CLUB . 

The antennal club 
comes next to the 
funicle as a bearer 
of important generic 
characters. The 
range in modifica- 
tions of form is 
from a narrow, 
somewhat c o m - 
pressed and distinctly annulated club like that of Hypothenemus and 
PityopUhorus to the broad, thinly compressed, thickened at base, 
obliquely truncated, solid conical, or separated joint form. 

In 179 genera 123 have a more or less compressed and annulated 
club, m 44 the club is thickened at base, and in 12 it is conical. In 
170 genera 144 are annulated, 36 not annulated, and in 9 the joints 
are separated. In 136 genera 52 have sutural septa, 74 are without 
sutural septa, and 10 have chitinized sutures. 

It appears that the chitinized septum of the club, while variable 
and paralleled in different genera, is a good generic character. The 
septa vary m number from one to four and usually occur toward 
the ventral margin, but may also occur toward the dorsal margia 
of one or two sutures. The range of modification appears to be 
59026°— 15 2 




Fig. 97.— Abnormalities in the antennal fxmicle of Xylehorus tachy- 
graphus: a. Anterior aspect of right antenna of female; 6, posterior 
aspect of club of a; c, anterior aspect of left antenna of male; d, pos- 
terior aspect of club and funicle of c; e, anterior aspect of right club 
and funicle of same specimen as c;f, posterior aspect of e; g, posterior 
aspect of right antenna of another specimen; ft, anterior aspect of 
g. (Original.) 



174 THE SCOLYTID BEETLES. 

from a club with one or more septa to an annulated club with chiti- 
nized sutures and without septa, to a solid club without annulations 
or septa, or to a conical club with chitinized joints. 

The septum is evidentlj^ the remnant of the chitinized elements 
of a movable joint, indicating that the progressive modification of 
the antennee toward the opposite extreme has been by the process 
of reduction or fusion of two or more joints, while the modification 
of the funicle has evidently been along the Ime of accession by divi- 
sion as is indicated in the funicle of the retrograde sexual forms 
(fig. 97) and in the nymphs of certain Hemiptera and Isoptera. 

Thus we have in a single organ evidence of progressive modifica- 
tion by reversed processes which is not unreasonable and does not 
necessarily conflict with the facts and prmciples of other evolu- 
tionary processes. 

ANTENNAL SCAPE. 

The antennal scape is variously modified from simple and slender 
to short, stout, dilated, and fringed, but is of less importance as the 
bearer of generic characters than either the funicle or the club. 

The Eyes. 

The eyes are variously modified and range from simple, elUptical, 
round or oval, to emarginate or completely divided and from widely 
separated on the dorsal or ventral area to approximate on one or 
on both areas. In 114 genera the eyes in 65 are simple, in 98 emargi- 
nate, and in 10 divided. Among those with simple eyes, 4 have them 
approximate on the dorsal or ventral areas. 

The Mouthparts. 

The characters of the mouthparts have been quite extensively 
used by systematists m the definition of genera, but while the writer 
recognizes that some excellent characters are to be found in the 
mouthparts, he is convinced that they are by no means essential for 
the definition or classification of the genera. The principal objection 
to their use, as every systematist has doubtless recognized, is that 
they are not available for interpretation without mutilating the spec- 
imen, which in the case of rare or unique specimens and those 
from other collections is out of the question. Another serious objec- 
tion is in the fact that no two balsam mounts of a maxilla of the same 
species present the same contours and angles, and, therefore, these 
may appear to be quite different except, perhaps, in the number and 
relative lengths of the joints of the palpus. 



PEELIMIISrARY CLASSIFICATION OF SCOLYTOIDEA. 175 

EPISTOMA. 

The epistoma in all of the species examined shows more or less 
important generic and specific characters. There is a wide range of 
modification, from a simple, transverse, chitinous piece with few or 
no epistomal bristles to the epistomal process of Dendroctonus and 
the exceedingly long epistomal horn of Cactopinus, or the flattened 
labral-like form in Pycnarthrum. 

HYPOSTOMA. 

The hypostoma is also quite variable and can be used to good 
advantage in the more detailed definition of a genus or larger group. 

LABRUM. 

The absence of a true labrum in the adult scolytoid beetle appears 
to be universal. It has been stated by Eichhoff that it was present 
in Pijcnarthrum, but an examination of a balsam mount has convinced 
the writer that this is only a produced median area of the epistoma. 

CHARACTERS IN GENERAL. 

The Body. 

The size, form, color, vestiture, sculpture, and armature of the 
body represent characters of more or less importance in indicating 
lines of progressive modification and as aids in generic and specific 
definition. The size ranges from 0.4 or 0.5 mm. in the males of some 
Hypothenemus to 13 mm. in Phloe-oborus. While there is more or 
less variation in the size of the individuals of a species, the length is 
important not only as a guide to the recognition of a species but as 
an index to its proper position in the minor section of the genus to 
which it belongs. The writer has found that, as a rule, the smaller 
species of a division, subdivision, or section of a genus will, according 
to other correlated characters, occupy a position opposite to that of 
the larger species. In the system adopted by the ^i^Titer the genera 
with the average smaller species come first in the subfamily and in 
each primary and minor division, and in the genus the smaller forms, 
as a rule, precede the larger. There are, of course, exceptions to this 
rule, especially in genera with few and widely separated species. 

FORM. 

The form of the body ranges from elongate and slender to short and 
stout, and it would appear that the range in progressive modification 
is from the slender to the stouter forms. 



176 THE SCOLYTID BEETLES. 

COLOR. 

The color, except in a few genera, is of little taxonomic importance. 
It ranges from pale yellow through yellowish red, reddish brown, 
brown, and black. In a few genera the chitinous integument of the 
elytra or other parts of the body is bicolored; in others the varie- 
gated color is confined to the vestiture. Metallic and iridescent 
colors are rare. 

VESTITURE. 

The vestiture is of considerable taxonomic importance. It con- 
sists of scales, stout hairs, barbed hairs, plain hairs, fine pubescence, 
gummy exudations, or adherents. The range in progressive modi- 
fication of the vestiture appears to be from scales to stout hairs, from 
barbed hairs to simple hairs, and from a sparsely pubescent to an 
entirely glabrous body. 

SCULPTURE. 

In the sculpture of the body there is endless variety. It may be 
rugose or smooth, the rugosity fine or coarse, the punctures sparse or 
dense, arranged in rows or confused, regular or irregular in size, 
irregularly distributed on given areas, etc. Some of the elements 
of sculpture, such as the rugose or smooth pronotum, are of value 
in separating the major and minor groups of the famihes, but the 
characters of the rugosities and punctures are of special value in 
defining the smaller groups and species. Concavities and convexi- 
ties of the front of the head and of the apical dechvity of the elytra are 
often important generic, specific, and secondary sexual characters. 

ARMATURES. 

The armatures of the pronotum, head, and elytra are important 
in the definition of genera, species, and sexes. The armature of the 
head reaches its extreme development in the epistomal horn of 
Cactopinus. The armature of the declivity is strongly developed 
in Xylehorus, Xylodeptes, and Eccoptopterus of the Cryphahnse; in 
Pityogenes and Ips of the Ipinse; in Hylocurus in the Micracin?e; in 
Monarthrum and AmpJiicranus of the Corthylinse, and in most of 
the genera of the Platypodinse. As a rule the modification from a 
simple unarmed body to one with moderately or strongly armed 
parts is correlated with other elements of progressive modification. 
The serrate armature of the anterior margin of the pronotum is of 
considerable taxonomic importance. The apical serrations are com- 
mon in the Cryphalinse and Ipinse, but rare or absent in the other 
subfamihes. The extremes in apical armature are found in Hijpo- 
thenemus miles Lee, and in some other species, as, for example, the 



PEELIMINAKY CLASSIFICATION OF SCOLYTOIDEA. 177 

males of certain species of Xyleborus, and in Ampliicranus fastigatus 
Blndfd. The modification of the apex appears to progress from an 
apically serrate to a simple one or to one with an apical process. 

HEAD. 

The head is the most important part of the body in representing 
the greatest number of taxonomic characters. Its modification is 
from a short and narrow or broad and globular form concealed from 
above by the pronotum, toward a narrow elongate form exposed 
beyond the apical margin of the pronotum. The tendency toward 
the prolonged subrostrate form indicates a line of progressive modi- 
fication which is characteristic of the major and minor groups of all 
of the Rhynchophora, the extremes of which are found in the small, 
short, globular head of the Ipidse and the exceedingly prolonged 
beak of Apion, Balaninus, and other genera. In the Scolytoidea, 
Cosmoderes to StepJianoderes of the Cryphalinse have the shortest, 
simplest head, while the extreme is reached in certain genera of the 
Hylesininse, Hexacolinse, and Chapuisinae, with the extreme, or nearest 
approach to CurculionidaB, in Hylurgops and Hylastes. With a few 
exceptions the concealed head is characteristic of the first division 
of the Ipid« and the Scolytidse, while the exposed head is charac- 
teristic of the second division of the Ipidse and the Platypodidse. 
The antennae, eyes, and mouthparts have been discussed in pre- 
ceding pages. The front of the head is of special importance in 
defining the species and. sexes, and ranges from convex and glabrous 
to deeply concave and densely pubescent. 

PROTHORAX. 

The prothorax (Part I, figs. 16 and 17, pp. 23-24) represents a 
wide range of progressive modification from long and narrow to 
short and broad; the pronotum with sides and base not margined 
to margined and simple or to emarginate, while the anterior dorsal 
area ranges from closely and finely rugose to coarsely asperate, and 
from alutaceous or opaque to smooth and shining. The pleura 
range from convex to flat and concave, the anterior coxae from con- 
tiguous to widely separated, and the sternal, sterneUar, and post- 
sterneUar areas vary with the form of the prothorax and the size 
and position of the coxae. The vestiture and sculpture of the an- 
terior median and posterior dorsal and lateral areas often represent 
generic and specific characters of special importance. 

MESOTHORAX. 

The meso thorax (Part I, figs. 18, 19) is exceedingly variable, 
conforming to the variable form of the body. It doubtless contains 
many group, generic, and specific characters which may settle difficult 



178 THE SCOLYTID BEETLES. 

questions of distinction and definition and should be considered in all 
detailed studies of single genera and allied groups of genera. The 
scutellum has been extensively used by systematists, and a detailed 
comparative study of this element in a wide range of genera and 
species should be made to determine its real significance m taxonomy. 
The pleurum (Part I, fig. 19) is also exceedingly variable in the form 
and sculpture of the epimeron, episternum, and preepisternum, while 
the remarkable structure designated as the preepisternal process 
(Part I, p. 29) is of special taxonomic importance since its presence 
or absence is peculiar not only to groups and divisions of the Scoly- 
toidea but to many other groups of Coleoptera, and evidently represents 
an extreme in progressive modification which, in connection with 
other progressively modified elements, will doubtless serve as an index 
to the systematic position of genera and species. The elements of 
the sterna (Part I, fig. 18) are also variable, but, like the prosterna, 
their modification conforms to the variable form of the body, and they 
are of less importance than the more independent structures like the 
scutellum and preepisternal process. 

METATHORAX. 

The metathorax (Part I, figs. 20, 21) is quite variable in general 
characters and especially so in some of the elements of the tergum, 
probably due to the variable form cf the body and the requirements 
of flight. There is considerable variation of this element within the 
species of a genus and often there are wide differences m allied genera. 
Perhaps the element subject to the greatest variation is the postscutel- 
lum, which ranges from obscure or rudimentary to almost the length 
of the combined anterior elements. The scutellar groove, the trans- 
verse sutures, and the entothoracic ridges or apodemes are also quite 
variable. The writer has examined the metatergum of quite a large 
number of species and it would appear that there may be in it some 
important characters peculiar to ininor groups cf genera, but the 
taxonomic value of the variations is largely limited to the species. 
It would therefore be difficult to trace correlated lines of progressive 
modification. Nevertheless, a special comparative study of the 
metathorax should be made of a very large number of examples repre- 
senting all of the genera to determine whether or not there are special 
taxonomic elements or recognizable lines of modification. The epi- 
sternum (Part I, fig. 20) is quite variable in length, mdth, sculpture, 
and vestiture and is of considerable taxonomic importance. The 
metasterna are also variable in conformity with the variable form of 
the body. The sternum always occupies the greater part or nearly 
all of the area. The greatest range of variation between the elements 
of the metathorax is to be found in the short and broad forms of the 
stouter species of the Ipidse and Scolytidge and the exceedmgiy long 
and narrow forms peculiar to the Platypodidse. 



Tech. Series 17, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate XIII. 




TiBi/E AND Tarsi of Typical Species of the Subfamilies of Scolytoidea. 

Fig. l.— Cryphalus asperatus (Cryphalinfe), left tibia, dorsal aspect. Fig. la.— Left tibia of same, 
ventral aspect. Fig. 16.— Tarsus of same, lateral aspect; c, third tarsaljoiat, dorsal aspect. Fig. 2. — 
Ips typjographus (Ipinse), left tibia, dorsal aspect. Fig. 2a. — Left tibia of same, ventral aspect. 
Fig. 26. — Left tarsus of same, lateral aspect. Fig. 3. — Corthylus columbianus (Corthyliufe), left 
tibia of same, dorsal aspect. Fig. Sa. — Left tibia of same, ventral aspect. Fig. 36. — Left tarsus of 
same, lateral aspect. Fig. 4. — Micracis suturalis (Micracinse), left tibia, dorsal aspect. Fig. 4(i. — 
Left tibia of same, ventral aspect. Fig. 46.— Left tarsus of same, lateral aspect. Fig. 5.— Webbia 
dipterocarpi (Webbinse), left tibia, dorsal aspect. Fig. 5a.. — Left tibia of same, ventral aspect. 
Fig. 56. — Left tarsus of same, lateral aspect. Fig. 7. — Crypturgus pusillus (Crypturginse), left tibia, 
dorsal aspect. Fig. 7a.— Left tibia of same, ventral aspect. Fig. 76. — Left tarsus of same, lateral 
aspect. Fig. 8. — Phlaotribus oleas (Phloeotribina;), left tibia, dorsal aspect. Fig. 8o. — Left tibia of 
same, ventral aspect. Fig. 86. — Left tarsus of same, lateral aspect; c, third tarsal joint of same, 
subdorsal aspect. Fig. 9. — Hylesinus venatus (Hylesininse), left tibia, dorsal aspect. Fig. 9a. — Left 
tibia of same, ventral aspect. Fig. 96. — Tarsus ofsame,sublateral aspect. (Original.) 






o S2 




0. a-o 




w »g- 




9 gS 






<^ 


H •»"- 


^ 


\i ^J. 


\ 


o ^7 
g If 












< -3 


1 y^ 


:p\ ^ -s 



Tech. Series 17, Part II, Bureau of Entomology, U. S. Dept. of Agriculture, 



Plate XV. 






TiBi/E AND Tarsi of Typical Species of the Subfamilies of Scolytoidea. 

Fig. 12.— Hexacolus sp., left tibia, dorsal aspect. Fig. 12a.— Left tibia of same, ventral aspect. Fig. 
12&.— Left tibia of same, lateral aspect. Fig. n.—Bothrosternus sculpturatus (Bothrosternma), 
lefttibia, dorsal aspect. Fig. 13a.— Left tibia of same, ventral aspect. Fig. 13&.— Left tarsus of sanie, 
lateral aspect. Fig. 16.— Camptocems leneipennis (Camptocerinse), left tibia, dorsal aspect._ Fig. 
15a.— Left tibia of same, ventral aspect. Fig. 15b.— Left tarsus of same, dorsal aspect. (Origmal.) 



Tech. Series 17, Part II, Bureau of Entomology, U. S. Dept. of Agriculture. 



Plate XVI. 





TiBi/E AND Tarsi of Typical Species of the Subfamilies of Scolytoidea. 

Fig. 16. — Scolytus scolytus (Scolytiiise), left tibia, dorsal aspect. Fig. 16a. — Left tibia of same, ventral 
aspect. Fig. 166. — Left tarsus of same, dorsal aspect. Fig. 17. — Scolyfoplatypus sp. (Scolytoplaty- 
podinee), left tibia, dorsal aspect. Fig. 17a. — Left tibia of same, ventral aspect. Fig. 176. — Left 
tarsus of same, dorsal aspect. Fig. 18.— Platypus ci/lindrus (Platypodina;), left tibia, dorsal aspect. 
Fig. 18a.— Left tibiaof same, ventralaspect. Fig. 186.— Lefttarsusof same, lateralaspect. (Origmal.) 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 179 

LEGS. 

There is a wide range of variation in the form, color, sculpture, and 
relative proportions of the coxa, trochanter, tibia, and tarsus of the 
anterior, median, and posterior legs. (Part I, figs. 1, 3, 26-29.)^ 
The anterior tibia and tarsus are of special taxonomic importance in 
distmguishing the primary and secondary divisions of the superf amily 
and, to a less extent, in distinguishing the primary or minor groups of 
the families, subfamilies, and genera. (Pis. X, XIII-XVI.) The 
variation in the tibia is from a simple, compressed, slightly dilated 
form with the outer margin serrate, as in Hypothenemus, to a short, 
broad form with parallel, smooth margins, as in Micracis and Scolytus, 
or broader at the base, as in Wehhia and Hypohorus, and to extreme 
and odd forms as in Platypus. The character of the vestiture usually 
conforms to that of the body, such as scales, barbed hairs, and simple 
hairs, varying in distribution and density on the ventral and dorsal areas 
and the margins. The sculpture ranges from smooth to imbricate 
and from irregular rugosities to prominent ridges, the latter reaching 
a maximum development in Platypodidas. The tarsi vary in form 
from slender to stout and the joints in relative lengths, widths, and 
vestiture. The third joint varies from simple to emarginate and 
deeply bilobed, with the ventral surf ace ranging from nearly glabrous 
to pubescent and to densely padded, which latter extreme is found in 
PKloRoborus . 

ELYTRA. 

The elytra, or anterior wings, are exceedingly variable in form, 
vestiture, and sculpture, ranging from the simple types with scales, 
fine punctures, and obscure striae which are not impressed, as in 
Hypothenemus, to the forms \vith hairs and with distinctly impressed 
stri^ and elevated interspaces, the latter with rugosities and rows of 
punctures or smooth and without punctures; the base from plain to 
strongly elevated and serrate; the sides from parallel to converging 
posteriorly or strongly rounded; the dorsal area from flat and straight 
to convex and strongly rounded from base to apex; the declivity 
from plain, steep, and convex to retase or armed and strongly oblique 
from base to apex, and the side margins from serrate to straight or 
emarginate. With all of these almost endless variations and their 
different combinations of elements there is available a profusion of 
characters for the definition of groups of genera and species. 

POSTERIOR WINGS. 

The true functional wings, as pointed out by Niisslin (1911), are 
quite variable in form, proportions, and character of the venation 
and represent two specified types, one with and the other without 

o Figures 26-29, Part I, represent the reverse faces of the right instead of the left 
tibia, a mistake which was unfortunately overlooked in the manuscript and proofs. 



180 THE SCOLYTID BEETLES. 

a basal lobe. The writer has given considerable attention to the study 
of the wings of the Scolytoidea and other insects, but he has failed 
to find any constant and readily recognizable characters in the 
Scolytoidea which appear to be of sufficient taxonomic value to 
justify giving them special attention. If there is any particular 
fine of progressive modification in the wings it is to be found in the 
subfamilies and minor groups, in which the range appears to be from 
a simple type, like that of Hypothenemus, CrypJialus, and Cryp- 
turgus, with a narrow, simple base, long fringe, and simple venation, 
toward a broader base, lobed or not, and with an increasing number 
and complexity of veins. The writer reahzes that the complex 
type of venation is generally supposed to be more primitive than 
the simple type with few or no veins, but he is by no means convinced 
that this is the correct interpretation as applied to the wings of all 
insects. The wings in different orders of insects may be, in spite of 
the prevailing opinion, the result of independent origin from simple 
types of primitive winglike processes, and their evolution may have 
been influenced by two primary factors: (1) A dominant tendency 
to perpetuate and promote lines of modification pecuhar to and in 
conformity with the dominant morphological characters peculiar to 
the order, and (2) adjustment of this modification to the peculiar 
mechanical needs of the varying related forms, with frequent examples 
of parallel modifications in unrelated species. 

In Niisslin's table (1911, pp. 302-304) the wings without basal 
lobes are found in widely separated genera, representing, according 
to the present writer's classification, two families (Ipidse and Scolytidse) 
and five subfamilies, while the wings with basal lobes are found.infive 
subfamilies of the Ipidse. It is evident that whenever the wings 
are studied in their relation to other of the more important taxonomic 
characters and characteristics of the species, the variations noted 
by Niisslin will be of considerable value in the definition of minor 
gi^oups of genera and species, but the difficulty of spreading the 
wmgs and securing good balsam mounts wiU preclude their general 
use.' 

ABDOMINAL TERGITES. 

The general character of the abdominal tergites is shown in Part I, 
figures 22, 23, and 24. While there is considerable variation in the 
fii'st to sixth tergites, inclusive, in the same individual and between 
individuals of different species, the seventh and eighth are the ones 
of special importance in the identification of the genus or sex. Ac- 
cording to Niisslin (1911), who examined 16 genera, the eighth ter- 
gite is not covered in either sex in 11 genera, and is exposed in the 
male and covered in the female in 5 genera. The writer has found 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 



181 



the eighth to be uncovered in both sexes in three genera, uncovered 
in the male and covered in the female in 13 genera, and covered in 
both sexes in 3 genera. 

According to the writer's classification, the 31 genera in which 
species were examined by Niisslin and the writer represent the 
following subfamilies : 

Table III. — Abdominal tergites in the subfamilies of Scolytoidea in which species were 
examined by Niisslin and the writer. 



Subfamily. 



Cryphalinae 

Ipinse 

Corthylinae 

Micracinse 

Crypturginse... 
PMoeotribinas . 

HylesiniiiEe 

Scolytinse 

Platypodinffi . . 



Eighth ter- 
gite uncov- 
ered in both 
sexes. 



Genera. 



Total. 



Eighth ter- 
git« uncov- 
ered in male, 
not in female. 



Genera. 



Eighth ter- 
gite covered 
in both sexes. 



Genera. 



From the foregoing it will be seen that the characters of the seventh 
and eighth tergites are paralleled in widely separated genera and sub- 
families. The writer has found that the eighth tergite may or may 
not be exposed or covered in the same genus or in individuals of the 
same species, especially in Platypus, where the eighth tergite is of the 
same or similar form in both sexes and may be covered or not, de- 
pending upon the expansion or contraction of the abdomen. It is 
evident that a much more extensive study of these elements is neces- 
sary before any conclusions are warranted as to their taxonomic 
value or fines of progressive modification. In a number of genera 
the seventh or eighth tergite, or both, is exposed beyond the apex 
of the elytra and is either oblique, declivous, or vertical. The ninth 
and tenth tergites are absent as such or are modified into elements 
of the genital organs. 

ABDOMINAL SPIRACLES. 

According to Ntissfin (1911) and Fuchs (1912) the number of func- 
tional abdominal spiracles (stigmata) ranges from five to seven. The 
larvae and pupse, so far as studied by the writer, have eight. 

The spiracles are variable in structure and doubtless, upon further 
investigation, will furrnsh excellent taxonomic characters. The 
writer, however, is doubtful as to the phylogenetic significance of the 
variability in number. The genera given by Ntissfin (1911, pp. 2-5) 
and Fuchs (1912, pp. 13-14) which are represented by species having 



182 



THE SCOLYTID BEETLES. 



from five to eight spiracles fall into the following subfamilies as recog- 
nized by the writer : 

Table IV. — Abdominal spiracles in the subfamilies of Scolytoidea in which species were 
examined by Niisslin and Fucks. 



Subfamily. 



Eight 
spiracles.a 



Seven 
spiracles. 



Six 
spiracles. 



Five 
spiracles. 



Cryphalinae . . . 

Ipinee 

Corthylinse . . . 

Micracinse 

Crypturginse . . 
Pliloeotribiiiffi. 
Hylesininse . . . 
Scolytinse 



Genera. 



Genera. 



Genera. 



Total . 



a The eighth spiracle is rudimentary. 

In the genus Hylesinus (Fuchs, ibid., p. 13) there are from five to 
seven spiracles with the sixth and seventh rudimentary, while in 
Dendrodonus and Hylurgops the eighth is rudimentary. Doubtless 
if the number of abdominal spiracles were determined for all of the 
species, some good evidence would be furnished as to hnes of modi- 
fication. 

ABDOMINAL STEENITES. 

The abdominal sternites 1 and 2 are fused and concealed in the 
coxal cavity, 3 to 7 are exposed, and the eighth is covered by the 
seventh, while the ninth and tenth evidently are represented by the 
genital organs. (See Pt. I, pp. 25 and 38, and Technical Series 20, Pt. 
I, Pis. VII, VIII, and IX, as also the discussion of the reproductive 
organs m the present paper.) The exposed sternites are quite vari- 
able in the different genera and species, and certain characters are of 
value in designating minor groups of genera, but few, if any, are com- 
mon or pecuhar to a subfamily. 

The modifications range from the simple type fomid in the Crypha- 
linse to the steep, excavated, armed, and odd forms of the Scolytidae 
and PlatypocUdse. There is a wide variation, which is apparently 
of specific importance, in the eighth ventral segment. The palpi of 
the ninth ventral segment (Niisslin, Fuchs, and others) fomid in the 
females of Scolytus and Hylesinus, may after all represent the tenth 
tergite or sternite, one or both of which may be represented by the 
ovipositor as in certain Curcuhonoidea (Pissodes) , Cerambycoidea 
(Cyllene), and many other insects. But this is a matter requirmg 
more comprehensive investigation. 

STRIDULATING ACCESSORIES. 

The stridulating accessories have not been investigated by the 
writer in many genera but they appear to be confined largely to the 
seventh abdominal tergite and the inner sub apical area of the elytra 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 



183 



(Part I, figs. 23, 3.1, and 33) and to the anterior margin of the pro- 
notum and the posterior dorsal area of the head. 

Internal Anatomy. 

Considerable study has been made by Lindemann, Ntisslin, FuchS; 
and others, of certain elements of the internal anatomy, especially 
the proventriculus of the digestive system and the male and female 
reproductive organs. It is evident, however, from a review of the 




Fig. 98.— Proventric- 
ular plate of Ips 
emarginatus, inner 
aspect: a, Entire 
plate; 6, divided an- 
terior plate; c, poste- 
rior or masticatory 
plate; d, median 
lonfritudinal sutm-e; 
e, sutural teeth; /, 
median transverse 
suture; g, transverse 
or apical teeth of an- 
terior plate; h, lat- 
eral margin or mar- 
ginal suture; i, lat- 
eral or masticatory 
teeth; ia, closing 
bristles or teeth; ./, 
femora of the mas- 
ticatory teeth; k, 
femoral teeth or 
ridge; I, mastica- 
tory brush; m, mar- 
ginal bristles or 
fringe. (Original.) 




Fig. 99. — Proventricular 
plate of Scolytus scolytus: 
b, Divided anterior plate; 
d, median suture; /(.lateral 
margin; gl, apical lami- 
nate teeth of posterior 
plate. (Original.) 




results, that as a sufficient basis for 
general or specific conclusions a far 
more comprehensive study is necessary. 



DIGESTIVE SYSTEM. 



Fig. 100. — Masti- 
catory plate of 
Crossotarsus le- 
contei: c, Masti- 
catory plate; la, 
apex "and lateral 
serrations o f 
masticatory 
teeth. This fig- 
ure should be 
viewed in a re- 
versed position 
in order to rec- 
ognize the ele- 
vated character 
of the median 
longitudinal 
area. (Original.) 



The general type of the digestive sys 
tem is shown in Part I, figures 35 and 36, for the 
adult and figure 43 for the larva. Lindemann (1876, pp. 148-169), 
Ntisslin (1912, pp. 85-87, figs. 135-143), and Sedlaczek (1902, pp. 
241-263, figs. 1-20) show that there is a wide variation in some of 
the elements and especially in the proventriculus and the median 
and posterior section of the midintestine, but to what extent these 
variations are of value in taxonomy has not been determined. 

Proventriculus . 

The proventriculus (figs. 98-100; Part I, figs. 35-38) is of special 
interest, and the work of Lindemann, Fuchs, Niisslin, and others 
has contributed much valuable information on the structural ele- 
ments. 



184 



THE SCOLYTTD BEETLES. 



There is a wide range of variation in the proventricular plates and 
their armatures, such as the various divided and undivided forms 
with and without the "brush" of fine masticatory teeth. These 
variable forms appear to be of considerable taxonomic importance 
when correlated with other internal and external characters. 

The classification proposed by Lindemann and Niisslin, as based 
on the structural characters of the proventricular plates, with a few 
plainly evident exceptions, correlates in a striking manner with the 
external characters adopted by the writer for the major and minor 
groups of genera. Wlien the principle of parallel modification is 
considered, the genera which seem to be out of place in the classifi- 




-20 



'^ 



Fig. 101. — Dendroctonus valens: Mem- 
branous and nonchitinous elements of 
the male reproductive organs. See ter- 
minology under Divisions 2, 3, 4, pp. 
192-193. (Original.) 




Fig. 102. — Xyleborus saxeseni: Chitinous ele- 
ments of male reproductive organs. See ter- 
minology, Divisionl, pp. 192-193. (Original.) 



cations based on proventricular characters alone mil, when correlated 
with a combination of characters, fall into their natural positions. 

REPRODUCTIVE ORGANS. 

The Reproductive Organs op the Male. 

The reproductive organs of the male (figs. 101-110) have been 
studied and figured by several investigators as follows: 

Table V. — Summary of previous studies of the male reproductive organs by various authors. 



Author. 


Year of 
publi- 
cation. 


Number 
of species 
figured. 


Number 
of genera 
repre- 
sented. 




1875 
1896 
1911 
1912 
1912 
1912 


32 
3 

20 

24 

30 

2 


15 


Verhoefl . . 


3 




4 




22 




27 




2 







PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 



185 



In addition to the foregoing, the writer, during the years from 1892 
to 1912, has studied the male reproductive organs of 68 species, repre- 
senting 43 genera. The total species studied by all authors, with- 
out dupHcation, is 147, 
representmg 57genera. /<- -./- ■ -/—-/?<? 

The reproductive or- 
gans of the male repre- 
sent four primary divi- 
sions or elements :( 1) The 
posterior chitinized di- 
vision (fig. 102); (2) the 
posterior membranous 
division (fig. 101), (3) the 
median division, and (4) 
the anterior division. 
(See terminology, pp. 
192-194.) 



POSTERIOR ELEMENTS. 

The posterior chiti- 
nized elements ( divi- 
sion 1) consist of four 
primary sections, (a) the 
body, (5) the end plates, 
(c) the tegmen, and {d) 
the spicule. In addition 
to these more constant 
elements there are (&7) 
the seminal valve and 
{1)2) the seminal rod, 
both of which appear to 
be more intimately asso- 
ciated with element & 
than with element a; also, 
there are (e) the connect- 
ing membrane and (/) 
the muscles. From a 
somewhat comprehen- 
sive study of the chiti- 
nized elements in the 
scolytoid beetles in com- 
parison with those in 
other insects it seems plain that, so far as these beetles are concerned, 
the chitinized parts of the male reproductive organs represent ele- 
ments of the ninth and tenth abdominal segments, either as direct 
modifications of primitive sclerites or as independent developments 




Fig. 103. — lips emarginatus: Cliitinous elements of male repro- 
ductive organs. /, ventral aspect; //, dorsal aspect. See 
terminology, Division 1, pp. 192-193. (Original.) 



186 



THE SCOLYTID BEETLES. 




Fig. 104.— PityophtTiorussp.: Chitin- 
ous elements of male reproductive 
organs. See terminology, Divi- 
sion 1, pp. 192-193. (Original.) 



&2 




from the same fundamiental source as that from which the various 
segmental lobes, sclerites, and ajopendages have developed. It 
would appear best to refer to the chitinous element of the copulatory 

apparatus as repre- ^^,._ 

senting certain seg- 
mental elements 
rather than to as- 
sume that they have 
been derived through 
modification from 
sclerites or append- 
ages which had ex- 
isted as such in a 
primitive ancestor. 

It is quite evident 
that element a rep- 
resents the tenth sternite, h the tenth tergite, c 
the ninth tergite, and d the ninth sternite, while hi 
and 1)2 appear to represent elements of either 
the tenth tergite or of both the tenth tergite and 
tenth sternite. It might also be weU to con- 
sider in future investigations the possibility of 
their representing the tenth pleurites or even 

an additional eleventh segment. The 
wide range in the variation of these ele- 
ments, the absence of some of them in 
certain species, and the joining or fusion 
of two or more in other species render 
it exceedingly difficult properly to in- 
terpret the primary and secondary ele- 
ments, especialty in the more complex 
and in the apparently simple forms. 

The body, or element a, is present in 
all rpecies. It is more or less distinct 
from the other elements and is nearly 
always suggestive of a modified sternite; 
the femora (al ) appear to represent the 
produced posterior angles, or apodemes, 
of a typical abdominal sclerite, and the 
lateral folds (aS) appear in some cases to 
represent the hypopleurites, especially 
in Ips, where they resemble end plates 
and were so identified by Lindemann. The function of element a is that 
of a sheath or tube for the ejaculatory sack(^a) and for the seminal valve 



Fig. lOo.—Pityophtho- 
rus bellus: End 
plates and seminal 
rod of male repro- 
ductive organs, lat- 
eral aspect. Note 
contrast between 
this and fig. 104. 
See terminology, Di- 
vision 1, pp. 192- 
193. (Original.) 




Fig. 106. — Micracis suturalis: Posterior 
elements of male reproductive organs, 
dorsal aspect. See terminology, Di- 
vision 1, pp. 192-193. (Original.) 



PEELIMIIsrAEY CLASSIFICATION OF SCOLYTOIDEA. 



187 




03 (E 
■3 03 



IS 



c3 O 






03 c3 



Pi-- 



.. "^ - 

828.3 

§■& - 

t, CM +J 

2 - o 

•00 CQ 



188 THE SCOLYTID BEETLES. 

(hi ) or the seminal rod (b2), or both, as the case may be. The end plates 
(b) are commonly present. They arc sometimes fused with a3 but 
are more often separated or more directly connected with hi or 1)2."' 
The end plates proper appear to function as accessories to a in forming 
the sheath or outer tube; hi may function as a valve to close the 
seminal duct while muscular or blood pressure is brought to bear on 
the ejaculatory sack to force the seminal fluid into the copulatory 
pouch, or they may function, as indicated by Lindemann, as a furrow 
or troughlike support for the posterior end of the seminal duct. The 
end plates are subject to great modification, from simple chitinous 
pieces, as in Pissodes, to the more complex structure with many parts, 
as in Dendrodonus (fig. 107), and especially Hylesinus, or into a long 
slender troughlike rod, as in Ips (fig. 103), a stouter rod with apical 
dilation and a long flagellum, as in Xyleborus, or a brushlike form, as 
in Micracis (fig. 106), and Xylocleptes. The function of the slender 
rod or apical filament is not known, but it may serve to conduct the 
seminal fluid directly into the spermathecal duct. 

When we consider the enormous range of possibilities in the modifi- 
cation of tergal and pleural sclerites, as manifested in the tergum and 
pleura of the metathoracic segment, we can readily understand that 
the most complicated and complex copulatory apparatus yet found 
in insects is comparatively simple and that the possibilities of further 
modification have not been exhausted. 

The tegmen, or ring (c), is generally present but may be obscure or 
absent. There is a wide range of variation, from a simple and ven- 
tral plate (Scolytus rugulosus, fig. 108) or fork {S. quadrispinosus, 
fig. 109, and Crossotarsus, fig. 110) to a continuous simple ring or band 
(fig. 102), and from a plain or forked dorsal piece to a forked or plain 
ventral piece. In nearly all cases it functions as an apodeme for the 
attachment of the primary and accessory muscles for the posterior 
extension and movements of elements a and h. The sections of the 
various forms of the tegmen may be referred to as (cl) the posterior 
section, {c2) the median section, and (c3) the apodemal process. 
Section cl may be either dorsal or ventral; in whichever case section 
c2 will occupy the opposite position. 

The spicule or rod (element d) is usually present in a more or less 
distinct form which may vary from a curved forked rod to a simple 
rod, and in its various modifications and functions may occupy a 
dorsal, sublateral, lateral, or subventral position. ■ The sections of 
the various forms may be referred to as the anterior section (dl) and 
the posterior section (d2). It functions as an apodeme for the attach- 

aVerhoeff (1896) and Ntissliii (1912) considered tlie end plates as belonging to the 
body. 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 



189 



ment of the primary and. accessory muscles for the retractile move- 
ments of the tube {a and h). 

The muscles (e) are more or less complicated and variable in size, 
number, and attachments. In some cases, as in DeTidroctonus, the 




Fig. 108. — Scolytus rugulosus: Posterior elements of 
male reproductive organs. /, lateral aspect of 
body; //, lateral aspect of body and accessories 
in situ; ///, body accessories separated. See ter- 
minology, Division 1, pp. 192-193. (Original.) 



Fig. 109.— Scolytus quadris- 
pinosus: Posterior ele- 
ments of male reproduc- 
tive organs, lateral aspect. 
Seetermiaology , Division 
l,pp. 192-193. (Original.) 



extensor and retractile muscles are prominent, the former (el) at- 
tached posteriorly to the anterior edge of element c, while the sup- 
plementary muscles (eS) are attached anteriorly to the posterior edge 
of c and posteriorly to the inner surface of the eighth sternite, as in 
Dendroctonus, or to the anterior section of element c and the eighth 
59026°— 15 3 



190 THE SCOLYTID BEETLES. 

tergite, as in Grossotarsus. In Bendroctonus the retractile muscles 
(e£) are attached to the anterior end of the spicule (d) and to the 
basal angles of the body (a), while in Grossotarsus they are attached 
to e^ and to the sides of a ventral groove in a. 

The connecting membrane (f) is very difficult to locate and follow 
to its primary connections, but if it could be accurately traced it 
would doubtless furnish good evidence as to the proper assignment of 
the chitinous elements to their respective tergal and sternal origins, 
unless, as is the case with muscles, the attachments are changed or 
even reversed to harmonize and economize the requirements of func- 
tion. The connecting membrane between the posterior section of 
element d and the eighth abdominal sternite in Dendroctonus seems 
to furnish quite conclusive evidence that the spicule represents the 
ninth sternite. 

ANTERIOR AND MEDIAN ELEMENTS. 

The anterior (4) and median elements (3) of the male reproduc- 
tive organs have received special attention by Niisslin, who bases a 
classification on the length of the ductus ejaculatorius {2h) and the 
character of the testes {Jjh) , seminal vesicles {3) , vas deferens {^a) and 
mucous glands {2c). 

VARIATIONS AND COMBINATIONS OF THE ELEMENTS. 

Various combinations of the elements of the copulatory apparatus 
are found in the species of allied genera. Quite a wide range of varia- 
tion is also found in the character of the elements in the species of 
the same genus. Even species which in all other respects appear to 
be closely allied have very different characters, either in the form 
of one or more elements or in different combinations of the elements. 

In Division I of the family Ipidse the absence of the seminal valve 
and the presence of the seminal rod appear to predominate. The 
valve, as a definite part, is absent in 21 genera and 52 species and 
present in 10 genera and 16 species. The rod is present in 25 genera 
and 55 species and absent in 8 genera and 14 species. Both the rod 
and valve appear to be absent in 6 genera and 10 species, representing 
4 subfamilies, and both are present in 10 genera and 15 species, repre- 
senting the same four subfamilies, and especially in Corthylinse. The 
striking feature of the Subfamily Ipinse is the apparent " separation 
of the femora from the body, especially in Ips and the closely allied 
genera. In Pityo'plitliorus (figs. 104, 105) they are found to be fused 
with the body in some species and separated in others; within still 
other species there is a supplementary band {aS) connecting them 
at or near the base. 

« The writer has found that the femora are connected to the body by ligaments. 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 



191 



In Division II of the Ipidse the presence of the seminal valve and 
absence of the seminal rod predominate. The valve is present in 20 
genera and 44 species and the rod is absent in 20 genera and 43 spe- 







cies. The valve is absent in 2 genera and 4 species and the rod is 
present in 6 genera and 12 species. The combination of rod and 
valve is found in 4 genera and 10 species. 



192 THE SCOLYTID BEETLES. 

In the family Scolytidse there appears to be a wide range of varia- 
tion. In ErineopMlus schwarzi Hopk., of the subfamily Hexacol- 
inse, the rod is absent, the valve present, and the femora are long 
and slender. In some species of the genus Scolytus the body is 
greatly modified, somewhat resembling a seminal rod. The valve 
is absent and the femora are rudimentary in 8. muticus Say and 
S. quadrispinosus Say (fig. 109). In S. rugulosus Katz., however, 
(fig. 108) the femora are long, the valve is represented, and there are 
some additional parts; in fact, all of the elements are radically differ- 
ent from those of the other two species. 

