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LELAND STANFORD JUNIOR UNIVERSITY PUBLICATIONS
UNIVERSITY SERIES

THE CALIFORNIA SPECIES
OF MEALY BUGS

BY
GORDON FLOYD FERRIS

Instructor in Entomology

STANFORD UNIVERSITY, CALIFORNIA
PUBLISHED BY THE UNIVERSITY
1918

1, }
Wo

RUA Ta

lan Nie yung
ia is ae

CONTENTS

A. Introduction : : ; ‘ ; ; Senate.
I. Purpose and scope of the paper 5

II. Acknowledgments and material . : BRAS e
Ill. Types . : : . : ; : ‘ Bee
B. Methods of Study 8

C. Taxonomic value of morphological characters, secretions, and

biology , : ; : ‘ ‘ ‘ ee

I. Morphology ; j : 5 : ‘ rhe 8

1. Adult female . , ; : : ‘ ; ' ere

(1). Antennae. : ‘ P ; : : 1)

(2). Les. : ; 4 : ( : Pet Wea

(3). Mouthparts . . : ; : . ; « 20

(4). Dorsal ostioles . : ‘ ; ; ‘ a 20

(5). Cerarii : . i A j , : ony

(6). Anal lobes . I : ; : ; : a ae

(7). Pores and ducts . ; , ; : . ae

(8). Body setae . : me AS

(9). Anal ring and anal lobe setae ; ; ; ae,

(10). Anal ring . , p ; : : 4

(42). Spiracles ; : ; : : : ae

2. Larvae . ; ; ; : ; : ; ; aes.

3. Adult male : : ; : - aye

II. Secretions . , : , ; a

III. Biology ‘ § : : : ; : ong”?

D. Systematic treatment . , : : : i ; : a0

I. Definition of the group : ‘ : ‘ ; ; ee

II. Generic concepts . , é hack 56 : gee

III. Synonymical list of names applied to California species 2 aie
IV. Key to genera represented in California ; . ; BO

CONTENTS

V. Description of genera and species

A,

. Phenacoccus

y

OCNA ww ff W WH

Pseudococcus .

Puto’

. Heterococcus .
. Trionymus

. Ripersiella

. Cryptoripersia

Erium

Antonina

FE. References cited

INTRODUCTION

PURPOSE AND SCOPE OF THE PAPER

The systematic literature dealing with certain groups of the Coccidae
or scale insects is extremely unsatisfactory, this condition being especially
pronounced in the case of those groups commonly called the “soft scales.”
The present author (4)+ has stated in an earlier paper that “. . . of the
nearly 100 species of mealy bugs thus far described from North America,
including some 35 from California, not more than three or four are
recognizable at all on the basis of the published descriptions if taken apart
from their typical host and their type locality,” nor at the time of the
present writing does this statement need any extensive modification. A
few species have been described since the statement was originally made
and to some of these it may not apply, but so far as the species known
at that time are concerned, the additional claim may safely be made that
even under the optimum conditions cited, the identification of many of
our species can not be accomplished with reasonable certainty.

The actuating motive of the present paper has been the hope of
remedying this condition as far as may be for those species of mealy
bugs occurring in California. However, progress toward this goal has
been made possible only by the utilization of certain methods not ordi-
narily employed, at least by American entomologists, and by the empha-
sizing of certain morphological features of which some have been en-
tirely overlooked and of which the full significance of none seems at any
time to have been realized. The scope of the paper has therefore been
extended to include an explanation of these methods and a description
of these morphological features.

The restriction of the paper to those species occurring in California
has not been due to any desire to keep within the boundaries of some
artificially defined political unit nor to any belief that the California
mealy-bug fauna is at all separable as a group from that of the rest of
North America. The limitations imposed have been purely those of can-
venience. It so happens that most of the species known from west of
Colorado and New Mexico have been described from California and that
the types of these are for the most part in western collections. It also
happens that most of the California species have been taken from two

+Numbers in the text refer to the list of references cited at the end of the
paper.

7

6 THE CALIFORNIA SPECIES OF MEALY BUGS

rather restricted localities, thus making it a comparatively simple matter
to procure topotypic material.

While it has therefore been convenient to treat of the California
species as a separate group, the disadvantages of so doing have been fully
realized. The range of most of our species is by no means so restricted
as would appear from the published records and it is highly probable that
at least a part of the species herein recorded as known only from Cali-
fornia occur in the eastern states as well. In fact it is definitely estab-
lished that at least three species have been described under different
names from California and from the eastern or central states and it is
almost certain that the same thing is true of some others. For the sake
of stability in nomenclature it is highly desirable that the mealy-bug
fauna of the entire continent be treated as a unit, but it has not seemed
wise to wait until this can be done. A beginning must be made some
place.

ACKNOWLEDGMENTS AND MATERIAL

Acknowledgments are due first of all to Mr. E. O. Essig of the
University of California, who has most kindly placed his entire collection,
including types, at my disposal with the fullest permission to remount
any of the specimens as I saw fit. Without this assistance it would lit-
erally have been impossible to proceed. Scarcely less am I indebted to
Mr. E. M. Ehrhorn of Honolulu, who has made numerous comparisons
of specimens with types in his collection and who has lent material of
critical importance. Mr. C. P. Claussen, of the California State Board
of Horticulture, has also lent types of manuscript species in his collec-
tion and has assisted in procuring other material. Mr. A. F. Swain of
the Citrus Experiment Station at Riverside, Mr. S. P. McClenahan of
Palo Alto and Professor R. W. Doane of Stanford University have aided
in the procuring of several species which could not otherwise have been
obtained. Mr. Harold Morrison of the Bureau of Entomology has ren-
dered invaluable aid by comparing specimens with types in the National
Collection of Coccidae at Washington and by notes which he has gen- ©
erously furnished. Mr. E. E. Green has supplied me with material of
several genera and species having a bearing upon the general problems
connected with this group. To all these gentlemen are due the fullest
thanks for their assistance.

With the material obtained from the above sources, together with
that in the Stanford collection when I began work and with that procured
by my own efforts, I have been enabled to examine types, topotypes or
other authentic material of nearly all the species recorded from the state.
Two species have escaped me entirely, and of a few I have not been able

INTRODUCTION 7

to see authentic material, although I believe I have satisfactorily identi-
fied them. Two species which have been recorded from the state, but
which were certainly introduced and are not now known to occur here,
have been passed over with a mere mention.

TYPES

The types of all the new species herein described are in the Stanford
collection. In all cases a holotype has been selected. It is perhaps not
out of place to urge that this method be adopted by all students of the
Coccidae, for even in this limited study three cases have been met with
in which more than one species had been included in material labeled type.
There is really no more reason for failure to name holotypes in this group
than in any other.

METHODS OF STUDY

The satisfactory study of any of the scale insects demands the use
of special methods in the preparation of specimens for microscopic ex-
amination and such methods are especially useful in the group here under
consideration. I feel no hesitation in saying that by far the greater part
of the difficulty that has been experienced in dealing with this group has
been due solely to the unsatisfactory methods usually employed in the
making of microscopic mounts. Satsifactory work can not possibly be
done on the basis of tangled masses of antennae, legs and spines, or of
specimens so transparent that they can scarcely be located on the slide.
Yet it is exactly upon such preparations as these that most of the sys-
tematic work on this group has been based. The possession of first-class
preparations is as essential for the proper study of these insects as is the
possession of a microscope.

In the past, the difficulty has been due for the most part to the lack
of satisfactory methods by which good preparations may be obtained and
to some extent to the failure to use those methods which were available.
Within recent years the problem has been attacked by various investigators
and there no longer exists any excuse for the employment of the crude
and unsatisfactory methods with which authors, until very recently, have
been content. Green (5), Brain (2), Stafford (10), and Dietz and Mor-
rison (3) have all recommended methods, the simplest and least cumber-
some of which is perhaps that described by Stafford. This, with certain
modifications, is the one which I have employed.

The general method is simple enough. Specimens may be preserved
either dry or in alcohol, preferably the latter, as it is sometimes, quite
difficult to straighten out individuals which have become wrinkled in dry-
ing. The loss of the secretionary covering in the alcohol is a minor point,
for the final determination of species cannot be made from this covering
and it is usually of incidental assistance only. To prepare the specimens
for study they should be boiled in a weak solution of caustic potash. This,
for reasons which I shall later explain, should not be more than about
an 8% solution. The boiling should be carried merely to the point where
the body contents are thoroughly disintegrated, this point usually being
reached before the color has been entirely lost. The specimens should
then be removed to clean water in a watchglass under a binocular micro-
scope (one of the most indispensable aids in procuring first-class prepara-
tions), a slit cut in one side of the body and the body contents pressed

METHODS OF STUDY 9

out through this opening. For this purpose a straight needle and one
with the tip bent at right angles to the shank are most useful. Upon the
care with which this operation is performed, including the careful flat-
tening out of the body in the most favorable position for study, will
depend much of the value of the mount. It should be remembered that
the body contents will take the stain and if not thoroughly removed will
more or less obscure the characters that it is necessary to see.

Trouble is frequently experienced at this point due to the formation
of hard, white lumps either within the body or partially encasing it. Ap-
parently these lumps are formed by the combination of the caustic potash
with the body fats and there seems to be no way by which their formation
can be prevented. However, they are readily soluble in carbol-xylene
and may be removed by placing the specimens in a small amount of 95%
alcohol containing a few drops of this reagent. This should not be done,
however, until the specimens have been’‘cleaned as thoroughly as possible,
as the alcohol hardens the body contents and makes them difficult to re-
move. The carbol-xylene should be washed out in 95% alcohol before
the specimens are placed in the stain. If difficulty is not experienced with
the formation of such lumps the specimens may be removed to the stain
directly from the water in which they are washed.

The stain which I have used is Magenta and I cannot speak in regard
to the action of any of the others that have at times been recommended.
This stain may be procured in liquid form. The strength of the solutions
obtained seems to vary somewhat, but there appears to be very little, if
any, difference in their staining qualities and they can sometimes be much
diluted with water. Staining is best carried on in deep hollow slides
having a ground-glass surface upon which data may be written. These
should be covered with a coverglass and the stain allowed to run out
under this. The evaporation of this stain under the coverglass will seal
a slide so effectively that it may sometimes remain for days without
drying out.

Usually a half hour is sufficient to give a stain as good as can be
obtained, this having been demonstrated by numerous experiments. The:
specimens should then be removed to 95% alcohol in which the excess
stain is washed out, then placed for a moment in carbol-xylene (three
parts xylene to one part carbolic-acid crystals) and then mounted in
balsam. If the staining has been effective the resulting mount should
show all spines and chitinized areas of a bright red, while the remainder
of the derm is perfectly clear.

As I have said, the general method is simple enough, but care in
numerous details which one can only learn from experience is necessary
to get the best results. Above all care must be taken that the specimens

10 THE CALIFORNIA SPECIES OF MEALY BUGS

do not become unduly heated, as this destroys their staining qualities,
the resulting preparation being of a uniform bluish color and showing
little or no differentiation between the various parts. It is for this
reason that a weak solution of caustic potash should be used, for it ap-
pears that the boiling point of a strong, “soupy” solution is somewhat
higher than the critical point.

It is especially noteworthy that the method outlined above may be
used for the purpose of rendering old mounts suitable for study. The
results obtained are sometimes not as good as with fresh material, but
they are usually very much worth while. The specimens should be re-
moved from the mount if possible without the use of heat, or if heat is
necessary, it should be very carefully applied. The specimens should
then be freed from the medium in which they have been mounted, the
method of doing so depending upon what medium has been used. If the
medium is balsam they should be soaked in carbol-xylene or a similar
reagent, if glycerine jelly they may be boiled in caustic potash. In any
case this boiling should not be omitted, as it renders the specimens flex-
ible. Following this the treatment proceeds in the usual manner. The
only risk encountered in attempting to stain old mounts lies in the pos-
sibility that they may have been overheated when they were originally
made and this can only be discovered by experiment. However, this risk
may partially be avoided by staining but a part of the specimens con-
tained in the mount.

TAXONOMIC VALUE OF MORPHOLOGICAL CHAR-
ACTERS, SECRETIONS, AND BIOLOGY

MORPHOLOGY

The morphology of the mealy bugs has been dealt with to some
extent by Berlese (1) who has described some of the structures that are
available for taxonomic purposes, although he has not himself carried out
any taxonomic studies; Marchal (7) seems to have been the first to
make use of these characters. Smith (9a) has investigated the value of
the characters ordinarily employed and in a later paper (9b) has dis-
cussed the possible value of others. Brain (2) has also pointed out the
possible value of many characters not ordinarily employed, although, curi-
ously enough, he does not himself make any particular use of these char-
acters. In no case does there seem to have been any real appreciation of
the full value of the characters observed and there still remain some that
do not appear to have been noted at all. |

In view of the fact that no wholly satisfactory discussion of the tax-
onomic value of the various morphological features of this group exists
it is necessary to consider the matter at some length. Unfortunately the
material available for study is somewhat limited and most of the conclu-
sions presented must be regarded as but tentative. However, they apply
to the material examined by me (which includes a number of species not
coming within the scope of this paper), and very probably apply to a
considerable degree to the remainder of the group.

Adult Female

_ Antennae.—Almost since the serious study of the Coccidae began the
antennae have been relied upon to furnish both generic and specific cri-
teria. The present generic concepts in the group here under consideration
are based in large part upon the number of segments in the antennae of
the adult female and the relative or actual lengths of the individual seg-
ments have been much used for purposes of specific differentiation. The
value of the antennae in defining generic groups will be discussed in a
later paragraph on generic concepts and we shall here consider merely
their value as specific criteria.

If we compare a limited number of individuals of a few species it
may not only appear that there are certain rather obvious differences in
the actual lengths of the individual segments and of the antennae as a
whole, but there may also be differences in the relative lengths of the

12 THE CALIFORNIA SPECIES OF MEALY BUGS

segments, differences which are somewhat difficult to express. Attempts
to present these differences in a tangible way have led to the introduction
of the familiar “antennal formulae” and the more recent “antennal
graphs.” The weakness of the antennal formulae is now fully realized
and they are at present but little used, yet their pernicious effects remain,
for few, if any, of the species established largely upon them have ever
been redescribed and many of these must stand in our literature as doubt-
fully recognizable, or even not recognizable at all, until their types have
been re-examined. I cannot but consider the original introduction of
these antennal formulae as the most unfortunate mistake that has been
made in the study of the Coccidae.

The antennal graphs have been but recently introduced and have
not received any extensive trial. It will therefore be desirable to consider
them at some length. It may be said in their favor that if the antennae
are at all to be used as specific criteria no better method than this can
be devised by which to present the facts: The weakness of the method
will be in large part due to the unsatisfactory character of the material
with which it must deal. It can at the best do nothing more than transfer
difficulties from the slide to the paper before us where they may perhaps
be analyzed. | |

The method of constructing these graphs is quite simple. The actual
measurements of the individual segments are obtained by the use of an
ocular micrometer. Then upon a sheet of co-ordinate paper certain units
are laid out along one axis to represent the individual segments and upon
the other axis are plotted the lengths. Connecting up the points thus
obtained we have a graph which will give a very clear indication of the
relative and actual lengths of the segments. Plotting the graphs of sev-
eral antennae will indicate the range of variation. The method of con-
structing these graphs has been discussed at length by Brain (2) and
Hollinger (6) and for the purpose of this paper a combination of the
methods used by both of these authors has been employed.

In order that these graphs may be of value for taxonomic purposes
it must be shown that the graphs of different species are sufficiently dif-
ferent and sufficiently constant to permit of differentiation between the
species. Brain and Hollinger have considered that this condition is met,
the former having relied very heavily upon these graphs and the latter
having stated that with him the graphs “formed the working basis” for
specific indentification. :

I may say without reserve that I consider these graphs to be fully
as illusory and fully as dangerous as were the now wholly discredited
antennal formulae. ;

The objections to these graphs are both general and specific.

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 13

The graphic method is essentially a method of averages. Even
though it be true that the average graph of any species is relatively con-
stant this does not aid in the identification of non-average individuals.
We must deal with individuals and not with averages.

The graphs are neither sufficiently distinctive nor sufficiently con-
stant to be of value in separating species. In those cases where two or
more species occur together on the same hosts the graphs will not, in
general, reveal the fact that more than one species are present.

The range of observed variation in those species which have been
sufficiently studied is so great as to preclude the use of the graphs.

In proof of these contentions I shall present certain specific instances.

The two species, Pseudococcus crawu (Coq.) and P. quercicolus
n. sp. occur in the vicinity of Stanford University upon the same hosts
and have at times been found together. In life they are so very similar
that they are scarcely distinguishable, but structurally they are very
different, as a reference to the figures illustrating the two will show.
Text figures 1 and 2 present the graphs of these two species. These

4o
= Nw
SN
Ns x vA
INN =
\NY
2 3 4 5 6 7 8

Fig. 1—Pseudococcus crawii (Cog.); antennal graph drawn from specimens taken
from Quercus chrysolepis in the vicinity of Stanford University.

graphs were obtained from individuals taken from the same host plants in
the same vicinity. The two graphs coincide practically throughout and
it is obvious that no amount of contemplation of a graph, including both
species, would reveal the fact that two species are present.

Again, let us consider the case of Pseudococcus ryani (Cog.) and
P. sequoiae (Coleman). These two species occur together on Sequoia
sempervirens in the vicinity of Stanford University and both have been
recorded from various species of Cupressus. The two species are suf-

14 THE CALIFORNIA SPECIES OF MEALY BUGS

ficiently different in life to indicate that they are distinct and structurally,
while very similar, are sufficiently well marked to be readily separable in

=

4a

i |

WV
i

£4
KL

\
VET

2 3 4 5 6 7 8

Fig. 2.—Pseudococcus quercicolus n. sp.; antennal graph drawn from specimens
taken from Quercus chrysolepis and Pasania densiflora in the vicinity of
Stanford University.

7
Ut
LH
|
[|

aeRO FA
Se SE ay “hy PS Stk Be
“SSS Se a
SESE =
a ee
2 3 4 5 6 7 8

Fig, 3.—Pseudococcus ryani (Cog.) and P. sequoiae (Coleman) ; antennal graphs
drawn from specimens taken from Sequoia in the vicinity of Stanford Uni-
versity; graphs of ryani drawn in solid lines, those of sequoiae in broken
lines; shaded area indicates the gap between the two.

good preparations. Text figure 3 represents the graphs of the two spe-
cies obtained from individuals taken from Sequoia in the same locality.
It will be noted that the two graphs are in part quite distinct and that
there is an evident tendency for the average of one to lie well below the

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 15

average of the other. Still it is fair to ask to what extent these graphs
would be of aid did we not already know the two species to be distinct.

4
/
' ; J
/
ooh caN Z y ||
NS
WN 7 WN yy
Sa ee
= a J = — -_
— =
=e a
2 3 4 5 6 7 8

Fig. 4.—Pseudococcus ryani (Cog.) and P. sequoiae (Coleman) ; antennal graphs
drawn from specimens taken from various hosts and localities; position of
graphs as in fig. 3.

Text figure 4 presents the graph obtained by using all the available speci-
mens of these two species from different hosts and localities. It will be
noted that the gap between the two graphs has entirely disappeared,
although the average graphs would still lie quite far apart. Again it is
fair to ask if anyone would separate these species on the basis of such a

40
/
>.
> SN
~ es eS ied
aed ~N
\ bars
We a eM
20) =
aN Yt
Sw“ > al Ss
SS — i, Se
4) —
2 3 4 S 0 7 8

Fig. 5.—Pseudococcus longispinus (Targ.) ; antennal graph drawn from specimens
taken from various hosts and localities.

16 THE CALIFORNIA SPECIES OF MEALY BUGS

graph did he not already know them to be distinct, or, knowing them to
be distinct, where he would draw the line between them.

40
Y
SS
— —
vel
= a
a St
- 3 4 5 6) 7 8

Fig. 6—Pseudococcus citri (Risso); antennal graph drawn from specimens taken
from citrus.

Hi
fl

2. a 4 /

Fig. 7.—Pseudococcus citrophilus Claussen; antennal graphs drawn from speci-
mens taken from citrus and other hosts.

