THE PAN-PACIFIC ENTOMOLOGIST

EDITORIAL BOARD

E. G. Linsley P. D. Hurd, Jr., Editor R. L. Usinger

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THE SUCKING LICE by G. F. Ferris $6.00

A 320-page book which summarizes the knowledge on

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The Pan-Pacific Entomologist

Vol. XXXV January, 1959 No. 1

GORDON FLOYD FERRIS— 1893-1958

Robert L. Usinger

University of California, Berkeley

For some individuals, a full biography is justifiable only as a

matter of record. With Gordon Floyd Ferris the situation is

different. Here was a man of originality with much to tell to the

world and a gifted pen with which to tell it. In a life devoted to

principles as he saw them and to ideals as he conceived them, there

emerged a profound truth which, unfortunately, he never put into

words. It is that a scientist, endowed with ability and placed in a

favorable environment, has an overwhelming obligation to produce

for the society that supports him. This obligation, as Ferris saw

and filled it in fullest measure, was not necessarily to produce

practical results but rather to seek the truth in all things and to

make the results known. It happened that some of the work that

Ferris did with lice and scale insects was of direct importance to

mankind. But he insisted that his interest in these insects had no

connection with their actual or potential economic importance.

Instead, he was motivated solely by the fact, mystical or otherwise,

that he liked them. In a moment of self analysis (Canad. Ent. 56:25,

1924) he wrote on this as follows:

“I have often wondered what it is that determines our special

interests. Why does one entomologist prefer to work on dragon-

flies, another on butterflies and a third on beetles? ... If we were

to study man as we do other animals we would doubtless devise

a special terminology to ‘explain’ these things. We would say that

so-and-so is positively hemipterotropic or papiliotropic, or what-

ever else it may be, and let it go at that with a rather comfortable

feeling that we have disposed of the matter! ... As for myself,

I am irresistably drawn toward the smallest and most intrinsically

uninteresting of insect forms. I can appreciate the pleasures of

the collector who looks with gloating eyes upon his well-filled

cabinet of glorious butterflies or gleaming beetles, but I can not

expect him in turn to do more than wonder why I also should look

with the same feelings upon a well-filled cabinet of little pieces of

glass, each one with one or two minute specks in the centre of it.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Gordon Floyd Ferris (1893 - 1958 )

But it is so, and my knowledge of ancient languages is not sufficient

to enable me to coin a word behind which it is possible to take

refuge. I merely know that if an insect is too large to go on a slide

1 leave it for someone else, and if it is small enough to go on a

slide 1 have an impulse to put it there.”

Throughout his life Ferris, who was a large man, maintained

this interest in small insects.

Early life. The following quotation is from a brief biographical

sketch prepared by Ferris shortly before his death;

“Born January 2, 1893, at Bayard, Allen County, Kansas, a “tank town”

where his father was a gandy dancer on the railway. He was the fifth child

and the fourth boy in a family of hve. When he was between two and three

January, 1959]

years old the family moved to a 40-acre farm near Monticello, Cedar County,

Missouri, where they lived in a one-room log cahin. When he was a little

more than three years old his mother died after giving birth to another son,

who also died while still young.

“ . . . He and his sister went to live with his paternal grandmother and

an unmarried uncle on a farm near La Harpe, Allen County, Kansas. At the

age of about 10 years the grandmother died . . . His father thereupon moved

to La Harpe with the two brothers who had remained with him, obtained

employment as a day laborer in a zinc smelter . . . and . . . established a

“bachelor” home in La Harpe with the two brothers.

“Gordon remained with his father until he was 13, except for two

summers spent as a hired hand on a farm near lola, Kansas. In the meantime

his oldest brother, Leslie, . . . had enrolled at Ottawa University, a small

Baptist college at Ottawa, Kansas, where he acted as the local circulation

agent for the Kansas City Star. When Gordon graduated from the eighth

grade at La Harpe, at the age of 13, Leslie had him come to live with him.

With the support of Leslie and such money as Gordon could make by

carrying papers ... he remained at Ottawa until 1909, having in the

meantime graduated from the “academy” at Ottawa University . . . Leslie

graduated from the college in 1909 and secured a position as a teacher in

the high school at Telluride, Colorado. He left Gordon as circulation agent

for the Kaiisas City Star in Ottawa and Gordon entered Ottawa University

as a freshman in the fall of 1909. He was a complete failure at this, withdrew

from the college, and proposed joining the Navy as soon as he became 17.

“It was necessary to obtain the consent of his father to join the Navy.

The papers were sent to his father, who refused to sign them without first

consulting Leslie. Leslie refused to give his consent and instead secured a

place for Gordon with the Telluride Power Company at Telluride.

“ ... In its early days the Telluride Power Company had trouble securing

trained men, and under the leadership of L. L. Nunn, its founder, had

established the practice of employing young men at reduced salary and after

a time sending them to college. Out of this grew the Telluride Association,

an endowed institution, entirely apart from the power company ... In the

spring of 1911 Gordon was transferred to the station of a power company

at Olmstead, Utah, where the Telluride Association ran a school for its

members.

“The Telluride Association maintained a house at Cornell University,

where most of its men who were selected for “preferment” went. Gordon

disapproved of the social emphasis at this house and asked instead to be

sent to Stanford University. He had seen Kellogg’s American Insects while

he was at Ottawa, and Kellogg — ihen a professor at Stanford — was the only

entomologist he had ever heard of. In the summer of 1912 he was granted

$450 by the Telluride Association and came to Stanford. With the continued

support of the Association — never amounting to over $500 a year — he finished

his work for the degi'ee of M.A. in 1917.

“He was then appointed a teaching assistant in entomology at Stanford

and further data will be a matter of record.”

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Field work. To those whose only contact with Ferris was in the

laboratory or classroom it will come as a surprise to learn that

he was a dedicated field naturalist. He had heen at Stanford for

only one year when, doubtless through an interest in ectoparasites

kindled by Vernon Kellogg and the contact with Joseph Grinnell,

he joined a field party of the University of California Museum of

Vertebrate Zoology. The trip covered the north coast counties of

California. According to C. L. Camp, who was a member of the

party, (personal comm.) Ferris’ main job was to comb the verte-

brates for ectoparasites, but the bound collection of field notes in

the MVZ files shows that the 20 year old student’s interests ranged

widely and his handwriting, which he always deprecated, was

already, at that early age, nearly illegible. On July 24, 1913, three

miles south of Covelo, Mendocino County, he wrote: “While out

hunting butterflies in the afternoon 1 heard a loud squeak in the

oak tree above my head and looked up just in time to see a small

wood rat come hastily out of a pile of sticks on a limb and run

rapidly down the tree. I could see the folds of a snake in the sticks

and upon shooting into them, a large gopher snake, perhaps three

and a half feet long, fell to the ground. It lay beneath some sticks

and produced a rather loud buzzing sound by beating the tip of

its tail against a stick. It escaped into a hole when I attempted to

capture it.” Two years later Ferris joined another MVZ team and

collected ectoparasites from birds and mammals in the Yosemite

region.

Other extensive field trips included: Lower California, with

J. R. Slevin (1919) ; Southwestern United States investigating lac-

producing scale insects with Roxana S. Ferris (1918) and daughter

Beth and C. D. Duncan (1920) ; Mexico (Guggenheim Grant)

(Fall of 1925 to summer of 1926) visiting Sinaloa, Nayarit, the

Tres Marias Island, Colima, thence south by foot to Acapulco,

the east and west coasts of the Isthmus and Mexico City; Lower

California, with Shreve, Mallory and R. S. Ferris, crossing over

to Sonora on return (March and April, 1934) ; Panama including

the Volcan de Chiriqui, Puerto Armuellas, David and Boquete

(summer of 1938) ; Arizona, with L. P. Wehrle and R. S. Ferris

(1940) ; China (Fulbright Award) including Yunnan, Hong

Kong, and New Territories, Kwangtung and Taiwan (Fall of 1948,

spring, 1949). Also Ferris spent a year studying Anoplura at the

January, 1959]

Molteno Institute, Cambridge University and at the British Museum

(Natural History) (1930—1931).

As in other facets of his life Ferris was unpretentious in his

field work. He pointed this up in an unusual article in the Scientific

Monthly titled “Scientific Exploration, A Phantasy” (Sci. Mon.

27 :537— 541, 1928) . In the story a lone and weary collector pitches

camp and falls into a fitful slumber broken by bouts with tropical

fever. He dreams of a super expedition supplied by airlift with

the most elaborate equipment and the most luxurious food. Being

depedent on money in large amounts brought in by public rela-

tions men it is finally decided that the scientists, who are merely

an impediment, must go. Although facetious, the lone scientist is

obviously Ferris and the great expedition could be any one of a

number organized in recent years.

Systematics. Ferris was primarily a taxonomist and in this, as

in other activities, he brought a fresh approach and a plan for

work. Starting with the lice under Kellogg and the Coccidae under

Doane he soon set a pattern from which he deviated but little in

later years. Foremost was his insistence on detailed drawings —

doubtless as a reaction to the shocking state of knowledge of lice

and coccids based largely on inadequate descriptions. In his first

paper, with Vernon Kellogg, the drawings are not divided by a

vertical line into ventral and dorsal halves, but he soon adopted

this economical method and gave his philosophy on drawings in

1923 (Sci. 58:266): “A scientific illustration is not intended

merely as a pretty picture and it has nothing to do with art. Its

purpose is merely to present in the simplest and most accurate

manner the things that it is desired to show and its production

involves nothing more than good draftsmanship. If in addition to

these qualifications it is also artistic — whatever that may mean —

so much the better.” The arguments for divided drawings were

given as follows: 1) they save in cost of blocks; 2) they save in

the time of making drawings; 3) they save space in printing;

4) they are more convenient because both sides can be readily

compared.

Once started on the course of drawing details of minute insects

instead of describing them, Ferris was committed to a compound

microscope and pen and ink for the rest of his life. With prodigious

energy and usually with several projects running concurrently he

turned out monumental works. He provided the foundations for

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

our present knowledge of the Anoplura, Mallophaga, Coccoidea,

Diptera-Pupipara, Cimicidae and Polyctenidae. Probably no man

has ever made so many original detailed drawings of insects.

The cost of reproduction of illustrations was a constant problem

in his papers which reversed the usual ratio between text and

figures. To solve this Ferris turned to off-set lithography in place

of conventional letterpress. The result was Microentomology, his

own journal, in the Foreword (Microent. 1:1, 1936) of which he

states his “intention to speak as far as is practicable through the

medium of illustrations of which the text will be merely explanatory

and confirmatory.” This emphasis was shown in the papers that

we published together on Polyctenidae and Cimicidae. Ferris

devoted himself exclusively to the illustrations (which of course

could stand alone) and left the text entirely to me.

Although the off-set method reduced costs, the size and number

of Ferris’ publications raised problems of financing. Curiously,

the Atlas of Scale Insects was his greatest problem and required

the largest outlay of private funds. Here was a work of great

economic importance, a standard reference for economic ento-

mologists, yet such was the attitude in support of scientific work

before and during World War II that Ferris announced his inten-

tion (fortunately not carried out) to abandon the project after

Series IV. He reasoned that, “If the world wants this Atlas of the

Scale Insects of North America completed the world must find

some way to pay for it.” That the world did find a way to pay

for it is no great credit to the “world” because it was only after

Ferris had contributed not only the scientific work but also a

substantial amount of money that the work was continued. As

early as 1919 Ferris wrote an article (Science, 50:543) : “Why

not government-maintained fellowships?” but it took a cold war

to bring us to the present level of support for the things that

Ferris strove for.

The first and only “book”, as distinguished from monographs,

atlases, etc., written by Ferris was “The Principles of Systematic

Entomology” (Stanford University Publications, University Series,

Biological Sciences, Vol. 5, No. 3, with reprints bound and sold

separately) . Here again we see freshness and originality and some-

thing else that came to be a trademark. Sometime during the 1920s

Ferris developed a style of writing that became stately but not

pompous. As a first statement of the principles of a science his

January, 1959]

book stands as a classic. The principles stated in this book are

sound and were a part of Ferris’ own life and work. However, it

is necessary to point out that his strongest asset — originality — led

to a weakness which will probably not be serious because it can

be supplied by later workers. Ferris assumed that all work previous

to his own in a group was of dubious value and not worth the

trouble to look up. As he put it: (Microent. 13(3) :52, 1948) “We

would seem to be justified in the assumption that a very large

part of what was done in the earlier years is likely to be either

wrong or inadequate.” Continuing (Microent. 15(4) :127, 1950) :

“What is to be gained by laborious attempts to disassociate fiction

from fact until some basis for judgment has first of all been

established? Let us begin with first things first, and proceed then

step by step.” This, to my knowledge, has been the main basis for

criticism of Ferris’ work, a small but unnecessary price to have

paid for such an original and productive life.

The Plant Quarantine Controversy. Ferris was a fighter in the

cause of freedom. He saw in the inspection of personal belongings

at the borders of California an infringement of the rights of

citizens. This led to a frontal attack on the government agencies

dedicated to eradicating the Mediterranean Fruit Fly from Florida.

It also led, unfortunately, to personal feuds with friends and

colleagues in entomology but the outcome, looking back after 20

years, was beneficial to everyone. Ferris’ position was stated as

follows (Science, 70:452, 1929) : “Theoretically, perhaps, the fruit

fly can be eradicated [it was ! ] . Theoretically any insect can be

eradicated — at a price. A country can be converted into a desert.

Its inhabitants can be ruined. It can be depopulated . . . Even

should the fruit fly be for the time being ‘eradicated’ — from a

biological point of view a most improbable outcome — at whatever

cost, it will inevitably come again [it did!] and the whole per-

formance must again be gone through. Those who believe that

any quarantine can guarantee future immunity are leaning upon

a hollow reed.” In a later article (Science, 71:606, 1930) he states

that, “It would seem desirable for the supporters of these measures

to enlarge the range of their view sufficiently to include a larger

proportion of the facts of the situation.”

Summarizing his case against the “agricultural bureaucrats”

for their “kill or cure policy” and for wasting millions of dollars

of the taxpayers’ money on what he regarded as an unsound

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

biological principle, he suggested that a commission be appointed

“to consider with a cold and critical eye the whole question of

plant quarantines from biological, sociological and legal points

of view.” This was done and the final report, produced after several

years work by a team of highly competent university and govern-

ment scientists, was reviewed by Ferris with a final comment as

follows : “It is a tribute to the intellectual honesty of the members

of the committee that the reviewer — an avowed enemy of the quar-

antine system — can think of no critical phrase to apply to the

report more condemnatory than ‘unduly cautious’.”

Scientific Affiliations. Ferris was not an “organization man” or

“joiner”, yet he had certain deep seated loyalties. His scientific

“home” away from the Stanford campus was the Pacific Coast

Entomological Society. He was elected a member of the Society

at the 74th meeting on December 13, 1919, at which time he

reported on his recently completed trip to Lower California. My

first acquaintance with Ferris was at Society meetings in the late

1920s, held in Room 10 of the San Francisco Ferry Building. At

first it was a mystery to me why Ferris attended because the meet-

ings were dominated by talk of beetles and butterflies and the

philosophy of the long-time president, E. C. Van Dyke. I remember

in the depression years sitting on the publication committee of the

Pan-Pacific Entomologist when the “hat was passed” to make up

each year’s deficit. Ferris always contributed what impressed me

as large amounts.

The most important part played by Ferris in Society activities

was his association with successive generations of students from

neighboring universities. Hundreds of students outside his own

Stanford group came to know him in this way. Usually, he would

bring a folder of drawings and lay them out on a table for inspec-

tion. Or, in later years, he would present a controversial point in

morphology.

Morphology. Ferris liked to tell this story of his entry into the

field of comparative morphology. He was invited to do the chapter

on morphology for the book “Biology of Drosophila” as originally

planned by A. F. Heutter in 1939 and finally published under the

editorship of M. Demerec in 1950. According to Ferris this com-

mitment led him back to the primitive Diptera (Tipulidae), thence

to the Mecoptera and finally back to the segmented worms

(Annelida). Whether the sequence is literally true, I cannot say.

January, 1959]

but this much is certain: Ferris had always been interested in

structure as revealed by the techniques used for preservation of

his small insects. Then, in the mid-1930s, he was profoundly in-

fluenced by the breadth of thinking of a newly formed discussion

group in California, the Biosystematists, made up of active students

of evolution representing various disciplines in the biological

sciences. Thus began another of the major developments in his

varied career. Although continuing with the detailed drawings for

the “Atlas of Scale Insects” it was clear that his main interest

and intellectual challenge was comparative morphology in its

broadest sense.

In typical fashion he started with a firm set of principles and

with a plan, almost a schedule, for production. Some idea of the

scope of his thinking may be obtained from the following quota-

tion: (Microent. 13(3) :55— 56, 1948) “When — far in the future —

we really have a sound and broad knowledge of insect morphology,

we shall be able to see, upon the stage of that magnificent theater

of biological processes which have produced the greatest of all

animal phyla, the Arthropoda, the unfolding and flowering of the

processes of evolution with a clarity and degree of detail that can

nowhere else be attained.”

Characteristically, Ferris found himself almost immediately

upon entering the field of morphology, embroiled in controversy.

He found the field occupied by static concepts and dominated by

principles that he regarded as unsound. Accordingly, he advo-

cated the following methodology for comparative morphologists:

1. Every structure that is present in the organisms with which

we are concerned and which is pertinent to our immediate problem

must be seen and seen correctly. This implies the search for suitable

material ; tbe proper preparation and manipulation of the material ;

the possession of and the ability to use adequate optical equip-

ment and associated techniques ; the training of the eye to see and

of the mind to appreciate what is seen; and the rigorous testing

and checking of what is believed to have been seen.

2. Every structure which is seen and which can be pertinent

to the investigation in hand must be taken into consideration and

must be accounted for morphologically. No structure may be dis-

missed as having no morphological significance until its origin

and significance have been carefully explored. Nothing may be

dismissed as “merely there.”

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

3. It should be clearly recognized that comparative morphology

is not merely the comparison of the end products of evolution.

Little is to be gained by the direct comparison of highly specialized

forms. These end products must first be related to the less

specialized ancestral forms, from which alone an understanding

of them is to be derived, before they can be compared with each

other.

4. Discrepancies must be accounted for. In attempting to

account for them, the first assumption on which to proceed is that

there has been an error of observation. The next is that there has

been an error of interpretation. And if the discrepancy still per-

sists, it is not to be accepted as representing a condition that may

be dismissed. It still remains an unsolved problem that calls for

solution.

Ferris maintained that most students of morphology violated

these principles either occasionally or consistently. His opponents,

in turn, challenged his facts and conclusions. Criticism was

especially directed toward his interpretation of the insect head

(Manton, S.M. 1949, Phil. Trans. Royal Soc. London, B, Biol. Sci.

233 :483— 580) . Ferris replied in a fiery note (Microent. 15(4) .T27,

1950) as follows: “This idea of the adaptational shifting of the

mouth is one which could have been tenable in the days of

Lankester but which now is about on the intellectual level of a

belief in witchcraft. Not until such remnants of folklore have been

entirely discarded can the problems of comparative morphology

be reduced to a logical basis and a consistent system be developed.

They are like believing in ghosts without reflecting upon the source

of energy for these ghosts and their manifestations.”

The last word has not been said on the Ferris school of com-

parative morphology but a few generalizations seem to be war-

ranted. First it is clear that statements in the literature on absence

of homologies in genitalia, mouthparts, or other obviously com-

parable structures in insects are naive, unwarranted, or unsoud.

Whether Ferris, on the other hand, has provided sound interpreta-

tions on such subjects can only be determined by time and the

work of many investigators. Second, it seems clear that morpho-

logical literature contains many errors of observation and in-

terpretation. Whether Ferris, or his students have also erred is a

question for future morphologists to determine.

Meanwhile, Ferris has given us six hypotheses which he re-

January, 1959]

garded as basic to his philosophy of comparative morphology.

“1. We postulate that evolution has been continuous and orderly,

each change being based upon a stage that has gone before. To

assume otherwise, to assume that evolution has been discontinuous

and has proceeded by steps of great magnitude and unrelated to

anything that has gone before, is to surrender at the very beginning

any possibility of testing either assumption and of determining

whether or not such changes of great magnitude have actually

occurred.

“2. We postulate that all evolutionary changes have arisen from

changes in the genetic materials and are transmitted only in accord

with the laws of genetics, whether those laws are now completely

known or not.

“3. We postulate that evolutionary changes are first to be ac-

counted for by modifications of pre-existing structures ; by fusions

of pre-existing structures; or by loss of pre-existing structures.

Only after these possibilities have been exhausted will we assume

that a completely new structure has been developed. This is in

accord with the principle of parsimony ; the validity of the postu-

late has thus far been substantiated in practice.

“This is a direct reversal of procedures that have frequently

been followed by investigators, some of whom, when confronted

by a difficulty, have immediately resorted to the assumption that

even a whole new system has been evolved to replace a previous

system. This has been done time after time and is fatal to the

development of any consistent and coherent system of comparative

morphology.

“4. We postulate that evolution has not been merely endless

change involving all structures of the body. There are structures

which have remained stable while everything about them changed.

It is these structures which constitute the bases upon which the

student of comparative morphology depends for the tracing of

continuity. These are the ‘landmarks’, the importance of which is

known as a matter of experience, and search for them is one of

the primary activities of comparative morphology.

“5. We postulate the necessity for and the justification of a

faith that somewhere in the great mass of material offered to us

by the Annulata there will be an answer to any morphological

question that may arise. We shall assume that, if the answer is

not immediately forthcoming this is most likely due to the lack

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

of suitable material, a failure to interpret available facts correctly,

or the inadequacy of our own mental processes. To begin with

the assumption that the answer does not exist is to admit defeat

before the matter has been put to a real test.

“6. We postulate that it is not the business of the student of

comparative morphology to inquire into the function of a structure.

It is his business merely to trace the history of the structure. We

are not concerned with whether a leg is used for walking, or for

digging, or for leaping, or for grasping, or for swimming. We

are concerned only with the question of the morphological origin

of that leg and its parts.

“Thus armed with the general principles that have proven valid

and useful in the development of other types of scientific investiga-

tion, with a methodology, and with a set of at least preliminary

hypotheses that are subject to test, we are in a position to proceed.

We may pass on to the consideration and the pursuit of individual

problems which may offer some answer to the broad question

with which we began, that of whether or not the comparative

morphology of the Annulata can contribute something to the

understanding of the general problem of evolution.”

General Characteristics. It is interesting and perhaps useful to

consider the characteristics that make for greatness. Foremost

among these, perhaps, is motivation or drive coupled with a sense

of social responsibility. No less important for a teacher is personal

humility, sympathy and generosity to one’s fellow men. Then, in a

scientist, there is a need for technical competence in his special

field, imagination to conceive large works, patience to carry them

out and a critical attitude toward his own work and the work of

others. Gordon Floyd Ferris had all of these in generous measure.

January, 1959]

GORDON FLOYD FERRIS, THE TEACHER

Ira L. Wiggins

Stanford University, Stanford, California

Gordon Floyd Ferris possessed a rare combination of talents,

those of the highly productive research scientist, and at the same

time a flair for, and a liking of, teaching that made him an

exceedingly valuable member of three different departments during

his 42 years of service at Stanford University. He began his

career in teaching in the fall of the year in which he obtained

his A.B. degree at Stanford, serving as a laboratory assistant in

the Department of Entomology and Bionomics during the acedemic

year of 1916-1917. Those who were his superiors in that depart-

ment are long since gone, but there is little doubt that his per-

formance in the laboratories during his year of work toward the

Master’s degree was not only satisfactory but well above average,

for he was appointed to an instructorship in the Department of

Entomology and Bionomics in the autumn of 1917, and served in

that capacity until the end of the academic year of 1920—21. During

that four year period he was engaged in supervising the laboratory

work in the beginning courses in Entomology, with comparatively

little opportunity to deliver lectures, that duty being performed

by Vernon Kellogg, Professor Rene Doane and other members

in the department senior to the young Ferris.

Perhaps it was this initial term of close contact with the stu-

dents, working with first one and then another on an individual

basis as problems in the laboratory were discussed and debated,

that gave Gordon Ferris the foundation for such effective work in

dealing with his students on a totally informal, person to person

basis later in his career. During World War I and for several

years immediately afterward, classes in the biological sciences

were smaller than now, so one might expect that laboratory sections

were then smaller and handled on a more leisurely basis than they

are at present, but that assumption is not warranted. There were

times when Ferris had supervision of 30 to 35 students in a

beginning laboratory and many days found him still in the

laboratory at six in the evening, helping the slower students com-

plete their assignments. There were times when he resented the

extra effort required, the time taken from the research problems

on which he himself was engaged, but he did not shirk his duties

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

and responsibilities toward the young people in his laboratories.

He did comment pungently on the situation at times, however.

That he found time for a goodly amount of research is attested by

the list of published papers appearing during those early years

of his academic endeavors, and by the fact that he completed all

requirements for the Master of Arts degree in the spring of 1918.

Even at that time Ferris was developing some of the traits

that were to set him apart as an independent and courageous

thinker, for he disagreed with two or three other members of the

Zoology Department, with which Entomology was merged in 1921.

The present writer was first introduced to the Stanford campus

and its personnel in the autumn of 1924, three years after Gordon

Ferris had been promoted to an Assistant Professorship, but even

at that time the students in the departments of Botany and of

Zoology (then two separate departments) told their colleagues

with great glee the manner in which Ferris stuck to his own view-

points on controversial subjects and argued vigorously, and usually

successfully, with some of his colleagues.

The basis for much of his popularity with the students was

the breadth and variety of his reading. He kept up with the litera-

ture in entomology to a remarkable degree, and missed little in a

number of journals in other fields of zoology as well. This ability

to keep his ideas and his arguments up to the minute served him

well in his classes, for he quoted from the current research papers,

brought new ideas to the laboratory, and discussed many of these

papers frankly, often critcially with the students. He was, neither

then nor later, at all hesitant about approaching a particular paper

critically and objectively. If it seemed to him to be well written,

based on sound investigative work, and accurate in its conclusion,

he praised it; if he detected carelessness or unsupported statements,

he quickly pointed to the defects and many times made constructive

suggestions for overcoming the deficiencies apparent to him.

Nor was he content to teach his courses, do his laboratory

research, and read widely in scientific and non-scientific fields.

He spent weeks and months in the field, collecting insects of many

kinds, but principally scale insects and lice, for he began to

specialize in these two major groups comparatively early in his

career. Early in the 1920s he made huge collections of both lice and

scale insects in Mexico, walking many long weary miles with his

meager field equipment packed on one burro and accompanied

January, 1959]

by a timid peon often greatly in fear of his life from “bandits”

or “tigers.” After each such field trip his lectures and his dis-

cussions during laboratory sessions were enriched by his exper-

iences in such a way that the students gained a broader, deeper

interest in entomology than they would had he confined his remarks

to elaborations of the text or matter-of-fact consideration of

dissections or drawings of various types of arthropods. Often he

could associate a particular species of insect with a precise spot

where it, or a close relative, had been encountered in the field.

By bringing this type of personal touch into the classroom he

enlivened and vitalized the subject for the students, often leading

members of a class to elect entomology as a college major without

suggesting such a course verbally.

It would be a mistake to assume that because Ferris kept

members of his classes intensely interested in the subject that he

was an easy taskmaster. He required conscientious work, close

attention to detail, and a high degree of accomplishment in execut-

ing drawings or preparing insects for insertion into a collection.

He was an excellent draftsman and experimented extensively with

techniques for depicting insects accurately and yet with a minimum

of time and effort. He was among the first to admit that all students

were not equal in ability with a pencil or an inking pen, so did

not insist that each acquire excellency as a draftsman, but he did

insist that every member of his class could and should learn to

make recognizable drawings that were accurate as to outline and

scale. He was incensed by carelessness and by “fuzzy thinking.”

He wanted, and obtained, reasonably clear-cut English in examina-

tion papers and written reports if those exercises were to obtain

good grades or his support when a candidate was seeking a

position. He held high standards for his own professional achieve-

ment, and expected each student to strive diligently and constantly

for improvement — not necessarily for perfection. He was well

aware that the latter goal is rarely — if ever — attained.

He served as Assistant Professor from 1921 until 1927, and

became Associate Professor with the beginning of the academic

year of 1927-28. He remained at that grade for ten years, and was

promoted to a full professorship in 1937. During all of these years

his teaching activities claimed approximately one-half of his time,

or during some years, more than that proportion of each working

week. He taught general entomology, a course in histology and

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

microtechnique for several years, classification of insects, a spe-

cialized course on aphids, one on economic entomology with

special emphasis on Coccidae, principles of nomenclature, and

year after year supervised the advanced training of several gradu-

ate students working toward the Masters or Ph.D. degrees. He was

a cooperative member of the department, and when the depart-

ments of Botany and Zoology were merged in 1934, and together

with the course in General Biology constituted the Department of

Biological Sciences, he accepted greater responsibility for course

work in Entomology, and took over supervision of several sections

of the General Biology laboratories.

With the passing of the years his penchant for philosophical

contemplation of biology and biological activities increased and

he developed advanced courses in the principles of classification,

turned his attention to the internal anatomy of insects with the

conviction that such studies could help solve some of the problems

involved in the evolution of insects and provide stimulating teach-

ing material. When other members of the department accepted

appointments elsewhere and temporarily left the Department of

Biological Sciences short of offerings in elective courses for upper

division students, he accepted the responsibility for keeping such

offerings available. In one such situation he developed a course in

the philosophy and developments of the concepts associated with

evolution and was pleased greatly that it gained a high degree of

popularity, was greatly chagrined when his failing health neces-

sitated its termination.

When Professor Doane retired and Ferris was the sole remain-

ing entomologist in the Department of Biological Sciences, he

accepted even heavier teaching burdens by giving some courses in

alternate years, by reducing the time he could spend on his own

research, and by almost eliminating recreational activities in the

field of yachting and serving as a leader of a Sea Scout Ship. Thus,

with the flight of time his teaching activities took a larger pro-

portion of his time, his thought, and his energy, but these inroads

on his time for research and extracurricular activities prompted

very little protesting and no serious grumbling on his part. He

saw the need for courses for the training of biology majors in

certain aspects of entomology, and he filled those needs in a whole-

hearted, public-spirited manner.