In one genus and one species of Scolytoplatypodidae the rod is 
absent and the femcra are large and very broad, differing in this 
respect from anything yet observed in the entire superfamily. 

In two genera and five species of the subfamily Platypodinse the 
body is long and slender, but without the spicule, end plates, seminal 
valve, or rod, and the femora are represented by short hooks at the 
basal angles of the body. The tegmen is present in the form of a fork. 

A study of the available data relating to the primary and secondary 
elements of the male organs of reproduction shows that within the 
families, subfamilies, and genera there is a very wide range of varia- 
tion and that the same or similar elements individually, or in various 
combinations, are often paralleled in sDecies of widely separated 
genera and subfamilies, so that their principal taxonomic value 
appears to be restricted to the separation of species and minor divi- 
sions of the genus. 

It appears that if there is any line of progressive modification 
within the major and minor groups, it is from a simple form without 
seminal valve, seminal rod, or end plates, as in some of the Crypha- 
linse and in the Platypodinse, to the most complex forms with or 
without the valve and with or without the rod, the rod reaching its 
highest development in Xylehorus, Dryocoetes, Lymantor, Ips, and 
allied genera; while the valve without the rod reaches its highest 
development in Hylesinus and allied genera in the Hylesininse. 

Terminology of the PvEproductive Organs. 

In the following list it is intended that the numbers and letters 
should serve to designate the elements of the reproductive organs 
rather than names, because the names proposed by different authors, 
including the writer, do not agree in aU cases in designation or 
interpretation. 

Male Reproductive Organs. 

(Figs. 101-110.) 

Division 1. Posterior chitinous division. 

Division 2. Posterior membranous diAasion. 

Division 3. Median division. 

Division 4. Anterior division. 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 193 

Division 1. 

a. Body. 

al. Body apodenies (femora). 

alo-. Apodemal ligament. 

a2. Lateral folds. 

afa. Lateral plates (accessory pieces, Niisslin). 

aS. Supplementary body apodemes or transverse band. 

a4. Apical orifice or ejaciilatory canal. 

a5. Sensory area or pores. 
h. End plates. 

hi. Seminal valve and accessories. 

b2. Seminal rod. 

hS. Apical flagellum of M. 

b4. Apical brush of b2. 

h5. Sensory claspers (fig. 108, h5). 

b6. Apical lobe. 

c. Tegmen. 

cl. Posterior or dorsal section. 
c2. Anterior or ventral section. 
c3. Apodemal process. 

d. Spicule. 

dl. Anterior section. 

d2. Posterior section. 

c?f«. Lateral apodeme or minor prong. 

d2t>. Lateral barb. 

e. Muscles. 

el. Extensors. 

e2. Retractors. 

eS. Supplementary. 
/. Connecting membrane. 
g. Ligament. 

Division 2. 

2a. Ejaculatory sac or prasputial sac. 
2al. Chitinous base, or tube. 
2b. Seminal duct. 
2c. Mucous glands. 



3a. Seminal vesicle. 



4a. Vas deferens. 
4b. Testes. 



Division 3. 
Division 4. 



Female Reproductive Organs. 
(Fig. 111.) 



Division 1. Posterior division. 
Division 2. Anterior division. 



Division 1. 



la. Vagina. 

lb. Biirsa copulatrix. 

ib^. Accessory sac. 

Ic. Unpaired oviduct. 

Icf^. Apex and apical orifice of oviduct. 



194 



THE SCOLYTID BEETLES. 



Id. Spermatheca. 
le. Spermathecal gland. 
If. Seminal duct. 
Ig. Cement glands. 
Ih. Chitinous plates. 



2a. Paired o\T.ducts. 
2h. Ovaries. 



Division 2. 



Classifications Based on the Reproductive Organs. 

In the classifications of Niisslin (1911), Fuchs (1911), and others, as 
based on the male reproductive organs, we find, as we do in the classi- 
fications based on the elements 
of any single organ, that genera 
and groups which are plainly 
not closely aUied are brought 
together and those which by 
the majority of external and in- 
ternal characters are closely 
alhed have been placed in widely 
separated divisions or subdi^d- 
sions. When, however, the facts 
of parallel modification are 
taken into consideration and 
the principal elements are cor- 
related with those of other or- 
gans, the results are quite differ- 
ent and the true taxonomic 
value of the elements is recog- 
nized. It is e^ddent that a 
study must be made of the male organs of reproduction in a much 
larger number of species of all available genera before the true taxo- 
nomic value of any of their elements can be determined and correlated. 
Ntissfin (1912), who has given the subject of the female reproductive 
organs (Fig. Ill) of Scolytidse late consideration, calls attention to the 
taxonomic importance of the female genital organs in separating the 
Adephaga and Polyphaga of the order Coleoptera and in distinguish- 
ing the suborder Rhynchophora, which he claims is peculiar in having 
only two pairs of ovaria. 

The absence of a true ovipositor is apparently common to aU scoly- 
toid beetles, although in some species there are rudimentary parts 
which in other Rhynchophora and Coleoptera belong to the ovipositor, 
especially the genital palpi, which have been found only in Scolytus and 
Hylesinus. The presence of paired or single cement glands appears 
to be an important and more or less pecuHar element in the Scoly- 
toidea, although it is said to be absent in Scolytus and Emoporus, as 




Fig. III.— Dendroctonus valens: Female reproductive 
organs. See terminology, Division 1, pp. 193-194. 
(Original.) 



PRELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 195 

in other Rhynchophora. The bursa copulatrix, according to Nusshn, 
is present in Scolytus, Hylesinus, Polygraphus, Crypturgus, and Hypo- 
horus, much less evident in Carphohorus, Pityophthorus, Dryocoetes, 
TapJirorycJiUS, and Lymantor, and obscure or absent in CrypTialus, 
Xyloterus, Xylocleptes, Thamnurgus, Ips, and Pityogenes. 

Conclusions as to whether or not the presence or absence of a given 
element is primitive in the Scolytoidea, as based on morphological 
interpretations, are becoming much less reliable than formerly because 
of the frequency of parallel origin or disappearance of adaptive ele- 
ments. 

Ntisshn's (1911, pp. 333-338) classification as based on the female 
organs of reproduction relates primarily to the presence or absence of 
the cement gland and to its varying forms ; secondarily, to the presence 
and character, or the absence, of the bursa copulatrix, and the char- 
acter of the spermatheca and its seminal duct. 

When the characters of the female reproductive organs as given by 
Niisslin are correlated with the external characters on which the 
writer's prehminary classification is based, Scolytus falls into the sub- 
family Scolytinse; Hypoborus and Thamnurgus into the Micracinse; 
Crypturgus and Carphohorus into the Crypturginse ; Polygraphus into 
the Phloeotribinse ; Pityophthorus, Pityogenes, and Ips into the Ipinse ; 
and Xyloterus into the CorthyHnge; while Ernoporus, Cryphalus, 
Taphrorychus, Lymantor, Dryoccetes, Xyleborus, and Xylocleptes fall 
into the Cryphalinse. 

With a more comprehensive study of the female organs in num- 
bers of species representing all the genera it will evidently be found 
that there are some excellent taxonomic characters in the primary 
elements and in their lines of progressive modification, which in com- 
bination with other internal and external characters will be of special 
value in defining groups of alUed genera and in indicating relative 
positions of the groups in the classification. 

Secondary Sexual Characters. 

There is a wide range in the types and position of the secondary or 
external sexual characters, such as difference in the size of the body, 
as in Hypothenemus, Stephanoderes, Coccotrypes, Xyleborus, and allied 
genera, and the radical and contrasting differences in the structure, 
vestiture, and sculpture of various external parts and areas. The 
front of the head may be convex and glabrous in one sex and in the 
opposite sex it may be flat to deeply concave, smooth, and shining, 
punctured, and with dense and long pubescence or the margins fringed 
with long hairs. The armature of the decHvity and the character of 
the sutural impressions or broad excavation often vary to a remark- 
able extent in the two sexes. The scape of the antenna may be stouter 
to dilated and fringed with long hairs or not fringed. The funicle, 
club, mouthparts, tibia, tarsus, abdominal tergites, and stemites, in 



196 THE SCOLYTID BEETLES. 

fact almost every important element of the body, may be the bearer 
of characters for distinguishing one or the other sex. 

It would appear that as a rule there is a certain degree of con- 
stancy in the location and general appearance of a male or female 
character within the Hmits of a genus, but there are some remarkable 
differences, and even reversals, even in the major and minor divisions 
of a genus. In Dendroctonus, for example (Part I of this bulletin, 
p. 73), the females of subdivision A are distinguished by the presence 
of a transverse ridge on the anterior area of the pronotum, while in 
subdivisions B, 0, and D this character is not present. In sub- 
division B the elytral decHvity is more rugose in the female, in sub- 
division C this is reversed, and in subdivision D there is no difference 
in this respect. In some genera the pubescent or concave front is a 
female character, while in other genera it is a male character. Other 
reversals may not only occur in different, widely separated genera but 
in the same genus. Therefore it is not safe to conclude that because 
certain characters designate the female in one species this will hold 
true for the other species of the same genus or for alhed genera. This 
can only be definitely settled by dissection, which can be done without 
seriously mutilating the specimen if the abdomen is carefully removed 
from the body and the sternites are remounted on a card point after 
the examination is made. The presence of the chitinized sperma- 
theca in the female and the presence of the chitinized elements of the 
posterior section of the male organs are sufficient to settle the point, 
even in old dried specimens. 

The lines of progressive variation or modification in secondary 
sexual characters appear to range from absent and obscure to com- 
mon and prominent. 

THE PUP^. 

Comparatively very little study has been made of the pupse of even 
our common species of Scolytoidea, and until a comprehensive study 
has been made it is scarcely necessary to mention the characters in 
connection with general taxonomy. The description and figures of 
the pupa of Dendroctonus valens (Part I, pp. 53-57, figures 37-38; 
synopsis, pp. 73-74, and descriptions, pp. 81-152) will serve as an 
example of the general type, while the figures and terminology will 
serve as a guide to future study. 

The writer has examined the pupae of quite a large number of 
North American species and finds that there is quite a wide range 
of variation, but no attempt has been made to analyze the taxonomic 
characters except in the genus Dendroctonus, in which the form of 
the head and the character of the frontal, tergal, lateral, pleural, 
caudal, and femoral spines serve as important characters for identifi- 
cation and classification and, when correlated with the adult charac- 
ters, give the same or similar taxonomic result. 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 197 

THE LARV^. 

The structural and morphological elements of the larvae of Den- 
droctonus are shown in Part I, figures 39-43, and Plate VIII, figures 
1-23/, and these, with the terminology and descriptions, wiU serve 
as a guide to a greatly needed further study of the larval stage 
before we can have a basis for conclusions as to their importance in 
taxonomy. The writer has examined the larv^ of quite a large 
number of species, but no detailed study has been made except in 
Dendroctonus and of the labrum of a number of species in other genera 
(Hopkins, 1905, Plate I). This has been sufficient, however, to 
indicate the wide range of variation in some of the elements and the 
great importance of a more comprehensive knowledge of the subject. 
In Dendroctonus the important characters are found in the eighth and 
ninth abdominal tergites, and the front of the head. An example of 
progressive modification is found in the sculpture and armature of 
the eighth and ninth abdominal tergites from those without dorsal 
plates in Division I and Division II, section a^, to the unarmed plates 
of section a* and to the armed plates of subdivision D, which corre- 
lates so nicely with progressively modified characters in the adults and 
in the galleries. The larvas of the species of Platypus and Crossotarsus 
examined by the writer show radical differences in form and in some 
of the anatomical elements, as, for example, the labrum (Hopkins, 
1905). 

THE EGGS. 

While the eggs of many species have been observed by the writer, 
they have not been studied in detail. They appear to conform in 
general to an oblong, oval, or nearly globular shape, and are pearly 
white and smooth, with few elements of vestiture or sculpture to 
serve as taxonomic characters. However, this is a subject worthy 
of detailed study. The size of the egg in comparison with the size 
of the abdomen varies enormously in different species. In a species 
of CarjpJiohorus a fully developed single egg was found to be so large 
as to occupy almost the entire abdonfinal cavity. 

THE EMBRYO. 

The embryology of the scolytoid beetles is another subject which 
has not received much attention. While the writer feels that there 
is need of detailed study of the embryo to determine any additional 
facts which may be of value, he is inclined to the belief that more 
attention should be given to a comparative study and correlation of 
characters of the postembryonic stages (young to matured larvae) 
of a wide range of species in the order Coleoptera, in order that we 
may know something more of the fundamental facts and be better 
able to interpret their real significance. 



198 THE SCOLYTID BEETLES. 

PHYSIOLOGICAL CHARACTERISTICS. 

As pointed out in Part I (p. 64), the physiological characteristics 
are of special taxonomic importance when correlated with morpho- 
logical characters. The food, social, and sexual habits, character 
of the brood galleries, choice of host plants, and distribution of 
genera and species are all more or less rich in facts of taxonomic 
importance. 

GENERAL HABITS. 

The scolytoid beetles are distinguished from nearly all other 
Rhynchophora by their habit of excavating characteristic egg gal- 
leries in the living or dead plant tissue. The few notable exceptions 
are found in the genus Stenoscelis, of the CalandridsB, the adults of 
which excavate a primary egg burrow, but as a rule this habit within 
the suborder is peculiar to the Scolytoidea. The egg galleries of 
Scolytoidea are excavated in the bark or wood of trees and shrubs, the 
roots, stems, and leaves of herbaceous plants, the fruits or seeds of 
palms and other plants, young pine cones, the wood of barrels or 
casks containing water or spirituous liquors, etc. Some of the species 
excavate their galleries m decaying bark or wood or even in the 
fruiting bodies of fungi, while others confine their work to the bark 
or wood of weakened, dying, or recently dead plants, and still others 
prefer to enter the living and sound tissues. 

The food of the adults and larvse consists of the sugars, starches, 
and other nutritive elements of their host plants, or of fungi which 
grow in their brood galleries. 

CLASSIFICATION ACCOEDING TO HABITS. 

Any classification of the families or subfamilies based on food 
habits alone would not indicate a natural arrangement, as is plainly 
indicated by the parallel habits of groups of species in widely sepa- 
rated families, subfamilies, and genera. It is true that there are 
several well marked classes according to habits, such as bark beetles, 
twig beetles, seed beetles, cone beetles, and ambrosia beetles. It is 
evident, however, that food habits, like many other characteristics 
and structural characters, have evolved along parallel lines in alUed 
as well as in widely separated groups. There are many examples 
illustrating this principle. The genera Xyleborus, Corthylus, Scoly- 
to'platypus, and Platypus are, according to fundamental morpho- 
logical elements as well as groups of correlated characters, so widely 
separated that they each represent a different family or subfamily; 
yet the habit of excavating their galleries in wood and feeding on 
ambrosial fungi is common to them all. Between some of these genera 
there is also a more or less constant resemblance in certain morpho- 
logical characters, especially in the hairs and slender teeth of the 



PRELIMINARY CLASSIFICATIOISr OF SCOLYTOIDEA. 199 

lacinia of the maxilla, but this is evidently due to parallel adaptation 
to similar uses and not to common origin or phylogenetic descent 
from a common ambrosia-feeding ancestor. 

Food Habits of the Adults. 

As a rule the adults obtain their food from the substance in which 
they excavate their egg galleries or from the fungi growing on the 
walls of the galleries, but there are numerous examples of special 
food habit such as that found in Scolytus, Pteleohius, Phlmosinus, 
and Tomicus (see p. 220), which excavate food burrows in the living 
twigs of their host trees. 

Food Habits of the Larv^. 

There is a wide range of variation in the food habits of larvae of dif- 
ferent species, especially in the character of their food burrows or 
larval mines. Each species of a group of closely allied species may 
have similar habits, but, as shown in the genus Dendrodonus, there 
may also be a wide range of variation and some striking examples of 
progressive modification m this habit within a genus m which there 
is a restricted range in structural characters in the adults. In the 
genus Dendrodonus there is a tendency throughout for the larval 
mines to occur in groups of increasing numbers from the simple, isolated 
mine of Dendrodonus hrevicomis to closely placed groups in Dendroc- 
tonus simplex and D. piceaperda and to the large social chamber of 
micans, valens, and terebrans. (See figs. 73, 75, 79, 88, and 91, of 
Part I.) Thus the stage in the modification of the larval mine of a 
given species may mdicate, m connection with stages in the modifica- 
tion of structural characters, the natural position of the species. 

In the ambrosia beetles the larvae of some species and groups of 
genera, as Xyleborus, StepTianoderes, and Orossotarsus, live in the 
primary galleries in direct association with the eggs, larvae,- pupae, 
young adults, and parent adults, while in the subfamily Corthylinae, 
the genus Scolytoplatypus, and at least some of the species of Platypus, 
the larvae occupy separate chambers in the sides of the gallery, these 
chambers not extending beyond a size sufficient for the accommoda- 
tion of the body. 

Pupal Habits. 

Considerable variation exists in the habits of the pupae and in the 
cells occupied by them in transformation from larvae to adults. In 
perhaps the majority of species the transformation takes place at the 
end of the food burrow with or without a definite cell. In some ambro- 
sia beetles the pupation takes place in the social galleries occuped by 
different stages of the brood, in others it is in the lateral larval cell, 
and in Stephanoderes the transformation takes place in closely joined 



200 THE SCOLYTID BEETLES. 

cells at or toward the inner end of the social gallery, the walls and par- 
titions of these cells consisting of an ambrosia-like substance mixed 
with fine borings. 

Flight Habits. 

Further observations should be made on the flight habits of these 
beetles, but from what we know of a few species it would appear 
that in the same species flight may be either individual or collec- 
tive. Ip. one example noted by the writer (Hopkins, 1899a, pp. 346- 
348), a large number of species, together with some of their associates, 
scavengers, and predaceous enemies, were found in one great swarm. 
The periods of flight vary with the number of generations in a sea- 
son. Thus species with a single generation have but one definite 
period of flight, while those with more than one generation have two 
or more periods, or, when the generations overlap, there may be a 
continuous period of flight throughout the season. 

Social Habits. 

In the social habits we find some features of special interest both in 
their relation to taxonomy and to parallel lines of modification. In 
the relation of the sexes there is a wide range of variation from simple 
or unorganized and intensive polygamy to specialized or organized 
polygamy, and a gradual reduction in the proportion of the number of 
females, from 1 male and many females to 1 male and 2 females, and 
finally to specialized monogamy. 

In Hypotlienemus, StepJianoderes, Xyleborus, and allied genera the 
males are much smaller than the females and very rare. In certain 
species of Xyleborus as many as 60 females to 1 male have been found 
in one brood gallery, and the proportion appears to be even greater 
in Hypothenemus. In these groups there is no system in the relation 
of the sexes or in eggs of the brood galleries of the females, and all live 
together in the same galleries. In the other groups of genera of the 
subfamily Cryphalinse where there is no difference in the size of the 
sexes there is more evidence of separate egg galleries for the different 
females of the social group, and the galleries begin to take on more 
definite and characteristic forms or patterns. In the Ipidse the tend- 
ency toward a specialized polygamous relation of the sexes reaches 
its highest development in Pityogenes, while in Ips there is a tendency 
toward fewer females, the numbers of which in some cases are limited 
to 2 or 3 to the gallery. In Corthylinse the sexes are more equaUy 
divided, while m Crypturginas, Phloeotribinse, and Hylesininse there 
is a tendency toward 2 females and 1 male or to 1 of each. The last 
seems to prevail to a greater extent in the subfamily Scolytinae, or at 
least m the genus Scohjtus, than in other subfamilies. 



peeliminaky classification of scolytoidea, 201 

Galleries. 

There is a wide variation in the types or forms of the egg and brood 
galleries within the families, subfamilies, and the major and minor 
groups, and in some cases within the genus. Each species or group of 
allied species of a genus or group of allied genera is usually character- 
ized by some peculiar form or feature which in many cases is sufhcent 
in itself to indicate the species, genus, or group to which it belongs; 
therefore the galleries are of special taxonomic importance in indicat- 
ing the natural position and grouping of the species and genera. 

It has been supposed that a peculiar type of gallery was due to 
the character of the plant tissue in which it was excavated or that it 
had some relation to the species of plant. It is found, however, that 
the type of the gallery is the same, or similar, for the same species, 
regardless of the character of the substance or the species of plant in 
which it is excavated. In fact, the same species of spruce or pine, 
and the bark on the trunk or branches, or the wood of the same part 
of the tree, may have as many radically different types of galleries 
as there are different species of beetles to make them. 

It has also been supposed that the type of a gallery was due to the 
peculiar structure of the beetles, such as the retuse or concave and 
armed elytral declivity in I'ps, ascending or excavated abdominal 
sternites of Scolytus, etc., but it is found that certain species with the 
same or similar structures make very different types of galleries, 
while certain other species with very different structures make similar 
galleries. 

The fact that there is quite a definite relation between the type 
of the gallery and the systematic position of the species, genus, and 
group would indicate that the evolution of the gallery has been from 
the simple to the complex and that it has progressed with the evolu- 
tion of the beetles that make and inhabit it in a somewhat similar 
manner to that of the dommant tendency iji the evolution of human 
dwelling places from the simple cave to the modern palace. In other 
words, the simple and complex galleries represent evolution within 
the maximum and minimum limits of an mstinctive or dominant 
tendency common to all of the individuals of the superfamily Scoly- 
toidea and are expressed by each species of a genus in the varying 
degrees of simplicity or perfection accordmg to the varying stages 
in the evolution of the species. 

The fact that the same or a similar type of gallery is made by 
species of widely separated genera and subfamilies mdicates that 
we should not look for an explanation of the origin and evolution of 
types of galleries in the phylogeny of the species, but that a thorough 
consideration should be given to the explanation to be found in paral- 



202 THE SCOLYTID BEETLES. 

lei evolution due to a common tendency which may lead to the same 
or similar results during the same or similar stage in the evolution 
of the species. 

While the varying types of the egg and brood galleries furnish 
some very important taxonomic evidence, any attempt to classify 
the species of a family or subfamily accordmg to such characters 
alone would give the same heterogeneous results as are found in the 
different classifications based on the elements of a single external or 
internal part or organ of the body. On the other hand, if the galleries 
of the species of the major and minor divisions of a subfamily are 
studied separately, it will be found that the character of the gallery 
and the species of the host tree wilL serve as most important guides 
to the natural position of a species or group of allied species. 

TERMINOLOGY OF THE GALLERIES. 

Following is a revised list with defuiitions of the terms used to 
designate the different elements of the scolytoid gallery. 

Egg gallery. — The egg gallery is the burrow excavated by the parent 
beetles preliminary to depositing the eggs in niches along the sides 
or loosely in the gallery itself. 

Social gallery. — The social gallery is one in which all stages of the 
broods from the eggs to the matured individuals and the parents 
live, as in Xyleborus, certain species of Platypus, Crossotarsus, etc. 

Social cJiamber or hrood cliamber. — The social chamber, as in Xyle- 
horus saxeseni, is a dilate^ portion of the tubelihe gallery to accomo- 
date all stages of the brood. 

Death diamber {catacomb or garbage cliamber). — The death cham- 
ber (Hubbard, 1897; also Hopkins, 1898) is a section in the social 
chamber in which the dead individuals of the colony or the guests and 
enemies as well as other refuse matter are deposited and separated 
from the main chamber by a wall of the ambrosia fungus mixed with 
boring dust. 

Entrance burrow. — The entrance burrow is made by one or the 
other sex as a preliminary to the excavation of the egg gallery, and 
may connect in a direct manner or laterally with the base of the gal- 
lery in the case of single galleries, or with the middle in the case of 
double galleries. 

Nuptial (lateral or central) chamber. — The nuptial chamber is 
excavated by one or the other sex (probably in most cases by the 
male) at the base of the entrance burrow and the mouth of a single 
or many egg galleries. In Pityophthorus, Ips, etc., this chamber is 
short and broad, oblong, or rectangular. In some species of Scolytus, 
Phloeophthorus , Hylesinus, etc., the lateral entrance appears to serve 
the same purpose as the nuptial chamber, while in Phloeosinus the 
chamber is at the base of the egg galleries and entrance burrow and 



PRELIMIJSTAET CLASSIFICATIOlSr OF SCOLYTOIDEA. 203- 

extends to one or both sides. These chambers are usually occupied 
by the male of the polygamous colony or of the monogamous pair. 

Ventilating burrow. — The ventilating burrow is the vertical burrow 
which is located at more or less regular intervals in the roof of the 
egg gallery and extends to or near the surface. It may serve the 
purpose of ventilating the gallery, or, perhaps more frequently, as 
a place for the storage of boring dust or an opening through which 
this dust may be ejected. Short burrows in the roof or sides may 
be used as places in which the beetles turn around, or may serve the 
purpose of nuptial chambers. 

Branching gallery. — The branching gallery may branch from the 
central or nuptial chamber, or from the side of one of the main egg 
galleries. In the latter case it is referred to as a lateral branch. 

Connecting galleries. — The connecting galleries are those of one or 
more colonies which are connected either through the central cham- 
ber or by lateral and primary galleries, as in many species of Pityogenes, 
Pityophthorus, Carphoiorus, etc. 

Terminal burrows. — The terminal burrows are excavated usually 
by the female beetle at the farther end of a primary or a lateral gal- 
lery, after the egg gallery is completed and while the brood is devel- 
oping, as in Dendroctonus frontalis (Part I, figs. 51 and 52). 

Brood burrows. — The brood burrows are those excavated by the 
adults of a brood before the individuals emerge. They radiate from 
the respective pupal cells of the individuals, as in Dendroctonus, cer- 
tain species of Ips, etc. In the case of species with a single genera- 
tion annually, the developed brood may overwinter in the brood 
burrows. 

Hibernating burrows. — The hibernating or overwintering burrows 
are those excavated in places other than that in which the broods 
developed, such as those of Ips, in the twigs and branches, or in the 
thick corky bark at the base of the trees, and those of Phloeophthorus, 
in the outer bark on the trunks of the living host trees. 

Food burrows. — The food burrow is excavated by the adult in the 
same part of the tree in which it excavates its egg galleries, or in a 
different part, as in Scolytus rugulosus and S. quadrispinosus, which 
burrow in the living twigs at the base of a leaf stem or bud, and in 
PMcEosinus, which burrows at the base of living twigs. 

Trial burrows. — The trial burrows are those made by the parent 
beetles in the bark of living trees preliminary to the general attack 
and the excavation of successful egg galleries. 

Abandoned or failure gallery. — The abandoned or failure gallery 
is one which, through the resistance exerted by the vital part of the 
plant attacked, the beetles are compelled to abandon or be drowned 
in the resin or sap. 



204 THE SCOLYTID BEETLES. 

Exit burrow. — The exit burrow is that through which one or many 
individuals of a developed brood emerge. In the case of ambrosia 
beetles the entrance may also be utiHzed as an exit, but in the bark- 
beetles the exit is usually direct or indirect from the pupal chamber. 

Larval mine. — The larval mine is the food burrow excavated by 
the larvae from the point where it hatches from the egg. The indi- 
vidual mine may be widely separated from or closely approximate 
to those of other individuals of the same brood, and they may be 
arranged in groups or those of the entire brood may be connected to 
form one common larval chamber. 

Larval cell. — ^The larval cell is excavated by the larvae in the side 
of the gallery simply to accomodate the increasing size of the body, 
as in the case of many species of Corthylinse where tho food consists 
of ambrosial fungi provided by the parent. 

Pupal cell. — The pupal cell is formed by the prepupal larva or by 
the pupa itself and is usually located at the end of the larval mine or 
food burrow of the larva. This is especially true in the case of the 
barkbeetles or the wood-mining larvae, which latter, as in Micracis, 
TJiysanoes, Scolytus jnuticus, etc., extend their burrows for a long 
distance from the bark mines into the wood. On the other hand, 
certain species of Dendroctonus form the pupal cell in the outer 
corky bark {D . frontalis) or in the social larval chamber ID. valens), 
instead of at the end of the larval mine as in D. ponderosx. 

All of the named parts of the gallery have characters more or less 
peculiar to the species or group of alhed species. Therefore the gal- 
leries as a whole, or in their various elements individually, or in various 
combinations, are worthy of special attention in the search for taxo- 
nomic characteristics. 

CLASSIFICATION OF THE GALLERIES. 
Egg Galleries. 

In an attempt to classify the scolytoid egg galleries it is impor- 
tant to remember that the newly excavated galleries in which the 
first sets of eggs are deposited are more reliable in suggesting the type 
or group they represent than are the older ones, because in some 
species they may be so radicall}^ changed and confused by secondary 
branches and the interminglmg of two or more galleries that the 
characteristic type may be obscured. 

Larval Mines. 

The larval mines furnish, to a limited extent, evidence of pro- 
gressive modification, as is found in Dendroctonus. The larval mines 
in most of the Cryphalinae are without distinctive characters, while 
in Ipinae, Hylesininae, and Scolytinae their symmetrical arrangement 



PEELIMINAEY CLASSIFICATIOlSr OF SCOLYTOIUEA. 



205 



represents a high stage of progress, and consequently they are more 
characteristic of the species. 

The form of the egg gallery of any species consists of one or more 
of three primary elements in relation to the substance and its fiber 
in which the gallery is made, as follows: (1) The longitudinal gaUery, 
(2) the transverse gallery, and (3) the broad, irregular chamber. 

Types and Subtypes of Egg Galleries. 

The various modifications and combinations of the three primary 
elements seem to represent 8 general types or groups, which are 
designated by numbers, and 32 more specific subtypes or forms, 
which are designated by letters, as follows (see fig. 112) : 

GrouTpl. — The simple or generalized type, (a) Simple, longitudinal; (6) simple, 
transverse; (c) simple cavities; {d) various combinations of a and h, with lateral 
branches. 

Group 2. — The simple, irregular type branching from an irregular central or basal 
chamber, (a) Long, longitudinal, branching; (h) short, sublongitudinal, branching; 
(c) short, transverse, branching; {d) various combinations of h and c. 

Grouj) 3. — ^Ambrosia galleries. Division I, without lateral larval chambers, (a) Sim- 
ple, longitudinal, single or branching; (6) long, transverse, branching; (c) short, 




Fig. 112.— Classiflcation of galleries of Scolytoidea. (Original.) 

transverse, branched, dilated. Division II, tvith lateral larval chambers, (d) Double, 
transverse, branching; 

Group 4- — The specialized, intermediate, short type, branching from a regular cen- 
tral nuptial chamber, (a) Simple to complex, transverse; (6) simple to complex, 
sublongitudinal ; (c) intermediate combinations of a and 6; (d) complex, symmetrical 
com.binations of a and b. 

Group 5. — The specialized, intermediate, long, longitudinal type, branching from a 
regular central chamber, (a) Simple, irregular types with few branches; (6) special- 
ized, many-branched types; (c) specialized, with few branches, usually three; (d) 
specialized, double, longitudinal. 

Group 6. — The specialized, short, transverse type, (a) Simple, irregular, single or 
double, transverse or sub transverse, and sometimes branching; (6) intermediate, 
regular, double, transverse or subtransverse ; (c) specialized, regular, double or single, 
transverse, with or without lateral entrance chamber. 

Group 7. — The specialized, short, double, longitudinal type, (a) Irregular, 
branched ( Micracis) type; (6) irregular, without branches, but with lateral entrance 
chambers; (c) intermediate, more sj^ecialized, with or without lateral entrance chamb- 
ers; (cZ) highly specialized, without lateral entrance chambers. 

Group 8. — The specialized long or short, single, longitudinal type, (a) Long, 
irregular, winding, sometimes with lateral branches, with or without lateral entrance 
chambers at base; (6) short, irregular, without branches, but with lateral entrance 
59026°— 15 4 



206 



THE SCOLYTID BEETLES. 



chambers at base; (c) regular, short or long, with lateral entrance chambers at base; 
(d) the most highly specialized short or long type, without lateral entrance chambers at 



The foregoing classification and terminology is based on the observed 
galleries of a large number of species representing 57 genera and 
includes the galleries of European species figured by other writers. 

RELATION OF TYPES OF GALLERIES TO THE SUBFAMILIES AND 

FAMILIES. 

The relation of the eight groups of galleries to the subfamilies is 
shown in the following table. In some cases one genus may be repre- 
sented by several groups. In PityopJithorus 26 species are represented 
in groups 1, 4, and 5, and in Ips 39 species are represented in groups 
2, 4, and 5. 

Table VI. — Relation of groups of galleries to the subfamilies in th£ Scolytoidea. 



Subfamilies. 


Groups of gallery types and number of genera in each group. 


1 


2 


3 


J 


5 


6 


7 


S 


Ipidae: 

Division I— 

Cryphalinae. . 


Genera. 
4 

1 


Genera. 
4 

1 


Genera. 
3 


Genera. 


Genera. 


Genera. 


Genera. 


Genera. 


Ipinse 


3 


3 


2 






Corthylinse... 


5 














1 




2 




Division II— 

Cryptureinse . 


2 










1 






2 


1 


2 

8 




2 




2 


2 




2 




Scolytidse: 

Hexacolinae 






1 




Bothrosterninse. . 












1 
1 




Scolytinse 












1 


1 


Scolytoplatypodidse: 
Seolytoplatypod- 
inse 






1 
2 








Platypodidse: 

Platypodinae 

































This table shows that all of the simple types (1 and 2) are in the 
Ipidae and that most of them fall in the first part of Division I, while 
the more specialized types fall in the last part of Divisions I and II, 
with the far greater number in the latter; also, that in Scolytidse 
specialized types only have been found. It is probable, however, 
that simple types will be found in the Scolytidse when we know more 
about the habits of the species of the other genera of this family. 
It is also interesting to note from Table VI the number of subfamilies 
and genera in which the same group types are paralleled. 



I 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA, 207 

Table VII. — Relation of groups of galleries to the families of Scolytoidea. 



Groups. 


Ipidae. 


Scoly- 
tidse. 


Scolyto- 

platy- 

podidse. 


Platy- 
podidse. 


Division 
I. 


Division 
II. 


1 

2 

3 

4 

5 

6 

7 

8 


Oenera. 
5 
4 
9 
3 
4 
2 
3 



Genera. 
5 
3 

2 
1 
8 
2 
9 


Genera. 




1 
1 
2 
1 


Genera. 


1 







Oenera. 


2 








In Hypothenemus the types of galleries do not extend beyond 
group 1 and represent specific types a, l, c, and d; Cryphdlus is also 
confined to the same group and types, while PityopJithorus repre- 
sents group 1, a and d; group 4, a, h, c, and d; and group 5, a and d. 
Ips represents group 2, a, h, c, and d, and group 5, a, h, c, and d. 

The relation of types of galleries to species shows some striking 
examples of progressive modification within a genus and of parallel 
characters in different genera. It is not desirable to present a table 
to illustrate these relations in this connection, but it is intended to 
do so in subsequent parts dealing with the subfamilies. 

While considerable attention has been given to the subject, a far 
more comprehensive study of the egg and brood galleries than has 
yet been made is required as a basis for correlating their taxonomic 
characters with the morphological characters of the species. 

TAXONOMIC RELATION BETWEEN THE BEETLES AND THEIR HOST 

PLANTS. 

Among the scolytoid beetles there is often a close taxonomic 
relation between the species, genera, and groups of the beetles and 
the species, genera, or groups of plants they infest, so that we may 
often know the insect by the host, or the host by the insect. 

In other words, the host, together with the character of the gallery 
of a beetle, will often not only serve to identify the species to which 
it belongs but will indicate its systematic position. In a like manner 
the presence of certain species of beetles will serve to identify the 
species of plant and indicate its systematic position. 

Part of Plant Selected by the Beetles. 

The part of the plant in which the egg galleries are excavated is 
also of interest. In some species it is limited to the root or stem of 
an herbaceous plant; in others to the bark on the roots, main trunk, 
larger branches, and smaller branches, or to the twigs or fruit of a 



208 



THE SCOLYTID BEETLES. 



shrub or tree. Indeed it is found that the species of an entire genus 
of beetles may confine their breeding places to a restricted part of 
the plants of a single genus or closely allied group. Therefore even 
the part of the plant infested by a species may be of considerable 
taxonomic importance in indicating the natural position of a species 
or genus of heretofore doubtful position. 

Condition of the Plant. 

The condition of the plant or plant tissue at the time it is occupied 
by the beetles is of considerable systematic and economic importance. 
It ranges from young to old plants, living, declining, dying, and dead, 
and to different stages of decay of the plant or some part of its tissue. 
The fact that there is a relation between the species of beetles and 
one or more of the conditions mentioned shows that there is some- 
thing of taxonomic value in this phase of the subject. 

Relation of the Species of Beetles to the Systematic Posi- 
tion OF Their Host. 

The primary and minor divisions and groups of plant species repre- 
sented in which one or more species of beetles live range from the 
fruiting bodies of certain fungi of the Eumycetes to the liigher flower- 
ing shrubs and trees of the Angiospermse, the greater number of spe- 
cies and genera being confined to the Pinacese of the Gymnospermse 
and the shrubs and trees of the Dicotyledonese of the Angiospermse. 

Range of Host Plants. 

The range of host plants in the families of the Scolytoidea may be 
designated as follows : 

Table VIII. — Range of host plants in the families of Scolytoidea. 



Family in the Scolytoidea. 



Group of host plants. 



Fungi. 



Gymno- 
spermse. 



Monocoty- 
ledonese. 



Dicoty- 
ledonese. 



Ipidse 

Scolytidse 

Scolytoplatypodidse. 
Platypodidse 



Rare. 
None. 
...do.. 
..do.. 



Common. . 

Rare 

None 

Common. . 



Rare. 
None. 
..do.. 
..do... 



Common. 
...Do. 
Rare. 
Common. 



The range of host plants in the genera of true barkbeetles is usually 
more restricted and characteristic than in the genera of ambrosia 
beetles or in those species which live in dead or decaying bark or 
wood. 

Examples of restricted range of Jiost plants. — In the Cryphalinse we 
find that Cryphalus is partial to Abies and Picea; Trypophlcsus to 



PKELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 209 

Alnus, Salix, and Populus. In Ipinse the true Pityogenes and Ips 
are largely confined to Pinus and Picea. In Corthylinse the genus 
Gnathotrichus is confined to the Pinus group, while Monarthrum is 
partial to the dicotyledonous trees. In Crypturginse the species of 
Crijpturgus, Dolurgus, and Dendrodonus are confined to Pinus and 
allied genera. In Hylesininse Tomicus (MyelopJiilus), Hylurgus, and 
Hylastes are confined to Pinus, while the true Hylesinus and Pteleo- 
Mus are particularh^- associated with Fraxinus. 

Examples of wide range of host plants. — In Cryphalinse HypotJienemus 
has a range of host plants from fungi up through many genera of the 
Monocotyledonese and Dicotyledonese but is rarely found as an inhabi- 
tant of the Gymnospermse. Xyleborus has the widest range of all — 
through the ConifersB and Angiospermse. Dryocoetes is divided be- 
tween the conifers and allies of Quercus. In Corthylinss the genus 
Xyloterus is divided between the Gymnospermse and a wide range 
of the trees of the Angiospermse. Corthijlus is confined to a wide 
range of trees in the Angiospermae, as is also MonartJirum, except 
in the case of a few records from the Pinus and Juniperus groups. 
In Scolytinse the genus Scolytus as at present recognized is divided 
between the genera of the Finales (except Pinus) and the Amentales 
and Rosales. In Platypodinse there is a wide range of food plants 
in the trees of the Gymnospermss and Angiospermse. 

Associations of Species of Beetles and Species of Plants. 

In some of the genera which are restricted in the range of host plants 
as well as in those with the widest range there are many species which 
are restricted to a single species or group of closely allied species of 
plants. In Phloeotribinse there are species peculiar to Morus, Celtis, 
etc., and in PJilceo sinus , with but very few exceptions, each species is 
peculiar to, or prefers, a different species of cypress, cedar, or juniper, 
or group of allied species. In Scolytus we find Hicoria, Ulmus, Fagus, 
Betula, Celtis, Quercus, Ahies, Picea, Pseudotsuga taxifolia, etc., with 
species of beetles peculiar to each plant genus. 

Summary of Taxonomic Evidence Furnished by Host Rela- 
tions. 

In a study of the relations between the insects and their hosts some 
rather striking facts have been determined which have furnished evi- 
dence to clear much of the confusion in classification based on mor- 
phological characters alone and in which parallel or analogous char- 
acters have been mistaken for those of affinity. By the old method 
of morphological distinction closely allied species and genera have 
been widely separated in the classification and distantly related ones 
placed together. A number of such cases have been detected where 
the host plant and the character of the galleries have been studied. 



210 THE SCOLYTID BEETLES. 

In other words, the physiological characteristics gave the best clue to 
the natural affinities of the various groups and led to the discovery of 
heretofore overlooked morphological characters which furnished 
conclusive evidence of their true position. 