In text figures 5 to 8 are shown the graphs of the four species,
Pseudococcus longispinus (Targ.), P. citri (Risso), P. citrophilus

(Claussen) and P. maritimus (Ehrhorn). All four of these species occur f

on citrus, as well as on many other hosts, in California. They are readily
separable in life and structurally are very different and very easily recog-
nizable. Yet the graphs of the first three fall almost entirely within the
graph of P. maritimus nor is there sufficient difference between any of

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 17

them to “form a working basis” for their separation. It may be pointed
out that the use of more individuals would in all probability have in-
creased the difficulty in distinguishing between the graphs due to the
| introduction of extreme forms.

| 49

a ee fh,
y roe
fT ta NY
eae : =a AN Ng —
— WYNN — SS
AYA 4 R i;
2 s Wy
—T ae
N —
Qs “i
—= — | = cy
2 3 4 5 6 7 8

Fig. 8..Pseudococcus maritimus (Ehrh.) ; antennal graphs drawn from specimens
from various hosts and localities.

4
pe a
\ a's
ZN i
2 NN aN
SW B
NRE =
~4 N OY om
N
2 3 4 5 6 7 8

Fig. 9.—Pseudococcus maritimus (Ehrh.); antennal graphs of specimens taken
from Mesembryanthemum at La Jolla (above) and Rubus vitifolius at Stan-

ford University (below). The shaded area indicates the gap between the
two.

In text figure 9 are shown the graphs obtained from specimens of
P. maritimus taken from Mesembryanthemum at La Jolla, Calif., and
from Rubus vitifolius at Stanford University. There is more difference

18 THE CALIFORNIA SPECIES OF MEALY BUGS

between these two lots of specimens belonging to the same species than
there is between any of the species previously discussed. It may, of
course, be asserted that we are here dealing with distinct species. It is
true that structurally these individuals are slightly different but the dif-
ferences are at the best small and are connected by a complete .series of
intergrades. ;

40) ]
2 Lapeny |
Sib
is He
_ K q
Re ia oe % NS a at Gs Ve TALL
Sm Oe a VA
ENS ‘NS eT =e /
pS Na Zoe S> | y
NS \ —_ ——™~ <j
S11 — if
“NN we a, ey
10 Se —_— ——
2 3 4 5 6 7 8

Fig. 10.—Pseudococcus maritimus (Ehrh.) ; graphs of observed extremes (indicated
by heavy, broken line), including within them the graphs of individual an-
tennae of 13 other species (indicated by solid lines).

In figure 10 the heavy, broken line indicates the observed extremes
of P. maritimus and included wholly or in large part within these ex-
tremes are the graphs of individual antennae of thirteen other species, in-
cluding such diverse forms as Trionymus calceolariae, Pseudococcus citri,
P. citrophilus, P. comstocki, P. crawu, P. krauhniae, P. longispinus, P\
pini, P. quercicolus, P. ryani, P. salinus and P. virgatus. On the basis of
this graph any of these individuals might well be P. maritumus.

It seems hardly necessary to pursue the matter further. I have ex-
pended so much space in attempting to show the futility of these graphs,
not because of any personal prejudice against the method, but from a
belief that its use can do nothing but perpetuate the confusion now exist-
ing in the systematic literature of this group. The material with which
the method deals is of such a nature as to preclude its use and there are
other and fully reliable characters upon which determinations of species
may be based. A

Legs.—In many descriptions a considerable amount of attention has
been paid to the legs, their actual measurements in microns being given

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 19

and the character of their vestiture of hairs being minutely dealt with.
In general I have not found such characters to be of any special value.

A

Fig. 11—Pseudococcus maritimus (Ehrh.); outlines of legs, to show variation.
A, from Salix at Stanford University; B, from passion flower at Palo Alto;

C, from orange in southern California; D, from Mesembryanthemum at La
Jolla; E, from roots of clover at Medford, Oregon.

In text figure 11 are shown some of the variations in size found in P.
maritimus. Where extreme differences are found, such as in the case of
Erium lichtensioides and Ripersiella kelloggi where the legs are much
reduced in size, they may be of a certain value. Reference to the legs
has been omitted in this paper except in such cases.

A B

Fig. 12.—Tarsal claws of: A, typical Phenacoccus, and B, typical Pseudococcus.

The tarsal claw, however, affords certain very convenient generic
characters. In certain groups there is a minute tooth or “denticle” on
the face of the claw (figure 12) which affords a most valuable “key
character.” Taken by itself this character is probably of little importance,
for it has certainly been independently developed in widely separated
groups (for instance, it appears in some of the Margarodinae), but in
conjunction with other characters it is extremely useful. I have not been
convinced that the digitules merit special attention.

20 THE CALIFORNIA SPECIES OF MEALY BUGS

Mouthparts.—These seem to present no characters of taxonomic
value.

Dorsal Ostioles—Upon the dorsum of all the Pseudococcine forms
examined by me there are to be found at least one pair of curious pits
with raised, lip-like margins, the “eye-like glands” or “cicatrices” of
various authors, the “‘fossette ostioloformi labiate”’ or “‘foveole labiate”’
of Berlese (1, p. 82, figs. 8, 10). It is probable that two pairs of these
are always present, one pair being found just within and slightly behind
the eyes, the other being on the third segment from the posterior end of
the abdomen or between the second and third segments. The position of

en,

Fig. 13.—Pseudococcus longispinus (Targ.); outline of body to show position of
dorsal ostioles.

these organs is shown in figure 13. To these I apply the name “dorsal
ostioles,” for they appear to correspond in function to the lateral ostioles
of the Heteroptera. Berlese seems to have regarded these structures as
points of attachment for muscles, but this interpretation is probably
wrong. If an active mealy bug be disturbed it may aften be noticed
that one or more globules of liquid appear upon the dorsum. The posi-

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS Zh

tion of these globules, at the most four in number, corresponds to the

position of the dorsal ostioles and there seems to be no room for doubt

that it is from these ostioles that they arise. Whether they are secretions
from special glands or are of some other origin, I cannot at present say.
The connection of these globules with the dorsal ostioles has been noted
by Comstock in his Report of 1882.

In some cases the anterior pair of these ostioles appear to be much
reduced in size and they are usually somewhat difficult to see, due to the
fact that they lie directly above the anterior legs and are thus more or
less obscured in preparations. For this reason I am not prepared to say
that this pair is always present, but the posterior pair is to be found in
all the Pseudococcine forms that I have seen. I have not been able to
find them in the members of any other group and therefore regard them
as perhaps the most distinctive character of the group here under con-
sideration.

Ceraru.—This term seems first to have been used by Smith (9b,
p. 74)+ to designate the marginal groups of pores and differentiated
spines (the “lateral patches of bristles’ of some authors) from which
arise the tassels or “filaments” of wax that are such striking features of
many species of mealy bugs. While vague references to these structures
have been made by various authors, Berlese (1, p. 81) seems to have
been the first to describe them at any length, while Marchal (7) has
been the first to realize their taxonomic value. Smith (9b), who seems
not to have been familiar with Marchal’s paper, has independently con-
cluded that “the cerari as specific characters are very promising,’ and
Brain (2) and Hollinger (6) have made some slight use of them.

These are to be regarded as structures of the very highest taxonomic
value. They appear to be found only in the Pseudococcine group, al-
though not all of the species belonging to this group possess them. This,
together with the fact that they are sometimes so much modified as to
render them scarcely recognizable, reduces their value as a group char-
acteristic, but they still remain serviceable for generic and specific differ-
entiation.

A typical cerarius consists of a pair of spines (herein spoken of as
the “cerarian spines’), set close together at the margin of the body and
usually accompanied by a more or less distinct group of triangular pores

+The derivation of this word is not explained by Smith and he has used the
spelling “cerari” in both singular and plural. I am informed by Professor Elmore
of the Department of Latin of Stanford University that the word appears to be
incorrectly formed. He suggests that the spelling “cerarius” for the singular and
“cerarii” for the plural be adopted, with “cerarian” for the adjective form. I have
therefore used these forms throughout this paper.

22 THE CALIFORNIA SPECIES OF MEALY BUGS

and slender setae. This type, however, is subject to much modification.
In some cases the cerarian spines are quite large and stout and are readily
distinguishable from the body setae, while in others they are recogniz-
able only by their paired character and position. In some cases the
grouped pores are very numerous, in others entirely lacking. In some
species the cerarian spines are accompanied by a group of slender setae,
which I designate as “au-iliary setae,” while in others these are not pres-
ent. In some more than two spines are present in part or even all of the
cerarii.

In certain instances a part or even all of the cerarii are surrounded
by more or less well defined chitinized areas which it is usually difficult
or even impossible to see in unstained preparations. In the genus Puto,
and in certain exotic species at present referred to Phenacoccus, all the
cerarii are thus surrounded but in general these areas are to be found
only in connection with the last one or more abdominal pairs. The size
and shape of these areas are in some cases extremely constant and of
much assistance in recognizing species. I am not aware that these have
previously been noted except in connection with the genus Puto.

The number of pairs of cerarii ranges from none to as many as 24.
Within certain limits the number may be taken as of generic value,
although it is probably not always so. I have seen no species of Phena-
coccus (except certain exotic forms now referred here, but which I am
convinced are generically distinct) in which there are less than 18 pairs,
and no species of Pseudococcus in which there are more than 17, although
in some species there are less. In Trionymus the number is from one to
four, while in Antonina there are none. While I have made use of these
facts in this paper the matter needs to be much more fully investigated
before any extensive generalizations can be made.

Anal Lobes.—The form of the anal lobes is in general not such as to
be usable for taxonomic purposes, chiefly because of the difficulty experi-
enced in obtaining preparations of a uniform type. In some cases they
are quite prominent and in others practically lacking, but unless the
characters displayed are of an extreme nature it is hardly possible to
make use of them.

In certain species a more or less well defined chitinized area occurs
on the ventral side of each anal lobe, usually extending in from the base
of the anal lobe seta. In some species these areas are sufficiently con-
stant in shape and size to be extremely useful characters, although in
others where they are normally very small, they may vary even to the
point of being entirely lacking and hence may not be relied upon.

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 23

Pores and Ducts—wNo thorough investigation of the various wax-
secreting glands of the Coccidae and of their ducts and pores has ever
been made, yet it is probable that such an investigation would mark one
of the most significant steps that can be taken in the study of this group.
The existing information is very incomplete and in some instances quite
misleading. It is not desirable to enter into any extended discussion of
the subject here—it is deserving of a monograph by itself—but certain —
facts having a direct bearing upon the particular group under discussion
may well be presented. Unfortunately there exists no satisfactory term-
inology for the various types of ducts and pores and a certain degree of
circumlocution has been necessary in characterizing them, it being unde-
sirable to propose a definite terminology until a comprehensive study of
the whole subject has been made.

The wax-secreting glands are of several distinct types, each type
discharging its products through the body wall by means of a different
type of duct or pore. Certain of these types appear to be common to all
the Coccidae, while others are apparently characteristic of particular
groups. Their chief value will probably lie in the indications which they
give of the relationships of the various genera, although at times they are
useful even in the recognition of species.

The general type of gland opening through a duct (except in the
Diaspinae) is shown in text figure 14, which represents a section through

Fig. 14.—Section through a wax gland of Eriococcus adenostomae Ehrh.

a gland of Eriococcus adenostomae (Ehrhorn). It will be observed that
the duct is divided into two parts, a basal part of relatively large diam-
eter, the inner end of which is reflexed to form a sort of cup from the
rim of which rises the very delicate inner portion of the duct. Modifica-
tions of this type are found in the Coccinae and in certain groups of the
Dactylopiinae (as this subfamily is understood in the Fernald Catalogue),

24 THE CALIFORNIA SPECIES OF MEALY BUGS

some of these modifications being shown in text figure 15. In all the
Pseudococcine forms the inner end of the basal portion of the duct is
truncate and never'reflexed into a cup, a character which I consider to be
distinctive of this group.

i

A a

B C D

Od

Fig. 15.—Diagrams of optical, longitudinal sections through ducts typical of: A,
Eriococcus; B, Kermes; C, Pseudococcus; D, Lecanium.

These cylindrical ducts of the Pseudococcine forms present some
features that may be used as specific criteria.. Two rather distinct types
are commonly to be found. In one of these the ducts are relatively large
and the mouth is usually surrounded by a definite raised rim, while in
the other the ducts are smaller and lack this rim. The relative numbers
of these types is perhaps of specific importance, but is usually difficult to
express and therefore chiefly of value in the direct comparison of speci-
mens.

In some species the cylindrical ducts may be of a distinctive type,
examples of this being seen in such species as Phenacoccus stachyos,
Phenacoccus kuwanae and Pseudococcus virgatus. In Phenacoccus
stachyos and Pseudococcus virgatus these ducts are strikingly large and
in the first named species their mouths are borne at the apices of conical
projections. In both of these species a conspicuous character of the liv-
ing insect is the presence of numerous glassy, waxen threads, which
undoubtedly originate from these ducts. In Phenacoccus kuwanae the
cylindrical ducts are conspicuously short and stout.

The glands which do not possess cylindrical ducts discharge their
secretions through various types of dermal pores. Certain of these types
seem to appear in practically all of the Coccidae, while others seem to be
confined to particular groups. The extent to which they may be relied
upon in forming conclusions in regard to the limits of these various
groups can not be discussed here, for only a very extended study could
give such conclusions weight. However, they do appear to have certain
values within the group here under discussion that may well be con-
sidered.

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 25

The dermal pores in this group are of three quite distinct types
which are represented in text figure 16. Of these types, one (fig. 16a)
is trilocular and more or less triangular in shape. Throughout this paper

ES SESH

sg ees

Cia

Fig. 16—Dermal pores of Pseudococcine forms: A, triangular type; B, multilocular
type; C, quinquelocular type.

it is spoken of as the “triangular type.” Pores of this character are pres-
ent in all the species examined except one and occur for the most part
on the dorsum and in connection with the cerarii, although they may at
times appear on the venter as well. It is probable that the number of
these pores is more or less constant for each species, but, aside from those
occurring in the cerarii, it appears impracticable to make use of them.
They are too numerous to count and a very slight difference in the treat-
ment of specimens makes a marked difference in the visibility of the
pores and hence in their apparent numbers. The number of these pores
occurring in the cerarii, however, is of much significance.

The pores of the other two types are circular. Of these types, one,
herein spoken of as the “multilocular type” (fig. 16b), occurs for the
most part on the venter of the abdomen and appears to be especially con-
cerned with the production of the ovisac. ‘In a few species, notably in
certain of those herein referred to the genus Trionymus, these pores are
found upon the dorsum of the abdomen as well. Aside from this they
appear to present few possibilities of use.

The third type, herein spoken of as the “quinquelocular type” (fig.
16c), occurs in but one species (Heterococcus arenae n. gen. and sp.),
where they entirely replace the pores of the triangular type.

Body setae.—The body setae, especially those of the dorsum, fre-
quently afford good specific characters. Their number, shape, size, and
distribution are all worthy of consideration. The ventral setae are much
less significarit and I have not attempted to use them.

Anal ring and anal lobe setae—In some species these afford fairly
good characters which may perhaps best be expressed in terms of their
actual length, and their relative length as compared with each other.
The actual length, however, is difficult to obtain with accuracy, due to the
fact that the setae are usually bent and for the purposes of this paper |
have made but little use of this character.

26 THE CALIFORNIA SPECIES OF MEALY BUGS

Anal Ring.—The character of the anal ring has in the past been
used to some extent in defining certain of the larger groups, but it is
questionable whether the grouping thus obtained is at all natural. On
the basis of the compound setiferous anal ring the Ortheziinae are more
closely related to the Pseudococcine group than to the Monophlebinae,
yet the probabilities are that the reverse of this is true. To some extent
the anal ring is of generic value, but it needs to be supplemented by other
characters. In one case I have found it of real specific value and in a
few other cases it is of some assistance.

But little reliance should be placed upon the number of hairs on the
anal ring in forming conclusions as to the relationships of genera. Brain
(2) has referred the genus Puto to the Eriococcini because of the pres-
ence of eight hairs on the anal ring, yet this genus most certainly has
nothing to do with Eriococcus. The genus Ceroputo was originally sep-
arated from Puto largely because of a difference in the number of hairs
on the anal ring, yet other evidence indicates that this division is unde-
sirable. Lachnodius has as many as 20 hairs on the anal ring, yet in
other respects is clearly Pseudococcine.

Spiracles——I have not been able to see that the spiracles merit any
very special consideration in this group except in cases such as the genus
Antonina, where they are extraordinarily large.

Larvae

The first stage larvae of all the species examined by me have six-
segmented antennae except in the genus Puto, in which the antennae are
seven-segmented. Except in the genus Antonina the larvae of the female
differ from the adult female in the reduced number of antennal seg-
ments, the somewhat stouter character of the legs and in a general reduc-
tion of the number of pores and spines. There appear to be four instars.
Specimens in the first two instars are hardly recognizable specifically, but
specimens in the third instar can usually be recognized as easily as can
the mature females. Specimens in these stages may be distinguished
from the adult female by the apparent absence of the vaginal opening.

The larvae of the males of Pseudococcus, and in all probability of
most of the other genera, do not differ in the first instar from the larvae
of the female, but in the second instar the anal ring is simple and hair-
less, the mouthparts are entirely lacking, and the cerarii are represented
merely by paired, slender setae. A notable exception is presented by the
genus Puto, in which the second stage larvae of the males are indistin-
suishable from the second stage larvae of the females. In all cases
investigated the larvae form a small waxen cocoon in which pupation

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 27

takes place. An examination of larvae of the males of several species
has revealed no characters by which the species can be determined.

Adult Male

The adult male is alate in some species and apterous in others,
although the wings are usually present. Except for this character most
of the males present few features that can be utilized for systematic pur-
poses, and the difficulty of procuring specimens effectually precludes their
use. Attempts to found genera or subgenera upon the character of the
males alone, as has in one or two instances been done, are hardly to be
commended. The primary purpose of this paper being to aid in the de-
termination of species, the males have been used only in cases where
such action could not well be avoided.

SECRETIONS

The nature and disposition of the waxy secretions have been much
relied upon in this group for specific differentiation and even to some
degree in the formation of generic concepts. The fact has consistently
been ignored by some authors, that these secretions are not the insect
itself, and little or no attempt has been made to correlate them with mor-
phological characters. Furthermore the fact has frequently been over-
looked that similarity in secretionary covering does not of necessity imply
similarity in structure. We thus have Erium lichtensioides redescribed
as Eriococcus artemisiae and Cryptoripersia salina likewise referred to
Eriococcus solely on the basis of the fact that both are enclosed in a com-
plete sac, although neither has anything to do with the genus Eriococcus.
For the same reason we have the published record of the occurrence of
a certain species of Eriococcus in Japan, while the specimens upon which
the record was based are of a species of Antonina. We have Pseudo-
coccus eriogoni referred to the genus Erium because it is enclosed in a
sac, although it certainly has little else in common with the type of that
genus. Nor is this by any means the complete list of such errors.

In this connection we may recall the words of Maskell (8) which
might well serve as the creed of Coccidologists, “On the other hand
(rightly as it seems to me), I have always insisted that true Coccid class-
ification should depend upon the anatomical characters of the insects
themselves, and that mere external features, visible to the naked eye or
an ordinary lens, are but secondary.” It is only through the unreserved
acceptance of this idea with the adoption of the methods necessary in
order to carry it out that progress in the study of this group can be made.

All of this is not to be interpreted as a contention that the secretions
are entirely valueless for taxonomic purposes. There unquestionably
exists a very close correlation between the nature of the secretions and

28 THE CALIFORNIA SPECIES OF MEALY BUGS

that of the structures by which they are produced, a correlation that in
many cases can be directly traced. It is in fact possible to gain a very
accurate conception of the appearance of a species in life from a con-
sideration of its morphology alone and, conversely, it is possible to gain
some idea of the morphology of a species from a consideration of its
appearance in life. It is only in the light of this that most of the existing
descriptions are at all intelligible.