Among the courses he gave after Doane’s retirement was the

January, 1959]

one Professor Doane had taught for a number of years in economic

entomology, entitled, “Insects and Man” and known among the

students as “Bugs and Bites” after Ferris assumed responsibility

for it. No doubt his personal interest in the biting and sucking lice

led him to emphasize the importance of these ectoparasites and

partially set the tenor of the course. Be that as it may, the course

was one of the more popular elective courses offered within the

Department of Biological Sciences, despite the fact that it was a

lecture course and one that required considerable reading from

sources outside the text. He organized the material presented in

such a way that the course could be taken without prerequisites,

and many students at Stanford University had little or no other

contact with college biology courses. To Ferris this placed a heavy

responsibility on his shoulders, and he insisted on giving the

course for several years after his health had begun to fail, and

when the thrice-each-week lectures exacted a tremendous toll of

his strength. Nor was this course considered a “pipe” by the

students who elected it. Rather, they recognized it as an opportunity

to become acquainted with a great teacher who, incidentally to

them, was interested primarily in insects.

In this course as well as in his advanced ones, he drew from

his rich experience while doing field work as a Guggenheim Fellow

in Mexico during 1925—26, a year spent at the Molteno Institute

in England in 1930 and 1931, while on an extended expedition

into Lower California and Panama in 1938; and late in his teach-

ing career, from nearly a year spent in China as a Fulbright Fellow-

in 1948-49. When he described the malarial menace in the tropics,

he had personal knowledge of the disease and its vectors, for he

had suffered severly from malaria after his Mexican Trip in 1926.

He had seen natives of several countries existing under the burdens

of numerous ecto- and endoparasites and gave graphic and accurate

word pictures of his observations. Apparently, Gordon Ferris saw

little in the world of insects that he was unable to utilize in his

teaching.

One of the striking characteristics of many of his lectures was

the absence of notes when he confronted his classes. His ability to

organize such a tremendous amount of information in his mind,

and present it logically, interestingly, and with a personal slant

without notes was a constant stimulus and source of admiration

among his students. Not that he was oratorical in his presentation

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

— ^he was not. He presented the subject informally, in a rather

high-pitched voice, but with such obvious personal interest in

what he had to share, that his listeners remembered what he said

long after they would have forgotten flights of rhetoric. Ferris

did, however, have the ability to coin apt phrases that caught the

attention of his audience and helped to impress them with the

worth of his offerings. Perhaps, though, the greatest advantage he

possessed was the ability to make his students or the members of

an audience feel that they were sharing his experiences and his

enthusiasm with him.

Important as his contributions to the intellectual life of the

undergraduates who took his courses in entomology or related

subjects were, his greatest service in teaching was among his own

and other faculty members’ graduate students. His technique in

handling such instruction was almost wholly that of the informal

conference. During such conferences he usually smoked incessantly,

lighting one cigarette from the butt of the previous one. The

student often had to supply the match with which to light the first

one, and not infrequently, the subsequent cigarettes. The confer-

ence might last ten minutes, or it could continue for hours. Many,

many times the discussion involving the student’s research work,

his problems in other courses, or general philosophical questions

with numerous ramifications, would be interrupted long enough

for them to drive to the Ferris home where the discussion could

continue through the evening meal and well into the night. On

such occasions the student was made to feel at ease and was given

encouragement if that seemed to be called for. He could be, and

sometimes was, subjected to rigorous verbal chastisement. But

always he was urged to think his own thoughts, to approach every

question as objectively as possible, and to keep an open, inde-

pendent mind.

Several times each day Ferris would lay aside his pencil, his

pen, or his dissecting needles, stroll into the cubicle of one of the

graduate students, chat with him for a few minutes about the work

that the student was doing or about a particular research project

in which Ferris himself was engaged. These little interchanges,

totally unscheduled and delightfully informal served to keep the

graduate students aware of Ferris’ work as well as to stimulate

them to extended efforts toward solving their own problems.

One of the results of such a practice was that many students who

January, 1959]

would otherwise have become discouraged and dropped out of the

struggle toward an advanced degree, continued on to either the

Masters or Doctors degree — and some to both. He did not pamper

his students, but he brought out their better qualities, often helped

them find unexpected capacities, and won their life-long loyalty

and respect.

Particularly during the years following World War II, can-

didates for the Ph.D. degree were encountering difficulties in

passing the language examinations. Ferris decided that much of

this difficulty stemmed from the lack of concerted, continuous effort

toward really understanding the language involved, and in lack

of practice in reading papers in a foreign tongue. To remedy the

situation he organized a seminar in the reading of scientific

German. For nearly ten years he met with any and all graduate

students interested in biology who needed practice in reading

German, but who could not arrange their schedules so as to take

regular courses offered in the Department of Germanic and

Romanic Languages. He spent from two to four hours per week

in this service. He was not obliged to carry this burden, nor did

he receive compensation for the time and effort expended. He

received great satisfaction, however, from the knowledge that short-

ly after this seminar was instituted, there was a marked increase in

the number of candidates who passed the examination in reading

German — given by another member of the Department— on the

first trial. Throughout the period of this seminar the often men-

tioned but seldom practiced scheme of running a university by

having the “student on one end of the log and the professor on

the other” was followed, for Ferris and the members of his seminar

group tackled each reading assignment together, puzzled over

German syntax and idioms, and struggled with complex German

sentences. On rare occasions they had to consult a member of the

instructional staff in German, but as time passed such consultations

became less and less frequent, for Ferris’ own command of the

language improved with the repeated seminar sessions. This

seminar, like his informal chats with candidates for advanced

degrees, gave encouraging aid to a number of students who

otherwise might have abandoned their efforts to secure training

beyond the A.B. and kept them striving until the goal was finally

attained. He can be credited personally with having applied the

stimulus and the aid necessary to carry several students through

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

to one or the other of the advanced degrees so necessary for their

professional improvement.

The full extent of Gordon Ferris’ influence on modern ento-

mology and contemporary entomologists is difficult to estimate.

Some indication of the esteem in which his colleagues held him

and his work is found in letters from workers living in many parts

of the world. Many of them asked for professional advice or for

opinions bearing on the writer’s entomological problems. Their

general spirit is that of men who recognized Ferris as an authority

in his field, one whom they respected and whose opinions they

could trust.

Another key to the scope of his influence is reflected in the

number of graduate students who worked with him toward M.A.

and Ph.D. degrees. Forty-five of his candidates successfully met

the requirements for the M.A. and at least 19 individuals earned

the Ph.D. under his supervision. The late P. N. (Perk) Annand,

for a number of years Chief of the U. S. Bureau of Entomology

and Plant Quarantine, was the first Stanford student to complete

the requirements for the Ph.D. degree under Ferris’ direction.

Among the other men and women who have obtained advanced

degrees with Ferris’ guidance, some have continued in teaching

and research in various colleges and universities, some have found

employment with state and federal agencies, a number entered

teaching positions in high schools and junior colleges, a few have

accepted positions involving applied entomolgy or basic research

with industrial concerns. Their impact on entomology and ento-

mologists continues the philosophy and spirit of independence that

Gordon Ferris so ably represented.

Still another group of scholars entered Ferris’ laboratory for

specialized or advanced training and experience. Entomologists

who had previously obtained advanced degrees came from various

parts of the United States and from abroad to work with him;

to learn at first hand his techniques in preparing and illustrating

insects; to collaborate with him in particular research problems

or to absorb the highly regarded brand of tolerance, and paradox-

ically some of the intolerances, that characterized the atmosphere

in his laboratory and in his classrooms. Ferris himself would

probably have been one of the first to disclaim any idea that he

had a “mission” to teach these visitors anything special, or even

that he felt he was teaching many of them anything at all. He

January, 1959]

just gave each of them generously of his time and an opportunity

to discuss many subjects during the course of the days spent

with him. Each became, for varying periods, a guest in his home.

To many of them the social hours there were as much of an

education as were the days spent in the laboratory and in the

field collecting specimens. Thus, Ferris came to know and respect

entomologists scattered through many parts of the world, and

they, in turn, highly regarded him. For years he kept up a terse

correspondence with scores of them, and through this corres-

pondence, liberally larded with flashes of beautiful phraseology,

continued his influence in the field of entomology.

Ferris experimented with numerous approaches and methods

in carrying on his teaching activities. Discussion in small groups

was his forte but was by no means the only teaching technique he

mastered. He assigned specific problems to individuals or to small

groups in connection with laboratory courses, or placed material

in the laboratory with general instructions to find out as much

as possible by observing the organisms provided. Such an assign-

ment did not mean neglect during several laboratory periods;

rather, frequent but brief inspections of progress were made, with

hints — rarely more than hints — ^about promising lines of attack

or sources of information. His ability to keep in mind the respec-

tive tasks assigned to different members of a class, and his facility

in picking up or temporarily dropping his own research at

these frequent interruptions permitted him to accomplish large

tasks under conditions that would have appalled many who demand

protracted periods of uninterrupted concentration in order to

get anything done. The knowledge that he was doing important

research at the same time he was conducting his classes gave him

stature in the eyes of the students, so he was held in high repute

by them, equally as a research scientist and as an outstanding

teacher.

In one of his advanced courses particularly, that in the Philos-

ophy of Biology, Gordon Ferris utilized a technique that brought

to his audience not only his own philosophy but laid behind it a

background of the thinking and writing of many others. During

part of almost every class period he would read aloud from the

writings of both modern and older biologists until he had covered

a particular point or series of ideas, then spend the time remaining

to expound his own views about the passage read. In that way

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

he presented each ‘‘lecture” with a freshness of outlook and full

familiarity with the subject that was most stimulating to the class.

A streak of impishness that came to the surface at unexpected

times was a source of amusement to many and of embarrassment

to others when they were the object of his teasing. Those with a

well developed sense of humor usually could detect the approach

of one of his sly digs or near-shocking comments, but to the overly

serious or the person who sees little of humor if he is the butt of

a joke, these incidents were always a surprise. He used the Theory

of Evolution to lampoon pomposity by referring to the compar-

atively few generations that separate men from monkeys. From

time to time he expressed skepticism about the immaculate con-

ception or other aspects of Christian theology. Such deprecating

statements were more often than not delivered when some member

of a religious order was present in his class. At first such indi-

viduals were genuinely shocked and at times outraged, but usually

more prolonged acquaintance with the man and his deep human

sympathy revealed that his chief object was not to destroy another’s

faith, but only an effort to get his students to think for themselves

and to approach questions objectively and open-mindedly.

Ferris believed that every human individual should be himself,

think his own thoughts, find his own satisfactions in such manner

as to do so without violating the dignity and rights of others, and

determine his own philosophy and mode of life. He did not en-

courage conformity for the mere sake of conforming. He did not

favor mediocrity through compliance of all members of society to

the “norm” of that society. Often he astonished students seeking

advice by hearing only part of the details of a tangled situation

and then telling the man or woman to think out what he or she

thought was the right thing to do and then go do it, regardless of

what others might think. One may consider this type of advice

dangerous, but he did not — provided one always tried to avoid

harming another individual in the exercise of his own freedom.

He advocated direct action, although not violent action. He man-

aged to bring so much of his personal philosophy and convictions

into his lectures and the discussions he held with smaller groups

that no one could justly accuse him of neglecting the humanistic

side of his science or his obligations as a citizen in his community.

Even in the discharge of his duties as a citizen he continued

to teach effectively and apparently effortlessly. For a number of

January, 1959]

years he was active with the Sea Scouts, and every Scout in his

Ship learned a great deal about navigation, the rules of the sea,

small boat handling and repairs, and other phases of Sea Scouting.

But they also learned much about human relations, the place of

the individual in the community, self-reliance, and numerous other

traits and attributes of an American citizen. At one time he had

the son of a city official and one of a Chinese restaurant proprietor

in his Ship. The offspring of the city official had a bullying streak

that did little to endear him to his associates. After being the

subject of much heckling from the other, the Chinese boy turned

on his tormentor and with a totally unexpected ferocity gave him

a thorough trouncing. Ferris had seen the direction the squabble

was taking and conveniently had an errand to perform elsewhere

just before the argument reached the physical combat stage.

Obviously, his lessons were by no means confined to the class-

room or the laboratory, and those dealing with Entomology were

only a part of his genuine contribution to the total fund of

knowledge obtained by the men and women, the boys and girls,

with whom he lived and worked. Others might advocate teaching

school children the art of getting along together, of social adjust-

ment, even if the subject matter of scholastic endeavor were

neglected, but not Ferris. He believed that sound study habits and

a solid core of fundamental factual knowledge came first, that

the social graces, if they were to be learned at all, should be

learned at places other than in the classroom. Some of the “social

graces” impressed him not one whit, although there was no more

gracious host or companion within the membership of his depart-

ment. He practiced what he preached in that he lived his own life

according to his own lights and beliefs. He tried to avoid infringing

on the rights of others while following his own modes of living,

but he reserved unto himself the right to determine what his line

of actions should be.

Gordon Ferris was so intensely aware of his obligation as a

teacher that he made personal sacrifices that seemed, to some of

his friends, unreasonable. For example, he took sabbatical leave

only three times during the 42 years he served on the teaching

and research staff of Stanford University, once to do arduous

field work in Mexico, once to spend a year at the Molteno Institute

(in 1930-3 1-) in England, and once to spend nearly a year

(1948-49) under a Fulbright Fellowship in China. Summer after

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. ]

summer, when he was under no legal or moral obligation to remain

on campus he spent practically every week day in his office-

laboratory in order to be available to graduate students trying to

complete requirements for advanced degrees. He received no

additional remuneration nor academic preferment from such extra

services, but was content to know that his students were benefltting

from his presence. He refused to accept a flattering fellowship

which would have permitted him to do field work and research

simply because he felt obligated to teach his courses.

That such generous devotion to other’s needs made deep im-

pressions on the students whom he helped and befriended has been

attested by scores of letters addressed to Mrs. Ferris, to other

graduate students, and to the writer of this paper after the news

of his death had been circulated. Typical of the eulogistic, but

wholly sincere tributes are a few that read “To me Ferry’s value

as a teacher was not so much in the field of didactic instruction

but rather as a living example. He lived in strict accordance with

his principles. He was intellectually honest. He was honest with

himself, his colleagues, and his students.” Another writes; “I have

been, and always will be, proud to be known as one of Professor

Ferris’ students.” From another: “It has been said that the influence

of a teacher never ends. In my courses in biology. Ferry will

always be the most important influence. In the day-to-day practice

of biology, things he said long ago, continue to guide me. Ferry

had a strong and infectious spirit of radicalism about scientific

‘authority’ that I hope I never let down.” And from a student

whose major interests were in a segment of biology quite removed

from entomology came this: “You must know that your loss is

shared by every student who has worked in the Museum ... I re-

garded Professor Ferris as a personal friend. He helped me many

times with problems in my work and what was much more im-

portant, he was interested in what I was doing and what I was

thinking ... I took two formal courses from Professor Ferris . . .

in which he didn’t teach us about insects, but instead taught us to

observe, to accurately record our observations and to trust nature,

not books.” Finally, another student wrote: “ . . . please remember

that the instructions he gave at the University and his spirit as

a scientist may survive at least in one of his students. After all,

Professor Ferris was the greatest teacher for me.”

January, 1959]

GORDON FLOYD FERRIS AS A STUDENT OF THE

SCALE INSECTS

Howard L. McKenzie

University of California, Davis

The passing of Gordon Floyd Ferris on May 21, 1958, brought

to a close the career of one of the most versatile Coccidologists of

our time. It is evident that the things which can be properly said

about Professor Ferris with respect to his work on scale insects

involve the man and his approach to entomology rather than any

special relationship with coccids as a group. He approached any-

thing he undertook with a singleness of purpose, and an energy

drive that was probably unmatched in any other entomologist

I have ever known, and he worked with marked efficiency on any

group which kindled his interest. In the scale insects he was pro-

ductive in describing new taxonomic units, and in developing and

publishing new terminology and new ideas about their morphology

and classification. It is generally felt that most of his work on

scale insects will stand indefinitely, though some of his proposals

on the classification will doubtless be modified as new forms are

discovered and others are studied more critcally.

Professor Ferris helped and encouraged many contemporary

and younger workers, both in this country and abroad, a few of

which include A. S. Balachowsky, G. De Lotto, W. J. Hall, Ray-

mond Mamet, V. P. Rao, J. B. Steinweden, D. J. Williams, and

the writer. In this respect he was most generous and hospitable

in receiving fellow scientists into his laboratory and home in

Palo Alto, making available his collection and library, and in

lending valuable specimens for study. His uppermost thought was

sharing his findings to advance the scientific knowledge of Coc-

coidea. His correspondence was world-wide, and he remained in

touch with Coccidologists in many countries by prompt and

generous exchange of specimens and helpful notations.

The attitudes of several foreign workers on scale insects about

Profesor Ferris are abstracted as follows from certain letters

received after his death.

D. J. Williams, London, England, May 28, 1958:

I remember his welcome help when I first took up the study of

the Coccoidea and since then he has given every assistance which,

I understand, was his way with everyone.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

His works have always been a source of encouragement and his

wit often enlivened our subject. We will, no doubt, remember that

he made life easier for us and that he has set us a standard which

we feel bound to try and follow.

I am sorry that I never met him, but to you it will be a great

personal loss.

Raymont Mamet, Mauritius Island, Indian Ocean, June 2, 1958 :

The work of Professor Ferris on the Coccoidea and other insects

will last for very many decades and we who have known him will

always remember his simple and kind manners.

A. S. Balachowsky, Paris, France, June 2, 1958:

I was extremely pained to know that Dr. Ferris died from his

heart disease May 21 last. It was a very great loss for the world’s

science. He was the most famous coccidologist of any time. He had

had a very strong and original personality and contributed to

establish coccidology on a modern level.

W. J. Hall, London, England, May 28, 1958 :

His loss will he deeply felt by entomologists the whole world

over and in particular by Coccidolo gists. No one has done more to

advance our knowledge of the Coccids than Professor Ferris, and his

work in this field will remain a tribute to his memory for many years

to come. But he also made great contributions to other aspects of the

subject, and by his passing we have lost an outstanding entomologist

of world- wide repute.

Something should be said about Professor Ferris’ ideas on

scientific illustration or, as be preferred to call it, “entomological

drafting.” He felt that the average worker should content himself

with good draftsmanship. That is, to record with severely simple

means, and with utmost accuracy, what he sees. In Professor

Ferris’ opinion, there is no excuse for sloppiness in entomological

illustration. He believed that accuracy and exquisite care are essen-

tial in entomology as in the field of mechanical design.

One of the most practical compromises he made in coccid

delineation was that of the divided drawing which shows one half

of the dorsal and one half of the ventral side of the insect. Such

drawings are possible because these insects are bilaterally sym-

metrical. Even though divided drawings were used as early as

1904 by J. G. Sanders, in his work on Coccidae of Ohio, I., we

can credit Professor Ferris with a considerable role in applying

the method, not only to the Coccoidea but also to a number of

other insect groups. He developed this technique to its present

January, 1959]

state of outstanding utility, including the precision of detail

demanded by the highest standards of work. Professor Ferris’

brilliantly executed illustrations have gained for him the un-

questioned reputation of being the world’s foremost illustrator of

Coccoidea,

The first publication of Professor Ferris on scale insects was

done in 1916, as a junior author, with Professor R. W. Doane.

It included the descriptions of three new species of Samoan scale

insects, and the excellence of Ferris’ drawings of these insects will

be noted in this first article.

In 1918, he produced his first rather extensive paper on Coc-

coidea, “The California Species of Mealybugs,” published in the

Leland Stanford Junior University Publications, University Series.

The period from 1918—1935 was marked by the publication of

a wide variety of papers on scale insects, among which were a

series of 11 articles on “Notes on Coccidae (Hemiptera)” appear-

ing in various numbers of the Canadian Entomologist. During

this period other of his outstanding papers on scale insects ap-

peared mostly in the Stanford University Publications, University

Series, Biological Sciences, and several book reviews on scale

insect works by other authors were published in certain numbers

of Canadian Entomologist and Entomological News.

During 1936, and until the time of his death. Professor Ferris

made tremendous strides in developing the taxonomy of Coccoidea.

The publication “Microentomology” was founded by him in

1935, and, as has already been pointed out by Dr. Ira Wiggins

in a Memorial Number of that same Journal (vol. 23, pt. 2, p. 71,

1958), he “edited each number, contributed many important

papers to its pages, and, more frequently than he really could

afford, gave financial support to insure its publication.”

Many issues of Microentomology were devoted to clarification

of generic types, particularly of the armored scale insects (Diaspid-

idae). In this regard. Professor Ferris records the following in

one of the earlier issues of this journal:

It is sufficient to say that the student of the Coccoidea is pre-

sented with two alternatives. On the one hand he may cling to the

admittedly inadequate system of an earlier day and on the other he

is forced to wander in a maze of generic names the application of

most of which can not be determined from the existing literature.

We are thus at the moment in something of an impasse in the study

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

of this group. But the way of escape is evident, though laborious.

The first step is to examine critically and illustrate adequately the

types of the named genera so that students may have some basis for

forming their own conclusions as to the characters which are avail-

able for the recognition of generic groups.

This he did in his inimitable manner, and these works have

been the basis for critical world-wide revisions of many of these

scale groups.

Less than a year before his death. Professor Ferris published

in Micro^ntomology, “A Brief History of the Study of the Coc-

coidea.” His comments regarding certain workers on scale insects

are quite critical, although justifiably so. The introduction ex-

presses his feelings regarding the high quality work he believed

necessary in Coccoidea taxonomic research. He comments:

The author can quite truthfully say “much of this I saw and

some of it I was.” In writing this history toward the end of his life

when there is no longer anything to be gained by undue avoidance

of objectivity or of undue consideration for the feelings of others,

the writer feels it incumbent upon him to express the truth as he

feels it to be. Such truth requires a severe judgment upon various

men who have been in the past or still are his colleagues. His

estimates of the work done by some of his predecessors, and even

by some of his colleagues is extremely unfavorable, but they are

estimates made as honestly and sincerely as possible. For that he

beg’s no one’s pardon.

Perhaps the most outstanding contribution on scale insect

taxonomy which the world has ever known, is his monumental

treatise entitled “Atlas of the Scale Insects of North America,”

The first four volumes of this Atlas, published during 1937—1942,

were devoted to a comprehensive treatment of some 245 species

of armored scale insects (Diaspididae) , all of which were mag-

nificently illustrated by Professor Ferris. Volumes V and VI of

the Atlas, published in 1950 and 1953 respectively, included the

systematics of the mealybugs (Pseudococcidae) of North America.

In these two volumes 206 species and 38 genera of mealybugs

were considered, and beautifully delineated by Professor Ferris.

Volume VII of the Atlas, published in 1955, dealt with the

Coccoidea families, Aclerdidae, Asterolecaniidae, Conchaspididae,

Dactylopiidae, and Lacciferidae. In this volume taxonomic treat-

ments for 168 coccids in 21 genera, together with keys to all, were

presented.

January, 1959]

BIBLIOGRAPHY OF GORDON F. FERRIS

Hugh B. Leech

California Academy of Sciences, San Francisco

A bibliography of the scientific papers of the late Gordon

F. Ferris appeared in Microentomology, Volume 23, Part 2,

pages 75—80; September 26, 1958. The present one adds some

titles; nearly all have been checked with the original sources.

Attention was paid to the actual dates of publication whenever

they could be verified, and some changes in the sequence of

titles have resulted.

I am indebted to Dr. Ira Wiggins and Dr. Laura Henry for

supplying me with lists of most of the papers by G. F. Ferris.

It is difficult to know how to list the numerous cases in which

Professor Ferris’ (paraphrased?) remarks at meetings of the

Pacific Coast Entomological Society appear in the published

Minutes. There is hardly a doubt that the information therein

should be attributed to him, rather than to the persons who

recorded and signed the minutes, and I am sure the recorders

would have been the first to agree to this. Since all appeared as

untitled notes in The Proceedings of the Society, they are given

below as a unit, with my explanatory comments in brackets.

It should be noted that some of the Minutes which concern

us (those of the 71st to 116th meetings) are in the two volumes

of separately published Proceedings, and cannot be dated exactly.

For details see the summary by J. W. MacSwain in the Pan-

Pacific Entomologist for 1951 (27 (3) :105— 109) . The remainder

are in the Proceedings, published annually in the Pan-Pacific

Entomologist.

It is quite possible that Professor Ferris has been similarly

reported in the minutes of other entomological societies, but

there has not been an opportunity to make the necessary search.

1915

1. (Junior author with V. L. Kellogg) Anoplura and Mallo-

phaga from Zululand. Annals of the Durban Museum,

1(2) :147-158; pis. XV-XVI. Published May 15.

2. (Junior author with V. L. Kellogg) The Anoplura and

Mallophaga of North American mammals. Leland Stanford

Junior University Publications, University Series. 74 pp.;

18 figs., pis. 1—8. Published May 25.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

1916

3. (Junior author with R. W. Doane) Notes on Samoan

Coccidae with descriptions of three new species. Bulletin

of Entomological Research 6(4) :399— 402; figs. 1—3. Pub-

lished in January.

4. Mallophaga and Anoplura from South Africa with list of

mammalian hosts of African species. Annals of the Durban

Museum, 1 (3) :230— 252; figs. 16—27. Published April 20.

5. Cervophthirius crassicornis (N.) (Anoplura). Entomologi-

cal News, 27(5) :197— 200; 1 fig. Published May 2.

6. A catalogue and host list of the Anoplura. Proceedings of

the California Academy of Sciences (Fourth Series),

6(6) :129-213. Published May 12.

7. Notes on Anoplura and Mallophaga, from mammals, with

descriptions of four new species and a new variety of

Anoplura. Psyche, 23(4) :97— 120; figs. 1—12. August num-

ber.

8. Some generic groups in the mallophagan family Meno-

ponidae. Canadian Entomologist, 48(9) ;301— 311; figs.

10—15. Published September 15.

9. Anoplura from sea-lions of the Pacific Ocean. Entomological

News, 27 (8) :366— 370; figs. 1—4. Published September 30.

10. Some ectoparasites of bats (Dipt.). Entomological News,

27(10) :433— 438; pis. XXII— XXIII. Published December 6.

1917

11. Methods for the study of mealy-bugs. Journal of Economic

Entomology, 10(3) :321— 324. Published June 19.

12. A new genus and species of Coccidae (Hemip.— Homop.) .

Canadian Entomologist, 49(11) :375— 378; figs. 36-39. Pub-

lished November 1.

1918

13. A note on the occurrence of abdominal spiracles in the

Coccidae (Hemiptera). Canadian Entomologist, 50(3) :85—

88. Published March 15.

14. An apparently new species of Leptinillus (Coleoptera,

Leptinidae). Canadian Entomologist, 50(4) : 125— 128; figs.

4^6. Published April 13.

January, 1959]

15. Notes on Coccidae (Hemiptera) . Canadian Entomologist,

50(7) :221— 225; pi. IV. Published July 10.

16. The California species of mealybugs. Leland Stanford Junior

University Publications, University Series. 78 pp.; 16 text

figs., pis. I— III. (Received at the California Academy of

Sciences Library on July 24.)

17. Notes on Coccidae II (Hemiptera). Canadian Entomologist,

50(10) :323— 332; figs. 12—16. Published October 10.

18. The alleged occurrence of a seasonal dimorphism in the

females of certain species of mealy bugs (Hemiptera;

Coccidae). Entomological News, 29(9) :349— 352. Published

November 2.

1919

19. A remarkable case of longevity in insects (Hem., Horn.),

Entomological News, 30(1)27—28. Published January 4.

20. Two species of Phylloxera from California (Hemiptera;

Aphidae). Entomological News, 30(4) :103— 105; text figs.

A— D. Published April 3.

21. A contribution to the knowledge of the Coccidae of south-

western United States. Leland Stanford Junior University

Publications, University Series. 68 pp.; figs. 1—38. (Received

at the California Academy of Sciences Library on May 17.)

22. Notes on Coccidae — HI (Hemiptera). Canadian Entomolo-

gist, 51(5) :108— 113; figs. 17—19. Published May 20.

23. Observations on some mealy-bugs (Hemiptera; Coccidae).

Journal of Economic Entomology, 12(4) :292— 299; figs.

15—17. (Received at the California Academy of Sciences

Library August 27.)

24. Lac-producing insects in the United States (Hemiptera;

Coccidae). Journal of Economic Entomology, 12(4) :330—

333. (Date of receipt as for previous item.)

25. Anoplura of the Canadian Arctic Expedition, 1913—18.

Report of the Canadian Arctic Expedition 1913—18. Vol. 3,

Part D:lld.

26. Notes on Coccidae — IV (Hemiptera). Canadian Entomolo-

gist, 51(11) :249— 253; figs. 33—36. Published November 20.

27. Why not government-maintained fellowships? Science, (N.

S.) 50 (1302) :543— 544. Published December 12.

28. A new species of Pseudodiaspis (Hemiptera; Coccidae).

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Entomological News, 30(10) :275— 276; 1 fig. Published

December 20.

1920

29. Contributions toward a monograph of the sucking lice.

Part I. Leland Stanford Junior University Publications,

University Series, Biological Sciences, 2(1) :1— 52 [p. 52 is

blank] ; figs. 1—32. Dated 1919, but actually published

January 14, 1920.

30. Clarke nutcracker at sea. Condor, 22(1) :39. Published

January 26.

31. Notes on Coccidae — V (Hemiptera) . Canadian Entomologist,

52(2)29—32; figs. 7—11. Published February 28.

32. The first stage larva of Cuterebra americana (Fabr.)

(Diptera; Oestridae). Psyche, 27(1):13— 14; fig. 1. Febru-

ary number.

33. Notes on Coccidae — VI (Hemiptera). Canadian Entomolo-

gist, 52(3) :61— 65; fig. 12. Published March 31.

34. Some records of Polyctenidae (Hemiptera). Journal of the

New York Entomological Society, 27 (4) :261— 263 ; pi.

XXIV. Published April 17.

35. Scale insects of the Santa Cruz Peninsula. Stanford Univer-

sity Publicaitons, University Series, Biological Sciences,

1(1) :1— 57 ; figs. 1—35. (Received at the California Academy

of Sciences Library on July 30.)