The close relationship between some of the existing representatives 
of ancient groups of plants and representatives of evidently ancient 
types of the beetles indicates that the beetles and plants may have 
been closely associated in their evolution from their respective primi- 
tive forms. 

GEOGRAPHICAL DISTRIBUTION IN ITS BEARING ON TAXONOMY. 

Superfamily SCOLYTOIDEA. 

It is evident from our present knowledge that the superfamily 
Scolytoidea is represented to a greater or less extent in every section 
of the world where woody plants grow. 

Family IPIDiE. 

The family Ipidse is also represented by species in all of the great 
faunal regions. The subfamily Cryphalinae, with its widely dis- 
tributed Hypothenemus, Steplianoderes , and Xylehorus, has a wider 
range of distribution perhaps than is found in any of the other sub- 
families. The Ipinse are more restricted to the Holarctic regions and 
to the distribution of Pinus and its allies, Ahies, Picea, Larix, etc. 
The Corthylinse are more restricted to temperate, subtropical, and 
tropical America except in the genus Xyloterus, which extends 
through the Palsearctic and Nearctic regions. The Micracinse, with 
the exception of Liparthrum and Hypohorus, are largely restricted to 
north temperate America. In Crypturginae the genus Aphanar- 
thrum is restricted to Madeira and the Canary and Cape Verde Islands, 
while Crypturgus has a wide range through the Palsearctic and Nearc- 
tic regions, and Dendroctonus, with one exception, is confined to North 
and Central America. The Phloeotribinse and Hylesininse are widely 
distributed throughout the regions of tree growth. 

Family SCOLYTID^. 

The family Scolytidse, as represented by the genus Scolytus, extends 
over a wide range of the Palsearctic and Nearctic regions, but the 
greater number of genera and species are evidently to be found in the 
subtropical and tropical regions of Central and South America. 

Family SCOLYTOPLATYPODID^. 

The family Scolytoplatypodidse, so far as known, is restricted to 
small sections of the eastern Palsearctic and of the Ethiopian regions. 



PRELIMIISrAKY CLASSIFICATIOlSr OF SCOLYTOIDEA. 211 

Family PLATYPODID^. 

The family Platypodidse has a wide range through the temperate, 
subtropical, and tropical regions of both continents. 

There are many features in the distribution of genera and species 
which are of special interest, but there is need of information on many 
species from the different countries which at present are poorly or 
not at all represented in collections. 

PAIRED SPECIES. 

The subject of so-called paired or parallel species is of special 
interest in connection with a study of the distribution. There are 
some striking examples of paired species in the genus Dendroctonus 
which, if they occupied the same local faunal area, would be difficult 
of separation on account of their close resemblance in structural 
characters. These paired species are hrevicomis and harheri, frontalis 
and arizonicus, mexicanus and parallelocollis, monticolse and ponde- 
rosse, piceaperda and engelmanni, punctatus and micans, and terebrans 
and valens. In each case the pairs are more or less widely separated 
from each other in their geographical distribution, as, for example, 
micans of northern Europe and punctatus of the Appalachians of 
North America, frontalis of the Southern States and arizonicus of 
Arizona, harheri of Arizona and New Mexico and hrevicomis of the 
Pacific Slope States, Idaho, Montana, and part of Wyoming, ponde- 
rosse of the central and southern Rocky Mountains and monticolse in 
the Northern Rocky Mountains and Pacific Slope. 

There are any number of similar cases of so-called paired species 
in other genera, and the supposition that some of them are one and 
the same species has led to considerable confusion concerning the 
true range of a species. Between North America and Europe we 
have several examples, such as Anisandrus pyri of America and 
Anisandrus dispar of Europe, Dryoccetes autogro.pJius of Europe and 
Dryoccetes septentrionis of the western coast and Alaska of America, 
Xyloterus lineatus of Europe and Xyloterus hivittatus of America, 
Hylurgops glabratus of Europe and Hylurgops pinifex of America. 
There are many others common to two or more countries which super- 
ficially seem to be the same. 

REVIEW OF TAXONOMIC CHARACTERS AND CHARACTERISTICS. 

In a review of the foregoing discussion of the morphological char- 
acters to be found in the different stages of the scolytoid beetles and 
of the physiological characteristics in their habits, it is shown that 
there is a wide range of taxonomic elements by which to distinguish 
species and genera and to indicate or fix their position in the classifica- 
tion. Indeed the vast number of these characters and characteristics 



212 THE SCOLYTID BEETLES. 

and the wide range in their variation, together with the disturbing 
factors, or reversals, and parallel modifications, are enough to over- 
whelm, confuse, and discourage anyone who attempts to study and 
utilize them in a comprehensive classification. It is plain, however, 
that it is only through such a comprehensive treatment that we can 
hope to approach the ideal natural classification. 

MORPHOLOGICAL CHARACTERS. 

In a review of the morphological characters it is found that the 
tarsus and tibia are of special value in the classification of the super- 
family; the head, pronotum, elytra, and third joint of the tarsus for 
the family; the head, tarsi, pronotum, elytra, anterior coxae, abdomi- 
nal sternite, and pygidium for the subfamily; the antennae, eyes, 
pronotum, elytra, tarsi, tibiae, body, abdominal sternites, anterior 
coxae, and mouthparts for the genus, and for the species there is such 
a wide range of characters of varying, and sometimes reverse, value 
in different genera that they can not be specified except for limited 
groups. However, the size, form, color, vestiture, and sculpture 
of the body, the front of the head, elytral declivity, etc., are in gen- 
eral among the most important bearers of specific characters. 

In the digestive system the masticatory plates of the proventric- 
ulus appear to be of special taxonomic value when applied to sub- 
famihes, genera, and species. 

In the reproductive organs of the male there is a very wide range 
of variation in the primary and secondary elements of the posterior 
or chitinized section, but with a few exceptions their taxonomic 
value appears to be restricted to major and minor divisions of the 
genus and especially to the separation of the species. In the anterior 
section of the organs the length of the ductus ejaculatorius and the 
form of the testes, seminal vesicles, vas" deferens, and mucus glands 
appear to represent the principal taxonomic elements, and they are 
of varying value as applied to major and minor groups. 

The female reproductive organs, like the proventriculus, appear to 
possess a number of taxonomic characters of family, subfamily, and 
generic value, such as the presence or absence of the cement glands 
and their varying characters and the presence and character of the 
spermatheca with its seminal duct. 

In the secondary sexual characters we find a few which are peculiar 
to a genus or to groups of allied genera, but m general their principal 
value appears to be m distmguishing the species. 

In the pupae it is found that the head and the tergal, lateral, pleural, 
caudal, and femoral spmes appear to be the most unportant taxo- 
nomic elements. 

In the larvae there is a wide range of variation in the form of the 
body and in some of the primary elements of the head and posterior 



peelimijStary classification of scolytoidea. 213 

abdomiiial segments, spiracles, etc., which are of value in separating 
the families, but the majority of these characters are of greatest value 
in connection with the genera and species. 

Tlie egg and embryo doubtless bear some important characters, 
but they have not been sufficiently studied by the writer to justify 
their discussion in this connection. 

PHYSIOLOGICAL CHARACTERISTICS. 

A review of the evidence relatmg to the physiological character- 
istics shows that the feeding, breeding, flight, and social habits, the 
galleries, larval mines, host plants, the species of plant, the part of 
the plant occupied, conditions of the plant, restricted or wide range of 
host relations, and features in geographical distribution have some- 
thing to contribute in facts and evidence of more or less taxonomic 
importance. 

CORRELATION OF MORPHOLOGICAL AND PHYSIOLOGICAL ELEMENTS. 

In order to arrive at satisfactory^ conclusions in regard to the proper 
correlation of the mass of taxonomic data mentioned m the foregomg 
review one must have a far more comprehensive and first-hand 
knowledge of the subject as applied to the species of the world than is 
possessed by any investigator at the present time. Therefore it is the 
object of the writer to call attention to the need of further investi- 
gation of this broader prmciple of zoological taxonomy rather than 
to attempt to draw conclusions. It is evident, however, that at some 
future time the essential facts will be correlated uito a sj^'stem which 
wiU not only indicate true natural relations but perhaps give. a better 
clue to the action of natural laws and a better mterpretation of the 
fundamental principles involved m the evolution of life in general. 

In order to arrive at sound conclusions the whole subject must be 
mvestigated without prejudice for or against any theory as to phylo- 
genetic origin of the organism, or as to the primitive or recent char- 
acter of an organ or element. The problem must be attacked with 
the view of locating the more important or essential facts relating to 
the distinguishing characters and characteristics as applied to the 
species and their various aggregations into genera, subfamilies, and 
families and the major and minor divisions of each. In other words, 
a given group of organisms should be studied in all of its morphological 
and physiological aspects with the view of locating, by the process of 
elimination, the elements of primary importance until one or more 
characters in each of the principal morphological and physiological 
groups of taxonomic elements are found to correlate in the formation 
of a harmonious taxonomic compound. 

It has been shown in the foregoing pages that a classification based 
on any limited set of external or internal elements of the adult body, 



214 THE SCOLYTID BEETLES. 

such as those found in the proventriculus or ui the male or female 
organs, will suggest a phylogenetic system, but when it is found 
that each system differs from the other to such an extent that the 
same genus or species will occupy a radically different position in 
the different systems it is perfectly plain that the true taxonomic 
value of the elements has not been correctly interpreted or applied. 
On the other hand, it is equally plain that if characters can be found 
m all or a majority of the groups of external and internal elements 
which point in the same direction, we may safely assume that we 
have more nearly approached the true principle involved and the 
ideal classification. 

Examples of an attempt by the writer to correlate and harmonize 
the various morphological and physiological elements will be found 
in the synoptic tables of Dendroctonus, Part I of this bulletin, and of 
Pissodes in Technical Series 20, Part I. 

PARALLEL MODIFICATIONS. 

Parallel modification in morphological and physiological elements 
is without doubt an important factor to be considered in taxonomy. 
It is evident from a comparative study of the various systems of 
classification that the failure of taxonomists fully to realize its im- 
portance has m many cases led to wrong conclusions. It has been 
shown in the foregomg discussion that there are a great many ex- 
amples of parallel characters and characteristics in widely separated 
species and genera and that if they are not recognized and properly 
interpreted as such by the taxonomist, radically wrong positions 
will be assigned to many species and genera. 

REVERSALS. 

The reversal of characters and characteristics in different species 
of the same genus or m different genera and larger groups is another 
important fact to be kept in mind, especially as related to secondary 
sexual characters. Therefore it is never safe to conclude that be- 
cause a given character or a group of characters is of special value 
in distinguishmg one group, genus, species, or sex, it will hold in aU 
cases. There have been numerous examples of wrong determina- 
tion of the sexes from a failure to recognize this principle, as has been 
pointed out by the writer (Hopkins, 1894, pp. 274-280) and Bland- 
ford (1895, pp. 83-86). As has been shown on preceding pages, 
reversals are also found in specific, generic, and group characters, 
even to the subdivision of the superfamily, where we find a most 
striking example in the reversal of the apical spine or process of the 
anterior tibia from the mner to the outer angle. 



PRBLIMINAKY CLASSIFICATIOlsr OP SCOLYTOIDEA. 215 

PROGRESSIVE MODIFICATIONS. 

The discontinuous yet more or less progressive change or variation 
in the modification of morphological and physiological elements 
along definite lines within the minor to major groups is very evident 
in every group. The more this subject is studied the more we are 
convinced that there are certain important facts involved in this 
principle that have not been satisfactorily explained by any theory 
of the processes of evolution. The recognition and application in 
taxonomy of those unexplained features does not, however, neces- 
sarily require the acceptance or rejection of any theory of ortho- 
genesis or phylogenesis. It is only necessary to correlate them with 
other more easily explained elements of distinction or to utilize them 
as guides to the position a species or group should occupy in a given 
series. 

Examples of Progressive Modification. 

The examples of progressive modification in morphological char- 
acters and physiological characteristics which have been noted by 
the writer in the scolytoid beetles may be summarized as follows: 

Morphological characters. 

Body small to large. 

Body slender to stout. 

Body with scales to hairs, to glabrous. 

Head concealed to exposed. 

Head short and broad to narrow and subrostrate . 

Head with front convex, glabrous, to concave and pubescent or fringed. 

Head with eyes oblong, elliptical and not emarginate, to short, oval, and deeply 
emarginate or divided. 

Antennal joints of funicle increasing in number to the limit cf seven. 

Antennal joints of club decreasing in number through fusion or disappearance of 
sutures. 

Prothorax long and narrow to short and broad. 

Prothorax with sides not margined to margined, or not emarginate to emarginate. 

Prothoracic pleurum convex to flat and concave. 

Tarsi with third joint simple to emarginate and bilobed. 

Tarsi with first joint short to long. 

Elytral declivity convex and smooth to rugose and armed; retuse to concave, with 
the margin unarmed to strongly armed. 

Sexes of unequal size and the males rare to equal size and the males common. 

Secondary sexual characters obscure to prominent. 

Galleries. 

Simple cavities in decaying bark and wood, to complex designs and regular forms of 
egg galleries and larval mines. 

Excavated in bark to excavated in wood, seeds, etc. 

Social habits. 

Unorganized polygamy to organized polygamy, to highly organized monogamy. 
Independent larvae, procuring their own food, to dependent larvae, with the food 
provided by the maternal parent. 



216 THE SCOLYTID BEETLES. 

STATISTICAL TAXONOMY. 

The subject of statistical taxonomy has received considerable 
attention by the writer and an attempt has been made (Hopkins, 
1911, pp. 28-30, PI. II) to show its successful application in the genus 
Pissodes. The detail and accuracy required for this method and the 
difficulty of determining a specific and taxonomic formula are so great, 
however, as to be rather trying on the patience of the investigator. 
Therefore it is the opinion of the writer that it should not be resorted 
to except in cases in which other methods fail to give satisfactory 
results and then only when the elements to be included are of such a 
nature as to be readily available for the mathematical determination 
of relative proportions. There is m this, as m many subjects relating 
to taxonomy, a need of a more comprehensive investigation in order 
that the most reliable basis for conclusions may be reached. 

THE SPECIES. 

Reference has been made (Part I, p. 66) to the WTiter's views on 
specific distinction and the range and limits of specific variation. In 
addition it may be said that no species can be established beyond dis- 
pute without a knowledge of both the morphological and physiological 
elements of distinction. This does not, however, preclude the recog- 
nition andnammgof imperfectly defined and poorly represented species 
and of retaining them as long as they can be readily distinguished 
from other allied forms. Such names and definitions serve as a basis 
for study until it is more definitely shown whether or not the forms are 
worthy of permanent specific distinction. There is necessarily a wide 
range in the relative prominence of the distinguishmg characters as 
between a species which is the onh^ representative of a genus and sub- 
family and one of a group of closely allied species. It is found, how- 
ever, that for the purpose of systematic and economic investigation 
the isolated species may be of no more value for study and ma}^ con- 
tribute no more to the advancement of knowledge than the one which 
is with difficulty separated from its congeners. 

Since we now insist upon a single specimen as the author's designa- 
tion of the type of a described species it is deemed by the writer to be 
perfectly proper to base the description of a new species on a single 
specimen, provided the author of the description is sufficiently 
familiar with the previously described allied species and genera to 
enable him to recognize the characters and characteristics which are 
of real specific importance. 

VARIETIES, RACES, ETC. 

There is in all species a greater or lesser range of variation from 
the type, and often some groups of individuals may be so different as 
to indicate a distinct varietv or race. If such variant forms are 



PEELIMIJN^AEY CLASSIFICATION OF SCOLYTOIDEA. 217 

found intermingled with the broods of the typical forms and the 
characters are not safhciently constant to distinguish them as separate 
species they should' simply be considered as coming within the range 
of specific variation, but in the writer's opinion these varieties should 
not be distinguished by a trinomial. 

In some genera and species the sexes may so differ in w^hole groups 
of characters as to indicate, m some cases, vdifferent genera. But 
there is always some character or set of characters common to both 
sexes which would point to the same species even if they w^ere not 
found associated in the same brood. 

There is evidence that in some of the species of Xyleborus and 
allied genera in the Cryphalmse there may be occasional individuals 
which represent a degenerate form or a caste in the social relations of 
a brood or colony, with uniform but radically different characters 
from those of either sex, and that on account of the radical differences 
w^hich separate them from the species with which they have been 
found they have heretofore been recognized as good species. Xyle- 
horus planicollis Zimm. may be such an odd member of the Xyleborus 
inermis colony and Xyleborus viduus Eichh. an odd type of the 
Xylehorus fuscatus colony, but further observations will be needed 
to settle the question. If this should be true in these cases it may 
hold with isolated cases in other social species, like those of HypotJie- 
nemus, Steq)hanoderes, Dryoca;ies, etc. 

THE GENUS. 

There is more latitude, perhaps, for the selection of distinguishing 
characters of the genera than there is for the species, but the same 
principle applies. There mast be some single character or group of 
characters common to a group of allied species which will serve to 
distinguish the group readily from all other allied groups. The 
range of departure or variation from the tj^pe of the genus is restricted 
in some genera, but in other genera with many species there may be 
a very wide range, so that the species mil fall in distmct divisions 
and subdivisions, which are designated by some authors as subgenera. 

Unfortunately there is a wide range in the opinions of different 
authors as to the limits of a genus. Some go to the extreme in 
restricting it to closely allied species w^hile others go to the opposite 
extreme (Hagedorn, 1910) and include a large number of genera or 
so-called subgenera. It is plain to the writer that of the two extremes 
the latter is the more objectionable because it will certainly contrib- 
ute more than the former toward retarding than advancmg knowl- 
edge. The writer believes that there is a middle ground on which 
systematists should endeavor to get together in order that there may 
be more uniformity in the conception and definition of the genus. 



218 THE SCOLYTID BEETLES. 

NOMENCLATURE. 

Nomenclature, as applied to taxonomy, is a subject on which there 
are wide differences of opinion, and it is evident that until there is 
more uniformity there will be continued contributions to the con- 
fusion of knowledge along with those which contribute to its advance- 
ment. 

It seems to the writer that the subject of designating by name 
varieties, forms, races, subspecies, and subgenera should demand the 
special attention of systematists with the view of coming to an under- 
standing as to the limitmg of such names to the more definite con- 
cepts, as those of the species or genus. It would seem that if we 
should limit the names to the s])ecies, genera, subfamilies, families, 
and superfamilies within an order and designate the major and minor 
divisions of each as divisions (I, II), subdivisions (A, B, C, D), sections 
(al, a2, etc.), subsections (bl, b2, etc.), series (cl, c2, etc.), and sub- 
series (el, e2, etc.), it would be in the line of progress toward a con- 
sistent, practical, and uniform method of expressing the varying 
ranks as interpreted by different authors. It would avoid, at the 
same time, the use of names for divisions and subdivisions which 
have different meanings in the systems proposed by different authors 
and prevent the accumulation of obsolete names with every change or 
important advance in the classification. 

In the systematic treatment of the scolytoid beetles the writer fails 
to see the need of recognizing subspecies or subgenera. The species 
and the genus are the two most important biological concepts on 
which to base both systematic and economic investigations. They 
serve, also, as the most important units on which to base a classifica- 
tion. Therefore the writer holds that the individual represents a 
species and that a species represents a genus; hence the introduction 
of the trinomial for a subspecies and the naming of a subgenus are 
unnecessary. Moreover, he believes that a general practice of giving 
names to such divisions of these taxonomic units will ultimately lead 
to endless confusion and retard rather than advance the spirit of 
research and the acquisition of knowledge. At best the designation of 
the species and the genus to which a given individual should be refer- 
red is an arbitrary interpretation of a concept. Therefore, when an 
author designates an individual or a group of individuals as repre- 
senting a subspecies, or a group of species as representing a subgenus, 
it involves the assumption that the concept is a fact and that he has 
sufficient knowledge of this fact to enable him to analyze it into com- 
ponent categories the relations of which are so definitely determined 
as to justify the subordination of one part to another. 

There is such a wide range for the interpretation of specific and 
generic distinctions and such a vast difference in the relative rank of 
such distinctions, between isolated survivors of highly specialized and 



PKELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 219 

ancient groups and common, closely allied, and variable forms, that 
the difference between two species in one genus may be equivalent 
to or greater than that between two allied genera. In a Uke manner 
the difference between two genera may be almost equivalent in rank 
to that between two allied subf amihes. Therefore the fact that a given 
group of individuals or a group of species appears to represent a position 
of lower rank than that of an allied species or genus is not sufficient rea- 
son that it should be designated as a subspecies or subgenus. In other 
words, it is the writer's opinion that if a group of individuals is suffi- 
ciently distinct from allied groups to justify its designation by a name, 
it should be recognized as a species ; and if, on the other hand, the group 
is connected with allied forms by such a number of intermediate forms 
as to render its specific distinction doubtful, it should be included with 
other variable types under a species name, and that the same principle 
should hold in regard to the genus. 

AU of the purposes served by the subspecific designation can be 
served just as well, if not better, by classifying the individuals of a 
species into major and minor divisions, sections, etc., and letting these 
represent the same conception as that represented by the named sub- 
species, and on the same principle the divisions and subdivisions of a 
genus will serve the same purpose as if designated by names. 

TYPES OF GENERA. 

Difference of opinion among systematists who have worked on the 
scolytoid beetles, guided as they have been by different rules and con- 
ceptions as to what constitutes a vafid genus, has led to much uncer- 
tainty and confusion as to some of the older names for the genera of 
our common species. Therefore it seems necessary that the author 
should present the evidence which appears to him to be conclusive in 
regard to the generic names and the synonymy of Scolytus, Ips, Tomi- 
cus, Cryphalus, Hy pothenemus , etc. 

Genus SCOLYTUS Geoflfroy. 

It has seemed to the writer that there is not sufficient reason or 
authority for the suppression of the name Scolytus as proposed and 
clearly defined by Geoffroy (1762, p. 309). The single "species" of 
the "genus" recognized by him was at the same time (p. 310) indi- 
cated by a number, the genus name (Scolytus), and a reference to a 
figure (Vol. I, pi. 5, fig. 5), and the characters were more clearly 
defined in a description. 

Mtiller (1764, p. xiv) recognized the genus Scolytus Geoff, and re- 
ferred to the original description and figures. 

Schaeffer (1766, Tab. CXII, figs. 1, 2, 3, 4) redescribed the genus 
Scolytus and described and figured the species indicated by Geoffroy, 
but did not name it. 



220 THE SCOLYTID BEETLES. 

Linnaeus (1767) failed to mention the genus or to refer to Geoffroy, 
Miiller, or Schaeff er. 

Fabricius (1775, p. 59) recognized tlie species described by Geoffroy 
and redescribed it under the name Bostrichus scolytus, with the cita- 
tion ''Geoff. Ins. 1-310, [No.] 1, Tab. 5, fig. 5, Mai." Thus Fabricius 
recognized the species indicated by Geoffroy on page 310, but did not 
refer it to the genus described on page 309 under the name Scolytus 
because he (Fabricius) evidently considered it synonymous with Bos- 
trichus. Consequently, the name proposed by Geoffroy for the genus 
should stand with Bostrichus scolytus (Fab.) as the type. 

Sulzer (1776, Th. 1, p. 21: Th. II, Tab. II, f. 13k), under the name 
Dermestes scolytus, described and figured the species indicated by 
Schaeff er, 1766. 

Mtiller (1776, p. 57) published a description under the name Scoly- 
tus punctatus but referred it doubtfully to Geoff roy's figures. 

Goeze (1777, p. 143), under the name "Dermestes scolytus Geof- 
froi," cited Bostrichus scolytus Fab. (1775), Scolytus Geoff. (1762), 
and Scolytus punctatus MiiJl. (1776). Goeze evidently did not men- 
tion "Geoffroi" as a specific name but merely to indicate that 
Geoffroy was the author of or authority for the name Scolytus. 

Linnaeus (178S, 1793, p. 1602) recognized Bostrichus scolytus Fab. 
and cited Fabricius, 1787, Geoffroy, 1762, Sulzer, 1776, and Schaeffer, 
1766. 

Herbst (1793, p. 124) described the genus Eccoptogaster with 
Bostrichus scolytus Fab. as the type. 

Olivier (1795, No. 78, p. 5, PI. I, fig. 4 a, b, c) adopted the name 
Scolytus for the genus and cited Geoffroy, 1762, and Fabricius, 1775, 
but substituted for the species the name destructor in the place of 
Scolytus Fab., evidently concluding, as other contemporary writers 
did, that the generic and specific names could not be the same. In 
fact this opinion evidently influenced the action of Fabricius, Miiller, 
Herbst, and others. 

Curtis (1824, p. 43) designated the type of the genus Scolytus as 
Bostrichus scolytus Fab. 

Genus IPS De Geer. 

There appears to be no room for doubt that the genus Ips of De 
Geer (1775, p. 190), with Dermestes typographus Linn., 1758, as the 
type, has j)riority over Ips of Fabricius (1776, p. 23). 

Genus TOMICTJS Latreille. 

Latreille (1802, p. 203) described the genus Tomicus with Hyle- 
sinus piniperda Fab. (1801, p. 392) as the type, but H. piniperda of 
Fabricius is, as cited by him (1801, p. 392), the same as Bostrichus 
piniperda (L.) Fab. (1775, p. 60; 1792, p. 367) and Dermestes pini- 



PEELIMINAEY CLASSIFICATION OF SCOLYTOIDEA. 221 

'perda Linn. (1758, p. 355 and 1767, p. 563). Therefore it appears 
that the name Tonnicus should stand, with Hylesinus piniperda (L.) 
Fab. as the type. The slight difference in the description by Fabri- 
cius of the insect he identified as Linnseus's species does not warrant 
the conclusion that it was different, because the species has the 
characters mentioned. 

The species piniperda L. 

piniperda L., Dermestes Linnseus, 1758, p. 355; 1761, p. 143; 1767, p. 363. 
piniperda L., Bostrichus Fabricius, 1775, p. 60; 1792, p. 367. 
piniperda L., Hylesinus Fabricius, 1801, p. 392. 
piniperda L., Tomicus Latreille, 1802, p. 203. (Type of genus.) 
piniperda L., Blastophag^iis Eichhoff, 1864, p. 25. (Type of genus.) 
piniperda L., Myelophilus Eichhoff, 1878, p. 400. (Type of genus.) 

Genus HYLASTES Erichson. 

Erichson (1836, p. 48) described the genus Hylastes to include 
Bostrichus ater Paykull, Bostrichus angustatus Herbst, etc., stating 
that Bostrichus ater Fab. was not a scolytid. Therefore, since 
Bostrichus ater Paykull is a good species, it stands as the type of tha 
genus Hylastes Erichson, as designated by Thomson, 1859, p. 146. 

Genus CRYPHALUS Erichson. 

Erichson (1836, p. 61) described the genus Cryphalus to include 
Apate tilise Panz., Apate fagi Fab., and Bostrichus asperatus Gyll. 
Thomson (1859, p. 146) designated {Bostrichus) Cryphalus asperatus 
Gyll. as the type and (p. 147) referred Cryphalus tilise Panz. to his 
monobasic subgenus Ernoporus and (1865, p. 360) Cryphalus fagi 
Fab. to Ernoporus. 

Hypothenemus Westw., 1834, p. 34; Ernoporus Thorn., 1859, p A 4:7; 
Trypophlmus Fairm., 1868, p. 105; Stephanoderes Eichh., 1871, p. 
132, and Cryphaloides Form., 1908, p. 91, are all sufficiently distinct 
from Cryphalus Erich, and from one another to stand as separate 
and distinct genera; therefore they should not, in the opinion of the 
writer, be considered as subgenera of Cryphalus. 

Genus HYPOTHENEMUS Westwood. 

Westwood (1834, pp. 34, 36, PI. VII, fig. la to h) described the 
genus Hypothenemus with eruditus Westw. as the only species and 
mentioned and illustrated a three- jointed funicle of the antenna as 
one of the distinguishing characters. 

Duvall (1868, p. 105, pi. 33, fig. 161) examined specimens from the 
type series and redescribed and figured the species, and referred to 
the funicle as having but three joints. 

Eichhoff (1879, p. 165) referred the genus Hypothenemus Westw., 
and species eruditus Westw. to synonymy under Stephanoderes 
59026°— 15 5 



222 THE SCOLYTID BEETLES. 

ariccse (Hornung, 1842, p. 117 [not 115]) because he fonnd that cer- 
tain species previously referred to Hypoihenemus had five joints in 
the funicle of the antenna instead of three, as designated by West- 
wood. 

Fauvel (1884, p. 315) examined specimens of H. eruditus Westw. 
from the type series and compared them with examples of BostricTius 
hoieldieui Perroud, 1864, p. 188, and B. ariccsp, Hornung, 1842, p. 117, 
identified by Eichhoff, and concluded that all three were identical, 
and that HomceocrypJialus Lind., 1876, p. 168, as represented by 
Stefhanoderes elilersi Eichh., was identical with Hypoihenemus Westw. 

Newberry (1910, p. 83) determined that the funicle had four 
joints. 

Through the kindness of Prof. Poulton, the writer had the oppor- 
tunity of examining a specimen of H. eruditus Westw. from the type 
series, and the funicle was found to be four-jointed. It is not improb- 
able, however, that the antenna examined by Westwood was from a 
naale, which, in this genus, is smaller than the female and has but 
three joints in the funicle. Therefore there can be no doubt as to the 
validity of the genus Hypoth-enemus Westw. as distinguished from 
the genus StepTianoderes Eichh., which has a fi\'e-jointed funicle. 

Genus POLYGRAPHUS Erichson. 

Erichson (1836, p. 57-58) described the genus PolygrapTius, citing 
Dermestes polygrapJius L. (1758, p. 355) as synonymous with P. 
jpubescens Fab. (1792, p. 368) ; but since P. polygrapJius L. has priority, 
the genus stands with Dermestes polygrapJius Ij. as the type. 

Genus LEPISOMTJS Kirby. 

Earby (1837, p. 193) described Lepisomus as a subgenus of Apate 
Eab., including three species, one of which, Apate {Lepisomus) 
Tiigriceps, has been referred to synonymy (Lee, 1868, p. 169) while 
Ap)ate (Lepisomus) hrevicornis is not recognizable, probably a synonym. 
Therefore Apate {Lepisoinus) ruf.pennis Kirby is the type of the 
genus Lepisomus, which, on account of the six-jointed funicle, is a 
good genus. For the same reason PolygrapJius grandiclava Thom. 
(1886, p. 62) must also be referred to this genus. 

DESCRIPTION OF A NEW GENUS AND SPECIES. 

WEBBIA n. gen.Q 

.\ntennal funicle four-jointed, the fourth broad; club narrowed 
irom middle to base, broader than long, with one sinuate chitinous 
suture on the obliquely truncate anterior face, the posterior face 

a This genus and species is described here in order that the subfamily may be 
included in the classification. 



PEELIMIXAEY CLASSIFICATION OF SCOLYTOIDEA. 223- 

without sutures; eyes oblong, elliptical, deeply emargiiiate ; anterior 
tibia strongly narrowed to apex, with submarginate row of closely 
placed teeth on the ventral side; elytral declivity with many closely 
placed marginal' teeth ; lateral margin of pronotum acute, anterior 
margin mthout serrations. Type of genus, Wehhia dijAerocarpi new 
species. 

This genus is named for Mr. J. L. Webb on account of the large 
number of scolytoid beetles collected by him during his brief employ- 
ment in the Philippine servdce. 

Webbia dipterocarpi n. sp. 

Length, female tvpe, 3.1 mm. Snbelongate ; pronotum and all 
but the decHvity of the elytra light ferruginous, the declivity black. 
Pronotum with anterior area swollen, opaque, nearly smooth; apex 
steep, subtruncate, and finely rugose; median and posterior areas 
smooth, subopaque, and very fuiely punctured. Front moderately 
narrow, subconvex, opaque, with fine median line. Elytra to near 
declivity smooth, shining, with fine, closely placed, and confused 
punctures with no trace of strise, narrow, and near vertex slightly 
swollen, densely opaqtie, nearly black; declivity steep, flat, opaque; 
interspace 1 elevated, smooth; interspaces 2 and 3 each with straight 
rows of granules; margin from vertex to apex armed each side with 
10 closely placed serrations or teeth, becoming slightly larger toward 
apex. These teeth evidently represent the 10 interspaces of the 
normal elytra. ISTear Pagbilao, Philippine Islands, in wood of dead 
log of Dipterocarpus grandiflorus Blco., August 7, 1903. J. L. Webb, 
collector. Under his number 94c. 

Type.— C2it. No. 7406, U. S. National Museum. 

Length, male type, 2.6 mm. Subelongate; pronotum slightly 
broader than elytra which are narrowed toward declivity, light 
ferruginous throughout. Head narrow, shining, with deep epis- 
tomal impression and a slight posterior impression. Pronotum 
opaque, fuiely rugose throughout, with distinct median impressed 
shining line to. anterior declivity which is steep and strongly retuse 
to anterior margin, sides slightly narrowed from anterior angles to 
base, lateral margins subobtuse; elytra with sides slightly narrowed 
toward declivity, shining, punctured as in female but with striae evi- 
dent and faintly impressed; declivity steep, flat, opaque, rugose, 
but ^vithout elevated interspaces, margin with but seven marginal 
teeth, coarser toward vertex and arranged in groups of 3-2-2. In 
a lateral aspect the dorsal line forms a broad curve from the mandi- 
bles to the apex of the abdomen. From same colony as the female. 

The smaller size and general appearance of the male suggests 
affinities with the Xyleborus group but it is radically different in all 
of the more imnortant characters. 



224 THE SGOLYTID BEETLES. 

The fact that specimens were collected in the wood indicates that 
the food habits are similar to those of Crossotarsus lecontei, found in 
the same log. 

The pupa is peculiar in the absence of caudal spines, the presence 
of tergal and pleural hairs instead of tubercles, and the length of the 
wing-pads, which extend to the apex of the abdomen. 

The larva, as shown by a dried specimen, appears to have the 
posterior part of the body stouter, then narrowed toward the apex 
of the abdomen. 

PRELIMINAEY CLASSIFICATION OF THE SUPERFAMILY 
SCOLYTOIDEA. 

Key to the Families. 

(See PI. IX.) 

I. Anterior tarsi witli joint 1 shorter than 2, 3, and 4 together. 

A. Anterior tibia without prominent process on the outer apical angle. 

Family Ipidae. 

B. Anterior tibia with prominent process on the outer apical angle. 

al. Anterior tibia without prominent rugosities on ventral area. 

Family Scolytidse. 
a2. Anterior tibia with prominent rugosities on ventral area. 

Family Scolytoplatypodidae. 
II. Anterior tarsi with joint 1 longer than 2, 3, and 4 together. 

C. Anterior tibia with prominent apical process and with rugosities on the 

ventral area Family Platypodidee. 

Family IPIDiE. 

Key to the Subfamilies. 

(See Pis. X, XI, XIII, XIV.) 

I. Pronotum with anterior dorsal area commonly rugose; head concealed from above; 
anterior tarsi with joint 3 simple. 
A. Abdominal sternites 5-7 not strongly ascending. 

al . Anterior tibia broader toward apex or serrate on outer margin, 
bl. Abdominal sternite 7 with posterior margin always rounded. 

1. Subfamily Cryphalinse. 
b2. Abdominal sternite 7 with posterior mai'gin rarely rounded. 

cl. Pronotum and elytra clothed with scales orhaii-s, very rarely 

glabrous 2. Subfamily Ipinae. 

c2. Pronotum and elytra without scales, commonly glabrous or 

sparsely pubescent 3. Subfamily Corthylinse. 

a2. Anterior tibia not distinctly broader toward apex or not serrate on 
outer margin. 
b3. Anterior tibia not strongly naiTowed toward apex, apical tooth 
stout; antennal club compressed ; elytra with scales. 

4. Subfamily Micracinse. 
b4. Anterior tibia strongly narrowed toward apex, apical tooth small; 
antennal club thickened at base, with- anterior face obliquely trun- 
cate; elytra with hairs 5. Subfamily Webbinae. 

B. Abdominal sternites 5-7 strongly ascending; tibia broadly compressed, outer 
margin sen-ate •- 6. Subfamily Xyloctoninse. 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA, 225 

II. Pronotum with anterior dorsal area commonly smooth; head exposed or rarely 
concealed from above; tarsi with joint 3 simple or bilobed. 

C. Antenna! club subglobose, compressed to laminate, never conical. 

a3. Body slender to moderately stout; pronotum longer than broad or not 

broader than long 7. Subfamily Crypturginse. 

a4. Body stout; pronotum always broader than long. 

8. Subfamily Phloeotribinse. 

D. Antennal club usually conical, rarely compressed -.9. Subfamily Hylesininse. 

10. Subfamily Phlceoborinae. 

Family SCOLYTIDiE. 

Key to the Subfamilies. 
(See Pis. XII, XV, XVI.) 

I. Pronotum constricted toward the middle; anterior tarsi with joint 3 simple. 

11. Subfamily Coptonotinse. 
II. Pronotum not constricted toward the middle ; anterior tarsi with joint 3 simple or 
bilobed. 

A. Anterior tibia with small tooth on inner apical angle extending beyond the 
tarsal insertion; tibia with at least one tooth on the outer margin, in addition 
to the apical one. 

al. Eyes not divided; tibia with outer margin armed. 

bl. Pronotum with transverse rugosities on anterior area. 

12. Subfamily Hexacolinse. 
b2. Pronotum without transverse rugosities on anterior area. 

13. Subfamily Bothrosterninse. 
a2. Eyes divided; tibia with outer margin unarmed. « 

14. Subfamily Hyorrhynchinse. 

B . Anterior tibia with srnall tooth on inner apical angle not extending beyond the 
tarsal insertion. 

a3. Abdominal sternum convex throughout; antennal scape not very short, 

15. Subfamily Camptocerinse. 
a4. Abdominal sternum not convex throughout; antennal scape very short. 

16. Subfamily Scolytinse. 

Family SCOLYTOPLATYPODIDiE. 
(See Pis. XII, XVI.) 

Anterior tibia with prominent lateral process at outer angle; third tarsal joint 
simple 17. Subfamily Scolytoplatypodinse. 

Family PLATYPODIDiE. 

Key to the Subfamilies. 
(See Pis. XII, XVI.) 

I. Third tarsal joint simple 18. Subfamily Platypodinse. 

II. Third tarsal joint bilobed 19. Subfamily Genyocerinse. 

20. Subfamily Chapuisinse. 

« Uncertain from description whether or not inner angle of anterior tibia is produced 
beyond the tarsal insertion. 



226 



THE SCOLYTID BEETLES. 



POSITION OF THE PRINCIPAL DESCEIBED GENEEA IN THE 
PRELIMINARY CLASSIFICATION. 



Family Ipidae. 

Subfamily Cryphalinae. 
Cosmoderes Eichh. 
Cryphalomorphus Schauff. 
Hypothenemus Westw. 
Stephanoderes Eichh. 
Ernoporus Thoms. 
Trypophlceus Fairm. 
Cryphalops Reitt. 
Cryphaloides Form. 
Cryptarthrum Blndfd. 
Cryphalus Erich. 
Eidophelus Eichh. 
Kyrtogenius Stroh. 
Lepicerus Eichh . 
Lymantor L0v. 
Dendroterus Blndfd. 
Xylodeptes Ferr. 
Taphrorychus Eichh. 
Thamnurgus Eichh. 
Coccotrypes Eichh. 
Ozopemon Haged. 
Bryocoetes Eichh. 
Xyleborus Eichh. 
Anisandrus Ferr. 
Eccoptopterus Motsch. 
Cnestus Sampson. 
Subfamily Ipinae. 

Pityophthorus Eichh. 
Olonthogaster Motsch. 
Acanthotomicus Blndfd. 
Pityogenes Bedel. 
Ips De Geer. 
Subfamily Corthylinse. 
Metacorthylus Blndfd. 
Monarthrum Kirsch. 
Cosmocorynus Ferr. 
Phthorius Eichh. 
Anchonocerus Eichh. 
Glochinocerus Blndfd. 
Tricolus Blndfd. 
Amphicranus Erich. 
Steganocranus Eichh. 
Corthylus Erich. 
Brachyspartus Ferr. 
Gnatliotrichus Eichh. 
Premnobius Eichh. 
Xyloterus Erich . 
Subfamily Micracinse. 
Poecilips Schauf. 
Thysanoes Lee. 



Family Ipidse — Continued. 

Subfamily Micracinee — Continued. 

Micracis Lee. 
Hylocurus Eichh. 
Styracopterus Blndfd. 

Liparthrum WoU. 

Daa'yostactus Schauff. 

Hypohorus Erich. 

Glochiphorus Stroh. 

Cactopinus Schwarz. 
Subfamily Webbinse. 

Webbia n. gen. 
Subfamily Xyloctoninse. 

Scolytogenes Eichh. 

Scolytomimus Blndfd.' 

Scolytodes Ferr. 

Xyloctonus Eichh. 

Gtonoxylon Haged. 
Subfamily Crypturginse. 

Aphanarthrum Woll. 

Triotemnus Woll. 