The lateral waxy tassels or “filaments”? which are so conspicuous in
many species of this group, are produced by the cerarii and their form
and size are directly dependent upon the nature of the cerarii. If the
cerarii are lacking there will be no tassels, and if there are no tassels the
cerarii are lacking or very weakly developed. If the cerarii contain a
large number of pores, cerarian spines or auxiliary setae, the tassels will
be strongly developed, and if the tassels are of unusual length or size
we may predict that the cerarii will present a corresponding development,
although we cannot say exactly what form this development will take.
In other words, similar tassels may be produced by a quite different ar-
rangement of the various elements of the cerarii as is shown in the case
of Pseudococcus maritimus and P. citrophilus.

The presence of glassy waxen threads such as appear in Pseudo-
coccus virgatus or Phenacoccus stachyos is unmistakable evidence of
the presence of some sort of large pore or duct, although the exact nature
of this pore or duct can not be predicted. I have been unable, however,
to trace the correlation between the presence of dorsal lumps of wax,
such as appear in Pseudococcus nipae and P. aurilanatus, and any mor-
phological elements, nor have I been able to account for the formation of
a complete sac in some species and not in others. It is possible that this
is dependent upon the physiology of the insect as is the case when the
wax is yellow instead of the usual white. The correlation between the
ovisac and the ventral glands by which it is produced is likewise too
subtle to be traced. I cannot say why the ovisac of one species is
loose and fluffy while that of another is firm, or why one is short and
another is long. It is by no means impossible, however, that sufficiently
careful and extended studies may throw some light upon these difficulties.

BIOLOGY

There has always been more or less of a tendency among students
of the Coccidae to name species on the basis of differences in host plant
and habitat. In a group such as this, where tangible structural differen-
ences are supposedly or actually few, it is difficult to resist this tendency,
yet it must be resisted if we are to avoid confusion of the most serious
kind. Essig, in separating his Pseudococcus bakert from his previously

TAXONOMIC VALUE OF MORPHOLOGICAL CHARACTERS 29

described Pseudococcus obscurus, stated that “. . . it did seem very im-
probable that the same species found on the roots of Opuntia at Los
Angeles, could be synonymous with a species found working on the new
bark of Sambucus glauca (Elder) at Santa Paula, 60 miles away.” On
this basis it would seem even more improbable that the species found on
Opuntia at Los Angeles should be the same as a species found feeding
on the bark of wild cherry in New York. Yet the evidence now avail-
able indicates that both the species on Opuntia and the species on Sam-
bucus are identical with one found in New York on wild cherry.

The apparent anomalies which we encounter if host and biological
differences be disregarded will usually disappear as our knowledge in-
creases and at the worst are as nothing in comparison with the anomalies
which appear if the morphological evidence be disregarded in the same
manner. It is certainly possible, if not even probable, that there exist
forms behaving for all practical purposes as distinct species, yet which
are not separable by any morphological characters that we are able to
appreciate. Indeed, it may eventually be worth while to designate some
of these forms by names, but it seems hardly desirable to take such a step
until all other possibilities have been exhausted.

For the purposes of the present paper all evidence other than that
of morphology has been strictly disregarded. This action has been taken
not from a belief that biological evidence is always entirely valueless,
but from a conviction that it is extremely dangerous when not fully un-
derstood. It is too often merely misleading.

SYSTEMATIC TREATMENT

DEFINITION OF THE GROUP

The vernacular term “mealy bugs” has been used in the title of this
paper chiefly because of a doubt as to the scientific term that should be
applied. Whether the group of genera commonly included under this
term should be regarded as a tribe, the Pseudococcini, of the subfamily
Dactylopiinae, or as a subfamily, the Pseudococcinae, or as a family, the
Pseudococcidae, of a superfamily, the Coccidoidea, is open to debate.
Whatever status may be assigned to them it appears quite certain that
the genera herein discussed, together with certain others, form a natural
group quite distinctly separated from the remainder of the Coccidae, a
group that does not include such forms as Dactylopimus, Eriococcus,
Asterolecanium and other genera of these types with which it is asso-
ciated in the Fernald Catalogue. However, this is a question which as
far as this paper is concerned is a secondary issue, and I have chosen
to employ a vernacular term which is in general quite well understood.

The group of genera included may be defined by the following char-
acters. Coccidae with at least one pair of dorsal ostioles; with entally
truncate cylindrical wax ducts; commonly with at least one pair of
cerarii; the adult female either with well developed legs or antennae or
with these organs vestigial or entirely lacking, commonly more or less
covered with cottony or mealy secretion or enclosed within a cottony
or felted sac; without abdominal spiracles; posterior end of the abdomen
neither cleft nor pygidiform; never with 8-shaped pores.

The extent of the group thus defined is somewhat problematical, but
from the material available to me for study and from the published de-
scriptions it appears that the following genera may be regarded as be-
longing here: Antonina, Cryptoripersia, Erium, Geococcus, Lachnodius,
Natalensia, Phenacoccus, Pseudococcus, Rhizoecus, Ripersia, Ripersiella,
Trabutina, Trionymus and Tylococcus. The genera Cryptococcus and
Fonscolombia (at least F. fraxini) are almost certainly Eriococcine al-
though their position in the Fernald Catalogue would indicate relation-
ship with Pseudococcus. Through the kindness of Mr. E. E. Green I
have been enabled to see specimens of both of these genera and in neither
Cryptococcus fagi nor Fonscolombia fraxim are the dorsal ostioles pres-

SYSTEMATIC TREATMENT 31

ent, while in both species the ducts are distinctly of the Eriococcine type.
Certain other genera remain to be elucidated before their position can
be determined.

GENERIC CONCEPTS

As I have stated in an earlier paragraph the present generic con-
cepts in this group are based largely upon the number of segments in
the antennae of the adult females. The splendid simplicity of this
arrangement is indeed attractive. We need only name a genus to corre-
spond with each different number of antennal segments, with perhaps a
few others for the reception of “aberrant” forms, and the system is com-
plete. This is in fact practically what has been done. However, the
unnatural character of the groups thus obtained has in part been recog-
nized and the system has not been strictly adhered to, with the result
that its simplicity has been somewhat marred. The genus Ripersia was
originally established for the reception of forms with six-segmented an-
tennae, but the majority of the species now referred to it have seven.
Pseudococcus was intended for forms with eight-segmented antennae,
but it now includes species with seven segments as well. As a conse-
quence of this it appears that in several instances a species has been
referred to one genus or another simply because “it looks like a Ripersia”’
or a Pseudococcus as the case may be.

I have attempted in the present paper to utilize other characters than
the antennae for the recognition of generic groups. For this purpose
the cerarii have seemed most useful, the number of pairs present being
considered as diagnostic. It must be admitted that the groupings thus
obtained are not entirely satisfactory, even with as few species as are
here considered. It is not at all improbable that a reduction in the num-
ber of pairs of cerarii might occur in groups that are but little related,
and in fact this appears to have happened. Nevertheless at its worst a
system based fundamentally upon this character is much more satisfying
than one based upon the antennae, and when supplemented by the other
characters that are available seems to approximate a natural arrangement.

I wish specifically to disclaim any attempt to limit or define the
genera here dealt with except as they concern the limited fauna discussed.
The number of species available to me for examination is sufficient
merely to indicate something of the difficulties that must be overcome
before any final definitions can be formed. The task is one for a student
who has before him a very large proportion of the known species and
must not be undertaken on the basis of any purely local fauna.

6 4 THE CALIFORNIA SPECIES OF MEALY BUGS

SYNONYMICAL LIST OF NAMES APPLIED TO CALIFORNIA SPECIES

Note: synonyms are in italics; the genus Dactylopius (as formerly under-
stood) being a synonym of Pseudococcus is not included in the list.

Genus PSEUDOCOCCUS Westwood.

agrifoliae Essig.
Pseudococcus quercicolus n. sp. (in part).
Trionymus villosa (Ehrh.) (in part).
andersont (Coleman).
Pseudococcus ryani (Cog.).
affinis (Maskell).
artemisiae (Essig).
Erium lichtensioides (CkIl.).
aurilanatus (Maskell).
azaleae (Tinsley).
bakeri (Essig).
Pseudococcus maritimus (Ehrh.).
calceolariae (Maskell).
Trionymus calceolariae (Maskell).
citri (Risso).
citrophilus Claussen.
comstocki (Kuwana).
crawii (Coq.) (as to female).
Pseudococcus quercus (Ehrh.).
crawtt (Coq.) (as to male).
Puto yuccae (Coq.).
cupressi Coleman.
Puto cupressi (Coleman).
cupressicolus n. sp.
dudleyi (Coleman).
Pseudococcus ryani (Coq.).
eriogoni (Ehrh.).
| Erium eriogom (Ehrh.).
Pseudococcus yerba-santae Essig.
ephedrae (Cogq.).
krauhniae (Kuwana).
longisetosus n. sp.
longispinus (Targ.).
maritimus (Ehrh.).
Pseudococcus bakeri Essig.
Pseudococcus obscurus Essig.
nipae (Maskell).
Pseudococcus pseudonipae (CkIl.).
obscurus Essig.
Pseudococcus maritimus (Ehrh.).
omniverae Hollinger.
Pseudococcus maritimus (Ehrh.).

SYSTEMATIC TREATMENT 33

quercicolus n. sp.
Pseudococcus agrifoliae Essig (in part).
quercus (Ehrh.).
Pseudococcus crawii (Coq.).
ryani (Cog.).
Pseudococcus andersoni (Coleman).
Pseudococcus one (Coleman).
salinus (CkIl.).
sequoiae (Coleman) (as to female).
Pseudococcus ryani of authors (not of Cogq.).
sequotae (Coleman) (as to male).
Puto cupressi (Coleman).
smithi (Essig).
Trionymus californicus Ehrh. (in part).
Trionymus smithii (Essig) (in part).
solani of authors (not of (CkIl.).
Phenacoccus solani n. <3
timberlakei Ckll.
yerba-santae Essig.
Pseudococcus eriogoni (Ehrh.).

Genus PHENACOCCUS Ckil.
artemisiae Ehrh.
bahiae Ehrh.
Puto yuccae (Cogq.).
colemani Ehrh.
eriogoni n. sp.
kuwanae Coleman.
osborni Sanders.
Phenacoccus stachyos Ehrh.
ramonae Essig.
Puto yuccae (Cogq.).
simplex King.
solani n. sp.
_ Pseudococcus solani of authors (not of Cklil.).
stachyos Ehrh.
Phenacoccus osborni Sanders.
Phenacoccus pettiti Hollinger.
yuccae (Coq.).
_ Puto yuccae (Cogq.).

Genus CEROPUTO Sulc.

Here regarded as a synonym of Puto.
arctostaphyli (Coleman) (Nomen nudum).
Genus PUTO Signoret.

ambigua (Fullaway).
Ceroputo ambigua Fullaway.

34 THE CALIFORNIA’ SPECIES OF MEALY BUGS

cupressi (Coleman).
Pseudococcus sequoiae (Coleman) (as to male).
Pseudococcus cupressi Coleman.
yuccae (Coq.).
Ceroputo bahiae (Ehrh.).
Phenacoccus bahiae Ehrh.
Pseudococcus craw (Coq.) (as to male).
Phenacoccus ramonae Essig.
Ceroputo yuccae (Cog.).
Phenacoccus yuccae (Coq.)
Ceroputo yuccae var. ceanothi Ckll.
yuccae var. ceanothi Ckll.
Puto yuccae (Cogq.).

Genus HETEROCOCCUS new genus.

arenae n. Sp.

Genus TRIONYMUS Berg.

bromi n. sp.
calceolariae (Maskell).
Pseudococcus calceolariae (Maskell).
californicus Ehrh.
Ripersia smithu Essig (in part).
distichlii n. sp.
festucae (Kuwana).
Ripersia festucae Kuwana.
grindeliae n. sp.
smithii (Essig).
Ripersia smithiu Essig (in part).
Pseudococcus smith (Essig) (in part).
villosa (Ehrh.).
Ripersia villosa Ehrh,
Pseudococcus agrifoliae Essig (in baat,

Genus CRYPTORIPERSIA Ckll.

salinus (Ehrh.).
Eriococcus salinus Ehrh.

Genus RIPERSIELLA Towns.
kelloggi Ehrh.

Genus ERIUM Maskell.

eriogonit (Ehrh.).

Pseudococcus eriogoni (Ehrh.).
lichtensioides (CkIl.).

Eriococcus artemisiae Kuwana.

Eriococcus artemisiae var. catalinae Ehrh.

Pseudococcus artemisiae Essig.

Pseudococcus hymenocleae of Essig (not of CkIl.).

Genus ANTONINA Signoret.
crawii Ckll.

SYSTEMATIC TREATMENT 35

KEY TO GENERA REPRESENTED IN CALIFORNIA

Note: This and succeeding keys are based upon carefully stained material.
The key to genera will apply to non-stained specimens, the keys to species will
not. It is assumed that satisfactory preparations will be available and no attempt
has been made to provide for the determination of species upon the basis of any
other.

1. Posterior end of the abdomen of the adult female
deeply invaginated, the anal ring at the inner end
of this invagination ... . wt i! Jy hootonine
Posterior end of the abdomen not ‘nvuandted mid: ee Cat Bala cites
2. Dorsum of the abdomen with circular pores only
Heterococcus
Hata ot thes ahdpnikt width ibatanhie Dare tie) Ah el de ee haga

3. With a denticle on the face of the tarsal claws .
Without a denticle on the face of the claws .

wn

4. With a continuous series of conspicuous, chitinized
areas about the margin ofthe body. . . . . Puto
Chitinized areas, if present, confined to the posterior
end of the abdomen . .. . . .. . Phenacoccus
5. With more than four pairs of cerarii . . Pseudococcus
With not more than four pairs of cerarii. . . . . .... 6
6. Antennae of the adult female very short, 5-segmented,
set close together at the front of the head. Ripersiella
Antennae of the adult female at least 6-segmented, not
set close together at the front of the head. . . . ... 7
7. Dorsum of the abdomen beset with short, stout “spearhead-
shaped” spines. . . wie. 4 as vent! ee ee
Dorsum of the abdomen with sleades spines or setae jaws. | 8
8. Anal ring on the dorsum at some distance from the poste-
rior end of the abdomen, antennae in all stages 6-

SEOMIORIEE Ci ri, . . « « Cryptoripersia
Not so: antennae of the ss dute female 7-8-segmented
Trionymus

Genus PSEUDOCOCCUS Westw.

Coccidae of the type described above; without a tooth or denticle
on the face of the tarsal claw; with not more than 17 pairs of cerarii and
sometimes with not more than 5 distinct pairs; antennae of the adult
female 7- or 8-segmented, usually the latter ; male minute, the male larvae

36 THE CALIFORNIA SPECIES OF MEALY BUGS

after the first molt without mouth-parts or cerarii and with a simple, non--
setiferous anal ring.
Type of the genus Pseudococcus longispinus (Targ.)

Notes: As the genus is thus defined the species found in California form a
quite homogeneous group except for four species in which there has been a reduc-
tion in the number of pairs of cerarii. In P. eriogoni the cerarii are lacking on
the head and also behind the eyes, there being in all but 9 or 10 pairs instead of
the usual 17. In P. salinus certain ot the cerarii along the lateral margins just
behind the eyes are lacking. Both of these species, however, seem clearly to belong
to this group. In P. nipae there is a reduction in the number of cerarii accom-
panied by a tendency for the cerarian spines to be so widely separated that in.
the adult their identity is obscured. In P. aurilanatus there are not more than
five or six pairs of recognizable cerarii and in the anteriormost of these the spines
are so small and so widely separated that they are scarcely to be recognized. This
species, in fact, departs so widely from the typical form of the genus that I am
inclined to believe it should be removed. Nevertheless such action would be at
present unjustifiable and I retain it in Pseudococcus in spite of the discord that
it causes.

The distinction that I have drawn between Pseudococcus and Trionymus is
clear enough as far as the type species of the two genera are concerned, but grad-
atory forms may make it difficult to maintain.

KEY TO CALIFORNIA SPECIES

1. Anal lobe cerarii with as many as 10 cerarian spines
craw ( Coq.)

Anal lobe‘ cerarii with ot more ian 2cerarian spines . . . . 2
2. Anal ring very weakly chitinized, almost simple, pre-
senting a crescentic appearance . . salinus (CkIl.)
Anal ring strongly chitinized, distinctly compound, not
presenting a crescentic: appearance 00 a
3. With no auxiliary setae in the cerarii anterior to the
last two abdominal pairs. . . eA MANA RUPAS IMINO
With auxiliary setae in most or all of Lika! Were ee cA ee see
4. With no definite cerarii anterior to the last 5-6 abdom-
inalsegments. . . . . . . aurilanatus ( Mask.)
With cerarii on the head and ‘thorax :!.0200.0.. °V0 ae

5. Cerarian spines in all except the anal lobe cerarii
tending to be widely separated, dorsal body setae
short, stout and conical . . . . . mtpae (Mask.)
Cerarian spines close together in all the cerarii, dorsal
body setae slender or at least not short and conical. . . . 6

6. With a large, well defined, chitinized area surround-
ing thé anal lobe cerarii . . . timberlakei (CkIl.)

10.

$i:

12.

13.

14.

15.

16.

SYSTEMATIC TREATMENT

Without a definite chitinized area about the anal lobe
cerarii

. Pores of the anal lobe cerarii numerous, arranged in

a crowded, circular area about the cerarian spines
Pores of the anal lobe cerarii few, not thus arranged .

. With no cerarii on the head anterior to the eyes, in all

with not more than 9-10 pairs . . eriogoni (Ehrh.)
With cerarii on the head anterior to the eyes, in all
with 17 pairs .

With a narrow, chitinized bar on the ventral side of the
anal lobes, lobes extending in from the base of the
anal lobe seta Pa he Chr tutti oka! pc) itt Gh

Without such a structure . . . . sequoiae (Coleman)

Tips of cerarian jae and dorsal body setae flagellate

4 : Noe krauhniae (Kuw.)

Tips of cerarian spines and dorsal body setae not

flagellate . citrt (Risso)

v

With not less than 15 pairs of cerarii. . . ryant (Coq.)
With not more than 11 pairs of cerarii cupressicolus n. sp.

With a well defined, chitinized area surrounding the
anal lobe cerarii

With no chitinized area orGatae the val ‘pbs
cerarii yt Beg papel CNTR CAEL RNAS ae et Ayal

With a chitinized area on the ventral side of the anal
lobes

Without a chieuised: area on lite Genital aay of the
anallobes . .... . ... cupressicolus n. sp.

Pores of the anal cerarii arranged in a crowded, cir-
cular group; id setae short, inconspicuous .
maritimus (Ehrh.)
Pores of the nee cerarii aeabhered, body setae numer-
ous and long, giving the body a conspicuously
hairy appearance . . . . . . Jongisetosus n. sp
With chitinized areas surrounding both: penultimate
and anal lobe cerarii
With a chitinized area Se omiaine the ‘gua toe
cerarii only

Pores of the anal lobe cerarii hepa in a crowded,

circular group. . . . . maritimus (Ehrh.)
Pores of the anal lobe cerarii scattered

37

10

15

13

14

18

16

17

38 THE CALIFORNIA SPECIES OF MEALY BUGS

17. With a chitinized area on the ventral side of the anal

lobes seo MW Ge ae! commstocks ( Kitw.o
Without such a structure . . . . . quercicolus n. sp.
18. Pores of the anal lobe cerarii arranged in a crowded,

circular group . .. . .. . Jlongispinus (Targ.)
Pores of the anal lobe cerarii scattered . Tega
citrophilus (Claussen )

Pseudococcus affinis (Maskell).

1914. Pseudococcus affinis (Maskell) ; Essig, Mon. Bull. Calif. State Com. Hort.
3 :106.

Essig states that the U. S. Dept. of Agriculture has recorded this
species as attacking Anemone sp. in California. However, there is no
other record of its occurrence in the state and it is not considered in the
present paper.

Pseudococcus aurilanatus (Maskell).
1889. Dactylopius aurilanatus Maskell, Trans. New Zealand Inst. 11:151.
1915. Pseudococcus aurilanatus (Mask.); Essig, Inj. & Benef. Ins. Calif., 2d ed.,
125, fig. 106.