36. Insects of economic importance in the Cape Region of

Lower California, Mexico. Journal of Economic Entomology,

13 (6) ;463— 467. December number.

1921

37. Notes on Coccidae — VH (Hemiptera). Canadian Entomolo-

gist, 53(3) :57— 61. Published April 4.

38. Review of “Monografia delle Cocciniglie Italiane.” Ento-

mological News, 32(5) :157. Published May 3.

39. Notes on Coccidae — VHI (Hemiptera). Canadian Ento-

mologist, 53(4) :91— 95. Published June 8.

40. Some Coccidae from eastern Asia. Bulletin of Entomological

Research, 12(3) :211— 220; figs. 1—7. November number,

published in September.

41. The Coccidae of Ceylon \in re funds for publication].

January, 1959]

Science, (N.S.) 54 (1397) :330. Published October 7.

42. Contributions toward a monograph of the sucking lice.

Part II. Stanford University Publications, University Series,

Biological Sciences, 2(2) :53— 134 [pp. 58 and 134 are

blank] ; figs. 33—89. Published October 14.

43. Report upon a collection of Coccidae from Lower Cali-

fornia. Stanford University Publications, University Series,

Biological Sciences, 1(2):59— 132; figs. 1—52. (Received at

the California Academy of Sciences Library on November

18.)

44. A new species in the Hormaphidinae (Hemiptera, Aphid-

idae) . Entomological News, 32(10)289—291; pi. VI. Pub-

lished December 16.

1922

45. Concerning lice. Journal of Mammalogy, 3(1) :16— 18. Pub-

lished February 8.

46. The mallophagan family Trimenoponidae. Parasitology,

14(1) :75— 86; figs. 1—8. Published April 25.

47. (Senior author with F. R. Cole) A contribution to the

knowledge of the Hippoboscidae (Diptera, Pupipara),

Parasitology, 14(2) :178— 205; hgs. 1—20. Published June

15.

48. Review of “The Coccidae of Ceylon,” by E. E. Green.

Science, (N.S.) 56 (1446) : 3 12— 313. Published September

15.

49. Notes on Coccidae — IX (Hemiptera). Canadian Entomolo-

gist, 54(7) :156— 161; figs. 1—4. Published October 12.

50. Contributions toward a monograph of the sucking lice. Part

III. Stanford University Publications, University Series,

Biological Sciences, 2(3) :135— 138 [p. 138 is blank] ; figs.

90—118. (Received at the California Academy of Sciences

Library on October 19.)

51. A note on Timema calif ornicum Scudder (Orthoptera;

Phasmidae). Entomological News, 33 (9) :282— 283. Pub-

lished November 3.

52. De anopluris. Scientific Monthly, 15(6) : 55 1—556.

53. Two new Coccidae from Cyrenaica. Bollettino del Labora-

torio di Zoologia generale e agraria della R. Scuola superiore

d’Agricoltura in Portici, 16:207—210; figs. 1—2. This parti-

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

cular article was published on December 15.

54. Notes on Coccidae— X (Hemiptera). Canadian Entomolo-

gist, 55(11) :246— 248. Published December 30.

1923

55. Observations on the larvae of some Diptera Pupipara, with

description of a new species of Hippoboscidae. Parasitology,

15(1) :54r-58; figs. 1—4. Published March 22.

56. (Senior author with Persis Hyatt) The life history of

Euphyllura arbuti Schwarz (Hemiptera; Chermidae). Can-

adian Entomologist, 55(4) :88— 92; pi. 2. Published May 10.

57. The place of the systematist in modern biology. Scientific

Monthly, 16(5) :514— 520. May number.

58. Mallophaga. In: Biological Survey of the Pribolof Islands,

Alaska. North American Fauna, No. 46, Part H, p. 141.

Published June 20.

59. (Senior author with J. B. Kelly) Some Coccidae from about

the Gulf of California. Proceedings of the California Acad-

emy of Sciences (Fourth Series), 12(14) :315— 318; 1 fig.

Published July 10.

60. Mallophaga. In : Report of the Scientific Results of the

Norwegian Expedition to Novaya Zemlya 1921. 1(8) : 11—

12. Published in July.

61. Entomological illustrations. Science, (N.S.) 58(1501) :265—

266. Published October 5.

62. Observations on the Chermidae (^Hemiptera; Homoptera).

Part I. Canadian Entomologist, 55(11) :250— 256; pi. 12,

and text fig. 1. Published November 29.

63. Contributions toward a monograph of the sucking lice.

Part IV. Stanford University Publications, University Series,

Biological Sciences, 2(4) : 179— 270; figs. 119—172. (Received

at the California Academy of Sciences Library on January

4, 1924.)

1924

64. Two Diptera Pupipara from Philippine bats. Philippine

Journal of Science, 24(1) :73— 79; figs. 1—3. Published

January 22.

65. The mallophagan family Menoponidae. Part I. Parasitology,

16(1) :55— 66; figs. 1—5. Published January 31.

January, 1959]

66. [General comments on the immature stages of the Psyllidae.]

As quoted in the Minutes of the 87th meeting, Proceedings

of the Pacific Coast Entomological Society, 2(2) :24. Pub-

lished January 31.

67. The study of minute insects. Canadian Entomologist,

56(2) :25— 28. Published February 29.

68. Report upon a collection of insect ectoparasites from Austra-

lian and Tasmanian mammals. (Diptera Pupipara. Siphon-

aptera). American Museum Novitates. No. 110:1—7; figs.

1—5. Published April 21.

69. The New World Nycteribiidae (Diptera Pupipara). Ento-

mological News, 35(6) :191— 199; pi. 3, and 1 text fig. Pub-

lished June 4.

70. The nymphs of two species of Chermidae (Hemiptera).

Pan-Pacific Entomologist, 1(1) :24^28; figs. 1—2. Published

June 25.

71. A note on some Hippoboscidae (Diptera Pupipara). Ento-

mological News, 35(7) :234— 235. Published July 3.

72. Some Diptera Pupipara from the Philippine Islands. Philip-

pine Journal of Science, 25(4) :391— 403 [p. 402 is blank] ;

figs. 1—7. Published November 17.

1925

73. Observations on the Chermidae (Hemiptera; Homoptera) .

Part II. Canadian Entomologist, 57(2):46— 50; figs. 1—3.

Published February 28.

74. On two species of the genus Halarachne (Acarina; Gama-

sidae). Parasitology, 17 (2) :163— 167 ; figs. 1—2. Published

May 18.

75. The content of systematic biology. Scientific Monthly,

20(6) :653— 658. June number.

76. Third report upon Diptera Pupipara from the Philippine

Islands. Philippine Journal of Science, 27 (3) :413— 421 ;

figs. 1—5. Published August 3.

77. Notes on Coccidae — XI (Hemiptera) . Canadian Entomolo-

gist, 57 (9) :228— 234; figs. 1—4. Published September 26.

78. (with L. E. Myers) The generic types of the Diaspidae

(Hemiptera). Bulletin of Entomological Research, 16(2):

163—167 ; pis. 12—13. Published in October.

79. Fourth report upon Diptera Pupipara from the Philippine

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Islands. Philippine Journal of Science, 28(3) :329— 341 [p.

340 is blank] ; figs. 1—5. Published November 6.

80. Systematic problems and work [a short general statement

apparently quoted in full, in the Minutes of the 97th meet-

ing] . Proceedings of the Pacific Coast Entomological Society,

2(4) :59— 60. Published afted mid-June, 1925 (see notes

following this numbered biblography) .

1926

81. The mallophagan family Menoponidae. Part II. Parasit-

ology. 18(1) :1— 3; 1 fig. Published January 22.

82. Observations on the Chermidae (Hemiptcra: Homoptera).

Part III. Canadian Entomologist, 58(1):13— 20; figs. 1—5.

Published January 30.

83. Indians’ Hair [congregations of Phalangida on tree trunks

in Mexico.] Nature Magazine, 8(4):222; 1 fig. October

number.

84. Report upon a collection of Hippoboscidae (Diptera Pupi-

para) from Borneo. Sarawak Museum Journal, 3 (Part III,

No. 10) :279— 286; pi. 11 and text figs. 1—2. December

number.

1927

85. (Senior author with E. W. Nissen) The larva of a species

of the Cassididae (Coleoptera) . Pan-Pacific Entomologist,

3(4) : 169— 172; 1 fig. Published June 9.

86. The generic types of the Diaspidae (Hemiptera). (Intro-

ductory note to Part II) . Bulletin of Entomological Research,

17(4) :341. Published in June.

87. Hippoboscidae. British Museum (Natural History) : Insects

of Samoa, and other Samoan terrestrial Arthropoda. Part

VI, Diptera; fascicle 1, pp. 10—21; figs. 1—6. Published

July 23.

88. Mealybugs. Monthly Bulletin of the Department of Agri-

culture, State of California, 16(6) :336— 342. June number

(received at the California Academy of Sciences Library

on August 10.)

89. Some American Hippoboscidae (Diptera Pupipara). Can-

adian Entomologist, 59(10) :246— 251; figs. 1—4. Published

in October.

January, 1959]

90. Fifth report upon Diptera Pupipara from the Philippine

Islands. Philippine Journal of Science, 34(2) :207— 233 ;

figs. 1—19. Published November 9.

91. Notes on an entomological enigma. Canadian Entomologist,

59(12) :279— 281; 1 fig. Published December 11.

1928

92. The larva of Olfersia vulturis Van der Wulp. (Diptera:

Hippoboscidae) . Entomological News, 39(2):36— 37; 1 fig.

Published February 3.

93. The genus Myialges (Acarina: Sarcoptidae) . Entomological

News, 39(5) :137— 140; pi. III. Published May 3.

94. Observations on the Chermidae (Hemiptera: Homoptera).

Part IV. Canadian Entomologist, 60(5) ; 109— 117; figs. 1—4.

Published June 2.

95. The genus Brachypteromyia Williston (Diptera Pupipara;

Hippoboscidae). Pan-Pacific Entomologist, 4(3) :140— 142;

figs. 1—2. Published June 26.

96. (Senior author with J. C. Chamberlin) On the use of the

word “Chitinized.” Entomological News, 39(7) :212— 215.

Published July 2.

97. The mallophagan family Menoponidae. Part III. Parasit-

ology, 20(2) :221— 227 : figs. 7—9. (Not seen by compiler.)

98. Observations on the Chermidae (Hemiptera; Homoptera).

Part V. Canadian Entomologist, 60(10) :240— 245; figs. 1—3.

Published October 29.

99. The principles of systematic entomology. Stanford Univer-

sity Publications, University Series, Biological Sciences,

5(3) : 101— 270 [also separately paged 1—170; pp. 106, 108

and 270, or 6, 8 and 170 are blank], figs. 1—11. Published

in December.

100. Review of “A Classification of the higher groups and genera

of the coccid family Margarodidae,” by H. Morrison.

Entomological News, 39(10) :325— 327. Published Decem-

ber 17.

101. The wax-secreting organs of the Coccidae. Pan-Pacific

Entomologist, 5(2):67— 70. Published December 29.

102. Scientific exploration, a phantasy. Scientific Monthly,

27 : 537— 541. December number.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

1929

103. (Junior author with J. C. Chamberlin) On Liparocephalus

and allied genera (Coleoptera; Staphylinidae) . Pan-Pacific

Entomologist, 5(3) : 137— 143. Published April 18.

104. (Junior author with J. C. Chamberlin) On Liparocephalus

and allied genera (Coleoptera; Staphylinidae). Pan-Pacific

Entomologist, 5(4) :153— 162; figs. 1—5. (Continuation of

previous item.) Published May 31.

105. Systematic biology and the mutation theory. Quarterly Re-

view of Biology, 4(3) :389— 400. Published in September.

106. Concerning the Mediterranean fruit fly. Science, (N.S.)

70(1819) :45 1—453. Published November 8.

107. Review of “A Manual of External Parasites,” by H. E.

Ewing. Entomological News, 40( 10) :337— 341. Published

December 19.

1930

108. Observations on the genus Ornithoica (Diptera: Hippo-

boscidae). Canadian Entomologist, 61 (12) : 280— 285; figs.

1—4. The December, 1929 number, but actually published on

January 4, 1930.

109. The effectiveness of a plant quarantine. Science, (N.S.)

71(1829) :68— 69. Published January 17.

110. Some New World Hippoboscidae (Diptera Pupipara).

Canadian Entomologist, 62(3):62— 70; figs. 1—5. Published

March 31.

111. The plant quarantines once more. Science, (N.S.) 71(1850) :

606—607. Published June 13.

112. Some African Diptera Pupipara. Parasitology, 22(3) :275—

282; figs. 1—6. Published July 30.

113. The puparium of Basilia corynorhini (Eerris) (Diptera:

Nycteribiidae) . Entomological News, 41(9) :295— 297, 1 fig.

Published November 7.

114. Report upon certain ectoparasites of mammals. lii\ Report

of the Harvard-African Expedition upon the African Re-

public of Liberia and the Belgian Congo, 2:1022—1038; figs.

12—24. (Not seen by compiler.)

115. Plant quarantines run wild. New Republic, 63:335—338.

(Not seen by compiler.)

116. Sixth report upon Diptera Pupipara from the Philippine

January, 1959]

Islands. Philippine Journal of Science, 43(4)537—553; figs.

1—7. Published November 10.

1931

117. The louse of elephants. Haematomyzus elephantis Piaget

(Mallophaga; Haematomyzidae) . Parasitology, 23(1) :112—

127; text figs. 1—5, pis. 4^5. Published January 31.

1932

118. Contributions toward a monograph of the sucking lice.

Part V. Stanford University Publications, University Series,

Biological Sciences, 2(5):271— 414 [p. 414 is blank; this

item is also separately paged 1—143] ; figs. 173—251. Pub-

lished October 15.

119. New species and other records of Mallophaga from the

Marquesas. B. P. Bishop Museum, Bulletin No. 98, pp. 53—

72; hgs. 8—20. (Marquesan Insects — 1. Pacific Entomologi-

cal Survey Publication I, article 5.)

120. Ectoparasites of Marquesan rats. B. P. Bishop Museum,

Bulletin No. 98, pp. 117—127; figs. 35—39. (Pacific Ento-

mological Survey Publication I, article 12.)

121. (With F. D. Klyver) Report upon a collection of Chermidae

* (Homoptera) from New Zealand. Transactions and Pro-

ceedings of the New Zealand Institute, 63(1):34— 61; pis.

7—16. Published October 30.

122. Mallophaga from Tahiti. B. P. Bishop Museum, Bulletin

No. 113, pp. 7—12; figs. 1—4. (Society Island Insects. Pacific

Entomological Survey Publication 6, article 2.) Published

December 23.

1933

123. A new species of Polyplax (Anoplura). Parasitology,

25(1) : 127— 129; figs. 1—2. Published March 4.

124. The mallophagan genus Trichophilopterus. Parasitology,

25 (4) :468— 471 ; figs. 1—2. Published December 7.

125. Contribution toward a monograph of the sucking lice. Part

VI. Stanford University Publications, University Series,

Biological Sciences, 2(6) :415— 470 [p. 418 is blank; this

item is also separately paged 1—56] ; figs. 253—276. Pub-

lished December 15.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

1934

126. Review of “The efficacy and economic effects of plant quar-

antines in California.” Science, (N.S.) 79(2036) :13— 14.

Published January 5.

127. A summary of the sucking lice. (Anoplura.) Entomological

News, 45(3) :70— 74. Published March 9.

128. A summary of the sucking lice (Anoplura), Entomological

News, 45(4) :85— 88. (Continuation of previous item.) Pub-

lished April 10.

129. Contributions toward a monograph of the sucking lice. Part

VII. Stanford University Publications, University Series,

Biological Sciences, 2(7) :471— 526 [p. 474 is blank; this

item is also separately paged 1—56] ; figs. 277—305. Pub-

lished June 1.

130. Setae. Canadian Entomologist, 66(7) : 145— 150; 2 figs.

Published August 31.

1935

131. The prothoracic pleurites of Coleoptera. Entomological

News, 46(3) :63— 68; fig. 1. Published March 7.

132. The prothoracic pleurites of Coleoptera. Entomological

News, 46(4) ;93— 95. (Continuation of previous item.) Pub-

lished April 3.

133. Scale insects (Hemiptera: Coccoidea) from the Marquesas.

B. P. Bishop Museum, Bulletin No. 142, pp. 125—131. figs.

1—3. (Marquesas Insects — III. Pacific Entomological Sur-

vey, Publication 8, article 9.) Published May 29.

134. An apparently undescrbed mealybug (Hemiptera: Pseudo-

coccidae) from Tahiti. B. P. Bishop Museum, Bulletin No.

142, pp. 133—135; fig. 1. (Marquesas Insects — III. Pacific

Entomological Survey Publication 8, article 10.) Published

May 29.

135. Contributions toward a monograph of the sucking lice.

Part VIII. Stanford University Publications, University

Series, Biological Sciences, 2(8) :527— 634 [also separately

paged 1—108] ; text figs. 306—338, pis. I— III, plus an

Erratum slip relating to pis. I and II. Published July 2.

1936

136. Foreword [to the first issue of the new journal Micro-

January, 1959]

entomology]. Microentomology, 1(1) :1. Published Janu-

ary 27.

137. Contributions to the knowledge of the Coccoidea (Homop-

tera). Microentomology, 1(1);2— 16; figs. 1—10. Published

January 27.

138. (Senior author with Michael Doudoroff) Taxonomic prob-

lems in Lepidoptera. Entomological News, 47(5) : 124^128.

Published May 8.

139. Review of “Bibliography of Australian Entomology 1775—

1930 with bibliographical notes on authors and collectors”

by A. Musgrave. Pan-Pacific Entomologist, 12(2) :64. Pub-

lished May 15.

140. Contributions to the knowledge of the Coccoidea (Homop-

tera). 11. Microentomology, 1(2) :17— 92; figs. 11—74. Pub-

lished May 22.

141. (Senior author with G. E. Murdock) Contributions to the

knowledge of the Coccoidea (Homoptera) . IIL Micro-

entomology, 1 (4) :115— 122 ; figs. 83—85. Published Decem-

ber 9.

1937

142. Atlas of the scale insects of North America. Series 1,

Numbers 1—136. Stanford University Press. (No page

numbers, each Number to be considered as a separate publi-

cation.) Published in January.

143. Contributions to the knowledge of the Coccoidea (Homop-

tera). IV. Microentomology, 2(1) :1— 45; figs. 1—36. Pub-

lished March 24.

144. On certain words used in connection with the Coccoidea

(Homoptera). Entomological News, 48 (5) : 141— 143. Pub-

lished May 13.

145. Contributions to the knowledge of the Coccoidea (Homop-

tera). V. Microentomology, 2(2) :47— 102; figs. 37—79. Pub-

lished June 15.

146. Contributions to the knowledge of the Coccoidea (Homop-

tera). VI. Microentomology, 2(3) :103— 122; figs. 80—95.

Published November 24.

147. On nomenclatorial and other problems in the systematics

of the Coccoidea (Insecta: Homoptera). Annals and Maga-

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

zine of Natural History, (Tenth Series) 20(119) :525— 530.

November number.

1938

148. Contributions to the knowledge of the Coccoidea (Homop-

tera). VII. Microentomology, 3(2) :37— 56; figs. 17—32.

Published April 28.

149. Atlas of the scale insects of North America. Series 2, Num-

bers 137—268. Stanford University Press. (No page num-

bers, each Number to be considered as a separate publica-

tion.) Published in November.

150. Contributions to the knowledge of the Coccoidea (Homop-

tera). VIII. Microentomology, 3(3):57— 75; figs. 32—44.

Published December 10.

151. In forests of hair and feathers. Where insects live in virtual

darkness and are adapted to their environment. Nature

Magazine, 31(10) :591— 592, 1 fig. December number.

1939

152. Review of “The Genus Septobasidium. By John N. Couch.”

Entomological News, 49(10) ;297— 298. Number for Decem-

ber, 1938, but not actually published until January 11, 1939.

153. (Senior author with R. L. Usinger) The family Polyctenidae

(Hemiptera; Heteroptera) . Microentomology, 4(1) :1— 50;

figs. 1—25. Published February 21.

154. (Senior author with B. E. Rees) The morphology of Panorpa

nuptialis Gerstaecker (Mecoptera: Panorpidae). Micro-

entomology, 4(3):79— 108; figs. 36—51. Published October

17.

155. (Senior author with P. Pennebaker) The morphology of

Agulla adnixa (Hagen) (Neuroptera: Raphididae). Micro-

entomology, 4(5) : 121— 142; figs. 59—72. Published Novem-

ber 22.

156. (Junior author with B. E. Rees [and C. P. Alexander, see

following] ) The morphology of Tipula reesi Alexander

(Diptera: Tipulidae). Microentomology, 4(6) :143, 145—

178 [the last part of p. 143 and all of p. 144 are by C. P.

Alexander] ; figs. 72—91. Published December 20.

1940

157. The morphology of Plega signata (Hagen) (Neuroptera:

January, 1959]

Mantispidae) . Microentomology, 5(2) :33— 56; figs. 6—20.

Published May 15.

158. The myth of the thoracic sternites of insects. Microentomo-

logy, 5(3) :87— 90. Published June 4.

1941

159. Atlas of the scale insects of North America. Series 3, Num-

bers 269—384. Stanford University Press. (No page num-

bers, each Number to be considered as a separate publica-

tion.) Published March 27.

160. Contributions to the knowledge of the Coccoidea( Homop-

tera). IX. A forgotten genus of the family Margarodidae.

Microentomology, 6(1) :6-10; fig. 3. Published March 28.

161. Contributions to the knowledge of the Coccoidea (Homop-

tera). X. Illustration of eleven genotypes of the Diaspididae.

Microentomology, 6(1) :11— 24; figs. 4—14. Published March

28.

162. Contributions to the knowledge of the Coccoidea) Homop-

tera). XL A new genus in the Pseudococcidae (Homoptera;

Coccoidea). Microentomology, 6(1) :25— 28; figs. 15—16.

Published March 28.

163. A new species of Stomacoccus (Homoptera; Coccoidea;

Margarodidae), Microentomology, 6(1) :29— 32; figs. 17-’

18. Published March 28.

164. The genus Aspidiotus (Homoptera; Coccoidea; Diaspi-

didae). Microentomology, 6(2):33— 70; figs. 19—26. Pub-

lished November 28.

1942

165. (Senior author with E. G. Linsley and R. L. Usinger) An

American Entomological Code. (In the Minutes of the 169th

meeting, Proceedings of the Pacific Coast Entomological

Society) in\ Pan-Pacific Entomologist, 18(1) :44— 45. Pub-

lished February 19.

166. Insect morphology and evolution. Abstract of an address

for the Entomological Society of America. Proceedings of

the Thirty-Sixth Annual Meeting, in: Annals of the Ento-

mological Society of America, 35(1) :114— 116. Published

April 2.

167. Atlas of the scale insects of North America. Series 4,

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Number 385—448. Stanford University Press. (No page

numbers, each Number to be considered as a separate

publication.) Published May 28.

168. The needs of systematic entomology. Journal of Economic

Entomology, 35(5) :732— 738. October number.

169. Observations on some ectoparasitic mites (Arachnida:

Acarina: Dermanyssidae) . Microentomology, 7(3):77-83;

figs. 36—38. Published December 28.

170. Some North American, rodent-infesting lice (Insecta: Ano-

plura). Microentomology, 7(3):84-90; figs. 39—42. Pub-

lished December 28.

1943

171. Some fundamental concepts in insect morphology. Micro-

entomology, 8(1) :2— 7. Published May 6.

172. The basic materials of the insect cranium. Microentomology,

8(1) :8— 24; figs. 1—6. Published May 6.

173. Additions to the knowledge of the Diaspididae (Homoptera:

Coccoidea). Microentomology, 8(2)58—79; figs. 24^32.

Published September 22.

174. The genus Targionia Signoret and some of its allies (Homop-

tera: Coccoidea: Diaspididae). Microentomology, 8(3) :81—

111; figs. 33—43. Published December 30.

1944

175. On certain evolutionary tendencies in the heads of insects.

Microentomology, 9(2) :78— 84; fig. 40. Published December

176. (Senior author with R. L. Usinger) Notes and descriptions

of American Polyctenidae (Hemiptera). Pan-Pacific Ento-

mologist, 21 (4) :121— 124. Published November 26.

1946

177. Information concerning the genera Chortinaspis and

Aspidiotus (Homoptera: Coccoidea: Diaspididae). Micro-

entomology, 11(1) :37— 49; figs. 16—20. Published April 30.

1947

178. (Senior author with V. P. Rao) The genus Pinnaspis

January, 1959]

Cockerell (Homoptera; Coccoidea: Diaspididae) . Microento-

mology, 12(2) :25— 58; figs. 10—12. Published October 22.

179. The contradictions of the insect head. Microentomology,

12(3) :59— 64. Published November 10.

1948

180. The principles of comparative morphology. Microentomo-

logy, 13(3) :49— 56. Published September 15.

1949

181. (Senior author with L. M. Henry) The nervous system and

a problem of homology in certain Crustacea (Crustacea:

Copepoda: Caligidae). Microentomology, 14(4) :113— 118;

figs. 66—67. Published December 20.

1950

182. Report upon scale insects collected in China (Homoptera:

Coccoidea). Part I. Microentomology, 15(1) :1— 34; figs. 1—

20. Published February 21.

183. Report upon scale insects collected in China (Homoptera:

Coccoidea). Part H. Microentomology, 15(3):69— 97; figs.

30—47. Published July 27.

184. A reply to criticism. Microentomology, 15(4) :126— 128.

Published October 20.

185. Atlas of the scale insects of North America. Series 5, The

Pseudococcidae. Part I. Stanford University Press, vii -|-

278 pp.; figs. 1—108. (Not seen by compiler.)

186. External morphology of the adult. Chapter 5, pp. 368—

419 in: Demerec’s “Biology of Drosophila.” John Wiley

and Sons, Inc., New York. (Not seen by compiler.)

1951

187. The sucking lice. Memoirs of the Pacific Coast Entomologi-

cal Society, San Francisco. Memoir 1: x 320 pp.; figs.

1—124. Published October 19.

1952

188. Some miscellaneous Coccoidea (Insecta: Homoptera).

Microentomology, 17(1) :2— 5; figs. 1—2. Published Febru-

ary 8.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

189. Report upon scale insects collected in China (Homoptera;

Coccoidea). III. Microentomology, 17(1) :6— 16; figs. 3—9.

Published February 8.

190. (Junior author with V. P. Rao) The genus Andaspis Mac-

Gillivray (Insecta: Homoptera: Coccoidea). Microentomo-

logy, 17(1):17— 32; figs. 10—19. Published February 8.

1953

191. On the comparative morphology of the Annulata. A sum-

ming up. Microentomology, 18(1) :2— 15. Published Janu-

ary 1.

192. (Senior author with R. L. Usinger) A new species of

Aphraniola from Cambodia [Hemiptera, Cimicidae] . Revue

frangaise d’Entomologie, 20(2) : 138— 139. Published July 25.

193. Illustrations of three species of sucking lice, with notes on

a fourth species (Insecta: Anoplura). Microentomology,

18(2):52— 57; figs. 19—22. Published August 31.

194. Report upon scale insects collected in China (Homoptera:

Coccoidea). Part IV. Microentomology, 18(3) :59— 84; figs.

23—39. Published September 22.

195. Atlas of the scale insects of North America. The Pseudo-

coccidae. Part H. Volume VI. Stanford University Press,

vii -|- pp. 279—506; figs. 109—193. (Not seen by compiler.)

1954

196. A new species of Anoplura. Annals of the Natal Museum,

13(1) :91— 94; 1 text fig. Published June 30.

197. New species of Diaspididae from Florida and the Caribbean

Islands (Homoptera: Coccoidea). Microentomology, 19(2) :

41—50; figs. 26—31. Published July 15.

198. Report upon scale insects collected in China (Homoptera:

Coccoidea). Part V. Microentomology, 19(2) :51— 66; figs.

32—42. Published July 15.

1955

199. Atlas of the scale insects of North America. Volume VII.

Stanford Univeristy Press, iii 233 pp. ; 94 figs. (Exact

date not known to compiler.)

200. On some genera of the Pseudococcidae (Homoptera: Coc-

coidea). Microentomology, 20(1) :1— 19; figs. 1—13. Pub-

lished January 28.

January, 1959]

201. Some miscellaneous Coccoidea (Insecta: Homoptera).

Microentomology, 20(2) :21— 29; figs. 14—17. Published

April 15.

202. Report upon a collection of scale insects from China. VI.

(Insecta: Homoptera). Microentomology, 20(2) :30— 40;

figs. 18—24. Published April 15.

203. The genus Phenacaspis Cooley and Cockerell. Part I. (In-

secta: Homoptera: Coccoidea). Microentomology, 20(3):

41—82; figs. 25—52. Published August 1.

204. The contribution of natural history to human progress. Pp.

75—87 in: A Century of Progress in the Natural Sciences,

1853—1953. Published in Celebration of the Centennial of

The California Academy of Sciences. California Academy of

Sciences, San Francisco. Published October 20.

205. Some minute insects: Anoplura, Mallophaga and the scale

insects. Pp. 517—523 in: A Century of Progress in the

Natural Sciences, 1853—1953. Published in Celebration of

the Centennial of The California Academy of Sciences. Cali-

fornia Academy of Sciences, San Francisco. Published

October 20.

1956

206. The genus Phenacaspis Cooley and Cockerell. Part H.

(Insecta: Homoptera: Coccoidea). Microentomology,

21(2) :67— 83; figs. 35—43. Published April 13.

1957

207. (Senior author with R. L. Usinger) Notes on and descrip-

tions of Cimicidae (Hemiptera). Part I. Microentomology,

22(1) :1— 37; figs. 1—25. Published April 15.

208. Two species of Diaspididae new to the Hawaiian fauna

(Homoptera: Coccoidea). Proceedings of the Hawaiian

Entomological Society, 16(2) :212— 215; figs. 1-2. Published

May 13.

209. (Senior author with R. L. Usinger) Hemiptera: Heterop-

tera: Cimicidae. South African Animal Life, 4:374—376;

1 fig. (Exact publication date not known to compiler.)

210. A brief history of the study of the Coccoidea. Microento-

mology, 22(2) :39— 57 ; fig. 26. Published July 1.