Crypturgus Eichh. 

Cisurgus Reitt. 

Dolurgus Eichh. 

Dendroctonus Erich. 
Subfamily Phloeotribinse. 

Phlaeophthorus Woll. 

Eulytocerus Blndfd. 

Phloeotribus Latr. 

Dryotomus Chap. 

Renocis Casey. 

Chsetophlceus Lee. 

Chramesus Lee. 

Chortastus Schauff. 

Carphoborus Eichh. 

Cladoctonus Stroh. 

LissoclastiLS Schauf. 

Tiarophorus Schrein. 

Phrixosoma Blndfd. 

Lepisomus Kirby. 

Polygraphus Erich. 

Spongotarsus Haged. 
Subfamily Hylesininse. 

Xylechinus Chap. 

Kissophagus Chap. 

Hylurgus Latr. 

Pachycotes Sharp. 

Tomicus Latr. ( Myelophilus 
Eichh.) 

Hylastinus Bedel. 

Scierus Lee. 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 



227 



Family Ipidae — Continued. 

Subfamily Hylesininee — Continued. 

Hylesinus Fab . 

Pteliohius Bedel. 

Dendrosinus Chap. 

Acanthophorus Stroh. 

Hylurgops Lee. 

Hylastes Ericli. 
Subfamily Phloeoborinse. 

Phloeotrupes Erich. 

Phlmohorus Erich. 

Dactylipalpus Chap. 
Family Scolytidse. 

Subfamily Coptonotinse. 

Craniodicticus BLadfd. 

Microhorus BLadfd. 

Coptonotus Chap. 
Subfamily HexacoUnse. 

Erineophilus Hopk. 

Hexacolus Erich. 

Epomadius Bhidfd. 

Problechilus Eichh. 

Aricerus Blndfd. 

Pycnarthrum Eichh. 

Prionoceles Bhidfd. 

Rhopalopselion Haged. 

Strombo])horus Haged. 

Sphxrotrypes Blndfd. 

Diamerus Erich. 

Bothryperus Haged.- 
Subfamily Bothrosteminse. 

Pagiocerus Eichh. 

Cnesinus Lee. 

Meringopalpus Haged. 



Family Scolytidae — Continued. 

Subfamily Bothrosteminse — Contd. 

Eupagiocerus Blndfd. 

Bothrosternus Eichh . 
Subfamily Camptocerinse. 

Ceratolejns Chap. 

Cnemonys Eichh. 

Camptocerus Dej. 

Loganius Chap. 
Subfamily Scolytinae. 

Scolytopsis Blndfd. 

Scolytus Geoff. 
Family Scolytoplatypodidae. 

Subfamily Scolytoplatypodinae. 

Scolytoplatypus Schauff. 

Spongocerus BLadfd. 

Txniocerus Blndfd. 
Family Platypodidee . 

Subfamily Platypodinse. 

Spathidiceres Chap. 

Periommatus Chap. 

Tesserocerus Saund. 

Symmerus Chap. 

Cenocephalus Chap. 

Mitosovia Chap. 

Diapus Chap. 

Cylindropalpus Stroh. 

Platypus Herb St. 

Crossotarsus Chap. 
Subfamily Genyocerinse. 

Genyocerus Motsch. 
Subfamily Chapuisinse. 

Chapuisia Duges. 



GENERA DESCRIBED IN 1911-12, NOT INCLUDED IN FORE- 
GOING LIST, BUT HERE PROVISIONALLY ASSIGNED TO THE 
SUBFAMILIES. 

Cyclorhipidion Hagedorn Cryphalinae. 

Xyleborites Wickham Do. 

Pseudothamnurgus Eggers Do. 

Neotomicus Fuchs Ipinae. 

Xestips Hagedorn Do . 

Pityokteines Fuchs Do. 

Trigonogenius Hagedorn Do . 

Allarthrum Hagedorn Micracinse. 

Acacicis Lea Hylesininse. 

Hapalogenius Hagedorn Do . 

Hylesinosoma Lea Do. 

Chsetophorus Fuchs Do. 

Fields Lea Do . 

Minulus Eggers Hexacolinae. 

Mesoplatypus Strohmeyer Platypodinse. 

Notoplatypus Lea Do. 



228 THE SCOLYTID BEETLES. 

GENERA OF DOUBTFUL POSITION. 

Mesoscolytus Blndfd. Styphlosoma Blndfd. 

Phthorophlceus Rey. Aaantus Broun. 

Hylosq/llus Schauff. Inosomus Broun. 

Dendrotrupes Broun. Araptus Eichh. 



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1802. Marsham, T. — Entomologia Britannica, sistens insecta Britanniae indigena 

secundum methodum Linnaeanam disposita, Tom. I, Coleoptera, pp. 51-60. 
1802. Latreille, P. A. — Histoire naturelle, generale et particuliere des crustacee 
et des insectes, Vol. 3, p. 204. 

229 



230 THE SCOLYTlD BEETLES. 

1802. Illiger, J. K. W. — Zusatze, Berichtigungen und Bemerkungen zu Fabricii 

Systema Eleutheratorum, Tomus. I. Magazin fiir Insektenkunde, Bd. 1, 

pp. 306-425, with special reference to p. 367. 
1807. Illiger, J. K. W. — Vorschlag zur Aufnahme im Fabricischen Systeme fehlen- 

der Kafergattungen. Magazin fiir Insektenkunde, Bd. 6, pp. 318-349, with 

special reference to p. 367. 
1807. Latreille, P. A. — Genera crustaceorum et insectorum secundum ordinem 

natui'alem in familias disposita, iconibus exemplisque plurimus explicata, 

Vol. 2, pp. 273-280. 
1813. Gyllenhal, L. — Insecta suecica descripta, Tom. I, Pars. Ill, pp. 335-372. 
1824. Curtis, John — British entomology, Vol. I, No. 43. 
1834. Westwood, J. 0. — Description of a minute coleopterous insect, forming the 

type of a new subgenus allied to Tomicus, with some observations upon the 

affinities of the Xylophaga. Trans. Ent. Soc. London, Vol. I, Pt. I, pp. 

34-36, PL VII, figs, la-lh. 

1836. Erichson, W. F. — Systematische Auseinandersetzung der Familie der Bor- 

kenkafer (Bostrichidse). Archiv fiir Naturgeschichte v. A. F. Wiegmann, 
Jahrg. II, Bd. I, pp. 45-65. 

1837. Ejeby, W. — Fauna Boreali-Americana; or the zoology of the northern parts 

of British America. Part 4. The insects, pp. 191-195. 
1839. Ratzeburg, J. T. C— Die Forst-insecten, Th. I, pp. 168-232, Pis. 7-15. 
1842. HoENUNG, A. — Ueber einige in den Betelniissen vorkommende Kafer. Stettin 

Ent. Ztg., Jahrg. 3, pp. 115-117. 
1859. Thomson, C. G. — Skandinaviens Coleoptera synoptiskt bearbetade, Tom. I, 

pp. 146-147. 
1864. Perroud, B.P.,& Montrousier, A. — Essai sur la faune entomologique de 

Kanala (Nouvelie-Caledonie) et description de quelques especes nouvelles 

ou peu connues. Ann. Soc. Linn. Lyon, Tom. II, pp. 46-257, PI. I. See 

p. 188. 

1864. EiCHHOFF, W. — Ueber die Mundtheile und die Fiihlerbildung der em'opaischen 

Xylophagi sens strict. Berlin Ent. Ztschr., Jahrg. 8, pp. 17-48. 

1865. Thomson, C. G. — Skandinaviens Coleoptera synoptiskt bearbetade, Tom. 7, 

pp. 345-378. 

1866. Lacordaire, J. T. — Histoire natm'elle des insectes. Genera des coleopteres, 

Tom. 7, pp. 358-376. 
1868. Jacquelin Du Val, P. N. C, & Fairmaire, L. — Manuel entomologique. 

Genera des coleopteres d'Europe, Vol. 4, pp. 97-108, Pis. 31-34. 
1868. Zimmermann, C. — Synopsis of the Scoh'tidse of America north of Mexico. 

Trans. Amer. Ent. Soc, Vol. II, pp. 141-149. 

1868. Le Conte, J. L. — Appendix and notes to Zimmermann, C. Synopsis of Scoly- 

tidse of America north of Mexico. Trans. Amer. Ent. Soc, Vol. II, pp. 
150-178. 

1869. Chapuis, F. — Synopsis des scolytides. Mem. Soc. Sci. Liege, Ser. 2, Vol. 3, 

pp. 213-269 (author's extra issued 1869). 

1871. Eichhoff, W. — Neue exotischeTomiciden-arten. Berlin Ent. Ztschr., Jahrg. 

15, pp. 131-137. 

1872. Gemminger, M., & Harold, B. de. — Catalogus Coleopterorum, Vol. 9, pp. 

2669-2702. 

1875. LiNDEMANN, C. — Vergleicheud anatomische Untersuchung iiber das msenn- 

liche Begattungsglied der Borkenkafer. Bui. Soc. Imp. Nat. Moscou, Tom. 
49, No. 1, pp. 196-252, Pis. I-V. 

1876. Le Conte, J. L. — The Rhynchophora of America north of Mexico. Scolytidse. 

Proc. Amer. Phil. Soc, Vol. 15, pp. 341-391. 



PRELIMINARY CLASSIFICATION OF SCOLYTOIDEA. 231 

1876. LiNDEMANN, C. — ^Monographie der Borkenkafer Russlands. Die crypha- 
loiden Tomiciden. Bui. Soc. Imp. Nat. Moscou, Tom. 51, No. 4, pp. 
320-380, figs. 6-55. 

1878. EiCHHOFF, W. — Ueber die Borkenkafer-Gattungen Hylm'gus Latr. und Blasto- 

phagus Eich. Stettin Ent. Ztg., Jahrg. 39, pp. 399-400. 

1879. EiCHHOFP, W. — Ratio, descriptio, emendatio eorum Tomicinorum, p. 165. 

1880. RuPERTSBERGER, M. — Biologie der Kilfer Europas, Scolytidae, pp. 224-232. 

1881. EiCHHOPF, W. J. — Die europaischen Borkenkafer, pp. 315, figs. 104. 

1884. Fauvel, a. — Sur I'identite des genres Hypothenemus, Stephanoderes et 

Homoeocryphalus. Revue d'entomologie, Tom. 3, p. 315. 
1886. Thomson, C. G. — [Note on Polygraphus grandiclava Thorns.] Bui. Soc. 

Ent. France, p. LXII. 
1888. Bedel, L. — Faune des coleopteres du Bassin de la Seine, Tom. VI, pp. 385-421. 
1894. Hopkins, A. D. — Sexual characters in Scolytidee. Canad. Ent., Vol. 26, 

pp. 274-280, October. 
1894. Reitter, E. — Bestimmungs-tabelle der Borkenkafer (Scolytidse) aus Em-opa 

luid den angrenzenden Landern, Hft. 31, pp. 40-97. 

1894. RuPERTSBERGER, M. — Die biologische Literatur fiber die Kafer Europaa von 

1880 an, mit Nachtragen aus fruherer Zeit und einem Larven-Cataloge, pp. 
236-243. 
1895-1905. Blandford, W. F. H. — Scolytidse. Biologia Centrali-Americana. 
Insecta. Coleoptera, Vol. 4, Pt. 6, pp. 81-298, Pis. IV-IX. 

1895. JuDEiCH, J. F., & NiTSCHE, H. — Lehrbuch der mitteleiiropaischen Forstin- 

sektenkunde. Vol. I, pp. 435-556, figs. 142-178. 

1896. Verhoeff, C. — Ueber das Abdomen der Scolytiden, ein Beitrag zur verglei- 

chenden Morphologic des Hinterleibes der Coleopteren. Archiv fur 
Natm-geschichte, Jahrg. 62, Bd. I, pp. 109-144, figs. A-D. 

1897. Hubbard, H. G. — The ambrosia beetles of the United States. Bui. 7, n. s., 

Div. Ent., U. S. Dept. Agr., pp. 9-30, figs. 1-34. 

1898. Hopkins, A. D. — On the history and habits of the "wood engraver" ambrosia 

beetle — Xyleborus xylographus (Say), Xyleborus saxeseni (Ratz.) — with 
brief descriptions of different stages. Canad. Ent., Vol. 30, pp. 21-29, Pis. 
2-3, January. 

1899. Hopkins, A. D. — Report on investigations to determine the cause of unhealthy. 

conditions of the spruce and pine from 1880-1883. Bui. 56, W. Va. Agr. Exp. 
Sta., pp. 197-461, figs. XGIX, April. 

1901. Barbey, Augusts. — Les scolytides de I'Europe centrale. pp. 121, Pis. 18. 

1902. Sedlaczek, W. — Ueber den Darmcanal der Scolytiden. Centralblatt flir das 

gesamte Forstwesen, Jahrg. 28, Hft. 6, pp. 241-263, fold. PL, Juni. 

1903. Ganglbauer, L. — Systematisch-koleopterologische Studien. Miinchener 

koleopterologische Zeitschrift, Vol. I, pp. 271-319. 

1905. Hopkins, A. D. — Notes on scolytid larvae and their mouth parts. Proc. Ent. 

Soc. Wash., Vol. 7, No. 2-3, pp. 143-149, Pis. 4-5, October. 

1906. Heyden, L. von, Reitter, E., & Weise, J. — Catalogus Coleopterorum 

Europae, Caucasi et Armenias rossicae. Ed. 2, column 707-714. 

1907. Tredl, R. — Nahrungspflanzen und Verbreitungsgebiete der Borkenkafer 

Em-opas. Entomologische Blatter, Bd. 3, pp. 2-4, 18-22, 37-42, 53-56, 69- 
72, 87. 

1908. FoRMANEK, Roman. — Eine neue Borkenkafer-gattung. Entomologische Blat- 

ter, Jahrg. 4, p. 91, May 15. 

1909. Hopkins, A. D. — Contributions toward a monograph of the scolytid beetles. 

I. The genus Dendroctonus. Tech. Ser. No. 17, Pt. I, Bur. Ent., U. S. Dept. 
Agr., pp. 1-164, figs. 1-95, Pis. I-VIII, June 30. 



232 THE SCOLYTID BEETLES. 

1909. SwAiNE, J. M. — Catalogue of the described Scolytidse of America north of 

Mexico. 24th Report of New York State Entomologist. Appendix B. 
New York State Education Dept. Bui. 455, New York State Museum, 
Museum Bui. 134, September. 

1910. Hagedorn, M. — Ipid^. Schenkling, S., ed. Coleopterorum Catalogus, Para. 

4, p. 134. 
1910a. Hagedorn, M. — Coleoptera. Fam. Ipidse. Wytsman, P. Genera Insectorum, 

fasc. Ill, pp. 178, Pis. 14. 
1910. Newberry, E. A. — On Hypothenemus eruditus, Westwood. Ent. Mo. 

Mag., Vol. 46 (Ser. 2, Vol. 21), pp. 83-84, April. 

1910. Lea, A. M. — On Australian and Tasmanian Coleoptera, with descriptions of 

new species. Part I. Proc. Roy. Soc. Victoria, Vol. 22 (new ser.), Part II, 
pp. 133-150. 

1911. Hopkins, A. D. — Contributions toward a monograph of the bark weevils of 

the genus Pisodes. Tech. Ser. No. 20, Pt. I, Bur. Ent., U. S. Dept. Agr., 
pp. 1-68, figs. 1-9, Pis. I-XXII, January 7. 
1911; NiJssLiN, 0. — Phylogenie und System der Borkenkafer. Ztschr. Wiss. Insek- 
tenbiol., Bd. 7, No. 1-12. 

1911. FucHS, G. Morphologische Studien iiber Borkenkafer I. Die Gattungen Ipa 

De Geer und Pityogenes Bedel, pp. 1-45, figs. 1-39. 

1912. NtJssLiN, 0. — Phylogenie und System der Borkenkafer. Ztschr. Wiss. 

Insektenbiol., Bd. 8, No. 1-7. 
1912. FucHS, G. — ^Morphologische Studien Tiber Borkenkafer. II. Die euro- 

paischen Hylesinen, pp. 1-13, figs. 1-85, 3 fold. pis. 
1912. Sharp, D., & Muir, F. — The comparative anatomy of the male genital tube 

in Coleoptera. Proc. Ent. Soc. London, Pt. Ill, pp. 477-642, Pis. XLII- 

LXXVIII, December. 

Note. — The next contribution toward this monograph, entitled "Preliminary 
Classification of the subfamily Cryphalinse, with descriptions of new genera and 
species," will be published as Report No. 99, Office of the Secretary, U. S. Depart- 
ment of Agriculture. 



ADDITIONAL COPIES 

OF THIS PUBLICATION MAT BE PROCUEED FROM 

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Technical Series, No. 18. 

U. S. DEPARTMENT OF AGRICULTURE, 

BTJRE^kXJ OF EISTTOMIOLOO-Y. 

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

C^ 



THE ANATOMY OF THE HONEY BEE. 



R. E. SNODGRASS, 

Agent and Expert. 



Issued May 28, 1910. 




WASHINGTON: 

GOVERNMENT PRINTING OFFICE. 
1910. 



s rs. 



\<^\< 



BUREAU OF ENTOMOLOGY. 

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

C. L. Marlatt, Assistant Entomologist and Acting Chief in Absence of Chief. 

R. S. Clifton, Executive Assistant. 
W. F. Tastet, Chief Clerk. 

F. H. Chittenden, in charge of truck crop and stored prod^ict insect investigations. 

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

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

F. M. Webster, in charge of cereal and forage insect investigations. 
A. L. Quaintance, in charge of deciduous fruit insect investigations. 
E. F. Phillips, in charge of bee culture. 

D. M, Rogers, in charge of preventing spread of moths, field work. 
RoLLA P. Cureie, in charge of editorial loork. 

Mabel Colcord, Uorarian. 

Investigations in Bee Cltlture. 
E. F. Phillips, in charge. 

G. F. White, J. A. Nelson, B. N. Gates, R. E. Snodgrass, A. H. INIcCkay, agents 
and experts. 

Ellen Dashiell, preparator. 
Jessie E. Marks, clerk. 
T. B. Symons, collaborator for Maryland. 
H. A.. Surface, collaborator for Pennsylvania. 
J. C. C. Price, collaborator for Virginia. 
2 



LETTER OF TRANSMITTAL 



U. S. Department of Agriculture, 

Bureau of Entomology, 
Washington, D. (7., October 19, 1909. 
Sir: I have the honor to transmit herewith a manuscript entitled 
" The Anatomy of the Honey Bee," by Mr. R. E. Snodgrass, agent 
and expert, of this Bureau. It embodies the results of detailed 
studies made by Mr. Snodgrass and should prove of value as bring- 
ing to the bee keeper reliable information concerning an insect of 
such great economic importance, and also as furnishing a sound 
basis in devising new and improved practical manipulations. I 
recommend its publication as Technical Series, No. 18, of the Bureau 
of Entomologj^ 

Eespectfully, L. O. Howard, 

Entomologist and Chief of Bureau. 
Hon. James Wilson, 

Secretary of Agriculture. 

3 



CONTENTS. 

Pag-e. 

I . Introduction 9 

II. General external structure of insects. 10 

III. The head of the bee and its appendages 26 

1. The structure of the head 26 

2. The antennae and their sense organs 32 

3. The mandibles and their glands 39 

4. The proboscis 43 

5 . The epipharynx 51 

IV. The thorax and its appendages - 53 

1 . The structure of the thorax 53 

2. The wings and their articulation .59 

3. The legs 66 

V. The abdomen, wax glands, and sting 69 

VI. The alimentary canal and its glands 84 

1. The general physiology of digestion, assimilation, and excretion. 84 

2. The salivary glands 87 

3 . The alimentary canal 90 

VII. The circulatory system 107 

VIII. The respiratory system 112 

IX. The fat body and the oenocytes 119 

X. The nervous system and the eyes - 122 

XI . The reproductive system 130 

1. The male organs 132 

2 . The female organs 134 

Explanation of the symbols and letters used on the illustrations 139 

Bibliography 148 

Index 151 

5 



ILLUSTRATIONS, 



Page. 

Fig. 1. Median longitudinal section of body of worker 8 

2. Diagram of generalized insect embryo 12 

3. Example of generalized insect mouth parts 17 

4. Diagram of generalized thoracic segment 19 

5. Typical insect leg 21 

6. Diagram of generalized insect wing and its articulation 22 

7. Diagram of terminal abdominal segments of a female insect and early 

stage in development of gonapophyses 25 

8. Example of a swordlike ovipositor 7 25 

9. Head of worker bee 27 

10. Heads of worker, queen, and drone 29 

11. Median longitudinal sections of heads of worker and drone 30 

12. Antennal hairs and sense organs 36 

13.- Mandibles of worker and drone , 40 

14. Internal mandibular gland of worker 42 

15. Mouth parts of worker 43 

16. Median section through distal half of mentum and base of ligula of 

worker 50 

17. Epipharynx and labrum of worker 51 

18. Sense organs of epipharynx 52 

19. Median longitudinal section of head of worker 52 

20. Dorsal view of ventral walls of body of worker 53 

21. Thorax of worker 54 

22. Lateral view of mesotergum of worker 56 

23. Thoracic terga of worker 57 

24. Upper part of left mesopleurum of worker 58 

25. Wings of Hymenoptera _ 60 

26. Basal elements of wings of Hymenoptera 61 

27. Median section through thorax of drone 64 

28. Internal view of right pleurum of mesothorax of drone 65 

29. Legs of worker, queen, and drone 67 

30. Claws and empodium of foot of worker 68 

31. Tai'sal claws of worker, queen, and drone 69 

32. Lateral view of abdomen of worker 70 

33. Ventral view of abdomen of worker 70 

34. Dorsal view of abdominal sterna of drone 70 

35. Sixth abdominal sternum of worker, queen, and drone 72 

36. Semidiagrammatic view of left side of sting of worker 75 

37. Ventral view of sting of worker 76 

38. Section of small piece of wall of poison sac 79 

39. Sections of alkaline gland of sting 79 

40. Details of sting of worker 81 

41. Tip of abdomen of worker with left side removed 82 

6 



ILLUSTRATIONS. 7 

Page. 

Fig. 42. Alimentary canal of worker 85 

43. Details of pharyngeal and salivary glands 88 

44. Honey stomach of worker, queen, and drone 94 

45. Longitudinal section of honey stomach and proventriculus of queen . 97 

46. Histological details of alimentary canal of worker 103 

47. Dorsal diaphragm of drone, from one segment 108 

48. Small part of dorsal diaphragm of drone 110 

49. Pericardial chamber of one segment in worker Ill 

50. Tracheal system of worker 113 

51. Tracheal system of worker 117 

52. Nervous system of worker 123 

53. Brain and sub oesophageal ganglion of worker 125 

54. Horizontal section of compound eye and optic lobe of worker 127 

55. Histological details of compound eye of worker 128 

56. Reproductive organs of drone 133 

57. Reproductive organ and sting of queen 135 



THE ANATOMY OF THE HONEY BEE. 



I. INTRODUCTION. 

The anatomy of the honey bee has been for years a subject of much 
interest to those engaged in bee keeping both for pleasure and for 
profit. This interest is due not only to a laudable curiosity to know 
more of the bee, but to the necessity of such information in order 
to understand fully what takes place in the colony. All practical 
manipulations of bees must depend on an understanding of the be- 
havior and physiology of bees under normal and abnormal circum- 
stances, and those bee keepers who have advanced bee keeping most 
by devising better manipulations are those, in general, who know 
most of bee activity. In turn, a knowledge of bee activity must rest 
largely on a knowledge of the structure of the adult bee. 

Studies on the anatomy of the bee have not been lacking, for 
many good workers have taken up this subject for investigation. 
The popular demand for such information, however, has induced 
untrained men to write on the subject, and most accounts of bee 
anatomy contain numerous errors. This is probably to a greater 
extent true of the anatomy of the bee than of that of any other 
insect. Frequently the illustrations used by men not trained in 
anatomical work are more artistic than those usually found in papers 
on insect anatomy, and they consequently bear the superficial marks 
of careful work, but too often it is found that the details are in- 
accurate. It has therefore seemed the right time for a new presenta- 
tion of this subject based on careful work. 

The drawings given in the present paper are original, with the 
exception of figures 12, 54, and 55, and have been prepared with 
a thorough realization of the need of more accurate illustrations of 
the organs of the bee, especially of the internal organs. Mistakes 
will possibly be found, but the reader may be assured that all the 
parts drawn were seen. Most of the dissections, moreover, were 
verified by Dr. E. F. Phillips and Dr. J. A. Nelson, of this Bureau, 
before the drawings were made from them. An explanation of the 
abbreviations and lettering is given on pages 139-147. 

It is hoped that the work will furnish the interested bee keeper 
with better information on the anatomy of the bee than has hereto- 
fore been offered to him, that it may provide a foundation for more 
detailed work in anatomy and histology, and, finally, that it will be 



10 THE AKATOMY OP THE HONEY BEE. 

of service to future students of the embryology and pliysiologj^ of 
the bee. With this last object in view the writer has tried to sum 
up under each heading the little that is at present known of insect 
physiology in order to bring out more clearly what needs to be done 
in this subject. 

II. GENEHAL EXTERNAL STRUCTURE OF INSECTS. 

"Wlien w^e think of an animal, whether a bee, fish, or dog, we uncon- 
sciously assume that it possesses organs which perform the same vital 
functions that we are acquainted with in ourselves. We know, for 
example, that an insect eats and that it dies when starved ; we realize 
therefore that it eats to maintain life, and we assume that this involves 
the possession of organs of digestion. We know that most insects see, 
smell, and perform coordinated actions, and we recognize, therefore, 
that they must have a nervous system. Their movements indicate to 
us that they possess muscles. These assumptions, moreover, are en- 
tirely correct, for it seems that nature has only one way of producing 
and maintaining living beings. No matter how dissimilar two 
animals may be in shape or even in fundamental constitution, their 
life processes, nevertheless, are essentially identical. Corresponding- 
organs may not be the same in appearance or action but they accom- 
plish the same ends. The jaws may work up and down or they may 
work siclewise, but in either case they tear, crush, or chew the food 
before it is swallowed. The stomach may be of very different shape 
in two animals, but in each it changes the raw food into a soluble and 
an assimilable condition. The blood may be red or colorless, con- 
tained in tubes or not, but it always serves to distribute the prepared 
food which diffuses into it from the alimentary canal. The situa- 
tion of the central nervous system and the arrangement of its parts 
may be absolutely unlike in two organisms, but it regulates the func- 
tions of the organs and coordinates the actions of the muscles just 
the same. . 

Hence, in studying the honey bee we shall find, as we naturally 
expect to find, that it possesses mouth organs for taking up raw food, 
an alimentary canal to digest it, salivary glands to furnish a digestive 
liquid, a contractile heart to keep the blood in circulation, a respira- 
tory system to furnish fresh oxygen and carry off waste gases, ex- 
cretory organs for eliminating waste substances from the blood, a 
nervous system to regulate and control all the other parts, and, finally, 
organs to produce the reproductive elements from which new indi- 
viduals are formed to take the places of those that die. 

The study* of anatomy or the structure of the organs themselves 
is inseparably connected with a study of physiology or the life 
functions of the animal. While physiologj^ is a most interesting 
and important subject, and, indeed, in one sense might be said to be 



GENEEAL EXTERNAL STRUCTURE OF INSECTS. H 

the object of all anatomical research, yet the mere study of the 
structure of the organs alone, their wonderful mechanical adapta- 
tions, and their modifications in different animals forms a most fasci- 
nating field in itself, and besides this it gives us an insight into the 
blood relationships -and degrees of kinship existing between the 
multitudes of animal forms found in nature. In the study of com- 
parative anatomy we are constantly surprised to find that structures 
in different animals which at first sight appear to be entirely differ- 
ent are really the same organs which have been simply changed in 
a superficial way to serve some new purpose. For example, the 
front wing of a bee and the hard shell-like wing cover of a beetle are 
fundamentally the same thing, both being front wings — that of the 
beetle being hardened to serve as a protection to the hind wing. 
Again, the ovipositor of a katydid and the sting of a bee are identical 
in their fundamental structure, differing in details simply because 
they are used for different purposes. Hence, in the study of anat- 
omy we must always be alert to discover what any special part cor- 
responds with in related species. In order to do this, however, it 
is often necessary to know the development of an organ in the 
embryo or in the young after birth or after hatching, for many 
complex parts in the adult have very simple beginnings in an imma- 
ture stage. 

Thus it becomes evident that the structural study of even one 
organism soon involves us in the subjects of anatomy, physiology, 
and embryology, and, if we add to this a study of its senses, its 
behavior, and its place in nature, the field enlarges without limit. 
The student of the honey bee realizes that a lifetime might be spent 
in exploiting this one small insect. 

The differences between animals are much greater on the outside 
than on the inside. In the descriptions of the organs of the honey bee 
anyone will know what is meant by the " alimentary canal," the 
" nervous system," or the " respiratory system," but the external 
parts are so different from those of animals with which we are more 
familiarly acquainted that no general reader could be expected to 
know what is meant by the names applied. Moreover, the bee and its 
allies are so modified externally in many ways that, at first sight, 
their parts look very different even from those of other insects. 
Hence, we shall give a preliminary account of the external structure 
of insects in general, for it is hoped that the reader will then more 
easily understand the special structure of the honey bee, and that the 
application of the terms used will appear more reasonable to him. 

Since all animals originate in an egg, the change into the adult 
involves two different processes: One is growth, which implies 
merely an increase in size, the addition of material to material; the 
other is development, which means change in shape and the produc- 



12 



THE ANATOMY OF THE HOISTEY BEE. 



tion of a form with complex organs from the simple protoplasmic 
mass of the egg. The part of development that takes place in the 
eggshell is known as embryonic develo-pment ; that which takes place 
subsequent to hatching is known as postembryonic development. In 
insects there are often two stages in the postembryonic development, 
an active one called the larval stage and an inactive one called the 
pupal stage. During the first of these the young insect is termed a 
larva; during the second, a pupa. When there is no resting stage the 
immature creature is often called a nymph. The final and fully de- 
veloped form is an adult, or imago. 

Since this paper is to deal only with the anatomy of the adult, the 
attractive fields of embryonic and postembryonic development must 
be passed over, except for a few statements on 
fundamental embryonic structure, a knowledge 
of which is necessary to a proper understanding 
of the adult anatomy. 

When the embryo, in its course of development, 
first takes on a form suggestive of the definitive 
insect, it consists of a series of segments called 
metameres., or somites, and shows no differentia- 
tion into head, thoracic, and abdominal regions. 
Typically, each segment but the first is provided 
with a pair of latero-ventral appendages, hav- 
ing the form of small rounded protuberances. 
These appendages are of different sizes and take 
on different shapes in different parts of the 
body, for some of them are destined to form the 
antennae, some the mouth parts, others the legs 
and perhaps the cerci, while the rest of them 
remain very small and finally disappear. A^^iat 
we know of the embryology of insects is based 
on the observations of a number of men who 
have worked mostly on the development of dif- 
ferent species. Their observations are not all 
alike, but this is probably due in large part to the fact that the 
embryos of different insects are not all alike. Embryos have a very 
provoking habit of skipping over or omitting little and yet im- 
portant things m their development, but fortunately they do not 
all omit the same things. Therefore, by putting together all the 
reliable information we possess, we can make up an ideal embryo 
which would be typical of all insects. Such a generalized embryo is 
represented diagrammatically by figure 2. 

The first six or seven metameres very early begin to unite with 
one another and continue to fuse until their borders are lost. These 
consolidated embryonic segments form the head of the adult insect. 




Pig. 2. — Diagram of a 
generalized insect em- 
bryo, showing the seg- 
mentation of the head, 
thoracic, and abdom- 
inal regions, and the 
segmental appendages. 



GENERAL EXTERNAL STRUCTUEB OF INSECTS, 13 

Observers differ concerning the fate of the seventh segment, but it 
is most ]Drobable that a part of it fuses with the sixth segment, thus 
taking part in the formation of the head, and that a part of it forms 
the neck or some of the neck plates of the adult. 

The appendages of these first seven segments form the antennae 
and mouth parts, except one or two pairs that disappear early in 
embryonic life. It is not certain that the first segment ever possesses 
appendages, but from it arise the large compound eyes and appar- 
ently also the upper lip, or labrum {Lin). The appendages of the 
second segment form the feelers, or antennae {lAnt) of the adult, 
those of the third {2Ant) disappear in insects, but they correspond 
with the second antennae of shrimps and lobsters. The appendages 
of the fourth segment form the mandibles (Md). Those of the 
fifth segment (SUn), when present, fuse with a median tonguelike 
lobe (Lin) of the following segment, and the three constitute the 
hypopharynx, or lingua of the adult. The next pair {IMx) form the 
maxillae, while the last (^ilfa?), or those of the seventh segment, 
coalesce with each other and constitute the adult labium, or lower lip. 

The bodies of the head metameres fuse so completely that it is 
impossible to say positively what parts of the adult head are formed 
from each. The last, as already stated, possibly takes part in the 
formation of both the head and the neck. Some embryologists at- 
tribute the plates which usually occur in this region to the last em- 
bryonic head segment, while others believe they come from the next 
segment following. Sometimes these plates are so well developed 
that they appear to constitute a separate segment in the adult, and 
this has been called the microthorax. If this name, however, is 
given to the embryonic segment from which these plates are said to 
be derived, it must be remembered that it is not " thoracic " at all 
and belongs partly to the head. The name cervicunh has been ap- 
plied to the neck region with greater appropriateness since it does 
not imply any doubtful affiliation with adjoining regions. What 
we really need, however, is not so much a name as more information 
concerning the development of the rear part of the head and the 
neck plates in different insects. 

The next three segments remain distinct throughout life in nearly 
all insects, but, since they bear the legs and the wings, the}^ become 
highly specialized and together constitute the thorax. The indi- 
vidual segments are designated the fvothorax^ the mesotTiorax^ and 
the metathorax. The legs are formed from the embryonic ap- 
pendages (fig. 2, iZ, ^Z, 3L) of these segments, but the wings are 
secondary outgrowths from the mesothorax and metathorax and 
are, hence, not appendages in the strict embryological sense. 

The remaining segments, nearly always 10 in number, constitute 
the abdomen. The appendages of these segments, except possibly 



14 THE ANATOMY OF THE HONEY BEE. 

those of the tenth, disappear early in embryonic life in all insects, 
except some of the very lowest species, in which they are said to form 
certain small appendages of the abdominal segments in the adults. 

An adult insect is often described as being " divided " into a head, a 
thorax, and an abdomen, but this is not true in most cases. While all 
insects consist of these parts, the divisions of the body are usually 
not coincident with them. The prothorax in the adult is separated 
from the head by the neck and is very commonly separated from the 
mesothorax by a flexible membranous area. On the other hand, the 
mesothorax and metathorax are almost always much more solidly at- 
tached to each other, while, in most insects, the mgtathorax is solidly 
and widely joined to the first abdominal segment, though in the flies 
these latter two segments are usually separated b}^ a constriction. In 
such insects as ants, wasps, and bees a slender, necklike peduncle 
occurs between the first and second segments of the abdomen, the 
first being fused into the metathorax so that it appears to be a part 
of the thorax. This is the most distinctive character of the order 
Hymenoptera, to which these insects belong. 

The body wall of insects is hard on account of the thick layer of 
chitin which exists on the outer side of the true skin. Chitin is a sub- 
stance similar to horn, being brittle, though tough and elastic. It 
gives form and rigidity k) the body and affords a solid attachment for 
the muscles within, since insects have no internal framework of bones 
such as vertebrate animals have. The skin between the segments is 
soft and unchitinized and thus forms a flexible intersegmental mem- 
hrane which is often very ample and, in the abdomen, allows each seg- 
ment to telescope into the one in front of it. 

The chitin of each segment is not continuous, but is divided into 
plates called sclentes. The most important of these are a tergum 
above and a sternum below, but, in the case of the thorax, these two 
plates are separated on each side by another called the pleurum^ which 
lies between the base of the wing and the base of the leg. Pleural 
plates are sometimes present also on the abdominal segments. These 
principal segmental plates are usually separated by membranous 
lines or spaces, which permit of more or less motion between them. 
Such lines are called sutures in entomology, though strictly this term 
should be applied only to the lines of fusion between adjoining parts. 

The terga, pleura, and sterna of each segment are furthermore 
subdivided into smaller sclerites, which may be termed tergites^ pleu- 
rites, and ste'niites, respectively. The sutures between them are 
sometimes membranous also, but most frequently have the form of 
impressed lines or narrow grooves. In such cases they are generally 
nothing more than the external marks of ridges developed on the 
inside of the body wall to strengthen the parts or to give attachment 
to muscles. Since these sutures are conspicuous marks on the outside 



GENERAL EXTEENAL STEUCTUEE OF INSECTS. 15 

of an insect, tliey are usually regarded as morphologically impor- 
tant things in themselves, representing a tendency of the tergum, pleu- 
rum, or sternum to separate into smaller plates for some reason. The 
truth about them would appear to be just the opposite in most cases — 
they are the unavoidable external marks of an internal thickening 
and strengthening of the plates. In a few cases they may be the 
confluent edges of separate centers of chitinization. Hence, most of 
the sutural lines in insects appear to signify a bracing or solidifying 
of the body wall rather than a division of it. 

Since the body wall of insects is continuous over all the surface it 
contains no articulations of the sort that occur between the bones in 
the skeleton of a vertebrate. Although insects and their allies be- 
long to the class of animals known as the Articulata, yet an articu- 
late articulation is simply a flexibility — two chitinous parts of the 
exoskeleton are movable upon each other simph^ by the intervention 
of a nonchitinized, flexible, membranous part. ^¥liile there are often 
special ball-and-socket joints developed, these are always produced 
on the outside of the membranous hinge and simply control or limit 
the movement of the articulation. 

The head of an adult insect is a thin-walled capsule containing the 
brain, the ventral head ganglion of the nervous system, the pharynx 
and anterior part of the oesopliagus, the tracheal tubes, and the 
muscles that move the antennae and the mouth parts. Its shape varies 
a great deal in different insects, being oval, globular, elongate, or 
triangular. In some it is flattened dorso-ventrally so that the face is 
directed upward and the mouth forward, but in most, including 
the bee, it is flattened antero-posteriorly so that the face looks for- 
ward and the mouth is directed ventrally. In a few it is turned so 
that the face is ventral. The walls of the head are usually divided 
by sutures into a number of sclerites, which in general are located 
and named as follows: The movable transverse flap forming the 
upper lip is the Idbrum. Above it is a sclerite called the clypeus^ 
which is a part of the solid wall of the head and carries the anterior 
articulations of the mandibles. The clypeus is sometimes divided 
transversely into an anteclypeus (" clypeus anterior," " epistoma ") 
and into a post-clypeus ("clypeus posterior"). Above the clypeus 
is the front., a plate usually occupying the upper half of the face 
between the compound eyes and carrying the antennae. The top of 
the head is called the vertex., but does not constitute a separate scle- 
rite. The sides of the head below the compound eyes are often sepa- 
rated by sutures from the anterior and posterior surfaces and are 
known as the genae. The back of the head is formed by the occiput. 
which surrounds the large opening or foramen magnurrh that leads 
from the cavity of the head into that of the neck. The parts pos- 
terior to the gense, carrying the posterior mandibular articulations, 



16 THE ANATOMY OF THE HONEY BEE. 

are sometimes separated from both the occiput and the gense and are 
known as the postgena?. In a few insects, especially beetles, one or 
two median plates occur in the ventral wall of the head posterior to 
the base of the labium. These are the gular sclerites. Finally, small 
plates are sometimes found about the bases of the antennae and be- 
tween the bases of the mandibles and the gense. The latter have 
been termed the trochantins of the mandibles. The term epicranium 
is often used to include all the immovable parts of the head, but is 
frequently applied only to the dorsal parts. Most of these sclerites 
preserve a pretty definite arrangement in the different orders, and 
they are probably homologous throughout the entire insect series, 
though they are in some cases very much distorted by special modi- 
fications and are often in part or wholly obliterated by the disap- 
pearance of the sutures. Embryologists are coming to the conclu- 
sion that the sclerites of the head have no relation to the primitive 
segments. The latter very earlj^ consolidate into a head with a con- 
tinuous wall, while the sutures defining the sclerites are formed 
later. Some of the older entomologists were led, from a study of 
the sclerites, to suppose that the head consisted of a number of seg- 
ments, but it has been shoAvn that these anatomical segments do not 
correspond with the embryonic ones. 

The appendages growing from the front of the face are the 
antennae (fig. 9A, Ant) or " feelers " and consist of a series of joints 
or segments. 

At the lower edge of the face is the front lip or labrinn (fig. 9 A, 
Zm) , behind which are the median epipharynx^ the paired mandihles 
{Md) and maxillce, the median hypopharynx^^ and the labium or under 
lip. All these organs together constitute what are known as the 
mouth parts or trophi. They vary greatly in shape and appearance 
in different insects according to the nature of the food, but their 
typical form is usually taken to be that shown by the lower insects 
which feed on solid food and have biting mouth parts. Figure 3, 
representing the jaws and lips of the common black cricket, is given 
as an example of generalized insect mouth parts. 