In life. Of a deep red or purple color, the waxy secretion of a
bright yellow color and arranged in tufts on the dorsum and along the
margins. Ovisac short, quite firm, white.

Morphological characteristics. Body contents black; derm blue-
green; cerarii present on the last five or six abdominal segments only,
each with two cerarian spines, those in the anteriormost pairs are so small
and so widely separated as to be scarcely recognizable. Anal lobe cerarii
surrounded by a small, indefinite, chitinized area and accompanied by
two or three slender setae and a few pores. Remaining cerarii with
neither grouped pores nor auxiliary setae. Dorsal body setae small, con-
ical, somewhat constricted at the base, scattered, few in number except
on the head. Tubular ducts few, very small, without a raised rim about
the mouth. Anal lobe setae about equal to the anal ring setae. Antennae
7-8-segmented.

Type host and locality. From Araucaria in Australia.

Host and distribution. Essig states that this species occurs on dif-
ferent species of Agathis as well as on Araucaria. Its out-of-door range
in this state is not definitely known, but it occurs on Araucaria out of
doors at Mountain View in Santa Clara County.

Authentication. Not authenticated. Agrees with original descrip-
tion.

Notes: In many respects this species departs rather widely from typical

Pseudococcus and in part it is morphologically much nearer Erium or even Tri-
onymus. I shall not, however, attempt to remove it from Pseudococcus.

SYSTEMATIC TREATMENT 39

, Pseudococcus azaleae (Tinsley).

1898. Dactylopius azaleae Tinsley, Can. Ent. 30:319,
1914. Pseudococcus azaleae (Tins.); Essig, Mon. Bull. Calif. State Com. Hort.
3 :108.
This species was originally described from specimens taken from
azalea in a Japanese nursery at San Jose. It is not now known to occur
in this state and is not here considered.

Pseudococcus citri (Risso).
Plate 1, figs. 7, 9.

1910. Pseudococcus cttri (Risso.); Essig. Pomona Col. Jn. Ent. 2, no. 4:289-320,
figs. 111-119.

Note: The literature in regard to this species is so voluminous that only the
reference to the above admirable article is given.

In life. Covered with thick white cottony wax, the marginal tassels
short but distinct, becoming somewhat longer posteriorly with the caudal
pair equaling perhaps one-fourth the length of the body. Eggs deposited
in large, more or less irregular masses. .

Morphological characteristics. With 17 pairs of cerarii, none of
which contain more than two conical spines, these quite large, the anal
lobe pair not exceeding in size those preceding it. All the cerarii without
auxiliary setae and with very few pores. Anal cerarii without a definite
chitinized area, but occasionally showing a very slight chitinization.
Ventral side of each anal lobe with a narrow, very distinct, chitinized
area extending in from the base of the anal lobe seta and bearing a small
hair at about midway of its length. Dorsal body setae scattered, quite
small and slender. Cylindrical ducts very few in number, of the small
type, without raised rim at the mouth. Anal lobe setae somewhat longer
than anal ring setae.

Immature female differing but little from the adult.

Hosts and distribution. In the northern part of the state this species
appears to be confined to greenhouses, although I have at hand specimens
taken from citrus at Marysville. In the vicinity of Stanford University
it does not appear to live out of doors. Its distribution and status in the
southern part of the state are too well known to require comment here.

Authentication. Common repute.

Notes: The only California species with which this is likely to be confused
are P. krauhniae (Kuwana) and P. eriogoni (Ehrh.). It is indeed very similar
to the former, but may readily be distinguished by the fact that the tips of none
of the spines are prolonged into a sort of flagellum and by the much scantier cov-

ering of hairs. From the latter it may always be distinguished by the presence
of cerarii between the eyes.

40 THE CALIFORNIA SPECIES OF MEALY BUGS

The chitinized area on the ventral side of the anal lobes is remarkably con-
stant in size and shape. I have examined specimens from cacao and an unknown
host from Samoa, greenhouse plants from Ohio and Stanford University and from
citrus from southern part of California and find practically no variation, except
for a very weak chitinization about the anal cerarii of the Samoan specimens.

Pseudococcus citrophilus Claussen.
Plate 1, fig. 6.

1915. Proudopocmes citrophilus Claussen, Calif. Agr. Exp. Sta. Bull. 258:30-35,
figs. 1b, 6, 7.

In life. Covered with white wax except for four longitudinal rows
of conspicuous impressed dots on the dorsum. Marginal tassels very
short and quite stout. The single pair of elongated caudal tassels are
from one-fourth to one-third the length of the body and are likewise
quite stout. Oviparous.

Morphological characteristics. 17 pairs of cerarii present, the an-
terior 3-4 pairs with three conical spines, the remainder with but two,
those of the anal pair considerably larger than those of the others. Each
lateral cerarius with numerous pores and four or more auxiliary setae.
Penultimate pair surrounded by a small circular chitinized area contain-
ing numerous pores. Anal lobe pair surrounded by a large, well defined,
more or less oval chitinized area with many pores, which are more or
less scattered, and with several long, slender setae. Ventral side of
each anal lobe with a small but distinct chitinized area extending in from
the base of the anal lobe seta, bearing two slender setae at its posterior
tip. Dorsal body setae small and slender, not numerous. Cylindrical
ducts of two sizes present, the smaller without an elevated rim at the
mouth predominating. Anal lobe setae equaling anal ring setae.

Immature female in general resembling the adult, differing in the
weaker development of the cerarii.

Type host und locality. These have not been distinctly stated (the
species has not been formally described), but are probably to be taken
as citrus, at Upland, Calif.

Hosts and distribution. The species is chiefly known as a pest of
citrus fruits in the southern part of the state. However Claussen has
recorded it from a wide range of hosts in that section. It also occurs on
oranges at Irvington in Alameda County and has been taken from laurel
in a greenhouse at Oakland.

Authentication. Metatype material examined.

Notes: The longitudinal rows of impressed dots in the living insect are

quite distinctive. Morphologically the species most nearly resembles P. comstocki
(Kuwana) from which it can be distinguished only by the presence of a chitinized

(

SYSTEMATIC TREATMENT 41

area about the penultimate cerarii as well as about the anal lobe pair. It also
somewhat resembles P. quercicolus n. sp., from which it differs in the same man-
ner as from P. comstocki and further by the presence of a chitinized ventral area
on the anal lobes.

Pseudococcus comstocki (Kuwana).
Plate 1, fig. 2.
1902. Dactylopits comstocki Kuwana, Proc. Calif. Acad. Sci. (3) 3:52.

In life. Almost identical with P. maritimus (Ehrh.), the lateral tas-
sels short and slender, the caudal pair one-third or one-half the length of
the body. Ovisac not seen.

Morphological characteristics. With seventeen pairs of cerarii, of
which the first six or seven pairs have three or four conical spines, the
remainder having but two. The spines of the anal lobe cerarii are con-
siderably larger than any of the others. Lateral cerarii with three or
four auxiliary setae and a small cluster of pores. Anal lobe cerarii sur-
rounded by a large, oval, chitinized area, which is attenuated posteriorly,
extending back to the anal lobes, and which bears several slender setae
and numerous pores, the latter showing some tendency to concentrate _
about the cerarian spines. Ventral side of the anal lobes with a small,
but constant, triangular, chitinized area extending in from the base of the
anal lobe setae. Dorsal body setae numerous, slender and rather long.
Triangular pores very abundant over the body, mingled with many tubu-
lar ducts, of which the majority are without a raised rim about the mouth.
Anal lobe setae about equaling anal ring setae or perbaps a trifle longer.

Immature female in general resembling the adult female.

Type host and locality. From mulberry, Akabane, Japan.

Hosts and distribution. Not previously recorded from the U. S. I
have specimens taken by Professor Doane from mulberry and maple on
Staten Island, N. Y., and also specimens from Pimus radiata on the
campus of Stanford University.

Authentication. Type specimens examined.

Notes: It is reasonable enough to find this species on mulberry and maple
in Japan and on the same hosts in New York, for we need only assume that it has
been introduced on one of these hosts, but it is somewhat difficult to believe that
the species taken from pine in California is the same. However, I can find not
the slightest ground upon which to effect a separation and I do not propose to
brave the pitfalls which one will surely encounter if he disregard the evidence of
morphology and venture to discriminate between species upon the shifting and
inconsistent basis of hosts and biology alone.

I have taken this species in California only upon Pinus radiata upon the
_ Stanford campus, where it is quite common, although by no means abundant.

. This is very near P. citrophilus Claussen, from which it differs only in the
absence (in life) of the longitudinal rows of impressed dots on the dorsum and
the absence of a chitinized area about the preanal cerarii.

42 THE CALIFORNIA SPECIES OF MEALY BUGS

Pseudococcus crawii (Coq.).
Plate 1, fig. 1.

1889. Dactylopius crawit Coquillet, West. Amer. Scientist. 6:123. “(As to the
female. )

1900. Dactylopius quercus Ehrhorn, Can. Ent. 32:220.

1914. Pseudococcus crawii (Coq.); Essig, Mon. Bull. Calif. State Com. Hort.
3:117, fig: 32.

In life. “The bodies of the females are light yellow and covered with
thick plates of white cottony material which entirely hides the color. The
lateral filaments are very distinct, being about one-fourth as long as the
width of the body. The anal filaments vary from one-third to one-half
the length of the body. They usually form a sharp angle at the posterior
end and in some individuals the ends are curved inwardly.” (Essig.)
The females are viviparous, the young being placed in a pad-like ovisac.

Morphological characteristics. Seventeen pairs of cerarii present, all
with more than two conical spines, the anal lobe pair containing 15 to 20,
the others from 3 to 6. The lateral cerarii are without auxiliary setae
and are accompanied by a small cluster of pores. The anal lobe pair are
borne upon a large, nearly circular and somewhat protuberant chitinized
area, the spines being accompanied by numerous scattered pores and
several slender setae. In some cases the penultimate cerarii are also sur-
rounded by a more or less distinct area. Ventral side of the anal lobes
without chitinized area. Dorsal body setae very few, scattered and small.
Numerous triangular pores scattered over the body, mingled with rela-
tively few, large, cylindrical ducts. Anal lobe setae equaling anal ring
setae. .

Immature female differing but little from the adult.

Type host and locality. From Ramona polystachya (white sage)
near Los Angeles, Calif.

Hosts and distribution. Common on the white sage throughout the
southern part of the state, and according to Essig, occurring on the Cali-
fornia sage (Artemisia californica) also. Also from Quercus chrysolepis
and Pasania densiflora in the vicinity of Stanford University, and from
Adenostoma fasciculatum in Marin County.

Authentication. Not authenticated. Specimens from the type host
and location agree with the original description and belong to what is
commonly accepted as this species.

Notes: Specimens of two species of Pseudococcus taken by me from Quercus
chrysolepis near Stanford University were sent to Mr. Ehrhorn who identified one
of them as his Pseudococcus quercus. I can find no basis for separating this from
P. crawii in spite of the apparent anomaly presented by the wide difference in

hosts. It is perhaps the most distinctive species occurring in the state and cannot
be confused with any other.

-

SYSTEMATIC TREATMENT 43

I do hot know that any notes on the male other than those in the original
description by Coquillet have ever been published and I have not seen the male
myself. However, it seems almost certain that the male described by Coquillet
really belongs with Puto yuccae (Coq.), a species that also occurs upon the white
sage, for this description applies throughout to a male of the latter genus.

Pseudococcus cupressicolus n. sp.

Plate 1, fig. 11.

In life. Living specimens have not been examined, but judging from
the morphology of the species the insect will bear a pair of slender caudal
tassels of perhaps one-half the length of the body, a series of much shorter
tassels along the lateral margins of the abdomen with perhaps two or
three very short pairs on the head.

Morphological characteristics. Antennae 7-8-segmented, both num-
bers frequently appearing in the same individual. Not more than 11
pairs of cerarii present, these appearing in a quite definite arrangement
as follows: a pair on each abdominal segment, a pair opposite each ante-
rior spiracle and each anterior leg, an ocular pair and a pair anterior to
the eyes. There is a tendency for the cephalic and thoracic pairs to be
lacking and any of these pairs may at times be present on one side of the
body and not on the other. The cerarian spines are small in all except
the anal lobe pair, in which they are conspicuously larger. The anal lobe
cerarii include a circular cluster of many crowded pores which are not
surrounded by a chitinized area. Remaining cerarii with few pores, either
with or without two or three auxiliary setae, for the most part with but
two cerarian spines, but at times with three. No chitinization on the ven-
tral side of the anal lobes. Dorsal body setae numerous, very slender and
quite long. Tubular ducts not strikingly abundant, all small and without
a raised rim about the mouth. Anal lobe setae about equaling the anal
ring setae.

Immature specimens differing from the adult in the reduction of the
number of pores in the anal cerarii and in a tendency for the cerarian
spines to be slender and elongated.

Type host and locality. From Cupressus guadelupensis at Riverside,
Calif. A. F. Swain, collector.

Hosts and distribution. From the type host and from Cupressus
arizonicus at Riverside.

Notes: Morphologically this most nearly resembles P. ryani, but it may
readily be separated by the reduced number of cerarii, by the much more numerous
dorsal body setae and by the marked tendency toward the development of auxiliary
setae in connection with the cerarii.

‘44 THE CALIFORNIA SPECIES OF MEALY BUGS

Pseudococcus eriogoni Ehrh.

1899. Dactylopius eriogoni Ehrhorn, Can. Ent. 31:103.
1902. Erium eriogoni (Ehrh.) ; Cockerell, Ann. Mag. Nat. Hist., (7), 10:465.
1913. Pseudococcus yerba-santae Essig, Pomona Col. Jn. Ent. 2:85.

In life. Adult female enclosed in a more or less definite sac. Lateral
tassels very short, confined to the posterior part of the body.

Morphological characteristics. Antennae of the adult female nor-
mally 8-segmented, sometimes 7-segmented. Cerarii reduced in number,
there being but eight or nine pairs, counting forward from the anal lobes,
and usually an ocular pair, but none on the head between the eyes. The
ocular pair may contain several cerarian spines, the remaining cerarii
have but two, all being quite small, the anteriormost almost merging with
the body setae. In none are there any auxiliary setae except in the anal
lobe pair, which has 3-5 small hairs, and in none is there any grouping of
the pores. In some specimens there is a very small chitinized area on
the ventral side of the anal lobes which in extreme cases somewhat re-
sembles that found in P. citri, but this is variable in size and sometimes
entirely lacking. Dorsal body setae quite numerous, slender. Triangular
pores abundant, mingled with many large cylindrical ducts with a raised
rim about the mouth, these in many specimens being quite striking by
reason of their abundance. Anal lobe setae about twice as long as the
anal ring setae.

Immature female in all essential respects resembling the adult.

Type host and locality. From the roots of Eriogonum sp. Stevens
Creek, Santa Clara County, Calif.

Hosts and distribution. From Eriogonum nudum, Stevens Creek,
Santa Clara County; Viola sarmentosa at the reservoir on Big Creek,
Santa Cruz County; Stachys sp., Corte Madera Creek, near Stanford
University; Eriodictyon californicum, Sespe Canyon, Ventura County;
Eriodictyon tomentosum, Del Mar; ragweed, Upland, Calif. ; Ceanothus
cuneatus, southern California; Erigeron sp., locality unknown.

Authentication. Material from the type host and locality, agreeing
with the original description.

Notes: I have not seen the types of this species, but I have at hand speci-
mens taken from the roots of Eriogonum in the type locality and agreeing in all
respects with the original description, which was probably based upon immature
specimens. I have also examined the types of P. yerba-santae Essig and of P.
leptodontis Claussen MSS. and am unable to regard either of these as in any way
distinct.

While this is not a typical Pseudococcus its affinities appear to be entirely
with that genus. The absence of cerarii between the eyes and along the anterior
portion of the lateral margins should be sufficient to prevent confusion with any

other species. The sac is by no means so definitely formed as the original de-
scription would indicate, the insects leaving it readily when disturbed.

SYSTEMATIC TREATMENT 45

Pseudococcus ephedrae (Coq.).
1890. Dactylopius ephedrae Coquillet, West Amer. Scientist 7:43.

In life. “Adult female elongate-ellipsoidal, from two and a half to
three times as long as broad, dark olive, almost black, thinly covered with
a snow-white mealy powder not entirely concealing the ground color;
cottony appendages confined to posterior end of body, the longest less
than half the length of the body... .’ “The adult female secretes a
layer of white cottony matter from the lower part of her body, and this
is gradually extended upward until finally the entire insect is enclosed in
a cottony sac.” (Coquillet.) © ~

Morphological characteristics. According to Coquillet’s description
the antennae are 8-segmented and the claw is without a denticle.

Type host and locality. From Ephedra californica in Los Angeles
County.

Hosts and distribution. Known only from the original description.

Notes: This species has not been seen since it was originally described, and
nothing can be added to the description. It may possibly be an Erium.

Pseudococcus krauhniae (Kuwana).

Plate 1, fig. 8. (Also see plate 1, fig. 7.)
1902. Dactylopius krauhniae Kuwana, Proc. Calif. Acad. Sci. (3), 3:55, pl. 9,
figs: 39-40.

In life. “Enclosed in a.cottony sac of irregular shape,” (Kuwana).

Morphological characteristics. Seventeen pairs of cerarii present,
all with but two spines, without auxiliary setae and with but very few
pores. The cerarian spines are all practically of the same size and in all
the tip is produced into a sort of flagellum. No chitinization associated
with any of the cerarii. Ventral side of the anal lobes with a narrow chit-
inized bar extending in from the base of the anal lobe setae. Dorsal body
setae very numerous, stout at the base, quite long and with the tips more
or less flagelliform. Triangular pores numerous, intermingled with many
small cylindrical ducts without a raised rim about the mouth. Anal lobe
setae perhaps one and a half times as long as the anal ring setae.

Immature female not seen.

Authentication. Types examined.

Type host and locality. From Krauhnia floribunda, Yokohama,
Japan.

Hosts and distribution. Among the material received from Mr. Essig
is one slide labeled “From wisteria and persimmon,” Nordhoff, which is
unmistakably this species, the types of which are in the Stanford collec-
tion. It has previously been recorded only from Japan and once from
quarantine in New Jersey, the latter record needing verification.

46 THE CALIFORNIA SPECIES OF MEALY BUGS

Notes: Morphologically this is very close to P. citri, but the curiously flagel-
liform spines and hairs, together with the abundance of dorsal body setae, is suffi-
cient to separate it at once. It might perhaps be regarded by some as a “variety”
of P. citri, but I believe it to be sufficiently distinct. | ?

Pseudococcus longisetosus n. sp.

Plate 1, fig. 10.

In life. Thickly covered with white powdery wax, with short, stout
lateral tassels which become somewhat longer posteriorly. Ovisac not
observed.

Morphological characteristics. Antennae either 7- or 8-segmented in
the adult female. With from 15 to 17 pairs of cerarii, the normal num-
ber, 17, generally being obscured either by the dropping out of one or two
pairs or their merging with the body setae. The first three or four cerarii
have usually three conical spines and the remainder two, although there
is considerable variation, but the normal number for most of the cerarii
is two, those of the ultimate pair being somewhat larger than any of the
others. All cerarii with prominent auxiliary setae and numerous pores.
The anal lobe cerarii have many scattered pores, and several long, stout
hairs. Ventral side of the anal lobes with a small and rather irregular
chitinized area, bearing two or three slender setae at its posterior margin.
Dorsal body setae very numerous and unusually long, especially on the
abdomen. Derm thickly beset with triangular pores and small cylindrical
ducts without a rim about the mouth. Anal lobe setae about equal to the
anal ring setae in length and accompanied by several minor setae of nearly
their own length.

Immature female in general resembling the adult, but somewhat less
hairy.

Male not definitely recognized, due to the association of this with
another species, but an apterous male was observed that may perhaps be-
long with this.

Type host and locality. From beneath a stone, associated with
ants, (Cremastogaster lineolata) on the mountain road above Woodside,
San Mateo County, Calif., April, 1917.

Hosts and distribution. From the type station and from the under-
ground stems of Castilleia foliolosa and Orobanche tuberosa, Coal Mine
Ridge, near Stanford University, most abundant upon the latter host, and
from roots of Armeria vulgaris on the cliffs at Pacific Grove.