211. Notes on some little known genera of the Coccoidea (Hcmop-

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

tera). Microentomology, 22(3) :59— 79; figs. 27—38. Pub-

lished September 4.

212. A review of the family Eriococcidae (Insecta: Coccoidea).

Microentomology, 22(4) :81— 89; figs. 39—43. Published

October 15.

1958

213. The nervous system of insects. Proceedings of the Tenth

International Congress of Entomology, Montreal, August

17—25, 1956. 1:469—472. Published in December.

214. The geographic distribution of the Coccoidea. [A short

abstract only.] Proceedings of the Tenth International Con-

gress of Entomology, Montreal, August 17—25, 1956. 1:829.

Published in December.

1959

215. (Senior author with R. L. Usinger) A new species of

Stricticimex from Kenya (Hemiptera: Cimicidae). Annals

of the Entomological Society of America, 52(1) : 8 1—82; figs.

1—2. Published in February.

216. (With R. L. Usinger) Heteroptera: Cimicidae. In: Insects

of Micronesia. B. P. Bishop Museum. [In press.]

217. Sections Insecta, Mallophaga and Anoplura. In'. Encyclo-

pedia of Science and Technology. McGraw-Hill Book Co.,

Inc. [In press.]

Untitled Notes Presented in the Proceedings of the

Pacific Coast Entomological Society

A. Appearing in the two separately published volumes of

The Proceedings of the Pacific Coast Entomological

Society and as reported in the Minutes of the meetings.

1919. [Reference to fleas on mountain beaver, and a polyctenid

on bats] Proceedings of the Pacific Coast Entomological

Society, 1 : unnumbered page carrying the Minutes of

the 7Tst meeting.

1920. [Comment on the scarcity of Lepidoptera and Odonata

in Lower California during the summer of 1919.] Loc. cit.,

1 : unnumbered page carrying the first part of the Minutes

of the 74th meeting.

1922. [Very general note on the sucking lice.] Minutes of the

January, 1959]

83rd meeting. Loc. cit., 2(1) :7. Published November 15.

1927? [Note on some California scale insects.] Minutes of the

106th meeting. Loc. cit., 2(6) :93. Published between late

June, 1927 and the beginning of March, 1928.

1927? [Report on Mr. Meyers’ studies on the parasitism of

swallows by a species of Cimex.] Minutes of the 107th

meeting. Loc. cit., 2(7) :97. Published between late June,

1927 and the beginning of March, 1928.

1928? [On the nymphs of the Psyllidae, and on the then-

unidentified larva of the beetle Brachypsectra ; see also

loc. cit. p. 105.] Minutes of the 112th meeting. Loc. cit.,

2(7) :109. Published between the beginning of May, 1928,

and early February, 1929.

1929? [On the staphylinid beetle genus Liparocephalus.] Min-

utes of the 116th meeting. Loc. cit., 2(8) :121— 122. Pub-

lished between late April, 1929, and late January, 1930.

1929? [On the larva of Brachypsectra.] Minutes of the 116th

meeting. Loc. cit., 2(8) :122. Published between late April,

1929, and late January, 1930.

B. Appearing in The Proceedings of the Pacific Coast Ento-

mological Society, as published in The Pan-Pacific

Entomologist, and as reported in the Minutes of the

meetings.

1936. [General comments on the sucking lice.] Minutes of the

129th meeting. Proceedings of the Pacific Coast Ento-

mological Society, in: The Pan-Pacific Entomologist,

11(4): 186. This, the October, 1935 issue of the journal

was published on January 10, 1936.

1938. [Announcing the preparation of an atlas of scale insects.]

Minutes of the 145th meeting. Loc. cit., 14(4) :192. Pub-

lished November 15, 1938.

1939. [Comment on the stability of species, with a definition.]

Minutes of the 147th meeting. Loc. cit., 15(1) :46. Pub-

lished February 10, 1939.

1940. [General remarks on a trip to Panama.] Minutes of the

153rd meeting. Loc. cit., 16(1) :39. Published February

17, 1940.

1940. [On the principles of geographic distribution.] Minutes

of the 155th meeting. Loc. cit., 16(1) :42,

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

1940. [General note.] Minutes of the 158th meeting. Loc. cit.,

16(1) :45.

1941. [General remarks.] Minutes of the 162ncl meeting. Loc.

cit., 17(1) :42. Published February 21, 1941.

1942. [Comment on ectoparasites.] Minutes of the 166th meet-

ing. Loc. cit., 18(1) :40— 41. Published February 19, 1942.

1943. [Movement for an American Commission on Scientific

Nomenclature in Entomology.] Minutes of the 173rd meet-

ing. Loc. cit., 19(1) :37. Published March 22, 1943.

1943. [More on the American Commission (see preceding

item) ; comment on his Atlas of Scale Insects.] Minutes

of the 175th meeting. Loc. cit., 19(1) :38, 39.

1946. [General comments on the mealybugs of North America.]

Minutes of the 187th meeting. Loc. cit., 22(1) :36— 37.

Published March 29, 1946.

1946. [Labor-saving devices in entomological drawing.] Min-

utes of the 188th meeting. Loc. cit., 22(1) ;39.

1948. [Resume of an address on “The Mealybugs of North

America.”] Minutes of the 193rd meeting. Loc. cit.,

24(1) :39— 40. Published March 31, 1948.

1949. [A comment on Dr. Henry’s work on the nervous system

of the lower invertebrates, as relating to the hypopharynx

of insects.] Minutes of the 199th meeting. Loc. cit.,

25(1) :42. Published March 17, 1949.

1953. [Comment on the principles of comparative morphology

of arthropods.] Minutes of the 223rd meeting. Loc. cit.,

29(1) :64. Published March 23, 1953.

1957. [Resume of a talk on the family Margarodidae.] Minutes

of the 247th meeting. Loc. cit., 33(1) :47. Published

March 22, 1957.

1957. [Comment on the 10th International Congress of Ento-

mology.] Minutes of the 250th meeting. Loc. cit.,

33(1) :49.

January, 1959]

PACIFIC COAST ENT. SOC.

PACIFIC COAST ENTOMOLOGICAL SOCIETY

R. L. Doutt D. D. Jensen D. P. Flrman

Vice-President President Secretary

Gordon Floyd Ferris (1893 - 1958)

Proceedings

Two Hundred and Fifty-seventh Meeting

The two hundred and fifty-seventh meeting of the Pacihc Coast

Entomological Society was held February 14, 1958, at 7:40 p.m. in the

Morrison Auditorium, California Academy of Sciences, San Francisco, Cali-

fornia. President D. D. Jensen presided at the meeting.

The following members were present: A. E. Michelbacher, E. 0. Essig,

P. H. Arnaud, Jr., G. F. Ferris, L. M. Henry, R. L. Doutt, K. P. Shea, D. H.

Groves, S. W. Hitchcock, M. S. Wasljauer, J. A. Powell, E. L. Kessel, F. E.

Skinner, B. Barichievich, D. Breedlove, M. Kenny, J. G. Edwards, C. J.

Worthington, A. Ross, P. F. Torchio, D. J. Burdick, P. D. Ashlock, D. D.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Linsdale, L. A. Ruud, T. S. Acker, D. M. Maddox, W. E. Ferguson, Jane

MacSwain, J. W. MacSwain, E, G. Linsley, W. W. Middlekauff, R. L.

Langston, W. A. Russell, D. G. Denning, K. S. Hagen, D. D. Jensen, D. P.

Furman, H. B. Leech. Visitors were Martha Michelbacher, Marie Essig, F. D.

Bennett, S. M. Sager, Frances Powell, Carl Kaufeldt, Phil Capitate, William

Tuft, Beverly Ehreth, Bertha B. Kessel, Terry Seeno, Dolores Damiano,

Virginia Ashlock, Barry E. Pullen, Walter E. Rathgen, Nancy MacSwain,

John MacSwain, Juanita Linsley, Katherine Furman, Phillis Middlekauff,

Evelyn Langston, Elsa P. Russell, Loretta Denning.

The minutes of the meeting held December 7, 1957 were read and

approved.

The President announced appointment of the following members to

serve as a committee to judge the merits of the entomological exhibits at

the annual science fair: Kenneth Hagen, Chairman; Laura Henry, Otto

Graph.

Dr. Hagen commented briefly on the nature and mechanics of the award.

The first place award consists of an entomological book and a one year

honorary membership in the Society. The Society will send letters of com-

mendation to the recipients of the first three places in the judging. The

exhibits need not be of the systematic collection type to compete for an award.

The President announced the following changes in membership of

standing committees :

Historical — E. 0. Essig, Chairman, E. S'. Ross, R. L. Usinger, J. W.

Tilden, H. B. Leech.

Editorial Board — P. D. Hurd, Jr., Editor; J. A. Powell, H. B. Leech,

E. G. Linsley, E. S. Ross, R. L. Usinger, J. E. Swift,

R. C. Miller.

Membership — Herbert Ruckes, Jr., Chairman; J. Gordon Edwards,

Howard L. McKenzie.

The President discussed briefly types of membership open in the Society

with particular emphasis on the student membership category which enables

students to enjoy membership at a reduced cost.

Dr. Edwards nominated the following for regular membership: Mr.

Johnson C. Montgomery, Dr. Leland R. Brown, Miss Dolores Damiano, Mr.

Keith Radford. They were elected by unanimous vote.

The President announced the reinstatement of C. Don MacNeill as a

regular member of the Society.

Dr. K. S. Hagen exhibited equipment devised for airplane collections of

of Hippodamia convergens Guerin. The device is to be used in checking the

hypothesis that migration flights of the beetles from their concentration sites

in transition zone areas are made in the upper air where easterly winds blow

them westward. Previously it was believed that dispersion flights followed

down canyons and into the valley. Earlier attempts to follow flights of marked

beetles have been unsuccessful. The present airplane trap, which looks

superficially like a wing tip fuel lank, has ten discs which may be dropped

individually to an exposed position at will. A screen on the disc is supplied

with an adhesive substance containing rosin and castor oil; any beetles

contacting the disc are expected to adhere to it.

January, 1959]

PACIFIC COAST ENT. SOC.

Mr. H. B. Leech discussed the current status of historical files of the

Society. Authorization has been granted to obtain another filing cabinet to

relieve the congestion in the single file now available for such material. It

will be maintained in the Entomological Museum of the California Academy

of Sciences, Mr. Leech read excerpts from letters and field notes of early

entomologists demonstrating the value of many of the documents on file.

Drawings, published and unpublished, are included in the material. Photo-

graphs of entomologists and other “curiosities” were placed on exhibit.

Dr. Usinger commented on the data available in the historical files of

the academy. He suggested that information on the existence of this material

be published in some detail in older to increase its usefulness to interested

persons.

Dr. R. L. Usinger reported on interesting habits of reduviid bugs known

as resin bugs in Thailand. Kodachrome slides were shown to illustrate

Dipterocarpus tree-holes with bugs waiting in position near the edge of the

resin pools.

Jerry A. Powell showed a vial containing the remains of a caterpillar

of a gelechiid moth, Pseudochelaria manzanitae Keifer, and some nematodes

of the family Mermithidae which had destroyed it. Apparently Mermithids

have never been recorded from any North American Gelechiidae. The habitat,

manzanita bushes on the very dry hillsides west of Alpine Lake, Marin

County, seems to present an interesting problem. Presumably the life-cycle

of the worm includes a free living stage in the soil and a certain amount of

moisture is necessary to facilitate the parasite’s mobility in leaving the host

and entering the soil. However, the larvae, which were collected June 6, 1957,

normally pupate and transform by late June or early July. Therefore at a

locality where one would not expect any June rain, the nematodes apparently

must depend upon dew or some other unexplained source of moisture for

their return to the coarse, well-drained soil. Six larvae were found infested

among 40 collected at random from terminals three to four feet above the

ground.

H. B. Leech reported as follows;

“On February 10, 1957, several greenish bees were noticed flying around

just outside my kitchen window, in Mill Valley, California. They settled

on a galvanized iron pipe to sun themselves, and stayed so persistently within

reach that finally one was collected. It was not the common Osmia lignaria

Say, but another species of the typical subgenus. In Sandhouse’s revision

of 1939 it keyed to 0. ribifloris Cockerell, of which the California Academy

of Sciences has only a few from California (Mokelumne Hill, Calaveras

County, ‘Feb.’, ‘Mar.’, F. E. Blaisdell collector; and Altadena, Los Angeles

County, also in February, C. D. Michener collector), and none from the San

Francisco Bay region. Sandhouse gives merely New Mexico, Arizona, Cali-

fornia, Oregon, and does not comment on the date of emergence.

“On February 12, 1958, my son Bill collected another male, within six

feet of where mine was taken the year before. Examination shows the species

to be nesting in the cracks between cedar shingles, on the south wall of

the house.” . . .

Mr. Leech also exhibited illustrations of the national butterfly of Japan,

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Sasakia charonda Hewitson, together with Japanese stamps picturing the

butterfly.

Dr. Kessel introduced a guest, Mrs. Beverly Ehreth, who is a naturalist

and teacher at Novato, California, as well as director of the entomological

society of the 4-H Club in her area. Dr. Kessel commented on Mrs. Ehreth’s

recent collection of two Jerusalem crickets which seemed abnormally slug-

gish. They proved to be infected with horsehair worms, class Gordiacia. Five

specimens were released from one insect when it was dropped in water.

Dr. R. C. Miller presented an address on the subject of the Ninth

Pacific Science Congress held at Bangkok, Thailand, from November 18

to December 9, 1957.

The first Pacific Science Congress was held in 1920 at Honolulu, where

it was organized by the director of the Bishop Museum, H. E. Gregory. Since

that time the congresses have been held about every four years. Eight

hundred and fifty persons attended the ninth Congress, many more than

anticipated. Numerous sections were held to cover fields and problems of

interest to the people in and around the Pacific area. Dr. Miller noted a

resolution passed at the Congress which urged establishment of a National

Muesum of Science in Thailand. Action is expected within a couple years.

Numerous interesting color slides were shown of scenes and events covered

while attending the Congress.

The meeting was adjourned to the “Coffee Social” at 9:10 p.m. — Deane

P. Furman, Secretary.

Two Hundred and Fifty-eighth Meeting

The two hundred and fifty-eighth meeting of the Pacific Coast Ento-

mological Society was held March 15, 1958, in the Morrison Auditorium,

California Academy of Sciences, San Francisco, California. President D. D.

Jensen called the meeting to order at 2:05 p.m.

The following members were present: H. B. Leech, G. F. Ferris, L. M,

Henry, P. H. Arnaud, Jr., S. W. Hitchcock, D. D. Linsdale, J. A. Powell,

J. A. Chemsak, D. Breedlove, H. Ruckes, Jr., D. Burdick W. W. Middlekauff,

D. MacNeill, R. L. Langston, W. A. Doolin, J. G. Edwards, L. A. Ruud, Jr.,

W. E. Ferguson, D. M. Maddox, K. Innes, F. E. Skinner, E. L. Kessel, D. D.

Jensen, D. P. Furman. Visitors were: Bruce Hudson, Ben Feingold, Thomas

Leech, Barry E. Pullen, Phyllis Middlekauff, Grace MacNeill, Nora Mac-

Neill, Daren MacNeill, Margaret M. Hanna, Karen Engelhart, Nancy Doolin,

Peter Westigard, S. Smith.

The minutes of the meeting held February 14, 1958 were read and

approved.

Dr. Ruckes nominated Dr. Dennis Hynes of California State Polytechnic

College and Dr. Leslie Smith, of the University of California at Davis, for

regular membership. He nominated S. M. Sager of the University of Cali-

fornia for student membership. The candidates were elected unanimously to

the membership categories for which nominated.

The President announced that of the two remaining meetings scheduled

for the spring months one is to be held at San Jose State College and the

other is to be a field day. He announced the following committee to decide

January, 1959]

PACIFIC COAST ENT. SOC.

on a date and arrange for a place for the annual field day meeting: Hugh

Leech, Chairman; Herbert Ruckes Jr. and Laura Henry.

Program Chairman Frank Skinner announced April 26 as the probable

date for the meeting at San Jose State College. Dr. Tilden and Dr. Edwards

have arranged the program for the day.

Dr. Ed Kessel described highlights of his last summer’s trip to Alaska,

illustrated by a series of beautiful color slides.

Bruce Hudson presented a lecture on the mass culture of fleas for

production of flea-bite antigen. He is working on the project as an entomolo-

gist with Kaiser Foundation Hospital, under a grant from the U. S. Public

Health Service. Dr. Ben Feingold, director of the project was also present.

Dr. Hudson mentioned the early history of flea nuisance in the San

Francisco area, known as the “Puebla de las Pulgas” of the Portola Expedi-

tion. He presented color slides of immediate and delayed types of reactions

in man following flea bites.

The cat flea, Ctenocephalides felis (Bouche), appears to be the most

common flea annoying man in the bay area, but the human flea, Pulex

irritans Linnaeus causes the most trouble by its bite.

Methods were described by which 250—500 thousand cat fleas are reared

per week at the laboratory.

An interesting by-product of the investigation has been the finding that

what at first appeared to be morphologically identical strains of Pulex irritans

contain at least two, and perhaps three distinct species. These differ physio-

logically, among other ways, in adaptation to different hosts. The San Fran-

cisco form is the typical Pulex irritans, well adapted to man. It is the only

one of the complex studied which has been maintained successfully on

human blood.

Following a brief discussion the meeting was adjourned to the “Coffee

Social” at 3:50 p.m. — Deane P. Furman, Secretary.

Two Hundred and Fifty-ninth Meeting

The two hundred and fifty-ninth meeting of the Pacific Coast Entomologi-

cal Society was held April 26, 1958, in the Science Building, San Jose State

College, San Jose, California. President D. D. Jensen called the meeting

to order at 2:00 p.m.

The following members were present: D. D. Jensen, W. E. Ferguson,

R. F. Schoeppner, W. H. Lange, F. R. Cole, J. W. Tilden, P. H. Arnaud, Jr.,

H. B. Leech, K. S. Hagen, P. S. Bartholomew, R. P. Allen, E. G. Linsley,

H. Ruckes, Jr., D. J. Burdick, D. D. Linsdale, K. Radford, R. C. Whitney,

R. L. Doutt, K. P. Shea, F. E. Skinner C. J. Worthington, W. W. Middlekauff,

J. G. Edwards, D. Damiano, D. Breedlove, T. H. Lauret, D. P. Furman.

Visitors were: J. P. Figg-Hoblyn, Pauline L. McMasters, Stephanie S.

Ferguson, Douglas W. Price, James M. Code, Fred J. Santana, Vernon W.

Nellis, Carl H. Lindroth, John Harville, David Bartholomew, May S. Drei,

Joan Gerdts, Perry Allen, Peter Westigard, Carol Radford, A. Sanchez,

Myrnadel Guzman, Wesley L. Bond, M. Susan Gardner, Lorance W. Har-

wood, June Haslett, Julia Hoagland, Robert E. Dolphin, S. Smith, Fred

Bennett, L. E. Caltogirone, Robert Rolling, Robert Fox, Roger Lasch, Ronald

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

S. Daniel, Albert E. Crane, R. N. Ito, David Huntzinger, Terry Seeno, Leta

Rae Lauret, Homer H. Eastman, Robert V. Gauthier.

Dr. Ellis, Acting Chairman of the Department of Biological Sciences of

San Jose State College, read a welcoming message from Dr. C. D. Duncan,

who was absent due to illness.

The minutes of the meeting held March 15, 1958 were read and approved

following correction.

Dr. J. G. Edwards nominated Mr. Terry Seeno of San Jose State College

for a membership in the Society. Mr. Seeno was elected to regular member-

ship by unanimous vote.

The President announced as winner of the best entomological collection

of the Fifth Annual Bay Area Science Fair, Charles Cushner of Lincoln

High School, San Francisco. His project: Coleopteran survey. His teacher:

W. F. Monahan. Two other collections chosen for honorable mention were

those of :

1. David Cavagnaro of Tamalpais High School, Mill Valley. His project:

Preserving insect larvae. His teacher: Katherine M. Flanagan.

2. Bradford S'. Perry of Vallejo Senior High School, Vallejo. His pro-

ject: Lepidoptera of California. His teacher: C. G. Aldrich.

Dr. Kenneth Hagen and his committee were commended by the President

for their efforts in judging the entomological exhibits at the Science Fair.

Hugh Leech announced that the annual field trip of the Society would

he held May 24 at Russelman Park, Contra Costa County.

In response to the President’s call for notes and exhibits Vernon W.

Nellis exhibited specimens of Cerambycidae, Necydalis barbarae Rivers and

Necydalis cavipennis LeConte, both species of which were collected at his

home in San Carlos, California. The collection area is near the hills with

nearby chaparral, eucalyptus, bay, and oak trees as well as many other plants.

Four specimens of the uncommon species, N. barbarae, were found in mid-

August of different years. The feeding place is unknown. The specimens

are almost times as large as the ones described in Dr. Linsley’s key to

the tribe Necydalini.

David Huntzinger, of San Jose State College, exhibited a wingless

tipulid collected from a snow bank at Crater Lake Park, Oregon.

Dr. J. G. Edwards discussed the collection of insects above timberline,

illustrating his account with a series of beautiful color slides.

Mrs. Pauline McMasters described her work and conclusions on the

question of segmentation in the head of the polychaete annelid. Nereis.

Dr. John Harville discussed work of the San Jose State College Field

Station, with particular reference to studies on food habits of trout in Cali-

fornia mountain lakes. An interesting series of color slides enhanced the

presentation.

Dr. J. W. Tilden talked about the alpine butterfly associations at Tioga

Pass, Yosemite National Park. He stressed three main points as follows:

Firstly, there are several plant associations in the vicinity of Tioga Pass

and each has its own characteristic butterflies. Some associations have only

one characteristic species, others have several. One aspect of the study he

January, 1959]

PACIFIC COAST ENT. SOC.

hopes to make, is to map more completely the components, both plant and

butterfly, of each association. As an example, the morraines near the pass

are covered with white-bark pines and with several herbaceous plants

characteristic of these areas. Icaricia shasta Edw., Thoryhes nevada Scud.,

Euphydryas nuhigena Behr and Lycaena cupreus Edw. are butterflies typi-

cally found there. Of course they also may be found higher in similar areas.

But let one walk a few feet down into the meadows, and one finds Plebeius

saepiolus Bdv., Colias behrii Edw., and other species not found in numbers

on the morraines. Dr. Tilden interprets this to be due to the drainage and

soil conditions, favoring the food plants of the respective butterflies.

Secondly, there is a definite succession of species during the summer.

Drier areas warm up sooner. The meadows even when not excessively

elevated are colder, and butterflies appear there later than they do on the

drier areas. This is evident even when the dry areas are much more elevated.

So we get the interesting phenomenon of elevational reversal. Butterflies are

flying around the peaks before they appear in the meadows below. Each

species seems to have a definite season of flight earlier or later, just as at

lower elevation. This is in contrast to the usual concept of the uninitiated,

who might suppose that everything comes out at the same time at higher

elevations. Data as to how this succession takes place is one aim of the study.

Thirdly, the time of emergence of a given species is evidently different

in different years, depending on the lime the season opens up. This seems to

be due to amount of snowfall, as well as temperatures leading to melting of

the snow. The extent to which emergence varies in different years could be

correlated with known weather for the year if sufficient data were obtained.

He added that weather data if given as daily temperatuie and wind

velocity, might be used to show that free flight of these alpine butterflies

depends on two factors: direct sunlight and wind velocity. Observation will

tell you that these butterflies will cease flight when a cloud passes over the

sun. Likewise they cease flight at high wind velocities. Because of the low

ground temperatures, they seem more sensitive to these factors than, for

instance, butterflies under desert conditions, where ground temperatures are

high.

In commenting upon Dr. Tilden’s talk, Dr. E. G. Linsley stated that he

and Dr. J. W. MacSwain had observed the emergence of Vanessa cardui L.

over several hundred square miles of the Mojave Desert which appeared to

be the major source of the butterflies in the widespread flights in the

Central Valley during the second week of April. Apparently the extensive

fields of annual plants which appeared in response to suitable late autumn

rainfall provided unusually favorable conditions for the build up of Vanessa

populations. He stated that Boraginaceae were particularly subject to attack

by larvae and that in many sections of the desert nearly all of the Cryptantha

was defoliated. Amsinckia, Lupinus, and Sphaeralcea were also heavily

attacked in local areas following defoliation of Cryptantha. In some places

it was difficult to walk without crushing pupae. Residual populations had

produced a large larval population, most of which would probably not mature

because of lack of food. On the Colorado desert, evidence was found of a

large but earlier emergence, perhaps a generation ahead of that on the

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Mojave. During the April flights the migrating butterflies provided a

definite hazard in the San Joaquin Valley because they obscured windshields.

Blackbirds and English Sparrows feasted on the dead butterflies on the

shoulders of the highway. The general trend of the flights was northerly or

northwesterly, although the pattern varied from time to time.

Dr. Hagen and Dr. Middlekauff commented on the recent abundance

of Vanessa in the Kettleman Hills and Marysville areas of California.

Dr. John P. Figg-Hoblyn presented an address on the interpretation of

the “dual innervation” of the apterygote antenna with reference to cephalic

segmentation.

Due to the absence of Dr. Duncan his address entitled “Entomology at

San Jose State College,” was read by Dr. Tilden. The growth of the ento-

mology program at the college from 1930, when general entomology first

was placed on its curriculum, to Its current, varied offerings and facilities i-s

indicative not only of the increasing significance of the field but also of the

enthusiasm of a dedicated staff. The meeting was adjourned at 4:10 p.m.

— Deane P. Furman, Secretary.

Two Hundred and Sixtieth Meeting

The annual field meeting of the Pacific Coast Entomological Society

was held at Russelman Park, Contra Costa County, May 24, 1958. The

following members were present: Jane MacSwain, W. W. Middlekauff, Robert

L. Langston, Jon L. Herring, John A. Chemsak, Deane Furman, Ed Swift,

Hugh B. Leech, D. D. Jensen, E. G. Linsley, J. W. MacSwain, Herbert Ruckes,

Jr., J. W. Tilden, Wm. A. Russell, Paul H. Arnaud, Frank E. Skinner, D. G.

Denning, A. Earl Pritchard, Bill Ferguson, Donald M. Maddox, Charles

Cushner. Visitors were Juanita Linsley, Kay, Lynne, Bryan and Phil Furman,

Phyllis and Dave Middlekauff, William and June Simonds, Dick Gardiner,

Evelyn and Ann Langston, Mrs. Jon L. Herring, Mrs. Jerry Powell, Mary Ann

MaeHenry, Riley and Tim Swift, Patricia and Carol Jensen, Nancy, Lyn and

Ginger Ruckes, Hazel, Jimmy, Bernice and Jan Tilden, Marguerite Arnaud,

David Skinner, Loretta Dennintr, Marie Moor, Stephanie, Rickie and Robin

Ferguson, Ann and Debbie Maddox, Mary, Thomas and Bill Leech.

The pleasant, sunny afternoon was spent in picnicking, visiting with

old and new friends, collecting, and enjoying the recreational facilities of the

park. — Deane P. Furman, Secretary.

Two Hundred and Sixty-first Meeting

The two hundred and sixty-first meeting of the Pacific Coast Entomologi-

cal Society was held October 17, 1958, in the Morrison auditorium at the

California Academy of Sciences, San Francisco, California. President D. D.

Jensen called the meeting to order at 7:40 p.m.

The following members were present: E. S. Ross, 0. Bryant, G. A.

Samuelson, J. Gordon Edwards, Terry Seeno, W. A. Doolin, E. 0. Essig,

A. E. Michelbacher, Paul A. Harvey, Richard Dahl, Kenneth S. Hagen, Y.

Tanada, Don Burdick, Ibrahim K. Kaddou, Paul H. Arnaud, Jr., Frank E.

Skinner, Evert E. Lindquist, Jerry A. Powell, Otto W. Graft Jr., Laura M.

Henry, T. S. Acker, William Russell, K. P. Shea, R. L. Doutt, J. W. Mac-

Swain, Jane MacSwain, Herbert Ruckes, Jr., Paul D. Hurd, Jr., Williaiu E.

January, 1959]

PACIFIC COAST ENT. SOC.

Ferguson, D. D. Jensen, Dick Wilkey, Edwin Colt, Dennis Hynes, W. W.

Sampson, John A. Chenisak, Dennis Breedlove, W. D. Murray, Don Mac-

Neill, J. W. Tilden, J. H. Freitag, Richard Bushing, C. J. De Mars, D. P.

Furman. Visitors registering were: Edgar A. Smith, L. Caltagirone, Sterling

Bunnell, M. W. Bunnell, M. Ayroti, G. Pronin, David H. Huntzinger, Patricia,

Carol and Anita Jensen, Nancy Doolin, Marie W. Essig, Martha Michelbacher,

Kay M. Furman, Tom Leech, Ed. W. Kirschbaum, D. C. Rentz, S. D. Smith,

K. W. Brown, Mr. and Mrs. R. C. Elliott, Mr. and Mrs. Peter Rubtzoff,

Dorothy and Lynda Dahl, R. H. Van Zwalenburg, Mr. and Mrs. G. L. McGill,

Philip S. Barker, Theo F. Watson, Barry E. Pullen, F. X. Brace, L. Brace,

Margaret Cooper, Vlaxine Lindquist, Frances C. Powell, Anne Graf, G. J.

Haughey, M. Haughey, Elsa P. Russell, Ricky and Jeff Doutt, Ralph J.

Black, Art Smith, Lois B. McNally, James H. McNally, Keith A. Scott, Jen

Scott, Jeanette and Calvin Rogeis, Catherine Tosski, Mrs. W. D. Murray,

Nancy MacSwain, Mrs. C. G. Noack, John MacSwain, Elias Tuma, Paul

Kraemer, George P. Willsey, Gerard M. Thomas, David L. Wood, Glenn

Kissinger, Doris Eujimoto, Judy T amada, Ron Hall, John E. Henry, Meloi'a

Hilbert, Leon Spreyer, P. Watson, Dick Halbert, Richard H. Whitsel, Fred

J. Santana, Dorothy De Mars, Kathleen Conrey, David Bettencourt, Onkar

Singh Bindra, K. R. Thakare, R. Toschi, Nancy Ruckes, Grace Hurd, Kathryn

Hurd, W. W. A.llen, Frank B. Hicks, Stephanie Ferguson, Mrs. R. F. Wilkey,

Jose Rutilio Inezada, Woody Hilbert, D. Giuliani.