The labium (fig. 9A, Zm) is usually a simple transverse flap in front 
of the mouth, being developed, as already shown, from a similarh^ 
situated lobe on the first segment of the embryo (fig. 2, Lin). 

The epipharynx (fig. 19, Ephy) is a sort of dorsal tongue, and is 
situated on the membrane leading into the mouth from behind the 
labrum. 

The mandibles (figs. 3A; 9A, Md) are typically formed for 
biting, being heavy organs situated immediately behind the labrum 
and working sidewise on a hinge articulation with the head. Their 
cutting edges are usually notched and toothed, though smooth m the 
worker bee. 



GENEEAL EXTEENAL STEUCTUEE OP INSECTS. 



17 




A 




Hphy 



The maxillae (fig. 3 B and B) are complicated appendages in their 
typical form. Each consists of a principal piece called the stipes {St) , 
which is hinged to the head by means of a smaller basal piece, the 
cardo (Cd). Terminally the stipes bears an outer lobe, the galea 
{(ra), and an inner lobe, the lacinia (Lc). On the outer side, at the 
base of the galea, it carries a jointed appendage called the maxillary 
palpus {Pip). 

The hypopharynx (fig. 3 C and D, Hphy) is a median, ventral, 
tonguelike organ, called also the lingua^ situated either on the upper 
surface of the labium or on the membrane between this organ and the 
mouth. It is de- 
veloped principally 
from a median lobe 
of the head of the 
embryo behind the 
mouth (fig. 2, Zm), 
but some entomol- 
ogists claim that it 
is compounded of 
this lobe and two 
smaller lateral ones 
developed from the 
appendages of the 
fifth embryonic 
head segment (fig. 
2, Slin), the super- 
lingum. 

The labium (fig. 
3 C and D) consti- 
tutes the under lip 
of the adult, but it 
is formed from the 
two appendages of 
the seventh segment in the embryo, which fuse with each other. For 
this reason it is often called the second maxillm. It consists of a basal 
submenfum {Smt) bearing the mentum {Mt), which in turn carries 
three parts, a median ligula {Lg) and two lateral palpigers {Pig). 
The latter support the lahial palpi {Pip) , while the ligula bears four 
terminal lobes, of which the median ones are called the glossce {Gls) 
and the lateral ones the paraglossm {Pgl). If we should cut the 
labium into two parts along its midline we should see that even in 
the adult stage each half is very similar to one maxilla. The only 
discrepancy to be noticed in the example given (fig. 3) is that there 
22181— No. 18—10 2 




Fig. 3. — Example of generalized insect mouth parts, from 
common black cricket {Gryllus pennsylvanicus) : A, man- 
dibles ; B, B, maxillae, ventral view ; C, labium or second 
maxillse, ventral view ; D, labium, lateral view. 



18 THE ANATOMY OF THE HONEY BEE. 

is no maxillary palpiger, but many insects possess a corresponding 
part in the maxilla, frequently distinguished as the palpifer. 

The neck or cervicum is usually a short membranous cylinder which 
allows the head great freedom of motion upon the thorax. In nearly 
all insects its lateral walls contain several small plates, the cervical 
sclerites^ while, in many of the lower species, dorsal, ventral, and 
lateral sclerites are present and highly developed. As already stated, 
the origin of these plates is doubtful. Some entomologists would 
derive them from the prothorax, others think they come from the 
last head segment, while still others think that they represent a 
separate segment. Only pure anatomists, however, entertain this 
last view and call this supposed segment the " microthorax," for 
embryologists have not yet reported a metamere between the labial 
segment and the prothoracic segment. Most embryologists who have 
studied the subject admit that some of the cervical sclerites may be 
formed from the last embryonic head somite which carries the labium 
and probably forms a part of the back of the head. Therefore, if 
it is desirable to retain the word microthorax as a name for a true 
segment, it can be applied only to this labial metamere.'^ 

The thorax, as has already been stated, is a distinct anatomical 
region of the bod}^ rather than a " division " of the body, since it car- 
ries both the legs and the wings and contains the large muscles for 
each. Since the prothorax does not possess wings, it is not so highly 
developed otherwise as the two wing-bearing segments, and is, indeed, 
generally reduced in some ways, some of its parts being frequently 
rudimentary. Therefore we shall base the following description of 
a typical segment on the structure of the wing-bearing segments. 

A typical thoracic segment, then, presents four surfaces, as does also 
the entire body. These are a dorsum above, a venter below, and a 
latus ^ on each side. From these names we have the terms " dorsal," 

" In a former paper on the thorax of insects (Proc. U. S. Nat. Mus., XXXVI, 
1909, pp. 511-595) the writer probably drew a too definite conclusion on the 
subject of the " microthorax." The origin of the neck sclerites has probably 
never yet been actually observed. Comstock and Kochi (Amer. Nat., XXXVI, 
1902, pp. 13-45), in summarizing the segmentation of the head, accredited 
the gular and cervical sclerites to the labial segment, but did not recognize the 
latter as taking part in the formation of the true head capsule. Riley, how- 
ever, in his study of the development of the head of a cockroach (Amer. Nat., 
XXXVIII, 1904, pp. 777-810), states that in Blatta the labial segment does 
form a part of the back of the head and that the posterior arms of the 
tentorium are derived from it. Bonier (Zool. Anz., XXVI, 1903, pp. 290-315) 
and Crampton (Proc. Acad. Nat. Sci. Phila., 1909, pp. 3-54) believe that the 
cervical sclerites are derived principally from the prothoracic segment. The 
notion that they constitute a separate segment, the " microthorax," equivalent 
to the maxilliped segment of the centipedes, has been elaborated principally 
by Verhoeff in his numerous writings on the Chilopoda and Dermaptera. 

* The writer introduces this word here because he knows of no other term 
applied to the side of the segment in this sense. 



GENERAL EXTEENAL STRUCTURE OF INSECTS. 



19 



" ventral," and " lateral." The chitinous parts of the dorsum con- 
stitute the terguTYi; of the venter, the sternum; and of the latus, the 
'plev.TUUfh. 

The tergum of the wing-bearing segments usually consists of 
two plates— a front one or true notuTn (fig. 4, N) carrying 
the wings, and a posterior one, which the writer has termed the 
postnotum or pseudonotuwi {PN)^ having no connection with the 
wings. The first is often more or less distinctly marked into three 
transverse parts called the prescutum (Psc), scutum (Set), and scu- 
tellum {Scl). In such cases the exposed part of the postnotum is 
called the postscutellum (Pscl). From either the anterior or the pos- 
terior margin of the tergum, or from 
both, a thin transverse plate projects 
downward into the interior of the 
thorax for the attachment of muscles. 
These plates are the phragm,as (Aph 
and Pph). The notum supports the 
wing on each side by two small lobes, 
the anterior and posterior notal iving 
processes {ANP and PNP). Behind 
the latter is the attachment of the 
axillary cord (AxC) or basal ligament 
of the wing. A large V-shaped ridge 
on the under surface of the notum hav- 
ing its apex forward is the " entoclor- 
sum." (A better name would be 
entotergum.) 

The pleuruin consists principally of 
two plates, the epistemum (fig. 4, Eps) 
and the epimerum {Epm) lying before 
and behind a vertical groove, the pleural suture (PS), which extends 
from the pleural coxal process {CxP) below to the pleural wing 
process {WP) above. The pleural suture marks the position of a 
heavy internal ridge, the pleural ridge or entopleurum. The epi- 
merum is connected with the postnotum {PN) behind the base of the 
wing. These parts occur in almost all insects. In some of the lower 
ones another plate is present in front of the episternum which may 
be called the preepisternum {Peps)."' Lying along the upper edge of 

"' Objection may be made to the use of the term " preepisternum " on the 
ground that it combines a Latin prefix with a word compounded of Greeli ele- 
ments. The same may be urged against " prephragma," " postphragma," " pre- 
paraptera," and " postparaptera," words introduced by the present writer in a 
former paper on the thorax (Proc. U. S. Nat. Mus., XXXVI, 1909, pp. .511-595). 
However, we are barred from making up equivalent terms with the Greek pre- 
fixes pro and meta because these are used to designate the first and the third 




TnC 

Fig. 4. — Diagram of generalized 
thoracic segment, left side. 



20 THE ANATOMY OF THE HONEY BEE. 

the pleurum and associated with the under surface of the wing base 
are several small plates known as the paraptera {P).°- Two lie above 
the episternum in front of the pleural wing process and are the 
episternal paraptera or pfeparaptera {IP and 2P)^ while one or 
occasionally two are similarly situated behind the wing processes 
and are the epimeral paraptera or postparaptera {3P and ^P) . The 
preparaptera afford insertion for the muscle concerned in the exten- 
sion and pronation of the wing. 

The coxa (Cx), or basal segment of the leg, is hinged to the seg- 
ment by a dorsal articulation with the pleural coxal process (CxP), 
and by a ventral articulation (TnC) with a plate called the trochan- 
tin {Tn) lying in front of it and connected above with the lower 
end of the episternum {Eps). Hence, while the leg is of course con- 
tinuous all around its base, by means of membrane, with the body- 
wall, its movement is limited to a hinge motion hj these two special 
articulations of the chitin. 

The sternum or ventral plate of the segment is not so complicated as 
are the tergum and pleurum. • It is often divided transversely into 
three parts, however, and some authors say 4:ypically into four. These 
parts have been named the presternum {Ps), sternum proper (S), 

segments of the thorax or their respective parts. Entomologists have already 
established the system of referring a part to the front or back of any individual 
segment by the Latin prefixes pre (or prw) and post as used in " prescutum," 
" presternum," " postscutellum," and " poststernellum." Furthermore, pre and 
post are so indiscriminately used in English combined with Latin, Greek, and 
even Anglo-Saxon words that they may be regarded as general property. 
Hence, in order not to sacrifice an anatomical system, which certainly needs 
to be fostered in every way, the writer has preferred to sacrifice strict gram- 
matical rules by applying pre and post, regardless of the origin of the noun 
in the case, to designate anterior and posterior parts of the same segment. We 
already use such hybrid terms as " presternum," " mesotergum," and " meta- 
tergum." 

The name " preepisternum " has been applied by Hopkins (Bui. 17, Pt. I, 
technical series. Bur. Ent., U. S. Dept. Agr., 1909) to a part of the mesepister- 
num of Dendroctonus — a plate apparently not homologous with the preepisternal 
element of the thorax in primitive insects. 

''The name " parapterum " is taken from Audouin's term paraptere (Ann. 
des Sci. Nat., I, 1824, pp. 97-135, 416-432), and its application, as used by the 
present writer, is based on Audouiu's definition given in his Chapter III, 
'' Considerationes generates sur le Thorax,'' where he says (p. 122) : "Finally 
thei'e exists a piece but little developed and seldom observed, connected with 
both the episternum and the wing. It is always supported by the episternum 
and is sometimes prolonged ventrally along its anterior margin, or again, 
becoming free, passes in front of the wing and may even come to lie above 
the base of the latter. At first we designated this sclerite by the name of 
Hypoptere but on account of its change of position relative to the wing base 
we now prefer the name of Paraptere." The first part of his description leaves 
no doubt that Audouin referred to the little pleural plate beneath the front 
of the wing which is usually very inconspicuous except in carefully dissected 




GENEEAL EXTERNAL STRUCTURE OF INSECTS. 21 

stemeUuin (SI), and porststernellum {Psl). In some of the lower 
insects a plate (x) occurs at each side of the presternum or of the 
sternum which seems to fall in line with the preepisternum of the 
pleurum. This has been variously called a part of the prestermtm, 
the coxosternum, an accessory sternal plate, and the sternal laterale. 
The inner surface of the 
sternum carries a large 
two-pronged process 
called the furca or ento- 
sternum. 

This plan of structure 
for the mesothorax and 
the metathorax prevails 
throughout all insects. 
The honey bee probably 
presents the greatest de- Emp' 

i j> -x 1 X Fig. 5. — Typical insect leg. 

parture rrom it, but even 

here the modification consists principally of a suppression of the 

sutures of the pleurum resulting from a condensation of the parts. 

The leg (fig. 5) of an adult insect consists of a number of joints 
or segments. It is attached to the body, as just described, by a thick 

specimens. lu such preparations, however, one finds that there are in most 
cases two sclerites here instead of one. and, furthermore, that one or occa- 
sionally two others are similarly situated beneath the rear part of the wing 
base behind the pleural wing process. The present writer has, therefore, 
made the term " paraptera " cover this whole row of little plates, distinguish- 
ing those before and those behind the pleural wing process by the designations 
given above. 

In the latter part of Audouin's definition it would seem that he may have 
confused the rudimentary tegula as it exists in some insects with the parapte- 
rum, but even this is not probable since he says it is always connected with 
the episternum, which is never true of the tegula. In his description of the 
thorax of beetles, Dytiscus, Garatus, Buprestis, and Gurculio, it is evident 
that he regards the anterior upper part of the episternum as the parapterum 
fused with the latter plate. In fact, in each case he definitely states that such 
is the case and, in describing Dytiscus circumflexus, he says (p. 420) : "The 
episternum, the parapterum, and the epimerum all fuse dorsally and constitute 
a support for the wings and tergum." While Audouin is undoubtedly mis- 
taken in this homology, especially in the mesothorax, he at least shows that 
his " paraptere " is a part of the pleurum. Hence modern writers such as 
Packard and Folsom who make the term " paraptera " synonymous with 
" tegulcie " are certainly wrong. The tegula is a dorsal scale or its rudiment 
at the humeral angle of the wing, while the parapterum is a co-existent scle- 
rite below this part of the wing base. The present writer agrees with Comstock 
and Kellogg, who, in their Elements of Insect Anatomy (first edition), define 
the little sclerite in front of the base of the wing in the locust, articulated to 
the dorsal extremity of the episternum, as the " parapteron," though in this 
insect there are here really two of these parapteral plates instead of one. 



22 



THE ANATOMY OF THE HONEY BEE. 



basal joint called the coxa {Cx). Beyond this is a smaller joint 
called the trochanter {Ti")^ this is followed by a long and strong 
segment, the femur (F) , which extends outward from the body, while 
bending downward from its distal end is the long and slender tibia 
(Th), followed finally by the foot, or tarsus (Tar). The tarsus itself 
consists typically of five small segments of which the last bears a pair 
of claws (Cla). The under surfaces of the tarsal joints are often 
provided with small cushions or pads called pulviUi Those between 
the claws are generally specially prominent and are called the 
empodia {Emp). The leg varies greatly in shape in different in- 
sects but usually preserves all of these parts. The segments of the 
tarsus, however, are frequently reduced in number. 

The adult wing is a thin expanse of membrane supported by hollow 
branching rods called veins. It originates as a hollow outgrowth of 
the body-wall, but soon becomes flattened out dorso-ventrally and the 




Fig. 



-Diagram of 



generalized insect wing and its articulation to first plate (.V) of 
the tergum. 



contained tracheae or air tubes mark out the courses of the veins. 
These veins form various patterns in different insects, but they can all 
be derived by modification from one fundamental plan. This plan is 
shown diagrammatically by figure 6. The first vein, which usually 
forms the anterior margin of the adult wing, is the costa (C). The 
next vein is the subcosta (Sc), which in typical cases divides into 
two branches (/S'6\ and Sco). The third and usually the principal 
vein is the i^aclius {R). It divides dichotomously into five branches 
{R^ to i?5), the anterior branch of the first fork remaining single. 
The next vein is the ineclia (M), which forms four branches (J/^ to 
iJ/4). The fifth is the cubitus {Cu), which again is two-branched. 
The remaining veins are called the anals and are designated indi- 
vidually as the first anal (^A), second anal {^A), etc. 

Several cross- veins of common recurrence should be noted. The 
first is situated near the base of the wing between the costal and 
subcostal veins and is known as the humeral cross- vein. A second 



GENERAL EXTERNAL STRUCTURE OP INSECTS. 23 

occurs between the radius and the media near the center of the wing 
and is called the radio-medial cross-vein. Another one, the medio- 
cuMtal^ is similarly located between the media and the cubitus, 
while a fourth, called the median, occurs between the second and 
third branches of the media. The areas of the wing surface inclosed 
by the veins, the cross-veins, and the margins of the wing are known 
as the cells. 

A great many different names are applied by different entomolo- 
gists to the veins of the wings, both of the same and of different 
insects. The nomenclature here given is the one first consistently 
applied by Comstock and Needham and now used by a large number 
of entomologists working in different orders of insects. 

The wing is articulated at its base (except in mayflies and dragon- 
flies) to the anterior and posterior wing processes of the notum 
(fig. 6, ANP and PNP) and to the wing process of the pleurum (fig. 
4, WP) by several small articular sclerites called axillaries. Two 
of these, the first {lAx) and the fourth {JfAx) , form a hinge with the 
anterior and the posterior notal wing processes, respectively, while 
the second {2Ax) articulates below with the wing process of the 
pleurum, constituting thus a sort of pivotal element. The third axil- 
lary {3 Ax) intermediates between the bases of the anal veins and the 
fourth axillary — excej^t when the latter is absent (as it is in nearly 
all insects except Orthoptera and Hymenoptera), in which case it 
articulates directly with the posterior notal process. The thin mem- 
brane of the wing base may be called the axillary membrane {AxM). 
On its anterior edge is a hairy pad, the tegula (Tg), which is some- 
times a large scale overlapj)ing the humeral angle of the wing. The 
j)osterior margin of the axillary membrane is thickened and may be 
called the axillary cord (AxC) or basal ligament of the wing. 

The base of the costa is not directly associated with any of the 
axillaries, but is specially connected by tough membrane below with 
the episternal paraptera. The subcosta abuts against the end of 
the curved neck of the first axillary. The radius is either attached 
to or touches upon the anterior end of the second. The media and 
cubitus are usually associated with each other at their bases and also 
more or less closely wuth one or two median plates (m) in the wing 
base. These plates, hov/ever, are not of constant shape and occur- 
rence as are the articulating axillaries. The anals are generally 
attached to the outer end of the third axillary, which acts as a lever 
in the folding of the wing. 

A few insects have a generalized wing almost identical with the 
diagram (fig. 6), but most of them depart from it in varying degrees. 
Few go so far, however, as the honey bee, whose venation is very 
different, but yet the fundamental basal structure is the same even 



24 THE ANATOMY OF THE HONEY BEE. 

here, as will be shown in the special description of the wing of the 
bee. 

The abdomen consists almost always of 10 segments. There are 
never any more than this number well developed in adult insects, and 
if there are fewer the reduction is due to a modification of the ter- 
minal segments to accommodate the external organs of reproduction. 
The posterior opening of the alimentary canal is at the end of the 
tenth segment, which carries also two small appendages at the sides of 
the anus. These are called the cerci (fig. 8, Cer) . In some insects they 
are short, styletlike processes, in others they are long and many 
jointed, while in many they are absent. The cerci are supposed to 
be developed from the embryonic appendages of the tenth segment, 
although, on the other segments, these appendages disappear before 
the embryo hatches, except in some members of the lowest wingless 
order of insects, which have a pair of cercuslike appendages on each 
segment of the abdomen. 

Each abdominal segment presents a tergum above and a sternum 
below; the former usually also reaches far down on the sides and 
overlaps the edges of the sternum. In some insects one or more small 
pleural plates intervene between the tergum and the sternum, but 
the abdominal pleura are never developed in any way suggestive of 
a thoracic pleurum. Very frequently there is present an upper 
pleural plate, or epipleurite, adjoining the edge of the tergum and a 
lower, or hypopleurite^ adjoining the edge of the sternum. The line 
separating these two sclerites, however, is horizontal and can not 
correspond with the vertical suture of a thoracic pleurum between the 
episternum and the epimerum extending from the base of the leg 
to the base of the wing. 

The most complicated structures on the abdomen are the external 
organs of reproduction. In the male these serve as clasping organs 
and take on a great variety of forms in different species. The organs 
in the female form an ovipositor and are of much more definite and 
constant structure. 

The ovipositor (fig. 8), in its most perfect development, consists of 
three pairs of long, closely appressecl bladelike processes called 
gonapophyses {IG, 2G^ 3G). These six pieces fit neatly together and 
form an organ by means of which the female makes a hole in the 
ground or in the bark of a tree, or punctures some other insect, and 
then places her eggs in the cavity thus produced. An interesting fact 
in this connection is that the sting of a wasp or bee is simply a modi- 
fied ovipositor. This can be proved by a comparison of the organs 
themselves or by a study of their development. Each is formed from 
six little peglike processes that grow out from the sterna of the eighth 
and ninth abdominal segments of the larva or young soon after hatch- 



GENERAL EXTERNAL STEUCTtJEE OF INSECTS. 



25 



ing (fig. 7, IG, 2G^ and 3G). At first there is only one pair of these 
processes on each of the two segments, but those on the ninth soon 
split each into two, thus producing two pairs on this segment. The 
opening of the oviduct {OvO) is on the 
eighth segment between the bases of the 
first gonapophyses. 

The ovipositor of the longhorned grass- 
hopper, shown by figure 8, may be taken as 
a typical example of this organ. The 
median pair of gonapophyses on the ninth 
segment {2G) remain slender and fuse at 
their bases into a small bulblike swelling 
open below (ShB). The pair from the 
eighth segment (IG) form two long blade- 
like pieces, which fit by sliding articula- 
tions upon the lower edges of the corre- 
sponding second gonapophyses {2G). The 
first can therefore be worked back and 
forth while they are braced and held in 
position by the second pair. The third 
gonapophyses {3G), or the outer pair of 
the ninth segment (the left one in figure 8 is shown as if cut off near 
its base), form two long flat blades which are closely appressed 
against the outer surfaces of the others. In the detailed study of 
the bee it will be shown how closely the structure of the sting corre- 
sponds in every way with that of this ovipositor. 




An ;^G 

Fig. 7. — Diagram of terminal 
abdominal segments of a fe- 
male insect and early stage in 
development of gonapophyses 
{IG, 2G, and 3G), from 
which is formed the ovi- 
positor of most insects and 
the sting of wasps and bees. 




Fig. 8 
cephalu 
bee, fig. 



Example of a swordlike ovipositor, from a longhorned grasshopper (Cono- 
sp.), Illustrating the fundamental similarity of structure with the sting of the 



Some entomologists have supposed that the original two pairs of 
gonapophyses represent the embryonic appendages of the eighth and 
ninth segments, and they would thus establish a homology between 
the ovipositor or sting and the legs and mouth parts. It has been 
shown, however, that the true appendages of the abdominal segments 
disappear in embryonic life while the gonapophyses appear much 
later, during early nymphal or larval life. Furthermore, each pair 



26 THE ANATOMY OF THE HONEY BEE, 

of gonapophj^ses arises in a median depression on the ventral side of 
the segment while the true appendages are latero-ventral. Hence, 
the evidence is very much against this theory and the gonapophyses 
appear to be special secondary processes of the body Avail. 

All insects do not have ovipositors of the sort described above. 
Flies, beetles, moths, and butterflies do not. Such insects simply 
drop their eggs from the orifice of the oviduct or deposit them in 
masses upon the external surfaces of various objects. In some of 
the flies, however, the terminal segments are long and tubular and 
entirely telescoped into one another. They are hence capable of 
being protruded in the form of a long tapering tube having the open- 
ing of the oviduct near the tip. This enables the insect to deposit its 
eggs in deep crevices, but the structure is not a true ovipositor — -it is 
simply the abdomen itself stretched out. 

Insects breathe through a series of small holes situated along each 
side of the body. These breathing apertures are called sjnracles and 
they lead into a system of internal air tubes called trachece. There 
are nearly always 10 spiracles present on each side of the body. Two 
are located on the thorax, the first between the prothorax and the 
mesothorax, the second between the mesothorax and the metathorax, 
while the other eight are situated on the first eight abdominal seg- 
ments. Some embryologists believe that the spiracles of the pro- 
thorax move forward in early embryonic life and unite with each 
other in front of the hj^popharynx to form the salivary opening, their 
tracheae constituting the salivary ducts. 

After this review of the general external structure of insects we 
may proceed to a more detailed account of the parts and organs of 
the honey bee. 

III. THE HEAD OF THE BEE AND ITS APPENDAGES, 

The head of an insect, as already explained, is a composite organ 
formed of six or seven primitive segments, each of which, except the 
first, typically bears a pair of appendages (fig. 2). The antennae are 
developed from the embryonic appendages of the second segment, 
the mandibles from the fourth, the maxillae from the sixth, and the 
second maxillse, or labium, from the seventh. The appendages of 
the third segment disappear in early embryonic life while those of 
the fifth segment, when the latter is present, fuse with a median 
tonguelike lobe of the next segment to form the hypopharynx of 
the adult. 

1. THE STRUCTURE OF THE HEAD. 

The general appearance and outline of the head of a worker bee 
are shown from before and behind by figure 9, A and B. In facial 
view the head is triangular, with the apex below. The side angles 



THE HEAD OF THE BEE AND ITS APPENDAGES. 



27 



are roimded and capped by the large compound eyes {E). In the 
opposite direction the head is very much flattened, the greatest diame- 
ter being crosswise through the middle of the eyes. The face is con- 
vex, while the posterior surface is somewhat hollowed out and fits 
snugly upon the anterior end of the thorax. 

The large lateral eyes (fig. 9 A, E) are called the compound eyes, 
because each is composed of a large number of separate eye elements 
forming the little hexagonal facets visible on the surface. All of 
these facets together constitute the cornea^ or the transparent outer 
surface of the eye, which in the bee is densely clothed with long hairs. 
The dark color of the eye is located in the deeper parts, but these will 
be described in the section dealing with the nervous system. On the 



Vx ten 




Gl/]/ Pgl 

Fig. 9. — A, front view of head of worker bee with mouth parts (Prl}) cut off a short 
distance from their bases ; B, corresponding view of posterior surface of head. 

top of the head between the compound eyes are the three simple eyes, 
or ocelli {0), arranged in a triangle with the median ocellus in front. 

Between the lower halves of the large eyes and near the center of the 
face arise the antennae {Ant) , each of which is inserted into a small, 
circular, membranous socket of the head wall, and consists of a long, 
basal, 1-segmented stalk carrying a terminal 11-jointed arm movably 
articulated to the stalk and generally hanging downward from it. 
(In the drone the terminal arm consists of 12 joints.) 

The mouth parts are attached at the lower part of the head, and 
coiLsist of the mandibles {31 d) laterally and the maxillce {Mx) 
and labium {Lb) mesially. The latter two include the set of elongate 
bladelike organs surrounding the protrusible " tongue," which to- 
gether constitute what is commonly known as the lyroboscis {Prb). 



28 THE ANATOMY OF THE HONEY BEE. 

When not in use the j)arts of the proboscis are bent back beneath 
the head. By referring to figure 9B, giving a posterior view of the 
head, it will be seen that the basal parts of both the maxillae (St) 
and the labium (Mt) are suspended in a large hollow on the back of 
the cranium. This may be called the cavity or fossa of the proboscis 
(PrhFs). Between the mandibles on the front of the head (fig. 
9A) is a transverse movable flap,- the lahrum (Lm), attached to the 
lower edge of the front wall of the head and constituting the upper 
lip. The mouth {31 th) lies behind the labrum and the mandibles 
close beneath it. 

Below the antennal sockets is a transverse, slightly arched suture 
(a) which turns downward on each side and extends to the inner 
angles of the bases of the mandibles. The area bounded by this 
suture is the clypeus {Glp) and the suture itself may be called the 
clypeal suture. 

On the posterior surface of the head (fig. 9B) is seen the pen- 
tagonal foramen Tnagnum {For) by means of which the cavity of 
the head communicates with that of the thorax and through which 
pass the nerves, oesophagus, blood vessel, and tracheal tubes. A 
small rod {ten) inside the head arches transversely over the fora- 
men magnum, cutting it into a dorsal and a ventral half. At each 
side of the foramen is a large pit {c) which marks the base of an 
internal chitinous beam of the head known as the mesocephalic pillar. 
The opposite end of this pillar unites with the front wall of the 
head on the clypeal suture below the antennae, where it produces 
another smaller pit (5). 

Below the foramen magnum and separated from it by a wide trans- 
verse bridge of the cranial wall is seen the large fossa of the proboscis 
(fig. 9B, PrhFs) having the shape of an inverted U. The side walls 
of this cavity are chitinous and from their upper edges are suspended 
the maxillae, while the base of the labium is contained in the mem- 
branous floor of the fossa. The base of the labium projects from the 
head beneath or behind the mouth opening and its dorsal surface 
forms the floor of a preoral cavity surrounded by the bases of the 
mouth parts and labrum. 

It will be seen from the above description that the head wall of the 
bee contains no suture except that bounding the clypeus and the one 
which separates the labrum from the latter. Many of the higher 
insects have the head wall completely continuous, showing no division 
at all into sclerites, but, in such forms as a grasshopper or cockroach, 
and, in fact, most of the lower insects, the head as well as the other 
parts of the body is made up of a number of plates. Hence this may 
be regarded as the primitive condition, and it is presumed that the 
head of the bee has been produced from one whose wall was divided 
by sutures into a number of distinct parts. Therefore the different 



THE HEAD OF THE BEE AND ITS APPENDAGES. 



29 



regions of the bee's head may be named according to the sclerites with 
which they correspond in other insects. Thus, the part of the face 
above the clypeus and between the compound eyes may be called the 
front (fig. 9A, Ft) , the parts below the compound eyes the gence {Ge) , 
and the top of the head the vertex 
{Vx). The area on the back of the 
head around the foramen magnum 
may likewise be termed the occijntal 
region (fig. 9B, Oc) and the parts be- 
hind the gense and the lower halves 
of the compound eves the 'postgenm 

{Pge)- 

The worker, queen, and drone differ 
conspicuously in the shape and size of 
the head, as will be seen by comparing 
A, B, and C of figure 10. In these 
drawings the front has been removed 
in order to show various internal 
parts, which will be described later. 
While the head of the worker (A) is 
triangular in facial view, that of the 
queen (B) is more rounded and wider 
in proportion to its length. The head 
of the drone (C) is much larger than 
that of the female and is nearly cir- 
cular in outline. In shape the head 
of the queen is intermediate between 
that of the worker and that of the 
drone, but in size it is somewhat 
smaller than the head of the worker. 
The eyes {E) of the worker and queen 
are about equal, but those of the drone 
are enormously enlarged and are 
broadly contiguous on the vertex and 
the upper part of the front. On this 
account the ocelli {0) oi the drone are 
crowded down on the front nearer the 
bases of the antennae and the front 
itself is very much narrowed above. 
The antennae of the drone consist of 
13 segments, while those of the females 
have but 12 segments. The mandibles are largest proportionately in 
the queen and are very small in the drone. Those of the worker have 
a smooth terminal edge, while this edge is notched in the queen and 
the drone. The parts of the proboscis are much longer in the worker 





Fig. 10. — A, anterior view of head of 
worker, with front, antenna?, and 
proboscis removed ; B, correspond- 
ing view of head of queen ; C, same 
of drone. 



30 



THE ANATOMY OF THE HONEY BEE. 



and capable of much more action than in the queen and drone, which 
are almost entirely dependent upon the workers for their food. 

The internal structure of the cranium may be studied best in a longi- 
tudinal section of the head (fig. 11). In order to prepare a section 
for this purpose imbed the head in paraffin and then carefully slice 
off one side with a sharj) knife or razor just outside of the bases of 
the mandible and antenna. Holding the remainder in the block of 
paraffin or fastening the whole in a dish of water or alcohol, care- 
fully dissect away the soft parts from the head cavity so as to expose 




Fig. 11. — A, longitudinal section through head of worker between the median plane and 
outer edges of mandibles (Md) and antennae (Ant) of left side, all internal soft parts 
removed ; B, corresponding section through head of drone, except that the pharynx 
(Phi/) and oesophagus (CE) are not removed. 

the internal chitinous parts shown in figure 11 A and B. These 
figures, however, represent a slice of the head taken from between the 
median plane and the outer edges of the antennal and mandibular 
bases of the left side. Thus only the parts on one side of the mid- 
line are shoAvn. Figure A is from a worker and Figure B from a 
drone. In the latter the phaiynx and oesophagus are retained and 
the neck is not removed. Figure 20 .shows the head cut open from 
above and the mouth parts removed. A specimen so cut and boiled 
a short time in caustic soda or potash to remove the soft parts will 
be found a valuable adjunct to this study. 



THE HEAD OF THE BEE AND ITS APPENDAGES. 31 

The principal parts of the internal skeleton of the head, or ento- 
cranium, consist of two large, oblique, strongly chitinous bars form- 
ing a brace between the anterior and the posterior walls of the head 
(fig. 11 A and B, Ten, showing the parts on the left side only, and 
fig. 19, Ten). These bars have been named by Macloskie (1881) the 
7nesocephaUc pillars. As already pointed out the base of each is 
marked externally by a conspicuous pit (fig. 9 B, e) laterad of the 
foramen magnum, and its facial end by a smaller pit (fig. 9 A, h) 
in the clypeal suture near the upper end of each side of the latter. 
The bases of these pillars are connected by the slender bar (fig. 11 A, 
ten), already noticed, arching over the foramen magnum (fig. 9 B, 
ten). This bar and the two pillars represent what is called in other 
insects the tentorium. In the embryo the tentorium is formed from 
tubular ingrowths of the head wall which unite internally and 
assume different shapes in different insects. Since the air tubes of 
the body also first appear as tubular ingrowths of the body wall, 
some entomologists have supposed that the hollow tentorial in- 
growths of the head represent the spiracular tubes of the head 
Avhich are, otherwise, lacking. However, there is not sufficient evi- 
dence to support such a view as this, and there is no reason why the 
tentorium should not have been originally designed simply to give 
greater rigidity to the walls of the head where the latter support the 
appendages. 

The usual form of the tentorium in the lower insects is that of an 
X, with a large central body, situated like a brace across the lower 
part of the head, having two of the arms directed anteriorly and 
laterally and two directed posteriorly and laterally, and while the 
former are said to be ingrowths from the mandibular segment, there 
is some difference of opinion concerning the segment to which the 
latter belong. Eiley states that they are formed in the labial seg- 
ment of the cockroach and Carriere and Burger describe the same 
thing for the mason bee. Other authors have ascribed them to the 
maxillary segment, but they may, in later stages, lie in this segment 
and thus appear to belong to it, while they originated in the one 
following, having moved forward on account of the condensation 
of the back part of the head. The tentorium of the honey bee, 
consisting as it does of the two great mesocephalic pillars (fig. 11 
A and B, Ten) and the small arched bar {ten) is so highly modified 
that it is hard to see just how its parts are to be homologizecl with 
the parts of an X-shaped tentorium. Probably the two pillars repre- 
sent the separated halves of the X, while the slender arch is an addi- 
tional structure. In any case we have not enough evidence to war- 
rant us in regarding the tentorial invaginations as modified tracheae, 
or their external pits as rudimentary spiracles. Similar processes 
extend inward from the w^alls of the thorax to strengthen it or to 
give attachment of muscles. Such processes in general form the 



32 THE ANATOMY OF THE HONEY BEE. 

eiitoskeleton and are individually called apodeTnes. Those of the 
head constitute the entocranium^ those of the thorax the entothorax. 

The side walls of the fossa of the proboscis form two high, thin, 
vertical plates, as seen from the interior of the head (fig. 11), in 
front of the mesocephalic pillars. The posterior edge {d) of each 
of these plates is so much thicker than the rest of it in the worker 
that it appears at first sight to be a separate rod. Its upper end 
projects above the body of the plate as a free arm (e) to which is 
articulated the basal piece of the maxilla (Cd). It thus constitutes 
the maxillary suspensorium. (Macloskie includes under this term 
both the arm of the cranial wall and the cardo of the maxilla.) 

The head of the drone (fig. 11 B) presents, besides the parts de- 
scribed, a thin plate (/) depending from the vertex of the cranium 
along the line between the compound eyes. 

Besides these apodemes of the cranial wall itself there are others 
which project into the head cavity from the bases of the appendages 
to afford points of insertion for their muscles. These are specially 
developed in connection with the mandibles and will be described in 
the discussion of these organs. Still other internal chitinizations are 
developed in the walls of the pharynx, but these likewise will be 
described later. 

2. THE ANTENNA AND THEIR SENSE ORGANS. 

The antennse of the bee are the two slender, jointed appendages 
movably attached to the center of the face, where each is inserted 
into a circular membranous area or socket just above the upper part 
of the clypeal suture. Their general shape and position are shown 
by figures 9 A, 11 A, and 19, Ant. Each is seen to consist of two 
parts, forming a prominent elbow with each other, and usually so 
held that the first or proximal part extends outward and upward 
from its frontal attachment and carries the other in a pendent posi- 
tion from its distal end. The first part thus forms a basal stalk, 
called the scape (figs. 9 A; 19, Sep), consisting of a single joint 
inserted into the antennal socket of the front by a prominent basal 
condyle bent toward the face. This articular knob is attached to 
the rim of the socket by a circle of membrane, but it is also pivoted 
on a slender peglike process projecting upward from the lower edge 
of the socket. Hence, while the flexible membrane allows each 
antenna to revolve freely in any direction, the latter is at the same 
time held firmly in position by the pivot. The antennae are moved 
by special sets of muscles inserted upon their bases within the head. 
The second or distal division of the antenna is cylindrical and longer 
than the first, forming a flexible fagelhim (fig. 9 A; 19, Fl) hanging 
downward from the distal end of the scape. It is composed of 11 



THE HEAD OF THE BEE AND ITS APPENDAGES. 33 

small joints in the worker and queen and of 12 in the drone. The 
male antenna thus consists of 13 joints in all, while that of the female 
has but 12. The first joint of the flagellum is freely articulated to 
the scape, but the others do not have much play upon one another, 
though they give flexibility to the flagellum as a whole. 

Each antenna is a hollow tube containing the large antennal nerve, 
minute extensions of the tracheal system, and the small muscles which 
move the segments upon one another. 

Popularly the antennae of insects are known as the " feelers," be- 
cause they are constantly moved about in all directions with a nervous 
kind of motion as if the creature were feeling its way along by means 
of them. In fact " feelers " is a better name for these appendages 
than the scientific term, for there can be no doubt that the sense of 
touch is very highly developed in them and that by means of them 
insects acquire a great deal of information concerning their surround- 
ings and their companions. Moreover, a large mass of evidence 
derived from experiments shows unquestionably that the organs of 
smell also are located upon the antennae in a great many if not all 
insects, while some investigators believe that in some species they 
carry in addition the organs of hearing. 

The study of the senses of insects is a most elusive subject, and 
becomes more so the more we ponder on the results of experiments. 
In the first place, it is manifestly impossible for us to acquire any 
real knowledge of an insect's sensations, for wdiat is to us an odor, 
a taste, a color, or a sound may be something quite different to such a 
differently organized creature. We can, however, by experiments 
determine that some things which give us the sensation of an odor 
are perceived also by insects when placed near them. Also it can be 
shown that some of them distinguish substances of different taste in 
their food, and likewise that they perceive movement and distinguish 
the colors and in a vague way the outlines of objects. Furthermore, 
it is known that some of their perceptions are more delicate than ours, 
and that some insects at least see color where we see none. They may 
even possess senses of which we have no conception. 

Hence, while it can be positively stated that insects perceive differ- 
ences of touch, taste, smell, sound, and light, and act accordingly, we 
can not say what the sensations they acquire are like. In fact we 
do not know that they have conscious sensations at all. What looks 
like an action due to intelligent perception may be purely a reflex one, 
unaccompanied by any sensation. This of course involves the ques- 
tion as to whether such creatures or insects are possessed of conscious- 
ness or not — a question which can not be answered one way or the 
other. 

Understanding, then, that our knowledge of insect senses amounts 
only to this, that what gives us the sensation of light, sound, taste, 
22181— No. 18—10 3 



34 . THE ANATOMY OF THE HONEY BEE. 

touch, or smell makes also some sort of an impression on the insect 
and varies in degree and kind much as it does in us, we may go on to 
a study of the senses located on the antennae. 

Here, again, however, we are confronted by a difficulty, for while, 
at first thought, it seems very easy to hold some strong-smelling sub- 
stance near the antennae of a beetle, ant, or bee and observe the evident 
displeasure with which the creature turns away, yet we may be en- 
tirely wrong if we conclude that the insect " smells " the substance 
that repels it. Strong-smelling, volatile liquids may simply produce 
pain in some of the delicate nerve endings of the antennae. Some 
other kind of a being, experimenting on our senses, might close up 
our nose and mouth and prove that we smell by means of our eyes 
on observing the blinking we should perform when strong formalin 
or ammonia was held close to the face. Furthermore, irritant gases 
and volatile liquids affect the mucous membranes of our noses and 
throats in a way quite independent from the odor that we perceive, 
and there is no reason why the same may not be true of insects. As 
pointed out by Forel, experiments on the sense of smell should be 
made with odorous substances that the insect meets Avith in a state of 
nature, which would be principally the materials it feeds on. In- 
sects are indifferent to almost every mildly odorous substance not 
used as food, which, however, does not prove that they do not smell 
them. 