Notes: In general this species most nearly resembles P. maritimus, differing,
however, in the rather striking vestiture of long hairs and in the less marked con-
centration of the pores about the conical spines of the anal lobe cerarii.

SYSTEMATIC TREATMENT 47

On the basis of hosts this might be P. aphyllonis Ckll. which is recorded from
Orobanche (Aphyllon) fasciculatum at Wentachee, Wash., but Mr. Morrison has
- furnished me with a sketch of the type of that species which proves that the two
are quite different. The description of P. formicarii Ehrh. suggests some resem-
blances to that species.

Pseudococcus longispinus (Targ.).
Text figure 13; plate 1, fig. 4.

Note: The literature and synonymy of this species are so extensive that it is
hardly desirable to present them here.

In life. Remarkable for the unusual development of the lateral and
caudal tassels, those on the margins of the body equaling in length nearly
half the width of the body and the caudal pair being at times actually
longer than the body. Oviparous, the ovisacs more or less loose and
elongated.

_ Morphological characteristics. Seventeen pairs of cerarii present,
the first three or four pairs of these with three or four conical spines, the
remainder with two. These increase gradually in size toward the pos-
terior end of the body until the penultimate pair are reached and of
these the spines are much larger than any preceding, while those of the
anal lobe pair are still larger, being extremely stout. All of the cerarii
with auxiliary setae and numerous pores. In the anal lobe and penulti-
mate pairs the pores are concentrated into a crowded circular area about
the cerarian spines. Penultimate cerarii surrounded by a circular chitin-
ized area and the anal lobe pair by a large well-defined oval area. Ven-
tral side of the anal lobes with a large, triangular chitinized area with the
apex directed forward and with a narrow thickening along its median
edge. Dorsal body setae few, slender. Triangular pores not numerous,
intermingled with scattered tubular ducts of which a few have a raised
rim about the mouth. Anal lobe setae shorter and more slender than
anal ring setae.

Immature females differing from the adult chiefly in a slight reduc-
tion in the number of pores.

Hosts and distribution. Upon a very considerable range of hosts,
out of doors in the southern part of the state. In the bay region it occurs
only in greenhouses, where it chiefly infests ferns.

Authentication. Common repute.

Notes: The characters of the anal lobes are such that this species can
hardly be confused with any other found in the state. I do not propose to enter
into the discussion as to whether this should be called longispinus or adonidum. I
retain the better known name.

48 THE CALIFORNIA SPECIES OF MEALY BUGS

Pseudococcus maritimus Ehrhorn.
Text figure 11; plate 2, fig. 13.

1900. Dactylopius maritimus Ehrhorn, Can. Ent. 32:315.

1909. Pseudococcus obscurus Essig, Pomona Col. Jn. Ent. 1:43.

1909. Pseudococcus bakeri Essig, Ibid. 2:334, fig.

1917. Pseudococcus omniverae Hollinger, Ann. Ent. Soc. Am. 10:271, fig. 31.

In life. Covered with thick, white, waxy powder, the lateral tassels
rather short and slender, equaling perhaps one-fourth the width of the
body, the caudal tassels longer, equaling at times half the length of the
body. Eggs laid in loose, white, cottony masses.

Morphological characteristics—With 17 pairs of cerarii of which
the first two or three pairs have three or four cerarian spines, the remain-
der but two, the spines becoming larger toward the posterior end of the
body. All cerarii with auxiliary setae and with numerous pores, the
pores in the anal lobe pair being concentrated into a crowded, circular
area about the cerarian spines. In some cases the anal lobe cerarii are
surrounded by a weakly indicated chitinized area, in others this is entirely
lacking. Ventral side of the anal lobes with a well-defined somewhat
triangular chitinized area, varying somewhat in size but always present.
Dorsal body setae few, slender. Triangular pores more or less numerous,
intermingled with a few tubular ducts without a raised rim about the
mouth. Anal lobe setae slightly shorter and slenderer than the anal ring
setae.

Immature female differing but little from the adult.

Type host and locality. From roots of Eriogonum latifolium on the
cliffs at Santa Cruz, Calif.

Hosts and distribution. I have at hand specimens of this species
from wild cherry near New York City (R. W. Doane coll.) and from
roots of clover at Medford, Ore., (M. N. Reeher coll.). It has been re-
corded from Missouri under the name of P. omniverae Hollinger. Within
the state of California the list of hosts and localities is limited merely ©
by the collecting that has been done. In the vicinity of Stanford Uni-
versity I have taken the species from tomato, roots of carnation, passion
flower, Quercus agrifolia, Magnolia, sp., maple, Crataegus and Pinus
radiata, In the vicinity of Fresno it is a pest on the grapes, and in the
southern part of the state it is a serious enemy of the citrus fruits. It is
probable that there is almost no cultivated or wild flowering plant upon
which it will not feed.

Authentication. Specimens compared with type by Mr. Ehrhorn.

Notes: Specimens taken by me from the type host at Santa Cruz were
forwarded to Mr. Ehrhorn, who identified them as his species, P. maritimus. I

SYSTEMATIC TREATMENT 49

have also examined the types of P. obscurus Essig and P. bakeri Essig, and can
find no basis upon which to separate them from maritimus. This conclusion is
supported by the fact that specimens taken from the type host and forwarded to
Mr. C. P. Claussen were by him reared on green lemons. He informs me that he
could not distinguish these from ordinary P. bakeri.

Mr. Claussen has proposed to separate the species occurring on grape at
Fresno, being moved to this action chiefly by the behavior of certain parasites. In
this I cannot concur.

It is unfortunate that a name so familiar as is P. bakeri should be replaced
by one which has appeared but once in our literature, but there is no alternative.
Even yet it is by no means certain that this name will stand, as the occurrence of
the species in the Eastern states suggests the possibility that it has been described
under some other name there. In fact I am inclined to suspect that the species which
has passed as the “summer form” of P. trifolii Forbes may be identical with this.

I have received specimens of P. omniverae Hollinger through the kindness
of Mr. Hollinger, and although the specimens are not strictly comparable due to
differences in treatment, I regard this also as a synonym of P. maritimus.

Pseudococcus nipae (Maskell).
Plate 2, fig. 12.

1892. Dactylopius nipae Maskell, New Zealand Trans. 25 :232.

1897. Dactylopius pseudonipae Cockerell, Science Gossip, N.S. S::189, 302.

1908. Pseudococcus nipae (Maskell) ; Marchal, Ann. Soc. Ent. Fr. (8), 8:236-39,
figs. 7-8.

1914. Pseudococcus pseudonipae (Ckll.); Essig, Mon. Bull. Calif. State Com.
Hort. 3 :120-1, fig. 34.

1915. Pseudococcus pseudonipae (Ckll.); Essig, Inj. & Benef. Ins. Calif. 2d ed.,
132.

1917. Pseudococcus nipae (Maskell); Green, Ent. Mon. Mag. 53:262-63, fig. 1.

In life. Covered with yellowish secretion arranged in conical humps.
Divested of secretion the body is of an amber or orange-yellow color.

Morphological characteristics. The number of pairs of cerarii is
somewhat difficult to determine, but there appear to be from 12 to 15 in
all of which the cerarian spines are stout and conical In all except the
anal lobe pair the cerarian spines are very widely separated and are rec-
ognizable only by their paired character, being unaccompanied by either
grouped pores or auxiliary setae. In the anal lobe cerarii the cerarian
spines are close together and are accompanied by two or three slender
setae and a few grouped pores. Anal lobes quite prominent and with a
small chitinized area on the ventral side extending in from the base of
the anal lobe seta. Dorsal body setae few, rather small, stout, conical.
Tubular ducts few, small without raised rim about the mouth. There
appear to be no definite structures corresponding to the lumps of wax.
Anal lobe setae somewhat shorter than the anal ring setae. Antennae
normally 7-segmented. Legs rather short and stout.

50 THE CALIFORNIA SPECIES OF MEALY BUGS

Immature female differing from the adult chiefly in the fact that the
cerarian spines are closer together.

Type host and locahty. From Nipa fruticans, Demerara, British
Guiana. .

Hosts and distribution. This is a common greenhouse species occur-
ring on palms. It is not known to occur out of doors in this state.

Authentication. Not authenticated. It is the species that commonly
passes as P. nipae and P. pseudonipae. It agrees fairly well with the
original description and very well with the eee of P. nipae given
by Marchal and Green.

Notes: I can see no basis for the separation of P. nipae (Maskell) and P.

pseudonipae (Ckll.). Specimens from Hawaii which pass there as P. nipae are
certainly not different from those which pass here for P. pseudonipae.

Pseudococcus quercicolus n. sp.
Plate 2, fig. 18.

1910. Pseudococcus agrifoliae Essig, Pomona Col. Jn. Ent. 2:147-148, figs. 60B,
60C. (Not of Essig, Ibid. 1:42.)

1911. Pseudococcus agrifoliae Essig; Smith, Ann. Ent. Soc. Am. 4:316.

1913. Pseudococcus agrifoliae Essig; Smith, Pomona Col. Jn. Ent. & Zool. 5:78.

In life. With rather long marginal tassels and a caudal pair of at
least half the length of the body. Ovisac not observed.

Morphological characteristics. With 17 pairs of cerarii of which the
anterior 3-4 pairs have 3-4 cerarian spines and the remainder but 2, those
of the anal lobe pair much larger than any of the others. Lateral cerarii
each with a small cluster of pores and with three or four auxiliary setae.
Anal lobe cerarii surrounded by a distinct chitinized area, which does not
extend back to the base of the anal lobe hair and which contains numerous
scattered pores. Ventral side of the anal lobes without a chitinized area.
Dorsal body setae, few, slender. Anal lobe setae about equaling anal ring
setae.

Immature female in general closely resembling the adult.

Type host and locality. From Quercus chrysolepis, Stevens Creek,
Santa Clara County, Calif.

Hosts and distribution. From the type host and from Pasania densi-
flora in the vicinity of Stanford University and from Quercus agrifolta
near Claremont, Los Angeles County, Calif. There appears to be but
one generation a year, this maturing in May or June in the vicinity of
Stanford.

Notes: The problem of the proper name for this species presents a curious

tangle. I have at hand the type slides of Pseudococcus agrifoliae Essig. The slide
labeled “Type” contains mature females of the species herein described as P. quer-

SYSTEMATIC TREATMENT 51

cicolus n. sp. Another slide labeled “cotype” contains immature specimens of a
species of Puto. Other slides not indicated as type material contain specimens of
Trionymus villosa (Ehrh.) either alone or in company with immature specimens of
P. quercicolus. None of the slides bear any date, but I am informed by Mr. Essig
that they were not all collected at the same time.

The original description of the species cannot apply to the specimen labeled
“type,” for this description indicates that the antennae were 7-segmented. It does
apply to Trionymus villosa (Ehrh.). The so-called type was evidently selected sub-
sequent to the publication of the original description from the specimens described
by Essig as the “summer form” of P. agrifoliae. In view of this it appears that the
name agrifoliae cannot be used for this species, and for it I propose the name quer-
cicolus. . ‘ae

It further remains to be decided what is to be done with the name agrifoliae.
It probably is a synonym of T. villosa (Ehrh.), although none of the specimens of
this species are included in the material bearing any type designation. The name
might be used for the species of Puto, but there can be but little if any question
that this is either P. yuccae (Cogq.) or P. koebelei Ehrh., so that in any case the name
becomes a synonym. Inasmuch as the original description applies especially to T.
villosa I regard agrifoliae as a synonym of this and place the species of Puto as a
misidentification.

Pseudococcus ryani (Cogq.).
See plate 1, fig. 11.

1889. Dactylopius ryani Coquillet, West Amer. Scientist 6:122-123.
1903. Dactylopius andersoni Coleman, Jn. N. Y. Ent. Soc. 11:62-3, pl. 5, figs. 3-4.
1903. Dactylopius dudleyi Coleman, Ibid. 11:63-4, pl. 5, figs. 5-8.

In life. Covered with white, powdery wax, the lateral tassels very
short, the caudal pair from one-half to one-third the length of the body.

Morphological characteristics. With 15-17 pairs of cerarii, of which
the anterior 2-3 pairs have three conical spines and the remainder but
two, these spines in all the cerarii being very small except for those of
the anal lobe pair which are conspicuously larger than the others. All
the cerarii without auxiliary setae and with a cluster of but a half-dozen
pores, except the last which has several small hairs and numerous pores
concentrated into a circular area about the conical spines, this area being
slightly chitinized. Ventral sides of the anal lobes sometimes with a very
small isolated chitinized area which may, however, be entirely lacking.
Dorsal body setae few, small and slender. Triangular pores few; tubular
ducts few, small, for the most part without raised rim about the mouth.
Anal lobe setae about equaling anal ring setae.

Immature female not observed.

Type host and locality. From Cupressus macrocarpa near Los An-
geles.

Hosts and distribution. Recorded by Coquillet from Thuya orientalis
and Araucaria excelsa as well as from the type host near Los Angeles.

s

52 THE CALIFORNIA SPECIES OF MEALY BUGS

Also from Cupressus goveniana, southern part of Lake County; Cupres-
sus macnabiana near Shasta postoffice, Shasta County; Cupressus
macrocarpa at Pasadena; Libocedrus decurrens, Scott Valley, Siskiyou
County; Sequoia sempervirens near Stanford University.

Authentication. Specimens compared with the type by Mr. Mor-
rison.

Notes: This name has ordinarily been employed for the species which must
stand as P. sequoiae (Coleman). Mr. Morrison’s comparison of specimens with
the type, together with the fact that the specimens seen by me in life agree with
the original description, seems to establish the identity of the species satisfactorily.
The types of P. dudleyi (Coleman) and P. andersoni (Coleman) are at hand and
I can see no basis for separating them.

This species is near P. sequotae (Coleman), but is distinguishable in life by
the rather long caudal tassels and in preparations by the clustered pores of the anal
lobe cerarii.

Pseudococcus salinus Ckll.
Plate 1, fig. 5.
1896. Dactylopius salinus Cockerell, Ann. Mag. Nat. Hist., (7), 9:21.

In life. “Grey, with some white secretion, six caudal tassels and two
cephalic ones.” (Ckll. 1896.) Found on the upper side of the leaf blades
of the host.

Morphological characteristics. Apparently not more than 11 pairs
of cerarii present, four of these being on the head and seven at the poste-
rior end of the body. In the posterior seven pairs the cerarian spines of
the anal lobe pair are the largest, the others becoming successively smaller
anteriorly. First two cephalic pairs with three spines, the remainder with
but two. Anal lobe pair with a group of rather numerous pores, the re-
mainder with fewer. Auxiliary setae present in conjunction with the last
two pairs only. Dorsal body setae very few, small, slender. Triangular
pores quite numerous, mingled with a few large tubular ducts with rim
about the mouth. Anal ring very weakly chitinized, simple except for
the posterior half which is weakly chitinized and shows faint indications
of the usual cellular structure. Due to the chitinization of the end of the
alimentary canal, the ring usually presents a distinctly crescentic appear-
ance. The position of the ring is quite unique, inasmuch as it lies be-
hind the anal lobe cerarii and in all of the numerous specimens examined
by me appears on the ventral side of the abdomen when the specimens are
flattened out on the slide. Anal lobe and anal ring setae all noticeably
short, the former about half the length of the latter.

Type host and locality. From grass (probably Distichlis) on the
cliffs at La Jolla, Calif.

SYSTEMATIC TREATMENT 53

Hosts and distribution. From Distichlis spicata in the salt marshes
at Palo Alto. Probably occurs throughout the range of this host at least.

Authentication. Specimens compared with the type material by Mr.
Morrison. |

Notes: I have examined specimens (from Essig collection) collected at La
Jolla and determined by Cockerell as P. salinus, but which are evidently not P.
salinus, belonging instead to the species herein described as Trionymus distichlii
n. sp. The most striking features of P. salinus and those which at once separate
it from any other are the almost simple anal ring, the extremely short anal lobe
and anal ring setae, and the fact that the anal ring is borne upon the ventral side
of the abdomen instead of on the dorsal side.

Pseudococcus sequoiae (Coleman).

Plate 1, fig. 3.

1901. Dactylopius sequoiae Coleman, Proc. Calif. Acad. Sci., (3), 2:410-418,
pl. 27. (As to the female.)

1910. Pseudococcus ryani (Coq.); Essig, Ponoma Col. Jn. Ent. 2:148, fig. 61.
(Not of Coquillet.)

In life. Thinly covered with powdery secretion, the lateral and
caudal tassels practically lacking. Oviparous, the eggs placed in a very
long, slender and somewhat fluted ovisac.

Morphological characteristics. With 17 pairs of cerarii the first
three or four pairs with 3 spines, the remainder with but 2, all very
small, those of the anal lobe pair somewhat the largest. The lateral
cerarii without auxiliary setae and without grouped pores, so obscure as
to be distinguishable only with difficulty in the adults. Anal lobe pair
with three or four slender setae, the base of each of these and of each
of the conical spines being surrounded by a little chitinous ring; with
very few pores, these not grouped. Ventral side of the anal lobes without
chitinized area. Dorsal body setae few, very small. Triangular pores
few. Small cylindrical ducts without a raised rim about the mouth ex-
tremely abundant, especially at the lateral margins of the body. Anal
lobe setae somewhat shorter than anal ring setae, accompanied by two
very small minor setae.

Immature female differing but little from adult.

Type host and locality. From Sequoia sempervirens, near Stanford
University.

Hosts and distribution. On Cupressus macrocarpa throughout the
bay region, at Santa Cruz and at Cypress Point near Monterey. From
Sequoia sempervirens near Stanford University. There appears to be
but one generation a year, this, in the vicinity of Stanford, maturing
early in June.

Authentication. Types examined.

54 THE CALIFORNIA SPECIES OF MEALY BUGS

Notes: An examination of the types of Pseudococcus sequoiae (Colem.) in
the Stanford collection shows that this species was based upon the male of a
species of Puto and.the female of a species of Pseudococcus. As this male was
later redescribed by Coleman under the name of Pseudococcus cupressi the name
sequoiae must be retained for the female of the original description. This is the
species which has ordinarily passed as P. ryani (Coq.), but that name appears
rightfully to belong to another species.

Pseudococcus timberlakei Ckll.
Plate 2, fig. 19.
1916. Pseudococcus timberlakei Ckll., Jn. Econ. Ent. 9:312.

In life. “The lateral and caudal secretions of the active females
closely resemble those of Pseudococcus citrophilus Claussen, figured on
page 20, Calif. Exp. Sta. Bull. 258” (Timberlake’s notes, quoted by
Cockerell).

Morphological characteristics. With 17 pairs of cerarii, all with
but two cerarian spines which are quite small except in the anal lobe
cerarii in which the spines are large and stout. Lateral cerarii with few
or no grouped pores and, except for the posterior one or two pairs, with-
out auxiliary setae. Anal lobe cerarii surrounded by a conspicuous defi-
nite chitinized area bearing numerous long setae and many scattered
pores. Ventral side of anal lobes with a small irregular chitinized area.
Dorsal body setae numerous, quite long and slender. Triangular pores
very few. Dorsum with many small cylindrical ducts without a raised
rim about the mouth. Anal lobe setae and anal ring setae about equal,
quite long. Anal ring of the usual type.

Type host and locality. From “salt marsh grass’ (probably Dts-
tichlis spicata) at Milbrae, San Mateo County, Calif., October.

Hosts and distribution. Known only from the San Francisco Bay
region. I have taken it from Dvstichlis spicata in the salt marshes at
Palo Alto.

Authentication. Specimen compared with type by Mr. Morrison.

Notes: This differs markedly from P. salinus (Ckll.) which occurs upon
the same host, the well developed anal ring alone being sufficient to distinguish it
from that species. It really most nearly resembles Trionymus californicus Ehrh.
although the two are regarded as generically separate because of the difference in
the number of cerarii. A further discussion of this resemblance will be found
under the description of T. californicus. The species appears to be rather rare,
at least much more so than either P. salinus (Ckll.) and Trionymus distichliu
n. sp., which are found with it. Like these species, there is but a single brood,
which matures in September or October.