The minutes of the meetings held April 26 and May 24, 1958 were read

and approved.

The President announced appointment of a Nominating Committee to

select a slate of nominees for officers for 1959. The committee consisted of

Dr. Kenneth Hagen, Chairman; Dr. E. S. Ross and Dr. J. W. Tilden. He also

announced the appointment of an Auditing Committee composed of Dr. E. L.

Kessel, Chairman; Dr. J. W. VlacSwain and Dr. J. Gordon Edwards.

The President reported the death of one of the Society’s most valued

members. Professor G. F. Ferris. A special issue of the Pan-Pacific Entomolo-

gist is to be devoted to the memory of Professor Ferris.

Laura Henry announced that contributions are still being accepted for

the Gordon F. Ferris Memorial Fund, which will be used to provide a

scholarship in entomology.

Mr. H. B. Leech stated that Mrs. Ferris has donated drawings and other

items of interest to the Society’s historical file. The Society owes thanks to

Vlrs. Ferris for her generosity.

Dr. Ed. Smith, who recently returned from Indonesia, announced that

twenty to thirty foreign service positions are available for medical entomolo-

gists. Entomologists, either with or without previous field experience, are

being considered. The program is sponsored by the Internationa] Cooperation

Administration.

Dr. Ruckes nominated the following individuals for membership in the

Society:

Leopoldo Caltagirone, Entomology student from Chile, currently at the

University of California.

F. F. Bibby, Arizona Fertilizer Company, Phoenix, Arizona.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

Dr. George D. Butler, Jr., Dept, of Entomology, University of Arizona.

Mr. Van Zwalenburg, retired entomologist of the Hawaiian Sugar

Planters’ Association, now residing in Santa Rosa, California.

Dr. Edwards nominated for membership:

Mr. David H. Huntzinger, student at San Jose State College.

The nominees were elected unanimously to regular membership.

In response to the President’s call for notes and exhibits Dr. Doutt

described the life history of a gall wasp which has an alternation of genera-

tions. This is the first proven case of heterogony among California cynipids

and was worked out simultaneously, independently, and without knowledge

of the other’s research by Mr. Robert Lyons of Los Angeles City College

and Dr. Doutt.

The species is Dryocosmus duhiosus (Fullaway). It produces a bisexual

generation in early spring which develops in the male aments of the black

oaks. The mated females oviposit in the veins of the leaves. The eggs are

slow in hatching for the first larvae appeared only after two months and

some eggs were found to be viable and to hatch as long as six months after

deposition. Dr. Doutt found that unlike other Hymenoptera the unfertilized

eggs of this species do not develop parthenogenetically, and he suggests this

may be unique.

The galls produced on the leaves are bicornate and from them emerge

a uni-sexual generation of females. Dr. Doutt found these to be of two types:

one type produces female progeny only, whereas the other produces males

exclusively. The wasps from these leaf galls were originally described as a

distinct species in a separate genus, and the present biological study shows

they must now be synonymized with D. duhiosus which has priority. Dr. Doutt

objects to the use of trinomials to designate alternate generations, and

furthermore believes that the common usage of the terms sexual and agamic

generations is erroneous for the agamic generation is really sexual. He

would propose that the binomial be used and tben followed by an appropriate

designation of either bisexual or unisexual generation which is both descrip-

tive and accurate.

Mr. Hugh B. Leech stated that in the minutes of the 232nd Meeting of

this Society, he recorded Stator limbatus (Horn) as occurring in some

numbers at Lucerne, Lake County, California, in August, 1953 (Proc.

Pacific Coast Ent. Soc., in Pan-Pacific Ent., 30 (l):85-86. 1954). In mid-

August, 1959, be and his family were camping on the grounds of Cottage

City Resort at Lucerne. Adults of S. limbatus were flying in numbers during

the heat of the day, and annoying people by settling on them. The beetles

were almost as persistent as small flies, and two of those which got on his

skin gave an appreciable nip with their mandibles. Seed pods were maturing

on adjacent black acacia trees {Acacia melanoxylon R. Br.) at the time,

and the bruchids were readily obtained by jarring branches. It was difficult

to get even fifty per cent of those knocked onto a beating sheet, though,

because they took flight almost as they landed.

Dr. J. W. Tilden noted that the summer season was longer this year

at high elevations in the Sierra Nevada as compared with the same season

of the preceding three years (1955, 1956, 1957). Whereas in 1957 no insects

January, 1959]

PACIFIC COAST ENT. SOC.

were flying after September 1, in the same area above Tioga Pass, Yosemite

National Park, this year many insects were flying on September 3.

Of particular interest was the presence of several specimens of Parnassius

clodius haldur Edwards. This insect normally flies earlier in the season and

at much lower elevations. On September 3, 1958, it was flying at an elevation

of 11,000 feet, in the Alpine Fell-fields Association, an area where Parnassius

smintheus hehrii Edwards is normally found in July and August. This

elevation is not the usual habitat of P. clodius.

Mr. D. H. Huntzinger exhibited two Cimex pilosellus (Horv.) and

four Basilia forcipata Ferris, taken from Myotis species collected in Oregon

Caves National Monument, September 8, 1958. He also noted the collection

of one Silpha surinamensis Fabricius taken at Crater Lake National Park

on September 4, 1958, under a deer carcass. This, eastern species of Silphidae

may be a new record for Oregon.

J. G. Edwards displayed a specimen of the brilliant gi'een buprestid,

Trachykele opulenta Fall, which was collected by Mrs. Vaughn in Cambria,

California. Ecological data are not yet known concerning this specimen, but

it is believed that this is a significant extension of the known range of this

species.

Edwards also displayed a reduviid of the genus Triatoma which had

hospitalized a San Jose rancher the previous week with severe nausea, wide-

spread rash, dizziness, and so forth. When he returned home he found the

bug resting beneath his pillow. It feigned death for two days but when

heated on the stove it jumped into action and was as lively as a spider.

When it cooled, it again feigned death, for two more days, until heated

again. This procedure has been repeated again, but the specimen now is

“looking dead” again.

Dr. Usinger commented on bites of Triatoma in California. He stated

that in an average year about four such cases of severe reaction to the bites

of the bug are reported. The severity of reaction seems associated with

increasing sensitivity following successive bites.

Dr. E. S. Ross next presented the main address of the evening, dealing

with his recent extensive collecting trip in Africa. His major research interest,

study of the insect order Embioptera, requires personal collection and

observations in the major tropical regions of the world. Photographic records

of the natural history of these areas, particularly that concerned with insects,

has become a major by-product of the research. Dr. Ross described highlights

of his 26,000 mile trip through Africa. Ample evidence of the endless

opportunities for filming little known fauna and flora of the area was

presented in the form of a series of wonderful color slides.

The meeting was adjourned to the “Coffee Social” at 10 p.m. — Deane

P. Furman, Secretary.

Two Hundred and Sixty-second Meeting

The two hundred and sixty-second meeting of the Pacific Coast Ento-

mological Society was held December 13, 1958, in the Morrison Auditorium

at the California Academy of Sciences, San Francisco, California. President

D. D. Jensen called the meeting to order at 1:10 p.m.

The following members were present: R. L. Doutt, Herbert Ruckes, Jr.,

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

D. P. Furman, D. D. Jensen, F. E. Skinner, E. L. Kessel, P. D. Hurd, Jr.,

R. L. Usinger, E. 0. Essig, K. S. Hagen, J. H. Freitag, R. W. Thorpe, D. D.

Linsdale, E. E. Lindquist, J. A. Powell, J. W. MacSwain, T. S. Acker, R. L.

Langston, J. W. Tilden, W. D. Murray, L. E. Caltagirone, P. H. Arnaud, Jr.,

C. D. MacNeill, C. Cuschner, P. A. Harvey, D. J. Burdick, W. A. Doolin,

A. E. Pritchard, J. A. Chemsak, E. S. Ross, M. Wasbauer, R. C. Miller.

Visitors registering were: H. Ruckes, Sr., John MacSwain, James Milstead,

Derham Giuliani, Woody Hilbert, H. Bikler, K. W. Brown, Marshall Reed,

Nancy Doolin, Jack R. Powers.

The minutes of the meeting held October 17, 1958 were read, amended

and approved.

Di-. Ross announced that Mr. Owen Bryant, a long time member of the

Pacific Coast Entomological Society, passed away late in October of this

year and left the bulk of his estate to the California Academy of Sciences

for the benefit of the Department of Entomology. This included the remaining

portions of his insect collection and library as well as an endowment fund.

Prior to his death Mr. Bryant had already donated a large part of his insect

collection to the California Academy of Sciences.

Dr. H. Ruckes, Jr., proposed the following nominees for membership in

the Society: Woody Hilbert, Derham Giuliani, Jack R. Powers, Robin Thorpe,

C. E. Kennett. They were elected by unanimous vote to full membership.

President Jenson announced that Mr. Van Zwalenburg had been appoint-

ed to fill the vacancy in the Publication Committee left by the death of Pro-

fessor Ferris. Dr. J. Gordon Edwards was appointed as a member of the

Special Committee on Nomenclature, and Dr. Usinger was appointed chair-

man.

The Chairman of the Auditing Committee, Dr. E. L. Kessel, reported

the books of the Society were found to be in order.

In response to the President’s call for notes and exhibits, Jerry A. Powell

exhibited specimens of an unusual tortricid moth, Synnoma lynosyrana

Walsingham. The species shows remarkable sexual dimorphism which is

a very aberrant characteristic among the Tortricidae. The females, having a

very large abdomen, seem unable to fly successfully. The genus and the

monotypic species were described from specimens collected in 1871 on the

plains northeast of Mt. Shasta by Lord Walsingham. The species has sub-

sequently been reared from Chrysothamnus from the same area and at least

as far north as southern Idaho. The specimens shown were reared from larvae

collected on both Chrysothamnus nauseosus and C. viscidiflorus near Weed,

Siskiyou County, California in August, 1958. The larvae live in tough, silken

tubes and are often so numerous that they draw up the branches of the host

plant into conspicuous bundles. Also shown were the black egg patches of

the species which had been laid in late September. Some of the eggs had

hatched, but it was suggested that this was probably due to the laboratory

conditions, as the larvae died without constructing hibernaculi.

Dr. J. W. Tilden exhibited the larva of a fly of the genus Microdon,

family Syrphidae, that had been taken by Paul Shafer, a student at San

Jose State College. The larva was collected on November 28, 1958, from

under the bark of a decayed stump of ponderosa pine, in an area that had

January, 1959]

PACIFIC COAST ENT. SOC.

been timbered over about fourteen years ago. The locality was three miles

below Georgetown, El Dorado County, California, at an elevation of 2300 ft.

Donald J. Burdick and Marius Wasbauer presented the following paper

entitled. Provisioning behavior of Methocha calif ornica Westwood.

“The only previous biological note concerning Methocha californica

Westwood was presented in 1912 at the 46th meeting of this society. At this

meeting J. C. Bridwell in a commentary on some rare and little known

Hymenoptera, which he had collected, mentioned that the species was

‘parasitic in the burrows of tiger beetles.’ The first account of Methocha

as a cicindelid parasite was by Adlerz in 1903 when he observed a female

of M. ichneumonides Latreille successfully attack and paralyze a large

tiger-beetle larva. Since this time a number of observations on this and

other species have been reported and in each case a' cicindelid larva was

used as prey.

“Our observations were made on two females collected by J. Powell and

Burdick as they crossed the nesting site of a species of a sphecid which

they had under observation. After establishing Omus calif ornicus Eschscholtz

larvae, admittedly abnormal hosts, in observation burrows, a Methocha

female would be placed in the jar and watched while she searched, found,

paralyzed and oviposited on the Omus larva. In all, twelve larvae were

paralyzed and eleven eggs were laid by the two Methocha females. When

first dropped into the jar, the female, moving rapidly and randomly, investi-

gated any depressions or burrows that she came across. Upon finding the

occupied burrow, she pointed her antennae down into the burrow and

became quiescent. Usually she would then move to the side of the burrow

that the host’s mouthparts were on and would walk down the burrow toward

the host. This orientation placed the female in position to reach the ventral

part of the host’s neck with her sting. The Omus larva, by this time aware

of the intruder, had prepared itself to strike. The approach of the Methocha

was very rapid, and she was able to place at least her first two pairs ol

legs on top of the flat shield formed by the head and prothorax of the host

neiore the larva lunged upward, raising its head and closing its mandibles

upon its attacker. At this time, the female dropped her abdomen down, and

the mandibles of the Omus closed harmlessly around the wasp’s narrow

waist. In this position, the female stung her host in the neck, temporarily

paralyzing it. She then lifted her abdomen through the now inactive mandibles

of her host and climbed out of the burrow. Here she wandered about for a

short time and then reentered the burrow and began preparing the host

for oviposition.

“After the initial sting delivered by the female Methocha, the tiger

beetle larva was usually rendered immobile. However, thei'e was some

variation in the degree of paralysis. The paralysis was of a very temporary

nature, and in most cases eighty to ninety percent mobility was regained by

the Omus larva within 24 hours.

“Following the initial sting and paralysis of the prey, there was always

a secondary stinging response which was altogether different from the first.

The female Methocha oriented herself so that she faced the venter of the

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 1

prey within the hurrow, grasped one of the fore or mid coxae in her man-

dibles, curled her entire body upwards and around and stung the Omus

larva in the membranous area just behind the head capsule. This was

repeated an average of four to five times during the course of a provisioning

cycle. We found that we could increase the number of subsequent stings

merely by moving the Omus larva in the burrow. Apparently movement on

the part of the host incited the stinging response.

“After the host larva was stung into quiescence, the wasp engaged in

host feeding. She accomplished this by maxillating the venter in the neck

region or at the base of one of the coxae. When she succeeded in puncturing

the skin she then fed on the exuding blood.

“In preparing the larva for oviposition, the wasp tilted the head capsule

of the prey back by pushing it with her head. She then climbed up and

grasped the head capsule with her fore and middle legs, moving her

abdomen around until it contacted the hind coxa of the host larva. The

abdomen was then moved back and forth several times and the egg extruded.

The egg was always placed mesally at the base of the hind coxa and its

position was relatively constant. This constancy is understandable when

we consider that the prey recovered rapidly and was able to move about

in the burrow. In this position the egg was relatively safe from being crushed

or dislodged.

“After oviposition, the wasp began to fill the burrow, usually first by

prying material loose from the upper sides of the burrow with her mandibles

and then walking out of the burrow to search for pieces of material of

suitable size, which she carried in her mandibles and placed on the cephalo-

thoracic shield of the prey. We noted no raking of material with the front

legs, a trait characteristic of so many aculeate wasps.

“The egg hatched in from four to five days and the first instar larva

remained in the same position as the egg had been, feeding suctorially on

the body juices of the host. This suctorial feeding continued up to the time

of death of the host, in about ten to twelve days. By this time, the Methocha

larva was quite large and moved about freely, often feeding inside the body

of the dead Omus larva. The active larval period of the parasite was com-

pleted in twelve to fifteen days. Two of the larvae formed cocoons. The

adults will probably not emerge until next spring.”

The talk was illustrated by a series of excellent color slides.

Dr. Hagen, as Chairman of the Nominating Committee, announced that

the following members had been nominated for officers for 1959: R. L.

Doutt, President; R. M. Bohart, Vice-President; D. P. Furman, Secretary;

R. C. Miller, Treasurer. No further nominations were I'eceived from the

floor. The nominees were unanimously elected to office.

President Jensen turned over the chairmanship of the meeting to incom-

ing President Doutt.

Dr. Jensen delivered the Presidential address entitled “Insects — Both

Hosts and Vectors of Plant Viruses.” The text of his stimulating address

is to appear as a separate paper in the Pan-Pacific Entomologist.

The meeting was adjourned to the “Coffee Social” at 3:40 p.m.— Deane

P. Furman, Secretary:

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Vol. XXXV APRIL, 1959

No.2

THE

Pan-Pacific Entomologist

CONTENTS

JENSEN — Insects, both hosts and vectors of plant viruses 65

PRITCHARD & POWELL — Pyramidobela angelarum Keifer on

ornamental Buddleia in the San Francisco Bay area 82

ABBOTT — The 1958 migration of the Painted Lady Butterfly,

Vanessa cardui (Linnaeus), in California 83

RUCKES — Two new records for the cone beetle genus Conoph-

thorus Hopkins in California 94

SCHUSTER — Notes on Morius occidens Casey with a description

of the male - 95

KORMONDY — A few Odonata from Yosemite 98

SMITH — The Japygidae of North America 1 — Provalljapyginae

and Nanojapyx 99

CLAUSEN — Releases of recently imported insect parasites and

predators in California, 1956—57 - 107

POWELL — Notes on the California species of the genus Pyla Grote 109

EATON — Observations on the survival of Arhopalus productus

(LeConte) larvae in Douglas-flr lumber 114

WILLIAMS — A preoccupied name in Solierella 116

FOURTH ANNUAL INSECT PHOTOGRAPHIC SALON 97

SAN FRANCISCO. CALIFORNIA • 1959

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

THE PAN-PACIFIC ENTOMOLOGIST

EDITORIAL BOARD

E. G. Ltnsley P. D. Hurd, Jr., Editor R. L. Usincer

E. S. Ross J. A. Powell, Acting Editor H. B. Leech

D. D. Linsdale, Asst. Editor

R. C. Miller, Treasurer J. E. Swift, Advertising

Published quarterly in January, April, July, and October with Society Proceed-

ing appearing in the January number. Papers on the systematic and biological

pha,gss of entomology are favored, including articles up to ten printed pages on

insect taxonomy, morphology, life history, and distribution.

Manuscripts for publication, proof, and all editorial matters should be addressed

to J. A. Powell, at 112 Agricultural Hall, University of California, Berkeley 4,

Calif. All communications regarding non-receipt of numbers, changes of address,

requests for sample copies, and all financial communications should be addressed

to the treasurer. Dr. R. C. Miller, at the California Academy of Sciences, San

Francisco 18, California.

Domestic and foreign subscriptions, $4.00 per year in advance. Price for single

copies, $1.00. Make checks payable to “Pan-Pacific Entomologist.”

MEMOIRS SERIES

of the

PACIFIC COAST ENTOMOLOGICAL SOCIETY

THE SUCKING LICE by G. F. Ferris S6.00

A 320-page book wiiicli summarizes the knowledge on

the Anopiura of the world. Published by the Society,

October, 1951.

THE SPIDER MITE FAMILY TETRANYCHIDAE by A. Earl

Pritchard and Edward W. Baker $10.00

This world-wide treatment deals with the systematics

identification, and economics of the “Red Spiders” and

includes descriptions of thirty-three new species. Pub-

lished by the Society, July, 1955.

Send orders to: Treasurer, Pacific Coast Entomological Society,

California Academy of Sciences, Golden Gate Park 18, San

Francisco.

Second-class postage paid at San Francisco, California.

The Pan-Pacific Entomologist

Vol. XXXV. April, 1959

No. 2

INSECTS, BOTH HOSTS AND VECTORS OF

PLANT VIRUSES ^

D. D. Jensen

University of Calif ornia, Berkeley

Man, in his attempts to explain the observed phenomena of

nature, travels from naive conjectures to the complexities of

experimental facts and then seeks generalized simplicity. In the

field of virus research we are still in the complex stage in which

every generalization proposed has at least one Achilles Heel.

A Newton of viruses has not yet appeared on the horizon. Never-

theless, the chase is on and in scores of laboratories throughout

the world the quarry is being glimpsed at closer and closer, though

still distant, range. Most avidly pursued of all is tobacco mosaic

virus which must scarcely dare call its nucleic acid its own.

Among the most complex and least understood of the plant

viruses are those which rely heavily or exclusively on insects for

their dissemination in nature. The relationships of viruses to their

vectors have been studied by a relatively small group of research

workers. Yet evidence is accumulating that the insect vector, far

from being merely a “flying needle” contaminated with virus, as

some have been inclined to consider it, may prove to be the

pristine host of some, if not all, plant viruses.

I should like to review some of the more interesting discov-

eries and hypotheses involving vector-virus relationships, parti-

cularly the recent ones and those dealing primarily with the leaf-

hopper transmitted viruses.

Some important discoveries have been made accidentally or

as by-products of planned investigations; others have required

the penetrating thought and imagination so perceptively described

by Schopenhauer (and recently quoted by a colleague) when he

said : “Thus the task is, not so much to see what no man has seen

yet, but to think what nobody has thought yet, about that which

everybody sees.”

Why is a Vector a Vector?

The vast majority of known plant virus vectors are insects.

Eriophyid mites are also proving to be important vectors of a

^ Presidential address, presented to the Pacific Coast Entomological Society, December 13,

1958.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

few viruses in recent years. This year for the first time, a non-

arthropod was demonstrated to transmit a plant virus when

Hewitt et al. (1958) reported Xiphinema index Thorne and Allen,

a plant parasitic nematode, to be a vector of grape fan leaf virus.

One of the basic questions in vector-virus relationships, and

one about which we are still almost completely ignorant, concerns

the factors which govern the transmissibility of viruses by vectors.

The vectors of most plant viruses show general but not inflexible

phylogenetic affinities. Thus, aphid-transmitted viruses have not

been shown conclusively to be transmitted also by leafhoppers,

beetles or other insect groups. Viruses associated with leafhoppers

are transmissible only by leafhoppers — except in the case of

Pierce’s disease virus of grapes which is carried by spittle bugs

of the family Cercopidae and by sharpshooters in the family

Cicadellidae, but not by other leafhoppers. The common denomi-

nator among all the vectors of this virus is their habit of feeding

in the xylem rather than the phloem of the plants.

There appears to be less group specificity among the vectors

of animal viruses. For example, the experimental vectors of the

virus causing western equine encephalitis include ticks as well

as mosquitoes.

Bennett and Wallace (1938) have shown that the curly top

virus of sugar beets, transmitted in the United States only by

Circulifer tenellus (Baker), can be acquired and carried in the

bodies of aphids, mites, thrips and other leafhoppers for long

periods of time, but is not transmitted by these species.

The specific cause for failure of a virus to be transmitted by

non-vector species or by a genetic strain of its normal vector has

been demonstrated only once. Storey (1939) bred a strain of

Cicadulina mbila (Naude) which consistently failed to transmit

maize streak virus in Africa. However, these “inactive” indi-

viduals could be made active vectors if, after they had fed on a

diseased plant, the stomach wall of the insects was punctured with

a fine needle to permit some of the infective plant juice to enter

the blood in the body cavity. This showed that the barrier to

virus passage existed in the wall of the intestine.

Present evidence suggests that the constitutional stability of

insects as vectors of plant viruses is much greater than are the

viruses which they transmit. However, this does not imply that

April, 1959] jensen — insects and plant viruses

vector species are homozygous in this respect and show uniformity

and consistency in transmitting ability.

Bjorling and Ossiannilsson (1958) have just published the

results of extensive experiments involving 10 asexually reared

strains of six different species of aphids as vectors of two persist-

ent viruses, beet yellows and potato leaf roll. The aphids were:

Myzus persicae (Sulzer), M. ascalonicus Doncaster, Aphis fabae

Scopoli, Aulacorthum circumflexum (Buckton), A. solani Kalten-

bach) and Macrosiphum euphorbiae (Thomas). Nearly all of the

strains of Myzus persicae were better vectors of both potato leaf

roll and beet yellows virus than were the best strains of the other

aphid species compared.

Most of the work dealt with 85 strains of Myzus persicae

which had been collected from widely different geographical and

ecological sources. Virus transmission tests over a three year

period revealed that these 85 strains could be grouped in a

continuous series which ranged from 10 per cent to 80 per cent

in virus transmitting ability. Moreover, there was no correlation

between the field source plant of the aphid strain, nor the plants

on which the aphids fed just prior to being used in virus tests,

and their efficiency as vectors.

Most of the strains were fairly efficient vectors with individual

aphids of 70 of the 85 strains transmitting beet yellows virus to

30 per cent or higher of the test plants. The absolute differences

in transmission efficiency were great between some strains and

significant in a number of cases. The authors were led to conclude

that “these differences between more or less effective vector strains

within the same species seem to be genetically determined.” This

was also supported by tests in which two aphid strains of similar

vector efficiency were crossed. The efficiency of the progeny was

markedly below that of the parents.

Loss AND Recovery of Insect Transmissibility by Viruses

There is now considerable evidence that the insect transmis-

sibility of many plant viruses is not constant, but varies sometimes

in relatively short periods of time. For many years it has been the

not uncommon experience of those who work with insects and

viruses to find that known vectors of certain viruses sometimes

transmit them inefficiently or not at all. Very often this is encoun-

tered after an interruption of transmission work for several

months or years. It is apparently often due to a reduction in virus

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

litre in a plant which has been infected for an extended period

of time. However, in other instances it may occur with viruses

whose titre has been maintained at a high level by frequent

transmission to new plants or animals by means of mechanical

inoculation.

Transmissibility by mosquitoes of certain strains of yellow

fever, encephalitides and dengue viruses is greatly reduced or

eliminated after repeated passage by mechanical means in the

vertebrate hosts (Reeves, 1958).

Viruses are known to mutate and it is probable that vectorless

mutants may appear which, in the absence of insect transmission,

might supplant the insect transmissible virus strain.

Black (1953) maintained three isolates of potato yellow dwarf

virus for 12-16% years in plants without insect transmission. At

the end of this period the leaf hop per vector was unable to transmit

these isolates whereas control leafhoppers transmitted fresh

isolates of the virus, collected in the field, with normal high

frequency. Very recently Black et al (1958) reported the same

phenomenon for wound-tumor virus and its leafhopper vector.

Similar loss of transmissibility by aphids was reported by

Hollings (1955) for tomato aspermy virus. One isolate, at first

easily transmitted by aphids, was transmitted only with great

difficulty by vectors after two years of mechanical transmission

to tobacco.

I worked with an aphid transmissible virus of garden

nasturtium, Tropaeolum majus, over a period of several years.

Vector work was then discontinued for approximately two years

during which the virus was transmitted by juice inoculation

every few months. When aphids were tried again as vectors trans-

mission occurred only rarely. Field collected Myzus persicae

were also tried as vectors but they succeeded no better than did

our greenhouse colonies.

In the above cases loss of transmissibility by insects followed

two or more years of virus maintenance or transmission in the

absence of insects. Swenson (1957) reported that one of two

isolates of bean yellow mosaic virus lost its ability to be trans-

mitted by three species of aphids within a period of one to

four months.

The most obvious explanation suggested by these results is

that the virus isolates in question lost their ability to be trans-

April, 1959] JENSEN INSECTS AND PLANT VIRUSES

mitted by their normal vectors as a result of mutations of the

virus. Since the tomato aspermy virus isolate became almost com-

pletely vectorless in a two-year period it may be inferred that the

alleged mutant strain was replacing the insect transmitted strain

gradually. In contrast to this, however, is the fact that Swenson’s

bean virus apparently became completely vectorless within one

to four months after having been transmitted readily by three

species of aphids.

That the apparent loss of insect transmissibility by a virus

may be reversible has been reported by Watson (1956, 1958).

Potato virus C, derived from potatoes, was not transmissible by

aphids. However, after mechanical inoculation into Nicotiana

glutiiiosa this virus could be transmitted by Myzus persicae.

When returned to potato by means of aphids, the virus ultimately

reverted to the vectorless strain although certain isolates remained

aphid transmissible through several passages in potato.

The first speculative explanations suggested by this phenomenon

were of two different types. One involved possible mutations; the

other required a quantitative rather than a qualitative hypothesis

and postulated vectorial and vectorless virus particles which varied

in ratio in the different hosts. Each of these hypotheses required

assumptions that are without precedent.

What may prove to be a more plausible explanation has just

been reported. Badami and Kassanis (1958) obtained a strain of

potato virus Y, from Solanum jasminoides from India, which in

potato closely resembled the potato virus C used by Watson.

However, after separation from two other viruses which were also

present in S. jasminoides, this virus was readily transmitted by

Myzus persicae. These previously undescribed viruses are reported

to decrease the multiplication of the virus Y strain and also its

aphid transmissibility.

This suggests that Watson’s potato virus C may not be aphid-

transmissable from potato because of the possible presence in

potato of another virus which reduces or prevents transmission of

virus by aphids. Tobacco may not be a host of this virus and thus

virus C would be available for transmission by aphids.

This phenomenon is just the opposite of that reported by Smith

(1945) for the virus complex known as tobacco rosette. One

component, mottle virus, is easily transmitted mechanically but

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

cannot be transmitted by Myzus persicae unless it occurs in the

same plant which carries vein-distorting virus.

Cross Protection Between Viruses in Insects

Although plants are not known to produce antibodies to infec-

tive disease agents such as we know to be common in animals,

plants, after infection by a virus, may acquire immunity to related

and sometimes more virulent strains of the same virus. The different

strains have been assumed to arise through mutation, but there

is now evidence that genetic recombination may also be involved.

The ability of one virus strain to prevent a second strain from

producing its own characteristic disease symptoms in the same

plant has been considered evidence that the two strains are elosely

related. However, it is also possible to have closely related strains,

such as those of curly top virus of sugar beets, which do not protect

against each other either in the plant or in the insect vector

(Giddings, 1950) .

Kunkel (1955) showed that the California and the eastern U.S.

strains of aster yellows virus protect against each other in some

of their common host plants. He also obtained evidence that the

same strains protect against each other in the aster leafhopper,

Macro steles fascifrons (Stal) . Working with groups of leafhoppers,

he found that those which fed for two weeks on plants infected

with one strain of virus and then fed two weeks on plants carrying

the other virus strain, always transmitted only the first strain.

Experiments of several different types have demonstrated that

aster yellows virus multiplies in the aster leafhopper as well as in

the plant. Although the mechanism of cross protection among

viruses is not known, it has been inferred that related strains need

the same materials for multiplication and that the first virus to

invade a host occupies the sites of virus multiplication and uses

up the materials available, thus denying to the challenging strain

the products it needs for multiplication.