Again, in many cases, it would be difficult to decide whether the re- 
sults of an experiment should be accredited to smell or sight. For 
example, every bee keeper knows that hungry bees are attracted to 
honey a long distance from their hives, and it would seem almost self- 
evident that they are guided by a sense of smell. Yet one might con- 
tend that they find the honey by sight, as, indeed, is claimed by a 
number of entomologists who have made experiments on the olfactory 
powers of bees. This question has been decided in some other insects 
by painting the eyes with some opaque substance or by removing the 
antennge, but the evidence is not conclusive on either side in the case of 
bees. 

Experiments made by a large number of competent investigators, 
including Lubbock, Schiemenz, and Forel, have proved conclusively 
that the organs of the sense of smell in insects are located principally 
on the antennae. The most interesting of these experiments are per- 
haps those which Forel (1903) made on carrion-feeding beetles. He 
found the dead and putrid bodies of a hedgehog and a rat infested by 
a swarm of these beetles belonging to several genera. He collected 
more than 40 specimens from the carcasses and removed their an- 
tennae. Then he placed them all at one place in the grass and moved 
the dead bodies to a distance of 28 paces from the beetles where he 
concealed them in a tangle of weeds. Examination the next day 



THE HEAD OP THE BEE AND ITS APPENDAGES. 35 

revealed the fact that not one of the mutilated beetles had found the 
carcasses. Repeated experiments gave the same results — no beetle 
without its antennae was ever found on the dead animals, although at 
each examination new individuals of the several species were present. 
It might be supposed that the mutilation itself distracted the beetles 
to such an extent that they did not care to eat. In order to test this 
point Forel next cut off all the feet on one side of the body from a 
dozen intact beetles and changed the location of the dead bodies again. 
The next day five of this lot were found on the carcasses. 

The same results have been obtained from experiments on other 
insects. Ants distinguish between their comrades and enemies by 
means of their antennal sense organs. Males of the silkworm moth 
and many other moths and butterflies perceive the presence of 
the females and are guided to them by an evident sense of smell 
located on the antennae, for they fail completely to find them when 
these appendages are removed, although one immediately recognizes 
a female when j^laced in contact with her. 

Similar experiments have been made on the bee, testing the ability 
of the workers to find honey hidden from their sight. The results, 
according to Forel, seem, curiously enough, to indicate that bees can 
j)erceive odors but a very short distance from their heads. Forel 
found that hungry bees in a cage would pass and repass hundreds of 
times within a few millimeters of some honey concealed from their 
sight by a lattice without discovering it. They ate it greedily, how- 
ever, when the lattice was removed, though it had been perfectly 
accessible to them all the time. Forel believes that " bees guide them- 
selves almost exclusively by vision," and Lubbock holds the same 
opinion. At the same time it would probably be a very difficult mat- 
ter to convince many practical bee keepers that bees do not " smell " 
from long distances. It is a well-known fact that at times when nec- 
tar is scarce bees are attracted in large numbers to the houses of an 
apiary where honey is stored, though, when the natural flow is suf- 
ficient, they pay no attention to it. Tests of the olfactory sense should 
undoubtedly be made under natural conditions. Bees inclosed in a 
box with some honey concealed from their sight might not be able to 
locate it in such close quarters though they might be smelling it all 
the time. An odor in a room may so fill the air that it does not seem 
to come from any particular direction and we ourselves would have 
to exert our intelligence to discover its source. 

While, then, it does not seem probable that bees have such limited 
olfactory powers as some investigators claim their experiments indi- 
cate, it may be accepted as proved that the organs of smell are located 
principally on the antenna?. It has already been stated that the sensQ 
of touch also is very highly developed on these organs, although in a 
less sensitive degree it is distributed over most of the other parts of 



36 



THE ANATOMY OF THE HONEY BEE. 



the body. It is again specially developed on the palpuslike append- 
ages of the sting. (See figs. 36 and 37, StnPlp.) Sections of a bee's 

antenna show that there are 
on its surface a great number 
of minute structures of sev- 
eral different kinds, though 
all apparently are to be re- 
garded as modified hairs, 
whidi are undoubtedly the 
sense organs. Now the diffi- 
culty arises of deciding which 
of these to assign to the sense 
of touch and which to the 
sense of smell. Different au- 
thors have made such differ- 
ent interpretations of the 
sense organs of insects that 
the student attempting to get 
information on the subject 
from books must soon be dis- 
couraged by their conflicting 
statements. Bul it must be 
realized that only intelligent 
guessing is possible where 
several senses are located on 
the same part. In the case of 
the bee some authors have 
ascribed even a third sense, 
that of hearing, to the an- 
tennae, but there is little evi- 
dence that bees possess the 
power of hearing. The senses 
of taste and touch are pos- 
sessed by the mouth parts, 
and some entomologists think 
that they contain organs of 
smell also. Thus, the organs 
of sight are apparently the 
only ones that can not be con- 
fused with some other sense. 
The best account of the 
antennal sense organs of the 
bee is that of Schiemenz (1883), whose drawings are here reproduced 
(fig. 12) and whose text is the basis of the following descriptions. 
The organs consist, as before stated, of modified hairs and their basal 




Fig. 12. — Antennal hairs and sense organs 
(after Schiemenz). A, example of antennal 
hairs {Hr) imbedded in cuticle (Ctl) but 
having no nerve connection ; B, hollow hair 
containing prolongation of special cell (CI) ; 
C, D, straight and curved tactile hairs con- 
nected with basal cells (CI) and nerve libers 
(Nv) ; E, conical hair (Hr) sunken in a pit 
(Pt) of the cuticle, probably an olfactory 
organ ; F, closed sac shut in by thin disc 
(hr) on surface of antenna and containing a 
delicately poised cell (CJ) with nerve con- 
nection (Nv). 



THE HEAD OF THE BEE AND ITS APPENDAGES. 37 

insertions which are connected with the ends of nerve fibers. Some 
of them stand exposed on tlie surface of the cuticle while others 
are sunken into, or entirely concealed within, pits of the integument. 
In addition to these, there are two other kinds of special hairs on 
the antennae which have no nerve connections, while, finally, the ordi- 
nary hairs, such as are found on all parts of the body, occur also on 
them, especially on the scape. 

The special hairs not provided with nerve endings are of two 
sorts. One is a solid curved or hooked hair (fig. 12 A, Hr) which 
is simply articulated into a socket of the cuticle (Ctl), wdiile the 
other (B) is hollow and is situated over a channel through the cuticle, 
and contains a prolongation of a speciall}^ enlarged epithelial cell 
(CI) \\ing beneath it. These hairs can not be regarded as sensory, 
since they have no communication with the central nervous system, 
and it is not clear just what purpose they do serve. 

The simplest sensory organ is a short, hollow, conical hair (C, 
Hr) arising directly from the surface of the cuticle, over a wide 
opening through the latter, and containing the end of a sensory cell 
(CI) connected with a nerve fiber (IVv), which goes into the main 
trunk of the axial antennal nerve. A modified form of this organ 
consists of a curved hair (D, Hr) set into a small depression over 
the cuticular channel. Such hairs are probably tactile in function; 
that is to say, by means of them the bee can perceive that its antenna? 
are in contact with some surface. The general integument is too 
thick and dense to allow of any sort of delicate touch sensation being 
communicated through it, but if one of these movable hairs brushes 
against an object the nerve within it must be at once stimulated. 
Tactile or touch hairs are distributed especially over the outer sur- 
face of the antennae and at its apex, but occur also scattered over 
the other parts of the body and on the mouth parts. 

Microscopic sections of the antennae reveal still other organs 
which are not so apparent on the surface as the hairs just described. 
One of these is shown at E of figure 12. It consists of a small pit 
(Pt) in the integument, widened basally, and having a small papilla 
on its floor, in whose summit is the opening of a still deeper cavity 
which also expands toward its deeper end. This inner cavity is 
almost filled up by a conical plug (Hr) which arises from its floor 
and ends just below the aperture into the outer pit. The plug con- 
tains a thick nerve ending Avhich arises from a ganglion cell con- 
nected with the antennal nerve by a nerve fiber. Ten or more of 
these sense organs occur on the terminal and the first three segments 
of the flagellum. It is evident that each is simply a sensory hair 
which has been doubly sunken into a cavity of the integument. 



38 THE AK ATOMY OF THE HONEY BEE, 

As before stated, it lias been conclusively proved by several investi- 
gators that bees perceive odors, and it is said that if the antenna? 
are covered Avith shellac, bees can distinguish between distasteful 
substances only by means of the proboscis. Schiemenz and most 
other writers on the subject therefore conclude that the sunken cones 
are the organs of smell, since, being below the surface, they could not 
be organs of touch. Some other authors, however, among whom are 
Cheshire, regard these inclosed cones as hearing organs. They sup- 
pose that the sound waves of the air enter the pit, as into an ear 
cavity, and these set up a vibration in the cone which stimulates the 
attached nerve ending. However, the appearance of one of these 
cones would suggest that it is too stable a structure to be affected 
by sound waves, so the olfactory theory seems much more probable. 

Finally, Schiemenz describes the most specialized of all the anten- 
nal sense organs as a closed cavity {Pt) in the cuticle {Ctl) extend- 
ing into the hollow of the antenna as a long, curved, tapering sac. 
This is shown at F of figure 12. A nerve {Nv) enters the lower . 
extremity of the pouch, expands slightly into a nucleated ganglion 
cell {01)^ and then extends toward the top as a delicate spindle 
drawn out into a fine tapering point. The surface covering of the 
pit is a thin layer of chitin presenting several concentric light and 
dark rings surrounding a central disc (Ar). Sections show that this 
appearance of rings is due to circular thickenings of the membrane, 
and Schiemenz points out that the central disc is probably a modi- 
fied hair, while the whole structure is to be regarded simply as a 
modification of a tactile organ such as that shown at D with the 
nerve-ending and its ganglion inclosed in a sac. These organs are 
most abundant on the antennae of the drones, where they are situ- 
ated, especially on the under surface, so close together that but little 
space is left between them for the tactile hairs, while in the workers 
and queens they are farther a^^art and are interspaced with many 
tactile hairs. Hence, whatever sense they accommodate must be 
much more highly developed in the males than in the females. 
Schiemenz described these organs, as well as the sunken cones, as 
organs of smell. He ascribed only the senses of touch and smell to 
the antenna3, and both Cheshire and Cowan concur in his view of the 
closed pits. Arnhart (1906), however, argues that an organ of smell 
must be open to the air in order to permit the ingress of odor par- 
ticles. Such an organ is constituted by the sunken cones, but the 
closed pits have nothing to recommend them for an olfactory func- 
tion. Arnhart then further points out that the buried sacs, inclosing 
a delicately poised nerve-ending and covered by an external tym- 
panum, have all the mechanical elements of an organ of hearing. 
He finally argues that bees must hear, since they produce special 
sounds such as the piping of the queens, and that, since no possible 



THE HEAD OP THE BEE AND ITS APPENDAGES. 39 

organs of hearing have been discovered on any other part of the body, 
some of the antennal sense organs must be auditory in function. His 
conckision from these premises is, of course, inevitable that the 
closed sacs on the antennte are the hearing organs of the bee. What 
invalidates the argument, however, is the fact that no one has yet 
produced any actual evidence that bees perceive sound. 

The following, then, may be stated as a general summary of the 
evidence concerning the antennal senses and their sense organs in 
the bee: (1) The antenna^ are highly sensitive to touch and are the 
seat of the sense of smell. (2) They are covered by several kinds 
of minute structures which are modified hairs containing special 
nerve-endings. (3) By inference, it would seem certain that these 
are the sense organs, but we can only form an opinion, based upon 
their structure, as to which are tactile and which olfactory. (4) One 
set of organs does not appear to belong to either of these categories 
and their structure suggests an auditory function, but, in the absence 
of evidence that bees hear, the purpose of these organs must be re- 
garded as problematical. 

3. THE MANDIBLES AND THEIR GLANDS. 

The mandibles (fig. 9 A, Md) are the dark, strongly chitinous 
appendages of the head, commonly called the jaws, situated at each 
side of the mouth, anterior to the base of the proboscis. In all in- 
sects with biting mouthy parts the jaws work sidewise, each being 
attached to the head by an anterior and a posterior articulation. 
They can thus swing in and out on a longitudinal axis in such insects, 
as the bee, that carry the head with the mouth directed downward, 
or in the same way on a vertical axis in those that carry the head 
with the mouth forward. 

Both mandibular articulations are of the ball-and-socket type, 
although in the bee the socket is a very shallow one, the anterior 
consisting of a condyle on the outer angle of the clypeus fitting 
against a facet on the mandible, and the position of a facet on the 
lower edge of the postgena receiving a condyle from the mandible. 
The motion of the mandible is thus reduced to a hinge- joint moA^e- 
ment, and, on this account, insects can only bite and crush their 
food; they can not truly chew it, since their jaws are incapable of 
a grinding motion. Each mandible is, of course, as pointed out in 
the introduction, really suspended from the head by a continuous 
membrane between its base and the cranium, being simply a modified 
saclike outgrowth of the head wall. The two articulations are pro- 
ductions of the chitin on the outside of this membrane. 

Figure 9 A shows the location and shape of the mandibles {Md) 
of the worker as seen in a facial view of the head. Figure 11 A 



40 



THE ANATOMY OF THE HONEY BEE, 



shows the appearance of the left mandible in side view, while the 
right one is shown detached from the head in fignre 13 A. The 
mandibles differ conspicuously in size and shape in the three forms 
of the bee as already described and as shown in figure 10 A, B, and C. 
That of the worker is hollowed out somewhat on the distal half of 
its inner face (fig. 13 A, Md) forming a sj^oon-shaped organ, the 
edge of which is smooth and rounded. The mandibles of both the 
queen (fig. 10 B) and the drone (C), however, are pointed at the 
apex and have a conspicuous subapical notch. Those of the drone 

■TQ, I are smaller than those of 

either form of the female, 
but appear to be especially 
small on account of the 
great size of the drone's 
head. The mandible of the 
worker is undoubtedly to 
be regarded as the special- 
ized form, since the notched 
mandible of the drone and 
queen is of the ordinary 
Hymenopteran type. Both 
the drone and the queen 
are, under normal circum- 
stances, fed almost entirely 
by the workers, and they 
probably never have any 
use for their jaws as feed- 
ing organs. The queen 
needs her large, sharp- 
pointed mandibles for bit- 
ing her Avay out of the 
thick wax cell in which 
she is reared, but the 
drone, on the other hand, 
being reared in an ordinary cell resembling that of a worker, except 
in size, is easily able to cut through the thin cell cap with his com- 
paratively weak jaws. The workers, however, have numerous uses 
for their mandibles, such as biting through the cell caps, eating 
pollen, and modeling wax. The last is the especial function of 
the worker mandible, and probably it is to accommodate this pur- 
pose that it has acquired its specialized spoonlike shape. 

Each mandible is moved by two sets of muscles within the head. 
The outer one constitutes the extensor muscle (fig. 13 A, EMcl) and 
the inner the flexor muscle {RMcl). The latter is the stronger of 




Fig. 13. — A, right mandible of workei% anterior 

view, witli extensor and flexor muscles (EAIcl 

and RMcl) and mandibular glands (iMdGl) at- 

tacbed ; B, corresponding view of mandible of 

drone, with muscles cut oft a short distance from 
their bases. 



THE HEAD OE THE BEE AND ITS APPENDAGES. 41 

the two, since all the work of the mandible falls upon it, the ex:tensor 
being used simply to open the jaw. While these muscles have their 
origins on the walls of the head, they are not inserted directly upon 
the mandibles, but on large apodemes (fig. 13 A, EAp and RAp) 
attached to the edges of the mandible. 

A gland opens at the inner margin of each mandible between the 
anterior articulation and the base of the apodeme of the flexor 
muscle (fig. 13 A and B, IMdGX). In the worker it consists of a 
. large sac covered with secreting cells lying within the front part of 
the head between the clypeus and the compound e^^e (fig. 10 A, 
IMdGT). These mandibular glands may be most easily studied by 
removing the front as shown in figure 10 A, B, and C. In order to 
do this, pull the head from the thorax and allow the prothoracic legs, 
which will usually come off with the head, to remain attached to it. 
Next melt a small hole in the bottom of a paraffin dish with a heated 
needle and fasten the head face upward into this, the attached legs 
helping to anchor the head in the paraffin. Cover the specimen with 
weak alcohol and by means of sharp needles remove the part of the 
front on either side between the clypeus and the lower half of the 
compound eye in the worker and drone and the entire front of the 
queen. In figure 10 the whole front is removed in all three forms in 
order to expose other internal parts of the head. 

The mandibular gland {IMdGT) is of greatest size in the queen 
(fig. 10 B), though it is large in the worker (fig. 10 A and fig. 13 A), 
but it is reduced in the drone (fig. 13 B) to a very small oval sac, 
which is hidden by another gland {%Gl) in front (fig. 10 C). It was 
first described by Wolff (1875) as an olfactory mucous gland {Riech- 
schleimdrilsse) and was supposed by him to secrete a liquid which 
was poured upon the roof of the mouth in order to keep this surface, 
on which Wolff thought the olfactory organs were located, in a moist 
condition capable of absorbing odor particles. There is absolutely 
no evidence, however, of the presence of organs of smell in the mouth, 
and furthermore, as pointed out by Schiemenz (1883), the gland 
varies in the three forms of the honey bee according to the size of the 
mandible, which is proportionately largest in the queen and smallest 
in the drone. Of the three, we should expect the drone or the worker 
to have the sense of smell most highly developed, and hence, even if 
we did not know that the sense of smell is located in the antennae, 
it would seem more reasonable to suppose that the glands of the 
mandibles are connected in some way with the functions of these 
organs themselves. 

The mandibles, as already stated, are used for eating pollen and as 
tools for manipulating and modeling wax. Therefore, according to 
Arnhart (1906), since the queen does not eat raw pollen, the product 



42 



THE ANATOMY OF THE HONEY BEE. 



of the mandibular glands must be intended for softening the wax 
when it is w^orked in the jaws. The secretion of the glands is said 
to be very volatile and strong smelling and to have an acid reaction. 
It is iDrobabl}^ entirely possible that it may have a solvent etfect upon 
the wax, or even, when mixed with it, change somewhat the chemical 
composition of this substance; hi fact, some investigators claim that 
the wax of the comb differs chemically from that freshly taken from 
the wax plates. Even this explanation, however, does not seem en- 
tirely satisfactory, for the only occasions on which the queen has any- 
thing to do with wax is when she gnaws her way out of her cell after 
hatching or bites her way into the cells of young queens in order 
to sting them. Howes^er, these occasional uses by the queen of her 
mandibles appear to be imjDortant enough to maintain the large size 
of these organs in the queen, and it ma}^ be reasonable to assume that 
the demand upon their glands is likewise a large one when it does 

occur. Yet the mandibles of the 
queen are toothed and sharp 
pointed, Avhich should provide her 
with sufficient cutting power both 
to emerge from her own cell and to 
enter the cells of other queens, and 
so, on the whole, the opinion of 
Schiemenz that the secretion of the 
mandibular glands is merely sali- 
vary in function would seem to be 
the simplest explanation and the 
most logical one. However, an 
actual test should certainly be made 
to determine whether the worker's 
manipulation of the wax with her mandibles produces any change in 
it, and to discover whether the queen simply bites her way mechan- 
ically through the wall of the cell or at the same time softens the wax 
by a secretion from her mouth. The male in any case has little use 
for his mandibles, and the glands are so small that they must certainly 
be functionless. 

A second mandibular gland (fig. 14, 2MclGl) is present in the 
worker. It consists of a delicate, flattened, racemose mass lying 
against the internal face of the wall of the fossa of the proboscis, 
whose duct opens into the mouth cavity at the posterior inner edge 
of the mandible. This gland was first described by Borclas (1895) as 
the internal mandibular gland. According to him, it corresponds 
with a similar gland in the Bombidae (bumblebees) and in the Ves- 
pidse (yelloAv jackets) and to the maxillary glands of other Hy- 
menojDtera. Nothing is known of its secretion. 




2MdGl 

Pig. 14. — Internal mandibular gland 
(2M4IGI) of worker, lying against inner 
wall of postgena (Pge) and opening 
(Dot) at inner edge of base of man- 
dible. 



THE HEAD OF THE BEE AND ITS APPENDAGES. 43 

4. THE PROBOSCIS. 

The conspicuous group of mouth appendages in the honey bee, 
forming what is commonly known as the prohoscis (fig. 9 A, Prh), 

Lbl 

Hr-m9 




Fig. 15. — Mouth parts of the worker: A, tip of glossa, showing labellum {Lhl), guard 
hairs (Hr), and ventral groove (fc) ; B, same, from above; C, small piece of glossal 
rod (r) with adjoining parts of walls {q) of glossal canal attached, showing ventral 
channel (I) guarded by rows of hairs. D, parts forming the proboscis, labium in middle 
and maxillse at sides, flattened out, ventral view ; Ei, cross section of glossa showing its 
invaginated channel (Luin) and position of rod (r) along its dorsal wall, and likewise 
position of channel (l) of rod along median line within the glossal channel; F, end of 
mentum {Mt) and bases of ligula (Lg) and labial palpi (LhPlp), showing opening 
of salivary duct (SalDO), dorsal view; G, lateral view of proboscis showing parts on 
left side; H, lateral view of glossa (Gls) with its rod (r) torn away at base showing 
attachment of retractor muscles {2RMcl). 

by means of wliich the bee takes up liquid food, consists of what cor- 
respond with the maxillae and .the labium of insects that feed on solid 



44 THE ANATOMY OF THE HONEY BEE. 

food alone. By separating the parts of the proboscis a little (fig. 
9 B) it will be seen that, while there are five terminal pieces present, 
three of them arise from one median basal sclerite {Mt)^ the two 
wider lateral appendages {Mx) being carried each by a separate lat- 
eral basal piece {St). The median group constitutes the lahmin and 
the separate lateral parts the maxillce. 

If the reader will now turn again to figure 3 C (p. 17) , which may 
represent any generalized insect labium, and compare with it the 
draAving of the bee labium, forming the median series of parts in fig. 
15 D, he will at once be able to identify the parts of the latter. The 
principal elongate median basal plate is the mentum {Mt) , the small 
triangular plate at its base is the siihmentnm (Smt) , and the two 
jointed lateral appendages of the mentum are the labial palpi 
{LhPl])), each carried by a basal palpiger {Pig). It is only the parts 
of the bee's labium that lie between the palpi which are actually 
different from those in the generalized diagram where they consist 
of the four lobes of the ligiila {Gls and Pgl). But even here it will 
be seen that the two small lobes {Pgl) in the bee's labium, partly con- 
cealed within the bases of the palpi, correspond with the paraglossa. 
Hence we have only the long median appendage to account for and it 
is unquestionably the representative of the glossce {Gls) which are 
here fused together and drawn out into this flexible tonguelike organ. 
In fact, a comparison with the mouth parts of other Hymenoptera in 
which the elements are much less modified leaves no doubt of this 
being the true interpretation of the bee's labium. It is simply an 
example of how nature constantly prefers to modify an already exist- 
ing part to serve some new purpose rather than to create a new organ. 

If, then, we bear in mind that the slender median appendage of 
the bee's labium represents the glossse of other insects, we may for 
convenience call it the " tongue," as it is popularly termed, or, since 
it is a single organ, there is probably no grammatical objection to 
calling it the glossa. The word " tongue," however, to use it prop- 
erly, should be applied to the true lingua or hypopharynx (fig. 3 C 
and D, Ilpliy) which arises from the upper surface of the labium. 
Many of the older entomologists, adopting the notion from Kirby 
and Spence, who defined the term in 1826, regarded the glossa of 
the bee as the homologue of the lingua in other orders. Even Pack- 
ard in his Text -book of Entomology calls the glossa the " hypo- 
pharynx," Cheshire named it the " ligula," and his mistake has been 
perpetuated by several other writers on bee anatomy, including Cook 
and Cowan. The term ligula properly includes both the glossa and 
the paraglossse, or should signify the basal piece from which these 
four lobes arise (fig. 3 C, Lg)-> so that it can not be applied to the 
fflossa alone. 



THE HEAD OF THE BEE AND ITS APPENDAGES. 45 

The derivation of anatomical names counts for nothing in their 
application — this must be determined by scientific usage and priority. 
Thus, glossa is the Greek word for " tongue," but it was first applied 
in entomology to the median lobes of the labium ; lingua is its equiva- 
lent in Latin and was first given to the true tongue or hypopharynx 
in insects ; ligula is a diminutive derivative from " lingua " and has 
come to be applied collectively to the terminal parts of the labium 
beyond the mentum but not including the palpi. Hence, all these 
words mean the same thing by their origins, but their anatomical 
applications should be carefully distinguished. In this paper there- 
fore the slender median appendage {Gls) of the labium will be 
called the glossa, or, for convenience, the tongue, but with the strict 
understanding that the organ in question is not the true tongue. 
This latter should be called the " hypopharynx," but, as will be shown 
later, it is absent in the bee. 

The glossa of the bee (figs. 9 B; 11 A and B, and 15 D, F, and G, 
Gls) is covered with long hairs which increase in length toward the 
end. The tip is formed of a small spoon-shaped lobe, the lahellum or 
bouton (Lbl), which is covered by short delicate processes branched 
at their ends (fig. 15 A and B, LM). The long hairs of the glossa 
are arranged in circles and the transverse rows of hair bases give 
the tongue a multiarticulate appearance. Surrounding the dorsal 
side of the base of the labella and forming two short subterminal 
rows on the ventral side of the glossa are a number of stiff, out- 
wardly curved, spinelike hairs (Hr). These hairs have been de- 
scribed as taste organs, but their appearance would suggest that they 
are simply protective spines guarding the delicate tip of the tongue. 
Between the two ventral rows of these spines is the termination of 
a groove (A, k) which extends along the midline of the under sur- 
face of the glossa (D, k) to its base (fig. 9 B, k). The cleft of this 
groove is covered by two fringes of converging hairs whose tips are 
inclined also toward the tip of the tongue. 

Let us now return to a study of figure 15 D. The series of lateral 
pieces as already explained are the maxillae. A comparison with 
figure 3 B representing a generalized maxilla will show that these 
organs in the bee have suffered a greater modification than has the 
labium, but the parts can yet be quite easily made out. The main 
basal plate (St) is the combined stipes, suhgalea, and palpifer, the 
basal stalk is the cardo {Cd), and the little peglike process {MxPlp) 
at the outer end of the stipes is the greatly reduced maxillary palpus. 
Hence, we have left only the terminal bladelike lobe {Mx) to account 
for, and it is evident that it must be either the galea or the lacinia 
(see fig. 3 B, Ga and Lc) or these two lobes combined. Here again 
a comparative knowledge of the mouth parts of Hymenoptera comes 



46 THE ANATOMY OF THE HONEY BEE. 

to our aid and shows clearly that the part in question is the outer 
lobe or galea^ for the inner one becomes smaller and smaller in the 
higher members of the order and finally disappears. 

The base of the submentum is connected in the bee with the upper 
ends of the cardines by a flexible, widely V-shaped band, the lorum 
(Lr). The posterior angle of the submentum rests in the apex of the 
lorum, while the tips of the loral arms are movably articulated with 
the distal ends of the cardines. The name " lora " was given to this 
structure by Kirby and Spence, but " lorum " is more correct, since 
this is the Latin form of the word (meaning a thong or lash). Some 
recent entomologists have spoken of the structure as consisting of 
two rods, thus making the word do duty as a plural, but the thing 
itself is all one piece. Cheshire and some others have incorrectly^ 
ajDplied the name to the submentum. 

The lorum is peculiar to the Hymenoptera, and the reason for it 
is clear when we examine the attachments of the parts of the proboscis 
to the head. As already stated, the maxillae and labium are sus- 
pended in a large cavity on the back of the head which may be' called 
the fossa of the 2)rohoscis (fig. 9 B, PrhFs). The maxillae are articu- 
lated by their cardines (Cd) to the maxillary suspensoria (fig. 11 
A, e) at the upper edges of the side Avails of the fossa. The labium, 
on the other hand, is not attached to the solid walls of the cranium 
but is suspended in the membranous floor of the fossa. This is to 
afford it freedom of movement during feeding, but, in order to 
giye it' more substantial support and to make the regulation of its 
motions possible, the submentum is slung to the ends of the cardines 
by the lorum. 

The terminal lobes of the labium and maxillae when not in use 
are ordinarily folded down beneath the head against the mentum 
and stipites (fig. 19). When, however, the bee wishes to imbibe a 
thick liquid such as honey or sirup in large quantity, these parts are 
straightened out and held close together so as to form a tube between 
them leading into the mouth, the terminal joints of the labial palpi 
alone diverging from the rest (fig. 11 A). 

The action of the mouth parts while feeding may be observed quite 
easily if some bees are given a small amount of honey and then 
watched through a lens while they are eating. A most convenient 
method is to put a few workers in a small screen-covered cage, such 
as are used for queen nurseries, spread a small drop of honey on the 
wire, and then place the cage under a simple microscope. It will be 
seen that the maxillae are held almost stationary but that the base 
of the labium slides back and forth between the maxillary bases 
with a very regular to-and-fro movement as if the honey were being 
either pumped or sucked up into the mouth. It is probable that there 
is a sucking force exerted by the pharynx (fig. 11 B, Plit/) but not 



THE HEAD OF THE BEE AND ITS APPENDAGES. 47 

by the honey stomach (fig. 44, H/S), which latter, as Cheshire re- 
marks, could no more suck honey through the oesophagus than a 
balloon could suck gas from a pipe. The liquid undoubtedly runs 
up the temporary tube betAveen the blades of the mouth parts first 
by capillary attraction, but it must be greatly assisted along its way 
to the mouth by the retraction of the labium. The load brought up 
when this is pulled back is then sucked into the mouth by the 
pharynx while the labium immediately goes out again after more. 
It acts thus as a sort of mechanical feeder and this function is prob- 
ably derived from the lapping motion of the under lip in wasps and 
hornets. 

The mentum (fig. 15 D and G, 3It) is hinged freely upon the 
submentum {Smt) ^ the latter, as already described, is set into the 
socketlike angle of the lorum, while, finally, the arms of the lorum 
are articulated to the distal ends of the carclines of the maxillae. 
Now, when the labium is retracted by means of muscles attached to 
the mentum, the submentum turns in the loral socket and assumes a 
position at right angles to the mentum while the lorum itself turns 
somewhat on its articulations with the cardines. This great freedom 
of motion is permitted b}^ the loose membrane of the fossa in which 
both the maxillae and the labium are suspended. 

The observer, however, can not fail to note that beside this motion 
of the entire labium the tongue itself, or glossa (Gls) , performs a 
conspicuous independent movement of its own. It is b}^ far the most 
active member of the mouth parts during feeding, being actively 
thrust out and retracted while its tip is constantly moved about in 
a way suggestive of its being delicately perceptive of taste or touch 
or perhaps to both of these senses. So great is the retractile power 
of the tongue that its tip, which normally extends far beyond the end 
segments of the labial palpi, can be drawn back entirely within the 
latter. This contractile activity appears at first sight to be due to 
elasticity, but a closer examination will show that the entire ligula, 
i. e., the paraglossse {Pgl) as well as the glossa {Gls) , moves back 
and forth and that the action is due to a retraction of the base of the 
ligula (fig. 15 F, Lg) into the anterior end of the mentum (Jit). 
The ligula is supported on a membranous cone at the end of the 
mentum whose walls are strengthened by three thin chitinous plates, 
two above (F, p) and one below (D, o). By the contraction of 
muscles situated within the mentum (fig. 16, IRMcl) and inserted 
upon the base of the ligula the latter is pulled into the end of this 
cone whose walls, including the chitinous plates, simply turn inward. 

But the tongue does possess also a contractile power of its own by 
means of which it actually shortens its length. A flexible rod arising 
from the median ventral supporting plate (fig. 15 D, o) of the ligula 
extends throughout its length. The base of this rod is curved down- 



48 THE ANATOMY OF THE HOISTEY BEE. 

ward and has two muscles attached to it. This is shown by figure 
15 H, where the rod (r) is torn from the glossa {Gls) basally so as 
to show the muscles {2RMcl) inserted upon it and its connection 
with the plate {o). By the contraction of the muscles the rod bends 
at its base and is drawn back into the mentum. The glossa thus 
shortens and becomes bushy just as does a squirrel's tail when one 
attempts to pull the bone out of its base. 

The protrusion of the parts is due to the pressure of blood driven 
into the ligula from the mentum, while probably the glossa extends 
also by the straightening of its rod as the muscles relax. Wolff 
described a protractor muscle at the base of the ligula. The rod of 
the tongue is certainly not in itself contractile, as supposed by 
Cheshire, who looked for evidence of muscular striation in it. It has 
mostly a transparent and cartilaginous appearance, but is presumably 
chitinous. 

The mouth parts, their action in feeding, and the muscular mech- 
anism by which they are moved have been elaborately described 
and illustrated by Wolff (1875) in his monograph on the organs of 
smell in bees. Most unfortunately, however, Wolff's paper was 
written to show that the seat of the sense of smell is in the mouth, 
a most erroneous notion, and the title of his paper based on this 
notion has caused little attention to be paid to this work on the mouth 
parts of the bee, which is one of the best anatomical treatises ever 
produced on the mouth parts of any insect. 

It still remains for us to describe the details of the glossa and its 
particular function in feeding. The tongue is not a solid appendage 
nor yet is it truly tubular. A compromise is effected by the longi- 
tudinal groove (fig. 15 A and D, k) on its ventral surface which 
expands within the tongue into a large cavity occupying half of its 
interior (E, Lum). The glossal rod (r), which has already been 
mentioned, lies in the dorsal wall of this channel and is, hence, 
really not an internal but an external structure. The rod is itself 
grooved along its entire ventral length (E, Z) and this groove again 
is converted into a tube by two rows of short hairs which converge 
from its margins. The lips of the ventral groove of the glossa are 
so deejDly infolded that its cavity is almost divided along the midline. 
Hence, the glossa might be described as containing three channels — 
a small median dorsal one {I) and two large latero- ventral ones 
{Luin). 

The glossal rod (fig. 15 C, r) is very flexible but not contractile, as 
already stated, and is mostly clear and cartilaginous in appearance, 
its ventral groove {I) alone being lined by a deposit of dark chitin 
(fig. 15 C and E). Its shape in section is sufficiently shown by the 
figures. The walls of the large channels of the proboscis consist of 
a delicate membrane (C and E, q) covered with very small hairs. 



THE I-IEAD OF THE BEE AND ITS APPENDAGES. 49 

The entire ventral cavity {Lum) with the rod (r) can be evaginated 
through the ventral cleft {k) by blood pressure from within. As 
Cheshire points out, this permits of the channels being cleaned in 
case of clogging by pollen or any foreign matter. 

It is supposed that these glossal tubes are of especial service to the 
bee by enabling it to take up the smallest drops of nectar- -quantities 
that would be lost in the clumsy tube formed between the parts of 
the iabium and the maxillae. The suction must be in large part 
capillary attraction, but here again the shortening of the glossa by 
the retraction of its rod must squeeze the contained nectar out of the 
upper ends of the channels where it is received upon the ventral flaps 
of the paraglossse (fig. 15 F, Pgl), from Avhich it runs around the 
base of the tongue (Gls) within the paraglossse to the dorsal side of 
the mentum (Mt) and so on to the mouth. 

The maxillse and labium of both the queen and the drone (fig. 11 
B) are smaller and weaker than those of the worker, and neither of 
these two forms is capable of feeding itself to any extent. If a 
hungry queen be given some honey she attempts to eat it and does 
imbibe a small quantity, but at the same time she gets it very much 
smeared over her head and thorax. 

The mouth is hard to define in insects; practically it is the space 
surrounded by the bases of the mouth parts, but strictly speaking it 
is the anterior opening of the alimentary canal situated behind the 
bases of the mouth parts (fig. 19, Mth). Yet the enlargement of the 
alimentary canal (Phy) immediately following this opening is never 
spoken of as the mouth cavity but is called the pharynx. On the 
other hand the so-called epipharynx (Ephy) and hypopharynx 
(absent in the bee) are located in front of this opening and are con- 
sequently not in the pharynx at all, the former being attached to the 
under surface of the labrum and clypeus, while the latter is situated 
on the upper surface of the base of the labium. These and numerous 
other inconsistencies in the nomenclature of insect morphology have 
to be endured because the parts were originallj^ named for descrip- 
tive purposes by entomologists who were not familiar with scientific 
anatomy. In this paper the term mouth will be applied to the true 
oral opening (fig. 19, Mth). The space in front of it between the 
bases of the mouth parts may be called the preoral cavity. 

The duct of the salivary glands of insects in general opens upon the 
base of the labium in front of the hypopharynx. In the honey bee 
the salivary opening is on the dorsal side of the base of the ligula 
between the paraglossse (fig. 15 F, SalDO). This alone would show 
that the glossa is not the hypopharynx of the bee, as many authors 
have supposed, for otherwise the opening of the salivary duct should 
be ventrad to the base of the glossa. In fact, this makes it clear that 

22181— No. 18—10 4 



50 THE ANATOMY OF THE HONEY BEE, 

the bee does not possess a hypopharynx. There is, however, a con- 
spicuous chitinous plate located on the anterior part of the floor of the 
pharynx (fig. 19, s) having two terminal points hanging downward 
over the lower lip of the oral aperture, but, although this plate is truly 
hypopharyngeal in position, it is not the homologue of the organ 
called the hypopharynx in other insects. It is variously developed 
in all Hymenoptera, being simply a chitinization of the floor of the 
pharynx, and should be called the pharyngeal plate {Schlundhein of 
Wolff) . It will be more fully described in connection with the ali- 
mentary canal. If a hypopharynx were present it should be situated 
on the u]3per side of the labium (see fig. 3 D, Hphy) but there is here 
present only a plain arched membranous surface in the honey bee 
and other typical Hymenoptera. 

The external location of the salivary opening enables the saliva 
to be mixed with the food before the latter enters the mouth. This 
is necessary in insects since the jaws are also on the outside of the 



SaD 
^B---iRMcl 




Lum ^ /, ^^^^^^^i^^^^^^W-- 2 RMcI 



Fig. 16. — Median section tlirougti distal half of mentum (Ifi) and base of ligula (Lfir) 
of worlser, stiowing opening of salivary duct (SalDO), and muscles connected with 
ligula and the "salivary syringe" (t). 

mouth, and wdiatever chewing or crushing the food receives from 
them is consequently done in the preoral cavity. 

In some insects the saliva is used for other purposes than diges- 
tion. For example, the saliva of some preclaceous insects with pierc- 
ing mouth parts belonging to the order Hemiptera is poisonous, and 
when one of these insects " bites," the saliva is injected into the 
wound by a sjDecial j^ump. The bite of the mosquito is made painful 
likewise by an irritant secretion from a part of the salivary glands. 
Bees appear to have the power of letting their saliva run down the 
tongue when necessary to dissolve a hard substance like sugar and 
render it capable of being taken up in solution, for they, do not eat 
sugar with their mandibles. Moreover, there is even a sort of pump 
or so-called " salivary syringe " at the termination of the salivary 
duct in the ligula, bj^ means of which the secretion can be forcibly 
ejected from the opening. 

The salivary opening on the base of the ligula (fig. 15 F, SalDO) 
leads into a deep transverse pit with collapsible cartilage-like walls 
having its deepest part turned horizontally toward the base of the 



THE HEAD OF THE BEE AND ITS APPENDAGES. 



51 



labium (fig. 16, t). The salivary duct {SalD) bends downward in 
the anterior part of the mentum {Mt) and opens into the posterior 
end of the pit (z^). When the retractor muscles {IRMcl) of the 
ligula pull the latter back into the mentum the lips of the salivary 
pit must necessarily be closed. The simultaneous contraction of the 
elevator muscle (w) attached to the roof of the horizontal part of the 
pit must expand the latter and suck the saliva from the salivary duct. 
"\^'lien, finally, these muscles relax and the ligula is driven out by 
blood pressure in the mentum, probably produced in part by the 
contraction of its dorsal transverse muscles {T3Icl), the saliva -in 
the temporarily formed bulb must be squirted out upon the base of 
the tongue. Wolff (1875) calls each dorsal longitudinal muscle- of 
the mentum {IRMcl) — the two inserted upon the basal hooks {n) of 
the glossa (fig. 15 H and fig. 16) — the retractor linguce longus. The 
large ventral retractor muscle of each side {2RMcl) he calls the 
retractor linguoe, hicejys since its anterior end divides into two parts, 
one of which is inserted by a tendonous prolongation upon the base 
of the glossal rod (fig. 15 H and fig. 16, r) and the other upon the 
base of the ligula. The use of the word " lingua " in these names is 
objectionable because, as already explained (page 45), the lingua is 
properly the true tongue or hypopharynx. " Ligulse " should be sub- 
stituted for " linguse." The dilator muscle (fig. 16, u) of the salivary 
pit {t) is termed the ijrotractor linguce, by Wolff because, as. he sup- 
poses, when the ligula is pulled back into the mentum the position 
of this muscle is reversed, so that a contraction of its fibers would 
help to evert the ligula. 