SYSTEMATIC TREATMENT 55

Genus PHENACOCCUS Cockerell.

Pseudococcine forms with not less than 18 pairs of cerarii; with a
denticle on the inner face of the tarsal claw; with normally 9-segmented
antennae, although sometimes with 8 segments. Without a continuous
series of chitinized areas about the margin of the body. Male as in Pseu-
dococcus. Dermal pores of the triangular type present.

Type of the genus P. aceris (Sign.).

I have before me specimens determined by E. E. Green as P. acerts
(Sign.). These, together with the excellent redescription of this species
given by Marchal (7), permit somewhat more definite conclusions as to
the nature of the genus than does the original description given by Cock-
erell, who merely stated that the genus consists of forms with female
having “9-segmented antennae and secreting an ovisac.”’

The presence of 18 pairs of cerarii appears to be quite constant and
quite characteristic of the group. In some cases the cerarii are difficult
to count, due to the small size of the spines, but it is usually possible to
determine their number with sufficient surety from the immature stages.
In the case of an apparently undescribed species on beech in Canada this
number is increased to as high as 24 by the development of what may be
called secondary cerarii, but I am inclined to believe that the immature
stages of this species would show the normal number. I consider that
such species as Phenacoccus mangiferae Green cannot be regarded as
congeneric with P. acerts.

The genus Tylococcus Newstead was based entirely upon the fact
that the cerarii in the type species are elevated upon prominent tubercles
and according to the original definition of this genus and the later char-
acterization of it by Brain, certain species included by me in Phenacoccus
should be referred to Tylococcus. However, I do not believe that the
genus is tenable on the basis of this character alone, for I] have observed
that there is often a tendency for the cerarii to be more or less prom-
inent in the immature stages and in Phenacoccus artemisiae Ehrh. they
are borne upon very prominent tubercles which entirely disappear in the
adult. In the case of the undescribed species from beech, of which I have
spoken above, the tubercles persist in the adult female.

The genus as I have defined it here appears to be a perfectly natural
group, although doubtless the examination of more species will disturb
this apparent harmony. Some species are included in which the specific
characters are extremely obscure and I anticipate that the separation of
any considerable series of species will be attended by much difficulty.

56 THE CALIFORNIA SPECIES OF MEALY BUGS

KEY TO SPECIES OCCURRING IN CALIFORNIA

1. With numerous very large tubular ducts, the mouths
of which are borne at the apices of prominent,

conical projections .... 0... . . stachyos Ehrh.
Without: sich Geb ati eh Te de oe ee
2. Ducts very short and stout. . . . . kuwanae, Colem.
Ducts: Not 80 ane ees a i aba eat le Vi) er
3. Anal lobe cerarii with as many as g. 10 cerarian
Spas) Ail! . 4% erlogont Nn. sp.
_ Anal lobe cerarii Nath at the dascsct 3-4 cerarian spines . . .. 4
4. Anal lobes prominent, with the cerarian spines borne
pt their tip... !.5) . . . artemisiae Ehrh.
Anal lobe cerarii with at the bak 3-4 cerarian spines . . . . 4
borne on the dorsum. . . 2 Nah Oa Bad a aU Rca

5. Antennae of the adult female Suscotwented ;
AS ST OA eM ROM San! UES OVI BL fet os colemant Ehrh.
Antennae of the adult female 8-segmented . solani n. sp.

Phenacoccus artemisiae Ehrh.
Plate 2, fig. 15.

1900. Phenacoccus artemisiae Ehrhorn, Can. Ent. 32:313.

In life. Very slightly dusted over with waxy powder, without notice-
able lateral or caudal tassels. Oviparous, the ovisac firm, quite long and
slender. On the leaves and small twigs of the host.

Morphological characteristics. First three or four pairs of cerarii
with three to four spines, the remainder with but two, all without aux-
iliary setae. In all, the spines are quite small and are associated with
but two or three grouped pores, the cerarii being so obscure that they
are difficult to trace along the lateral margins in the mature insect. Con-
ical spines of the anal lobe cerarii borne at the tip of the prominent, teat-
like anal lobes, associated with no auxiliary setae and with no especially
grouped pores and with but the slightest indication of chitinization.
Dorsal body setae extremely minute and very few in number. Triangular
pores scattered sparingly over the body, mingled with a few tubular ducts
without a raised rim about the mouth. Anal lobe setae perhaps slightly
shorter than the anal ring setae.

Immature female very much resembling the mature female except
that the cerarii of the anterior half of the body are borne at the tips of
prominent tubercles.

Type host and locality. From Artemisia californica, Stevens Creek,
Santa Clara County, Calif.

SYSTEMATIC TREATMENT 57

Hosts and distribution. Essig has recorded this species from the
type host in Ventura County. Otherwise it is known only from the
original locality. It very probably occurs throughout the range of this
‘ host, at least.

Authentication. Specimens from type host and locality, agreeing
with original description.

Notes: The only really distinctive feature of this species is the curiously
prominent character of the anal lobes. In some species the anal lobes are more
or less prominent in immature individuals, this prominence disappearing in the
adult, but in P. artemisiae the prominence remains. Otherwise the species is
scarcely distinguishable from P. coleman.

Phenacoccus colemani FEhrh.
Plate 2, fig. 20.
1906. Phenacoccus colemani Ehrhorn, Can. Ent. 38 :332-3.

In life. Thinly dusted with waxy powder, without noticeable lat-
eral or caudal tassels. Oviparous, the eggs laid in a very loose, fluffy
ovisac. The species is scarcely, if at all, distinguishable in life from
Phenacoccus solani n. sp., although structurally they are sufficiently dis-
tinct.

Morphological characteristics. First three or four pairs of cerarii
with three or four conical spines, the remainder with but two, except for
the anal lobe pair, in which there may be as many as four. In all, the
spines are quite small, slender, sharply pointed and somewhat constricted
at the base. In the typical form the anal lobe cerarii have but two con-
ical spines which are accompanied by three or four very small spines.
This form, however, grades over into another in which one or two of the
small spines are equal to the larger in size, and into another form in
which the small spines are replaced by a single stiff seta. In all the
cerarii there are but few pores, and no auxiliary setae. Dorsal body setae
very few and scattered, small, more or less conical, not constricted at the
base. Triangular pores few, mingled with a few tubular ducts. Anal
lobe setae nearly twice as long as anal ring setae.

Immature female differing but little from the adult.

Type host and locality. From Rubus sp., Pescadero road, south of
Palo Alto, Santa Clara County, Calif.

Hosts and distribution. I have taken this from Rubus vitifolius,
wild strawberries, Symphoricarpos racemosus, Castilleia sp. Eriophyllum
confertiflorum and from beneath a rock, associated with ants, in the hills
near Stanford University. Mr. Swain has sent me specimens from ge-
ranium at Riverside.

58 THE CALIFORNIA SPECIES OF MEALY BUGS

Authentication. Specimens from type host and locality, agreeing in
general with the original description.

Notes: I am by no means certain that all the forms ee by me to this
species are rightly placed. The extreme forms observed are sufficiently different
to arouse doubt, yet there appears to be an almost perfect gradation between these
and I can find no real basis upon which to effect a separation.

Phenacoccus eriogoni n. sp.
Plate 2, fig. 14.

In life. Thickly covered with white wax, the lateral tassels short
and broad, the caudal tassels somewhat longer. Ovisac not observed.

Morphological characteristics. All the cerarii with more than two
cerarian spines, the anal lobe pair having as many as ten, the remainder
three to six, these spines slender, sharply pointed and slightly constricted
at the base; no auxiliary setae. Pores of the cerarii scarcely exceeding
the spines in number and not at all concentrated. The anal lobe and
penultimate cerarii show more or less of a tendency toward the develop-
ment of a chitinized area, which is, however, never well defined. Ventral
sides of the anal lobes without chitinized area. Dorsal body setae almost
lacking, the few that are present being extremely minute. Triangular
pores numerous, mingled with a few large tubular ducts with raised rim
about the mouth. Anal lobe setae equal to or a trifle shorter than the
anal ring setae.

Immature female not differing materially from the adult.

Type host and locality. From crowns and stems of Eriogonum
nudum, Stevens Creek, Santa Clara County, Calif.

Host and distribution. Known only from the above host and local-
ity. It is common enough in this vicinity.

This is quite distinct from any other species seen by me, and I can-
not connect it with any described form.

Phenacoccus kuwanae Coleman.

1903. Phenacoccus kuwanae Coleman, Jn. N. Y. Ent. Soc. 11:62, pl. 5, figs. 1-2.

In life. “Length 1.6 mm.; color yellowish-white; body sparsely
covered with a yellowish-white powder; ...’ With ovisac, “length
about 2.5 mm.; ovisac yellowish-white, short, oblong, smooth.” (Cole-
man. ) |

Type host and locality. “On a species of lichen growing on the
weeping spruce, Picea breweriana, on the east side near the summit of
the Salmon Mountains, about ten miles west of Salmon Forks, Siskiyou
County, Calif., August 3, 1901.”

Hosts and distribution. Known only as above.

SYSTEMATIC TREATMENT 59

Notes: The types of this species are in the Stanford collection, but are in
such bad condition that it is impossible to base a redescription upon them. The
original description contains nothing that will give any particular aid in recognizing
the species. The only distinctive feature that the types reveal is the presence of
many short, stout, cylindrical ducts. In this character the species is evidently very
similar to P. piceae (Loew) which has been redescribed by Marchal (7, pp. 243-5,
figs. 11-12). It is entirely possible that they are identical.

Phenacoccus stachyos Ehrhorn.
Plate 2, figs. 17, 22.

1900. Phenacoccus stachyos Ehrhorn, Can. Ent. 32:313.

1904. Phenacoccus osborni Sanders, Ohio Nat. 2 :284.

1912. (?) Heliococcus bohemicus Sulc, Cas. spoleé. Ent. 9:39, figs.
1917. Phenacoccus pettiti Hollinger, Can. Ent. 49:281, fig.

1918. Phenacoccus stachyos Ehrh.; Hollinger, Ibid. 50:23.

In life. Covered with mealy secretion from which there arise many
delicate, glassy threads of wax. Lateral and caudal tassels very short.
Oviparous, not forming a definite ovisac.

Morphological characteristics. Ocular cerarii with three or four
cerarian spines, the remainder with but two, all small and slender, those
of the anal lobe pair but little larger than most of the others. No aux-
iliary setae and practically no concentration of pores about the cerarian
spines. Spines of the anal lobe pair borne at the extreme tips of the quite
prominent anal lobes. Ventral side of the anal lobes usually with a small
chitinized area extending in from the base of the anal lobe setae. Dorsal
body setae few, extremely small. Ventral setae long and slender. Tri-
angular pores few. Tubular ducts of a very peculiar type, many of them
extraordinarily large and all with their mouths borne at the apices of
conical prominences, about the base of which there may be from one to
four small spines. The base of many of these projections is surrounded |
by a small circular, chitinized area. These ducts appear to have a rather
definite arrangement, there being from two to four on each anal lobe, two
or three near each cerarius, an indefinite number on the head and an
irregular double row along the median line of the body. It is from these
ducts that the long, glassy threads of wax arise. Anal lobe setae about
one and one-half times as long as the anal ring setae. Antennae notice-
ably long and slender.

Immature specimens differing from the adult chiefly in a reduction
of the number of ducts.

Type host and locality. From Stachys bullata, San Franciscquito
Creek, near Stanford University, California.

Hosts and distribution. 1 have this species from Stachys sp., Woodside,
San Mateo County, and Big Basin, Santa Cruz County: Monardella sp.,

60 THE CALIFORNIA SPECIES OF MEALY BUGS

Big Basin; Diplacus glutinosus, Woodside; Solanum sp., Permanente
Creek, Santa Clara County; Toxicodendron (Rhus) diversiloba, Stevens
Creek and Corte Madera Creek, Santa Clara County. In the Stanford
collection from Catalpa at Columbus, Ohio. Recorded by Hollinger (as
Phenacoccus pettiti) from numerous hosts in Missouri and by Sanders
(as Phenacoccus osborni) from sycamore at Columbus, Ohio. What is
possibly the same species has been described by Sule as Heliococcus bo-
hemicus from Robinia pseudacacia in Bohemia and Moravia.

Authentication. Specimens from the type host and locality agreeing
with the original description.

Notes: This is one of the most distinctive species of the genus. Mr. Mor-
rison has kindly furnished me with copies of the figures accompanying the original
description of Heliococcus bohemicus Sule and these leave no doubt that this
species is extremely close to P. stachyos, if it be not the same. It is worthy of
note that Robinia pseudacacia is a native of Eastern North America, where Phe-
nacoccus stachyos appears to be quite common. Mr. Morrison also informs me
that Phenacoccus osborni Sanders appears to be the same.

I do not for the present accept the genus Heliococcus. The genus can be
maintained only on the basis of the peculiar ducts, for it is in all other respects
not different from typical Phenacoccus. On this basis Pseudococcus virgatus
(which is indeed not a Pseudococcus) would be included in Heliococcus, although
it does not belong to the Phenacoccus series. Until the group is better under-
stood it may be well to maintain the present arrangement.

Phenacoccus solani new species.

; Plate 2, fig. 21.
1914. Pseudococcus solani Ckll.; Essig, Mon. Bull. State Com. Hort. 3, No.
3:122, fig. 35. (Misidentification?)

In life. Dusted with powdery secretion, with no lateral or caudal
tassels. Ovisac not known.

Morphological characteristics. Antennae 8-segmented in the adult
female. Tooth of the tarsal claws usually very small. None of the cerarii
with more than two conical spines, these, in all the cerarii, small, sharply
pointed, accompanied by no auxiliary setae and by but the slightest group-
ing of the pores, the cerarii being so obscure that in the adult it is only
with difficulty they can be traced. Anal lobe cerarii sometimes with a
single small hair, sometimes with two or three very small spines in addition
to the cerarian spines; borne upon the dorsal surface of the anal lobes.
No chitinization either dorsally or ventrally. Dorsal body setae small,
sharply pointed, of a characteristic lanceolate shape.. Triangular pores
numerous, accompanied by a very few cylindrical ducts without a raised
rim about the mouth. Anal lobe setae about one and one-half times as
long as the anal ring setae.

Immature female in general resembling the adult.

SYSTEMATIC TREATMENT 61

Type host and locality. From roots of Hemizonia rudis (tarweed)
on the campus of Stanford University.

Hosts and distribution. I have this from potato, tomato and wild
radish in the vicinity of Stanford University, potato and Malva sp. from
near San Jose and immature specimens that I believe to be this species,
from roots of pansies at Salt Lake (R. W. Doane, coll.). Specimens are
in Mr. Essig’s collection from Malva rotundifolia and Malva parviflora,
in Ventura County. Recorded by Essig from these hosts and from
potato, tomato, wild sunflower, Aster, nightshade and purslane from
southern California and from potatoes at Stockton.

Notes: In spite of the 8-segmented antennae this is certainly a Phenacoccus,
although the generic characters are indeed obscure. The determination of this as
Pseudococcus solani Ckll., which was described from New Mexico, seems to have
been made upon the basis of the fact that both have been found upon potatoes.
It is entirely possible that this determination is correct, but it is equally possible
that it is not, and I am describing the species as new, keeping the name solani in
order that no change will be necessary if this determination is confirmed.

In life this might, perhaps, be confused with Pseudococcus eriogont (Ehrh.)
which it greatly resembles both in appearance and habitat. In preparations it much
more nearly resembles Pseudococcus than Phenacoccus, but the presence of 18 pairs
of cerarii and of a denticle on the claws will place it in the latter group.

Phenacoccus simplex King.
1902. Phenacoccus simplex King, Ent. News. 13:42-3.

In life. Oval in shape, 3 mm. long, 2 broad, of a reddish-brown
color. Body thinly covered with white secretion” (King).

Morphological characteristics. The only information of any value
conveyed by the original description is as follows: “Boiled in KOH,
cleared and mounted in balsam, practically colorless except around the
area of the grouped spines, which is tinged with yellowish-brown. These
groups are variable in size and the spines are conical in shape, short, stout
and placed close together. The dorsum is quite thickly beset with short
conical spines and thin, not at all long, hairs. These not uniform in
length.”

Type host and locality. Lone Pine, California. On Atriplex.

Notes: I have not seen this species. The description conveys a faint hint
that this may be a Puto, but beyond this little can be said.

Genus PUTO Signoret

1875. Puto Signoret, Ann. Soc. Ent. Fr., (5), 5:394.
1897. Ceroputo Sulc, Sitz. k. Bohm. Ges. Wiss., no. 61.

Pseudococcine forms with a tooth on the face of the tarsal claws
and with all the cerarii (from 16 to 18 pairs) surrounded by a definite

62 THE CALIFORNIA SPECIES OF MEALY BUGS

chitinized area and including numerous conical spines. Antennae of the
adult female 8- or 9-segmented. Anal ring with 6 to 8 hairs. Male
larvae resembling those of the female in all respects up to the time of
pupation. Adult males apterous or alate, extraordinarily large, with a
wing expanse of as much as 8 mm. and a body of 5 mm. Type of the
genus Puto antennata Sign.

I consider Ceroputo to be strictly a synonym of Puto. The former
genus was separated from the latter solely on the basis of the presence of
but six hairs on the anal ring and of non-clubbed digitules. The descrip-
tion of the type species of Ceroputo is extremely minute, and the author
of the genus also redescribes Puto antennata from the type specimens.
Of the species herein dealt with, P. cupressi (Coleman), except for the
8-segmented antennae, would ordinarily be referred to Puto, while the
remainder would be referred to Ceroputo, but I cannot consider the dif-
ferences to be sufficient to justify a generic separation, especially in view
of the close resemblance in other respects.

It may be noted that Brain (2) has quite thoroughly misunderstood
this genus and has placed it among the Eriococcine forms solely because
of the 8-haired anal ring. The species which he has described as P. afri-
canus undoubtedly does not belong to this genus.

The males of this genus are very different from the males of Pseu-
dococcus and the other genera herein dealt with. Until the formation
of the cocoon they differ in no respect from the larvae of the females,
and they are also notable for their large size. In the case of P. ambigua
(Full.) they are said to be apterous, while in other species they are alate.

But three species are known from California. These may be dis-
tinguished by the following key.

1. Antennae of the adult female 8-segmented, anal ring of the
adult female with 8 setae . . . cupressi (Coleman)

Antennae of adult female 9-segmented; anal ring of all

stages with 6 setae . daa neh, eee Woah pve it fea 2
2. Adult. male winged (sha 0.68 se a pecear: (Cogs) .
Adult male apterous . . . . . . . ambigua (Full.)

Puto ambigua (Fullaway).
1910. Ceroputo ambigua Fullaway, Proc. Davenport Acad. Sci. 12:233.

This species resembles P. yuccae (Coq.) in every respect except in
the fact that the males are apterous. There is the possibility that both
apterous and alate males are produced (as is the case with Gossyparia
spuria) and should this prove to be true, the species may be dropped.

Type host and locality. From Salicornia ambigua, salt marshes at
Palo Alto, Calif.

aye

SYSTEMATIC TREATMENT 63

Hosts and distribution. Known only from the above host and lo-
cality.
Authentication. ‘Topotype specimens examined.

Puto cupressi (Coleman).

1901. Dactylopius sequoiae Coleman, Proc. Calif. Acad. Sci, (3), 2:410, fig.
(As to male.)
1908. Pseudococcus cupressi Coleman, Jn. N. Y. Ent. Soc. 16:197-8, pl. 4, figs. 1-4.

In life. Adult female reaching the length of 6 mm.; dorsally bare
of secretion except for a dusting of fine powder; with very short, stout,
lateral tassels and with a somewhat longer caudal pair. Dorsum of a
light brown color, with marginal and submedian longitudinal rows of
dark spots. Ovisac more or less pad-like. Oviparous.

Morphological characteristics. Adult female with 8-segmented an-
tennae. The fundamental arrangement of 18 pairs of cerarii (indicated
in the first stage larva) is obscured by the presence of small secondary
cerarii and occasionally by the more or less complete suppression of cer-
tain of the cerarii. Chitinous areas surrounding the cerarii for the most
part rather small, the number of cerarian spines ranging from 5-12, the
anal lobe area being much larger than any of the others and being con-
tinuous with a small chitinized area on the ventral side of the anal lobes.
Dorsum with few spines, these short and conical. Anal ring with 8 hairs
(in the adult), one pair of these being considerably smaller than the
others.