However, more extensive experiments by Freitag (1958) in

California with three strains od aster yellows virus and the aster

leafhopper have shown that cross protection is not always stable

and permanent. Working with single insects and transferring them

daily over long periods of time, he found that under some circum-

stances the leafhoppers finally stopped transmitting the first strain

acquired and then transmitted the challenging strain. These were.

April, 1959] jensen — insects and plant viruses

however, exceptional cases. In most instances there was marked

cross protection in both insect and plant.

Maramorosch (1957, 1958a) has reported the same phenom-

enon with two strains of corn stunt virus and its vector Dalbulus

maidis (DeLong & Wolcott). In the studies conducted with aster

yellows virus and with corn stunt virus, one strain in each disease

complex showed greater dominance than the other in both the

insect vector and the host plant. In the leafhoppers, strain “A”,

acquired first, was sometimes replaced by strain “B”. However, if

strain “B” was acquired first, strain “A” failed to supplant it. This

indicates that the phenomenon of cross protection is more complex

than mere prior occupancy of virus multiplication centers in the

insect or plant by the first strain to enter.

Our perplexity regarding the nature of this apparent competi-

tion between plant virus strains within the body of the leafhopper

vector is compounded by the nature of strain replacement in the

insect. In a few instances, test plants fed on by single insects

showed symptoms of both virus strains. In most cases, however,

symptoms were entirely of one strain or the other. Also, replace-

ment of one strain by a more dominant strain within the same

leafhopper was not always permanent when first accomplished.

Sometimes the strains were transmitted alternately before the

dominant strain finally completely supplanted the other.

The presence and multiplication of two virus strains in the

same leafhopper vector have not yet been known to result in a

new hybrid virus. However, more extensive tests involving related

virus strains carried by the same vectors might produce new hybrid

plant viruses in insects as it has in plants and in vertebrate animals.

Recombinations of genetic units between related strains of

animal and plant viruses have already been reported. The first

work of this type was done by Burnet and Lind (1951) with

strains of influenza virus. Subsequently, Best and Callus (1955)

and Best (1956) achieved similar results with tomato spotted wilt

virus. Plants infected with mixtures of two strains of this virus

developed symptoms of both parent strains and also symptoms

of new strains. From these plants with mixed infections they

recovered both parent strains and also several new strains. The

latter were distinct from each of the parent strains but possessed

some of the characters of each parent. Further experiments led

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

to the conclusion that such new strains were genetically stable and

not mere mixtures of the two parent strains.

It would be of great interest to determine if parallel results

could be obtained by allowing the thrips vector to acquire two

or more of these same strains of spotted wilt virus. Multiplication

of spotted wilt virus in thrips has not yet been demonstrated, but

the 5—10 day incubation period of the virus in the vector, plus

long retention of the virus by the vector,' suggest multiplication.

Beneficial Effects of Plant Virus in Relation to

Insect Vectors

A few reports exist indicating that a plant virus may be of

indirect benefit to its own insect vector or to a non-vector species.

Carter (1939) found that infected Emilia sonchifolia, a weed host

of both spotted wilt virus and its vector Thrips tahaci Lindeman,

maintained a higher population of thrips than did healthy plants.

The reasons for this appeared to be that diseased plants had

curled leaves which provided improved shelter for the thrips vector

and that these plants also grew more slowly and survived longer.

A more direct beneficial effect of virus on Aphis fabae has

been reported by Kennedy (1951). He found that sugar beet

mosaic virus altered the physiology of the beet plant in such a

manner that the production of young aphids per mother averaged

higher on virus infected leaves of all ages than on healthy leaves

of comparable age.

Severin (1946), incidental to his search for new leafhopper

vectors of aster yellows virus, found that 9 species of leafhoppers

completed their nymphal development on celery or asters infected

with California aster yellows virus, but that the adults died within

a few days after transfer to healthy celery or asters. No data were

presented on the longevity of any control adults that may have

been held on diseased plants. The results given were interpreted

as demonstrating that aster yellows virus in some way altered

celery and aster plants so as to make them more suitable food hosts

for the leafhopper species tested.

Maramorosch (1958b) reported similar conclusions drawn

from experiments with Dalbulus maidis (DeLong & Wolcott), a

non-vector of aster yellows virus. This leafhopper, which transmits

corn stunt virus, was known to feed well and breed only on Zea

mays and Euchlaena mexicana. Eighty adult insects were caged

on healthy China asters and 80 were caged on asters infected with

April, 1959] jensen — insects and plant viruses

aster yellows virus. All insects on healthy asters were dead within

four days, but 63 of 80 were still alive on virus-infected plants

after 42 days. The survival on diseased asters was practically

identical with the survival on corn plants hut no eggs were

deposited in the aster plants. In subsequent tests D. maidis, after

five weeks on infected asters, were successfully maintained on

healthy asters. Adults and nymphs of a related leafhopper, D.

elimatus (Ball), with a limited host range identical to that of

D. maidis, were found to survive well on asters carrying virus, but

died rapidly on healthy asters. Maramorosch interprets his experi-

ments as indicating that the virus altered the chemical composition

of the aster plant in the direction of more adequate nutrition for

the leafhoppers which, in the case of D. maidis, became sufficiently

adapted to aster as to permit survival even on virus-free asters.

Among the. species discussed by Severin (1946) in his report

on the longevity of leafhoppers on virus-infected and healthy

plants was Texananus spatulatus Van Duzee. He stated that

individuals of this species which transmitted aster yellows virus

lived longer than those which failed to transmit virus. Only a

small part of the data on which this conclusion was based has

been published (Severin, 1945). Since the virus transmitting

efficiency of this species was exceedingly low (most individuals

carrying virus infected less than one percent of the plants fed

upon) and since it is not clear whether the transmitters and the

non-transmitters were maintained in the same manner, further

confirmation is needed for the generalization that the transmitting

insects lived longer than the non-vectors. Severin construed the

increased longevity to mean that the plants infected were better

hosts and prolonged the life of the leafhoppers. However, since

the insects were transferred daily to new test plants and trans-

mitted virus to only a few plants this explanation does not seem

feasible. If the transmitting insects lived significantly longer than

non-transmitters, it would be evidence that the virus per se was

beneficial to the insect.

No reports, parallel to those given for plant viruses, have been

found to indicate that animal viruses may render their hosts

suitable as food sources for arthropods if, in the absence of virus,

such hosts are unacceptable to the arthropod species. Virus-infected

animals sometimes carry abnormally high populations of arthropod

parasites but apparently this is only because such animals become

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

debilitated and therefore are less capable of ridding themselves of

the parasites than are healthy individuals.

In none of the situations just described was it suggested by

the research workers involved that the virus itself contributed

directly to the increased longevity of the insect vector.

A Virus Harmful to Both Animal and Plant

Although there have been many insect-borne plant viruses

known and a less lengthy list of insect-borne animal viruses, until

recently there has been no report of a virus that was harmful to

its insect vector.

It is true that we have such an example among the rickettsiae,

some of which have also been called viruses, but Snyder (1948)

classes these as “micro-organisms . . . intermediate in character-

istic between bacteria and viruses . . .” The body louse, vector of

typhus fever rickettsiae, is also killed by the infectious agent it

carries. Zinsser (1935), describes the plight of the body louse in

these words: “The louse shares with us the misfortune of being

prey to the typhus virus. If lice can dread, the nightmare of their

lives is the fear of some day inhabiting an infected rat or human

being. For the host may survive; but the ill-starred louse that

sticks his haustellum through an infected skin, and imbides the

loathsome virus with his nourishment, is doomed beyond succor.

In eight days he sickens, in ten days he is in extremis, on the

eleventh or twelfth his tiny body turns red with blood extravasated

from his bowel, and he gives up his little ghost.”

The first indication that a plant virus could cause tissue changes

in its insect vector was given by Littau and Maramorosch (1956).

They reported that aster yellows virus caused a higher percentage

of fat body cells to have stellate nuclei than occurred in virus-free

Macrosteles fctscifrons (Stal) . There is as yet, however, no evidence

that these effects are harmful to the leafhopper.

The effect of aster yellows virus on the longevity of this vector

was tested extensively by Severin ( 1947 ) . Both infective and non-

infective leafhoppers were held on Sacramento barley, which is

immune to the virus, and survival was recorded. Adult longevity

ranged from approximately 30 days to over 140 days. The 350

infective leafhoppers survived as long as did an equal number of

non-viruliferous individuals. He concluded that the virus itself is

neither beneficial nor injurious to adult aster leafhoppers.

A report from Japan (Yoshii and Kiso, 1957), indicates that

April, 1959] jensen — ^insects and plant viruses

in several respects the virus causing dwarf disease of orange alters

the metabolism of infected orange leaves and the metabolism of

the plant hopper vector, Geisha distinctissima WaL, in a similar

manner. For example, oxygen consumption and total phosphorus

were reduced in both host plant and vector. The publication does

not indicate whether or not these metabolic disturbances adversely

affected the infective insects in a way that was reflected in reduced

longevity, reproduction or activity.

Recently we have been able to demonstrate that the most im-

portant stone fruit virus in western North America, Western X-

disease virus, causes the premature death of at least one of its

leafhopper vectors.

Colladonus montanus (Van Duzee) had proved to be an exceed-

ingly inefficient vector of the virus from peach to peach. However,

after the discovery that celery is also a susceptible plant host of

the virus (Jensen, 1956), C. montanus was retested using celery as

the virus source. From these experiments it was determined that

C. montanus readily acquires the virus from celery. Groups of

leafhoppers, fed alternately on a series of healthy peach and celery-

plants, transmitted virus to 57 of 213 celery plants but to only

four of 238 peach trees (Jensen, 1957) . Because of the superiority

of celery over peach as both a virus source and a test plant and

because C. montanus lives well on celery but dies rapidly on peach,

experiments comparing the longevity of viruliferous and non-

viruliferous leafhoppers were carried out on celery. Longevity was

measured from the first day on test plants which was after com-

pletion of the average latent period of the virus in the vector.

The experiments, reported elsewhere (Jensen, 1958; 1959),

show that Western X-disease virus itself causes the premature death

of its insect vector. The leafhoppers which did not transmit virus

survived on the test plants approximately twice as long as did the

transmitters. The increased mortality was not due to altered plant

nutrition, because in many of the tests all insects, both transmitters

and non-transmitters, fed together on the same virus source plants.

The single variable was whether or not a given individual happened

to acquire virus from the diseased plant upon which all had fed.

Moreover, an alternative explanation cannot be found in inher-

ent differences between individual leafhoppers. The percentage of

insects proving to be infective was usually directly proportional

to the access time on the inoculum plant. Thus, in one experiment

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

insects from the same stock colony were caged on a diseased plant

and removed from the plant as three different groups each with

a different acquisition feeding time. Of those removed after three

days, 27 per cent transmitted virus. After seven days, 70 per cent

transmitted and after 16 days 90 per cent transmitted virus. In

each group, longevity was greatest among the non-transmitters.

Multiplication of Virus in Insect Vectors

Multiplication of persistent viruses in their aphid vectors has

not yet been demonstrated conclusively. Day (1955) reported

experiments which suggested to him that potato leaf roll virus

multiplies in the aphid Myzus persicae. However, Harrison (1958)

found that infectivity and retention of this virus by M. persicae

increased with feeding time on a virus source and that aphids

gradually lost their virus on immune hosts. He concluded that the

virus does not multiply in the aphid.

It has now been well-established that four plant viruses do

multiply in their leafhopper vectors as well as in their plant hosts.

These are the viruses of rice stunt, clover club-leaf, aster yellows

and wound tumor.

Fukushi (1939) demonstrated that rice stunt virus is trans-

mitted transovarially by its vector Nephotettix apicalis Motschulsky

var. cincticeps Uhler. Starting with a single infective female, the

experiment was carried through six leafhopper generations over

a period of 374 days. At the end of the experiment there was no

evidence of a decline in the number of insects per generation be-

coming infective nor in the percentage of plants infected.

Black (1950) carried out an experiment, similar to that of

Fukushi, with clover club-leaf virus and its vector Agalliopsis

novella (Say). Starting with a single viruliferous female, Black

maintained the insects for five years and through 21 generations

on alfalfa which is immune to the virus. At the end of this period

there was no reduction in virus infectivity in the insects. If there

was no multiplication in the insects the virus would have to retain

its infeetivity at a dilution conservatively estimated to exceed

1: 2.8X10^^* This is far beyond the dilution tolerance of the virus

and means that the virus multiplies in the vector.

Occasional transmission of plant virus through the egg of the

insect vector has also been reported by Black (1953) for wound-

tumor virus and potato yellow dwarf virus in their respective

vectors, Agalliopsis novella and Agallia constricta Van Duzee. Also,

April, 1959] JENSEN^ — insects and plant viruses

Grylls (1954) reported that rugose leaf curl virus is transmitted

through the egg of its leaf hopper vector, AustJ'oagallia torrida

Evans. It is not yet known whether these viruses can persist in

their vectors generation after generation, as has been demonstrated

for the viruses causing rice stunt and clover club leaf, without

being replenished by feeding on a diseased plant.

That aster yellows virus (Black, 1941; Maramorosch, 1952 j

and wound tumor virus (Black & Brakke, 1952) multiply in their

respective leafhopper vectors has been shown by serial passage

of the viruses through their vectors. Diluted virus was injected

mechanically into the bodies of the vectors. Later, virus was

recovered from the same insects, diluted, and injected into fresh

leafhoppers. Ten such serial passages were made with aster yellows

virus and seven with wound tumor virus, yet the concentration of

virus in the insects attained as high a level in the last passages as

in the first. In the absence of multiplication one would have to

postulate a dilutions of 10'^° for aster yellows virus and 10"^^ for

wound tumor virus.

It is probable that among the many other leafhopper trans-

mitted viruses, especially those having long incubation periods

in the vector, will be found additional plant viruses which also

multiply in insects. However, it should not be assumed that all

leafhopper vectors are also virus hosts. Freitag (1936) and Bennett

and Wallace (1938) have presented strong evidence that curly top

virus of sugar beets does not multiply in the beet leafhopper,

Circulifer tenellus (Baker). It should be added, however, that this

virus has a very short (approximately 24 hours) incubation period

in the vector.

Virus Origin

After tobacco mosaic virus was purified and shown to be a

crystalline protein (Stanley, 1935), viruses were considered by

some to be unrelated to living organisms. It was suggested that

plant viruses originated first from plant cell components and

secondarily developed the broad spectrum of specialization which

ranges from little or no dependence upon insect vectors, such as

in the case with tobacco mosaic virus, up to the leafhopper

transmitted viruses, most of which require insects for their

dissemination.

In opposition to this idea, is the organismal theory of virus

origin proposed by Green (1935) and supported by Laidlaw

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

(1938), Andrewes (1952) and others. According to this view

viruses originated by retrograde evolution from parasitic micro-

organisms possibly akin to protozoa and bacteria. The fact that

we now have several viruses which multiply in animals (insects)

as well as in plants has brought into greater prominence the

organismal theory of virus origin, and has resulted in its extension

(Maramorosch, 1954) to suggest that plant viruses were originally

arthropod viruses.

Viruses had been observed to cause disease symptoms in the

plant hosts, but not in the insect vectors. Until plant viruses were

demonstrated to multiply in the insect vector, their association

with insects had been considered somewhat fortuitous. However,

with the knowledge that some insects are hosts and not merely

vectors of these viruses the way was open to new interpretations

as to virus origin.

Among the widely held generalizations in biology is one stating

that those parasite relationships which are characterized by severe

damage to the host are of relatively recent origin whereas those in

which the parasite lives at the expense of the host but causes the

latter little or no damage are of much greater antiquity. Thus

plant hosts, such as those of aster yellows virus, which suffer

extreme damage, would be considered recently acquired hosts. The

leafhoppers in which this virus also multiplies without apparent

harm would be considered primitive hosts. Andrewes (1957) has

reviewed this subject extensively and suggests that arthropods may

have been the original hosts not only of the insect viruses, but

also of viruses commonly associated with plants and with verte-

brates. Insects play a critical role in all three of these general

groups of viruses. Originally viruses may have had insects as

their only hosts. Later the viruses may have made use of vertebrates

and plants to get from one insect to another. Lest we become too

carried away by this argument, however, it should be pointed out

that in the host range of some plant viruses and some vertebrate

viruses there are species which are as symptomless as are the

vectors which transmit the viruses. Nonetheless, this does not

invalidate the theory. It merely means that such viruses may have

been associated with the symptomless hosts much longer than with

those still suffering damage. Andrewes (1957) points out that

epidemics of yellow fever disease occur in South American

monkeys but not in African monkeys. The virus appears to be

April, 1959] JENSEN INSECTS AND PLANT VIRUSES

harmless to African monkeys and for this reason it may be inferred

that the virus has existed in Africa much longer than in the New

World.

Similar examples occur among plant viruses. The virus causing

Pierce’s disease in grapes is probably of New World origin because

its effect on the native American species of Vitis is mild whereas

it is lethal to the introduced V itis vinifera Linnaeus, the grape of

history, which is considered to be native from southeastern Europe

to western India. Also in America is a long list of apparently

symptomless host plants of Pierce’s disease virus, ranging from

grasses to woody shrubs. Moreover, in America are many species

and high populations of “sharpshooters,” the leafhopper vectors

of Pierce’s disease virus. This group of leafhoppers is almost

without representation in Europe.

The recent discovery of a plant virus which causes harm to its

insect vector adds support to the theory of an insect origin of plant

viruses. It provides more conclusive evidence that certain plant

viruses are also animal viruses. Moreover, it supplies a concrete

example of one stage in the postulated evolution of a virus which

until now existed only in theory. It may even raise a doubt, how-

ever microscopic, regarding the validity of one of our oldest

generalizations in virology, namely, that humans and other verte-

brates are immune to injury by plant viruses.

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1952. The place of viruses in nature. Proc. Roy. Soc. 6139:313-326.

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Badami, R. S. and B. Kassanis

1958. In: Report of the Rothamsted Experimental Station for 1957.

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Bennett, C. W. and H. E. Wallace

1938. Relation of the curly top virus to the vector, Eutettix tenellus.

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Best, R. J.

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Bjorling, K. and F. Ossiannilsson

1958. Investigations on individual variations in the virus-transmitting

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THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Black, L. M.

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1950. A plant virus that multiplies in its insect vector. Nature 166:

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Black, L. M. and M. K. Brakke

1952. Multiplication of wound-tumor virus in an insect vector. Phyto-

pathology 42:269-273.

Black, L. M., Sylvia Wolcyrz and R. F. Whitcomb

1958. A vectorless strain of wound-tumor virus. Vllth Internat. Congress

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Burnet, F. M. and P. E. Lind

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5:67-82.

Carter, W.

1939. Populations of Thrips tahaci Lind, with special reference to virus

transmission. Jour. Animal Ecology 8:261—276.

Day, M. F.

1955. The mechanism of the transmission of potato leaf roll virus by

aphids. Austral. Jour. Biol. Sci. 8(4) :498— 513.

Freitag, J. H.

1936. Negative evidence on multiplication of curly-top virus in the beet

leafhopper, Eutettix tenellus. Hilgardia 10(9) :303-342.

1958. Cross-protection tests with three strains of the aster yellows virus

in host plants and in the aster leafhopper. Phytopath. 48(8) :393.

Fukushi, T.

1939. Retention of virus by its insect vectors through several generations.

Proc. Imp. Acad. (Japan). 15(5) : 142-145.

Giddings, N. J.

1950. Some interrelationships of virus strains in sugar-beet curly top.

Phytopathology 40(4) : 377— 388.

Green, R. G.

1935. On the nature of filterable viruses. Science 82:443-445.

Grylls, N. E.

1954. Rugose leaf curl — A new virus disease transovarially transmitted

by the leafhopper Austroagallia torrida. Austral. Jour. Biol. Sci.

7:47-58.

Harrison, B. D.

1958. Studies on the behavior of potato leaf roll and other viruses in the

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6(1) :265-277.

Hewitt, W. B., D. J. Raski and A. C. Goheen

1958. Nematode vector of soil-borne fanleaf virus of grapevines. Phyto-

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April, 1959] JENSEN INSECTS AND PLANT VIRUSES

Rollings, M.

1955. Investigation of Chrysanthemum viruses. I. Aspermy flower dis-

tortion. Ann. Appl. Biol. 43(1) : 86-102.

Jensen, D. D.

1956. Insect transmission of virus between tree and herbaceous plants.

Virology 2:249—260.

1957. Differential transmission of peacb yellow leaf roll virus to peacb

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1951. Benefits to aphids from feeding on galled and virus-infected leaves.

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Kunkel, L. 0.

1955. Cross protection between strains of yellows-type viruses. Advances

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Laidlaw, P. P.

1938. Virus diseases and viruses. Cambridge Univ. Press. Cambridge,

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LiTTAU, V. C. AND K. Maramorosch

1956. Cytological effects of aster-yellows virus on its insect vector.

Virology 2:128-130.

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1952. Direct evidence for the multiplication of aster-yellows virus in its

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1958. Arthropods as vectors and reservoirs of animal pathogenic viruses.

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1945. Evidence of nonspecific transmission of California aster-yellows

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1945. Transmission by insects of a plant virus complex. Nature 155:174.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

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1935. Isolation of a crystalline protein possessing the properties of

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Storey, H. H.

1939. Transmission of plant viruses by insects. Bot. Rev. 5: 240—272.

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1957. Transmission of bean yellow mosaic virus by aphids. Jour. Econ.

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1956. The effect of different host plants of potato virus C in determining

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1957. Studies on the nature of insect-transmission in plant viruses (II).

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1935. Rats, Lice, and History. Boston, Little, Brown and Co.

PYRAMIDOBELA ANGELARUM KEIEER ON ORNAMENTAL

BUDDLEIA IN THE SAN FRANCISCO BAY AREA

(Lepidoptera: Ethmiidae)

Pyramidohela angelarum was described in 1936^ from southern

California. Specimens had been reared in Los Angeles, Santa

Ana, and Santa Paula from various introduced species of Buddleia,

and Keifer presumed that the moth had moved into the area

from a more tropical region. The species was believed to breed

continuously throughout the year, larvae having been collected

in April, July, and October.

In recent years the species has apparently become established

around the San Francisco Bay area, our first record having been

an infestation at San Bruno, San Mateo County in May 1949.

During the last four years adults have been taken at lights in

Berkeley, Alameda County, in October, December, February, and

April (Powell, J. R. Powers, G. 1. Stage) and in Pleasant Hill,

Contra Costa County, in May (P. A. Opler). — A. E. Pritchard

and J. A. Powell, University of California, Berkeley.

’ Keifer, H. H., 1936. Bull. So. Calif. Acad. Sci., 35(1) :13.

April, 1959]

ABBOTT BUTTERFLY MIGRATION

THE 1958 MIGRATION OF THE PAINTED LADY BUTTER-

FLY, VANESSA CARDUI (LINNAEUS), IN CALIFORNIA

(Lepidoptera: Nymphalidae)

Charles H. Abbott

Redondo Beach, California

The most recent migration of the painted lady butterfly,

Vanessa cardui (Linnaeus), was particularly notable for its long

duration, from November, 1957 through May, 1958. Observations

on this migration have yielded rather definite information on

several doubtful points and are the subject of this paper.

The previous reports by the present author on the migration

of this species (Abbott, 1950, 1951), based on the migrations of

1924, 1926, 1941, 1945, and 1949, determined various points

which will be referred to here only incidentally. It became apparent

from these studies that the breeding area of this species, in non-

migration years, is chiefly, if not entirely, south of the California-

Mexico border, probably in both Baja California and Sonora.

Comparatively few of the butterflies are found in California in a

non-migration year, and these few chiefly in the Colorado Desert.

In a migration year the number emerging in these breeding

areas is presumably much greater. One result is a very visible

diurnal migration, in a generally north-northwest direction, cover-

ing southern California and in some years north to the San

Francisco Bay region. Yet the final eggs laid in these regions

where it is an immigrant fail to result in a permanent occupation

of this more northern habitat.

As in the preceding studies, the present report results from

a cooperative study; in this case with the added advantage that

most of the contributors are professional entomologists. Thanks

to the continued interest of R. F. Smith in collecting data for

the project, valuable contributions were secured from the follow-

^ ing members of the staff of the Department of Entomology and

Parasitology, University of California, Berkeley: P. D. Hurd, Jr.,

E, G. Linsley, J. W. MacSwain, R. F. Smith, W. E. Ferguson,

J. A. Powell and from the following members of the Agricultural

Extension Service: G. Beards, J. Dibble, R. Emparin, G. L. Smith,

J. E. Swift. I am also indebted to R. C. Dickson of the Citrus

Experiment Station, University of California, Riverside, and

G. D. Peterson, Jr., Farm Adviser for Imperial County, for

descriptions of the winter and early spring invasion of the

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Colorado Desert, and to C. E. Strickler, Park Supervisor, and D.

E. Merkel, State Park Naturalist, for data for the entire season

from Anza-Borrego State Park. L. M. Martin of the Los Angeles

County Museum has been helpful with both data and criticisms.

Credit for individual contributions will be indicated in the text.

It should be understood that the author alone is responsible for

the conclusions drawn from the data.

The present status of our knowledge of insect migration in

general and of Lepidoptera in particular is summarized in two

recent papers by Williams (1949, 1957). The word “migration”

is defined by Williams (1949) as “a change of location which

is determined both in distance and direction by the insect itself,

and not a passive distribution of individuals by overpowering

forces such as strong winds.”

The present year, 1958, has presented a picture of a longer

continuing migration than any other in recent years. The season

was mild, with early rainfall resulting in extensive desert vegeta-

tion. Butterflies of this species emerged very early in their

normal breeding areas in northern Mexico in such numbers as

to result in migration. Numbers of them had already reached the

southern part of Anza-Borrego State Park by November, 1957

(Merkel), and Imperial Valley by late December (Peterson).

Peterson wrote : “As near as I could determine from actual flight

observations plus population density counts, the butterflies seemed

to be migrating generally from the desert areas south and east

of Imperial County and toward the north and northwest.”

Dickson wrote: “I first noticed the migration of painted lady

butterflies about the last week of January or possibly the week

before that. About that time I heard that automobiles coming

into Holtville from Yuma had the radiators practically clogged

by these butterflies. They were moving out of Sonora, flying

northwest to west, and more particularly numerous near Carpin-

teria. . .

“The painted ladies have laid eggs on suitable host plants

(particularly Cryptantha and Mailva) all over the deserts and

desert valleys. Large numbers of larvae are now (March 17)

feeding on these plants all over the Colorado Desert so that this

area may be a source of a later flight this season.”

The emergence of second generation adults from these meta-

morphosed larvae was recorded in February in Imperial County

April, 1959]

ABBOTT BUTTERFLY MIGRATION

by Peterson, who concluded that they joined the general migra-

tion which had been proceeding without interruption, and in

Borrego State Park by Merkel, who described increasing large

swarms somewhat farther north than the November— December

emergence, but did not observe the actual migration.

The height of the migration in the valleys of southern Cali-

fornia was in late February and early March. Under date of

March 17, Dickson wrote: “There has been some flight through

Riverside the past three weeks, but greater numbers flew through

Beaumont, Redlands, and Colton. Presumably others flew over

the Mojave Desert since I hear that some have reached the San

Joaquin Valley. I did not see any flight this week either on the

desert or in Riverside. This might indicate that the flight is over

for this year, or only that the cold weather has slowed emergence

of new butterflies in their areas of origin.” During this period the

“Redlands Facts” reported that “endless swarms” were flying-

through Redlands.

The “Los Angeles Times” reported on March 5 and 6 that a

“great invasion” had occurred in the Los Angeles area March

2—4. Particular mention was made of Hollywood, the Hollywood

hills, and the San Fernando and San Gabriel Valleys. The author

observed that they were still common at San Fernando, although

not in “millions,” on March 18 and 19. Many of them were

stopping to feed on wild radish flowers in vacant lots, before

continuing their flight to the north-northwest.

The western edge of this February— March migration was

recorded by the author at Redondo Beach. Individual V. cardui

were observed on garden flowers on January 4, 16, February 5,

19, 26, 27, March 9, 29; actual migrating butterflies were ob-

served February 21, daily March 1—6, 11, 16. The migration was

steady March 2—4, one or two per minute, the same days as the

maximum in Los Angeles.

After an interval in late March and early April, which was

marked by cold and rainy weather, a heavy migration wave

occurred in the middle of April in the San Joaquin and Salinas

Valleys. Summarized in brief:

San Joaquin Valley. — Heavy migration throughout valley from

south and east of Bakersfield to the region west and north of

Tracy. First flight recorded April 8, at height April 10, 11, began

to decline April 17, moderate flight April 20, a few stragglers

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO, 2

April 23 (R. F. Smith, Hurd, Linsley, MacSwain, Ferguson).

Stragglers to May 9 (R. F. Smith, Swift).

On April 13 they were abundant near Mt. Diablo (R. F.

Smith). On April 20 there was a heavy flight in the Walnut

Creek region (R. F. Smith, Swift).

Salinas Valley, April 17. — Migration reported (Dibble, Em-

parin) .

The author made one observation of three migrants at

Redondo Beach on April 19, but has no other report of this third

generation from the Los Angeles area.

Larvae, presumably from eggs laid by these April migrants,

were so abundant in the San Joaquin and Salinas Valleys from

about April 20 to May 9 that they were found not only on wild

plants, but damaging alfalfa, lettuce, and other crops (R. F. Smith,

Swift) . They were also abundant in clover fields in Glenn County,

farther north (reported by R. F. Smith). At Shafter, near the

southern end of San Joaquin Valley, the larvae were abundant

from about April 10, and in some cases caused enough damage

to cotton fields to require control measures (G. L. Smith, Beards).

The above description shows an apparent pattern of a built-up

migration through three generations rapidly succeeding each

Other. Most published descriptions of former spring migrations

of this species have assumed that the migrating adults were all of

one generation and in general from a common source. This has

applied not only to migrations in California and Utah, but also

to those in Europe. Campbell (1924) in California and Wood-

bury et al. (1942) in Utah described egg-laying along the way,

with the possibility that a new generation from these eggs took

part in a later migration the same year. It has, however, seemed

more usual that the larvae from these eggs either did not develop

to the adult stage, or, because of the smaller numbers, their

presence or possible later flight went unnoticed, except in rare

instances.