The glands that furnish the saliva lie within the head and the 
thorax and will be described later in connection with the alimentary 
canal and the process of digestion. 




Fig. 17. — Epipharynx (Ephy) and labrum (Lm) of worker: A, ventral view; B, 

anterior view. 



5. THE EPIPHARYNX. 

The epipharynx of insects in general may be described as a dorsal 
tongue, it being a median lobe developed on the roof of the preoral 
cavity from the under surface of the clypeus or labrum and situated 
opposite the hypopharynx. 



52 



THE ANATOMY OF THE HONEY BEE. 




The epipharynx of the bee is a large three-lobed appendage de- 
pending from the roof of the preoral cavity just in front of the mouth 
(fig. 19, Ephy). Seen from below it is triangular (fig. 17 A) with 
the apex forward. Its median lobe has the form of a 
high, vertical, keel-like plate, while the lateral lobes 
are rounded but have prominent elevated edges con- 
verging toward the front of the keel. The appearance 
in anterior view is shown by figure 17 B. Situated 
on the posterior parts of the lateral lobes are a num- 
ber of sense organs, each consisting of a small cone 
with a pit in the summit bearing a small hair (fig. 18). 
These are regarded as organs of taste. 
Wolff (1875) made a most thorough study of the epipharynx, 
which he called the "palate sail" {Gaumensegel) on account of the 
high median crest. His drawing is the standard illustration of the 
organ found in nearly all books on the anatomy of the honey bee 



Fig. 18. — Sense 
organs, prob- 
ably of taste, 
from epi- 
pharynx. 




Fig. 19. 



-Median longitudinal section of head of worker, but with entire labium attached, 
showing internal organs except muscles and brain. 



and in most works on general insect anatomy and the sense organs. 
Wolff, however, regarded the sensory cones as having an olfactory 
function, and this led him to erroneous conclusions regarding the 
functions of several other organs. For example, he thought that 
the mandibular glands jDOured a liquid upon the surface of the 



THE THORAX AND ITS APPENDAGES. 



53 



Ant 



ten 



epipharynx which kept it moist and capable of absorbing odor 
particles, while he ex- 
plained the inhalation 
of the latter into the 
preoral cavity as 
brought about through 
the contraction of the 
air sacs situated about 
the mouth. Wolff's 
anatomical researches 
are without doubt 
some of the best ever 
made on the bee, and 
it is due to his mis- 
taken idea of the loca- 
tion of the sense of 
smell, which, as al- 
ready explained, is on 
the antenna", that we 
have received from 
him a most excellent 
account and detailed 
drawings not only of 
the epipharynx but of 
the mandibular glands, 
the mouth parts, the 
salivary " pump," and 
the respiratory organs. 

IV. THE THORAX AND 
ITS APPENDAGES. 

1. THE STRUCTURE OF 
THE THORAX. 

The apparent thorax 
of the bee (fig. 20, 
T^-IT, and fig. 21)' 
and of most other 
Hymenoptera is not 
exactly the equivalent 
of the thorax in other 
insects. The middle 
division of the body, 
so conspicuous in this 
order, consists not only of the three leg-bearing segments, which alone 




Fig. 20. 



Dorsal view of ventral walls and internal skele- 
ton of body of worker. 



54 



THE ANATOMY OF THE HONEY BEE. 



constitute the thorax of all other insects, but also of the first ab- 
dominal segment. The conspicuous necklike constriction posterior to 
the base of the hincl legs (fig. 21, Pd) is, therefore, between the first 
and the second abdominal segments (fig. 1, 77". and IIT). 

The thorax of the honey bee at first sight looks entirely different 
in structure from that of all other insects except related Hymenoptera, 
in the higher families of which group it is more highly modified than 
in any other order of the whole series of insects. When, however, we 
examine the thorax of one of the lowest members of the Hymenop- 
tera, such as a sawfly, we are surprised to find that, in each segment, 
the structure agrees very closely with our ideal diagram of a general- 
ized thoracic 

i-2 segment (fig. 4). 

The three seg- 
ments are per- 
fectly distinct, 
and the first 
abdominal seg- 
ment, while it 
may be clearly 
separated from 
the rest of the 
abdomen, is not 
fused into the 
thorax so as to 
appear to be a 
part of it. If, 
now, we exam- 
ine representa- 
tives of several 
families inter- 
mediate between 
the sawflies and 

the bees, the line of specialization that has produced the bee thorax 
becomes perfectly evident. The principal features in these modifi- 
cations are the following: 

(1) The lateral and ventral parts of the prothorax (figs. 20 and 21, 
Ejjs^ and S-^) are suspended loosely in a large membranous area 
which is continuous anteriorly as the neck. They thus form a sort 
of suspensorium for the front legs, which appears detached from the 
rest of the thorax. (2) The protergum {T^) is solidly attached to 
the anterior edge of the mesothorax and its lateral parts extend 
downward till they meet on the venter behind the prosternum (figs. 
20 and 21). (3) The postnotum (postscutellum) of the mesothorax 
(figs. 22, PN ; 23 A, PN.) is entirely invaginated into flie cavity of 
the thorax and is reduced to the form of two lateral arms of the large 




Fig. 21. — Thorax of worker, left side, with intersegmental lines 
somewhat .exaggerated, showing prothorax (T^, Eps^, Gx{), 
mesothorax (To, Epso, Epnio, 82, Cx^) , metathorax (To, Pl^, 
pis, CXi) and propodeum or first abdominal segment (IT). 



THE THORAX AiSTD iTS APPENDAGES. 56 

internal poslphragma (Ppli) which has no median tergal connection 
at all. (4) The metatergum (figs. 21 and 23 A, 7'^) consists of a 
single narrow plate. (5) The metapleurmn (fig. 21, Pl.^ and pl.^) 
shows no trace of a division into episternuni and epimerum, but is 
divided into an upper (Pis) and a lower (pl^) pleural plate. (6) 
The first abdominal tergum (fig. 21, IT) is solidly attached to the 
metathorax and forms an intimate j^art of the thoracic mass. 

We shall now proceed with a more detailed account of the thorax, 
and the reader should occasionally turn back to figure 4 (p. 19) in 
order to keep clearly in mind the parts that make up a generalized 
thoracic segment. 

The parts of the prothorax are so separated from each other that 
they appear to belong to different segments. The protergum (fig. 21, 
7\) forms a collar completely encircling the front of the mesothorax. 
On each side a large lobe (it') projects posteriorly as far as the base 
of the front wing and constitutes a protective shield over the first 
thoracic spiracle. The tergum presents a median transverse groove, 
dividing it into an anterior and a posterior part, which parts may 
be called the scutum (fig. 23 A, T^, Set) and scutellum (Sd). The 
propleurum (figs. 20, 21, Eps.^^) consists of a large plate presenting 
both a lateral surface (fig. 21) and a ventral surface (fig. 20). On 
account of the position of the coxal articulation (fig. 21) this plate 
would seem to be the anterior pleural plate alone (see fig. 4), which 
IS the episternum. In some Hymenoptera the epimerum is repre- 
sented by a very small plate on the rear edge of the episternum. 
The anterior ends of the two episterna form knobs which loosely 
articulate with the occipital region of the head (figs. 11 B, 20, and 
21). Lying just ventrad of each is a slender cervical sclerite (fig. 21, 
7ni). The prosternum (^S^) is shown by figure 20. It carries a large 
entosternum {Fi(^), forming a bridge over the nervous system behind 
the prothoracic ganglion (fig. 62). 

The mesoterg-um, as seen in its natural position (fig. 21, T.,), consists 
of a large anterior scutum {Scto) and of a smaller but very prominent 
posterior scutellum (ScL,), separated by a very distinct suture (v). 
The scutellum has two latero-anterior areas partially separated from 
the median area by sutures. AVlien the mesotergum is detached from 
the rest of the thorax (fig. 22) it is discovered that there is attached 
laterally to the scutellum a large posterior internal part, which does 
not show on the surface at all. This is the representative of the 
postscutellum (Pscl) and its phragma {Pph) constituting the post- 
notum (PN) of our diagrammatic segment (fig. 4). The proof of 
this, again, is to be derived from a study of the lower Hymenopteran 
families. In some of the horntails (Siricidse) the postnotum or 
postscutellum is a j)rominent plate on the surface of the dorsum be- 
hind the scutellum. In Sirex (Siricida?) this plate is sunken below 



56 



THE ANATOMY OF THE HONEY BEE. 



the general surface and mostly concealed between the mesothorax 
and the metathorax. In higher families such as the Pompilidse the 
postnotum of the mesotergum is entirely concealed by invagination, 
but it still carries a very large phragma. When, now, we come to 
the highest members of the order we find that the median part of the 
postnotum in the mesothorax is gone entirely and that it is repre- 
sented only by the lateral arms (figs. 22, PN ; 23 A, PN„) carrying 
the large, jDurely internal postphragma {Pph). 

The mesopleurum is large and consists principally of the episternum 
(fig. 21, Ejis^)^ which, however, is continuously fused with the meso- 
sternum (figs. 20 and 21, S.^). The pleural suture (fig. 21, PS.) is 
short and sinuous and does not reach more than half way from the 
wing process to the base of the middle leg. The epimerum is reduced 
to a small double plate lying above the episternum and posterior to 
the wing process (figs. 21, Ejmi.^^ and 24 A, E-pm and Ejjm). The 

pleural ridge (fig. 24 B, PR) 
is weak, but the wing process 
(irP) is well braced by a num- 
ber of accessory internal ridges. 
One preparapterum {2P) and 
one postparapterum {3P) are 
present. Lying behind the 
postparapterum is another 
larger sclerite (fig. 24 A and 
B, />7i), whose anterior end is 

Fig. 22.— Lateral view of mesotergum of articulated tO the edge of the 
worker, removed from the rest of thorax to . . 

show large internal postscuteiium (post- epiiiierum and whose posterior 
notum, PA') and its phragma (Pph) not tapering end is looscly asso- 

visible normally in the bee from exterior. . ^ . ^ 

ciated with the terminal arms 
of the postnotum (fig. 22, PN and ;j»?i). This sclerite might be 
regarded as the fourth para^^terum, but it is much more probably 
the representative of a small terminal bar of the postnotum present in 
other Hymenoptera, such as Pepsis^ which connects this tergal plate 
with the epimerum, though in this genus it is not detached from the 
main postnotal sclerite. 

Both the mesosternum (fig. 20, S.j) and the metasternum (^„) con- 
tribute to the formation of a large entosternum {Euo+.^), wdiich forms 
a protecting bridge over the combined mesothoracic and metathoracic 
ganglia (fig. 52) and affords attachment for the ventral longitudinal 
muscles of the thorax (fig. 27, Imcl). 

The metathorax consists of a very narrow series of plates (fig. 21, 
T^, PZg, and pl^) compressed between the mesothorax and the first 
abdominal tergum {IT) . Its back plate is a single, narrow, transverse 
sclerite (figs. 21 and 23 A, T',) widening on the sides, where it carries 
the wings by the two wing processes (fig. 23 A, ANP and PNP) . The 




THE THORAX AND ITS APPENDAGES. 



57 



ordinary tergal divisions seem to be entirely obliterated. The meta- 
pleurnm consists of a dorsal plate (fig. 21, Pl^) supporting the hind 
wing and of a ventral plate {j^L) carrying the hind leg. These two 
functions certainly identify these two plates as constituting together 
the metaplenrum, but there is absolutely no trace of a division into an 
episternum and an epimerum. Once more, therefore, we have to go 
back to the generalized Hymenoptera to find out what has happened. 

X 



^VNK- 




FiGi 23. — A, thoracic terga of worker separated from one another, showing protergum 
(Ti), mesotergum {T2) and its internal postscutellum (postnotum PN2.) and phragma 
(Pplii), metatergum (7*3) and propodeum or first abdominal tergum {IT) • B, ventral 
view of principal or notal plate of mesotergum. 

The answer is simple. Sirex has a typical metaplenrum consisting of 
an episternum and epimerum separated by a complete pleural suture. 
In the higher forms this suture simply disappears, and consequently 
the pleurum shows no traces of its original component plates. The 
division into a wing-bearing and a leg-bearing plate is, therefore, a 
purely secondary one. 

None of the Hymenoptera has separate trochantinal sclerites (see 
fig. 4, Tn), but, since the coxse are articulated ventrally to knobs 



58 



THE AKATOMY OF THE HONEY BEE. 



(figs. 20 and 21, z) apparently belonging to the sterna, it might be 
supposed that the trochantins have fused with the latter plates. 

The posterior part of the thoracic mass (fig. 21) consists of the 
first abdominal tergum (IT), which fits into the deeply concave pos- 
terior edges of the metathorax and forms the peduncle (Pd) that 
carries the rest of the abdomen (fig. 32). It consists of a single large, 
strongly convex sclerite (figs. 21 and 23 A, IT) bearing the first 
abdominal spiracles laterallj^ i^^p) ^i"^cl having its surface divided 
into several areas by incomplete sutures. 

Many entomologists find it difficult to believe that this plate, which 
so apparently belongs to the thorax, is really derived from the abdo- 
men. But the proof is forthcoming from a number of sources. In 
the first place, the thorax is complete without it and the abdomen is 
incomplete without it, the latter having otherwise only nine seg- 
ments. Again, if the plate is reckoned as a part of the thorax we 



WP. 




WP, 




B 

Fig. 24. — A, upper part of left mesoplenrum of worker, external ; B, inner view of same. 

should have the anomaly of a thorax with three pairs of spiracles — 
there being the normal two on each side situated, as they always are, 
between the true thoracic segments. Furthermore, comparative anat- 
omy shows us that in some of the sawflies (Tenthredinidse) the first 
abdominal tergum, while separated by a wide membranous space 
from the second, is not at all incorporated into the thorax. In a horn- 
tail such as -jSirex (Siricidse) the entire first abdominal segment is 
fused to the posterior edge of the metathorax and is onl}^ loosely 
joined to the next abdominal segment by membrane. This insect 
affords, therefore, a most complete demonstration of the transference 
of this segment from the rest of the abdomen to the thorax. Finally^ 
we have absolute proof of its abdominal origin based on a knowledge 
of development, for it has been shown by Packard from a study of the 
bumblebee that the first abdominal segment of the larva is trans- 
ferred during the pupal metamorphosis to the thorax and forms the 



THE THORAX AND ITS APPENDAGES. 59 

part under discussion. We hence see that not only the first abdomi- 
nal tergum but the entire segment has undergone transposition, 
though the ventral part has disappeared in all the higher families. 
This transferred part has been named both the median segment and 
the ])TOfodeum, by writers who recognize it as belonging to the abdo- 
men and not to the thorax. 

The names current among systematists for the back plates of 
Hymenoptera afford an excellent example of the errors that ento- 
mologists may be led into through an ignorance of the comparative 
anatomy of insects. They recognize the protergum as such and then, 
knowing that there are yet two segments to be accounted for, they 
call the mesoscutum the " mesonotum," the mesoscutellum the 
" scutellum," the metatergum the " postscutellum " (being unaware 
that the true postscutellum is deeply concealed within the thorax), 
while the first abdominal tergum is called the metathorax.. Such 
a nomenclature assigns both pairs of wings to the mesothorax. Too 
many systematists working in only one order of insects do not care 
whether their names are applied with anatomical consistency or not, 

2. THE WINGS AND THEIR ARTICULATION. 

In the study of insects the wings always form a most interesting 
subject because by them insects are endowed with that most coveted 
function — the power of flight. It has already been stated that the 
wings are not primary embryonic appendages, but are secondary out- 
growths of the body wall from the second and third thoracic seg- 
ments. Therefore it is most probable that the early progenitors of 
insects were wingless, yet for millions of years back in geological time 
they have possessed these organs in a pretty well developed condition. 

Nearly all of the insect orders have some characteristic modifica- 
tion of the wing-veins and their branches. None of them, however, 
departs nearly so far from the normal type as do the Hymenoptera, 
even the lowest members of this group possessing a highly specialized 
venation. Before beginning a study of the Hymenopteran series 
which leads up to the bee the student should first iturn back to figure 
G (p. 22) and again familiarize himself with the generalized condi- 
tion of the veins and the articular elements of the wing. By com- 
paring, now, with this diagram the basal parts of the wing of a 
sawfly {Itycorsia discolor^ fig. 26 A) it will be easy to identify the 
parts of the latter. Vein C has two little nodules (6^, C) cut off from 
its basal end which lie free in the axillary membrane. Vein Sc articu- 
lates by an enlarged and contorted base {Sc) with the first axillary 
{lAx)^ while vein R is continuous with the second {2 Ax). The next 
two veins that come to the base and unite with each other are appar- 
ently not the media and cubitus but the first and third anals {lA and 



60 



THE AiSTATOMY OF THE HONEY BEE. 



oA), since they are associated ^ith the third axiUarv {3 Ax) . In this 
species the subcosta {Sc) is entirely normal, but in the related horntail 
{Sirtw favicornis, fig. 20 B) the enlarged basal part of the subcosta is 
almost separated from the shaft of the vein, while the latter (fig. :25A, 
Sc) is short and weak. A study of the Yenation of this wing leads 
us to belicYe that the Yem which arises from the radius a short dis- 
tance fi-om its base is the cubitus {Cii). Therefore the basal part 



C Sc R-N( 



M R R, 




Fig. 25. — Wings of Hymenoptera and their basal articular sclerites (lAx-iAx) : A, Sirex 
fiaiicornis, front wing; B, Pepsis sp., front wing; C, lionej- bee, front wing; D, honey 
bee, hind wing. 

of the media is either gone or is fused with the radius. Since we dis- 
coYer its branches in the distal field of the wing, arismg from the 
trunk of the radius, we conclude that the latter is the case. By this 
sort of reasoning we may arrive at the Comstock and Xeedham inter- 
pretation of the wing illustrated at A, fig. :2o. From this it is evident 
that the branches of both the radius and the media have been bent 
back toward the posterior margin of the wing. 



THE THOKAX AND ITS APPENDAGES. 

c 



61 




Fig. 26. — Basal elements of wings of Hymenoptera : A, base of front wing of a sawfly 
(Itycorsia discolor) showing comparatively generalized arrangement of veins and 
axlllaries : B, bases of anterior veins of front wing of a homtail iSirex 'flavicorni4\ , 
showing detachment of base of snbcostal vein i.S'ci from its shaft; C, corresponding 
view of anterior veins in front wing of a tarantula-tiller iPepsis sp. i. showing com- 
plete absence of shaft of subcosta. but presence of basal part (Set fused with base of 
radius {R) ; D. axlllaries of anterior wing of honey bee worker: E. tegula of worker; 
F. base of anterior wing of worker showing absence of shaft of subcosta but presence 
of scale f.S'ct derived from its base ; G. axiliaries of hind wing of worker, the fourth ab- 
sent in bee ; H, base of hind wing of worker, showing absence of costal and snbcostal veins 
and fusion of bases of subcosta (>'ci and radius (R> into large humeral mass: I. attach- 
ment of front wing to scutum *Sct-2< and scuteilum iScU) of mesotergum : .J. under view 
of end of mesoscutellum iscUi showing attachment of both first ilAxi and fourth 
axlllaries t^Ajc) to posterior wing process (PyP). an unusual connection for first axillarv. 



62 THE ANATOMY OF THE HONEY BEE. 

Taking this wing of Sirex as a foundation let us proceed a little 
higher and examine the wing of a Pompilid, such as Pepsis (figs. 
26 C and 25 B). We observed that in Sirex (fig. 26 B) the basal 
part of vein Sc is almost separated from the distal shaft. In Pejysis 
(fig. 26 C) it is entirely a separate piece, to which is fused also the 
base of vein R. Moreover, the shaft of Be has disappeared entirely 
(fig. 25, B). Thus there is at the humeral angle of the wing a large 
chitinous mass (fig. 26 C, Sc and R) representing the fused bases 
of both the subcosta and the radius, which is associated with 
both the first axillary {lAx) and the second axillary {'BAx'). 

If now we proceed to a study of the front wing of the bee we 
find that its basal characters (fig. 26 F) are more similar to those of 
Sirex (B), while its venation (fig, 25 C) resembles more closeh^ that 
of Pepsis (B). The subcostal scale at its base (fig. 26 F, Sc) is 
not fused with the base of the radius, but the distal part of the 
subcosta is gone (fig. 25 C), as in Pepsis. In the hind wing of the 
bee (fig. 26 H) the bases of the subcosta and radius are fused into 
one large humeral mass articulating with the first two axillaries 
{lAx and '2 Ax). The third axillary {3 Ax) is well developed but 
the fourth is absent. The venation (fig. 25 D) is reduced to a very 
simple condition, but to one just the opposite from primitive. 

The details of the axillaries in the two wings are shown by figure 
26 D and G. The fourth {Jj^Ax) is well developed in the front wing 
(D) and has a large accessory sclerite {y) connected with it, upon 
which is inserted a long slender muscle (fig. 28, 6c). A very small 
accessory sclerite {ax) occurs close to the muscle plate of the third 
axillary {3 Ax). These are called "accessory" sclerites because 
they are of irregular occurrence in the wing bases of insects generally 
and are developed in connection with the muscle attachments. Simi- 
lar ones occur in the hind wing (G, ax) in connection with the 
second {2 Ax) and third axillaries {3 Ax). 

The front wing is attached to the posterior half of the side of 
the mesonotum. The anterior notal wing process is bilobed (figs. 
22, 23 A, T'o, ANP) and is carried by the scutum, while the pos- 
terior process {PNP) is carried by the scutellum and is mostly 
hidden beneath the anterior wing process.. The two wing processes, 
in fact, are so close together that the first axillary articulates not 
only with the first but also with the second (fig. 26 J). The axillary 
cord (fig. 26 F, AxG) arises from a lobe of the scutellum overlapped 
by the lateral margin (I and J, AxG). In the hind wing, where the 
fourth axillary is absent, the third articulates directly with the 
posterior notal wing process of the metatergum (fig. 23 A, 7'., PNP). 

The base of the front wing is overlapped by a large scale (fig. 26, 
E and I, Tg) called the tegula. It is carried b^^ the axillary mem- 



THE THORAX AND ITS APPENDAGES. 63 

brane, to which it is attached between the humeral angle of the wing 
base and the edge of the notinn. The tegulse are present in most in- 
sects, generally on the base of each wing, but they usually have the 
form of small inconspicuous hairy pads, as shown in the diagram 
(fig. 0, Tg). In the flies, moths, butterflies, and Hymenoptera, 
however, the tegulaj of the front wings develop into large conspicu- 
ous scales overlapping the humeral angles of the bases of these 
wings. 

The motion of the wing in flight consists of both an up-and-down 
movement and a forward-and-backward movement, which two com- 
bined cause the tip of the wing to describe a figure-eight course if 
the insect is held stationary. Corresponding with these four move- 
ments are four sets of muscles. In the clragonflies nearly all of the 
wing muscles are inserted directly upon the base of the wing itself, 
but in other insects, excepting possibly the mayflies, the principal 
muscles are inserted upon the thoracic walls and move the Aving 
secondarily. In the lower insects, such as the grasshoppers, crickets, 
stoneflies, net-Avinged flies, etc., the two wing-bearing segments are 
about equal in their development and each is provided with a full 
equipment of muscles. In these insects the wings work together by 
coordination of their muscles, although each pair constitutes a sepa- 
rate mechanism. In such insects, however, as the true flies and the 
wasps and bees the metathorax, as we have seen in the case of the 
bee, is greatly reduced, and what is left of it is solidly attached to 
the mesothorax. In the flies the hind wings are reduced to a pair 
of knobbed stalks having no function as organs of flight, while in 
the bees the hind wings, Avhich are very small, are attached to the 
front wings by a series of booklets on their anterior margins (fig. 
25 D, Rh) which grasp a posterior marginal thickening of the 
front wings. Moreover, when we examine the interior of the bee's 
thorax we find that the muscles of the metathorax are greatly 
reduced or partly obliterated and that the great mesothoracic mus- 
cles serve for the movement of both Avings, thus assuring a perfect 
synchrony in their action. Hence, it is clear that the union and 
consolidation of the thoracic segments in the higher insects is for 
the purpose of unifying the action of the wings. 

The muscles of flight in the bee may be very easily studied by cutting 
the thorax of a drone into lateral halves. The cavity of the thorax 
is occupied almost entirely by three great masses of muscles. One 
of these is longitudinal, median, and dorsal (fig. 27, LMcl^)^ extend- 
ing from the mesoscutum {Set.,) and the small prephragma {Aph) 
to the large mesothoracic postphragma (Ppli.,). A small set of 
muscles {L3IcL) then connects the posterior surface of this phragma 
with the lower edge of the propodeum {IT). On each side of the 



64 



THE ANATOMY OF THE HONEY BEE, 



LMcl, 



anterior end of this great longitudinal muscle is a thick mass of 
dorso- ventral fibers (VMcl) extending from the lateral areas of the 
mesoscutum {/Set.,) to the lateral parts of the mesosternum {/S.J. A 
contraction of the vertical muscles must depress the tergal parts, 
at the same time expanding the entire thorax in a longitudinal direc- 
tion and stretching the longitudinal muscles. A contraction, then, 
of the latter muscles {LMcl) restores the shape of the thorax and 
elevates the tergal parts. Eemembering, now, that the wings are 

supported from be- 
low upon the 
pleural wing proc- 
esses and that each 
is hinged to the 
back by the notal 
wing processes, it 
is clear that a de- 
pression of the 
dorsum of the 
thorax must ele- 
vate the wings and 
that an elevation 
of the dorsum de- 
presses them — the 
pleural wing proc- 
esses acting as the 
fulcra. Hence, the 
chief up-and-down 
movements of the 
wings are pro- 
duced by these 
great thoracic mus- 
cles acting upon 
the shape of the 
thorax as a whole 
and not directly 
upon the Avings 
themselves. The vertical muscles are the elevators and the longi- 
tudinal the depresso7's. 

But besides being moved up and down the wings can also, as before 
stated, be extended and flexed, i, e., turned forward and backward in 
a horizontal plane upon the pleural wing process. The muscles 
which accomplish these movements lie against the inner face of the 
pleurum (fig. 28), and each wing is provided with a separate set. 
The extensor muscle {PMcl) is the most anterior and is inserted by 
a long neck upon the preparapterum {^P)- The latter is closely 




Fig. 27. — Median section through thorax of drone, showing 
longitudinal muscles (LMch) of mesothorax going from 
mesotergal scutum (Scto) and small anterior phragma 
(Aph) to posterior phragma (Pphn) of internal postscutel- 
lum (postnotum) of same segment, also showing vertical 
mesothoracic muscles {VMcl), and ventral longitudinal mus- 
cles (Imcl), and longitudinal muscles of metathorax 
(LMch) going from postphragma of mesothorax (Pplu) to 
posterior edge of propodeum or first abdominal tergum {IT). 
By alternate contraction of dorsal longitudinal muscles and 
vertical muscles, roof of thorax is elevated and depressed, 
causing wings to heat downward and upward respectively, 
being supported on fulcra formed by pleural wing processes 
(fig. 28, WP2) of side walls of thorax. 



i 



THE THORAX AND ITS x\PPE]SIDAGES. 



65 



WP, 2Ax 3AX 



connected with the anterior part of the base of the wing so that a 
contraction of the muscle turns the wing forward and at the same 
time dejDresses its anterior margin. For this reason the parapterum 
and the extensor muscle have been called the fronator ayijaratus^ and 
the muscle is known also as the pronator muscle. In some insects 
Avhich fold the wings back against the body this muscle is a great 
deal larger than in the bee. The flexor imiscle {RMcV) consists of 
three parts situated upon the anterior half of the pleurum and in- 
serted upon the third axillary (SAx) by long tendonlike necks. 
These muscles are antagonistic 
to the extensor and bj^ their 
contraction pull the wing- 
back toward the body. 

The mechanism which pro- 
duces the wing motion thus 
seems to be a very simple one 
and may be summarized as 
follows: Each wing rests and 
turns upon the wing process 
of the pleurum (figs. 24 and 
28, WP) by means of the 
pivotal sclerite or second axil- 
lary in its base (figs. 26 F and 
2S,2Ax). It is hinged to the 
back by the first and fourth 
axillaries (fig. 26 F, lAx and 
4Ax) which articulate with 
the anterior and posterior 
notal wing processes (fig. 23 
A, n, ANP and PNP), re- 
spectively. The large vertical 
muscles (fig. 27, VMcl) of 
the thorax depress the ter- 
gum, which pulls down with 
it the base of the wing and 
hence elevates the distal part — 
the fulcrum being the pleural wing process. The dorsal longitudinal 
muscle {LMcl) restores the shape of the thorax, elevates the tergum, 
and consequently depresses the wing. Extension and flexion of the 
wing are produced by special muscles (fig. 28, PMcl and RMcl) acting 
upon its base before and behind the pleural wing process, respectively. 

Besides these muscles there are several others (fig. 28) associated 

with the wing whose functions are less evident. Most conspicuous 

of these is a muscle occupying the posterior half of the mesopleurum 

{aa) and inserted upon the outer end of the scutellum. This may 

22181— No. 18—10 5 




Fig. 28. — Internal view of right ijleurum of 
mesothorax of drone, sliowing muscles in- 
serted upon parapteral plates (tP and SI') 
and upon third axillary (SAx). The wing 
rests upon wing process of pleurum (WP2) 
by second axillary (2Ax) ; it is turned for- 
ward and downward by the pronator muscle 
{PMcl), inserted upon anterior parapterum 
(2P) which is attached to costal head of 
wing, and is turned back toward body by 
flexor muscle (RMcl) inserted upon third 
axillary (3Ax). 



66 THE ANATOMY OF THE HONEY BEE. 

be simply accessory to the large vertical sterno-scutal muscle (fig. 27, 
VMd). Another is a long slender muscle {hh) attached to the upper 
end of the mesocoxa and inserted upon the postparapterum {3P). 
This is sometimes termed the coxo-axUlary muscle. A third {cc) is 
inserted upon the tip of the accessory sclerite {y) of the fourth 
axillary and is attached to the lateral arm of the large entosternum 
of the mesothorax and metathorax. 

3. THE LEGS. 

The legs of the honey bee are highly modified for several special 
purposes besides that of walking, but they are so well known and 
have been so often described that it will not be necessary to devote 
much space to them here. 

The front legs (fig. 29 A) have a structure formed by the adjoining 
ends of the tibia and the first tarsal joint, which is called, on account 
of its use, the antenna cleaner. It consists (fig. 29 C) of a semi- 
circular notch [del) in the base of the first tarsal joint {ITar) pro- 
vided with a comblike row of bristles. A specially modified, flat, 
movable spur (ee), shown in ventral view at B, is so situated on the 
end of the tibia tliat it closes over the notch when the tarsus is bent 
toward the tibia. By grasping an antenna between the notch and 
the spur and drawing it through the inclosure the bee is able to re- 
move from this sensitive appendage any pollen or particles of dirt 
that may be adhering to it. 

The middle legs (fig. 29 D) present no special modifications of any 
importance. It will be observed, however, that they, as well as the 
other legs (A and F), have the first joint of the tarsus {ITar) very 
greatly enlarged. 

The hind legs of all three forms, the worker (F) , the queen (E) , and 
the drone (H), have both the tibia and the large basal segment of 
the tarsus very much flattened. In the queen and drone there seems 
to be no special use made of these parts, but in the worker each of 
these tAvo segments is modified into a very important organ. The 
outer surface of the tibia (F, Tlj) is fringed on each edge by a row of 
long curved hairs. These constitute a sort of basket {Ch) in which 
the pollen collected from flowers is carried to the hive. The struc- 
tures are known as the pollen haskefs^ or coj'hhnla. The inner sur- 
face of the large, flat, basal segment of the tarsus {ITar) is pro- 
vided Avith several roAvs of short stiff spines (G) forming a brush by 
means of Avhich the bee gathers the pollen from its body, since it 
often becomes covered with this dust from the floAvers it \4sits for 
the purpose of getting nectar. When a sufficient amount is accumu- 
lated on the brushes it is scraped off from each over the edge of the 
tibia of the opposite hind leg and is thus stored in the pollen baskets. 
Hence the Avorker often flies back to the hive Avith a o'reat mass of 



THE THOJRAX AND ITS APPENDAGES. 



67 




Fig. 29. — A, left front leg of worker, anterior view, showing position of notch (ihl) of 
antenna cleaner on base of first tarsal joint (iTar) and of closing spine {cc) on end 
of tibia {Th) ; B, spine of antenna cleaner (ec) in fiat view; C, details of antenna 
cleaner ; D, left middle leg of worker, anterior view ; E, left hind leg of queen, anterior 
or outer view ; F, left hind leg of worker, anterior or outer view, showing the pollen 
basket (Cb) on outer surface of tibia (Tb) ; G, inner view of first tarsal joint of hind 
leg of worker showing rows of pollen-gathering hairs and the so-called " wax shears " 
iff) ; H, left hind leg of drone, anterior or outer view. 



68 



THE ANATOMY OF THE HONEY BEE, 



pollen adhering to each of its hind legs. The pollen baskets are 
also made use of for carrying propolis. 

Between the ends of the hind tibia {Th) and the first tarsal joint 
{ITar) is a sort of pincerlike cleft (F and G, ff) guarded by a row 
of short spines on the tibial edge. This is popularly known as the 
" wax shears " and it is supposed to be used for picking the plates 
of wax out of the wax pockets of the abdominal segments. The 
writer, however, has watched bees take the wax from their abdomen 
and in these observations they always poked the wax plates loose 



cu 




Fig. 30. — A, dorsal view of end of last tarsal joint of first foot (Tar), the claws (Cla), 
and empodium (Emp) of worker; B, ventral view of same; C, lateral' view of same, 
showing empodium in ordinary position when not in use. 

with the ordinary hairs or spines of the tibiae or tarsi and then by 
means of the feet passed them forward beneath the body to the 
mandibles. 

The last tarsal joint of each leg bears a pair of claivs (E, Cla^ and 
a single median empodium {Emp). Each one of the claws is bi- 
lobed, consisting of a long tapering outer point and a smaller inner 
one (figs. 30 and 31). The claws of the worker (fig. 31 A) and the 
queen (B) are only slightly diflerent in details of outline, although 
the claws of the queen are much greater in size than those of the 



THE ABDOMEN, WAX GLANDS, AND STING. 



69 



worker, but the drone's claws (C) are large and very strikingly 
different in shape from those of either the worker or the queen. 

The empodium (fig. 30 A, B, and C, Emp) consists of a terminal 
lobe bent upward between the claws (C) and deeply cleft on its 
dorsal surface (A), and of a thick basal stalk 
whose walls contain a number of chitinous 
plates. One of these plates is dorsal (A and 
C, hh) and bears five very long, thick, curved 
hairs projecting posteriori}^ over the terminal 
lobe, while a ventral plate (B and C, ii) is 
provided with numerous short thick spines. 
A third plate (A, B, and C, gg) almost 
encircles the front of the terminal lobe, its 
upper ends reaching to the lips of the cleft. 

\^'lien the bee walks on any ordinary' sur- 
face it uses only its claws for maintaining a 
foothold, but when it finds itself on a smooth, 
slippery surface like glass the claws are of no 
avail and the empodia are provided for such 
emergencies as this. The terminal lobe is 
pressed down against the smooth surface and 
its lateral halves are flattened out and adhere 
by a sticky liquid excreted upon them by 
glands said to be situated in front of them, 
the muscle that flattens the empodial lobes the latter spring back 
into their original position by the elasticity of the chitinous band 
{gg) in their walls. 




C ' 

Fig. 31. — A, outer yiew of 
hind claw of worker ; B, 
same of queen ; C, same 
of drone. 

On the relaxation of 



V. THE ABDOMEN, WAX GLANDS, AND STING. 

The abdomen of the worker and queen appears to consist of six seg- 
ments (figs. 1, 32, 33, //-F//), but it must be remembered that, as 
has already been explained, the thoracic division of the body in the 
Hymenoptera includes one segment, the propodeum or median seg- 
ment, which really belongs to the abdomen and is its true first seg- 
ment according to the arrangement in all other insects. Hence, 
counting the propodeum (figs. 21 and 32, IT) as the first, we find 
seven exposed abdominal segments in the worker and queen and 
nine in the drone. Each one except the first consists of a tergum 
{T) and a sternum {S)^ the former reaching far down on the side 
of the segment, where it carries the spiracle {Sp) and overlaps the 
edge of the sternum. The two plates of the last or seventh segment 
in the worker and queen are separated by a cleft on each side, and 
if they are spread apart it is seen that the tip of the abdomen 



70 



THE ANATOMY OF THE HONEY BEE. 



incloses a cavity which lodges the sting and its accessory parts. The 
end of the abdomen of the male (fig. 56 D) is quite different from 
that of the female, while in it parts at least of nine segments are 

in 
n 




Sp Stn 

Fig. 32. — Lateral view of abdomen of worker, showing the propodeum {IT) as a part 
of the abdomen, of which it is the true first segment. 

visible, the last is very much modified and is exposed only on the 
sides and below. 

An internal view of the ventral plates and the lateral parts of the 





Fig. .";;. — Ventral view of abdomen of 
worker, showing tip of sting (Stn) and 
palpiislike appendages (StnPlp) pro- 
jecting from sting chamber within 
seventh segment (T'7/). 



jCIsp-' 

Fig. 34. — Dorsal view of ab- 
dominal sterna of drone, 
showing clasping appendages 
(IClsp and 2Clsp) of ninth 
segment. 



terga in the worker is shown by figure 20, while a corresponding 
view of the male sterna is shown b}^ figure M. It will be seen that 
each sternum is very widely underlapped (viewed from above) by the 



i 



THE ABDOMEN, WAX GLANDS, AND STING. 71 

one next in front of it and that the intersegmental membrane (71/6) 
is reflected from the middle of the dorsal surface of each to the 
anterior edge of the following sternum. By removing an individual 
plate (fig. 35 A) this is more easily shown. It is also clearly seen 
that the transverse line of attachment of the membrane {Mh) divides 
the sternum into a posterior part {Rd)^ which is merely a prolonged 
reduplication underlapping the following sternum, and into an an- 
terior part underlapped by the preceding sternum. The posterior 
half is, hence, purely external while the anterior half forms the true 
ventral wall of the segment, its dorsal face being internal and its 
ventral face external. The anterior part is also very smooth and 
shiny and somewhat bilobed and for this reason it is sometimes called 
the " mirrors." Its edge is bounded by a thickened ridge giving off a 
short apodeme (^1/^) on each side. The mirrors of the last four 
sterna are also, and more appropriately, called the wax plates because 
the wax is formed by a layer of cells lying over them. It accumu- 
lates on the ventral side in the pocket between the wax plates and the 
posterior underlapping prolongation of the preceding sternum. Wax 
is formed only on the last four visible segments, i. e., on segments 
IV-VII, inclusive. 

In studying any part of the body wall of an insect it must always 
be borne in mind that the chitin is originally simply an external cutic- 
ular layer of a true cellular skin or epidermis (erroneously called 
"hypodermis" in insects), but that in the adult stage the latter 
almost everywhere disappears as a distinct epithelium. Thus the 
chitin comes to be itself practically the entire bod}^ wall, the cell layer 
being reduced to a very inconspicuous membrane. However, in cer- 
tain places the epithelium may be developed for special purposes. 
This is the case with that over the Avax plate:-; which forms a thick 
layer of cells that secrete the wax and constitute the so-called 'wax 
glands. The wax is first secreted in a liquid condition and is ex- 
truded through minute pores in the wax plates of the sterna, harden- 
ing on their under surfaces into the little plates of solid wax with 
which every bee keeper is acquainted. 

The secretion of the wax has l)een studied by Dreyling (1903), who 
made histological sections through the glands at different times in 
the life of the bee. He found that in young, freshly emerged workers 
the epidermis of the wax plates consists of a simple layer of ordinary 
epithelial cells. As the activities of the bee increase, however, these 
cells elongate while clear spaces appear between them and, when the 
highest development is reached, the epithelium consists of a thick 
hiyev of very long cells with liquid wax stored in the spaces between 
them. In old age most of the cells become small again and in those 
bees that live over the winter the epithelium degenerates to a simple 
sheet of nucleated plasma showing no cell boundaries. It is thus 
evident that the secretion of wax is best performed during the prime 



72 



THE ANATOMY OF THE HONEY BEE. 



of life, which in bees is at about IT days of age or before, and that 
old bees can only gather honey and pollen. Bees do not normally 
secrete wax while performing the other more ordinary duties of their 
life. When comb is needed a large number of young bees or bees 
that have not passed their prime hang together in vertical sheets 
or festoons within the hive and are fed an abundance of honey. After 
about twenty-four hours they begin to construct comb. During this 
time the wax is excreted through the wax plates and accumulates in 

the external wax pockets below. 
It is poked out of these pockets by 
means of the spines on the feet 
and is passed forward beneath the 
body to the mandibles. By means 
of these organs it is manipulated 
into little pellets and modeled 
into the comb. Dreyling describes 
the pores of the wax plates as ex- 
cessively fine, vertical, parallel 
canals only visible in very thin 
sections and under the highest 
power of the microscope. 