Immature female with 7-segmented antennae and with the chitinized
areas of the cerarii smaller than in the adult. The first stage larva pos-
sesses 7-segmented antennae. In all the immature stages of both male
and female I find but six anal ring setae. The male prepupa is enclosed
in a conspicuous white sac.

Adult male winged, with a wing expanse of 8 mm., of the type usual
to the genus.

Type host and locality. From Cupressus macrocarpa, Cypress Point,
Monterey County, Calif.

Hosts and distribution. From Pinus radiata, Pacific Grove; Torreya
californica, Stevens Creek, Santa Clara County (Ehrhorn, coll.) ; Se-
quoia sempervirens, vicinity Stanford University. At Pacific Grove and
Cypress Point the species becomes mature in December and January, in
the vicinity of Stanford University adult females are not found until May
and June.

Authentication. Cotype and topotype specimens examined, the for-
mer through the courtesy of Mr. Ehrhorn.

64 THE CALIFORNIA SPECIES OF MEALY BUGS

Notes: It has been proven beyond any room for doubt that the male de-
seribed by Coleman under Dactylopius sequoiae does not belong with the female
of that species. Neither is there any doubt that this male is the same as that
described by the same author under Pseudococcus cupressi. This=species is of
much interest, inasmuch as it effectually unites the two genera Puto and Ceroputo.

Puto yuccae (Coq.).

1889. Dactylopius crawii Coquillet, West Amer. Sci. 6:123. (As to male.)

1890. Dactylopius yuccae Coquillet, West Amer. Sci. 7:44.

1896. Phenacoccus yuccae (Coq.) ; Cockerell, Check List of Coccidae.

1900. Phenacoccus bahiae Ehrhorn, Can. Ent. 32:315.

1901. Ceroputo yuccae (Coq.) ; Cockerell, Proc. Biol. Soc. Wash. 14:166.

1901. Ceroputo bahiae (Ehrh.); Cockerell, Can. Ent. 33 :166.

1903. Ceroputo yuccae var. ceanothi Ckll., Ann. Mag. Nat. Hist., (7), 11:163.

1912. Phenacoccus ramonae Essig, Pomona Col. Jn. Ent. 1:26, fig.

1912. Ceroputo yuccae (Coq.); Essig, Pomona Col. Jn. Ent. 1:94, fig. 43.

1913. Ceroputo yuccae (Coq.); Essig, Mon. Bull. Calif. State Com. Hort. 2:95,
fig. 74.

In life. Thickly covered with white wax; with short, stout lateral
tassels, the caudal tassels being somewhat longer. Length 4-6 mm. Male
pupa contained in a whité cocoon of 3-4 mm. length. Adult male winged,
with a wing expanse of 7-8 mm. and with a pair of caudal wax tassels
as much as 6 mm. long.

Morphological characteristics. Antennae of the adult female 9-seg-
mented. Antennae and legs strongly chitinous. Cerarii varying in num-
ber from 15 to 18, each surrounded by a chitinous area, each containing
a dozen or more short, conical spines and a somewhat smaller number of
pores. Anal lobe area largest. No chitinized area on the ventral side
of the anal lobes. Dorsum with small, scattered, conical spines. Anal
ring with six setae, very rarely with eight.

Immature female resembling the adult except for the smaller size of
the chitinized areas. First stage larva with 7-segmented antennae.

Type host and locality. From Yucca whipplei, Los Angeles County,
Calif. He

Hosts and distribution. Recorded from several species of yucca,
from Diplacus glutinosus, Ramona polystachya, citrus and bananas in the
southern part of the state. In the vicinity of Stanford University I have
taken the species from Eriophyllum confertiflorum, Diplacus glutinosus,
Garrya elliptica and Stachys sp.

Authentication. Not authenticated. Spécimens from the type region
have been examined which agree with the original description and which
belong to a species that has generally been accepted as P. yuccae (Coq.).

Notes: I have examined topotypic specimens of Ceroputo bahiae (Ehrh.)
and can find no basis upon which to separate this from Puto yuccae. The variety

J

SYSTEMATIC TREATMENT 65

ceanothi, established by Cockerell because of the “considerably longer second and
fourth joints of the antennae,” is hardly to be taken seriously. Among the ma-
terial of Pseudococcus agrifoliae received from Mr. Essig is a slide labeled “co-
type” which contains specimens of larvae of a Puto taken from Quercus agrifolia
near Claremont. It is impossible to identify these specimens with certainty, but
it is possible that these are P. yuccae.

Genus HETEROCOCCUS new genus

Pseudococcine forms with quinquelocular dorsal pores instead of the
usual triangular pores; with cerarii on the posterior portion of the ab-
domen only; with a more or less distinct denticle on the tarsal claws;
with 9-segmented antennae in the adult female.

Type of the genus Heterococcus arenae n. sp.

Notes: Morphologically the species upon which this genus is based seems
to belong to the Phenacoccus series, the 9-segmented antennae and the presence
of a denticle on the claw being indicative of this. However, the presence of circular
dorsal pores is quite distinctive. It is possible that Pseudococcus poae (Maskell)
is congeneric with this, for it is said in the original description of that species that

“All over the body are small circular spinnerets.” Otherwise I know of no similar
species.

Heterococcus arenae n. sp.
Text figure 16C; plate 3, fig. 32.

In life. Found beneath the leaf sheaths of its host, where it is sur-
rounded by a small amount of secretion.

Morphological characteristics. Cerarii present on the last two ab-
dominal segments only; each with two slender and rather long spines
which are distinguishable from the body spines only by their position and
paired character ; without grouped pores. Anal lobe cerarii accompanied
by a few spines which are as large as the cerarian spines. Dorsal body
setae slender, scattered, ventral body setae few, slender. Dorsum with
numerous pores, all of which are circular and quite large. Tubular ducts
few and very small, all without a raised rim about the mouth. Tarsal
claws typically with a denticle, although in some specimens this appears
to be lacking. Anal ring large. Anal lobe setae slightly longer than the
anal ring setae. Antennae of the adult female 9-segmented.

Immature female differing from adult only in reduction of number
of antennal segments.

Type host and locality. From Poa douglasii, on the sand dunes near
Asilomar at Pacific Grove, Monterey County, Calif., December 1, 1917.

Hosts and distribution. Known only from above host and locality.

66 THE CALIFORNIA SPECIES OF MEALY BUGS

Genus TRIONYMUS Berg

Pseudococcine forms with normally not more than three or four
pairs of cerarii, these on the posterior portion of the abdomen. Derm
never pigmented. Dorsal body setae never conical or “spearhead-shaped.”
Dermal pores of the triangular type present. Dorsum frequently with
many multilocular pores. Claw without a denticle. Antennae of the
adult female 7- or 8-segmented. |

Type of the genus, Trionymus perrisi (Sign.).

Notes: The type species of this genus has been well described by Marchal
(7) and its nature is sufficiently clear. It is evidently very similar to T. califor-
nicus Ehrh. The genus was originally established chiefly upon the elongated form
of the female, but this can hardly be considered sufficient. If the genus be main-
tained at all it must be on the basis of the reduced number of pairs of cerarii, and
such an interpretation will necessitate its extension to include several species now
referred to the genus Ripersia. I cannot believe that such species as Trionymus
bromi n. sp. and Ripersia festucae Kuwana should be separated generically simply
because of the difference of a single segment in the antennae, nor, if this interpre-
tation be accepted, can I at present see any way by which such species as Ripersia
villosa Ehrh. or Pseudococcus trifolii (Forbes) can be kept out of this genus.

As a matter of fact such a step, even though it result in creating a somewhat
unnatural group, can produce no worse confusion than that at present displayed in
the genus Ripersia. The real nature of this genus cannot be determined, for it
was originally based simply upon the 6-segmented antennae of the adult female of
the type species and this is practically the only significant fact that we possess in
regard to this species. As at present understood the genus is made to include
almost any species with 7-segmented antennae occurring on grass, roots or with
ants. Even though the descriptions of practically every one of the North Ameri-
can species are hopelessly inadequate, it is obvious that the group thus obtained is
a most heterogeneous assemblage and is for all practical purposes meaningless.

KEY TO CALIFORNIA SPECIES.

1. With but one pair of cerarii, these on anal lobes . . . . .. 4
With at least two pairs of, cerarii . . . 6 1 1 se pee 8
2. Anal lobe cerarii surrounded by a chitinized area bearing
numerous pores and setae . . . californicus Ehrh.
Anal lobe cerarii.not so... .- : ; viene

3. Antennae of adult female 8-segmented calceolariae ( Mask.)
Antennae of adult female 7-segmented festucae (Kuwana)

4. Cerarian spines setiform . . . . . . distichin n. sp.
Cerarian spines ‘conical -),0)./).000) «pee 2 5
5. Cerarii without grouped pores . .. . : ‘ (oi

Cerarii with numerous pores . . . . grimdeliae n. sp.

SYSTEMATIC TREATMENT 67

6. Dorsum with short, broad, tubular ducts . villosa (Ehrh.)
Dorsum without such ducts

7. Anal ring setae scarcely more than one-half the length of
the anal lobe setae; antennae of adult female 7-seg-

amentem i ea Ok. smiths (Essig)
Anal ring setae equaling anal lobe setae; antennae of adult
female 8-segmented . .... . . bromin. sp.

Trionymus bromi new species.

In life. Either in the axils or on the upper surface of the leaves.
Surrounded by a fluffy mass of cotton.

Morphological characteristics. Antennae of the adult female 8-seg-
mented. But one pair of cerarii present, these on the anal lobes and con-
taining two short, sharp spines, two or three small setae and a very few
pores. Body setae few, inconspicuous, short, slender, curved. Body
with many triangular pores and small tubular ducts, and with a few mul-
tilocular pores on the dorsum of the abdomen. Anal lobe setae but little
if any longer than the anal ring setae. Anal ring rather small and heavily
chitinized.

Immature female not observed.

Type host and locality. From Bromus sp. Corte Madera Creek near
Stanford University, Calif.

Hosts and distribution. From the same host on the road above Con-
gress Springs, Santa Clara County; from Ammophila arenaria on the
sand dunes near Golden Gate Park, San Francisco; from the bark of
Ericameria fasciculata (a composite) on the sand dunes near Pacific
Grove, Monterey County.

Notes: This is quite abundant locally. I have but little doubt that it is a
described species, yet I can connect it with none and must give it a name. The
specimens taken from Ericameria had in all probability merely entered the crevices

of the loose bark in order to form their ovisacs and it is improbable that the
species feeds on this host.

Trionymus calceolariae (Maskell).
1878. Dactylopius calceolariae Maskell, Trans. New Zealand Inst. 11:218.

In life. Concealed beneath the sheathing bases of the leaves of its
host, surrounded by masses of wax.

Morphological characteristics. Antennae of the adult female 8-
segmented. Two pairs of cerarii present, the cerarian spines short, stout,
conical, not accompanied by grouped pores or auxiliary setae. Entire
body beset with many short hairs, of about equal length on dorsal and
ventral sides. Derm with many triangular pores and small tubular ducts

68 THE CALIFORNIA SPECIES OF MEALY BUGS

without a raised rim about the mouth. Large numbers of multilocular
pores present on the dorsum of the abdomen as well as on the venter.
Anal ring hairs slightly shorter and more slender than the anal lobe hairs.
Anal ring large and heavily chitinized.

Immature female much resembling the adult.

Type host and locality. From Calceolaria in Australia.

Hosts and distribution. As far as I am aware this species is known
only from “New Zealand flax” on the campus at Berkeley.

Authentication. Not authenticated. Is probably not the Dactylopius
calceolariae of Maskell.

Notes: Specimens from Berkeley have been compared by Mr. Morrison
with specimens in the Maskell collection at Washington and he has furnished the
following information: “We do not appear to have the type slides of calceolariae,
as that was described in 1878, and the first slide in the Maskell collection is dated
1880. However, your specimen appears to agree exactly with the one in the Mas-
kell collection. There are also two other specimens, about 1887, in the Maskell
collection which have three cerari on each side at the apex of the abdomen, in-
stead of two as have your specimens and the first one in the Maskell collection.
In all other respects they seem to be the same as the earlier specimens. I find
that the same situation exists in the Department collection as well, there being
two groups of specimens, one with three pairs of cerari and one with two.”

It is possible that these are the same species, for a similar variation occurs
in T. californicus Ehrh., but it is equally possible that two species are included.
In either case I can scarcely believe that the species at hand is the real Dactylopius
calceolariae of Maskell. In the original description it is stated that “the appen-
dages are short except at the posterior end, where there are two very long, and,
on each side of them, another somewhat shorter,” and the species is figured with
caudal tassels of nearly half the length of the body. In the specimens before me
there cannot possibly be any lateral tassels because of the entire absence of cerarii
and the weak development of the anal and preanal cerarii precludes the develop-
ment of any but the shortest tassels in this region. There is therefore every pos-
sibility that Maskell himself has confused two or more species under this name,
but toward the solution of this problem I can of course give no aid. For the pres- _
ent I am retaining the name for the specimens at hand rather than give a new
name for a species which in all probability already is supplied with one.

Trionymus californicus Ehrhorn.
See plate 2, fig. 19.

1910. Ripersia smithii Essig, Pomona Col. Jn. Ent. 2:219-221, fig. (In part.)
1911. Trionymus californicus Ehrhorn, Can. Ent. 43:279, figs. 17, 1-lb.

In life. Found beneath the sheathing bases of the leaves, enclosed in
more or less waxy secretion, or at times free upon the leaves. When
found free upon the host there are two or three pairs of caudal tassels
of which the last are quite stout and are nearly one- bri the length of the
body, the others minnie

SYSTEMATIC TREATMENT 69

Morphological characteristics. Antennae 8-segmented. Two to four
pairs of cerarii present at the posterior end of the abdomen, the cerarian
spines of the anal lobe pair large and stout, of the penultimate pair some-
what smaller, of the third and fourth pairs more or less hairlike, usually
indistinguishable from the body setae. Spines of the anal cerarii sur-
rounded by a rather large chitinized area, containing numerous pores and
bearing from 8-10 rather long hairs. Remaining cerarii with few pores
and no hairs. Dorsum with numerous slender setae which are nearly or
quite as long as those on the ventral side. Many triangular pores and
many tubular ducts without a raised rim about the mouth. Dorsum with
very few multilocular pores. Anal ring large and rather weakly chit-
inized, the anal ring hairs slightly longer than the anal lobe hairs.

Immature female differing from the adult chiefly in the smaller num-
ber of hairs and ‘pores and the weaker development of the chitinized area
about the anal cerarii.

Type host and locality. From grass at Lathrop, San Joaquin County,
Calif.

Hosts and distribution. In Essig’s collection from Elymus conden-
satus, Ventura County; in the Stanford collection from Elymus sp., We-
natchee, Wash. ; Elymus sp., edge of the salt marshes at Palo Alto, Calif. ;
Elymus sp., at Pacific Grove, Monterey County, Calif.

Authentication. Cotype slide examined.

Notes: I am indebted to Mr. Ehrhorn for the loan of a cotype slide of T.
californicus and to Mr. Essig for cotypes of Ripersia smithti. The slides labeled
as R. smithii contain two species, of which one will stand as Trionymus smith
(Essig). The other is identical with T. californicus, which is very near to, if not
identical with, specimens from Missouri that I have seen identified as T. ameri-
canus Ckll. T. californicus may possibly be a synonym of the latter.

The cotype of T. californicus examined by me has two pairs of cerarii, but
the specimens from Elymus at Palo Alto show a tendency to the development of
more. In several cases they show three pairs, in some four, and in one there
are several cerarii on one side of the body. Otherwise these specimens are typical
californicus, but further material may necessitate a separation. It should be noted
that except for the reduction in the number of cerarii this species is very similar
to Pseudococcus timberlakei Ckll., so much so in fact as to suggest a possibility
that they should not be generically separated.

Trionymus distichlii n. sp.
Plate 3, fig. 28.

In life. Occurring either upon the upper side of the leaves or at their
axils. Entirely covered by fluffy mass of wax. Insect of a purplish color
when denuded of secretion.

Morphological characteristics. Antennae of the adult female 7-seg-

70 THE CALIFORNIA SPECIES OF MEALY BUGS

mented. But one pair of cerarii present, these on the anal lobes, the
cerarian spines elongated and differing only in size from the body setae,
accompanied by two or three slender hairs and no grouped pores. Body
with relatively few pores and ducts. Dorsum of the abdomen with num-
bers of multilocular pores. Very few setae except on the last three or
four segments of the abdomen, where there are many slender setae of
about equal length on the dorsum and venter. Anal lobe setae slightly
shorter than the anal ring setae. Anal ring rather weakly chitinized.

Type host and locality. Quite abundant at times on the salt marsh
grass, Distichlis spicata, at Palo Alto, Calif.

Hosts and distribution. I have specimens from the type host at Ano
Nuevo Point, Santa Cruz County and from La Jolla. In the vicinity of
Palo Alto the species becomes mature in August and September.

Notes: This is distinguished chiefly by the elongated cerarian spines. I
have specimens of this from La Jolla collected by Cockerell and determined by
him as his Pseudococcus salinus, which they cannot possibly be, according to his
original description of that species.

Trionymus festucae (Kuwana).

1902. Ripersia festucae Kuwana, Proc. Cal. Acad. Sci., (3), 3:401, pl. 25, figs. 11-16.

In life. Concealed beneath the sheathing bases of the leaves, enclosed
in more or less mealy secretion.

Morphological characteristics. Antennae 7-segmented. Two pairs
of cerarii present, these on the last two segments of the abdomen, each
containing two small, sharp, conical spines, not accompanied by grouped
pores. Scattered about the anal cerarii are a few slender setae which do
not appear to be especially connected with them. Dorsum of the abdomen
with numerous multilocular pores. Body with short hairs dorsally and
ventrally, these somewhat more numerous toward the posterior end, but
not so strikingly so as in T. smithu. Setae of the anal ring quite long,
slightly longer than those of the anal lobes. Anal ring rather small and
heavily chitinized.

Immature female. The larva has 6-segmented antennae and but one
pair of cerarii.

Type host and locality. From Festuca sp., on Black Mountain near
Stanford University.

Hosts and distribution. I have taken this from Festuca sp., near
Woodside, San Mateo County, Calif., and have also found a species on
bamboo on the Stanford campus that I am unable to separate from it. —

Authentication. Types examined.

Notes: I have seen specimens from Missouri that I believe to be identical
with this species.

on a i ee

SYSTEMATIC TREATMENT 71

Trionymus grindeliae n. sp.
Plate 3, fig. 26.

In life. With a single pair of slender caudal tassels which attain
perhaps the length of the body. A root-infesting form.

Morphological characteristics. Antennae 7-segmented in the adult
female. With but one pair of cerarii, these on the anal lobes and con-
taining two or three slender, conical spines, several smaller spines, or
spines and slender setae, and a cluster of numerous and rather scattered
pores. Dorsum with no multilocular pores. Dorsal body setae few, vary-
ing in size and shape from small and slender to quite large and stout,
those near the margin of the next to the last abdominal segment being as
large as the cerarian spines. Tubular ducts all very small, almost lacking
on the dorsum, but abundant on the venter. Anal ring setae somewhat
stouter and larger than the anal lobe setae.

Immature specimens not observed.

Type host and locality. From roots of Grindelia robusta on the cliff
at Pacific Grove, Monterey County, Calif., Dec. 1, 1917.

Hosts and distribution. Known only as above.

Notes: This species is extremely close to Trinoymus trifolii (Forbes), of
which I have had the privilege of examining a cotype specimen, received through
the kindness of Mr. Forbes, and other specimens from Missouri, received from
Mr. Hollinger. In fact these two are so very similar that it is only with much
hesitation that I conclude to regard my species as distinct. The only difference
of any importance lies in the fact that the dorsal body setae of T. trifolti are very
small and slender, while in T. grindeliae they tend to be much larger and stouter.
Further collecting may show the two to intergrade.