The interpretation of numbers of generations involved is

complicated by the apparent great variation in the length of the

egg, larva, and pupa stages. L. M. Martin, after checking the

literature, wrote me (June 19, 1958) : “As far as I can find out,

there are no figures on the length of these stages. We know that

they vary, for instance, in the egg stage from 7—60 days. The

April, 1959]

ABBOTT — BUTTERFLY MIGRATION

larval stage varies from 13-41 days and the pupae have been

known to stay in that stage for eight months.”

The most distinct three generation pattern is found in the

records of D. E. Merkel, State Park Naturalist at the Anza-Borrego

State Park. As a background, Merkel notes that in the period

1956—57, prior to November 1, 1957, no painted ladies were

observed in the park. “If any were present, they were in such

small numbers as not to attract attention.” This appears to

eliminate the Park as part of the breeding area in non-migration

years.

As the earliest record of the 1958 migration, large swarms

were first noted in November, 1957, in the Mud Hills region in

the southern part of the Park. They had apparently emerged

farther south, probably in northern Mexico, and had flown north

to this secondary breeding area. The subsequent history is as

follows: January, 1958, in same Mud Hills region, many pupal

casts. February, 1958, at Borrego Campground area, about 25

miles farther north, increasing large swarms of adults. March,

1958. Larvae numerous throughout Borrego Valley, some begin-

ning to pupate. April, 1958. Adults numerous in valley. Empty

pupal cases abundant, probably of this species (also noted by

Ferguson) .

The above summary shows at least three generations. The

November generation probably came from last year’s eggs in

the normal breeding area in northern Mexico, following a long

pupal stage during the summer and fall. The second began to

emerge in late January, reaching a maximum in February, and

the third began to emerge in late March, reaching a maximum

in April.

The first and second of these generations were also observed

by Peterson in Imperial Valley. He recorded a steady migration

from about December 20, lasting six or eight weeks, correspond-

ing to the November and February periods of adult abundance

reported by Merkel. Peterson thinks it is probable that two

generations of larvae were abundant on desert hosts (mainly

Cryptantha spp.) during this period, although, because of con-

tinuous migration, he could not separate them. “There may have

been an extremely small third generation in the cultivated area of

Imperial Valley, but this is in doubt.” The decrease in numbers

could have been caused by most of the second generation adults

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

migrating before egg laying, or by a drying up of the food-

plants of the second generation larvae, or, as Peterson suggests,

parasitism of the larvae may slow down the numbers of each

successive generation.

The timing suggests that these second generation migrants are

those reported as numerous through the San Gorgonio Pass and

San Bernardino Valley regions in February and which were

numerous in the Los Angeles region in early March. These reached

the San Joaquin Valley, laid eggs, and from the eggs came the

very abundant migrants observed throughout the valley in the

middle of April. (Linsley and MacBride noted that these migrants

seemed to be nearly all fresh specimens). Also belonging to this

same third generation were those reported from Salinas Valley

April 17, Mt. Diablo April 13, and Walnut Creek April 20.

It is not so easy to interpret the observations made in the

desert in April, but the time element is again used. The names

of the observers are given, the assignment to generations is by

the author.

Colorado Desert, 18 miles west of Blythe, April 14—18. — Enor-

mous numbers of hatched and parasitized pupae (eggs laid by

second generation). Few adults left (Hurd, Powell). Most of the

adults which had emerged must have flown north-northwest (third

generation) .

Borrego Desert, April 11—18. — Emerged pupae (eggs laid by

second generation) abundant. Adults abundant, but not seen to be

migrating (third generation) (Ferguson, Merkel).

Eastern Mojave Desert, April 12, near Amboy. — Pupae abun-

dant, not emerged (eggs laid by second generation). Some worn

adults, which had probably flown from farther south. A few adults

near Topock, Arizona April 14 (Linsley, MacSwain).

Western Mojave Desert, April 12, 13, 14, 19. — Mature larvae

abundant (eggs laid by second generation). Some adults migra-

ting (third generation) (Hurd, Powell, Linsley, MacSwain).

These interpretations as to the principal generations involved

do not eliminate the probability that some members of each

generation flew farther than the ones here described, either before

depositing eggs or afterwards. The timing best fits the basic inter-

pretation here given, but there is probability of an overlapping

of generations in each locality.

The larvae from eggs laid by the third generation were those

April, 1959]

ABBOTT BUTTERFLY MIGRATION

found so abundant in the San Joaquin and Salinas Valleys from

about April 20 to May 9 that they were transferring from weeds

to vegetables. Larvae found at the same time in Glenn County

showed that adults of the third generation had reached there.

This appears to show that it was the third generation larvae which

were in such numbers as to be considered destructive, although

it may be because these reports were from regions which have

extensive field crops.

Reports of a fourth generation and on the final extent of the

1958 migration are fragmentary. J. A. and F. C. Powell collected

V. cardui in Siskiyou and Del Norte Counties on seven scattered

dates between June 23 and August 29, while G. Pitman found

them on September 12, also in Siskiyou County. Specimens

appeared fresh on June 24, 26, July 9, 18, August 4. This suggests

continuation of the migration to the Oregon border and at least

a fourth generation.

Based on preceding years of observation, it may be predicted

that there will be no increase in resident population of V . cardui

in 1959 in the regions invaded. The population control is dis-

cussed below under parasitism of larvae.

The butterfly Vanessa cardui (Linnaeus) received its specific

name from the thistle, and is commonly thought to lay its eggs

chiefly on thistles. But in California, where thistles are not

particularly abundant, other plants appear to be just as acceptable

as food plants for the larvae. This year’s observations emphasize

a number of these, as follows:

Colorado Desert: Cryptantha spp. (Peterson).

Colorado Desert: Cryptantha, Malva (Dickson).

Mojave Desert: Cryptantha, Amsinckia, Phacelia (Hurd, Powell).

Mojave Desert: Cryptantha, lupine, mallow (Linsley, MacSwain).

San Joaquin Valley: Amsinckia. Alfalfa, lettuce, and other crops

( Swift) .

San Joaquin Valley: Malva, Amsinckia, sunflowers, weeds, cotton

(G. L. Smith, Beards).

Glenn County: Yellow star thistle, Amsinckia, dock (reported by

R. F. Smith).

Salinas Valley: Nettle, lettuce (reported by R. F. Smith).

This shows that a considerable variety of wild plants are

utilized and that in a year of special abundance, cultivated crops

may be attacked. This favors large succeeding generations in the

same year and continuation of migration.

The causes of a population increase resulting in a heavy

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

migration have already been enumerated to include a mild winter,

a large amount of rainfall, and consequent abundant desert vege-

tation. In addition, in 1958, an unusually long favorable season

has resulted in a build-up of population through at least three

generations in one spring, with migration occurring in all of them.

Yet it is almost an axiom that such a migration does not result

in colonizing new areas or in increasing the permanent population

in the areas reached. Possible checks are too rigorous a winter

climate, effective toward the north, drying up of the food plants

on which eggs are laid, effective toward the south and in arid

climates, and control by parasitic attack. This last point will be

discussed further.

Elton ( 1927 ) , after extensive studies of population control,

concluded that the most effective end of an outbreak or plague,

whether resulting in a migration or not, is caused by some form

of parasitic attack.

Schrader (1928) emphasized that V anassa cardui (L.) is

subject to the attacks of “a considerable number of parasitic

wasps and flies.” Essig (1926) in his section on Tachinid flies,

names three species, Zenillia blanda, Frontina archippivora, and

Chaetogaedia monticola, which have been reared from larvae of

V. cardui. This statement in no way limits the number of possible

parasites.

The prevalence of parasitism is shown by several of this year’s

reports, as follows.

“Adult emergence within (Imperial County) was much less

than would appear to be indicated by the larval populations

present on the desert. This was undoubtedly due to the high rate

of parasitization by both insects (Tachinidae) and entomoph-

thorous fungi.” (Letter from G. D. Peterson, Jr., May 19, 1958.)

“At Topock, Arizona, on April 14 we observed relatively few

butterflies but enormous numbers of Chalcid wasps which had

apparently emerged in that area. These wasps were congregated

around a creosote bush and several thousand were taken in a

single swing of the net.” (Letter from J. W. MacSwain, May 23,

1958.) The possibility is suggested that many of these may have

emerged from V. cardui larvae and may account for the small

number of adults observed.

“. . . enormous numbers of hatched and parasitized pupae

present at the 18 miles west of Blythe locality in late April. The

April, 1959]

ABBOTT BUTTERFLY MIGRATION

few adults present could be stragglers, late emergences, etc., which

had not gone farther north yet.” (Letter from J. A. Powell, May

19, 1958.)

The fact that the direction of flight of migrating V. curdui

through California is mostly toward the north-northwest and

directly against the prevailing wind has been generally observed

in all the migrations on record. In the 1958 reports, this was

emphasized by Dickson and Peterson in the Colorado Desert, by

Dickson in describing the course taken through southern Cali-

fornia, by Linsley, MacSwain, and Ferguson in the San Joaquin

Valley, and by R. F. Smith and Swift in the Walnut Creek region.

The most important variant was in the San Joaquin Valley on

April 20, when the majority were flying northwest near Merced,

mostly west Merced to Tracy, and mostly north west of Tracy

(Linsley and MacSwain). It was not reported whether there was

any difference in the wind direction between the regions just

enumerated.

The author made a few observations on the direction of flight

of 1958 migrants at Redondo Beach, about 1000 feet back from

the ocean front. As the migrants came over singly, it was possible

to follow the flight of individual butterflies. Variable winds are

normal at Redondo Beach. The beach faces the west and the fair

weather winds vary from northwest to southwest and on any day

constantly fluctuate. The early morning winds are usually north-

east, and on days of desert winds there may be northeast gusts

all day. Because of the very small numbers of butterflies recorded,

any conclusions as to the effect of direction of wind on direction

of flight are very tentative. There was an observed tendency of

butterflies to fly into the northeast wind in the morning (observed

on five mornings at varying times from 7:45 to 11:20), and for

the flight to be chiefly northwest, but varying slightly toward the

north, during the afternoon. The limited observations support

the former observations that flight, whether individual or mass,

is usually straight into the wind, with little individual variation

(Abbott, 1951).

A survey of collected information on the migration of Vanessa

cardui (L.) in North America points to its classification as a

unidirectional flight with no return (Williams, 1930, 1938, Abbott,

1951). Study of it in California has the special advantage that

the compass direction of flight is clearly seen, the same prevailing

direction occurs in every migration year, and the numbers in-

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

volved are great enough to insure the accuracy of these two

statements.

The fact that years of general migration occur at intervals,

four years more or less, supports the theory that migration occurs

only when a population has been built up to what Chapman

(1939) calls “outbreak proportions.” Chapman describes this as

resulting from a relaxation of environmental resistance. This

might occur and take effect all in one generation, or it could be

cumulative in result, reaching outbreak proportions after a series

of generations. The occurrence of migrations at irregular intervals

of years favors the cumulative interpretation, whereas a great

migration in a year which is particularly favorable from an

environmental standpoint indicates that it might occur in one

generation.

Observations on the 1958 migration suggest another kind of

cumulative result, namely, pressure of population and resulting

migration continued through three generations, all of them devel-

oped under exceptionally favorable conditions. It may also be

emphasized that larvae of the third generation were so numerous

as to be destructive of cultivated crops.

In the reports of former years (Abbott, 1950, 1951) it was

noted that the migration was in a series of waves of abundance,

particularly in 1926 and 1941. It was interpreted that the flight

was periodically slowed down by unfavorable weather. No data

were available to show a succession of generations. Both factors

were apparently involved in the 1958 season. The weather might

affect metamorphosis of larvae as well as flight of adults.

Migration is only one of the results of an excessive increase

in population, and why it affects only certain species has not

been determined. It may be mentioned that Williams, after collect-

ing all available information on migration of Lepidoptera, con-

cludes that there are many more migratory species than the most

abundant and conspicuous ones which are well known. He states

(Williams, 1949) : “We have evidence today of migration in two

or three hundred species of butterflies from almost all parts of

the world. In about fifty of these there is a considerable weight

of evidence which makes it almost impossible to accept any other

explanation.” In another reference (Williams, 1957) he states

that 20 or 30 migrant species are known from the United States.

Summary

(1) The migration of the painted lady butterfly, Vanessa

April, 1959]

ABBOTT BUTTERFLY MIGRATION

cardui (L.), in California in the spring of 1958 was exceptional

in its duration from November, 1957 to May, 1958, as well as in

the great numbers of individual butterflies involved.

(2) The north-northwestward distribution during the period

was gradual and was correlated with the development of three

generations. The first generation invaded the Colorado Desert

from northern Mexico from November, 1957 to January, 1958.

The second generation emerged on the Colorado Desert from

eggs laid by the first generation and flew north in February and

early March through the San Bernardino Valley and the Los

Angeles area to the Mojave Desert and the San Joaquin Valley.

The third generation, emerging in the two last named regions,

but chiefly in the San Joaquin Valley, resulted in a great flight

through the San Joaquin and Salinas Valleys in the middle of

April. The large number of larvae resulting from this third wave

of the flight attacked various field crops.

(3) The continuance of a migatory flight through successive

generations is favored by the fact that several widely distributed

food plants serve as food for the larvae. This applies to both wild

and cultivated plants.

(4) Extensive parasitism during the larval and pupal stages

was noted by several observers in both the Colorado and Mojave

Deserts. This furnished evidence toward the theory of Elton

(1927) that parasitic attack is the most important control of an

insect outbreak, whether or not accompanied by migration.

Literature Cited

Abbott, C. H.

1950. Twenty-five years of migration of the painted lady butterfly,

Vanessa cardui, in southern California. Pan-Pac. Ent. 26(4) ;

161-172; 1 fig.

1951. A quantitative study of the migration of the painted lady butter-

fly, Vanessa cardui L. Ecology 32(2) :155— 171 ; 4 figs., 5 tables.

Campbell, R. E. :

1924. Painted lady butterfly. Insect Pest Survey Bull. (Washington)

4:70-71.

Chapman, R. N.

1939. Insect population problems in relation to insect outbreak. Ecol.

Monogr. (Duke Univ. Press) 9(3) :261— 269.

Elton, C.

1927. Animal Ecology, xxi -j- 207 pp. N.Y. Macmillan Co.

Essig, E. O.

1926. Insects of western North America, xi 1035 pp. N.Y. Macmillan

Co.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Schrader, W.

1928. Experiments on a species of migrating butterfly. Bull. So. Calif.

Acad. S'ci. 27(2) :67-70; 1 pi.

Williams, C. B.

1930. The migration of butterflies, xi -|- 473 pp. Edinburgh, Oliver and

Boyd.

1938. Recent progress in the study of some North American migrant

butterflies. Ann. Ent. Soc. Amer. 31 (2) :211-239 ; 7 figs.

1949. Migration in Lepidoptera and the problem of orientation. Proc.

Roy. Ent. Soc. (c) 13:70-84; 1 fig.

1957. Insect migration. Ann. Rev. Ent. 2:163—180.

Woodbury, A. M., J. W. Sucden, and C. Gillette

1942. Notes on migrations of the painted lady butterfly in 1941. Pan-

Pac. Ent. 18(4) : 165-176; 1 fig.

TWO NEW RECORDS EOR THE CONE BEETLE GENUS

CONOPHTHORUS HOPKINS IN CALIFORNIA

(Coleoptera: Scolytidae)

During the summer of 1956, a collection of cones of the

western white pine, Pinus monticola Douglas was made in the

Lassen Volcanic National Park area, Lassen County. Two speci-

mens of Conophthorus monticolae Hopkins were reared from

one of the cones. This species of cone beetle has been previously

known only from the cones of the western white pine in Idaho.

Washington and Canada (Keen 1958)^. During the same sum-

mer and following summers, collections of cones of the lodge-

pole pine, Pinus contorta var. latifolia S. Wats were made from

several locations in the Sierra Nevada mountains. In Calaveras

and Tuolumne Counties, the cones of this pine were found to

be heavily infested with Conophthorus contortae Hopkins. Until

now, this species has only been recorded from the cones of the

shore pine, Pinus contorta var. contorta S. Wats near Newport,

Oregon (Keen 1958). The author is indebted to Dr. S. L.

Wood of Brigham Young University in Provo, Utah for the

identification of these beetles. — Herbert Ruckes, Jr., University

of California, Berkeley.^

1 Keen, F. P., 1958 — Cone and Seed Insects of western forest trees. U.S.D.A. Technical

Bulletin No. 1169. pp. 48 & 55.

2 Studies of the cone and seed insect problems of the pines in California made possible by a

grant from the Gilbert M. Walker Fund.

April, 1959]

SCHUSTER — MALE PSELAPHID

NOTES ON MORIUS OCCIDENS CASEY WITH A

DESCRIPTION OF THE MALE

(Coleoptera: Pselaphidae)

Robert 0. Schuster

University of California, Davis

On the basis of a unique female, broken into many parts,

Thomas L. Casey described Morius occidens in 1893. Casey's

description is to be found in Bowman’s Pselaphidae of North

America and the genus is keyed into the North American

pselaphid fauna by Park ( 1953 ) . The male has not previously

been described.

The genera Morius and Rhinoscepsis are the only North

American representatives of the subtribe Rhinoscepsina, tribe

Euplectini. Morius was previously known only from the type

collected in Santa Cruz County, California.

What appears to be one species of Morius has been recovered

at a low frequency from leaf mold in the Coast Range from

Monterey to Mendocino Counties in California. The number of

specimens in the series now available for study is insufficient to

determine the nature of variation, mainly color and degree of

pubescence. The male is associated with Casey’s female because

(1) it is the predominant form recovered in Santa Cruz County,

(2) it is of the proper size and color, and (3) it agrees with the

type except for the male sexual characters. (Comparison with

the type, which I have not seen, was made by Gordon A. Marsh.)

Casey’s original description was good, and only a few char-

acters, mainly of the mouth parts of the female, and those pertain-

ing to the male are described here.

Morius occidens Casey

Additions to description of female. Head as in Casey’s description but

with weak median carina extending from neck to about the vertexal foveae.

Labrum straight across the front, rounded at lateral margins; 2 cone-shaped

structures occur medially on lower front margin and 5 or 6 in a row

behind the first 2; 4 macrosetae arise laterally, 1 on the dorsal surface

and 3 ventrally. Mandibles heavy, 6 teeth on each inner ramus; 1 lamellate

seta occurs midway on the dorsal surface along with a few smaller setae.

Maxillary palpus with segment I small, 1/3 length of II; II thin for basal

1/2 and swollen at distal 1/2; III slightly more than 1/3 length of II and

equally as wide; IV 3 times as long as III and noticably wider, terminated

by a long, thin palpal cone almost 1/2 as long as segment IV ; 2 specialized

setae occur midway on outer surface. Abdomen of 5 visible tergites and

6 visible sternites. Tergite I with fovea formed by 2 apodemes at each

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Explanation of Figures

Morius occidens: Fig. 1, adult male, dorsal aspect; Fig. 2, maxillary

palpus; Fig. 3, aedeagus, dorsal aspect.

April, 1959]

SCHUSTER — MALE PSELAPHID

anterio-lateral margin and a large pubescent excavation occupying the

median 1/3; II with a much smaller depression at the anterior margin;

other tergites simple; sternite I entire behind coxal lines although this

condition is obscured by pubescence on point-mounted specimens; II 3 times

as long as I with a deep, pubescent, transverse sulcus at the front margin.

Mesosternum with 2 carinae and 3 anterior pubescent foveae; mesocoxae

narrowly separated, the coxal cavities confluent but nearly closed by pro-

cesses of the meso- and metasterna; a large pubescent fovea occurs laterad

and one posterior to each coxa; metacoxae contiguous.

Male . — As in the female with the exception of the abdominal tergites.

Brachypterous males have been noted and fully winged populations may

eventually be found. Six sternites are readily visible, and a seventh, a

minute penal plate strongly attached to a genital segment and usually

removed with the aedeagus during dissection, is sclerotized distally and

may at times be visible externally. A small, semi-triangular projection occurs

at the middle of the posterior margin of the third visible sternite. Male

aedeagus 0.23 mm. long X 0.15 mm. wide; dorsal surface of capsule

membranous, ventral surface mostly so, depending to some extent on the

age of the individual; two parameres ventrally, extending to rear of

capsule and bridged by wide connection at about 1/2 the length of the

aedeagus; a membranous tube, apically bifid, is weakly sclerotized near

the apex of each paramere and again at the terminal dichotomy; numerous

small pores are present along the apical margin of the parameres and a

number of fine setae occur along their lateral margins.

Literature Cited

Bowman, John R.

1934. The Pselaphidae of North America. Pittsburgh (privately pub-

lished), pp. 1—149.

Casey, Thomas L.

1893. Coleop. Notices V. Ann. N.Y. Acad. Sci. 7 :433— 509.

Park, Orlando

1953. Discrimination of genera of pselaphid beetles of the United

States. Chicago Acad. Sci. Bull. 9:299—331, pis. 1—5.

FOURTH ANNUAL INSECT PHOTOGRAPHIC SALON

The Pacific Branch of the Entomological Society of America

announces the fourth annual insect photographic salon to be held

in conjunction with its meeting in Sacramento, June 23—25, 1959.

Photographs will be judged in three categories: monochrome,

color transparency, and color transparency sequence. Deadline for

all entries is June 16, 1959. Entry blanks may be obtained from

Dr. E. S. Ross, California Academy of Sciences, Golden Gate Park,

San Francisco 18, California.

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO, 2

A FEW ODONATA FROM YOSEMITE

Edward J. Kormondy

Department of Zoology, Oberlin College

A small collection of dragonflies taken in Yosemite National

Park (Tuolumne County, California) and deposited in the Uni-

versity of Michigan Museum of Zoology has been determined by

the author. The series was taken July 10—13, 1939, by fifteen

children on a “nature hike,” under the direction of C. L. Gibson,

who gave the specimens to the Museum. The locales represented

are Miguel Meadows and Sand Pit Lake in northwestern Yosemite,

about two miles southeast of Lake Eleanor at an elevation of

5200 feet. According to Chief Park Ranger Elmer Fladmark (in

litt., December 9, 1958) there is no “Sand Pit” Lake at Yosemite

and the excavation currently known as Gravel Lake may have had

the former name at one time.

In the only collective report on California Odonata to date,

Kennedy (1917) listed ten species as occurring in Yosemite:

Lestes dryas Kirby, Enallagma cyathigerum (Charpentier) , Amphiag-

rion abbreviatum (Selys), Aeshna interrupta nevadensis Walker, Cordulia

shurteffi S'cudder, Libellula pulchella Drury, Libellula quadrimaculata

Linnaeus, Sympetrum semicinctum (Say), Leucorrhinia glacialis Hagen,

Leucorrhinia hudsonica (Selys).

The following list of fifteen species, none of which constitutes

a new state record, includes eleven (indicated by an asterisk)

reported from Yosemite for the first time (Sand Pit Lake and

Miguel Meadows are abbreviated as SPL and MM respectively) :

Lestes dryas Kirby, 5 $ $ , 9 9 9, SPL and MM ; Enallagma cyathi-

gerum (Charpentier), 1^, SPL; Isclinura perparva Selys*, 3$ $, 29 9,

SPL and MM; Atiax junius (Drury)*, 2$ $, SPL; Aeshna palmata

Hagen*, 1^, MM; Aeshna multicolor Hagen*, SPL; lAbellula pulch-

ella Drury, 10 ^ ^ , 19, SPL and MM; Libellula quadrimaculata Lin-

naeus, 12 $ S , MM; Libellula saturata Uhler*, 11 ^ ^ , 19, SPL;

Sympetrum corruptum (Hagen)*, 19, SPL; Sympetrum madidum

(Hagen)*, 2$ $, 19, SPL and MM; Sympetrum occidentale californicum

Walker*, 19, MM; Sympetrum pallipes (Hagen)*, 1^, 29 9, SPL and

MM; Leucorrhinia intacta (Hagen)*, 1^, 19, MM; Tramea lacerata

Hagen*, 3$ SPL.

Literature Cited

Kennedy, C. H.

1917. Notes on the Life History and Ecology of the Dragonflies

(Odonata) of Central California and Nevada. Proc. U.S. Nat’l

Mus. 52:483-635.

April, 1959]

SMITH JAPYGIDAE I

THE JAPYGIDAE OE NORTH AMERICA 1—

PROVALLJAPYGINAE AND NANOJAPYX

Leslie M. Smith

University of California, Davis

The subfamily Evalljapyginae is characterized by the five

pectinate laminae of the lacinia and the bilateral asymmetry of

the forceps, as well as other characters. Silvestri (1949) studied

three specimens from the state of Sao Paulo, Brazil and noted

that they possessed a falciform distal lamina on the lacinia and

nearly bilaterally symmetrical forceps. He named this insect

Provalljapyx lanei new genus and new species and placed it in

the subfamily Evalljapyginae although it did not fit in the two

characters mentioned. Paclt (1957) placed Silvestri’s genus

Provalljapyx as a synonym of Evalljapyx but, as indicated below,

this is not justifiable.

In my collection are 63 specimens representing five species

which I assign to the new genus Nanojapyx. This genus from

California shows affiliation to Silvestri’s genus Provalljapyx from

Brazil. Neither of these genera can be placed logically in the

subfamily Evalljapyginae. In my opinion they represent an

evolutionary group between the Projapygidae and the Evalljapy-

ginae, and I therefore place them in the new subfamily, Provall-

japyginae. All of the Projapygidae known from the New World

have plumose body setae and bilaterally symmetrical anal cerci

which do not show sexual dimorphism. We may assume that the

Projapygidae arose from a primitive campodeid by the shortening

and thickening of the anal cerci, and later by the addition of

stout spines mesad on the cerci as a defensive mechanism for

protection against attack from the rear.

A further step in the evolution is represented by the Provall-

japyginae wherein the anal cerci are modified into unsegmented

forceps which are symmetrical and do not show sexual dimorph-

ism. The final evolutionary step is exhibited by the Evalljapyginae

wherein in the left forcep is different than the right forcep and the

forceps of the male are different from those of the female. This

line of evolution is also supported by anatomical features other

than the forceps.

Pro vail japyginae, new subfamily

Similar to the Evallj apginae except: antennae with 20 to

23 segments, distal lamina of lacinia falciform, styli each with a

100

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

single seta, tergite VII slightly lobed or bilobed, pleura VII not

heavily sclerotized or projected to the rear, forceps approximately

similar and without sexual dimorphism, length of body 2 to 4

mm.

Inasmuch as this subfamily is now known from eastern South

America and western North America, it seems probable that a

rich fauna exists in this group. They have undoubtedly been

overlooked by earlier collectors because of their small size and,

if seen, they were probably mistaken as nymphal stages of larger

species.

Key to the Genera

1. Stylus with a single seta, distal lamina of lacinia falciform, forceps

similar (Pro vail japyginae) 2

— Stylus with two setate, distal lamina of lacinia pectinate, forceps

not bilaterally symmetrical (Evall japyginae) 3

2. Two large median teeth on each forcep, no other teeth or denticles,

posterior seta on pleuron plumose, segment X tergite setae

4 -|- 1 + 4 Provalljapyx

— Four or more similar teeth on each forcep, posterior seta on pleuron

simple, segment X tergite plumose setae 3 -|~ 3 Nanojapyx

3. Setose frontal sinus Ctenjapyx

— Without setose frontal sinus Evalljapyx

Nanojapyx L. Smith, new genus

Type species: Nanojapyx paged L. Smith, new species.

Head. — Antenna with 22 ± 1 segments; vertex covered with a number

of and a lesser number of m ; distal half of antenna with segments

showing two fairly regular rows of m, a a little more to the base than to

the extremity of the segment, longer than m on the same segment; apical

lamina of lacinia falciform, shorter than the other four, pectinate laminae.

Thorax. — Pronotum 6 -{- 6 M, meso- and metanotum 7 to 9 7 to 9 M ;

legs, femora dorsal apex with two subequal small usually plumose setae

and one simple seta in a line; tarsus with two ventral rows of spine-like

setae, two or three setae per row. Abdomen. — Tergites with four to five

irregular transverse rows of M, and fewer m, M with 6 ± 2 pinnulae;

sternites with five irregular transverse rows of M and fewer m, M with

3 ± 2 pinnulae; tergite X with 3 3 plumose M between the carinae and

a variable number of m ; lateral subcoxal organs broad with about four

to seven plumose sensory setae each, in a single row, and two or three

irregular rows of distinct glandular setae; styli on segments I— VII, each

with one mesad seta, approximately one-third as long as the stylus ; abdominal

pleura ; anterior pleurite with two large setae, with the anterior one plumose ;

1 Abbreviations : M — • macrosetae, plumose; m — sub-macrosetae, usually simple but occa-

sionally plumose, a — dorsal sense-seta on the fourth antennal segment; LI, L 2, L 3, —

first, second, and third pair of legs; A, B, C, D — anterior, second, third, and posterior rows

of M on the abdominal sclerites; I-X — abdominal segments.

April, 1959]

SMITH — JAPYGIDAE I

101

pleuron with three large setae, posterior one simple; forceps apparently

symmetrical, differing only in number of teeth, teeth usually sharp pointed,

not rounded denticles; all setae on forceps simple, except one or two basal;

no sexual dimorphism in forceps.

This genus is close to the genus Provalljapyx but can be

distinguished from it by the presence of four or more teeth on

each forcep.

Nanojapyx pages! L. Smith, new species

Female. — Head: antenna with 22 segments; segments 13 to 22 with

all setae of same size, arranged in two transverse rows, about 16 setae in

distal circle, and about 20 setae in basal cricle; segment IV circled with

eight to ten larger setae anterior to a; labrum with 7 7 simple setae

of various sizes; mandible with three fused teeth and a fourth, distinct,

less heavily sclerotized, pointed tooth; first tooth (ventral) of mandible

largest and recurved at tip; galea of maxilla with sclerotized thumb at

apex of anterior lobe with five spatulate projections at tip of thumb; palpus

with ten simple setae; lacinia heavily sclerotized, arcuate, lamina 1 a rod,

shorter than lamina 2; laminae 2 to 5, typical combs; labium with numer-

ous simple setae; labial palpi short, conical, with two long and one short

simple setae; dorsum of head with about 24 -|- 24 setae, mostly all plumose.