Corresponding abdominal sterna 
present quite different shapes in 
the three forms of the bee (fig. 35 
A, B, and C). In the queen (B) 
the sterna are much longer than in 
the worker (A), while in the 
drone (C) they are shorter and 
have very long lateral apoclemes 
{Ap). 

The last three abdominal seg- 
ments — the eighth, nintli, and 
tenth — are \Qvy different in the 
two sexes on account of their 
modification in each to accom- 
modate the external organs of re- 
production and egg laying. In the female these segments are entirely 
concealed within the seventh, but, in the male, parts of both the 
eighth and ninth segments are visible externally. 

The seventh segment of the drone (counting the propodeum as 
the first) is the last normal segment, i. e., the last one having a com- 
plete terg-um and sternum resembling those of the anterior part of 
the abdomen (fig. 56 D, VIIT and VI IS). Behind the seventh ter- 
gum and partly concealed within it is the eighth tergum (VIIIT) 
carrying the last abdominal spiracles (Sp). The eighth sternum is 




Fig. 35. — Dorsal surface of sixth abdominal 
sternum : A, worker ; B, queen ; C. drone ; 
showing division of plate by line of at- 
tachment of intersegmental membrane 
(Mh) into anterior part with polished 
internal surface, in worker bearing wax 
glands, and into large posterior external 
part (R(l) underlapping anterior half of 
succeeding sternum. 



i 



THE ABDOMEN, WAX GLANDS, AND STING. 73 

almost entirely concealed within the seventh. It is very narrow 
below, but is expanded at the upper parts of its sides {VI I IS) , where 
it is partly visible below the eighth tergum and behind the seventh 
sternum. The dorsal part of the ninth segment is membranous except 
for a small apodeme-bearing plate on each side hidden within the 
eighth tergum. The ninth sternum, on the other hand, is a well- 
developed semicircular band {IXS) forming the ventral and ventro- 
lateral parts of the ninth segment. It bears on each side two con- 
spicuous lobes — one a small, darkly chitinized, dorsal plate {IGlsp) 
carrying a large bunch of long hairs, the other a large, thin, ventral 
plate {2Clsp). Between these four appendicular lobes is ordinarily a 
deep cavity, which is the invaginated penis (fig. 56 E), but in 
figure D this organ is shown partly evaginatecl {Pen). A^T^iile the 
penis is really an external organ, the details of its structure will be 
described later in connection with the internal organs of reproduction. 
The tenth segment is entirely lacking in segmental form. The anal 
opening is situated in a transverse membrane beneath the eighth ter- 
gum {VIIIT)^ and below it is a thin chitinous plate, which may 
belong to the tenth segment. 

In many insects the modification of the terminal segments of the 
males in connection with the function of copulation is much greater 
than in the bee. The ninth segment often forms a conspicuous 
enlargement called the JiyjjojjyghMn., which is usually provided with 
variouslj^ developed clasping organs in the form of appendicular 
plates and hooks. 

The development of the external genital parts of the drone has been 
described by both Michaelis (1900) and Zander (1900). A small 
depression first appears on the under surface of the ninth segment of 
the larva shortly after hatching. Soon two little processes grow 
backward from the anterior wall of this pouch and divide each into 
two. The part of the larval sternum in front of the pouch become? 
the ninth sternum of the adult, while the two processes on each side 
form the upper and lower appendicular lobes (the valva externa and 
the valva interna of Zander). The penis at first consists of two little 
processes which arise between the valvse internee, but is eventually 
formed mostly from a deep invagination that grows forward between 
them. These four processes arising on the ventral side of the ninth 
segment of the male larva are certainly very suggestive of the similar 
ones that are formed in the same way on the same segment of the 
female and which develop into the second and third gonapophyses 
of the sting. If they are the same morphologically we must homol- 
ogize the two clasping lobes of the ninth sternum in the male with 
the two gonapophyses of this segment in the female. Zander (1900) 
argues against such a conclusion on the ground that the genital pouch 
is situated near the anterior edge of the segment in the female and 



74 THE ANATOMY OF THE HONEY BEE. 

posteriorly in the male, while the parts in the two sexes develop 
later in an absolutely different manner. These arguments, how- 
ever, do not seem verj^ forcible — in the earliest stages the processes 
certainly look alike in the two sexes. 

The sting of the bee is situated in the sting cavity at the end of the 
abdomen, from which it can be quickly protruded when occasion de- 
mands. This sting chamber contains also the reduced and modified 
sclerites of the eighth, ninth, and tenth abdominal segments. In 
fact, the sting chamber is formed by an infolding of these three seg- 
ments into the seventh. It is consequently not a part of the true in- 
terior of the body or body cavity which contains the viscera, but is 
simply a sunken and inclosed part of the exterior, in the same sense 
that the oven of a stove is not a part of the real inside of the stove. 
Consequently the parts of the sting, though normallj^ hidden from 
view, are really external structures. 

A very gentle pull on the tip of the sting is sufficient to remove it 
from its chamber, but a sting thus extracted brings along with it the 
ninth and tenth segments, most of the eighth segment, the poison 
glands, and the terminal part of the alimentary canal. This is clue 
to the fact that the inclosed segments are attached to the surround- 
ing parts by very delicate membranes. For the same reason they so 
easily tear from the living bee as the latter hurriedly leaves its victim 
after stinging. The worker thus inflicts a temporary wound and 
pain at the cost of its own life. Undoubtedly, however, nature re- 
gards the damage to the enemy as of more importance to the bee 
community as a whole than the loss of one or a dozen of its members. 
The entire stinging apparatus with a bag of poison attached is thus 
left sticking in the wound while the muscles, which keep on working 
automatical^, continue to drive the sting in deeper and deeper and 
at the same time pump in more poison. Such a provision certainly 
produces much more effective results than would a bee giving a thrust 
here and another there with its sting and then rapidly flying away 
to escape from danger. 

The sting itself, when extracted from its chamber, is seen to con- 
sist of a straight tapering shaft with its tip directed posteriorly and 
its base swollen into a bulblike enlargement. In superficial appear- 
ance the shaft appears to be solid, although we shall presently show 
that it is not, but the bulb is clearly hollow and is open below by 
a distinct median cleft. Several plates of definite shape and arrange- 
ment always remain attached to the sting and overlap its base. The 
entire apparatus, including the base of the large poison sac, is shown 
somewhat diagrammatically in side view by figiire 36. The bulb of 
the sting (ShB) is connected with the lateral plates by two arms 
which curve outward and upward from its base. (Only the left side 
is shown in the figure.) Between these arms the two poison glands 



THE ABDOMEN, WAX GLANDS, AND STING. 



75 



{PsnSc and BGl) open into the anterior end of the bulb. From the 
posterior ends of the plates two whitish fingerlike processes (StnPlp) 
project backward. When the sting is retracted these lie at the sides 
of the shaft (figs. 33 and 37), but in figure 36 the sting is shown in a 
partly protracted position. These appendages, often called the sting 
palpi, undoubtedly contain sense organs of some sort by means of 
which the bee can tell when her abdomen is in contact with the object 
upon 'which she desires to use her sting. 

A close examination of the sting shows that it is a much more com- 
plicated structure than it at first sight appears to be. The shaft, for 
example, is not a simple, solid, tapering, spearlike rod, but is a hollow 
organ made of three pieces which surround a central canal. One of 
these pieces is dorsal (fig. 30, ShS) and is the true prolongation of 
the bulb (ShB), while the other two (Let) are ventral and slide 
lengthwise on tracklike ridges of the dorsal piece. Moreover, each 
basal arm of the 
sting is double, con- 
sisting of a dorsal 
or posterior piece 
(ShA), which is like- 
wise a prolongation 
of the bulb, and a 
ventral or anterior 
piece (Zc^), which is 
continuous with the 
ventral rod of the 
shaft on the same 
side. Hence the sting 
may be analyzed into 
three elements, which 

are characterized as follows: The dorsal piece, known as the sheath, 
consists of a prominent basal swelling or hidb (ShB) containing a 
large cavity, of a terminal tapering shaft (ShS), and of two curved 
7jasal arins (ShA). The ventral part consists of two long slender 
rods, called the lancets or darts (Let), which slide freely upon two 
tracks on the ventral edges of the sheath and diverge upon continua- 
tions of these tracks along the basal arms of the latter (ShA). The 
bulb is hollow, containing a large cavitj^ formed by invagination 
from below, where it is open to the exterior by a lengthwise cleft. 
This cavity continues also through the entire length of the shaft of 
the sting as a channel inclosed between the dorsal sheath and the 
latero- ventral lancets. This channel, as will be explained later, is 
the poison canal of the sting. 

Each arm of the sheath (ShA) is supported at its end farthest 
from the bulb by an ohlong plate (fig. 36, Oh), which normally over- 




FiG. .36. — Semidiagrammatic view of left side of sting of 
worker, accessory plates (Tri, Oh, Qd), sting palpus 
((?/)?P?/)), -alkaline poison gland (BQl), and base of large 
poison sac (I'snSc) of acid gland. 



70 



THE ANATOMY OF THE HONEY BEE. 



laps the side of the bulb, and which carries distally the palpi of the 
sting (StnPlp). Each lancet is attached at its base to a triangular 
plate {Tri) which lies latero-dorsad to the base of the oblong plate 
and articulates with a knob on the dorsal edge of the latter by its 
ventral posterior angle. By its dorsal posterior angle the triangular 
plate is articulated to a much larger quadrate plate {Qd) which 
overlaps the distal half of the oblong plate. A thick membranous 
lobe {IXS), concave below, where it is thickly set with long hairs, 
overlaps the bulb of the sting and is attached on each side to the 
edges of the oblong plates. All of these parts are shown flattened out 

in ventral view by 
figure 37. 

The presence of 
the two basal arms 
of the sheath might 
suggest that this 
part is to be re- 
garded as made up 
of fused lateral 
halves. In this case 
we should have six 
appendicular ele- 
ments, viz, the two 
lancets, the two 
halves of the sheath, 
and the tw^o pal- 
puslike organs. If 
now we turn back to 
figure 8, showing 
the component parts 
of the ovipositor of 
a longhorned grass- 
hopper, we can not 
fail to be struck at 
once by the great similarity between this organ and the sting of 
the bee (fig. 36). The first gonapophyses {IG) of the ovipositor are 
identical with the lancets {Let) of the sting, and their sliding connec- 
tion, by means of longitudinal tracks, with the second gonapophyses 
{2G) suggests at once that the latter represent the sheath of the 
sting {ShS). The identity is still more strongly suggested when we 
observe the small bulb (ShB) formed by the fused bases of these 
gonapophyses. The third gonapophyses (3G), which inclose between 
them the other parts of the ovipositor, represent the palpi of the 
sting {StnPlp). If , finally, Ave study the development of the parts of 
the sting we are convinced that this similarity between the sting and 
an ovipositor means something more than an accidental resemblance 




Fig. 37. — Ventrtil view of stinc; of worker and accessory parts 
flattened out. 



THE ABDOMEN, WAX GLANDS, AND STING. 77 

between two different organs — in fact we can not doubt that the sting 
is simj)ly an ovipositor which, being no longer needed for egg-laying 
purposes, has been modified into a poison-injecting apparatus. Zan- 
der (1899, 1900) and others have shown that the sting of the' bee 
arises from six little abdominal processes of the larva, two of Avhich 
arise on the eighth segment and four on the ninth. Those of the first 
pair develop into the lancets, those of the middle pair on the ninth 
segment fuse to form the sheath, while those of the outer pair be- 
come the paljDi. The ovipositor, it will be remembered, develops in 
the lower insects from two pairs of processes arising on the eighth 
and ninth abdominal sterna, the second pair of which very soon 
splits into four processes. The simultaneous aj^pearance of six on 
the bee larva is simply an example of the hurrying process or accelera- 
tion that the embryos and young of most higher forms exhibit in 
their development. 

It is only the higher members of the Hymenoptera, such as the 
wasps and the bees and their close relatives, that possess a true sting. 
The females of the lower members have ovipositors which closely re- 
semble those of such insects as the katydids, crickets, and cicadas, but 
which, at the same time, are unquestionably the same as the sting of 
the stinging Hymenoptera. It is said that the queen bee makes use 
of her sting in placing her eggs in the cells, but both the wasps and 
the bees deposit their eggs in cells or cavities that are large enough to 
admit the entire abdomen, and so they have but little use for an egg- 
placing instrument. But the females of the katydids and related 
forms like G onoce'phal'us (fig. 8) use their ovipositors for making a 
slit in the bark of a twig and for pushing their eggs into this cavity. 
The cicada and the sawfly do the same thing, while the parasitic 
Hymenoptera often have extremely long and slender piercing oviposi- 
tors for inserting their eggs into the living bodies of other insects. 

An examination of the sting in place within the sting chamber, as 
shown by figure 41, will suggest what the accessory plates represent in 
other less modified insects. It has already been explained that the last 
external segment of the female abdomen (fig. 32, F//) is the seventh. 
Within the dorsal part of the sting chamber is a slight suggestion of 
the eighth tergum (fig. 41, VI I IT) , which laterally is chitinizecl as a 
conspicuous plate bearing the last or eighth abdominal spiracle {Sp). 
The triangular plate (Tri), as Zander has shown by a study of its 
development, is a remnant of the eighth sternum, and the fact that it 
carries the lancet (Let) shows that even in the adult this appendage 
belongs to the eighth segment. The quadrate plate {Qd), since it is 
overlapped by the spiracle plates of the eighth tergum, might appear 
to belong to the eighth sternum, but Zander has shown that, by its 
development, it is a part of the ninth tergum. In many other adult 
Hymenoptera, moreover, the quadrate plates are undoubtedly tergal. 



78 THE ANATOMY OF THE HONEY BEE. 

for they are sometimes connected by a bridge behind the eighth 
tergmn. The oblong plate {Oh) and its stalk represent the ninth 
sternum, and since it carries both the arm of the sheath {ShA) and 
the palpus (StnPl) it still maintains its original relationships to the 
gonapophyses. The membranous lobe arising from between the 
oblong plates and overlapping the bulb of the sting (figs. 36 and 37, 
IXS) must belong to the median part of the ninth sternum. 

The tenth segment (fig. 41, X) consists of a short, thick tube having 
the anus {A7i) at its tip. It takes no part in the formation of the 
sting, but is entirely inclosed in the dorsal part of the sting chamber 
beneath the seventh tergum. 

In the accessory plates of the bee's sting we have a most excellent 
illustration of how the parts of a segment may become modified to 
meet the requirements of a special function, and also an example 
of how nature is ever reluctant to create any new organ, preferring 
rather to make over some already existing structure into something 
that will serve a new purpose. 

There are four glands connected with the sting, two of which 
are known to secrete the poison, which is forced through the canal 
between the sheath and the lancets and ejected into the wound made 
by the latter. It is this poison that causes the pain and inflammation 
in the wound from a bee's sting, which would never result from a 
mere puncture. The other two glands have been described as " lubri- 
cating glands," being supposed to secrete a liquid which keeps the 
parts of the sting mechanism free from friction. They lie within 
the body cavity, one on each side against the upper edge of the 
quadrate plate, where they are easily seen in an extracted sting, each 
being a small oblong or ovate whitish cellular mass. Transverse 
microtome sections through this region show that each of these 
glands opens into a pouch of the membrane between the quadrate 
plate and the spiracle-bearing plate of the eighth tergum. Each 
gland cell communicates with this pouch by a delicate individual 
duct. The secretion of the glands is thus poured upon the outer sur- 
faces of the quadrate plates and might easily run down upon the 
bases of the lancets and the arms of the sheath, but, for all that, the 
notion that it is lubricative in function is probably entirely conjectural. 

The large, conspicuous poison sac (figs. 3G, 37, 41, and 57, PsnSc) 
that opens by a narrow neck into the anterior end of the bulb of the 
sting is well known to everyone at all acquainted with bees. The 
poison which it contains comes from the delicate branched thread 
attached to its anterior end (fig. 57), a minute tube which, if traced 
forward a short distance from the sac, will be seen to divide into two 
branches, which are long and much coiled and convoluted, each ter- 
minating finally in a small oval enlargement (AGI). These terminal 
swellings are generally regarded as the true glands and the tubes 



THE ABDOMEN. WAX GLANDS, AND STING. 



79 




Fi( 38 — Section of 
&mall piece of wall of 
poison sac of sting. 



-Lum 



(AGID) as their ducts, but the epithelium of the tubes appears to be 

of a secretory nature also, and, if it is not, it is hard to see any reason 

for their great length. It also does not look 

probable that the two little end bodies could 

form all the poison that fills the comparatively 

enormous sac. 
The walls of the poison sac (fig. 38) are lined 

by a thick coat of laminated chitin {Int) thrown 

into numerous high folds. In the neck part of 

the sac the folds are arranged very regularly in 

a transverse direction and form interrupted 

chitinous rings, holding the neck rigidly open. 

The epithelium {Epth) contains nuclei (Nu), 

but the cell boundaries are very slightly marked. 

There is a distinct basement membrane (BJI), 

forming a tunica propria externalh^, but there 

are no muscle fibers of any sort present except 

a tew which are inserted upon the sac from some of the surrounding 

organs and which apparently act as suspensoria. 

The poison found in the sac has an 
acid reaction and is supposed to consist 
principally of formic acid. Hence its 
gland is known as the acid gland (AGl) 
of the sting. 

The other sting gland is a short, very 
inconspicuous, and slightly convoluted 
whitish tube (figs. 36, 37, 41, and 57, 
BGl) opening directly into the base of 
the bulb ventracl to the opening of the 
poison sac. Its walls consist of a thick 
epithelium of distinct cells (fig. 39, 
Epth) lined with a thin chitinous in- 
tima {lilt) and surrounded by a distinct 
basement membrane (i?J/), but, as in 
the other gland, there are no muscles 
present. The secretion of this gland is 
said to be alkaline and the gland is 
therefore known as the alkaline gland 
(BGl) of the sting. 

Experiments made by Carlet (1890) 
show that it is only the mixture of the 
products from the two poison glands 

that is fully effective in stinging properties. Carlet^s experiments were 

made upon houseflies and blowflies. He shows ( 1 ) that flies stung by a 

bee die almost instantly, (2) flies artificially inoculated with the secre- 




BM- 




,Lum 



Epth 



Pig. .39. — Sections of alkaline gland 
of sting. 



80 THE ANATOMY OF THE HONEY BEE. 

tion of either gland alone do not die for a long time even in spite of 
the necessary mutilation, while (3) successive inoculations of the 
same fly first from one gland and then from the other produce death 
in a much shorter time than when inoculated from one gland alone — 
presumably as soon as the two liquids mix within the body. 

The two secretions, one acid and the other alkaline, are poured 
together into the base of the sting bulb and mix within the cavity 
of the latter. The resulting poison is then driven through the chan- 
nel in the shaft to near the tip of the latter, where it makes its exit 
into the wound. Since the large poison sac is not muscular, the poison 
is not forced through the sting by it, as is often supposed. A glance 
at figure 57 (see p. 135) will show that the accessory plates of the sting- 
support several very compact sets of muscles on their inner faces. 
These muscles so act during the process of stinging that the triangular 
plates (figs. 36 and 37, Tri) turn upon their hinge- joint articulations 
with the oblong plates {Oh). By this motion of the triangular 
plates the attached lancets {Let) are moved back and forth along 
the tracks on the lower edges of the sheath and its arms {ShA) . 
Each of these tracks consists of a ridge with a constricted base which 
dovetails into a correspondingly shaped groove on the dorsal surface 
of the lancet. This structure, as seen in cross sections through the 
shaft and bulb of the sting, is shown by fig. 40 A, B, and C. The 
lancets are thus held firmly in place, while at the same time they may 
slide back and forth with perfect freedom. The figures show also 
that all three parts of the sting are hollow, each containing a pro- 
longation {he) of the body cavity. Between them, however, is in- 
closed another cavity through wdiich the poison flows. This is the 
'poison canal {PsnC). In the bulb (fig. 40 C) the body cavity is 
reduced to a narrow cleft {he) by the great size of the invaginated 
poison canal {PsnC). 

It will now be most convenient to describe the apparatus by means 
of which the poison is ejected from the sting. As before pointed out, 
the large poison sac can have no functions in this connection because 
its walls are entirely devoid of muscle fibers. On the other hand, 
there is an actual j^umping apparatus situated within the bulb. This 
consists of two pouchlike lobes, having their concavities directed 
posteriorly, attached to the upper edges of the lancets (fig. 40 D and 
G, Vlo) on the anterior ends of the parts of the latter which slide 
within the lower edges of the bulb chamber. The lobes lie side by 
side within the bulb (fig. 40 C, FZy), when the lancets are in the same 
position, and each has an accessory lamina against its own inner wall. 
When the lancets are pushed backward the walls of the lobes flare 
apart against the poison contained in the bulb and drive this liquid 
before them into the channel of the shaft, while at the same time they 
suck more poison into the front of the bulb from the glands. AYhen, 



THE ABDOMEN, WAX GLANDS, AND STING. 



81 



PsnC-, 



PsnC 



on the other hand, the lancets are retracted the pouches collapse so 
that they may be drawn back through the poison-filled bulb without 
resistance, but they are ready for action again as soon as the move- 
ment of the lancets is reversed. The whole apparatus thus consti- 
tutes an actual force pump in which the lobes on the lancets alter- 
nately act as a piston and as valves. The lancets need not work 
together ; in fact, 
they more often ^ -^^^ 

perhaps work al- 
ternately, the lobes 
being of such a 
size as to be ef- 
fective either when 
acting together or 
separately. 

The reader ac- 
quainted with 
other Avorks on 
the anatomy of 
the bee, such as 
those of Cheshire 
(18 8 6), Cook 
(1004), Cowan 
(1901), and Arn- 
hart (1906), will 
see often repeated 
the statement that 
the poison leaves 
the sting both by 
a ventral oj^ening 
between the lan- 
cets near their tips 
and by several lat- 
eral pores near the 
ends of the lancets 
opening from the 
poison canal upon 
the bases of the barbs. The writer, however, has never been able 
to observe the exit of the poison from any such lateral pores, while, 
on the other hand, it is very easy to watch it exude from between 
the lancets on the ventral side of the sting near the tip. If an 
excited bee is held beneath a microscope and the tip of the sting 
observed, the poison will be seen to accumulate in little drops near 
the tip on the ventral side. If, also, the bulb of an extracted sting 
22181— No. IS— 10 6 




Fig. 40. — Details of sting of worlvor : A, section througli tip of 
sting sliowing lancets (Let) and sliaft of sheath (ShS) sur- 
roimding central poison canal (PsnC), and each containing 
a prolongation of the body-cavity (be) ; B, section of same 
near base of bulb ; C, section of sting tlirough basal bulb, 
showing poison canal as large invaginated cavity (PsnC) 
in bulb of sheath (8hB) containing the two valves (Vlv) 
of lancets (Let) ; D, part of left lancet carrying valve (Vlv), 
dorsal view ; E, tip of lancet showing pores opening on 
bases of barbs (oo) coming from body-cavity (6c) of lancet — 
not from poison canal ; F, dorsal view of shaft of sheath 
showing lateral series of pores (oo) from prolongation of 
body-cavity (be) ; G, lateral view of left valve and part of 
lancet. 



82 



THE ANATOMY OF THE HONEY BEE. 



be squeezed gently between a pair of forceps the poison will be seen 
to emerge in the same way. In fact, it can be actually squirted out 
by a sudden compression wdien the bulb is well filled with poison, but 
there is never any evidence of its escape through the sides. 

An examination of the end of each lancet does reveal a number of 
oblique pores (fig. -iO E, oo) Avhich have been figured by other writ- 
ers, and they certainly open on the bases of the barbs as described, 
but their inner ends apparently communicate with the body cavity 
{he) of the lancet instead of passing clear through the lancet and 
opening into the poison canal. Furthermore, a paired series of 
exactly similar pores extends the entire length of the shaft of the 
sheath (fig. 40 F, oo)^ opening on its dorsal surface from the body 
cavity {he). No one could possibly claim that the poison emerges 




Tri Ob 

Fig. 41. — Tip of abdomen of worker with left side removed, showing right halves of sev- 
enth tergum (VIIT) and sternum (VIIS), containing the sting chamber (Iclc) cut open 
along the line bir, exposing the eighth tergum (VIIIT), the rudimentary tenth segment 
(X) carrying the anus (An), and the sting and accessory parts shown by fig. 36. 

also through these pores, which, very curiously, do not apjDcar to 
have been described before, although they are even more conspicuous 
as well as more numerous than those of the lancets. The writer has 
not been successful in preparing histological sections of the sting 
which show these pores, but they probably constitute the ducts of 
some kind of subcuticular glands. 

A cross-section through the sting a short distance in front of its 
tip shows that the lancets are here separated by a narrow cleft (fig. 
40 A), while elsewhere (B and C) thcA^ are contiguous. This cleft 
between the ends of the lancets forms the exit for the poison from the 
channel. 

The sting of the queen is much longer than that of the worker 
! nd is more solidly attachecl within the sting chamber. Its shaft is 



THE ABDOMEN, WAX GLANDS, AND STING. 83 

strongly decurved beyond the bnlb. The lancets have fewer and 
smaller barbs than those of the worker, but the two poison glands 
are well developed (fig. 57, AGl and BGl), while the poison sac 
{PsnSc) is especially large. 

A number of minute unicellular glands open upon the interseg- 
mental membrane between the seventh and eighth terga of the ab- 
domen. These are sometimes called the glands of Nassanoif, after 
their discoverer. Nassanoff suggested that they are sweat glands, 
while Zoubareff thought that they form small drops of liquid said 
to be excreted by bees during flight derived from the excess of water 
in the newly collected nectar. Their function, how^ever, has been 
much more carefully investigated by Slaclen (1902), who found that 
they are scent organs producing a strong odor even when the part 
of the back to which they are attached is removed from the rest 
of the abdomen. He furthermore identified this smell as the same 
that bees give off when a lot of them are shaken from a frame on 
the ground close to the front of the hive. Under such circumstances 
also, as in natural swarming or during the first flights in the spring 
or after a period of bad weather, bees are well known to produce a 
peculiar sound called the " joyful hum." Sladen observed that this 
was produced, in the case of bees shaken before the hive, by those 
individuals who first found the hive entrance, then by those next to 
them, until very soon all the others were informed of the location 
of the entrance and proceeded to make their way in. Also, when a 
swarm loses sight of its queen, those 'that find her first set up this 
" joyful hum " and immediately the rest of the swarm is attracted 
to the spot. In the springtime the young bees seem to be guided 
in their flights by this same hum of the old ones. Sladen, however, 
observing the ddor emitted at the same time, thinks that this and 
not the sound is the real means of information, the sound being 
simply incidental to the special movement of the wings produced 
for the jDurpose of blowing the odor away from the body. He argues 
that we have no evidence of an acute sense of hearing in bees, while 
it is well known that they possess a delicate sense of smell located on 
the antenna?. This argument certainly seems reasonable, and we 
may at least accept Sladen's theory as the best explanation of the 
function of the glands of Nassanoff. 



84 THE ANATOMY OF THE HONEY BEE. 

VI. THE ALIMENTARY CANAL AND ITS GLANDS. 

1. THE GENERAL PHYSIOLOGY OF DIGESTION, ASSIMILATION, AND 

EXCRETION. 

It is no exaggeration to say that eating is the most important thing 
that any animal does and that its alimentary canal is the most im- 
portant organ it possesses. The entire system suffers when there is a 
deficiency in the food supply or an impairment in the digestive appa- 
ratus. Every other function is either subservient to or dependent 
upon that which furnishes nourishment to the cells. The senses of 
sight, smell, and taste are all more or less concerned in the acquisition 
of food. The muscular system enables the animal to hunt for it, to 
dig for it, to climb for it, or to chase living prey either on the ground, 
in the water, or in the air, and to kill, tear, and chew it when ob- 
tained. The blood is the servant of the stomach, for its entire func- 
tion in insects is to carry the products of digestion to the body cells. 
The heart furnishes the motor power of the blood. The respiratory 
function is accessory to that of digestion, inasmuch as it furnishes the 
oxygen which unites with the waste materials ejected from the cells 
and renders them capable of being removed from the blood. This 
removal is accomplished partly by the respiratory system itself and 
partly by special excretory organs. Thus we see that the sense organs 
and the muscular system are the agents that cooperate in obtaining 
the raw food, the digestive tract is the kitchen of the body in which 
the food is prepared for use, the blood is the waiter that distributes 
it, while the respiratory and excretory systems are the refuse gath- 
erers that remove waste products. The nervous system holds the con- 
trolling power over all these organs. It regulates them in the per- 
formance of their duties and coordinates their actions so that they 
all work together. It makes a unified organism out of what would 
otherwise be simply a complex mass of variously specialized cells. 

The reproductive function alone contributes nothing to the indi- 
vidual. In fact, the production of spermatozoa by the male and of 
eggs by the female and the nourishing of the embryo and the young 
create a demand upon all the other organs for material which is 
separated from the individual that produces it. But this is what the 
organism exists for; this is its reason for being. At least this is 
what it amounts to in the case of the individual, though from a wider 
philosophical standpoint the real truth is probabl}^ just the reverse, 
viz, any species exists because its individuals reproduce themselves. 

The writer has already made frequent use of the word " cell," 
assuming that the reader is familiar with the meaning of this word 
as used in anatomy and physiology. The entire body of an animal 
or plant is made up of cells or their products. The word, however, is 
misleading, for a cell is not a small sac or empty space, as was at 



THE ALIMEJSTTAEY CANAL AND ITS GLANDS. 



85 



SInr 



\fentJ 




Fig. 42. — Alimentary canal of worker (Phy-Rect), together with pharyngeal glands (IGl), 
and salivary glands of head (201) and of thorax (SOD, as seen by cutting body open 
from above and pulling the ventriculus (Vent) out to left. 



86 THE ANATOMY OF THE HONEY BEE. 

first supposed from the study of plants, but is a little protoplasmic 
body or corjDuscle, visible only under the microscope, surrounded by 
a membranous cell wall and containing a small internal body called 
the nucleus. The different cells of the body are specialized in groups 
to do some one particular thing — the salivary cells secrete saliva, the 
muscle cells contract, the excretory cells pick out waste substances 
from the blood, and so on. But this specialization does not signify 
that each cell does not perform its own vital processes in addition to 
its specialt}^ The fact that it remains alive and works means that 
the complex chemical components of its body substance or frotoflasm 
are constantly being reduced to simpler compounds which are ex- 
pelled, while new protoplasm is built up from the supply of food 
material brought by the blood. This double process of destruction 
and reconstruction is known as metaholism^ while its two phases, the 
breaking-down process and the building-up process, are known as 
katabolism: and ancibollsm^ respectively. 

Now, while all the cells of the body must have nourishment, none 
of them, except those of the alimentary canal, is capable of utiliz- 
ing the raw food materials that an animal obtains in a state of nature. 
These materials must therefore be changed into some other form in. 
order that they may be assimilated by the cells. This change is called 
digestion. 

The single cell composing the body of a Protozoan, living free in 
nature, digests its own food and then assimilates the products of its 
own digestion. But, of the cells constituting the body of awj mul- 
ticellular animal, only those of the alimentary canal are capable of 
digesting raw foodstuffs, and, moreover, as digestion is the specialt}^ 
of these cells, thej^ have also to digest the food for all the other cells 
of the bodj^ 

The two most important changes that must be brought about in 
the natural food by digestion are those which make it soluble in the 
blood and which render it capable of passing through animal tissues. 
1)1 the first place, the food must diffuse through the walls of the 
alimentary canal as a liquid which mixes with the blood, for there 
are no pores or openings of any sort from the alimentary canal into 
the body cavitj^; and in the second place, it must pass through the 
walls of the cells themselves. The digestive changes result chiefly in 
a breaking down of the complex molecules of the raw food materials 
into more simple chemical substances. These are taken up by the 
cells and reconstructed into complex protoplasmic molecules Avhich 
can not escape through the cell membrane until thej^ are again broken 
down into simpler forms. 

The waste products of the cells consist j)rincipally of carbon, hy- 
drogen, and nitrogen. These are converted by the oxygen supplied 
by the respiratory system into carbon dioxid, water, and compounds of 



THE ALIMENTARY CANAL AND ITS GLANDS. 87 

urea. The first, being a gas, mixes with the air in the tracheal tubes 
and so reaches the exterior during exhalation. Much of the water is 
also given off through the tracheal system in the form of vapor which 
exhales from the spiracles, but, since insects are covered by their 
hard chitinous shell, it is probable that they do not " sweat." The 
compounds of urea, and probably also some water, are separated 
from the blood by the excretory glands, called Malpighian tubules 
in insects, which empty their products back into the alimentary 
canal, whence they are discharged with the fseces from the intestine. 

Digestion is brought about by substances called enzymes which are 
contained in the various liquids mixed Avith the food in the alimentary 
canal. These liquids are secreted by the salivary glands and by the 
cellular walls of the stomach. 

2. THE SALIVARY GLANDS. 

The opening of the salivary duct on the base of the proboscis has 
already been described (see pp. 49-51). The true salivary glands, or 
those corresponding with the salivary glands of other insects, are 
arranged in two pairs, one situated within the head (figs. 19 and 42, 
2GI) and the other within the thorax (fig. 42, 3GI). The four ducts 
unite into one median tube, wdiich enters the base of the labium (fig. 
19, SalD) and opens upon the upper surface of the ligula (fig. 15 F, 
and fig. 16, SalDO). The large and conspicuous glands lying within 
the anterior and upper parts of the head ancl opening into the 
pharynx will be described later in connection with this organ. They 
are special pharyngeal glands in no way homologous with the salivary 
glands of other insects, and are by many supposed to secrete the 
brood food instead of a digestive liquid like saliva. 

The salivary glands of the head {System No. 2 of Cheshire, "post- 
cerehral glands of Bordas) lie against the posterior walls of the 
cranium. In the worker each consists of a loosely arranged mass of 
pear-shaped follicles or acini whose individual ducts unite irregu- 
larly with one another and eventually form a common duct on each- 
side (figs. 19, 42, and 43 F, 2GI). Their two ducts unite with the 
median duct from the thoracic glands just before the bases of the 
mesocephalic pillars (fig. 19). In the drone these glands have a 
quite different appearance from those of the female, each consisting 
of a compact mass of very small follicles connected by minute ducts 
and flattened against the posterior Avails of the head (fig. 43 B and C, 
SGI). A large lobe of this gland in the drone extends forAvard on 
each side against the face, betAveen the compound eye and the clypeus 
(fig. 10 C, SGI)., thus occupying the position of the large mandibular 
gland in the worker (A, IMdGl) and in the queen (B, IMdGl). 
There is also a prominent triangular mass of glandular cells in the 
drone situated just above the ocelli (fig. 10 C, 2GT) Avhich has been 



88 THE ANATOMY OF THE HONEY BEE. 

described b}^ Bordas (1895) as a separate gland opening by two ducts 
into the oesophagus just behind the pharynx. The writer, however, 
has been utterly unable to discover any such ducts, though two sus- 
pensorial ligaments of the anterior end of the oesophagus are at- 
tached to the wall of the head at the posterior ends of these glands 
(fig. 11 B, ^) and might easily be mistaken for ducts. These " post- 
ocellar glands " of Bordas, moreover, appear to be simply detached 
lobes of the postcerebral glands. They are jDrominent also in the 
queen (fig. 10 B, 2GI) and are represented by a few follicles in the 
worker. 






^^ A ^^af4g ^^ 




mm 






D 



^ 




Fig. 43. — A, small piece of large lateral pharyngeal glands In liead of worker ; B, piece of 
postcerebral salivary glands in head of drone ; C. postcerebral glands (2G?) in normal 
position against posterior wall of head in drone ; D, pharyngeal plate {s) of worker, 
ventral view, showing bases of lateral pharyngeal glands (IGl) and their receptacula 
(»]»))■), and median ventral pharyngeal gland (^iQl) ; E, corresponding view of pharyngeal 
plate of drone, showing entire absence of lateral pharyngeal glands, and greater devel- 
opment of small median glands (liGl) ; F, part of postcerebral gland of worker. 

Bordas describes the follicles of the postcerebral glands in the 
worker as hollow sacs, each having a large lumen lined with a chiti- 
nous intima. Their secretion, he says, is a thin viscid liquid, pale 
yellow" in color and having a slightly alkaline reaction. According to 
Schiemenz (1883) each gland is developed as an outgrowth from the 
common duct of the thoracic glands. 

The salivary glands of the thorax in the bee {System No. 3 of 
Cheshire, tJioracir mliravy glavds of Bordas) are the ones that cor- 
respond with the ordinary salivary glands of other insects. Thej'' 
are described by Schiemenz (1883) as being formed inside of the 



THE ALIMENTARY CANAL AND ITS GLANDS. 89 

outer covering (tunica propria) of the first part of the larval silk 
glands. But it is of common occurrence in insects that the salivary 
glands are temporarily specialized as silk-producing organs in the 
larva. In the adult worker these glands lie in the ventral part of 
the anterior half of the thorax (fig. 42, ,^6^/). The two are widely 
separated anteriorly, but their posterior ends are contiguous. Each 
consists of a mass of small, many-branched, glandular tubes opening 
into several collecting ducts which empty into a sac near the ante- 
rior end of the gland {II) . From each of these reservoirs, then, a duct 
{Dct) runs forward and fuses with the one from the opposite side 
just within the foramen magnum of the head. The common duct 
thus formed turns downward within the head, receiving the two ducts 
of the postcerebral salivary glands and then enters the base of the 
mentum (figs. 19 and 43 C, SalD) , to open as already described on the 
upper side of the ligula at the root of the glossa and between the 
bases of the two paraglossae (fig. 15 F and 16, SalDO). The 
secretion of the thoracic glands is said also to be weakly alkaline. 
Therefore the entire salivary fluid poured out upon the labium is 
alkaline, and it must be designed to act especially upon the food 
taken through the proboscis. This action, furthermore, on account of 
the location of the salivary opening, may take place before the food 
enters the mouth. 

The food of the bee consists normally of pollen, nectar, and honey. 
The first is eaten entirely with the mandibles, while the other two are 
taken through the proboscis. The pollen is to the diet of the bee what 
meat is to ours; that is to say, it contains the jjroteid or nitrogen- 
containing ingredient of the food which is necessary to the sup- 
port of any animal, and also substances comparable with fat, called 
in general hydrocarhons. The nectar and honey consist principally 
of grape sugar, fruit sugar, and cane sugar, which belong to the class 
of chemical substances known as carhohydrates. Now, all of these 
foodstuffs, except the grape and fruit sugars, have to be changed 
chemically by the digestive process before they can be absorbed into 
the blood. The pollen, wdiich contains the proteids and hydrocarbons 
of the food, is taken directly into the mouth by means of tlie man- 
dibles and apparently is not digested until it reaches the small in- 
testine, and therefore it would seem that it is the cane sugar wdiich 
must be affected by the saliva. The change, or inversion, as it is 
called, of cane sugar, which has a very large molecule (CioHo.O^i), 
consists of its reduction to grape and fruit sugars which have smaller 
molecules (CpHijOe). Starch (CgH^oOs) must also be reduced to 
simpler and more soluble compounds before it is capable of absorp- 
tion. Its iuA^ersion is effected in us partly by the saliva, but starch 
appears to form a very inconsiderable element in the bee's diet. 



90 THE ANATOMY OF THE ITONEY BEE. 

3. THE ALIMENTARY CANAL. 

The alimentary canal is a tube Avhich extends through the entire 
length of the body and, on account of being more or less coiled, it is 
generally considerably longer than the length of the body in insects. 
It has no openings of any sort into the body cavity. The internal 
organs are packed closely about it, and the interstices are filled with 
the blood, there being no special arteries or veins in insects. The 
amount of space occupied by the alimentary canal varies according to 
the amount of food it contains, and for this reason it seldom looks 
exactly alike in any two individuals examined. 

The j)art of the canal immediately following the mouth forms an 
enlargement (fig. 42, Phy) called the pharynx. Succeeding this is 
a slender tube which leaves the head by the foramen magnum above 
the small transverse tentorial bar and traverses the entire length 
of the thorax. This is the cesophagus {(E). In the anterior part of 
the abdomen the oesophagus expands into a large thin-walled sac 
which is ordinarily called the croj) or inghivws^ but which, in the 
bee, is known as the honey stomach {IIS). Behind this is a short, 
narrow, necklike division, with rigid walls constituting the jwo- 
ventriculus {Pvent). Then comes a large U-shapecl part, with thick, 
spongy-looki