This is not the place to discuss at length a species not coming definitely within
the limits prescribed for this paper, but I may note that the species Trionymus
trifolti (Forbes) has been consistently misunderstood. Another species, in all
probability Pseudococcus maritimus (Ehrh.), has been described as the “summer
form” of trifolii, and, so far as I can determine, all the published records since the
original description refer, at least in part, to this.

Trionymus smithii (Essig).
Plate 3, fig. 27.

1910. Ripersia smith Essig, Pomona Col. Jn. Ent. 2:218-220. (In part.)
1916. Pseudococcus smithii (Essig) ; Cockerell, Jn. Econ. Ent. 9:312. (In part.)

In life. Concealed beneath the sheathing bases of the leaves of the
host. Slightly covered with secretion, without lateral or caudal tassels.
Morphological characteristics. Antennae 7-segmented in adult fe-
male. But one pair of cerarii present, these on the anal lobes and con-
sisting of but two small conical spines not accompanied by grouped pores

yp THE CALIFORNIA SPECIES OF MEALY BUGS

or slender setae. Dorsum with relatively few triangular pores and cy-
lindrical ducts and very few multilocular pores. Last two segments of
the abdomen with many short, slender, curved hairs both dorsally and
ventrally, the remainder of the body with few. Anal ring rather small,
the anal ring hairs quite short, the anal lobe hairs somewhat longer.

Immature female differing from the adult in having the antennae
6-segmented and in having the cerarian spines more or less setiform.

Type host and locality. From Elymus condensatus, Santa Paula,
Ventura County, Calif.

Hosts and distribution. I have specimens of this from an unknown
grass, Santa Clara Valley, Ventura County; from Elymus sp. at Alum
Rock Park, San Jose; from undetermined grass growing in the chap-
arral on Jasper Ridge near Stanford University.

Authentication. Cotypes examined.

Notes: This is one of two species included by Essig under his Ripersia

smithii. The other having been later described as Trionymus californicus Ehrh.,
the name smithi must be retained for the specimens herein described.

Trionymus villosa (Ehrh.).
Plate 3, fig. 31.
1899. Riprsia villosa Ehrhorn, Can. Ent. 31:6.

In life. “Female in clusters and single in the crotches of twigs of
oak. Sac loosely woven of long white wool, oval, about 2 mm. long and
1 mm. broad. Female when removed from the sac, bright crimson. . . .”
(Ehrhorn). The specimens here described were in all probability not
fully mature as the species reaches a much larger size.

Morphological characteristics. Antennae of adult female 7-seg-
mented. Cerarii present only on the anal lobes, the cerarian spines small
and slender but conical, accompanied by four or five small setae and with
no grouped pores. On the two or three segments preceding the last three
there is, at each margin, a single spine somewhat larger and stouter than
those of the dorsum, these perhaps representing the vestiges of the cerarii
of these segments. In the immature females these spines are quite prom-
inent, but they are somewhat difficult to distinguish in the adult. Dorsal
body setae few, scattered, quite small and slender. Derm with many tri-
angular pores and also with numerous short, stout cylindrical ducts with
a narrow raised rim about the mouth. Dorsum without multilocular
pores. Anal ring hairs quite small, scarcely exceeding one-half the length
of the anal lobe hairs.

Immature female not seen.

Type host and locality. From Quercus agrifolia at Berkeley, Calif.

Ee SyEEE———EE———=

a

SYSTEMATIC TREATMENT 73

Hosts and distribution. Known only from the above host. In the
Essig collection from Claremont and Berkeley; in the Stanford collection
from Mountain View, Santa Clara County, (Ehrhorn coll. and det.).

Authentication. Topotype and metatype specimens examined.

Notes: This is the species first described by Essig as Pseudococcus agrifoliae.
For a discussion of the nomenclatorial problem involved see the notes under
Pseudococcus quercicolus.

Genus RIPERSIELLA Tinsley.

Pseudococcine forms with the antennae set very close together at the
front of the head, 5- or 6-segmented.

Type of the genus Ripersia rumicis Maskell. |

Of this genus but one species is found in California. The material
of this species examined by me is not of such a character as to permit
any further explanation of the genus.

Ripersiella kelloggi Ehrh. & CkIl.
Plate 3, fig, 29.
1901. Ripersiella kelloggi Ehrh. & Ckll., Proc. Biol. Soc. Wash. 14:166.

In life. “Adult female generally attached to the roots of grass, pro-
ducing a small quantity of white, cottony secretion which generally en-
cases the body. Color creamy white, about 1144 mm. long and 1 mm.
wide, oval, sometimes pyriform, shiny.” (Ehrhorn.)

Morphological characteristics. Antennae very small and short, set
little more than the width of the basal segment apart, 5-segmented, the
fifth segment nearly as long as the other four together. Legs very small
and stout. Anal ring small, with short hairs.

Type host and locality. From roots of “bunch grass,” Stevens
Creek, Santa Clara County, Calif.

Hosts and distribution. Known only as above.

Authentication. Cotype slide examined.

Notes: I have not been able to find this species and know it only from a co-
type slide sent me by Mr. Ehrhorn. Unfortunately a close study of the species is
not possible without more material and I can say nothing in regard to the finer
details of its morphology. Whether or not it is really congeneric with the type
of this genus and with the other species included with it remains to be demon-
strated.

Genus CRYPTORIPERSIA CKkII.

1899. Ripersia (Cryptoripersia) Cockerell, Can. Ent. 31:5.
1899. Cryptoripersia Ckll., Ibid. 31 :278.

In defining Cryptoripersia as a subgenus of Ripersia Cockerell states

74 THE CALIFORNIA SPECIES OF MEALY BUGS

that it “will be distinguished by the possession of a complete subpyri-
form brittle sac.” This is a bit optimistic, but the description of the
genotype, C. arizonensis (Ehrh.) indicates that this species is very simi-
lar to the species later described by Ehrhorn as Eriococcus salinus. It
must be admitted that this is only a surmise, but the latter species is cer-
tainly not an Eriococcus and does seem to resemble C. arizonensis.
There being no other group into which it will fit, I regard it for the pres-
ent as a Cryptoripersia and base the following characterization of the
genus upon it.

Pseudococcine forms with but the anal pair of cerarii present, the
cerarian spines in these being long and slender, their homology with the
usual short conical spines being indicated only by their position and
paired character. Anal opening on the dorsum at some distance from
the posterior margin of the body. Antennae of the adult female and of
first stage larva, 6-segmented.

Cryptoripersia salinus (Ehrh.).

Plate 3, fig. 30.
1911. Eriococcus salinus Ehrhorn, Can. Ent. 43:276.

In life. Occurring on the roots of the host. The adult female en-
tirely enclosed in a white, pyriform sac as much as 5 mm. in length.
Oviparous.

Morphological characteristics. Adult female as much as 5 mm. long
on the slide, pyriform or ovoid when flattened out, the anterior end point-
ed. Antennae 6-segmented, quite small. Anal cerarii only present, each
with two slender hairs, no pores or auxiliary setae. Dorsal body setae
few, appearing only on the head and posterior end of the abdomen, very
small. Tubular ducts all very small, few in number. Dorsum of the ab-
domen without multilocular pores. Anal ring quite small, heavily chit-
inized, the anal ring setae slightly shorter than the anal lobe setae. In all
my specimens the body flattens out on the slide in such a manner that the
anal ring appears to lie on the dorsum at some distance from the pos-
terior end of the body.

Immature female not seen. First stage larvae with 6-segmented
antennae.

Type host and locality. From roots of “bunch grass,’’ Alameda
shore, Alameda, Calif.

Hosts and distribution. I have taken this species from the roots of
a grass (Stipa sp. or Poa sp.) growing in bunches on the dry hillside in
Stevens Creek Canyon, Santa Clara County, and from what is possibly the

SYSTEMATIC TREATMENT 75

same grass growing in the chaparral on Jasper Ridge near Stanford Uni-
versity.

Authentication. Cotype examined.

Notes: This species might very readily be confused with an apparently un-
described species (belonging to an unnamed genus) which I have taken from the
roots of grass at Pacific Grove. The unnamed species, however, does not belong

to this group and can readily be distinguished by the absence of triangular pores,
the non-Pseudococcine ducts and the absence of dorsal ostioles.

Genus ERIUM Maskell.

Pseudococcine forms with cerarii on the anal lobes only, without mar-
ginal paired spines of any character. Dorsum beset with stout, spear-
head-shaped spines, especially on the abdomen, without multilocular
pores. Antennae of the adult female 7-segmented. Legs short and stout.
Body more or less globular. Derm of the adult female blue-green. Adult
female partially or entirely enclosed in a felted sac.

Type of the genus, Erium globosum (Maskell).

Notes: I have before me specimens of the type species from Australia, de-
termined by Froggat, and this is certainly congeneric with our E. lichtensioides.
In fact the two species are so much alike it is even difficult to separate them spe-
cifically. The genus has usually been based upon the fact that the adult female
is entirely enclosed in a waxy sac, but this character is misleading. One species,
Pseudococcus eriogoni (Ehrh.), which has been referred to Erium on this basis,
certainly has nothing to do with it and possibly some of the others have also been
wrongly placed. On the other hand, some species now referred to Pseudococcus
appear to belong to Erium, among these being P. ledi Ckll. and P. hymenocleae
(Ckll.). It is possible that Pseudococcus ephedrae (Coq.) likewise belongs here.

Erium lichtensioides (CkIl.).
Plate 3, fig. 25.

1891. Dactylopius lichtensioides Cockerell, Science Gossip, n. d., 3:199.

1901. Eriococcus artemisiae Kuwana, Proc. Calif. Acad. Sci., (3), 3:399, figs.

1902. Erium lichtensioides (Ckll.) ; Cockerell, Ann. Mag. Nat. Hist., (7), 10:466.

1906. Eriococcus artemisiae var. catalinae Ehrhorn, Can. Ent. 38 :332.

1910. Pseudococcus artemisiae Essig, Pomona Col. Jn. Ent. 1:42.

1914. Pseudococcus hymenocleae (Ckll.); Essig, Mon. Bull. Calif. State Com.
Hort. 3:118. (Misidentification ?)

In life. In the earlier stages more or less covered with white secre-
tion, later becoming entirely enclosed in a tough felted white sac which
frequently is open along the dorsum. Insect black.

Morphological characteristics. Antennae 7-segmented, quite short.
Derm of a blue-green color before staining. Cerarii present only on the
anal lobes, containing as many as 10 “spear-head-shaped” spines, a few

76 THE CALIFORNIA SPECIES OF MEALY BUGS

pores and no auxiliary setae. Area about the cerarii more or less defi-
nitely chitinized. Dorsum of the abdomen and the posterior portion of
the thorax with numerous spines of the same type as those in the cerarii.
Toward the head these give way to short, slender setae. Derm, espe-
cially on the abdomen, with innumerable tubular ducts without a raised
rim about their mouths. Legs quite short and stout, the posterior coxae
‘with many ventral pores. Body form more or less globular.

Immature specimens resembling the adult except for a reduction in
the number of spines in the cerarii. Male larva of the same type as in
Pseudococcus.

Type host and locality. From Artemisia frigida, Fort Collins, Col-
orado.

Hosts and distribution. In California this species has been recorded
from the vicinity of Stanford University and from the southern part of
the state, the only recorded host being Artemisia californica.

Authentication. JI have examined the types of this in the National
Collection at Washington.

Notes: I am entirely in accord with Sasscer’s conclusion that Eriococcus
artemisiae Kuwana is identical with Erium lichtensioides. The types of the former
species are in the Stanford collection. I have not seen the types of Eriococcus
artemisiae var. catalinae Ehrhorn, but I can see no grounds for recognizing this
variety. The types of Pseudococcus artemisiae Essig are unmistakably E. lichten-
sioides, but the figure accompanying the original description appears to be of an-
other species occurring on Artemisia. I have this species, but do not possess
enough material to justify naming it. The published record of the occurrence of
Pseudococcus hymenocleae on Artemisia in California is probably based on a mis-
identification, as this species very much resembles E. lichtensioides.

Genus ANTONINA Signoret.

Pseudococcine forms without legs in the adult female; with the
antennae in adult female reduced to mere stubs; without recognizable
cerarii; with the posterior end of the body more or less invaginated, this
invagination forming a tube at the inner end of which is the anal ring.
Spiracles very large and conspicuous. Circular, multilocular pores pres-
ent on the dorsum.

Type of the genus Antonina purpurea Sign.

Notes: Although the species of this genus depart widely in form and ap-
pearance from the general type of the group, there is, I believe, no doubt that its
affinities are with the Pseudococcine forms. The cerarii are not recognizable, but
the tubular ducts are of the same type as in Pseudococcus and there are two pairs
of dorsal ostioles, although it is sometimes difficult to distinguish the anterior pair.

But one species of this genus has been recorded from California, although
there are in all probability other species on our native grasses.

SYSTEMATIC TREATMENT 77

Antonina crawii Ckll.
Plate 3, figs. 23, 24.

1900. Antonia crawit Cockerell, Psyche 9:70.
1902. Antonina crawit Ckll.; Kuwana, Proc. Calif. Acad. Sci. (3), 3:57, fig.
1915. Antonina crawii Ckll.; Essig, Inj. & Benef. Ins. Calif., 2d ed., 132-3.

In life. “The full grown female scales are covered with a thick,
compact white cottony coat which makes them very conspicuous. The
body proper varies from 1/8 inch to nearly 1/4 inch long, is broadly
oval or rounded and deep purplish-red in color. The females collect in
colonies, forming large cottony masses in the leaf axils of the canes.”
(Essig.) From the posterior end of the body there usually arises a con-
spicuous white, waxen thread.

Morphological characteristics. General form elongate, tapering
somewhat posteriorly. The adult female is apodous and the antennae
are reduced to mere conical stubs with three segments. Last two seg-
ments of the abdomen quite heavily chitinized and bearing numerous
short, spike-like spines, the remaining segments with just a few very
small spines at the margins. Entire body with numerous small cylin-
drical ducts. A longitudinal area of circular pores at the lateral margins.

Immature stages. I have seen only the first stage larva. In this
the antennae are of the usual type, 6-segmented. At the lateral margin
of each of the last three abdominal segments is a group of two or three
stout spines which may be the homologues of the cerarii. The anal ring
is very heavily chitinized and bears six short, stout, spine-like setae.

Hosts and distribution. An introduced species found on bamboo, in
various parts of California. I have examined species from Ventura re-
ceived from E. O. Essig. |

Notes: It is by no means certain that this is really A. crawii Ckll. The
California specimens agree with certain specimens taken at Yokohama, Japan, by
Kuwana and identified by him as this species. They do not agree with other speci-
mens from Tokyo, Japan, likewise identified as this species by Kuwana. In the
latter lot the antennae are 2-segmented and the spines on the posterior end of the
abdomen are much longer and more slender.

This differs from A. indica Green, of which I have specimens from Bermuda
grass in Hawaii (det. Ehrhorn), in the much more elongate form and the much

shorter and stouter abdominal spines. It appears to differ in this latter character
from A. nortoni Parrott & Ckll. and A. gramnis Parrott.

7%

REFERENCES CITED

Berlese, A.
1893. Le Cocciniglie Italiene viventi sugli agrumi. Revista Patol-
ogia Vegetale, 2:70-187, pl. 3-5.
Brain, C. K. |
1915. The Coccidae of South Africa. Trans. Royal Soc. South
Africa, 5, pt. 2.
Dietz, H. F., and Morrison, H.

1916. The Coccidae or Scale Insects of Indiana. Rept. State
Ent. Indiana, 8 :207-212.

Ferris, G. F.

1917. Methods for the study of mealy bugs. Jn. Ec. Ent., 10:
322-324.

Green, E. E.
1914. On the preparation of Coccidae for microscopical examina-
tion. Annals Applied Biology, 1 :98-106.
Hollinger, A. H.
1917. Taxonomic value of antennal segments of certain Coccidae.
Annals Ent. Soc. Am., 10:264-271, pls. XITX-XXII.
Marchal, P.
1908. Notes sur les Cochenilles de l’Europe et du Nord de
VAfrique. Annales Soc. Ent. France, 77 :223-309.
Maskell, W. M.
1894. Remarks on certain genera of Coccidae. The Entomologist,
27 :44-6.
Smith, P. E.

a. 1911. Specific characters used in the genus Pseudococcus.
Annals Ent. Soc. Am., 4:309-327.

b. 1913. A study of some specific characters of the genus Pseudo-
coccus. Pomona Col. Jn. Ent. & Zoology, 5:71-84.

PLATES
WITH EXPLANATIONS

77

10.

Li,

PLATE I.

Pseudococcus crawii (Coq.); penultimate and anal lobe cerarii.

Pseudococcus comstocki (Kuw.); penultimate and anal lobe cerarii (left) and ventral side
of anal lobe (right).

Pseudococcus sequoiae (Coleman); penultimate and anal lobe cerarii.

Pseudococcus longispinus (Targ.); penultimate and anal lobe cerarii (left) and ventral side
of anal lobe (right).

Psenudococcus salinus (Ckll.); posterior end of abdomen. Penultimate and anal lobe cerarii
(left) and ventral side with half of anal ring (right).

Pseudococcus citrophilus Claussen; penultimate and anal lobe cerarii (left) and ventral side
of anal lobe (right).

Pseudococcus citri (Risso); penultimate and anal lobe cerarii (left) and ventral side of
anal lobe (right). Except for the shape of the spines this figure will apply for P.
kraunhiae (Kuwana).

Pseudococcus kraunhiae (Kuwana); cerarian spine (above) and dorsal body seta (below).
Pseudococcus citri (Risso); cerarian spine (above) and dorsal body seta (below).

Pseudococcus longisetosus n, sp.; penultimate and anal lobe cerarii (left) and ventral side
of anal lobe (right).

Pseudotoccus cupressicolus n, sp.; penultimate and anal lobe cerarii. Except for the very
slightest details this figure will apply equally well for Pseudococcus ryani (Coq.).

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Piugocdsnae ibd (Maskell); penultinjate. aad Avia’ “lobe cerarii (left) and ventral side of
anal lobe (right). pire he es

Pseudococcus irises. (Ehrh.); Heniitimate and anal lobe cerarii (left) and ventral side
of anal lobe (right).

Phenacoccus eriogoni n. sp.; penultimate and anal lobe cerarii.

Phenacoccus artemisice Ehrh.; penultimate and anal lobe cerarii (left) and ventral side of
anal lobe (right).

Phenacoccus artemisiae Ehrh.; cephalic margin of the bad of immature specimen.

Phenacoccus stachyos Ehrh.; penultimate and anal lobe cerarii (left) and ventral side. of
anal lobe (right).

Pseudococcus quercicolus SP. penultimate and anal lobe cerarii.

Pseudococcus timberlakei Ckll.; penultimate and anal lobe cerarii (left) and ventral side of
anal lobe (right). Except for the ventral side of the anal lobe this figure will apply
equally well for Trionymus californicus Ehrh.

Phenacoccus colemani Ehrh.; a, penultimate and anal lobe cerarii (left) and ventral side of
anal lobe (right) of the typical form; b, anal lobe cerarius of an extreme form.

Phenacoccus solani n. sp.; penultimate and anal lobe cerarii; a, disproportionately enlarged
dorsal body seta.

Phenacoccus stachyos Ehrh.; various modifications of the ducts,

PLATE Hf.

Antonina crawti Ckll.; dorsal aspect of posterior portion of abdomen of adult female.

Antonina crawii; dorsal aspect of posterior portion of abdomen of first stage larva.

Erium lichtensioides (Ckll.); anal lobe cerarius. r
Trionymus grindeliae n. sp.; anal lobe cerarius.

Trionymus smithii (Essig); anal lobe cerarius and half of anal ring.

Trionymus distichlii n. sp.; anal lobe cerarius and anal ring.

Ripersiella kelloggi Ehrh.; antenna of adult female.

Cryptoripersia salinus (Ehrh.); dorsal aspect of posterior end of abdomen.

Trionymus villosa (Ehrh.); anal lobe cerarius and anal ring.

Heterococcus arenae n. sp.; penultimate and anal lobe cerarii and half of anal ring.

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