Thorax. — Pronotum with 6 + 6 M and 5 + 4 m ; mesonotum, prescutum

with 1 -f 1 M and 1 + 1 m ; scutum with 7 + 8 M and 7 + 7 m ; metano-

tum with 8 + 9 M; legs short, setae at dorsal apex of femur, numbers 1

and 2 plumose, number 3 simple, number 3 longer than number 1, number 2

shortest; large ventral setae on tarsus in two rows of two setae each, tarsal

claws simple sub-equal. Abdomen . — Tergite I: prescutum with 1 + 1 M and

1 + Im, scutum with 6 + 6 M and 6 + 6 m; sternite I — prescutum with 4 +

5 M, and no m, scutum Ar=4 + 4M, B — 2+ 2 M and 2 + 2 m, C = 4 +

4 M, D =: 9 + 9M ; lateral subcoxal organs with 7 + 7 M and numerous

glandular setae arranged in two irregular rows; stylus simple, conical with

slightly bulging base, one simple curved seta; tergite II, A — 4 + 4 M, B — 2

+ 2 M, C = 2 M and 3 + 3 m alternating in row with M; segments III to

VII inclusive similar to segment 2; six ventral setae associated with stylus

VII, as illustrated, setae 1, 3, and 5 with 3 or 4 pinnulae, setae 2 and 4

present. Segment VIII width 0.16 mm, length 0.18 mm; tergite, A = 4 + 4

M and 1 + 1 m, Br=2+2M and 1+ 1 m, C=:3 + 3M and 3 + 3 m

alternating in row with M ; sternite, A = 3 + 3 M and 1 + 1 m, B

3 + 3 M and 1 + lm, C = 3+ 3M and 3 + 3 m ; spermatheca between

B and C, bulbous with a long anterior spike; genital opening with many

simple setae, no palps. Segment IX tergite, 3 + 3 M and 3 + 3 m alterna-

ting; sternite with 3 + 3 M and 3 + 3 m; segment X, width 0.14 mm,

length 0.20 mm; tergite, (between crenulae) A = 2 + 2M, B = 1 + 1M,

C = 3 + 3 M of which the four mesad setae are simple, about six m

scattered over the tergite; sternite A = 3 + 3 M followed by a row 2 + 2

m, B = 2 + 2 M and 1 + 1 m, C=:1 + 1M, D — 3 + 3M and 2 + 2

m; acropygidium distinct, rounded, edge crenulate; carinae distinct;

forceps short and broad (as illustrated) and strongly bent upward toward

102

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

the tips; right forcep with seven sharp, recurved teeth, left forcep with

five similar teeth, each forcep with ten large setae of which the outer,

basal one is plumose; and nine smaller setae of which four arise near the

dental margin; length of forcep 0.14 mm; length of body, including

forceps 2.89 mm.

Male. — Similar to female except no spermathecae ; large median internal

setose sac opening ventrally on scutum III near the suture between the

prescutum and scutum, tip of sac extending anteriorly to posterior margin

of second urosternite, setae near the mouth of the sac plumose, the rest

simple; lateral subcoxal organs with 6-1-6 plumose sense-setae and

numerous hyaline glandular setae almost as long as the sense setae; right

forcep with six teeth, left forcep with five teeth ; length of body 2.72 mm.

Holotype female and allotype male (California Academy of

Sciences) and paratype female and male (University of California,

Davis) were collected in leaf mold near Oakland, Alameda

County, California, in January, 1953 by Mr. Robert 0.

Schuster.

I take pleasure in naming this species after M. L. Pages of

the Labor atoire de Biologie Animale, Universite de Dijon, France,

in recognition of his fine work in the taxonomy of the Japygidae.

Nanojapyx pricei L. Smith, new species

Female. — Similar to N. pagesi L. Smith except setae at dorsal apex

of femora, number 3 equal in length to number 1 ; setae on labial palp

4; meso-and metanotum 8 -f- 8 M; setae number 1 on forceps usually simple

or minutely plumose; length of forceps 0.18 mm; right forcep with seven

teeth, left with six teeth ; tips of forceps strongly curved inward, as

illustrated; lateral subcoxal organs each with six or seven plumose sense

setae; ventral setae associated with stylus VII as in pagesi but with three

pinnulae each ; abdominal segment X, width 0.20 mm, length 0.28 mm ;

length of body, including forceps 3.83 mm.

Explanation of Figures

Fig. 1, lacinia of Nanojapyx pagesi L. Smith showing rod-like structure

of the first lamina, e = 0.05 mm; fig. 2, male sac in III of pagesi dorsal

view, e n: 0.05 mm ; fig. 3. spermathecae of pagesi dorsal view ; fig. 4,

eighteenth antennal segment of pagesi female, dorsal view, e = 0.05 mm ;

fig. 5, apex of femur of pagesi dorsal view showing plumose setae 1 and 2,

and simple seta number 3 ; fig. 6, tarsus of L 2 of pagesi ventral view show-

ing two rows of two stout setae each, e = 0.05 mm ; fig. 7, fourth antennal

segment of pagesi female, dorsal view showing dorsal sense seta at a;

fig. 8, labial palpus of pagesi female, ventral view; fig. 9, pleuron of left

side of pagesi female showing setal pattern, left = pleurite, right —

pleuron; fig. 10, tergite X of pagesi female, dorsal view, e 0.10 mm;

fig. 11, right stylus VII and associated setae of pagesi, ventral view, e =

0.05 mm; fig. 12, right stylus VII and associated setae of N. gentilei L.

Smith ventral view, e rr 0.05 mm.

104

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Male. — Similar to female, except right forcep with six teeth, left with

five teeth ; male sac with few setae mostly simple, but nine setae show

minute pinnulae near the opening of the sac; abdominal segment X, width

0.17 mm, length 0.22 mm; length of body including forceps 3.54 mm.

In addition to the six paratypes, I have examined a series

of eight metatype adults from Sharp Park, San Mateo County,

California. In some of these the females lack the basal tooth on

the right forcep.

Holotype female and allotype male (California Academy of

Sciences), and 18 paratypes (California Insect Survey; Univer-

sity of California, Davis; U.S. National Museum) were collected

in chaparral leaf mold in the San Bruno Mountains, San Mateo

County, California, in January, 1957.

I take pleasure in naming this species after Mr. Douglas W.

Price who collected it.

Nanojapyx gentilei L. Smith, new species

Female. — Similar to N. pagesi L. Smith except setae at dorsal apex

of femora, number 3 longer than number 1, number 2 shortest and not

plumose; setae on labial palp 4, setae on maxillary palp 7; meso- and

metanotum 7 -)- 8 M; seta number 1 on forceps minutely plumose;

length of forceps 0.12 mm; right forcep with six teeth, left with five teeth,

tips of forceps strongly curved inwards; lateral subcoxal organs each with

four or five plumose sense setae; ventral setae associated with stylus VII

as illustrated with setae number 2 and 4 missing; abdominal segment X,

width 0.11 mm, length 0.16 mm; length of body including forceps 2.72 mm.

Male. — Unknown.

Holotype female (California Academy of Sciences), paratype

female and juvenile (University of California, Davis) were

collected ten to twelve inches deep in sandy soil in a growth of

poison oak by L. M. Smith and R. 0. Schuster near Trenton,

Sonoma County, California, on August 7, 1957.

I take pleasure in naming this species for Mr. Adriano

Gentile who has been of great assistance to me by translating

Silvestri’s works on Japygidae from Latin.

Nanojapyx hamoni L. Smith, new species

Female. — Similar to N. pagesi L. Smith except setae at dorsal apex

of femora, numbers 1 and 2 plumose, with number 3 equal in length to

number 1; setae on labial palp 4, setae on maxillary palp 7 + 8; ventral

Explanation of Figures

Forceps of Nanojapyx females, dorsal view, reversed left to right, setal

pattern of all as shown in fig. 13. All figures at same magnification, e rz

0.10 mm. Fig. 13, pagesi', fig. 14, hamoni, a, right, and b, left forcep;

fig. 15, pricei; fig. 16, gentilei', fig. 17, coalingae.

April, 1959]

SMITH JAPYGIDAE I

105

106

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

setae pei’ row on tarsus 3 -j- 3 ; meso- and metanotum 1 1 M ; seta

number 1 on forceps minutely plumose; length of forcep 0.14 mm; left

forcep with eight teeth with distal tooth almost hidden, right forcep with

five teeth, all teeth pyramidal in shape, not recurved ; lateral subcoxal

organ with 6 + 6 plumose sensory setae; ventral setae associated with

stylus VII with setae 2 and 4 present, these become progressively smaller

anteriorly, setae 1, 3, and 5 with 4 pinnulae each; segment X, width 0.19

mm, length 0.22 mm; length of body including forceps 3.40 mm.

Male. — Unknown.

Holotype female (California Academy of Sciences) collected

in redwood leaf mold near Half Moon Bay, San Mateo County,

California on July 21, 1957 by Mr. R. 0. Schuster. Three female

paratypes were collected in clay-loam soil and humus under

Monterey Cypress, by W. H. Lange and R. Schoeppner at Wolf

Ranch, San Mateo County, California, April 24, 1958 and are

deposited in the collection of the U.S. National Museum, Cali-

fornia Insect Survey and the University of California, Davis.

Nanojapyx coalingae L. Smith, new species

Female. Similar to V. pagesi L. Smith except antenna with 23 seg-

ments; antennal segment 18 with eight setae in the distal whorl; setae at

dorsal apex of femora, numbers 1 and 2 plumose with number 1 equal

in length to number 3 ; setae on labial palp 3, setae on maxillary palp 8 ;

meso- and metanotum 7 + 7 M ; ventral setae per row on tarsus L 1

2 + 2 ; L 2 and L 3 3 + 3 ; seta number 1 on forceps clearly plumose ;

tergum X setae between carinae, row A as in pagesi, the two pair of m

posterior to A strongly developed, row B anterior of middle, C irregular,

with one additional pair of m just mesad of the lateral setae, median pair

of M posterior, simple, all other M plumose; length of forceps 0.14 mm,

right forcep with one distinct sharp pointed recurved tooth and crenulations

suggesting reminants of six teeth, left forcep the same; lateral subcoxal

organ with 6 + 7 plumose sensory setae; ventral setae associated with

stylus VII, seta number 2 present but small, and seta 4 absent; setae 1, 3,

and 5 with four or five pinnulae; segment X width 0.14 mm, length 0.20

mm; length of body including forceps 3.06 mm.

Male. — Unknown.

Holotype female (California Academy of Sciences) was col-

lected in juniper leaf mold near Coalinga, Fresno County,

California on January 22, 1958, by Mr. H. L. Wilson.

Key to the Species of Nanojapyx L. Smith

1. Antennae with 23 segments coalingae

— Antennae with 22 segments 2

2. Three setae per row on tarsus... hamoni

— Two setae per row on tarsus (fig. 6).. 3

3. Three setae on labial palpus (fig. 8) pagesi

— Four setae on labial palpus 4

April, 1959]

CLAUSEN^ — -PARASITE RELEASES

107

4. Plumose seta at apex of femur shorter than adjacent simple seta

(fig. 5) gentilei

— Plumose seta at apex of femur same length as adjacent simple

seta pricei

Literature Cited

Paclt, J.

1957. Diplura. Genera Insectorum. Fascicule 212, p. 83.

SiLVESTRI, F.

1949. Contributo alia conoscenza degli Japygidae (Insecta Diplura)

Rend. Accad. XL, s. 3, 27:3—115.

RELEASES OF RECENTLY IMPORTED INSECT PARASITES

AND PREDATORS IN CALIFORNIA, 1956-57

C. P. Clausen

University of California, Riverside

The following list, reporting the first field releases of certain

imported species of parasites and predators by the Department

of Biological Control, supplements two preceding reports,^’ ^ cov-

ering the years 1952—53 and 1954—55. The year of first release

is 1957 unless otherwise indicated.

The species listed in the 1952—53 report as Platynaspis ( ? )

sp. has since been identified as Exochomus metallicus Korschefsky.

Area or County

Host and Parasites or Predators

Origin

of Release

Saissetia oleae (Bern.)

Coccophagus mexicensis Gir.*

Mexico

So. Calif.,

Tulare

Mesopeltis atrocyanea Masi

Mexico

San Diego

Metaphycus lichtensiae (How.)

Pakistan

Coastal

So. Calif.

Microterys consobrinus Comp.*

Mexico

San Diego

Microterys flavus (How.)

India

So. Calif.

(Black scale race)

Aonidiella aurantii (Mask.)

Aphytis sp. (Burma)

Burma

So. Calif.

Aphytis sp. (India)

India

So. Calif.

Aphytis sp. (Pakistan)

Pakistan

So. Calif.

Chilocorus tristis Fald.^

Japan

San Diego

San Bernardino

Cybocephalus sp.

India

So. Calif.

Pharoscymnus sp.

India

San Diego

108

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Area or County

Host and Parasites or Predators

Origin

of Release

Physcus sp.

Burma

San Diego

Riverside

Telsimia sp.

Burma

So. Calif.

Lepidosaphes beckii Newm.

Chilocorus discoideus Crotch*

Kenya

So. Calif.

Parlatoria oleae (Colvee)

Aphytis sp.

Pakistan

San Joaquin

Valley

Archenomus sp.

Pakistan

San Joaquin

Valley

Coccophagoides sp.

Pakistan

Fresno

Tulare

Planococcus citri (Risso)

Nephus sp.

Mexico

V entura

Therioaphis maculata (Buckton)

Adonia variegata Gze.^

India

Riverside

Fresno

Yolo

Chrysopa carnea Steph.^

India

So. Calif.

Yolo

Scymus nubilis Muls.^

India

Riverside

Yolo

Synharmonia conglobata (L. )*

Turkey

San Diego

Aphis spiraecola Patch

Aphelinus chaonia Wlkr.*

Hong Kong

Riverside

Tetranychidae and Eriophyidae

( various)

Stethorus sp. “P”

Pakistan

San Diegc

Stethorus sp. “D”

India

San Diegc

Riverside

Santa Barpa:

Phytophagous Snails

Gonaxis kibweziensis (Smith)*®

Agiguan

San Diego

* First releases made in 1956.

1 Pan-Pacific Entoni. 31(2):90-2. 1955.

2 Pan-Pacific Entom. 32(3) :125-7. 1956.

3 Received from Canadian Department of Agriculture.

4 Received from Entomology Research Division, U.S.D.A.

5 Received from Pacific Science Board and Hawaii Board of Agriculture and Forestry.

April, 1959]

POWELL PYRALID NOTES

109

NOTES ON THE CALIFORNIA SPECIES OF THE

GENUS PYLA GROTE

(Lepidoptera: Pyralidoidea)

Jerry A. Powell

University of California, Berkeley

The black moths of the genus Pyla Grote are dayflying

phycitids which, in western North America, inhabit boreal regions.

Carl Heinrich, in his magnificent monograph of the Phycitinae

(1956), divided the genus into two superficial species groups,

those with grey forewings and those with shining black-brown

wings. Four species are recorded from California, all of which

are members of the latter group, the typical Pyla of the earlier

authors.

During the course of the past few seasons, collecting in the

mountains of northern California^ has produced certain distri-

butional data which seem significant. It has become apparent that

P. sylphiella Dyar, previously only reported from Washington

and British Columbia extends southward and is continuous with

the California species, P. scintillans (Grote). In addition, a few

remarks on variation and a description of the female of P.

nigricula Heinrich are included in the present paper. Where not

otherwise indicated in the data, the collection records are my

own.

Pyla scintillans ( Grote

Nephopteryx scintillans Grote, Papilio, Vol. 1, p. 18, 1881.

Pyla scintillans (Grote), Heinrich, U.S. Nat. Mus., Bull. 207, p. 146,

1956.

Pyla sylphiella Dyar, Ins. Insc. Menstr., Vol. 9, p. 68, 1921; Heinrich,

U.S. Nat. Mus., Bull. 207, p. 147, 1956. (New synonymy.)

A shining, dark brown species with usually only faint indica-

tions of darker transverse banding on the forewing. Very dis-

tinctly marked specimens show dark markings on the forewings

as follows: a broad median band which angles slightly outward;

a thin subapical, usually somewhat sinuate, band; sometimes an

irregular blotch between the bands on the dorsal margin. It may

be easily distinguished from P. fasciella Barnes and McDunnough

and P. nigricula Heinrich, which it closely resembles, by its longer

palpi (exceed the head by about twice the diameter of the eye

as seen from side, fig. 1). Alar expanse 20—26 mm.

1 Field investigations during August, 1957 and July, 1958 carried out with the California

Insect Survey, University of California, Berkeley.

2 Complete synonymy has been given by Heinrich (1956).

110

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

Heinrich lists the distribution as Sierran, from Mineralking,

Tulare County northward to Cisco, Placer County, all at high

elevations. It is apparent, however, from the specimens I have

examined, that in California the species ranges throughout the

Sierras, the north central part of the state, and thence southward

along the inner North Coast Ranges for some distance. The

species is, in general, an early flier compared to the other Cali-

fornia Pyla, with almost all collections having been made in July.

Heinrich, in his treatment of the genus, recognized both P.

scintillans (Grote) and P. sylphiella Dyar as valid species even

though they are indistinguishable on the basis of external charac-

ters. His criteria for separation were: minor differences in the

male genitalia, the markedly different female genitalia, and the

allopatric distribution of the two species. However, the California

specimens of the complex are extremely variable in genitalic

characters and include intermediates between his interpretation

of P. scintillans and P. sylphiella. A series from Del Norte County

seems to be fairly uniform in representing a southern extension

of P. sylphiella in both male and female characters. A male from

southern Siskiyou County appears to be intermediate in the harpe

enlargements, while the aedeagus is of the P. sylphiella type. A

male from Shasta County, although having the harpe characters

of P. sylphiella, has one element of the aedeagus with one short

thorn, the other with three, a situation typical of P. scintillans.

The female genitalia in the specimens examined exhibit more

variability than was indicated by Heinrich. Typical P. sylphiella

forms occur in Del Norte and Siskiyou Counties, but not consist-

ently. One interesting variety lacks both the heavily sclerotized

deep folds of the lower part of the cup-shaped portion of the ductus

bursae characteristic of P. sylphiella and the oval, blade-like

projections of the cup evident in P. scintillans. This variant (fig.

3) has been found in random individuals from Del Norte County

and in single specimens from Plumas and Glenn Counties. This

may represent an intermediate form. Specimens from the southern

Sierras seem more typical of P. scintillans in both sexes, but I

have not seen any females really typical of either of the P. scin-

tillans varieties figured by Heinrich.

In view of the nature of the intergrading characteristics evi-

dent in the complex, I regard Pyla sylphiella Dyar as a synonym

April, 1959]

POWELL PYRALID NOTES

111

of P. scintillans (Grote), P. scintillans being a widespread species

(central California north to British Columbia).

Material examined: (See map) 1^,5 mi. W. Sonora Pass, Tuolumne

Co. VII. 21.56; 1$, Sonora Pass, Tuolumne Co. VII. 21.56; 1$, Glen Alpine

Cr., El Dorado Co. VII. 18.09 (F. X. Williams); 1$, Mohawk, Plumas

Co. VII.8.38 (W. R. Bauer) ; 1$, Chester, 6 mi. E., Plumas Co. VII.14.54;

1^, Hat Creek P. 0., Shasta Co. VII. 11.56 (J. W. MacSwain) ; 1$,

“Caslella” [prob. Castella, Shasta Co.] VII.’02 (no collector); 1^, 1$,

Mt. Shasta City, Siskiyou Co. VII. 2. 58; 3$ $, 11 $ $ , Little Grayback.

N.E. Del Norte Co. VII. 9.58; 1?, 5 mi. N. Black Butte, Glenn Co. 6200',

VI.19.56.

Pyla nigricula Heinrich

Pyla nigricula Heinrich, U.S. Nat. Mus., Bull. 207, p. 148, 1956.

Superficially P. nigricula is very much like P. scintillans, from

which it differs by the smaller palpi (exceed the head by only

about the diameter of the eye as seen from side, fig. 2), and

Explanation of Figures

Fig. 1, Lateral view of head of Pyla scintillans (Grote) ; fig. 2,

Lateral view of head of Pyla nigricula Heinrich; fig. 3, Female genitalia

(variant) of Pyla scintillans (Grote) ; fig. 4, Female genitalia of Pyla

nigricula Heinrich.

112

THE PAN-PACIFIC ENTOMOLOGIST [vOL, XXXV, NO. 2

apparently is almost indistinguishable from P. fasciella. All three

are easily separated by their genital characters. Heinrich described

P. nigricula from a unique male collected by A. H. Vachell at

Verdi, Nevada. 1 have encountered it at spots to the south of the

Lake Tahoe region, but only at elevations above 10,000 feet.

Verdi is on the eastern side of the Sierras, and it seems probable

that the specimen was collected in the mountains near Verdi to

the west.

The male exhibits some variation in wing color, and 1 have

a specimen which differs from the original description as follows:

both the antemedial and submarginal dark bands of the forewing

clearly indicated, the area between, as well as the margin, with

the same bluish overscaling as is present on basal area. Another

has just a faint suggestion of the median bluish, otherwise closely

resembling the type description. Alar expanse 23—25 mm. (the

type is 26 mm.) .

The unique type is a male, and the previously unknown

female is described at this time.

Head, palpi, other appendages, as in male. Forewing black, tinged

with bluish overscaling as follows: basal area, median area except for an

indistinct center blotch, a thin, distinct submarginal transverse line parallel

with margin, marginal area; remaining black bands as follows: antemedial

oblique line, irregular blotch at end of cell, two submarginal thin bands on

either side of the distinct blue line. (In older specimens the bluish tends

to become obscure, giving a less distinctly marked appearance). Hind

wing, shiny dark brown as in male. Alar expanse 23—24 mm. Genitalia as

in fig. 4 (three specimens examined), close to P. fasciella Barnes and

McDunnough but differing by the smaller ovipositor pads (ratio of ovipositor

pad length to: posterior apophyses distad of pads to: length of sclerotized

band of ninth segment about 1:6: 1.5; in P. fasciella about 1:4:1) and by

minor differences in shape of the sclerotization of the ninth segment and

paired plates of ductus bursae.

Neallotype femak: Sonora Peak, Tuolumne County, Cali-

fornia, August 10, 1957 (J. Powell) deposited in the California

Insect Survey collection. University of California, Berkeley.

I have not seen P. fasciella, but judging from the description

given by Heinrich, it is nearly indistinguishable from P. nigricula

in external characters. Although the genitalia are distinct, the

two entities are also geographically isolated (P. fasciella is known

only from Mt. Shasta in northern California), and perhaps the

status of P. nigricula should he reevaluated when more material

becomes available, especially from the intervening areas. It seems

April, 1959]

POWELL — PYRALID NOTES

113

a distinct possibility that P. nigricula Heinrich represents a

genitalic variant.

Material examined : 1 $ , Ruby Lake, Inyo County [11,250 ft.] VIII.13.57 ;

1$, near Mono Pass, Inyo County, 12,000', VIII. 11.57 (C. D. MacNeill) ;

1^, Sonora Pass, Tuolumne County, VII.21.56; 2$ $, near Sonora Peak,

Tuolumne County, 11,000', VII. 14.57 (C. D. MacNeill); 4$ $, Sonora

Peak, Tuolumne County, VIII. 10.57.

Pyla viridisuffusella Barnes and McDunnough

Pyla viridisuffusella Barnes and McDunnough, Can. Ent., Vol. 49, p.

406, 1917 ; Heinrich, U.S. Nat. Mus., Bull. 207, p. 149, 1956.

A smaller moth (expanse 17—20 mm.), quite distinct from the

Explanation of Map

Distribution of Pyla scintillans (Grote) (closed circles) and Pyla

viridisuffusella Barnes and McDunnough (open circles) in California ac-

cording to material examined and localities given by Heinrich (1956).

114

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

above mentioned species, with a shining metallic green or bronzy

overscaling and usually with rather distinct transverse bands on

the forewing. The intensity and quality of the color varies consid-

erably between individuals.

Heinrich gives its distribution as the southern high Sierras,

ranging from Miner alking, Tulare County, north to Kennick

Meadows, Tuolumne County. The present records extend a little

more northward. It was flying quite abundantly in northwest Inyo

County in mid-August, along with occasional examples of P.

nigricula Heinrich and another abundant Phycitine, Catasia histri-

atella (Hulst), which it resembles on the wing.

Material examined: (See map) 1^, 2$ ?, Ruby Lake, N.W. Inyo Co.

VIII. 13.57 (C. D. MacNeill), 1$ $, 3 9 9, same data (J. Powell); 3$ $,

29 9, nr. Mono Pass, Inyo Co. 12,000', VIII. (10-15) .57 (C. D. MacNeill) ;

3^ S, 4 9 9, same data (J. Powell); 1,$, Tuolumne Meadows, Tuolumne

Co. “VIII. (1—7)” (no collector); 3$ $, 19, Kennick Meadows, Yosemite

Nat’l. Park, alt. 9,250', VII.28.34 (E. 0. Essig) ; 19, nr. Sonora Peak,

Tuolumne Co. 11,000', VII. 10.57 (C. D. MacNeill), 29 9, same data (J.

Powell) .

I would like to acknowledge with thanks the help of C. Don

MacNeill of the Department of Entomology, California Academy

of Sciences, San Francisco, for the loan of specimens and for

assistance in the field, and Dr. P. D. Hurd, Jr., of the Department

of Entomology, University of California, Berkeley, for reading

the manuscript and suggestions.

Literature Cited

Heinrich, Carl

1956. American moths of the Subfamily Phycitinae. U.S. Nat. Mus.,

Bull. 207, 581 pp.

OBSERVATIONS ON THE SURVIVAL OF ARHOPALUS

PRODUCTUS (LeCONTE) LARVAE IN DOUGLAS-FIR

LUMBER

(Coleoptera: Cerambycidae)

Charles B. Eaton

California Forest and Range Experiment Station^

Forest Service, U.S. Department of Agriculture

Larvae of the roundheaded borer, Arhopalus productus

(LeConte), often are found in the wood of dying or dead

1 Maintained at Berkeley, California, in cooperation with the University of California.

April, 1959]

EATON CERAMBYCID SURVIVAL

115

Douglas-fir trees. They are known to be able to survive and to

continue to develop in recently cut lumber from such trees, even

after it has been used in buildings. Because the adults and larvae

of this beetle occasionally bore holes through floors, ceilings,

walls, and roofs of new structures, they sometimes are a problem

to lumber suppliers, builders, and home owners (Eaton and Lyon,

1955).

Emergence in buildings has almost invariably been reported

as occurring within a year after the buildings were constructed.

One might infer that the beetles all complete their development

in a single year. This is contrary to what we know or suspect

about the insect’s life span in nature. Kimmey and Furniss (1943)

report that the beetles, normally, probably take several years to

reach maturity. A more plausible explanation for the absence of

the beetles in buildings after the first year is that the environment

becomes unfavorable for those that are unable to mature in this

period. Consequently they die.

In their natural environment A. productus beetles develop in

fairly moist wood. In buildings constructed with infested lumber

that is green when first used, the drying that takes place would

result in conditions quite different from those of wood still in

the tree in the forest. These conditions probably would tend to

deter larval development, and it is conceivable that only the more

fully developed individuals would reach the adult stage. Some

support for this belief was found from observations on the survi-

val of a few larvae in lumber kept indoors at Berkeley, California,

over a 13-month period.

In November 1955, two pieces of infested Douglas-fir, about

2 inches by 4 inches by 4 feet in size, were obtained from recently

cut fire-killed trees near Jenner, Sonoma County, California.

When brought indoors, the wood was very moist, prominently

bluestained, and well riddled with roundheaded-borer galleries.

The larvae were located by cutting along their galleries through

the wood. After their size and condition were noted, the larvae

were sealed in their galleries with black masking tape. The tape

was removed briefly at intervals of one to two months so that the

condition of the larvae could be observed.

Eight larvae were found in the two pieces of lumber. One

additional larva escaped detection, but was discovered when the

wood was split up in September 1957. The larvae were kept under

116

THE PAN-PACIFIC ENTOMOLOGIST [vOL. XXXV, NO. 2

observation from November 16, 1955 to December 27, 1956. At

the beginning of this period they were thought to be between

one-half and three-fourths grown. However, head-capsule measure-

ments of those that died revealed that all but one were of the same

size, presumably full grown.

Only one of the eight larvae completed its development. The

remainder died at various lengths of time after observations were

begun. The one individual that survived was a male that emerged

as a full-grown beetle at seven months. Of those that died, one

succumbed after one month, one after four months, three after

seven months, one after ten months, and one after thirteen months.

The moisture content of green Douglas-fir sapwood is normally

100 per cent or more of its oven-dry weight. However, under

indoor conditions, such as those in which the samples were held,

moisture content would gradually decrease to the level commonly

reached in buildings in this area, i.e., about 10 per cent. During

the early part of the observation period, when the samples were

still fairly moist, most of the larvae continued their mines. How-

ever, as the wood dried, they became less and less active. The

frass they produced, which is normally fairly coarse, became very

fine. The larvae, instead of increasing in size, became smaller so

that in death their corpses were mummified. These observations

suggest that the wood became too dry for the larvae to survive,

and that they died from dessication. When infested lumber is

used in buildings, many larvae probably suffer the same fate. This

may explain why A. productus damage is seldom reported beyond

the first year after construction.

Literature Cited

Eaton, C. B. and Lyon, R. L.

1955. Arhopalus productus (Lee.), a borer in new buildings. U.S.

Forest Service, Calif. Forest and Range Expt. Sta. Tech. Paper

11, 11 pp., illus.

Kimmey, J. W. and Furniss, R. L.

1943. Deterioration of fire-killed Douglas-fir. U.S, Dept. Agr. Tech.

Bui. 851, 61 pp., illus.

A PREOCCUPIED NAME IN SOLIERELLA

(Hymenoptera : Sphecidae)

Solierella prosopidis Williams, new name, is proposed for

Solierella mandibularis Williams. Solierella mandihularis Williams

(Pan-Pac. Ent., 34:212, 1958) is preoccupied by Solierella

mandihularis de Beaumont (Bull. Soc. Sci. Nat. Phys. du Maroc,

36:147, 1957). — Francis X. Williams, La Mesa, California.

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Figs — For control of dried fruit beetles and vinegar flies. Use

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